SYSTEMATICS Species of Lygomusotima New and Neomusotima Yoshiyasu (: ) from and Southeastern Asia Feeding on Lygodium microphyllum (Schizaeaceae)

1 2 3 M. ALMA SOLIS, SHEN-HORN YEN, AND JOHN H. GOOLSBY

Ann. Entomol. Soc. Am. 97(1): 64Ð76 (2004) ABSTRACT Lygomusotima Solis & Yen, newgenus, and twonewspecies, stria and constricta, are described from Australia and southeastern Asia. L. stria was discovered feeding on Lygodium micro- phyllum (Cav.) R. Br. (Schizaeaceae) during exploration for biological control agents. Its immatures and biology are described. The newgenus is compared with Neomusotima conspurcatalis (Warren), newcombination, another species that wasdiscovered feeding on L. microphyllum. N. conspurcatalis is redescribed, and its immatures and biology are described for the Þrst time. fuscalis Snellen is designated as a newsynonym of N. conspurcatalis.

KEY WORDS Pyraloidea, Lygodium microphyllum, southeast Asia, Australia, pteridophagy

FROM 1997 TO 2002 EXPLORATION for biological control species of Neomusotima and Lygomusotima are pro- agents of Lygodium microphyllum, the Old World vided. climbing , has been conducted in Australia and Although pteridophagy is uncommon among Lep- southeast Asia (Fig. 1) by the Agricultural Research idoptera, there is a preponderance of fern-feeding Service Australian Biological Control Laboratory in records in . Previous explorations for Brisbane, . This fern is an invasive weed in biological control agents of invasive weeds have re- southern Florida where it threatens many wetland sulted in the discovery of fern-feeding musotimines, communities, including the Everglades (Pemberton for example, a species of Panotima Meyrick from South and Ferrier 1998); newevidence indicates that it may Africa was reared on bracken fern, Pteridium aquili- also threaten the citrus and timber industries (Wood num (L.) (Kuhn) (Dennstaedtiaceae) (Lawton et al. and Garcõ´a 2002). Three external fern-feeding Muso- 1988). Published historical musotimine feeding records timinae (Lepidoptera: Crambidae), tentatively re- until the late 1990s include Musotima dryopterisivora ferred to as Musotima conspurcatalis, Musotima spp., Yoshiyasu (1985) (orginally identiÞed as M. acclaralis and “Cataclysta” camptozonale were discovered and Walker) reared on Dryopteris prob. chinensis (Baker) tested for biological control purposes in Australia and Koidz. (Dryopteridaceae) in Japan (Nakamura 1977); Florida (J.H.G., T. Wright, M. Purcell, and J. Makin- Walker on Adiantum L. (Pteri- son, unpublished data). C. camptozonale belongs in a daceae) and Histiopteris incisa (Thunb.) J. Sm. new genus that will be treated elsewhere (Yen et al. (Dennstaedtiaceae) (Meyrick 1885; Philpott 1917); 2004); the other two species are treated in this article. and Musotima aduncalis Felder & Rogenhofer on Adi- We found that M. conspurcatalis should be transferred antum (Hudson 1928) from NewZealand. These his- to another genus, Neomusotima Yoshiyasu (1985), and torical records and recent research indicates that Mu- Musotima spp. (J.H.G., T. Wright, M. Purcell, and J. sotiminae may harbor more undiscovered fern Makinson, unpublished data) required a newgenus, feeders. Lygomusotima. In the process of searching collections Musotiminae consists of 20 genera and Ϸ200 named for additional specimens, another Lygomusotima spe- species occurring in all zoogeographical regions ex- cies was discovered from the Philippines. The imma- cept for Antarctica and the western Palaearctic. The tures of two species are described, and keys to the characters used to deÞne Musotiminae have yet to be fully investigated. But in general, the larvae of muso- 1 Systematic Entomology Laboratory, USDA, Smithsonian Institu- timines may be leaf miners or external feeders with a tion, P.O. Box 37012, National Museum Natural History, E-517, MRC 168, Washington, DC 20013Ð7012. E-mail: [email protected]. prognathous or semiprognathous head, and a round or 2 Department of Biological Sciences, Imperial College at Silwood dorsoventrally ßattened body, depending on their Park, Ascot, Berkshire SL5 7YP, UK, and Department of Entomology, feeding habit. Their pupae may have a smooth head The Natural History Museum, Cromwell Road, London SW7 5BD, with a medial dorsoventral depression or horned eye UK. 3 Australian Biological Control Laboratory, Agricultural Research caps; a pair of dorsolateral processes on the prothorax; Service, USDA, 120 Meiers Road, Indooroopilly, Queensland, Aus- spiracles normal or protruded on A1 to A4, or cone- tralia, 4068. shaped on A5 to A7; and cremaster present, usually January 2004 SOLIS ET AL.: FERN-FEEDING MUSOTIMINE PYRALOIDS 65

Fig. 1. Distributions of L. stria f, L. constricta Ⅺ, N. fuscolinealis Œ, and N. conspurcatalis F. with spiral spines. Several adult characters serve as will be discussed in a subsequent article (Yen et al. synapomorphies for Musotiminae, but the phyloge- 2004). netic signiÞcance of these characters has yet to be tested cladistically: 1) antennae laterally compressed Materials and Methods with ßattened segments, 2) R2 stalked with R3 ϩ 4,3) aedeagus with a reduced coecum, and 4) tympanal Biological Exploration Methods. Selection of search cases enlarged (Minet 1981, 1983, 1985; Yoshiyasu areas was guided by climate matching, habitat diver- 1985; Yen 1996, 1997; Munroe and Solis 1999; Solis et sity, herbarium locality data, and knowledgeable spe- al. 2004; S.-H.Y., unpublished data; Yen et al. 2004). cialists. The following countries were visited between Studies of , phylogenetic relationships, September 1997 and March 2002: Australia, China, and global diversity of Musotiminae are still in their Japan, Indonesia, Malaysia, NewCaledonia, Singa- infancy. Owing to the convergent wing patterns and pore, Taiwan, Thailand, and Vietnam. Each survey at genitalic structures, musotimines have been assigned each site was documented with a unique Australian to various crambid subfamilies, e.g., Nymphulinae, Biological Control Laboratory collection number, and Crambinae, Glaphyriinae, Pyraustinae, and Scopari- the following data were collected: date, time spent nae (Speidel 1984; Yoshiyasu 1985; Scoble 1992; Shaf- searching, site coordinates, temperature, plant growth fer et al. 1996; Munroe and Solis 1999; Speidel and Mey characteristics, and herbivore species. Most locations 1999). The phylogenetic relationships of Musotiminae were visited several times to account for seasonal within Crambidae are still in doubt. Yoshiyasu (1985) effects on herbivore biodiversity. Where possible, suggested that Musotiminae could be closely related other Lygodium species were surveyed for herbivores. to Scoparinae. Nuss (1999) suggested they could be Methods for collection included hand searches, closely related to Evergestinae. Recently, a prelimi- sweeping, beating of foliage, and dissection of under- nary cladistic analysis of the crambid subfamilies ground plant parts (J.H.G., T. Wright, M. Purcell, and based on adult characters also resulted in the unre- J. Makinson, unpublished data). solved relationship of Musotiminae to other subfam- Taxa Studied. Specimens were studied from the ilies (Solis and Maes 2002). A complete phylogenetic following collections, whose abbreviations are used analysis of the genera is lacking, and the relationship throughout the text: BMNH, The Natural History Mu- of Lygomusotima and Neomusotima to other genera seum, London, UK.; RMNH, National Natuurhisto- 66 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 1

Fig. 2. (A) Male adult of L. stria, Singapore, Bukit Timah Nature Preserve, Belukar. (B) Female adult of L. constricta, Philippines, Luzon, Mt. Makiling. (C) Female adult of N. conspurcatalis, Australia, Northern Territory, nr. Wangl, reared in Florida, Alachua Co., Gainesville, DPI, FBCL quarantine. (D) Live adult of N. conspurcatalis. risch Museum, Leiden, The Netherlands; and USNM, ton (1946), Klots (1970), Maes (1985, 1995, 1997), National Museum of Natural History, Smithsonian In- Yoshiyasu (1985), Phillips and Solis (1996), and Solis stitution, Washington, DC. Type material is deposited and Maes (2002). in these institutions as designated within the text. To determine the identity and/or placement of the taxa Results and Discussion feeding on L. microphyllum, the type species, and Lygomusotima Solis & Yen, New Genus other species if available, of all described genera cur- (Figs. 2A and B; 3A, B, and D; 4A, B, and D; rently in the Musotiminae, including several newun- 5A and B; 6AÐC; and 7AÐC) published genera by Solis et al. (2004), Yen et al. 2004 and S.-H.Y., unpublished data, were dissected, studied, Diagnosis. Area between postmedial and subtermi- and compared. nal line distinctively patterned with bright white tri-

Pinned specimens were examined and dissected angle between R5 and M1 that spans the two lines; after abdomens were soaked in 10% potassium hy- valva simple, sacculus slightly indented; juxta not in- droxide and wings soaked in bleach. Dissections were vaginated dorsally to sacculus. stained in chlorozal black for genitalia, and Eosin-Y for Description. Adult. Head. (Fig. 3A and B). Frons wings. Genitalia were placed in vials with glycerin and vertex white, brown between antennae; antennae and/or slide mounted in Canada balsam or Euparol white, brown dorsally; labial palpi three-segmented, (Clarke 1941; Holloway et al. 1987). Measurements Þrst segment white ventrally, brown dorsally, second were made with an ocular micrometer. Forewing segment mostly white ventrally, brown distally, third length is measured from the center of the axillar area segment brown, white distally; maxillary palpi four- to the apex of the forewing. Terminology follows Hin- segmented, white, all segments brown distally; fourth January 2004 SOLIS ET AL.: FERN-FEEDING MUSOTIMINE PYRALOIDS 67

Fig. 3. (A) Male head of L. stria, Singapore, Bukit Timah Nature Preserve, Belukar. (B) Female head, L. stria, Singapore, Sungai Buloh. (C) Male head of N. conspurcatalis, Indonesia, Sumatra, Tanah, Gambus Estate. (D) Tympanal organs of L. stria, Thailand, Luhbohlausa, Yi-ngo, Narathiwat, USNM slide #109,227. segment slightly longer and broader than other seg- proximal to postmedial line beige with scattered ments. Proboscis longer than labial palpi; scaled white brown-tipped scales; costa yellow or beige between basally. Antennae prismatic, laterally compressed. median and postmedial line; reniform spot white with Chaetosemata and ocelli absent. or without dark brown edge; area between postmedial Thorax (Figs. 2A and B, 4D). Thorax, patagium, and and subterminal line yellowdistinctively patterned tegula white, with brown-tipped scales. Forelegs with bright white between R2 and R3 ϩ 4 (streak), R5 white, tibia brown laterally; midleg white, brown at and M1 (triangle, spanning the two lines), and M3 and femur/tibia juncture; hindleg white, Þrst three tarsal 1Aϩ2A (longitudinal line of various thickness); sub- segments brown distally; midleg with one pair of tibial terminal line complete or incomplete, dark brown, spurs, hindleg with two pairs of tibial spurs. yellow distal to white areas; fringe double with short Forewing (Fig. 2A and B). Length 5.7 to 6.6 mm; white, brown-tipped scales and longer white scales margin incised or slightly crenulate. Costal swelling with black patches apically between M1 and M2 and absent. R1 from anterior margin of discal cell, R2, between CuA1 and CuA2. Underside beige with dark R3 ϩ 4, and R5 from anterior, distal apex of discal cell. brown lines slightly visible, and distal to postmedial Basal, median, and postmedial line dark brown; area line; color and pattern as on upperside of wing, but less 68 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 1 distinct. Retinaculum a short set of hooked setae lo- postvaginalis is sclerotized with a distinctive jagged cated basally on Cu. pattern, but in Lygomusotima the lamella postvaginalis Hindwing (Fig. 2A and B). Margin crenulate; beige is membranous. with scattered brown-tipped scales; medial and post- In the larvae of L. stria, the D and SD setal groups medial lines brown, barely visible; subterminal line have prominent chalazae on all abdominal segments, dark brown, yellow distally; fringe double with short but in N. conspurcatalis only A7Ð10 segments have white, brown-tipped scales and longer white scales prominent D and SD chalazae. In L. stria, the protho- with black patches at Þrst and second scallop. Under- racic shield has the anterior to lateral margin ridged side beige with dark brown lines slightly visible, and forming an L-shaped sclerotized pattern from the lat- distal to postmedial line; color and pattern as on up- eral view, but in N. conspurcatalis the margin is not perside of wing, but less distinct. Female frenulum ridged, and it forms a C-shaped sclerotized pattern. with two setae, male with one seta. The L group on A8 is trisetose in L. stria, and unisetose Abdomen. White with some brown-tipped scales. in N. conspurcatalis. The anal shield of L. stria has Tympanal organs (Fig. 3D): Tympanal cases (ϭbulla prominent D and SD chalazae and SD1 is on the outer tympani, caisses tympaniques) enlarged, tympanic margin, and in N. conspurcatalis the chalazae of D and frame, (ϭfornix tympani, cadre tympanique) highly SD are small and SD1 is not included on the anal sclerotized, processes tympani (ϭsailles tympa- shield. niques) absent, ramus tympani absent. In the pupae, the differences are striking. In L. stria Male Genitalia (Fig. 4A and B). Uncus elongate, the top of the head is truncate with a laterally pro- pointed, with short setae dorsally; gnathos (ϭpseu- truded clypeus and the pilifers are slightly visible. In dognathos of Maes 1997) with two lateral rows of N. conspurcatalis the top of the head is round with a sclerotized spinules dorsally, arms tapered to tegu- slightly protruded clypeus, and the pilifers are absent. men; tegumen continuous, forming an inverted “U” The prothorax of L. stria has no lateral projections, but with ventral sclerotized extension to costa of valva; in N. conspurcatalis paired prothoracic projections are juxta bifurcate, as long as vinculum, birfurcated pos- present. This structure is also shared by Musotima teriorly either less than or more than half the length drypterisivora (Nakamura, 1977; Fig. 9) and C. camp- of juxta; transtilla posteroventrally v-shaped with two tozonale. In L. stria, the proboscis extends to the mid- short lobes laterally, medially lightly sclerotized or dle of A4, but in N. conspurcatalis the proboscis only membranous; valva simple, apical margin rounded, extends to the middle of A1. In L. stria, the hind tarsus distally with short setae; sacculus simple, slightly in- is not visible, but it is in N. conspurcatalis. Dorsally, the dented medially; vinculum 3ϫ as long as tegumen. tergites of the abdominal segments and wing sheath Aedeagus simple without cornuti, distal half lightly are granulate in L. stria, but smooth in N. conspur- sclerotized; coecum absent. catalis. There is a paired lateral angulated protuber- Female Genitalia (Fig. 5A and B). Corpus bursae ance present along the margin of abdominal segment slightly rugose to distinctly rugose posteriorly and nine in L. stria, but in N. conspurcatalis the structure around signum, either slightly longer than seventh is more rounded. In L. stria, the cremaster has four segment or more than twice as long as seventh seg- pairs of curved setae, where as N. conspurcatalis has ment; signum a small circle dorsomedially; appendix two pairs of curved setae. bursae absent; ductus seminalis from ventroposterior area of corpus bursae; ductus bursae membranous, Type Species. L. stria Solis & Yen. constricted anteriorly or with sclerotized striae ex- panding in circumference anteriorly; ostium bursae Distribution (Fig. 1). Southeastern Asia from Sin- membranous; posterior apophyses either slightly gapore and Thailand of the Malay Peninsula, to the longer than anterior apophyses or apophyses of equal Philippine Islands and Pulo Laut, an island near the length; papillae analis dorsoventrally ßattened, in- southeastern corner of Borneo. dented mediodorsally. Etymology. The preÞx of the generic epithet is de- Comparison to other Taxa. In the adult forewing, rived from the generic name of the host, Lygodium, the area between the postmedial and subterminal line and “musotima” refers to its relatedness to this genus is distinctively patterned with a bright, white triangle in the subfamily. The gender of the genus is feminine. that spans R5 and M1 in Lygomusotima; in Neomuso- tima the bright white pattern is of varying thickness L. stria Solis & Yen, New Species between R and 1Aϩ2A and never spans the distance 2 (Figs. 2A; 3A, B, and D; 4A and D; 5A; 6AÐC; between the two lines. In the genitalia, Lygomusotima and 7AÐC) has a simple valva, with a round apical margin and the sacculus is membranous and slightly indented. In Diagnosis. Forewing (Fig. 2A) length mean 5.70 mm Neomusotima the apical margin of the valva can be (range 5.0Ð7.1 mm) (n ϭ 54); costa yellowbetween round or straight. The sacculus is sclerotized and mod- median and postmedial line, subterminal line dark iÞed with a thumblike structure or spine and the mar- brown, yellow distally. In the male genitalia (Fig. 4A) gin has an apical fold. Lygomusotima and Neomusotima juxta as long as vinculum, bifurcated posteriorly less both have a bifurcate juxta, but it is not invaginated than half the length of juxta. In the female genitalia dorsally to the sacculus. In the female genitalia of (Fig. 5A) ductus bursae with sclerotized striae ex- Neomusotima, the posterior margin of the lamella panding in circumference anteriorly; corpus bursae January 2004 SOLIS ET AL.: FERN-FEEDING MUSOTIMINE PYRALOIDS 69

Fig. 4. (A) Male genitalia of L. stria, Thailand, Luhbohlausa, Yi-ngo, Narathiwat, USNM slide #109, 227. (B) Male genitalia of L. constricta, Philippines, Luzon, Mt. Makiling, USNM slide #109,001. (C) Male genitalia of N. conspurcatalis, Australia, Northern Territory, 100 km SSE Darwin, Anniversary Creek, USNM slide #112,614. (D) Legs of L. stria, female, Singapore, Bukit Timah Nature Preserve, Belukar, USNM slide #110,450. 70 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 1

Fig. 5. (A) Female genitalia of L. stria, Singapore, Sungai Buloh, USNM slide #108,998. (B) Female genitalia of L. constricta, Philippines, Luzon, Mt. Makiling, USNM slide #109,001. (C) Female genitalia of N. conspurcatalis, Australia, Northern Territory, 100 km SSE Darwin, Anniversary Creek, USNM slide #112, 615. rugose, only slightly longer than seventh segment; Biology. Eggs are laid singly or in small clusters anterior apophyses and posterior apophyses of equal mostly on the newgrowth.Early instars skeletonize length. L. microphyllum leaves, whereas fourth instars con- Type Material. HOLOTYPE: 1 (, SINGAPORE: sume entire leaves. L. stria pupates above ground on Bukit Timah Nature Preserve, Belukar, 1Њ21.37Ј N the plant in a concealed location. The developmental 103Њ46.94Ј E, 1-III-2000, reared from larva feeding on period from egg to adult at 25ЊCisϷ25 d. This species pinnules of L. microphyllum, A.D. Wright (USNM). seems to be a specialist, having been collected only ALLOTYPE: 1 &, same data (USNM). from L. microphyllum. PARATYPES: 11 &,15(, same data (USNM; Egg. Translucent pale yellowin clusters. BMNH); 10 &,7(, SINGAPORE: Sungai Buloh, Larva (Fig. 6AÐC). Last instar length 13.0Ð13.5 mm 1Њ26.67Ј N 103Њ43.46Ј E, 10-X-99, reared from larva (n ϭ 27), cylindrical, tapered anteriorly and posteri- feeding on pinnules of L. microphyllum, A.D. Wright orly, prothoracic shield developed, primary setae (USNM); 1 (, Singapore, Swinhoe Collection, arising from distinct pinacula or chalazae (D and SD (BMNH); 1 &,3(, THAILAND: Luhbohlausa, groups). Head hypognathous, dark brown; epicranial Yi-ngo, Narathiwat, ex Lygodium microphyllum, 20- suture present; frontoclypeus and labrum dark brown, IV-99, A. Winotai; 2 &, Thailand, Nakhon Si Tham- Ϸ1.25ϫ as long as wide; six stemmata, C1 and C2 marat Prov., Sichol Dist., nr. Tech. School, 8Њ57.29Ј N approximate, C3 and C4 approximate, C5 anteroven- 99Њ53.92ЈE, ex larva on L. microphyllum, 15-XI-99, A. D. trad to C6; S2 1.3 ϫ as long as S1, SS3 longer than SS2, Wright (USNM); 1 ( (abdomen missing), MG1 short, F1 present near middle of frons, AF1 and MALAYSIA: Sarawak, Kuching (BMNH); 1 &,1(, AF2 on adfrontal area rather short, La and Aa present, INDONESIA: Borneo, Pulo Laut, Doherty (BMNH); P1 longer than P2, P2 base slightly elevated; labrum 1 &, Indonesia, Java, Prov. Pasuruan, Kalipari, 300Ð500 with three setae on each side externally and no seta on feet. W. Doherty 1891 (BMNH). each side internally; mandible with one row of serra- Distribution (Fig. 1). Southeast Asia from Singa- tions. T1Ð3 and A1Ð10 integument smooth, all seg- pore and Thailand of the Malay Peninsula, and Pulo ments with prominent chalazae of D and SD setal Laut, an island near the southeastern corner of Borneo. groups and small pinacula, setae with concolorous Etymology. The speciÞc name is derived from the pinacula. Prothoracic shield approximately twice as striae in the female ductus bursae. long as wide, anterior to lateral margin ridged forming January 2004 SOLIS ET AL.: FERN-FEEDING MUSOTIMINE PYRALOIDS 71

Fig. 6. (A) Lateral larval setal map of L. stria. (B) Frontral viewof L. stria larval head. (C) Lateral viewof L. stria larval head. (D) Lateral larval setal map of N. conspurcatalis. (E) Frontral viewof N. conspurcatalis larval head. (F) Lateral view of N. conspurcatalis larval head. an L-shaped sclerotized pattern from lateral view, D2 T2Ð3; L group bisetose, L2 1.5ϫ as long as L1; SV2 2.5 ϫ as long as D1, all of XD1, XD2, SD1, and SD2 ventrad to SV1, approximately equal in length; V1 located on prominent chalazae, XD1 very short, SD1 present, Ϸ1/3ϫ length of SV. Setae of A3Ð6 in general 3ϫ as long as SD2; L setae bisetose, anterior to spiracle, similar to those on A1Ð2, but D and SV group located L1 and L2 on same elongate pinaculum; SV setae on chalazae, L3 present, SV group trisetose at base of bisetose, on different small pinacula. T2-T3 with D prolegs. A7 with L2 much longer than those on A3Ð6. setae bisetose on a developed chalaza, D2 Ϸ2ϫ as long A8 with D1 and D2 located on conical chalazae; L as D1; SD setae bisetose on a similar chalaza as D setae; group trisetose, nearly equal in length. A9 with D1 on L setae trisetose, L2 on a larger pinaculum, Ϸ3ϫ as a large conical chalaza; SD group unisetose; L group long as L1 and L3; SV1 present, shorter than L2; V seta unisetose. Anal shield with prominent D and SD cha- short. A1-A2 with D setae very short on separate small lazae and D2, SD1, and SD2 at outer margin of shield. pinacula; SD group unisetose, SD1 as long as L2 of Prolegs with crochets biordinal in a complete circle. 72 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 1

Fig. 7. (A) Dorsal viewof L. stria pupa. (B) Ventral viewof L. stria pupa. (C) Lateral viewof L. stria pupa. (D) Dorsal viewof N. conspurcatalis pupa. (E) Ventral viewof N. conspurcatalis pupa. (F) Lateral viewof N. conspurcatalis pupa.

Pupa (Fig. 7AÐC). Length 5.5Ð6.5 mm (n ϭ 10). cremaster somewhat ßattened dorsoventrally with a Ventrally top of head truncate with a laterally pro- sharp bifurcate apex and four pairs of curved setae. truded clypeus; labrum narrow; pilifers slightly visible; maxillary palpi absent; labial palpi reaching middle of L. constricta Solis & Yen, New Species A3; proboscis extending to middle of A4; forefemur (Figs. 2B, 4B, and 5B) visible, prothoracic legs three-fourths length of fore- wing, reaching to middle of A3; mesothoracic legs Diagnosis. Forewing (Fig. 2B) length mean 6.6 mm extending beyond hind margin of A4, slightly longer (range 6.0Ð7.0 mm) (n ϭ 3); subterminal line not than antennal sheath; hind tarsi not visible; abdominal complete, dark brown and yellow distally distal to segments Þve and six with vestigial prolegs; abdominal white areas. In the male genitalia (Fig. 4B) juxta as segments 8Ð10 with genital oriÞce anteriorly; anus long as vinculum, bifurcated posteriorly more than posteriorly with lateral conical depressions. Dorsally half the length of juxta. In the female genitalia (Fig. prothorax smooth, with two dorsolateral hornlike struc- 5B) ductus bursae membranous and constricted an- tures that protrude only slightly; mesothorax without teriorly; corpus bursae only slightly rugose posteriorly seta; all spiracles on abdominal segments located on de- and around signum, more than twice as long as veloped conical protuberances; surface of all abdominal seventh segment. tergites and wing sheath granulate; a paired lateral pro- Type Material. HOLOTYPE: 1 (, PHILIPPINES: tuberance present along margin of abdominal segment 9; Luzon, Mt. Makiling, Baker (USNM). January 2004 SOLIS ET AL.: FERN-FEEDING MUSOTIMINE PYRALOIDS 73

ALLOTYPE: 1 &, same data (USNM). Forewing (Fig. 2C and D). Length mean 4.5 mm PARATYPES: 1 &, same data (USNM); 1 &, PHIL- (range 4.0Ð5.0 mm) (n ϭ 7); margin incised or slightly

IPPINES: Luzon, Palili, Benguet, 28 December 1912, crenulate. Costal swelling absent. R1 from anterior Ј & 2000 , gen. slide #21194, A. E. Wileman (BMNH); 1 , margin of discal cell, R2,R3 ϩ 4, and R5 from anterior, Philippines, Luzon, Klondyke, Benguet, 27 March, distal apex of discal cell. Basal, median, and postmedial 1912, 800Ј, A. E. Wileman (BMNH). line dark brown, area proximal to postmedial line Distribution (Fig. 1). Known only from Luzon in beige with scattered brown-tipped scales, reniform the Philippines. spot white, area between postmedial and subterminal Etymology. The speciÞc name is derived from the line yellow, distinctively patterned with bright white ϩ anterior constriction of the ductus bursae in the fe- of varying thickness between R2 and 1A 2A, never male. spanning distance between two lines; subterminal line Biology. Not known, but probably feeds on Lygo- dark brown, yellow distally; fringe double with short dium. brown scales and longer white scales with brown

patches apically, between M1 and M2, and between CuA and CuA . Underside beige with dark brown Key to species of Lygomusotima Solis & Yen 1 2 lines slightly visible, and distal to postmedial line color 1. Frenulum with one seta, male ...... 2 and pattern as on upperside of wing but markings less 1Ј. Frenulum with two setae, female ...... 3 distinct. Retinaculum short hooked setae located 2. Bifurcation more than half as long as juxta basally on Cu. (Fig. 4a) ...... L. stria Hindwing (Fig. 2C and D). Scattered with white 2Ј. Bifurcation less than half as long as juxta brown-tipped scales; medial and postmedial lines (Fig. 4b) ...... L. constricta brown, barely visible; subterminal line not marked; 3. Ductus bursae with sclerotized striae and ante- margin crenulate; with eight white and dark brown rior expansion to corpus bursae (Fig. 5a) . . spots along scallops; fringe double with short brown ...... L. stria scales and longer white scales with black patches at 3Ј. Ductus bursae membranous with anterior con- Þrst and second scallop. Underside beige with dark striction to corpus bursae (Fig. 5b) ...... brown lines and spots slightly visible, similar to up- ...... L. constricta perside markings but less distinct. Female frenulum with two setae, male with one seta. Abdomen. White with brown-tipped scales. Tym- N. conspurcatalis (Warren), New Combination panal organs: tympanal cases (ϭbulla tympani, caisses (Figs. 2C and D 3C, 4C, 5C, 6DÐF, and 7DÐF) tympaniques) enlarged, tympanic frame (ϭfornix Ambia conspurcatalis Warren, 1896: 202. Holotype, tympani, cadre tympanique) membranous, processes BMNH, without abdomen, examined, type locality: tympani (ϭsaillies tympaniques) absent, ramus tym- Khasis. pani absent. Musotima fuscalis Snellen, 1900: 298. Lectotype, Male Genitalia (Fig. 4C). Uncus elongate, pointed, RMNH, examined, type locality: Assam, newsyn- with few short setae dorsally, 1.5 ϫ longer than tegu- onym; Klima, 1937: 104; Munroe et al., 1958: 75, lec- men; gnathos (ϭpseudognathos of Maes 1997) with totype designation. scattered sclerotized spinules dorsally, arms tapered Diagnosis. Male genitalia with apical margin of to junction with tegumen; tegumen continuous, form- valva straight, sacculus sclerotized with a highly scle- ing an inverted “U” with ventral sclerotized extension rotized spine apically and margin with apical fold. to costa of valva; juxta simple, almost as long as vin- Female genitalia with a distinctive sclerotized pattern culum; transtilla straight, lightly sclerotized or mem- on the posterior margin of lamella postvaginalis. branous medially; valva apical margin straight, saccu- Redescription. Adult. Head (Fig. 3C). Frons and lar margin with apical fold; sacculus sclerotized with vertex with brown-tipped white scales; antennae a highly sclerotized spine apically; vinculum four x as white, each segment distally with brown-tipped long as tegumen. Aedeagus simple without cornuti, scales; labial palpi three-segmented, all segments distal half lightly sclerotized; coecum present and white ventrally, white with brown-tipped scales straight. dorsally; maxillary palpi four-segmented, white with Female Genitalia (Fig. 5C). Corpus bursae mem- brown-tipped scales; fourth segment slightly longer branous, equal in length to seventh and eighth seg- and broader than other segments. Proboscis longer ment; signum absent; appendix bursae absent; ductus than labial palpi; scaled white basally. Antennae pris- seminalis from ventroposterior area of corpus bursae; matic, laterally compressed. Chaetosemata absent and ductus bursae membranous; ostium bursae membra- ocelli very small. nous; lamella postvaginalis with a distinctive sclero- Thorax (Fig. 2C and D). Dorsally white with tized pattern and folded between seventh and eighth brown-tipped scales; patagium and tegulae white with segment; anterior apophyses slightly longer than pos- some brown-tipped scales. Forelegs white, tibia, fe- terior apophyses; papillae analis dorsoventrally ßat- mur, and, Þrst tarsal segment brown laterally; midlegs tened, not indented mediodorsally. and hindlegs white, femora and tibiae brown; midlegs Biology. Eggs are laid singly or in small clusters with one pair of tibial spurs, hindlegs with two pairs of mostly on the newgrowth.Early instars skeletonize tibial spurs. L. microphyllum leaves, whereas fourth instars con- 74 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 1 sume entire leaves. N. conspurcatalis pupates above cated on developed conical protuberances; tergites of ground on the plant in a concealed location. The all abdominal segments smooth; cremaster somewhat developmental period from egg to adult at 25ЊCis ßattened dorsoventrally with a sharp bifurcate apex Ϸ25 d. This species has been collected only from and two pairs of curved setae. L. microphyllum, so it seems to be a specialist on this Material Examined. INDIA: Sikkim, 2 (,J.G. fern. Pilcher (BMNH); Sangir: 1 (, II-III-1893, Doherty Egg. Translucent pale yellowin clusters. (BMNH); 1 (, FebruaryÐMarch 1892, Doherty Larva (Fig. 6DÐF). Last instar length mean 4.63 (BMNH); Khasis Hills, Assam, 1 ( (BMNH); Khasis: mm (n ϭ 31), cylindrical, tapered anteriorly and pos- 3 (, I-1895, Nat. Coll. (BMNH); 2 (, II-1895, Nat. Coll. teriorly, prothoracic shield developed, primary setae (BMNH); 1 (, Sept 1894, Nat. Coll.; 2 (, III-1894, Nat. arising from distinct pinacula or chalazae (D and SD Coll. (BMNH); Assam, 1 (, Hamilton (RMNH, lec- groups). Head hypognathous, dark brown; epicranial totype of M. fuscalis); INDONESIA: 4 &,1(, Sumatra, suture present; frontoclypeus and labrum dark brown, Tanah, Gambus Estate, 3Њ14.24ЈN99Њ24.20ЈE, 20-V-99, Ϸ1.2x as long as wide; six stemmata, C1 and C2 ap- LEP: Pyralidae reared from larva feeding on L. micro- proximate, C3 and C4 approximate, C5 anteroventrad phyllum, A.D. Wright (USNM); 1 ( (no abdomen), to C6; S2 longer than S1, SS3 longer than SS2, MG1 Sulawesi, S Celebes, AugustÐSept Õ91, W. Doherty short, F1 present near middle of frons, AF1 and AF2 (BMNH); 1 (, Java, Tegal, Kemanglen, VIII, Lucassen on adfrontal area rather short, La and Aa present, P1 (RMNH, paralectotype of M. fuscalis); EAST TIMOR: longer than P2, P2 base slightly elevated; labrum with 2 &,2(, Dili, V-1892, W. Doherty (BMNH); three setae on each side externally and no seta inter- AUSTRALIA: Northern Territory: 3 (, 100 km SSE nally; mandible with one row of serration. T1Ð3 and Darwin, Anniversary Creek, 13Њ19.28ЈS 131Њ08.21ЈE, A1Ð10 integument smooth, A7Ð10 segments with 27-IV-99, LEP: Pyralidae reared from larva feeding on prominent D and SD chalazae. Prothoracic shield pinnules of L. microphyllum, J. Goolsby (USNM); 1 &, Ϸ1.5ϫ as long as wide, anterior to lateral margin not nr. Wangl, on L. microphyllum, reared in Florida, Ala- ridged, forming a C-shaped sclerotized pattern from chua Co., Gainesville, DPI, FBCL quarantine, 25-IX- lateral view, D2 2.5ϫ as long as D1, XD1 very short, 01, G. R. Buckingham (USNM); 1 (, same date except SD1 3ϫ as long as SD2; L setae bisetose, anterior to 23-I-02. spiracle, L1 and L2 on same elongate pinaculum; SV Distribution (Fig. 1). Currently known from India, setae bisetose, on separate small pinacula. T2-T3 with Indonesia, East Timor, and Australia. D setae bisetose on separate pinacula, D2 Ϸ2ϫ as long as D1; SD setae bisetose on the same pinaculum; Discussion L setae trisetose, L2 on a larger pinaculum, Ϸ3ϫ as long as L1 and L3; SV1 present, shorter than L2; V seta N. conspurcatalis, sister species of N. fuscolinealis, short. A1-A2 with D setae very short on separate small has not been reported in the literature other than its pinacula, D1 pinaculum longer than that of D2; SD original description and those of its synonym, M. fus- group unisetose, SD1 as long as L2 of T2Ð3; L group calis. The holotype and other specimens were discov- bisetose, L2 1.5ϫ as long as L1; SV2 ventrad to SV1, ered in the BMNH as Ambia Walker, the original genus approximately equal in length; V1 present, Ϸ1/3ϫ for conspurcatalis and at one time a “dump” genus for length of SV. Setae of A3Ð6 in general similar to A1Ð2 almost all species of Musotiminae. The abdomen of the except L group bisetose on the same pinaculum, L3 holotype is missing, but we dissected several other present, SV group trisetose at base of prolegs. A7 and specimens collected from the same locality by the A8 with D setae on conical chalazae. A9 with D1 on a same collector. large conical chalaza; SD group unisetose; L group Neomusotima was described by Yoshiyasu (1985) unisetose. Anal shield with small D and SD chalazae, for N. fuscolinealis from Japan; subsequently, a new but SD1 not included in shield. Prolegs with crochets distribution record from Kurashiki was reported by biordinal in a complete circle. Uno (1992). N. fuscolinealis is known so far only from Pupa (Fig. 7DÐF). Length 5.0Ð5.5 mm (n ϭ 25). Japan and Hong Kong (Fig. 1). We discovered and Ventrally top of head round with slightly protruded examined four other specimens, one collected in Ja- clypeus and dorsal emargination at middle; labrum pan (Okinawa: Tengan, Ryukyu Is., 6Ð14-VII-1966, narrow; pilifers and maxillary palpi absent; labial palpi J.F.G. Clarke, USNM), and three collected in Hong reaching A1; proboscis extending to middle of A4; Kong ( Peak, VIII-1993, A. C. Galsworthy, forefemora visible, prothoracic legs three-fourths BMNH), a newdistribution record. N. fuscolinealis has length of forewing, reaching to middle of A3; meso- been reared from L. japonicum (Thunb.) Sw. in Tokyo thoracic leg extending to hind margin of A4, slightly and Tsu City, Japan by various researchers (Pember- longer than antennal sheath; hind tarsus visible, just ton et al. 2002). slightly beyond length of forewing; abdominal seg- Neomusotima fuscolinealis and N. conspurcatalis ments Þve and six with vestigial prolegs; abdominal share the following: a distinctive pattern of white on segments 8Ð10 with genital oriÞce anteriorly; anus forewing between the postmedial and subterminal posteriorly with lateral conical depressions. Dorsally line, a vinculum that is longer than the tegumen, prothorax smooth, with two dorsolateral hornlike valvae with a well-developed sacculus, both species structures that protrude only slightly; mesothorax with a large projection posteriorly, and a well-devel- without seta; all spiracles on abdominal segments lo- oped coecum in the aedeagus. In the female genitalia, January 2004 SOLIS ET AL.: FERN-FEEDING MUSOTIMINE PYRALOIDS 75 both species share a well-developed lamella postvagi- Lawton, J. H., V. K. Rashbrook, and S. G. Compton. 1988. nalis with minute spinules. Their distributions are al- Biocontrol of British bracken: the potential of two lopatric, N. fuscolinealis is the northern species and from South Africa. Ann. Appl. Biol. 112: 479Ð490. N. conspurcatalis is southern. Both N. conspurcatalis Maes, K.V.N. 1985. A comparative study of the abdominal and N. fuscolinealis feed on species of Lygodium. tympanal organs in Pyralidae (Lepidoptera). I. Descrip- tion, terminology, preparation technique. Nota Lepidop- Key to species of Neomusotima Yoshiyasu terol. 8: 341Ð350. Maes, K.V.N. 1995. A comparative morphological study of 1. Frenulum with one seta, male ...... 2 the adult Crambidae (Lepidoptera, Pyraloidea). Ann. 1Ј. Frenulum with two setae, female ...... 3 Soc. R. Entomol. Belg. 131: 383Ð434. 2. Sacculus with a highly sclerotized spine apically Maes, K.V.N. 1997. On the morphology of the gnathos in the (Fig. 4c) ...... N. conspurcatalis Pyraloidea (Lepidoptera). Entomol. Scand. 38: 381Ð390. 2Ј. Sacculus with a membranous, thumblike pro- Meyrick, E. 1885. Descriptions of NewZealand Microlepi- cess...... N. fuscolinealis doptera. 6. Pyralidina. Trans. Proc. N.Z. Inst. 17: 121Ð140. 3. Corpus bursae without appendix bursae Minet, J. 1981. Les Pyraloidea et leurs principales divisions (Fig. 5c) ...... N. conspurcatalis syste´matiques. Bull. Soc. Entomol. France (N.S.) 86: 262Ð 280. 3Ј. Corpus bursae with small appendix bursae . . . Minet, J. 1983. E´ tude morphologique et phyloge´ne´tique ...... N. fuscolinealis des organs tympanique des Pyraloidea. 1. Ge´ne´ralite´s et homologies. (Lep. Glossata). Ann. Soc. Entomol. Acknowledgments France (N.S.). 19: 175Ð207. Minet, J. 1985. E´ tude morphologique et phyloge´ne´tique des We thank the following for access to pyraloid collections: organes tympaniques des Pyraloidea. 2. Pyralidae, Cram- Kevin Tuck and Michael Shaffer (BMNH) and Erik J. van bidae, premie`re partie (Lepidoptera Glossata). Ann. Soc. Nieukerken (RMNH). We especially thank Linda Lawrence Entomol. France (N.S.). 21: 69Ð86. (SEL, USDA) who photographed the adults and their inter- Munroe, E., Diakonoff, A., and E. L. Martin. 1958. Cata- nal structures, enhancing them with Adobe Photoshop and logue of SnellenÕs types of Pyralidae, with selections of Jon Lewis (SEL, USDA) for dissections. We thank Jeff lectotypes. Tijdschrift voor Entomol. 101: 65Ð88. Makinson and Ryan Zonneveld (Commonwealth ScientiÞc Munroe, E., and M. A. Solis. 1999. The Pyraloidea, pp. 233Ð and Industrial Research Organization Entomology, Austra- 256. In Handbuch der zoologie, band IV, Arthropoda: lian Biological Control Laboratory, Indooroopilly, Queens- Insecta Teilband 35, Lepidoptera, moths and butterßies, land) for rearing and curation, and Tony Wright (Common- vol. 1: Evolution, systematics, and biogeography, N. P. wealth ScientiÞc and Industrial Research Organization Kristensen [ed.], Walter de Gruyter, Berlin, Germany. Entomology) for collecting in southeast Asia. We also thank Nakamura, M. 1977. Notes on the immature stage of Muso- G. R. Buckingham and B. Pemberton (Invasive Plant Re- tima acclaralis Walker (Lep.: Pyralidae). Kita-Kyushu no search Laboratory, ARSÐUSDA, Gainesville, FL) for provid- Konchu 23: 32Ð36. ing laboratory reared material for study. We are grateful to Nuss, M. 1999. Revision der Gattungen der Scopariinae, Anthony C. Galsworthy (BMNH), Roger C. Kendrick (Hong Lepidoptera: Pyraloidea, Crambidae. Nova Suppl. Kong University), and Yutaka Yoshiyasu (Kyoto Prefectural Entomol. 13: 1Ð151. 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