PROC.ENTOMOL.SOC.WASH. 105(4).200.^.pp. 1034-1042 STUDIES ON NEW GUINEA MOTHS. 1. INTRODUCTION (LEPIDOPTERA) Scott E. Miller, Vojtech Novotny, and Yves Basset (SEM) DepartmentofSystematic Biology,National MuseumofNatural History,Smith- sonian Institution, Washington, DC 20560-0105, U.S.A. (e-mail: [email protected]. edu); (VN) Institute ofEntomology, Czech Academy ofSciences and Biological Faculty, University ofSouth Bohemia, Branisovska31, 37005 CeskeBudejovice,CzechRepublic; (YB) Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Ancon, Panama — Abstract. This is the first in a series of papers providing taxonomic data in support ofecological and biogeographic studies of moths in New Guinea. The primary study is an extensive inventory ofthe caterpillar fauna ofa lowland rainforest site near Madang, Papua New Guinea, from 1994-2001. The inventory focused on the Lepidoptera com- munity on 71 woody plant species representing 45 genera and 23 families. During the study, 46,457 caterpillars representing 585 species were sampled, with 19,660caterpillars representing 441 species reared to adults. This introductory contribution is intended to provide background on the project, including descriptions of the study site, sampling methods, and taxonomic methods. Kev Words: Malesia, Papua New Guinea, Lepidoptera, biodiversity,rearing,community ecology A very large portion of tropical biodi- 1992 and 1993 (Basset 1996, Basset et al. versity consists ofherbivorous insects, and 1996). This paper represents the first in a among them, Lepidoptera are among the series of papers providing taxonomic doc- most amenable to study. To better under- umentation in support ofthe broader stud- stand the structureandmaintenanceoftrop- ies,andisintendedtoprovidegeneralback- ical biodiversity, we undertook a series of ground, including descriptions ofthe study related inventories of Lepidoptera in New site, sampling methods, and taxonomic Guinea. Our most extensive data set is an methods. inventory of the caterpillar fauna of low- During 2002, we sampled an additional land rainforests near Madang, Papua New 19 woody plant species near Madang, Guinea, from 1994-2001. Our ecological bringing the total sampling universe to 90 analyses focus on the Lepidoptera com- species representing 58 genera and 32 fam- munity on 71 woody plant species repre- ilies. The sampling effort per plant was re- senting45 generaand 23 familiesnearMa- duced according to the guidelines in No- dang. Ofthese species, 69 are native, while votnyetal. (2002c). The insects fromthese 2 species of Piper are not native. But for surveysarestill beinganalyzed. Atthetime taxonomic purposes we have evaluated of this writing, the Madang study is being specimens accumulated more broadly, in- expandedtoincludemontanesitesandase- cluding the material resulting from a study ries of lowland sites, and material from focused on 10 woody plant species con- these studies will be discussed and de- ducted near Wau. Papua New Guinea, in scribed in later papers in this series. Sam- VOLUME 105.NUMBER4 pling began atourfirst montanesiteinJune species are among the most important ones 2001 in primary and secondary forests and inlocalrainforests,withcombineddiversity partially deforested landscape around Mu of579 species in New Guinea (Hoft 1992). Village near Kundiawa town in Chimbu This selection offamilies includes all main Province (145°02'E, 6°05'S. 1.800 m). lineages of flowering plants, viz. gymno- sperms. monocotyledons, basal eudicots, Materialsand Methods euasterids and eurosids (APG 1998). Fur- Madang Study Area ther, locally common plants from all main habitats within the study area, including The study area is situated in Madang early and late stages offorest successionas Province, Papua New Guinea, extending well as riverine and seashore habitats, were Mforuonmtathiensc.oaAstvetroatgheeasnlonpueaslorfaitnhfealAldeinlbtehrits repPrleasnetnstewder(eLeipdsenettifaile.d2b0y01W).ayne Takeu- area is 3.558 mm. with a moderatedry sea- chi at Lae Herbarium and many of them son from July to September: mean airtem- were subsequently verified by the best perature is 26.5 °C (McAlpine et al. 1983). available international specialists. Plant The area is covered with species-rich ev- vouchers are deposited in Bishop Museum ergreen rainforest (152 species of woody (BISH). Rijksherbarium (L). Lae Herbari- plantswithdiameteratbreastheight ^5cm um (LAE)andSmithsonian Institution(US) perhectare; Laidlaw et al.. in press). Field- (herbarium acronyms follow Holmgren et work was concentrated in primary and sec- al. 1990). ondary lowland forests nearBaitabag.Ohu. and Mis villages, and in acoastal areanear Madang Insect Sampling and Rearing Riwo Village (145°41-8'E. 5°08-I4'S. ca. All externally feeding caterpillars (Lepi- 0-200 m). Specific localities are Baitabag doptera). including leafrollers and leaftiers, (145°47'E. 5°08'S, ca. 100 m). Ohu were collected by hand from foliage. Dur- (145°41'E. 5°14'S, ca. 200 m). Mis ing each sampling occasion, a collector (145°47'E. 5°11'S. ca. 50 m), Riwo Village spentoneday walkingthroughoutthestudy (145°48'E. 5°09'S. m). area searching the foliage ofthe target tree Madang Plant Sampling species for caterpillars. The sampling in- cluded only more accessible branches, i.e., Seventy-one species of trees and shrubs those which could be reached easily by from 45 generaand 23 families (seeappen- climbing or reached from the ground. Nu- dix), including 15 species of Ficiis and 1 meroustreesfrom variouspartsofthestudy speciesofArtocarpiis(Moraceae),6species area were sampled during each sampling ofMacaranga and 9 species representing9 occasion. The number of tree inspections, othergeneraofEuphorbiaceae,4speciesof that is, aparticulartree sampled at apartic- Psychotria and 12 species representing 12 ular time, was recorded, as well as the ap- othergenera ofRubiaceae. 3 species ofSy- proximateareaofthefoliagesampled.Each zigittm (Myrtaceae). 3 species ofPiper(Pi- tree species was sampled continuously for peraceae) and 18 species representing 18 the period ofat leastone yearbetweenJuly other families offlowering plants, were se- 1994 and December 2001. Sampling effort lected for the study oftheir associated cat- was equal forall plant speciesandainount- erpillars. Moraceae, Euphorbiaceae. and edto 1,500m-offoliageareaexaminedper Rubiaceae. whichwerestudiedindetail,are species, while the number of tree inspec- importantcomponentsoflowlandrainforest tionsexceeded 1,000perplantspecies.This flora in the Madang area and elsewhere in sampling effort represented approximately New Guinea (Oatham and Beehler 1998), 2.000 person-days offieldwork. hi the lab- The five genera represented by multiple oratory, each caterpillar was provided with — PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON fresh leavesoftheplantspeciesfromwhich and hosts, are recorded in acustom Access it was collected, and was reared to an adult database, described in Basset et al. (2000). wheneverpossible. Only the specimensthat Background data and images are available fed were considered in the analyses. Cat- at www.nmnh.si.edu/new_guinea. erpillars and adults were assigned to mor- Our taxonomic studies also incorporate phospecies. Moiphospecies were assigned material reared near Wau. Papua New seve—n character codes as permanent identi- Guinea by Yves Basset and assistants dur- fiers four lette—rs representing the family ing the precursor to the Madang project in and three digits which remain unchanged 1992-1993 (Basset et al. 1996). Morpho- even ifthe field identification ofthe family species numbers for these specimens are was incorrect. distinguishedby beginningwith "LE." The Oursamplingofthe 71 plantspeciespro- plants from that study were identified by duced 46,457 caterpillars representing 585 Robert Hoft and vouchers are deposited at species, with 19,660 caterpillars represent- LAE and L. ing 441 species reared to adults. Most of the field activities were carried out by par- Insect Identification ata.xonomists, as described in Basset et al. Inthelaboratory,themoiphospeciescon- (2000). The numbers reported in our ana- cepts were confirmed by disection ofgeni- lytical papers are often lower, because var- talia (Clarke 1941. Robinson 1976) andex- ious analyses included only a subset ofthe amination of other characters. Identifica- plant species or were adjusted to equalize tions were made using relevant literature, sample size per host plant. but especially by comparison to the collec- In addition to the basic locality data and tions of Smithsonian National Museum of adult morphospecies code, standard labels Natural History. Washington (USNM), including caterpillar morphospecies. host Bishop Museum. Honolulu (BPBM). and plant name, host plant abbreviation, and especially the rich historic collections of specimen number, were affixed to each The Natural History Museum. London specimen. The caterpillar morphospecies (BMNH). as well as less frequent compar- code begins with '"CAT" and is used on isons to collections of Australian National specimens after 1995. The original host Insect Collection. Canberra (ANIC) and plant identification on the label sometimes Nationaal Natuurhistorisch Museum, Lei- has changed with further study, so should den (RMNH), and types in other collec- be used with care. For this reason, begin- tions. Because most of the types of New ning in 1996. the labels include a three let- GuineamothsareatBMNH, ithasbeenthe ter host plant abbreviation that does not critical resource, and we are especially in- change with subsequent identifications. In- debted to their staff, as well as research as- dividual specimen numbers are assigned to sociates J.D. Holloway and M. Shaffer, for all specimens reared to adults. Early spec- unlimited access. imens bearnumbers in BishopMuseumda- General taxonomiccontextisprovidedby tabase series, while later specimens bear Holloway et al. (2001). although we follow numbers in Smithsonian database series Kristensen (1998) in recognizingCrambidae theuseofBishopMuseumandSmithsonian as a family. For macrolepidoptera. the on- on the labels provides a unique specimen going series "Moths ofBorneo" (Holloway numberand does not in itselfidentifyown- 1984-present) provides a vital foundation. ership of the specimen (e.g.. Thompson For pyraloids and microlepidoptera. Robin- 1994), sonetal.(1994)andDiakonoff(1952-1955) All data regarding specimens, theirrear- provide a general context. Nielsen et al. ing status, moiphospecies numbers, and (1996) provide a taxonomic framework for identifications, alonsz with imagesofinsects theAustralianfaunathathasbeenveryhelp- VOLUME 105. NUMBER4 1037 fill. Gressitt and Szent-Ivany (1968) provide sembled by the late J. J. H. Szent-Ivany. a bibliography of Lepidoptera systematics From 1968 to 1971, and again in 1974, literature for New Guinea, now updated by Szent-IvanycollectedGeometroideaatlight us and available online at www.nninh.si. and by rearing around Wau Ecology Insti- edu/new_guinea. tute, Papua New Guinea. He assembled a Taxonomic characters, and deHnitions of collection of some 300 species (listed in genera and species, follow tho.se in general Gressitt and Nadkarni 1978: 83-88) which use in Lepidoptera (e.g.. Miller 1994). as is now at BPBM. Szent-Ivany identified well asspecialistliteratureasavailable.The these during an extended visit to BMNH. reviews of many families in "Moths of Unfortunately, littleofhisrearingdatahave Borneo" have been especially important in been published, and many of the reared guiding generic and specific concepts. Be- specimens remain cryptically labelled. cause our immediate need is identification of the reared species, we often verified DiSCUS.SION identihcations by dissection of type speci- A prerequisite to investigation of the mens, but have not reviewed the variability ecology of species-rich insect taxa in di- across the entire geographical range of the verse habitats such as lowland tropical rain species. Cytochrome oxidase I (COI) se- forests is alarge sample size. Oursampling quences. DNA barcodes of Hebert et al. generated data on a scale that has rarely (2003).follow theprotocolsoutlinedin He- been achieved in the tropics (see Janzen bert et al. (2003). The primary set of Lep- 1988, Janzen and Gauld 1997, Janzen 2003 idoptera vouchers are deposited in USNM. for a similar exercise in Costa Rica). Our with representatives in the National Agri- data are now being used fora seriesoftax- cultural Research Institute (Port Moresby), onomic (e.g., Holloway and Miller 2003) BPBM. and other collections as appropri- andecologicalanalyses(e.g.,Novotnyetal. ate. 2002a, b, c). Some of the ecological con- clusions are reviewed below. Other Material Individual host-plant species sustained The historic collections of USNM. from 9 to 75 (median 25) species ofcater- BPBM, and especially BMNH (Frodin and pillars. Caterpillar communities were Gressitt 1982) provided the context for the strongly dominated by a single or few spe- Madang collections. USNM collections in- cies. The single most common species typ- clude material collected in Irian Jaya by ically represented 52% of individuals and Syuti Issiki May-August 1936, and in Pap- 50% of biomass while the five most com- ua New Guinea by Gary Hevel in Decem- mon species represented >80% ofindivid- ber 1976, ScottMillerand PamelaMillerin uals and biomass in the entire community July-August 1983, and Vitor Becker in (Novotny et al. 2002c). In addition tothese September-October 1992. The BMNH col- dominants, each community included a lection is especially richbecauseofaseries large numberofvery rare species(Novotny ofexcellent collectors sent to New Guinea and Basset 2000). Despite significant sam- by WalterRothschild fortheTring Museum pling effort, the species accumulation (Rothschild 1983). and also includes many curves for individual host plant speciesdid vouchersfromagriculturalandforestrypro- not approach an asymptote which suggests jects (e.g.. Bigger 1988). that the total species richness ofcaterpillar hi addition to the general collections at communities was not sampled (Novotny et BPBM (Frodin and Gressitt 1982) and ma- al. 2002c). terial collected by LarryOrsakaround Wau Caterpillars were mostly specialized to a and Madang in the early 1990s, there is an single plant family, and within families to important collection of Geometroidea as- a single genus, while capable offeedingon PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON multiple congeneric hosts (Novotny et al. investigators who have helped provide a 2002b). Only 15% ofthecaterpillarspecies context fortheproject, AllenAllison.Larry feeding on Ficits, Macaranga, and Psyclio- Orsak. George Weiblen. and Jan Leps. to trici strongly preferred a single host species postdoctoral researchers and graduate stu- (Novotny et al. 2002a). but even among dents Lukas Cizek. Pavel Drozd. Jiri Hulcr these species, none was strictly monopha- and Milan Janda, as well as Karolyn Dar- gous. Thus, it is conceivable that no. or row. Parataxonomists Mark Andreas. John very few. genuinely monophagouscaterpil- Auga. William Boen. Chris Dal. Micah lars feed on speciose plant genera in rain Damag. Samuel Hiuk. Brus Isua. Martin forests. A large overlap among caterpillar Kasbal. Richard Kutil. Max Manaono. Mar- communities on congeneric plants means kus Manumbor. Martin Mogia. Kenneth that the total number ofspecies feeding on Molem. and Elvis Tamtiai assisted with speciose plant genera is relatively small, in most aspects of the project in PNG. Nu- comparison with their size. These differ- merous collectors, acknowledged else- ences, in combination with low host speci- where, assisted with insect sampling. The ficity of herbivores with respect to conge- landowners Kiatik Batet. Ulai Koil. Hais neric plants, suggest that the average over- Wasel. and Sam Guru kindly allowed us to lap among herbivore communities on trop- collect on their lands. Botanical taxonomic ical trees may be higher than on temperate help wasprovided by L. Balun.C. Berg.B. trees (Novotny et al. 2002a). Bremer. K. Damas, John Dransfield, H.-J. Caterpillarcommunitieswerenotseason- Esser, Paul Katik. C. E. Ridsdale, George al, and the majority ofspecies were present Staples,George Weiblen, PetervanWelzen, almost continuously throughout the year T C. Whitmore, andespecially WayneTak- (Novotny et al. 2002c). Community com- euchi. Taxonomic help with Lepidoptera position was also constant spatially over was provided by David Adamski, Vitor distances <20 km (Novotny et al. 2002c). Becker. John Brown. David Carter. Ian The dominance of caterpillar communities Common. E.D. (Ted) Edwards. Marc Ep- by a small number of species, which also stein. Anthony Galsworthy. Robert Hoare, exhibited low spatial andtemporal variabil- Ronald W. Hodges. Jeremy D. Holloway. ity, permitted robust and reliable estimates Martin Honey. Marianne Horak, Ian J. of community composition and between- Kitching, Martin Lodl. Koen Maes,Jacque- community similarity from small samples, line Y. Miller, Lee D. Miller, Eugene G. typically <300 individuals per host plant. Munroe, Ulrich Paukstadt. Michael Par- In contrast, even considerably larger sam- sons. Richard Peigler. Robert W. Poole, ples were not sufficient for estimates of Gaden Robinson. Rikio Sato. Klaus Sattler. community species richness (Novotnyetal. Malcolm Scoble, Michael Shaffer,M. Alma 2002c). Solis, Kevin Tuck, and Richard Vane- The analyses produced from these data Wright. Technical assistance at BishopMu- show the importance of large samples col- seum was provided by Candida Cardenas, lected over multiple years in understanding Valerie Hedlund, June Ibara. Karin Kami, thestructureoftropical insectcommunities. Tracie Mackenzie, Gordon Nishida, and These large samples have only been logis- David Preston, and at The Natural History tically feasible with a team approach, uti- Museum, London by Maia Vaswani and lizing the skills of parataxonomists, ecolo- Shayleen James. Access to additional type gists. and systematists. specimens was providedby Wolfgang Mey, Humboldt University, Berlin: Erik van Acknowledgments Nieukerken and Rienk de Jong, Nationaal This has truly been a collaborative pro- Natuurhistorish Museum, Leiden; Chris ject. We are grateful to our other principal Burwell, Queensland Museum, Brisbane; . VOLUME 105.NUMBER4 1039 and D.J. Mann. Oxford University, Oxford. Guinea.ResultsoftheThirdArchboldExpedition JohnW. Brown,RonaldW. HodgesandDa- (1938-39).PartsI-V.VerhandelingenderKonink- vid Smith commented on the manuscript. lijke Nederlandsche Akademie van Wetenschap- The project has been funded by U.S. Na- p1e6n4.:s5e0r(.1)2.:419-(12)9:1;15-01(637);:419-(231):0.1-166:49(4): 1- tional Science Foundation (DEB-94-07297, Frodin. D.G. andJ. L. Gressitt. 1982. Biologicalex- 96-28840, 97-07928, 02-11591), National plorationofNewGuinea,pp.87-130.InGressitt, Geographic Society, Czech Academy of J. L.. ed. Biogeography and Ecology of New- Sciences (A6007106, Z5007907), Czech Guinea. W.Junk.TheHague. MGirnainsttrAygeonfcyEd(u2c0a6t/i9o9/n11(1E5)S,0I4n1t)e,rnaCtizoencahl GressiEKtcatio.nldoJig.:y.L.BWaaacnkudgrENoc.uonlNdoagdyktaorInnMsito.intut1tae9,n78eW.aNuG.ueiwdvieG-iut-oin1Me3t5a Centre of Insect Physiology and Ecology pp. (Nairobi), Otto Kinne Foundation, and Gressitt. J. L. and J. J. H. S/ent-lvany. 1968. Bibli- PNG Biological Foundation. National Sci- ography ofNew Guinea entomology. Pacific In- ence Foundation grant BSR 89-13871 pro- sectsMonograph 18: 1-674. vided curation support to the Bishop Mu- Hebert. P D. N.. A. Cywinska. S. L. Ball, and J. R. osinfeuPtmhNeGcoNlawltaeicsotniasolun,ppAoacrnatddedeMmiblyylerot'fhseSecBaiarelcniheceresFwuoanrndkd Hoft.doDfeRNW.LaAoa1n9rb9dda2o.r.nc2oP0dBl0eas3n2..t7s0PB:orifoo3lcNo1eeg3ei-dwci3an2Glgu1siidnoeefnattihfaeincdaRtotiyhoeanslStoShlorcooimueogtnhy the New England Biolabs Foundation. The Islands.DictionaryoftheGeneraandFamiliesof Smithsonian Institution, Bishop Museum Flowering Plants and Ferns. Wau Ecology Insti- andNatural History Museum(London)pro- tuteHandbook 13. vided critical facilities for the taxonomic HolloIswsauye,dJi.nDp.art1s98b4y-2M00a3l.ayaTnheNaMtoutrhesJoofurnBaolrnaeno.d work. Southdene,KualaLumpur Holloway.J.D..G.Kibby.D.Peggie.D.J.Carter,and Literature Cited S. E. Miller 2001. Families of Malesian Moths andButterflies. Brill.Leiden, xii + 456pp. APG (Angiospemi Phylogeny Group). 1998. 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Microlcpidoptera.EntomologistsGa/ette27: 127- Checklistofthe LepidopteraofAustralia. Mono- 132. graphsonAustralianLepidoptera4: xiv + 529. Robmson.G. S.. K. R.Tuck,andM. Shaffer. 1994. A Novotny. V. and Y. Basset. 2000. Ecological charac- held guide to the smaller moths of South-east teristicsofrarespeciesincommunitiesoftropical Asia. Malaysian Nature Society and Natural His- insect herbivores: Pondering the mystery ofsin- tory Museum. Kuala Lumpur and London. 309 gletons, OikosH9: 564-572. PP Novotny, V., Y, Basset. S. E, Miller, R Dro/d.and L, Roihschild. M. 1983, Dear Lord Rothschild; Birds, Ci^ek,20()2a,Hostspecializationofleaf-chewing ButtcrlliesandHistory. Balaban, Philadelphia,xx insects in a New Guinea rain forest. Journal of + 398 pp, Animal Ecology 71: 400-412, Thompson, F C. 1994. Bar codes for specimen data Novolny, V,,Y Basset,S.E.Miller,G.D.Weiblen,B. management. InsectCollections News9; 2-4. VOLUME 103. NUMBER4 1041 Appendix lull list ofthe*-)() species cit pkiius s;iiiipled nearMadang. alphabetieally by tainily. The three lettereode is usedon iiiseel specimen labelsand allowspositiveassociation withthecorrect name incasetheplantidentifi- cationhaschangedfromtheoriginalidentihcation.Novotnyetal.(2002b)providedaphylogcnyfor43ofthese species. Agavaceae C Ixlii ills V Beauv COR Agavaceae Drai iiiKiisiiloIni Roxb. DRA Apoeynaceae Tain I'liUnui(iiiniiuicciGaud. TAB Araliaceae (>Mii,>\xl,<ii .,vsilifloniin(Lauterb.)W.R. Phihpson OSM Arecaceae /ivdruiMcIciiiuii>\piiili.\(Becc.)Burret ARE Bignoniaceae Spailii'ilciiciiiii/iiiiiiiUiiii(L.) Kunth. SPA Caesalpiniaceae MaiiilUHicl.plitniiii^ii Merrill& Perry MAN Eaiphorbiaceae Brcyiiui.cniiui(Poir.) Much, Arg. BRE liuphorbiaceae Coduuiiiiiliulnvnuiiiiiiii Any .Shaw COD Euphorbiaceae Eiulo.spc'iiniiiiiliihiiis Schodde END Euphorbiaceae ExciiccariaaiiaUochaL. EXC EEuupphhoorrbbiiaacceeaaee HMoancuainlaiinuyhaiisiilncoiviiiiiiiniiiiiliiiswcEn.siMsue(lWla.rb.I K. Schuni. HMOAMA Euphorbiaceae Macanms^iihi/mciiui.1.1. Smith MAP luiphoibiaceae MiUiinmiiiiI'lculntriiIkiAny Shaw MAF l-nphoihi.Kce Mcuuiiiiif^ciilcnsitloraWarb. MAD l-.tiphorbiaceae Mcictiraiifiiiiiiniif^idneensisJ.J.Smith MAU liuphorbiaceae MiiciiiiiiiniiipiiulrigUmdulosaWarb. MAQ liuphorbiaceae Mallntiis niollisMiinis(Geisel.)AiryShaw MAL luiphorbiaceae Mchinolcpi-. iiiiiliiKliiiiiliilnsa(Reinw.ex Bl.lReichb.f.& Zoll. MEL luiphorbiaceae PhxIUiiiilnis liiiiipn.plixlhisMuell. Arg. PHY liuphorbiaceae Piiiu'loJciu/niiiaiiilioiiiiciini Hassk. PIM liuphorbiaceae EiiponuitiaUtiirina R. Br. EUP l-'abaceae Pleroviirinisiiulicu.sWilld. PTE I'lacourtiaceae Ciiscciiuicrvlliii'iiirpii.Sleum. CAS (inetaceae Ciicnim i^ncnioii 1,. GNE Heliconiaeeae HchcoiiuipapiuiiiiiW.J. Kress HEL Lecythidaceae Belliiiiiiltiiiuisp. BAR Leeaceae Lcciiiiu/icuMerrill LEE Loganiaceae Neiiburgiacoryiwcciipn(A.Gray)Leenh. NEU Malvaceae HibiscustiliaceiisL. HIB Malvaceae Sicniiliii s,liiiiiiiiniiiiiini (Lauterb.)Mildbr, STR Malvaceae Tinlif'spi-niiiiinpUniMigiiui(F. Muell.) Kostcrmans TRI Meliaceae A^^liiuicl ciniilliiici(Roxb.)Pellegr. AGL Monimiaceac Kihaiacl miuicca Hook.f.&Thoms. STG Moraceae AiiiHiiipii.scinimiiiiiisJ.R.etG.Forst. ART Moraceae Ficush&nuiysii King BER Moraceae FiiiisInitryocarpnMiq. EOT Moraceae FicusciniocephalifoliaRidley CON Moraceae FicusciipiiisaSteud. COP Moraceae FicusilaininaropsisDiels DAM Moraceae FicuscrythrospermaMiq. ERY Moraceae FicusgillK. Schum. & Laut. GUL Moraceae FicusliispidioiilcsS. Moore HIS Moraceae FicusinicnicarpaL. MIC Moraceae FicusiiKilliorF. Meull.ex Benth. MOL Moraceae FicusimdosaTeysm.& Binn. NOD Moraceae FicusjHichyrrhachisK. Schum. lVi Laut. PAR Moraceae Ficuspluiosxcc Lain c*;; K, Schum. PHA PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON Mitraceac Ficnspuiii^i'nsReinw.ex Bluine PUN Moraceae FicKSsepticaBurm. SEP Moraceae Ficussubtriner\'iaLaut. & K. Schuiii. PAS (= F.pachystemonWarb.) Moraceae FicuslernatanaMiq. TER Moraceae FicustinctoriaForst. TIN Moraceae FicKStrachypisonK. Schum. TRA Moraceae FicusvariegataBlume VAR Moraceae FicuswassaRoxb. WAS Myristicaceae Myristicacf.sepicaini D.B. Foreman MYL Myrtaceae Syzigiumlongipes(Warb.)Merrill & Perry SSW Myrtaceae Syzygiumsp. I SRS Myrtaceae Syzygiumsp. 2 SRB Piperaceae PiperaduncumL. PAD Piperaceae PipermacropiperPennant PMV Piperaceae PiperumbellatumL. PUB Rubiaceae AmaracarpusnymuniiValeton AMA Ruhiaceae Dolicholohiuino.wlohuin K. Schum. DOL Riihiaceae Gardeniahan.scnuiiiiiii K. Sclium. GAR Rubiaceae Morindahracteaia Roxb. MOR Rubiaceae MussaendascratclilcyiWernh. MUS Rubiaceae Naucleaorientalis(L.) L. SAR Rubiaceae NeonacleaclemensiiMeirill& PeiTy NEO Rubiaceae PavettaplatycladaLauterb. & K. Schum. PAV Rubiaceae PsycholrialeptothyrsaMiquel PSF Rubiaceae Psychotriamicralabaslra(Laut.& K. Schum.)Val. PSM Rubiaceae Psycholriamicrococca(Laut.& K. Schum.)Val. PSS Rubiaceae PsychotriaramuensisSohmer PSL Rubiaceae RandiaschumannianaMerrill& Perry MEN Rubiaceae Tarennaburiiensis(Miq.)Val. TAR Rubiaceae Timoniuslimon(Spreng.)Merrill TIT Rubiaceae Versleegiacaulijiora(K.Schum.& Laut.) VER Rutaceae LunasiaamaraBlanco LUN Sapindaceae PometiapinnalaForster POM Sapotaceae Pouteriasp. POU Sterculiaceae KleinhoviahospitaL. KLE Ulmaceae CellisphilippensisBlanco CEL Urticaceae Leucosykecapilellata(Poir.)Wedd. LEU Verbenaceae GeunsiafarinosaBlume GEU Verbenaceae PremnaobtusifoliaR.Br. PRE Verbenaceae Teijsmanniodendronsp. TEI Zingiberaceae Honistcdtiascoiliana(F. Muell,) K. Schum. HOR