PROC. ENTOMOL. SOC. WASH. 105(4), 2003, pp. 1043±1050

STUDIES ON NEW GUINEA . 2. DESCRIPTION OF A NEW SPECIES OF XENOTHICTIS MEYRICK (: : )

JOHN W. B ROWN,SCOTT E. MILLER, AND MARIANNE HORAK

(JWB) Systematic Entomology Laboratory, Plant Sciences Institute, Agricultural Re- search Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Washington, DC 20560-0168 (e-mail: [email protected]); (SEM) Department of Systematic Biology, National Museum of Natural History, Washington, DC 20560-0105; (MH) Australian National Collection, CSIRO Entomology, Canberra, ACT 2601, Australia

Abstract.ÐXenothictis gnetivora, new species, from Papua New Guinea, is described and illustrated based on morphological characters and DNA barcode. The type series consists of 124 specimens reared from leaf-rolling larvae primarily on Gnetum gnemon L. (Gnetaceae), but also on Celtis philippensis Blanco (Celtidaceae), Sterculia schuman- niana (Laut.) Mildbr. (Sterculiaceae), and other plants during a multi-year rearing project (1995±2001). The ®ve previously described species of Xenothictis are from Australia and Melanesia. Key Words: Malesia, Melanesia, Papua New Guinea, genitalia, pupa, Gnetum gnemon, Celtis philippensis, Sterculia schumanniana

The archipine genus Xenothictis Meyrick ] that appears to have radiated includes ®ve previously described species within New Caledonia and spread to other distributed from Australia north and east to areas of the Paci®c. The species are as- New Caledonia, Lifou (northeast of New signed at present to the closely related gen- Hebrides), Vanuatu (New Hebrides), and era Xenothictis and Xeneda and are being Fiji (see Appendix). Beyond brief original studied by Mr. K. Tuck [The Natural His- descriptions of the species, little has been tory Museum, London] . . . who has sug- published on the genus. Clarke (1958) il- gested they might be congeneric. The two lustrated the adult and male genitalia of X. genera together contain at least six species atri¯ora Meyrick and its synonym, X. me- in New Caledonia, two in the Loyalties and lananchis (Meyrick). Diakonoff (1961) il- one in each of the New Hebrides and Fiji.'' lustrated the adult and male genitalia of X. In addition, there are several undescribed nocti¯ua Diakonoff, presented brief com- species from New Caledonia in the collec- ments on Meyrick's (1910) description of tion of the National Museum of Natural the genus, and commented on the relation- History, Smithsonian Institution. ship with his new genus Xeneda Diakonoff. During a recent (1995±2001) insect ecol- Horak et al. (1996) transferred Barnardiella ogy project in Papua New Guinea (Basset sciaphila Turner to Xenothictis and synon- et al. 2000; Novotny et al. 2002a, b, c), a ymized Barnardiella Turner with the latter. large series of an undescribed species of Holloway (1979: 225) provided the fol- Xenothictis was reared from several plants. lowing comment: ``There is one group [of The purposes of this paper are to name the 1044 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1±2. Male and female of Xenothictis gnetivora. 1, Male, 2, Female. new species, provide descriptions and illus- meter under low power of a Leica MZ12᭧ trations of adult facies, male and female dissecting microscope. Terminology for genitalia, and immatures, and to present a genitalia structures follows Horak (1984). list of the described species of the genus. Cytochrome oxidase I (COI) sequences, Although considerable material, including DNA barcodes of Hebert et al. (2003), fol- other new species, has accumulated in ma- low the protocols in Hebert et al. (2003). jor museums worldwide, it is not within the SYSTEMATICS purview of this paper to review this infor- mation or describe additional new species. Xenothictis gnetivora Brown, Miller, and We have not seen any other species of Xen- Horak, new species othictis from New Guinea. (Figs. 1±9) Diagnosis.ÐSuper®cially, adults of X. MATERIALS AND METHODS gnetivora are characterized by a grayish General ®eld and laboratory methods for brown forewing, ®nely and faintly reticu- the project are described in Miller et al. lated with pale reddish brown throughout. (2003) and Novtony et al. (2002a, b, c). We Males have a bold, contrasting, blackish examined 124 pinned specimens of the new brown, short, slightly curved fascia origi- species, all of which were obtained during nating near mid-dorsum, terminating near the rearing project in Papua New Guinea. the middle of the discal cell; females lack Text descriptions are composite, based on the fascia. The male genitalia of X. gneti- all available specimens. Forewing measure- vora are distinguished by the narrow tegu- ments were made with an ocular micro- men, simple, obovate valva with costa and VOLUME 105, NUMBER 4 1045 ventral margin subparallel and with ex- late, arising relatively close because of nar- tremely slender sacculus not separated by a row tegumen; slightly and gradually con- shallow emargination from rest of valva, vergent distally; slightly upturned and slender gnathos arms with cross-bar close joined subdistally by transverse bar, free to tips, a narrowly ovate juxta, and a long, points short. Transtilla slender, extremely slender aedeagus with 1±2 cornuti. Among weak. Valva moderately long, obovate with the tortricids reared by the project team costa and ventral margin roughly parallel, near Madang, it is distinguished by the dif- attenuate distally, with ®ne hairs throughout ferential coloration of the fore- and hind- middle portion; sacculus simple, narrow, wings, maculation of the forewing, and the con®ned to ventral edge, extending ca. ½ large, shiny scales on the abdomen in both length of valva. Juxta a narrowly ovate 9 sexes. plate. Aedeagus slender, ca. ⁄10 length of 1 Description.ÐMale. Head: Lower frons valva, weakly curved, attenuate in distal ⁄10; pale tan to buff; upper frons slightly darker; vesica with one or two long, slender cor- vertex with overhanging tuft of scales short. nuti. Labial palpus short, ca. 1.0 times horizontal Female. Head: Essentially as described diameter of compound eye, robust, third for male, except antennal cilia short, un- segment very short; pale tan to buff mes- modi®ed. Thorax: Buff; tegula large, pale ally, pale reddish brown laterally. Antenna grayish brown. Forewing (Fig. 2) length with two rows of dorsal scales per ¯agel- 7.9±9.6 mm (xÅ ϭ 8.7 mm; n ϭ 10). Ground lomere, tan; cilia long, ca. 1.2±1.5 times color grayish brown, ®nely and faintly re- width of ¯agellomere. Chaetosema well de- ticulated with pale reddish brown through- veloped. Ocellus moderately large. Probos- out; 4±5 extremely short costal strigulae in cis present, presumably functional. Thorax: basal ½; a small brown spot near mid-costa, Buff; tegula large, pale grayish brown. an additional ill-de®ned spot ca. ¾ distance Forewing (Fig. 1) length 6.5±8.1 mm (0 ϭ from base to apex. Fringe concolorous with 7.7 mm; n ϭ 10). Ground color grayish ground color. Abdomen: Genitalia (Fig. 4; brown, ®nely and faintly reticulated with drawn from USNM slide 82244; 6 prepa- pale reddish brown throughout; a bold, rations examined). Papillae anales simple, short, slightly curved, blackish brown fas- slender slipper-shaped, unmodi®ed. Sterig- cia originating near mid-dorsum, terminat- ma an extremely slender, broadly U-shaped ing near middle of discal cell; 4±5 extreme- band, sometimes with patch of sclerotiza- ly short costal strigulae from base to mid- tion immediately posterad of ostium. Duc- costa; a small brown spot near mid-costa, tus bursae extremely long, narrow, gradu- an additional ill-de®ned costal spot ca. ¾ ally widening into corpus bursae; collicu- distance from base to apex; costal fold ab- lum present immediate anterad of ostium; sent; without modi®ed scales on underside. an irregularly-shaped sclerotized patch ca. Fringe concolorous with ground color. ⅓ distance from ostium to junction of duc- Hindwing uniformly gray brown, without tus bursae and corpus bursae, at which modi®ed scales. Abdomen: Pale gray point the ductus bursae is curved or weakly brown. Genitalia (Fig. 3; drawn from bent. Corpus bursae simple, ovoid, non- USNM slide 92771; 4 preparations exam- spiculate, with signum typically archipine ined) with tegumen narrow; a long, narrow, with a long, strong, internal curved spine medial spine extending anterad from dorsal from large, strongly sclerotized, elongate fusion of lateral halves of posterior portion diamond-shaped process on the surface of of tegumen. Uncus moderately slender, bi- the corpus bursae. furcate from base. Socius short, ca. ½ Pupa (Fig. 5). Typically tortricine with- length of uncus, pendant, weakly setose. out modi®ed head (Sohn 2002); no con- Gnathos slender, arms long, weakly undu- spicuous sculpturing; abdomen with one 1046 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 3±5. Genitalia and pupa of Xenothictis gnetivora. 3, Male genitalia, with valva spread and aedeagus removed (below genitalia capsule), 4, Female genitalia, 5, Pupa, venter on left, dorsum on right. row of spines dorsally on A3 and two rows posterior edge of segment; cremaster well on A4±8 in both sexes, posterior row con- developed with lateroposterior horns; 4 spicuously weaker, extremely weak on A4; pairs of long, hooked bristles, two laterally spines on A9 as a small irregular patch at and two mesally. VOLUME 105, NUMBER 4 1047

Gene sequence.ÐA male and a female, dang, and 195727 and 195728 are in the specimens 109973 and 120720, possessed Australian National Insect Collection, CSI- identical COI-5Ј sequences (DNA bar- RO, Canberra. Paratypes will be deposited codes), which have been deposited in in the following institutions: Bishop P. Ber- GenBank under the accession numbers enice Museum, Honolulu, Hawaii, U.S.A.; AY313945 and AY313944, respectively. The Natural History Museum, London, Type material.ÐHolotype, (, Papua England; Australian National Insect Collec- New Guinea, Madang Province, Ohu Vil- tion, CSIRO, Canberra; Nationaal Natu- lage, 5 July 1999, reared from Gnetum gne- urhistorisch Museum, Leiden, The Nether- mon, by the project team (Novotny, Basset, lands; National Agriculture Research Insti- Cizek, Auga, Boer, Dal, Hiuk, Isua, Kasbal, tute, Port Moresby, Papua New Guinea; Kutil, Manumbar & Molem). Deposited in Museum Zoologi Bogor, Cibinong, Indo- National Museum of Natural History, nesia; and National Museum of Natural Washington, D.C. Project specimen History, Smithsonian Institution, Washing- 107091. ton, D.C., U.S.A. Paratypes, 51 (,72&. Papua New Guin- Distribution and biology.ÐAll speci- ea, Madang Province, Mis, Baitabag, and mens are from a 25 km2 area, including the Ohu Villages, reared during 1995 to 2001 villages of Mis, Baitabag, and Ohu, near by the project team. All labeled with project Madang in Madang Province, Papua New morphospecies TORT039 (or one of the Guinea. Our project encountered 224 larvae synonyms, TORT101 and TORT119), and and successfully reared 124 adults (see including the following project specimens: Miller et al. 2003 and Novotny et al. 2002a 64808, 86216, 86817, 88484, 88170, for methods). The species feeds primarily 87565, 87554, 87700, 87545, 87719, on Gnetum gnemon (n ϭ 188) (Figs. 6±7), 87631, 87695, 87672, 87285, 87480, Sterculia schumanniana (Laut.) Mildbr. 87292, 87454, 109523, 109529, 109527, (Stericuliaceae) (n ϭ 25), and Celtis philip- 109538, 109521, 109542, 109130, 109317, pensis Blanco (Celtidaceae) (n ϭ 5). We 107600, 107359, 107596, 107461, 107454, also reared adults twice from Ficus nodosa 107370, 107419, 107598, 107323, 107364, Teysm. & Binn (Moraceae) and once each 107365, 107470, 108953, 108955, 108867, from Ficus variegata Blanco, Artocarpus 108915, 108857, 107931, 107987, 107870, communis J. R. Forst. & G. Forst. (Mora- 107864, 108796, 107829, 108918, 107849, ceae), Psychotria micralabastra (Laut. & 106999, 106960, 106990, 106973, 107037, Schum.) Val. (Rubiaceae), and Leucosyke 106938, 106834, 108459, 108262, 108482, capitellata (Poir.) Wedd. (Urticaceae). Be- 109973, 108367, 109995, 108585, 108579, cause of the low occurrence of larvae com- 108454, 108945, 121278, 121049, 120702, pared to the density of sampling, we do not 121101, 120737, 120577, 120995, 121425, consider the last ®ve ``normal'' hosts (see 120977, 120869, 122699, 122951, 122449, Novotny et al. 2002a for discussion). Lar- 122346, 121662, 122474, 122587, 122485, vae were collected year-round. 122383, 122080, 122591, 122058, 122477, Etymology.ÐThe species name is a noun 121602, 122081, 122400, 122374, 121579, in apposition, derived from the genus of the 122578, 122762, 122024, 122023, 122430, most common larval host (Gnetum) and the 121603, 122510, 121606, 125410, 125185, Latin ``to eat'' (vorare). 124501, 125172, 125182, 124979, 124887, Discussion.ÐXenothictis are character- 124820, 124880, 124597, 124648, 125489, ized by slight to moderate sexual dimor- 124661, 125116, 124757, 125024, 127025, phism in forewing pattern and length; a ve- 126975, 195727, and 195728. Project spec- nation with all veins separate and chorda imens 86216 and 86817 are in the PNG ref- present or at least indicated in forewing, in erence collection in the laboratory in Ma- hindwing Rs an M1 closely approximated 1048 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 6±7. Live larvae of Xenothictis gnetivora on Gnetum gnemon. 6, On leaf surface, 7, In leaf roll. to connate, M2 somewhat distant from con- female genitalia unambiguously associates nate M3 and CuA1; a very long labial pal- the genus with Archipini. The absence of pus with a greatly enlarged, triangular third the second row of spines on the dorsum of segment (not so in X. gnetivora); a ciliate abdominal segment 3 of the pupa has not antenna in male; a bifurcate uncus; and previously been reported in Tortricidae and long, subparallel gnathos arms joined by a may represent a synapomorphy for the ge- cross-bar. While the male genitalia of Xen- nus. othictis are moderately plesiomorphic, with Xenothictis gnetivora differs from all the valva, transtilla, and aedeagus little congeners by its short, unmodi®ed labial modi®ed, the well developed signum of the palpus in both sexes. However, wing ve- VOLUME 105, NUMBER 4 1049 nation, wing pattern, and genitalia of both tory, Washington, D.C. Parataxonomists sexes leave no doubt that X. gnetivora John Auga, Willian Boen, Chris Dal, Sam- forms a monophyletic group with all other uel Hiuk, Brus Isua, Martin Kasbal, Richard Xenothictis species. Its simple valva is more Kutil, Markus Manumbor, and Kenneth plesiomorphic than that of its three conge- Molem assisted with most aspects of the ners with known males (X. atri¯ora, X. noc- project in PNG. Numerous collectors, ac- ti¯ua, and X. sciaphila) and is nearly iden- knowledged elsewhere, assisted with insect tical to that of the Australian genus Thrin- sampling. The landowners Kiatik Batet, cophora Meyrick. Further apomorphies Hais Wasel, and Sam Guru kindly allowed linking Xenothictis to Thrincophora and its us to collect on their lands. Plant taxono- sister genus Acropolitis Meyrick are overall mists are acknowledged in Miller et al. very similar female genitalia and the same (2003). DNA barcodes were provided by tegumen, juxta, aedeagus, and transtilla Paul Hebert and Erin Penton, University of structure, although the latter is dentate in Guelph. The following provided helpful re- Thrincophora and Acropolitis. The short la- views of the manuscript: F. Christian bial palpus of X. gnetivora is similar to that Thompson and David Smith, Systematic of Thrincophora and Acropolitis, suggest- Entomology Laboratory, USDA, National ing that it represents the plesiomorphic Museum of Natural History, Washington, character state for the group as a whole. D.C., U.S.A; and Kevin Tuck, The Natural Given the combination of morphological History Museum, London, England. The characters present in X. gnetivora, it is even project has been funded by U.S. National more doubtful that Xeneda, with its modi- Science Foundation (DEB-94-07297, 96- ®ed, elongate labial palpus, should be treat- 28840, 97-07928, 02-11591), National ed as a separate genus simply on the Geographic Society, Czech Academy of strength of the two patches of bristles on its Sciences (GAAV 651106), Czech Ministry uncus base, leaving Xenothictis parapyletic of Education (ES 041), Czech Grant Agen- and without a generic synapomorphy. It is cy (653112), International Centre of Insect more likely that the patches of bristles rep- Physiology and Ecology (Nairobi), and resent a species-level autapomorphy. Otto Kinne Foundation. The Smithsonian The rearing data on the leaf roller X. gne- Institution, Bishop Museum, and The Nat- tivora are the ®rst comprehensive biological ural History Museum (London) provided information for the genus Xenothictis. critical facilities for the taxonomic work. There is only one host record in the Aus- LITERATURE CITED tralian National Insect Collection for the Australian X. sciaphila, with its larva found Basset, Y., V. Novotny, S. E. Miller, and R. Pyle. 2000. in or on the fruit of Musa sapientum at Quantifying biodiversity: Experience with para- taxonomists and digital photography in Papua Montville and Eumundi, Queensland, June New Guinea and Guyana. BioScience 50: 899± 1962 and August±September 1963 by D. A. 908. Ironside. Clarke, J. F. G. 1958. Catalogue of the Type Specimens of Microlepidoptera in the British Museum (Nat- ACKNOWLEDGMENTS ural History) Described by Edward Meyrick, Vol. 3. Published by the Trustees of the British Mu- Our colleagues Yves Basset, Vojtech No- seum, London, 599 pp. votny, and George Weiblen collaborated at Diakonoff, A. 1961. Records and descriptions of exotic all stages of the project. Kevin Tuck assist- Tortricoidea. Annales de la SocieÂte Entomolo- ed in identifying this species and com- gique de France 130: 49±76. mented on an early draft of the manuscript. Hebert, P. D. N., A. Cywinska, S. L. Ball, and J. R. deWaard. 2003. Biological identi®cations through Illustrations were skillfully prepared by Ka- DNA barcodes. Proceedings of the Royal Society rolyn Darrow, Department of Systematic of London B 270: 313±321. Biology, National Museum of Natural His- Holloway, J. D. 1979. A Survey of the Lepidoptera, 1050 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Biogeography and Ecology of New Caledonia. W. nardiella sciaphila Turner, 1925, by orig- Junk, The Hague, 588 pp. inal designation. Horak, M. 1984. Assessment of taxonomically signif- icant structures in Tortricinae (Lep., Tortricidae). atri¯ora Meyrick 1930 (Xenothictis), Mitteilungen der Schweizerischen Entomologisch- Exotic Microlepid. 3: 609. TL: Fiji en Gesellschaft 57: 3±64. (Vunidawa). HT (male): The Natu- Horak, M., I. F. B. Common, and F. Komai. 1996. Tor- ral History Museum, London. tricidae, pp. 123±136. In Nielsen, E. S., E. D. Ed- melananchis Meyrick 1933 wards, and T. V. Rangsi, eds. Checklist of the Lep- idoptera of Australia. Monographs on Australian (Xenothictis), Exotic Microle- Lepidoptera 4. CSIRO, Collingwood, Australia. pid. 4: 424. TL: Fiji (Vunida- Meyrick, E. 1910. Revision of Australian Tortricina. wa). HT (male): The Natural Proceedings of the Linnean Society of New South History Museum, London. Wales 35: 139±294. nocti¯ua Diakonoff 1961 (Xenothic- Miller, S. E., V. Novotny, and Y. Basset. 2003. Studies on New Guinea moths. 1. Introduction (Lepidop- tis), Ann. Soc. Entomol. France tera). Proceedings of the Entomological Society of 130: 66. TL: New Hebrides (Vanu- Washington 105: 1034±1042. atu). HT (male): MuseÂum National Novotny, V., Y. Basset, S. E. Miller, P. Drozd, and L. d'Historie Naturelle, Paris. Cizek. 2002a. Host specialization of leaf-chewing paragona Meyrick 1910 (Xenothictis), in a New Guinea rain forest. Journal of Ecology 71: 400±412. Proc. Linnean Soc. N. S. Wales 35: Novotny, V., Y. Basset, S. E. Miller, G. D. Weiblen, B. 280. TL: Lifu. HT (female): The Bremer, L. Cizek, and P. Drozd. 2002b. Low host Natural History Museum, London. speci®city of herbivorous insects in a tropical for- sciaphila (Turner 1925) (Barnardiel- est. Nature 416: 841±844. la), Trans. Roy. Soc. South Austral. Novotny, V., S. E. Miller, Y. Basset, L. Cizek, P.Drozd, K. Darrow, and J. Leps. 2002c. Predictably sim- 49: 50. TL: Australia (Queensland, ple: Assemblages of caterpillars (Lepidoptera) Toowoomba). HT (male): Austra- feeding on rainforest trees in Papua New Guinea. lian National Insect Collection, Can- Proceedings of the Royal Society of London (B) berra, Australia. 269: 2337±2344. semiota Meyrick 1910 (Xenothictis), Sohn, J.-C. 2002. A taxonomy study on the immature stage of leaf-rolling Tortricidae (Lepidoptera) on Proc. Linnean Soc. N. S. Wales 35: rosaceous fruit trees in Korea, Master of Science 280. TL: Lifu. HT (female): The Thesis, Kangwon National University, Korea, 151 Natural History Museum, London. pp. Xeneda Diakonoff APPENDIX Xeneda Diakonoff 1961, Ann. Soc. Ento- mol. Fr. 130: 62. Type species: Xeneda Xenothictis Meyrick coena Diakonoff, 1961, by original des- Xenothictis Meyrick 1910, Proc. Linnean ignation. Soc. N. S. Wales 35: 279. Type species: coena Diakonoff 1961 (Xeneda), Ann. Xenothictis paragona Meyrick 1910, by Soc. Entomol. Fr. 130: 64. TL: New original designation. Caledonia (environs of Noumea). HT Barnardiella Turner 1925, Trans. Roy. Soc. (male): MuseÂum National d'Historie South Austral. 49: 49. Type species: Bar- Naturelle, Paris.