Moreae, Artocarpeae, and Dorstenia (Moraceae): With Introductions to the Family and Ficus and With Additions and Corrections to Flora Neotropica Monograph 7(Flora Neotropica Mongraph No. 83)

FLORA NEOTROPICA MONOGRAPH 83

MOREAE,ARTOCARPEAE, ANDDORSTENIA (MORACEAE) With Introductions to the Family and Ficus and with Additions and Corrections to Flora Neotropica Monograph 7

CORNELIS C. BERG Of

c

CAN1CEl

FLORAt NEOTROPICA( '^

IOP'C

CAPRICORN

Publishedfor The Organizationfor FloraNeotropica by The New York BotanicalGarden Bronx, New York

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Library of Congress Cataloging-in-Publication Data Flora neotropica.-Monograph no. 1 -New York:Published for Organizationfor FloraNeotropica by The New York Botanical Garden,1968v.: ill.; 26 cm. Irregular. Eachissue has distinctivetitle. Separatelycataloguedand classified in LC before monographno. 40. ISSN 0071-5794 = Floraneotropica. 1. Botany - LatinAmerica- Classification- Collected works. 2. Botany - Tropics- Classification- Collected works. 3. Botany - Collectedworks.I. Organization Classification forFloraNeotropica.

II. New YorkBotanical Garden. QK205.F58 Libraryof Congress ISBN 0-89327-439-9

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[8508]

1 2 3 4 5 6 7 8 9 / 09 08 07 06 05 04 03 02 01 1V

AND DORSTENIA MOREAE,ARTOCARPEAE,

(MORACEAE) With Introductionsto the Family and Ficus and with Additions and Correctionsto Flora NeotropicaMonograph7 CORNELISC. BERG

CONTENTS .............................................................1 Abstract/Resumen/Resumo 4 .............................. Introductionto the Family Moraceae............................... 4 TaxonomicHistory............................................................. Morphology.............................................................4 12 Pollination............................................................ 14 Dispersal ............................................................. 15 ........................................... ................. DistributionandEcology 20 ............................. ClassificationandDiversification............................... 21 Use ............................................................. 22 ConservationStatus ............................................................ 22 TaxonomicTreatment............................................................ Key to the Tribes of the Moraceaein the Neotropics ............................ .................. 22 Key to the NeotropicalGeneraof the Moraceae...................................................... 22 24 Moreae............................................................ 67 Artocarpeae............................................................ 143 Dorstenieae............................................................ Dorstenieaep.p. ("Brosimeae")andCastilleae("Olmedieae"),Additionsand Correctionsto FloraNeotropicaMonograph7......................................................... 231 289 Ficeae ............................................................ 292 Acknowledgments............................................................ 292 LiteratureCited ............................................................. 300 Maps ............................................................ 313 NumericalList of Taxa ............................................................ 315 List of Exsiccatae ............................................................ 340 ............................................................ Names Index of Local 342 Index of Scientific Names ............................................................

ABSTRACT Berg, Cornelis C. (BotanicalInstitute,Universityof Bergen/TheNorwegianArboretum, N-5259 Hjellestad,Norway). Moreae,Artocarpeae,andDorstenia (Moraceae);with introductions to the family and Ficus and with additions and corrections to Flora Neotropica Monograph7. FloraNeotropicaMonograph83: iv+l-348. 2001.-The presentmonograph includesthe revisions of generaof the tribesMoreaeandArtocarpeae,the genus Dorstenia, correctionsand additionsto genera treatedin Fl. Neotrop. Monogr. 7, "Brosimeae,"pres-

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FLORANEOTROPICA

and"Olmedieae," entlyincludedin thetribeDorstenieae, presentlyCastilleae. Theyarepreto thefamilyMoraceae withbriefintroductions cededby anintroduction andsupplemented forthefirsttimeforFloraNeotropica tothetribeFiceaeandthegenusFicus.Theentitiestreated compriseninegenera:Bagassa (1 sp.), Batocarpus (3 spp.),Clarisia (3 spp.),Dorstenia (46 spp.),Maclura(2 spp.),Morus(2 spp.),Poulsenia(1 sp.), Sorocea (14 spp.),andTrophis (5 spp.)-77 speciesin total.Eightnewspeciesandtwonewsubspecieswerediscovered of themonograph, forthesegeneraduringthepreparation aswellasa secondyet-undescribed speciesforPoulsenia. Thisgroupof generacomprisesmanywidespreadand/orcommon speciesforwhichnumerouscollectionshavebeenexaminedandlisted.However,in Dorandthenumberof specimensexstenia, manyspeciesareverylimitedin theirdistribution aminedis oftenlimitedto a feworsometimes onlyone.Thegroupof theninerevisedgenera is in manyrespectshighlydiverse.Forthe groupof generatreatedin Monograph7 the additionsandcorrections include:revisedorextendeddescriptions of severalspeciesand dueto fusionof speciespreviouslyrecognizedasdistinct, subspecies,changeddescriptions anddocumentation forrangeextensions,somenewcombinations, fulltreatments indications of Monograph of taxadiscoveredsincethepublication 7 (12 speciesorsubspecies),supplementarycommentson generaandspecies,andpartlyorentirelyrevisedkeysto speciesfor severalgenera.Forthissecondgroup70 species(intheninegenera)arerecognized atpresent. to thefamilyincludeschaptersdealingwiththetaxonomichistory,morTheintroduction phology,pollinationanddispersal,distribution andecology,classification, diversification, ThegenusFicus (awaitingrevision)is to someextentincludedin the use,andconservation. inparticular inthechaptersorparagraphs introduction, dealingwithhabit,pollination,and a description of anda briefaccountof thisgenusareprovided.Inthe distribution; moreover, thenewcombinations presentcontribution, Clarisia Ruiz& Pavonsect.Acanthinophyllum (Allemao)C. C. Berg,Pseudolmedia Treculsect.Olmediopsis (Karsten)C. C. Berg,and Naucleopsis ulei subsp.amara (Ducke)C. C.Bergaremade,andthenewtaxonNaucleopsis ulei (Warburg) Duckesubsp.subandina C. C. Bergis described.

RESUMEN Berg, CornelisC. (BotanicalInstitute,Universityof Bergen/TheNorwegianArboretum,N-5259Hjellestad,Norway).Moreae,Artocarpeae, andDorstenia (Moraceae); with introductions to thefamilyandFicusandwithadditionsandcorrections toFloraNeotropica 7. FloraNeotropica Monograph 83:iv+1-348.2001.- Lapresentemonografia Monograph incluyelasrevisionesdelosgenerosdelastribusMoreaey Artocarpeae, el generoDorstenia, correcciones enlaFl.Neotrop.Monogr.7, "Brosimeae," y adicionesa losgenerospresentados actualmente incluidaenlatribuDorstenieae actualmente y "Olmedieae," Castilleae, precedidas a la familiaMoraceae porunaintroduccion y complementadas conbrevesintroducciones a la tribuFiceaey al generoFicus. Losgrupostratadosporprimeravez en FloraNeotropica nuevegeneros:Bagassa (1 spp.),Batocarpus(3 spp.),Clarisia (3 spp.),Dorcomprenden stenia (46 spp.),Maclura (2 spp.),Morus (2 spp.),Poulsenia (1 spp.),Sorocea (14 spp.)y Trophis(5 spp.)- 77 especiesen total.Ochonuevasespeciesy dos nuevassubespecies fuerondescubiertas paraestosgenerosdurantela preparaci6n de la monografia,asi como tambienunasegundaespecie,aumnodescritadePoulsenia.Estegrupodeg6neroscomprende muchasespeciesampliamente distribuidas y/o comunesde lascualesse hanexaminado y se citanmuchascolecciones. Sinembargo, enDorstenia,muchasespeciestienenunadistribucion muylimitaday el nu'mero de muestrasexaminadas se limitaa menudoa pocasy algunas veces solamentea una.El grupode los nuevegenerosrevisadoses, en muchosaspectos, extremadamente diverso.Parael grupode generospresentadosen la Monografia7, las adicionesy correcciones incluyen:descripciones revisadaso aumentadas de variasespecies y subespecies, cambiosenlasdescripciones debidoa fusi6ndeespeciespreviamente tratadas como diferentes,indicacionesy documentacionparaextensionesen los rangos,algunascom-

RESUMO

3

binaciones nuevas,tratamientos completosdetaxadescubiertos luegodelapublicacion dela Monografia 7 (doceespecieso subespecies), observaciones complementarias acercadelos generosy las especies,se revisanparcialo totalmente lasclaves,paralas especiesy varios g6neros.Paraestesegundogruposereconocen enla actualidad 70 especiesennuevegeneros. Laintroduccion a lafamiliaincluyecapitulosacercadelahistoriataxon6mica, morfologia, polinizaciony dispersi6n,distribucion y ecologia,clasificaci6n,diversificaci6n,usos y conservacion.El generoFicus (auinpor revisar)esta incluidohastaciertopuntoen la introduccion, enparticular encapituloso parrafosquese refierenal habito,polinizaci6ny distribuci6n; ademas,se proveeunadescripci6n y unapequenia relacionde esteg6nero.En la presentecontribuci6n, se hacenlas siguientescombinaciones nuevas:ClarisiaRuiz& Pav6n sect. Acanthinophyllum(Allemao) C. C. Berg, PseudolmediaTr6culsect. Olmediop-

sis (Karsten)C. C. Berg,andNaucleopsisulei (Warburg) Duckesubsp.amara(Ducke) C. C. Bergy se describeel nuevotaxonNaucleopsisulei (Warburg) Duckesubsp.subandinaC. C. Berg.

RESUMO Berg, CornelisC. (BotanicalInstitute,Universityof Bergen/The NorwegianArboretum,N-5259Hjellestad,Norway).Moreae,Artocarpeae, andDorstenia(Moraceae); with to thefamilyandFicusandwithadditionsandcorrections introductions to FloraNeotropica 7. FloraNeotropica 83:iv+1-348.2001.-Apresentemonografia Monograph Monograph incluirevis6esdosgenerosdastribosMoreaee Artocarpeae, do generoDorstenia,correcoes e acrescimosparaos generostratados naFl.Neotrop.Monogr.7, "Brosimeae" atualmente incluidanatriboDorstenieaee "Olmedieae," atualmente Castilleae.Saotodasprecedidas porumaintroducaoparaa familiaMoraceaee complementadas combreveintroducoes paraa triboFiceaee parao generoFicus.As entidadastratadaspelaprimeiravez paraa FloraNeotropicaabrangem novegeneros:Bagassa(1 sp.),Batocarpus(3 spp.),Clarisia (3 spp.), Dorstenia (46 spp.), Maclura (2 spp.), Morus (2 spp.), Poulsenia (1 sp.), Sorocea

(14 spp.)e Trophis (5 spp.)-num totalde 77 especies.Durante a preparaaodamonografia foramdescobertosparaestesgenerosoitonovasespeciese duasnovassubespecies,assim comoumasegundaespecieparaPoulsenia,aindanaodescrita.Estegrupode generoscompreendemuitasespecies comunse/ou amplamentedistribuidas,paraas quais foram e listadasnumerosas examinadas cole96es.Todavia,emDorstenia,muitasespeciespossuem limitadae o nuimero de amostrasexaminadas foi reduzidoou somenteum.0 distribuicao grupodos nove generosrevisadose, sobmultiplosaspectos,altamentediversificado.Os 7 incluem:descrie corre96esparao grupodosgenerostratados naMonografia acrescimos qoesrevisadasou ampliadasde variasespeciese subespecies,descric6esalteradaspor causadafusaodeespeciespreviamente reconhecidas comodistintas,indicac6ese documentratamento ta9aoparaa extensaodeocorencia,algumasnovascombina96es, completodos taxa descobertosdesde a publica9aoda Monografia7 (doze especies e subespecies), comentiriossuplementares paraos generose especiese chavesparaas especiesde varios revisados.Paraeste segundogruposao reconhecidas generosparcialou inteiramente atualmente70 especies(nosnovegeneros).A introdu9ao paraa familiaincluicapitulos dahistoriataxonormica, comabordagem morfologia,poliniza9aoe dispersao,distribuicao uso e conserva9ao.0 generoFicus geograficae ecologia,classifica9ao,diversifica9ao, em particular nos (aguardando revisao)e abordadocom algumaextensaona introdu9ao, capitulosou paragrafos quetratamsobreo hibito,polinizago e distribui9ao; alemdisso, umadescrigioe umasintesedessegenero.Napresentecontribui9ao sao saoacrescentadas feitasasnovascombinaq6es Clarisiasect.Acanthinophyllum (Allemao)C. C.Berg,PseudDucke olmediaTreculsect. Olmediopsis(Karsten)C. C. Berge Naucleopsis(Warburg) subsp.amara(Ducke)C. C. Berg,e e descritoo novotdxonNaucleopsisulei subsp.subandinaC. C. Berg.

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FLORANEOTROPICA

tinctfrombothMoraceae(s.str.)andUrticaceae(s.str.), Berg (1978a) proposedto treatthe group of six genera as a separatefamily,Cecropiaceae,a decisiongenerally The presentcontributionto FloraNeotropicacom- accepted,such as by Cronquist(1981), andconfirmed prisessectionswith generatreatedfor the firsttime and by Setoguchi et al. (1993) in a study on silicon-accusections with corrections and additions of genera re- mulating idioblasts in leaves of Cecropiaceae,which vised in FloraNeotropica Monograph7. This implies also supports the position of Poikilospermum as a thatall neotropicalMoraceaeexceptfor therepresenta- memberof the family.Forthe reducedMoraceae,Cortives of Ficus are included.However,in orderto make ner(1962) proposeda subdivisioninto six tribes:Artothe general survey of the family more complete, the carpeae,Brosimeae,Dorstenieae,Ficeae, Moreae,and genus Ficus is includedto some extentin the introduc- Olmedieae.Moreover,he reducedthe numberof gention to the family.Moreover,a briefintroductionto this era.The conceptsof the classificationanddelimitation genus is provided. of genera proposed by Cornerwere largely based on knowledge of the Asian andAustralasianmembersof thefamily.Classificationanddelimitationof generahave HISTORY TAXONOMIC been elaboratedby Berg (1973, 1977a, 1977b, 1986, 19th such as In most classifications of the century, those of Baillon (1875a) and Bentham and Hooker 1988, 1989b) by includingaccumulatedknowledgeof (1880), the Moraceae were included in an "order"or neotropical and African members of the family. The "family"with the subdivisions: Ulmaceae, Celtideae, results of these studies arediscussed underthe tribes. Accordingto presentconcepts,the familyMoraceae Cannabineae,Moreae, Artocarpeae,Conocephaleae, comprises 37 genera and 1050-1100 species worldandUrticaceae.However,in De Candolle'sProdromus, 19 wide, with generaandapproximately270 species in volumes 16 (De Candolle, 1869; Weddell, 1869) and 17 (Bureau, 1873; Planchon, 1873), the grouphad al- the Neotropics (Berg, 1998b). readybeensplitintofive related"orders"(Cannabineae, Urticaceae,Ulmaceae, Moraceae,andArtocarpeae),a MORPHOLOGY prelude to the subdivision of the order Urticales by The Moraceae are highly diverse in the morpholEngler(1889), in whichthe familyMoraceaecomprises ogy of bothvegetativeandreproductivestructures.The the subfamilies Moroideae (with the tribes Fatoueae, descriptionof the diversity is largely focussed on the Moreae,Broussonetieae,Strebleae,andDorstenieae), taxa treatedin the presentcontribution,but it includes Artocarpoideae (with the tribes Euartocarpeae, to some extentalso the genus Ficus andOld WorldrepOlmedieae,Brosimeae,andFiceae),Conocephaloideae, resentativesof the family. andCannaboideae.TheCannaboideaearemorerecently regardedas meriting the rank of family in the order Habit Urticales(cf. Cronquist,1981), butthe affinitiesto the In contrast to its sister family Urticaceae,the Moraotherfamilies of the Urticalesarenot quiteclear;a poceae are basically woody. However,nearly 10%of the sitioncloserto the Myricaceaecould be consideredand species are herbaceous. Most of the herbaceous speinvestigated. Inarevisionalstudy,Chew(1963) reducedtheAsian cies belong to the speciose genus Dorstenia and only genus ConocephalusBlume, the type of the subfamily two of them belong to the small paleotropical genus Conocephaloideae,to a subgenus of Poikilospermum Fatoua Gaudichaud.The herbaceousmembersof the Miquel, a genus which was accepted as a memberof genus Dorstenia show a wide range of habits and life the Urticaceae, e.g., by Engler (1889). This made the forms, as discussed in detail in the introductionto this weaknessof the demarcationof MoraceaeandUrtica- genus in the presentcontribution. About 30% of the species are basically hemiepiceae apparent.Chew proposed to transferthe microspermousgenera of the Conocephaloideae,Cecropia phytic and they belong to the genus Ficus. The ability Loefling, CoussapoaAublet,Musanga P. Brown, and to produce(aerial)adventitiousroots is not only comPoikilospennum(s.l.), to the Urticaceae, leaving the mon in Ficus (hemiepiphytesandsome groupsof climbmacrospermousgenera, MyrianthusP. Beauvois and ers,includingroot-climbers)butit is also foundin other PouroumaAublet,in the Moraceae.Comer(1962) made woody representativesof the order:it is characteristic a moreradicaldecisionby transferring the Conocephal- for the Cecropiaceae and occurs in the (Old World) oideae (includingPoikilospermum)to the Urticaceae, woody climbersof Urera Gaudichaud(Urticaceae).The which made the Moraceaemore homogeneousbut the remainderof the species (approx.60%) are terrestrial Urticaceae less so. As the Conocephaloideaeare dis- trees, shrubs,climbers, or (a few) subshrubs.

INTRODUCTION TO THE FAMILY MORACEAE

INTRODUCTION TO THE FAMILYMORACEAE

5

of the fig treeandthe supporting Mostof the speciesof thefirstcategoryaresmall the superstructure (= understory) to medium-sized (= canopy)trees,but cylinderof rootsarein lineorif thefig treemanagesto somecanattaina considerable height(to60 m), such senddownaerialrootsalongadjacenttreesandthereas speciesof Brosimum(subg. Ferolia), somespecies withproducesa broadcarryingbase.Onlya few speof Ficus sect.Pharinacosycea,andMaquiracoriacea. cies arepowerfulenoughto replacehosttreesin high Inthisgroupa numberof architectural modelsas de- forest,suchas in the Neotropicsspecieslike Ficus scribedby HalleandOldeman(1970)andHalleet al. gomelleira Kunth& Bouche, F nymphaeifoliaMiller, (1978)arerepresented, suchasthemodelsof "Corner" andF schultesiiDugand.Hemiepiphytic fig speciesare (discussedbelow), "Cook,"and"Roux."It is note- alsooftenfoundonrockysurfaces(ashemiepilithes). worthythatsometallormedium-sized treespeciescan However,severalspecies,suchas the ? lianescentF also be shrubs,or at least startfloweringas shrubs. schippiiStandley,areoftenterrestrial. Theshortstem Thefacultative frutescent habitmaybefoundinpartof and the broad crown, general features of the thespeciesrange,asinHelicostylistomentosa,Maquira hemiepiphytic Ficusspecies,allowdevelopment asterguianensis (subsp.costaricana), andthe OldWorld restrialtreesonlyin openhabitats. Antiaris toxicaria Leschenault, or almostthroughout Therepresentatives of theotherneotropical subdithespeciesrange,asin Perebeaangustifolia. visionof Ficus,about20 speciesof subg.PharmacoNumerouspaleotropical Ficus speciesareshrubs, syceasect.Phamacosycea, are(normally)terrestrial, someof themrheophytes. About20%of thenon-Ficus oftenelementsof riverineandmontaneforest,estabwoodyspeciesare(orcanbe) shrubs(orsubshrubs). lishingin ? disturbed places,a traitthatcanberelated of thesmallseeds. About 100 species are(or can be) climbers,the totheneedforlightforgermination majoritybeingroot-climbers of Ficus sect.Kalosyce Leaves (Miquel)Cornerandsect.RhizocladusEndlicher (CorThe leaves are usually altemateandthenarranged ner,1965),theothersof thestraggling type,asinMacTheleavesmaybe arranged in lura andFicus subsect.Palaeomorphe(King)Corner. in spiralsordistichous. spiralson the the mainaxes but in two rowson the InthegenusMaclura, thornsfacilitateclimbing. lateralbranches, asinthespeciesexhibitingthemodels Hemiepiphytism in Ficus of "Roux"and"Cook"(Halleet al., 1978).In some dueto shortSubgenusUrostigma (with about 280 species speciesof Dorsteniatheleavesarerosulate worldwide)is characterized by thehemiepiphytic life eningof the stem.Oppositeleavesareuncommonin form(also foundin a subdivisionof the OldWorld Moraceae.Theyarefoundin Bagassa,in the paleoandin a number Ventenat, "subgenus" Sycidium).Thislife formimpliesthatthe tropicalgenusBroussonetia Ficusspecies.In somepaleotropical plantsstartas epiphytes,whichusuallysenda leading of paleotropical aerialrootdownalongthebranches and/orthetrunkof Ficusspecies,theleavesmaybe (sub)verticillate. Thelaminais mostlybasallyattached, butitis peltate thehosttree.Whenthisroothasreachedthe soil and moreamplenutrients, theplantcanbegin in someDorsteniaspeciesorrarely(as anaberration) canpriovide morerapidgrowthandsoonerorlaterreachtherepro- in someotherspecies,suchas Castillaelastica. Thelaminais simpleandmostlyentire.Itis someductivephase.Thesemorerapidlygrowingplantssend orpalmately downadditional and timesmoreorlessdeeplyincised,pinnately aerialrootswhichcananastomose mayforma networkof rootsandeventuallya cylinder (orpedately),eitheronlyinjuvenilestagesof developof rootsaroundthetrunkof thehosttree.Atthatstage mentor alsowhenadult.Theincisionscanrundown thefig treemayoverpower (orkill)thehosttree.Such to the petioleandthe laminasegmentscan even be orstran- pseudopetiolulate(in Dorstenia bonijesu). fig speciesareoftenreferredto as stranglers The venation is mostly pinnate, sometimes glingfigs, suggestingthatkillingof thehosttreehapor in peltatelaminas pensby strangulation, cuttingoff the sapstreamsof (sub)palmate,(sub)trinervate, Thevenationis basicallybrochidodromthehosttreeby preventingthehosttree'strunkfrom (sub)radiate. thesubmarginal thesecItis morelikelythatinmostcasesfig trees ous,although loopsconnecting expanding. hosttreesby starvation, overpower by a moreefficient ondaryveinsaresometimesfaint.Insomespecies(of host Dorstenia),thelateralveinsmayrunstraightintothe andwater,andovershadowing uptakeof nutrients The treesby theirbroadcrowns.Killingof thehosttreeis margin(oftenendinginahydathode-like structure). (thetertiary oftena "suicidal" process,as the deadhosttreemay tertiaryvenationvariesfromscalariform andparallelbetweenthesectakethefig treedownin its fall.Thisneednotbe the veinsrunningtransverse case if the fig treestartedits life close to the soil, as ondaryveinsat? regulardistances)to reticulate,orin oftenhappensinriverinehabitatsorsavannas.Inhigh severalFicusspecieslargelyparallelwiththe(main) veins. forest,fig treescan(occasionally)remainstandingif secondary

6

FLORA NEOTROPICA

Theleafmarginis usuallyentirein sclerophyllous leavesandoftendentate,serrate orcrenateinmesophyllousones.Intaxawithentireleafmargins,themargin canbe dentateinjuvenilespecimens,andthelaminais thenoftenthinner. shortinleaveswithellipticto Thepetioleis normally lanceolate i coriaceous laminaswithanentiremargin. Itis oftenrelativelylonginmesophyllous leavesandin thosewitha cordiform to suborbicular lamina,suchas

forcertainfungiandmightsubstituteforpollenas attractant (seediscussion underPollination). Fungigrowingon densepluricellular trichomesarealsofoundin Pourouma(Berg,1990b;Berget al., 1990).

Glandular Structures Inaddition tothe"glandular" otherstructrichomes, tureshavebeenorcanbe indicatedas "glandular." In allFicusspeciesonecanfindglandular (almost?) waxy in Bagassa and species of Dorstenia andFicus. (orshining)spotson thelamina(andin paleotropical Scaleleavesarepresentas prophylls,as bractson taxaalsoonnodesof leafytwigs).Onthelaminathey inflorescences oramongflowpeduncles,subtending occuronthelowersurface,atthebaseof themidrib(in ers,astheproximalleaveson stemsandbranches, e.g., thehemiepiphytic monoeciousgroups,includingthe as scalesof lateralbuds,suchas in Morus.In several neotropical subg. Urostigma sect. Americana), only Ficus species,the prophyllsformcalyptratecovers in the axils of the basallateralveins (suchas in the enclosingtheyounginflorescences. neotropical subg. Pharmacosycea sect. Pharma-

Stipules

Thestipulesarefullyamplexicaulin manygenera Inothergeneratheyvaryfromnotfully (orsubgenera). to lateral.Thepairsof stipulesarefreeor amplexicaul fused.Theyvaryin lengthfrommorethan10 cm in someFicus speciesto less than1 mm,as in several Dorsteniaspecies.Theycanbe (sub)persistent, as in most Dorstenia species.

cosycea),and/or(also)intheaxilsof otherlateralveins andmajorfurcationsof thevenation(in variousOld Worldsubdivisions).As they are (almost?)always presentin oneplaceoranother,theyarelikelyto have somefunction,as suggestedby Coiner(pers.comm.) in emittingsubstances to attractpollinators. Thepositionof theglandsmayhavediagnosticvalue,as it has forthetwoneotropical subdivisionsof Ficus. Anothertypeof "glandular" structure is represented bysmallerorlarger"pockets" accumulating yellowdye,

of thestipules,suchaspersistence, Features fusion, areoftendiag- occurringin Maclura, Batocarpus, and Clarisia, emdimensions,shape,andindumentum, beddedin tepals,bracts,and/orconnectivesof anthers. nosticallyimportant. Inthesegenera,yellow-colored substances canalsobe foundelsewherein the plant,as in the barkof stems Indumentum androots.Theyellowsubstancemayfunctionas deThreetypesof trichomescanbe recognized: terrentagainstherbivory (seediscussionin thegeneric 1. Unicellular hairsof variouskindswithregardto description of Maclura). length,firmness,andcolor.Thesehairscanbe uncinate.Presenceorabsenceof uncinatehairscanbe diFlowers atvariouslevels,rangingfrom agnostically important The flowers are small to very smallandusually tribeto species.Uncinate hairscanbefoundonallhairy unisexual. bisexualbutfunctionally Morphologically itsvegetative partsof theplant,inparticular parts.They unisexual in flowers occur a paleotropical subdivision tendto be moreabundant injuvenilestagesthaninthe of Ficus, and bisexual flowers may develop occasionadultones. in taxanormallywithuni2. Pluricellularhairs,elongate,moniliform,and ally (andteratologically) oftenbrown(ish).Suchhairscanbe foundin several sexualflowers. Theflowersarepedicellate orsessile.Pistillate flowFicus species andin Poulsenia.Theyoccurpromiers are often connate and/or adnate the to of receptacle nentlyin somegeneraof theCecropiaceae (Cecropia the inflorescence. andPourouma). 3. Pluricellular globose-to ellipsoid-capitate trichomes("glandular hairs").Thepresenceorabsence andtheshapeof thesetrichomeshavediagnosticsignificanceinmanygenera;theshapee.g.to distinguish thesectionsof PerebeaandPseudolmedia andthepresenceto distinguish thetwosubgenera of Ficus.InseveralSoroceathese"glandular hairs"aredenseon the of pistillateflowers.Theyprovidea substrate perianth

Perianth Theperianthis tepaloid.In staminateflowersit is oftenmembranaceous andin pistillateones ofteni fleshy,morepronouncedly so in thefruitingstate. Themostcommonnumberof tepalsis four,less commonly five,threeortwo,andtheninstaminate flowerscommonlywithan equalnumberof antitepalous

TO THE FAMILYMORACEAE INTRODUCTION

7

stamens.Tetramerous and5-merousflowersarecommonthroughout theorderUrticales.Smallernumbers of tepalsarecommon;increaseof numbers is lesscommon(e.g., foundin Naucleopsis). Reduction oftheperianth innumber andsizeof tepals mayleadto vestigialperianths ornakedflowers,which occurin severalgenera.In the genusBrosimumthe wholerangefromwell-developed 4-merousstaminate flowersthroughreduction innumberandsizeoftepals to nakedflowers(witha singlestamen)is found.In thisgenustheindividual flowerscanstillberecognized.

Increaseof thenumberof tepalsis foundin some genera,suchasFicusandNaucleopsis.Itmayoccurin bothstamninate andpistillateflowers,suchasinNaucleopsis, in whichgenusit also leadsto the loss of the normalandclearpositionof thetepals,in particular in thepistillateinflorescences. ThefactthatinNaucleopsis thestaminate flowersareenclosedby theperipheral involucral bractsuntilanthesiscanberelatedto a looser arrangement of floralparts.Inthisgenusthenumber of stamensmaybereduced,whereasthetepalnumbers haveincreased. In other genera, such as Castilla and Pseudolmedia, Thetepalscanbe freeorpartlyto entirelyconnate. thereductionof the perianthof staminateflowersis In4-merousflowersthetepalscanbe decussatewith linkedto a processof loss of thenormalfixedrelative twoinnerandtwooutertepals.Thepairsareoftendifpositionof the floralparts,resultingin scatteredsta- ferentinshape,e.g.,inHelicostylis andSorocea.Inthese menson the receptacleamongbract-likestructures, generathe pairs of stamensmay be more or less apparently loosetepals,orwithoutanyremnantof the anisomorphic, oronepairevenstaminodial. perianth(in theAfricangenusTrilepisium Thouars). Thetepalsorthefreepartsof thetepalscanbe imDisorganization of thenormalfloralstructure is also, bricateor less often(suchas in Trophis)valvate.Imbut less pronouncedly, foundin staminateinflores- bricatetepals(of flowersnotenclosedbybractsand/or cences of Batocarpus and Clarisia. thereceptacle of theinflorescence) usuallyhaveciliolate Incasesin whichtheperianthlosesits functionto margins.Themarginsof valvatetepalsareprovided protectstamensbeforeanthesis,protection is provided withcurledhairs,thoseof oppositemarginsentangled otherwise, suchasbytheshapeof thereceptacle, deeply and closingthe slits in between.In Helicostylis,in to (sub)urceolate or(inCastilla)bivalvate. cup-shaped whichthetepalsof thepistillateflowersareonlybaBracts are often importantin this respect, either sallyconnateorentirelyfree(incontrasttomostother interfloral bracts,oftenpeltate(suchasinBrosimum), genera),stripsof densecurledhairsarefoundon the orbractsattheperiphery as(marginal) ofthereceptacle, innersurfaceof theinnertepalsandcloseoff theinteinvolucralbracts(suchas in the tribeCastilleae),or riorof theflower,barring accessto theovary(seeBerg, ostiolarbractsin Ficus.Theoccurrenceof staminate 1972:80, 84, and91). Thedifferencesbetweeninduflowersin pits in the receptaclein some speciesof mentum andvalvatemarginsis alsofound onimbricate Brosimum (subg.Ferolia)mayalsobe relatedto profor bracts (cf. Berg, 1990b). tectionof thestamens. flowersoftenhavea ? distinctly Tepalsof staminate Thestaminateflowersof the generaHelicostylis, havwell- cucullateapex.Thisfeatureandthetwotypesof induMaquira,andPerebeaof thetribeCastilleae, of developedandnormallystructured perianths,andthe mentumonthemarginscanbe relatedto protection or insect larvae. stamens insects against damage by staminateinflorescencesareopenbeforeanthesis.In theothergeneraof thetribetheperianths of thestaminateflowersaremoreor less disorganized and/orreducedandthe staminateflowersareenclosedby the receptacleand/ortherelativelylargeandbroadinner involucral bractsbeforeanthesis.Theinnerinvolucral of speciesof Perebea bractsof staminate inflorescences and sect.Noyeraarerelativelylong,narrow, incurved, coverthestaminate flowerswithwell-developed perifunctionthe anths,buttheymaynothavetheprotective broadened involucral bractsapparently have. Reductionof the perianthhappensmoreoftenin staminate flowersthaninpistillateones.Nakedpistillateflowerscanbe foundintheAfricangenusTreculia Decaisneandinsomepaleotropical speciesof Ficus.It is oftenuncertainwhethertheperianthis stillpresent in pistillateflowerswhichareentirelyfusedwiththe receptacleof theinflorescence (suchasinBrosimum).

Stamens Thestamensarebasicallyantitepalous andthustheir numberis equalto thatof the tepals,oftenfour,less commonlyfiveorthree.Thenumberof stamenscanbe innumbermaycoincide reducedtoone.Thereduction of theperianth, butit mayalsooccur withthereduction suchas in Bagassa,orit mayco-occur independently, withanincreaseof tepals,suchas in Naucleopsis. Twotypesof stamenscanbe distinguished: 1.Thosewithfilamentsthatareinflexedinthebud andbendoutwardssuddenlyandelasticallyat anthestasis,throwingoutthepollenin theair("urticaceous mens").Thistypeof stamenis foundin all generaof theMoreae(butnotin allspecies,seeunderMaclura); it is characteristic Suddenreleaseof of theUrticaceae. pollenis also reportedfor someUlmaceae:in some

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8

(Cuellar, 1967;VazdeArruda speciesof CeltisLinnaeus (Franco, & Sazima,1988)andinLozanellaGreenman asonly is somewhat different 1992);butthemechanism asinthe isinflexedandnotthewholestamen thefilament modeof explosive(or type.Theurticaceous urticaceous a well-developed periballistic)pollenreleaserequires undertension.The anthtokeepthelonginflexedstamens commonly foundinflowerswithsuchinflexed pistillode, stamens,seemsalsoto playa rolein themechanism, by keepingtheinflexedstamensin therightposition. 2. Thosewithfilamentsstraightin thebudor inflexedbeforeanthesis,suchas in Dorstenia,butbeAs straightfilamentsoften comingstraightgradually. elongateduringanthesis,theelongationleadingto inflexedstamenscanberegardedaspremature. to latrorse tointrorse. varyfromextrorse Theanthers of theantheris foundinBrosiA peculiarconstruction mumalicastrumsubsp. alicastrum.The antheris pelt-

canbe ateandthethecaearefused.Thisconstruction of anemophily (seeunderPollinarelatedto promotion tion,below). Pistillodes

forsomegroupsof taxa, arecharacteristic Pistillodes stamens, suchasthetribeMoreaewithits urticaceous andin some subdivisionsof Ficus, includingsubg. Pharmacosycea sect. Pharmacosycea and the paleo-

occurinsome group.Theyregularly tropicalSycidium speciesof Dorstenia,andarerareor absentin most othergeneraor species.However,in Helianthostylis presentwithstylesand spruceitheyareprominently stigmasto 2 cm long,andtheymightplayaninsectroleinpollination. attracting Pistil

Theovarycontainsa singleovule,witha subapical almostlaterattached position;theseedis occasionally ally(Castillatunu,cf. Berg,1972).Thepositionof the character whichdistinovuleis the only "technical" fromtherelatedfamiliesUrticaceae guishesMoraceae in thesefamiliesthe ovule has a andCecropiaceae; with position,whichseemsto be correlated (sub)basal thepresenceof singlestigma.In the Moraceaethere arebasicallytwostigmas,butthestigmaconnected with thesideof theovarynotbearingtheovulemaybe reducedin lengthorfinallyabsent,suchasfoundin some speciesof Dorstenia,in the genusMaclura,in the of thespeciesof Ficus,andin thesmallpaleomajority tropicalgenusFatoua. Thestigmasvaryfromfiliformto band-shaped to groupsof Fi(orin somepaleotropical tongue-shaped custo clavateorto infundibuliform).

Theovarycanbe freeorpartlyto fullyadnatetothe Theovarymaybe? stipitubular (partof the)perianth. of withthedevelopment tate,a featureoftencorrelated theovaryintoa dehiscentdrupe(seediscussionunder FruitsandInfructescences). Inflorescences

in Moraceae(aswellas inother Theinflorescences familiesof theUrticales)basicallyoccurin pairsinthe leaf axils. However,solitaryinflorescencesarealso andtheyarecharactercommon,suchasinBrosimum, istic for Ficus subg. Pharmacosycea sect. Pharma-

cosycea.Pistillateinflorescencesaremoreoftensolitarythanarestaminate ones(evenin thesamegenus). Pairedinflorescencesapparentlyarisefromthe two which (prophyll-bearing) basalnodesof alateralbranch, (cf.Eichler,1878).The mostlydoesnotdevelopfurther prophyllssometimessubtendinfloresences,onlyiniInseveralspeciesof Ficussubg. tiallyorpersistently. toabout (sometimes Urostigma, theyarewell-developed 2 cm long)andfused,forminga calyptratecoverenclosingthetwoyounginflorescences(figs). Inseveralgenera(e.g.,inthoseof thetribeCastilleae andinFicus),short-shoots (spurs)maydevelopin the leaf axils andproducemorethantwo inflorescences orsuccessively, bisexualones, (almost)simultaneously ones.It staminate ones,orbothpistillateandstaminate is notcertainwhethersuchleafless(butstipule-bearing) branchletsdevelopfromthe samemeristemsas bearingonlythepaired the(abortive) axillary branchlets or solitary"axillary" inflorescences.Ontheseshortmayalsooccurinpairs.These shoots,theinflorescences short-shoots maycontinueto produceinflorescences them afterthesubtending leafhasfallen,thusproducing on the lesserbranchesor even on the mainbranches andthetrunk.Theseshort-shoots arerarelybranched Inandtheyshowregularpattemsin theiroccurrence. florescencesmayalsobe bome(oftenin pairs)on anothertypeof leaflessbranchlet, oftenwithlongerinternodesand/oroftenbranched. Theseleaflessflowering branchlets oftendo notshowregularpatternsin their presenceandareusuallyfoundon themainbranches or on the trunk,even at the base of the trunk.Such branchletsoccurin Clarisiaracemosa(and in the in somespeciesof Artocarpus andin nupaleotropics merousspeciesof Ficus). Caulifloryis rarein theneotropicalMoraceae,restrictedtothegenusClarisia,andit occasionally found in Sorocea.Someneotropical Ficusspeciesareramiflorous(to subcauliflorous). Caulifloryis far more commonin thepaleotropical Moraceae andis foundin theAfricangenusTreculia,in theAsian-Australasian genusArtocarpus,andin particularin Ficus, which

INTRODUCTION TO THE FAMILYMORACEAE

9

includesthewholerangeof possibilities, fromaxillary lateones,andstaminateinflorescencescontainmore inflorescences to ramiflory to caulifloryto tronciflory flowersthanpistillateones.However,in bisexualinflowers florescences of Ficus,thenumberof staminate to flagelliflory(geocarpy). InMoraceae, theinflorescence hasdiversified con- is (in almostall species)farless thanthanthatof the siderably withregardto structure, shape,anddegreeof pistillateones, a traitthatcan be relatedto modeof inthisgenus.Thenumberof pistillateflowpollination condensation andfusion. Thebasicstructure of theinflorescence of theUrti- ersin unisexualandbisexualinflorescenesis to some calesis dichasially branched, bearing pistillate andstam- extentrelatedto the size of the fruits;the (African) speciesof Dorsteniahaveoneora few inateflowers,orsometimes bisexualflowers(cf.Berg, macrospermous butthemicrospermous speciesofthis pistillate flowers, 1977b).Themaintrendsinthedifferentiation fromthis genususuallyhavefarmore.Reductionof thenumber basicconstruction are: 1. Condensation by shorteningandfusionof the of flowersis in generaltheresultof reductionof the axesof theinflorescence, in accordance withthegen- size of theaxisof theinflorescence.Reductionof the eraltrendof crowdingof smallflowers,inparticular of numberof flowersto onlyonecanberelatedto protectionagainstherbivory.Uniflorouspistillateinfloresthosewithsimpletepaloidorreducedperianths. forPseudolmedia andare(in 2.Transition fromthebisexualtotheunisexual state. cencesarecharacteristic of Clarisia, Maquira, the Neotropics) found in species 3.Changefromanactinomorphic structure toazygobisexual Perebea, and Trophis. The (basically) inflomorphic one,through atransition tomonochasial branchoftencontaina rescences of macrospermous species ing(?)to elongateinflorescences asracemesandspikes, with inflorescences andsubsequently of therachisto globose singlepistillateflower.Staminate a shortening oconeora few flowersarerare;suchinflorescences ordiscoidcapitatestructures, beinginvolucrate (asby curin somePerebeaspecies. enlargement of someof thebracts)ornot.Discoidreceptacles canbecomeurceolate, asinFicusandtheNew Caledonian monotypicgenusSparattosyce Bureau. Distribution of Sexes Theplantscanbe dioeciousormonoecious.Inthe Twobasictypes of inflorescencescan be distinlattercasestaminate canbe andpistillateinflorescences guishedin Moraceae(cf. Berg,1977b): presentinthesameleafaxil,oftenso in somegeneraof 1.Basicallybisexualandbasicallycircularin outthe Castilleae(as Castilla, Helicostylis, andPerebea), line.Theymay(still)showdichasialelements. Thepisby several mostlya singlepistillateone surrounded tillateflowersareplacedinthecenterandthestaminate staminate of arrangement of pisones,thusreminiscent onesareeitherconfinedto theperipheryormayalso tilsandstamensin flowers.Inthesegenera(aswellas occurscatteredamongthepistillateones.Thistypeis in thepaleotropical trees genusAntiarisLeschenault) forthetribesDorstenieae andFiceaeand mayshiftfromdioeciousto monoecious,fromformcharacteristic is foundinthepaleotropical Blume inginitiallyonlystaminate generaBleekrodea inflorescences (orpistillate?) andFatoua,bothbelongingto thetribeMoreae.These to bothstaminateandpistillateoneslater.InPerebea func- guianensis,althoughnormallydioecious,staminate inflorescences maybecomesecondarily unisexual, theinflores- inflorescences tionallyormorphologically so. Moreover, and maybebomeontheupperbranches to pistillateoneson the lowerones of the sametree.In cencesmaysecondarily changefromactinomorphic zygomorphic, frequentlyso inDorstenia. Brosimumalicastrum subsp. alicastrum, a shift from 2. Basicallyelongate,racemoseorspicate,andusu- initialproduction to of (only)pistillateinflorescences A fewpaleotropical allyunisexual. species(inStreblus (only)staminate inflorescences later,whentreeshave havebisexualinflorescences. reachedthecanopy,hasbeenrecorded(Peters,1991); Loureiro andin Treculia) As a conse- thisphenomenon Theflowersshowanadaxialorientation. canbe relatedto promotionof wind quence,a sterilestripis foundattheabaxialsideof the pollination.InDorsteniacayapia,theonly speciesin axisof theinflorescence.Throughfurthercondensa- thegenuswithunisexualinflorescences, plants(examtionthe inflorescencescanbecomespadix-like,glo- inedundergreenhouseconditions,Hoen,1983)start orevendiscoid-capitate, andthenwith to producestaminateinflorescencesin theflowering bose-capitate, orwithoutaninvolucre.Thissecondtypeof inflores- seasonandcontinuewiththeproductionof pistillate andCastilleae, oneslater;bisexualinflorescences cenceis characteristic oftheArtocarpeae canbe found(only andit is commonin theMoreae. Itis notknownhowthesituationis in occasionally?). Thenumberof flowersperinflorescence mayvary nature,andto whatextentfloweringof staminateand is simultaneous. Judgingfrom fromoneto thousands(inFicus).Ingeneral,bisexual pistillateinflorescences tooccurinabout appears flowersthanpistil- labeldata,dioecyandmonoecy bearmorestaminate inflorescences

10

equalfrequencyinPoulseniaarnata,whichcouldbe causedby shiftsfromone stateto theother. Theneotropicalspeciesof Ficusareusuallyfunctionallydioecious,althoughtheinflorescences contain bothstaminateandpistillateflowers.Thissituationis createdby thefactthatindividual treesflowersimultaneouslyandthestaminate flowersareatanthesiswhen fromthepistillate thefruitsareripe,thusseparated phase of floweringby someweeks. Androdioecyis foundin Helianthostylis andalso occursintheAfricangenusBosqueiopsis DeWildeman (cf.Berg,1977c).Morphological gynodioecy (butfunctionaldioecy)is foundinabout500/ooftheFicusspecies (cf. Berg,1990a).A moreuncleartypeof gynodioecy is foundin Treculia afiicana Decaisne(cf.Berg,1977c). Ingeneral,dioecyandmonoecyappearto berepresentedaboutequallyin thefamily,asintheUrticaceae andtheUrticalesas a whole,butwiththenotableexinwhichallspeciesarestrictly ceptionof Cecropiaceae, inthedistribution dioecious.Theindicatedpatterns of sexescanberelatedto spatialseparation of staminate andpistillateflowerswithregardto protection against of maleandfemalefunctions orto separation herbivory withregardto themodeof pollination.Outbreeding as thedrivingforcetowardsdioecy(cf. Bawa& Opler, 1977)is notasevidentin Moraceae(andUrticales)as it maybe in otherfamilies(andorders).

FLORANEOTROPICA

ersandinflorescences andof whichthelarvaepenetrate in or amongflowers,wheretheycanfindbothfood andprotectionagainstpredators. However,insectlarvaearerarelyencountered in pistillateinflorescences andin staminate inflorescences beforeanthesis. Theprotection of stamensis primarily providedby theperianth. Theslitsbetweenfreepartsof theperianth areclosedoff by stiffhairson themarginsin flowers withimbricatetepals,orby entangledcurledhairson oppositemarginsin thosewithvalvatetepals.Slitsbetweenflowerscanbemoreorlessefficientlyclosedby of theflowers,(partial) crowding fusionof flowersand/ orhairsorscariousbractsinbetweentheflowers.Enclosurewithinbudcoversorcoveringbybracts,inparticularpeltatebracts(withimbricateorvalvateupper parts),mayprovideadditionalprotection.If theperianthbecomesdefunctduetoreduction, asoftenoccurs, theprotectivefunctionof theperianthis transferred to otherstructures. Thesecanbe bracts,as inBrosimum, andalsoflowersplacedin smalldepressions of thereceptaclein thisgenus.Inothercasesthewholeinflowithperescence,thereceptacle, oftenin combination as ripheral(involucral) bracts,canprovideprotection, canbe foundin Castilla,Naucleopsis,andPseudolmedia.

Therelativelysimpledesignof themoreorlesslax staminateinflorescencesin Moreaecanbe explained by thefactthattheflowersneedspaceanda well-developedperianthto performthe explosivereleaseof pollen.Theseinflorescences areusuallynotsuitablefor Patterns in Differentiation of Inflorescences insectbreeding.Pistillateinflorescences in thisgroup Themostlikelyfunctional factorto explainthepat- of taxaareoftenmorecondensedandmorecomplexor ternsinthedifferentiation of flowersandinflorescences variedin structure, anddesignsto protecttheinterior of Moraceaeis theprotectionof anthers(untilanthe- againstpenetration by insectlarvaeareoftenevident. sis) andof pistilsanddevelopingseeds/fruitsagainst Soroceais a remarkable genus,as it hasinfloresherbivoryinflictedin particularby larvaeof small cencesof thetypeoftenoccurring in theanemophilous beetlesandflies (Berg,1990b).Patternsanddesigns species,butwithoutanyadaptation to windpollination. whichcanberelatedtothatprotection inMoraceae (and Forpistillateflowersandinflorescences, thedesigns *otherUrticales)occurinnumerousotherplantgroups topreventpenetration by insectlarvaearemorevaried. withflowersdenselyset(incondensedinflorescences) Besidesthe featuresmentionedabovefor staminate or groupswith flowers with numerouspistils (as flowers,thefusionof thepistilwiththeperianthand Annonaceaeand Monimiaceae)or even in gymno- thepresenceof hairsinsidetheperianthcanbe added. sperms.Thisprotectionappearsto be oftenclosely Striking arethestripsof curledhairsoppositeeachother linkedto themodeof pollination,aswillbe discussed on theinnersurfaceof thetwoinnertepalsof thepisbelow.Theomnipresence of insects,atleastin humid tillateflowersof Helicostylis, whichincontrast to other tropicalhabitats,andtheircapacityasherbivorescan generaof thetribehasonlybasallyconnateorentirely be observedby dissectinginfloresencesof Moraceae freetepals.Entanglement of thehairsof theopposite conserved influid.Insectlarvaeare stripsclosesoff theslit abovetheovary. (andCecropiaceae) foundincondensed commonly staminate inflorescences As pistillateflowersdonotneedto openwidely,as atandbeyondanthesis.Judgingfromreportsonpres- do thestaminate to exposethepollen,theapicalaperenceof insectsandinsectlarvaeon andin staminate turescanbe verynarrow, just wideenoughto let the inflorescences of CecropiaLoefling(Andrade,1984; stylethrough.Thenarrowslitsbetweenthestyleand Fiebrig,1909;Wheeler,1942)andof Artocarpus (van theperianthareoftenbarredby hairson thestyleorin derPijl, 1953), the larvaeencounteredarethoseof theperianthapex.Forthesamereasonpistillateflowbeetlesandflieswhichlayeggsonthesurfacesof flow- erscanmoreeasilyfuse.

INTRODUCTION TO THE FAMILYMORACEAE

11

Uniflorouspistillateinflorescences arenotuncom- throughthe apexof the exocarpor ejectedwhenthe monin Moraceae.Thisreductionimpliesisolationof vascularbundlerunningthroughtheapexof thefruit flowersandcanalsobe relatedto protection,as it re- no longerkeepstheendocarp inplace.Largeendocarp moveshiding-places forlarvae.Features of thesurface bodiesaresmooth,blackish,andaresqueezedoutof of theperianthin pistillateflowers,likedenseindum- the exocarp,rarelyejected.Smallones areusually? entumoraverysmoothsurface(suchasinClarisiailici- tuberculate,whitish,grayishor brownish,and are withregardtoprotection. ejectedbytheexocarp.Smallendocarp folia),mayhavesignificance bodiesoccurin Moreover, chemicalsubstances, suchastheyellowdye mostDorsteniaspecies,as well as in thesmallpaleoaccumulated in perianthsandbracts(as in Maclura), tropicalgenusFatoua.Thedehiscentdrupeanddrupemayalsohaveabearing onprotection againstherbivory.let are found in some genera of the Moreae,the Inbisexualinflorescences, staminate andpistillate Artocarpeae, andtheDorstenieae. However,in several flowershaveto be separated in onewayoranotherto othergeneraof thesetribes,includingseveralsubdivipreventinsectlarvaefrompenetrating throughoralong sionsof Ficus,onecanfindfruitsshowingsimilarities thestaminateflowersintothepistillateflowers,dam- tothedehiscentdrupebyhavinga stipitatebaseand/or agingovariesandthedevelopingseeds.Thissepara- anunequallythickexocarp.Thedehiscentdrupesand tionis realizedinvariousways,inspaceandintime.In drupeletsaresubtended orenclosedby enlargedperiBrosimum thestaminate flowersareplacedontheouter anthswhichdonotorhardlybecomefleshyandmostly surfaceof thereceptacle, andthepistillateflower(s)are remaingreenish. Thesameappliestotheinflorescences completelyimmersedin thereceptacle.InDorstenia of Dorstenia;theydo notbecomereallyfleshystructhepistillateflowersaresessileandthestaminate ones turesanddonotchangecolor. Forthetaxawithdehiscentdrupesordrupelets, the pedicellate;theperianthsof thepistillateflowersare adnateto thepedicelsof thestaminate flowersandthus individual(dry)endocarpbodiesarethe entitiesfor closeoff thesurfaceof thereceptacle fromwhichonly dispersalandareonlyreleasedby forceexertedby the thestylesandstaminate flowersemerge(seeFig.33A). exocarp.Structuressurrounding themmay provide Theextendedprotogyny inFicuscouldberegarded protection duringthedevelopment of thefruit,butare asa wayto separateanthesisof staminate flowersand not(orhardly)involvedin dispersal.ThesmallpaleoandAntiaropsisK.Schumann, thatofpistillate flowers,afeature subsequently exploited tropicalgenera,Bleekrodea todevelopthepollination inthisrespect,astheperianth bracts areexceptional turnsredsystem.Theinterfloral andbristlesinthisgenusarelikelyto haveotherthana dishandprovidestogetherwiththewhiteexocarpand theblackishendocarp contrastcolorsand(deceiving?) function. protective Thefactthatbisexualinflorescences of Moraceae visualattraction. InFicusthefruitlets,beingeitherdrupaceous tendto be morecomplexthanunisexualonesmaybe (such orachene-like relatedto a morecomplexsituationwithregardtopro- asin severalpaleotropical subdivisions) tectionagainstherbivory. (suchasin theneotropical species),functionmostlyas seedsof a fleshy fruit;the syconiumthusbeingthe functionalentityfordispersalby fruit-eating animals. Fruits and Infructescences However,thefruitletsmayfunctionasindividualentiThedrupecanberegarded asthebasictypeof fruit tiesin caseof exozoochorous dispersalby seed-eating in Moraceae. Theendocarpis mostlycrustaceous, but animals(birds,suchaspigeons)removingfruitlets from Theexocarpcanbe verythin thesyconium.Thefruitletsusuallyhavea stickysurcanbe woodyorleathery. andjust providea stickysurfaceto fruitsor fruitlets facebywhichtheycanbeattached tothebodiesofthese which,therefore,areratherachenesthandrupesor seed-eatinganimals.Ina small(paleotropical) group SuchfruitsarefoundinmanyFicusspecies, of rheophyticspeciesof Ficus (subsect.Macrostyla drupelets. ones. includingtheneotropical Coiner)singlefruitletsareadaptedas entitiesfordisfruitsis oftenwhit- persal(see underDispersal). Theexocarpof freedrupaceous ish,butcanbeblackish,suchasinspeciesof thepaleoThe black drupes of Trophisinvolucrata are subThe tendedby greentepals,whichdo not enlargeduring involucrata. tropicalgenusStreblusandin Trophis freedrupaceous fruitsaremostlydehiscent.Dehiscent thedevelopment of thefruit.Inallothercasesthefruits drupesordrupeletsaremoreorlessdistinctlystipitate arepartof one-tomany-seeded comfleshystructures, andtheexocarpis alwayswhitishandunequally thick. prisingfleshyfruitingperianths ata minimum,butofItis thickestatthesidesthroughwhichthetwovascu- tenalsobractsandreceptacles. Thesimplestentityis a larbundlesleadto the stigmas.At the apexit is rela- fruitentirely(orpartly)enclosedbyanenlargedfleshy tivelythin.Thethickenedpartsforma kindof pincer perianth. Suchentitiescanbeindicated aspseudodrupes ontheendocarp puttingpressure body,whichis pushed (cf. Spjut,1994).However,the fruitsarenot always

12

FLORANEOTROPICA

orreceptacles Insomespeciesfruitingperianths can entirelyfusedwiththeperianth,butcanbe entirelyor orgreen(ish), partlyfreeinsidetheperianth such (suchas inHelicostylis). occurintwocolorvariants, red(dish) Pseudodrupes may occurseparately,pedicellate,or as in Clarisia biflora,Pseudolmedialaevis, andBrosisessile,on a moreorlesselongaterachis(e.g.,inSoro- mum alicastrum (subsp. alicastrum). The infructesFicusspeciesshowsimiinflorescences cea andTrophis)orinuniflorous (e.g.,in cencesof somepaleotropical Clarisia andPseudolmedia). Theyoftenoccuraggre- larcolorvariants. inflorescences(such gatedin few- to many-flowered asin manyspeciesof theCastilleae).Theseseed-conAnatomy tainingfleshystructures becomemorecomplexif the Several wood anatomicalstudieson systematic flowersarefusedwitheachotheras in someMaquira andUrticaleshavebeenpublishedin recent thatcanbeindi- Moraceae species,resultingin aninfructescence thoserelevantforneotropical taxa If theyarenotonly years.Inparticular catedas a "multiple pseudodrupe." fusedwitheachotherbuttheperipheral onesalsowith arelistedin a surveyof woodanatomicalcharacters the(fleshy)receptacle(as in Naucleopsis), thestruc- of Moraceaeby TerWelleet al. (1992)in theFloraof turesaremorecomplexthan"multiple pseudodrupes."theGuianas. A similardegreeof complexityis foundintheneotropicalwoodygeneraof theDorstenieae (Brosimum, POLLINATION Helianthostylis,andTrymatococcus).Here,thepistilIna limitednumberof taxaof theMoraceaeatleast lateflowers,one or severalperinflorescence,areenFor suggestanemophily. tirelyimmersedin andfusedwiththereceptacleand thedesignof floralstructures forma one-ormore-seeded wholeinwhich Ficus a uniquemode of entomophilyis well-docudrupaceous thosewith thereceptacle constitutes thefunctional exocarpwhich mented.Formanyothertaxa,in particular basedonpresentomophily maybearbractsand,in taxawithbisexualinflores- unisexualinflorescences, of thestaminate flowers.Struc- ence of breedingsites appearsto be the mostlikely, cences,alsoremnants mode;whereasforthe turallyevenmorecomplexorelaboratefruit-contain- althougha scarcelydocumented includingthemajority of taxawithbisexual ing structures,includingthose in whichinterfloral remainder, someothergeneralmodesof pollinabractsplayamoredominant role,canbefoundin some inflorescences, OldWorldtaxa(e.g., in Broussonetia, Treculia,and tioncanbe presumedorsuggested. Trilepisium).To avoidterminologyproblemsfor the variousmoreorlesscomplexfruit-bearing or-containAnemophily theindication"infructescence" ingstructures, appears Thetaxawiththeexplosivepollenreleasemechato be appropriate. Inmostcasesthesearetheentities nismas describedaboveare(predominantly) anemofordispersal,butnot so in Dorstenia. theindividualsof Theinflorescences areoften,suchasthoseof Arto- philous.Becauseof thisadaptation wherepollencanbetransported andin appearance thesetaxaneedhabitats carpus andFicus, "pseudocarpous" likeforestmargins, byairmovement, resemblingfruitsratherthanflowers. inparticular those Thecolorof theperianth partof thematurepseudo- alongstreams,orin ? openvegetation,mostly(semi-) drupeandof fleshyreceptacles containing ripefruitsis deciduousforest.Someare(deciduous)elementsof oftenred(dish), orange,oryellow,lesscommonlyturn- evergreenforest.However,sometaxawithexplosive ingblackish(suchas inSorocea), orremaining green- pollenreleasecanbe foundin evergreenforestunderish(suchasin Trymatococcus).IntheNeotropics, large growth,butthenasarulealongstreamlets whichcause infructescences (syconiaor figs) of Ficus tendto re- airmovement abovetherunningwaterthatcandisperse maingreenish,whereasthesmalleronesmaybecome thepollenthrownintomovingair.Thesameappliesto pinkish,red,orange,orwhitish,andthenusuallywith manytaxaof theUrticaceae. redorpurplishlongitudinal stripesorwithspots.The Insometaxapollenmaybereleasedfromlongpencolordifferencesof the infructescences in thisgenus dulousstaminate inflorescences movedby wind.This canbe relatedto dispersalagents,thered-colored ones couldbe thecasein, e.g.,Bagassa,whichcanbecome to birdsandthegreenishonesto bats. huge(emergent) treesandaredeciduous.InBrosimum Peripheral structures, involucresoronlytheinner alicastrum(at least in subsp. alicastrum)pollen is reinvolucralbracts,mayalso becomecolored(e.g., in leasedandtransported by wind(cf. Peters,1991).The Naucleopsis speciesandTrophiscaucana). InSorocea remarkable shapeof theanthers, peltateandwithfused therachisof theracemeandthepedicelusuallybecome thecae,aswellasa shiftto production of staminate infleshyandredororange,providingcontrastcolorsfor florescenceswhentreeshavereacheda considerable theblackfruitingperianths. height,couldberelatedto anemophily.

TO THE FAMILYMORACEAE INTRODUCTION

13

have thenopen anthers,or otherwisethey become through dustedbypollen.Theyleavetheinflorescences For the genus Ficus entomophilyis well-docutunnelsmadeby themaleinsectsin thewallof theinmented.Thepollinationsystemin thegenusis unique florescence(orthroughtheostioleof whichthebracts andcomplex.Itis brieflydescribedbelow. becomelesstightlypackedin somegroupsof Ficus). Thepollinatorsbelongto theAgaonidae(Chalcidof theostiole,theessential Exceptforthestructure Thisfamilyof fig waspscomoidae,Hymenoptera). relatedtothe andphenological requisites morphological prisesseveralgenera(cf. Wiebes,1994),thesegenera synstigma pollination systeminFicusare:heterostyly, beingmostlymoreorless clearlyassociatedwithone andextendedprotogyny.At anthesisof the pistillate of the subdivisionsof the genusFicus: in the Neoinonelayeratacertain flowersallstigmasarearranged tropicsthegenusTetrapus withsubg.Pharmacosycea distancefromtheinnerwallof theinflorescence. The sect.Pharmacosycea, andthegenusPegoscapuswith stigmasaremoreorlessstronglycoheringandconstiEachof thespecies tutea synstigmatic subg.Urostigma sect.Americana. layer,whichfunctionsasthe"platis in principleassociatedwitha singleFicusspecies. form"fromwhichthefemaleinsectsovipositandwhich However,in severalcasesmorethanonespeciesof fig mayallowunevendepositionof pollenandgrowthof witha singleFicusspecies,orsingle pollentubesfromcertainstigmasto ovulesof neighwaspis associated speciesof fig waspwithmorethanone speciesof Fi- boringflowers. cus(cf.Berg&Wiebes,1992;Rasplus,1996).Insome Inmonoeciousspeciesof Ficus(approx.50%of all cases,Agaonidaespeciesarenotinvolvedin pollina- speciesandall neotropicalones)the syconiacontain tion (cf. Berg & Wiebes, 1992). The females are staminate flowersanddenselypackedpistillateflowandeyes,weak erswithunequallylongstyles.Inorderto getthestigwinged,havewell-developed antennae bitingmouthparts,andoftenpockets(mostlyin the masatthesamelevel,theflowerswithshortstylesare thorax)to storeandtransportpollen.Themalesare oftenpedicellate andmayhavelongerovaries,whereas wingless,havepoorlydevelopedantennaeandeyes, thosewithlongstylesaremostlysessileandmayhave buthavestrongbitingmouthparts. shorterovaries.Inmonoecioustaxaof FicusthedifferThefemalewaspscartypollenfromaninflorescence entiationin stylelengthtakesplaceduringtheontog(syconium)releasingpollento anotherwithreceptive enyof theflowers,andall ovariescanproduceseeds. through Inthisgroupof taxa,eggsarelargelylaidin theovules stigmas.Theyentertheurceolate inflorescence theapicalopeningof theinflorescence (ostiole),being of theupperlayer(s)of ovaries(thusof theshort-styled ? tightlybarredby bracts.Thesespecializedinsects ones).Theseedsareformedin thelowerlayerof ovausuallymanageto get throughthebarrierandarrive ries(thusof thelong-styledflowers). (usuallydamaged,havingatleastlosttheirwings)in In(gyno)dioecious taxa(allin theOldWorld,sevthecentralcavityof theinflorescence, wheretheystart eralof them,includingFicuscaricaLinnaeus,introlayingeggs. The eggs aredepositedin theovulesof ducedin theNeotropics) therearetwotypesof syconia someof theflowers,namelythosewhereovipositorof ondifferent flowersand plants.Onecontainsstaminate theinsectmatchesthelength,diameter andshapeofthe onlyflowerswithshortstyles.Intheothertypeof systyle.Whileovipositing,pollenis removedfromthe coniathereareonlylong-styledpistillateflowers(and pollen-containing pocketsorpassivelyreleasedfrom oftenalso"neuter" flowers).Asthelengthof stylesdoes theinsectbodyanddepositedon a ? coherentlayerof notshowvariation, inonelayer theovariesarearranged stigmas(synstigma).Aftera numberof weeks(when to getthestigmasin onelevel,atleastatanthesis.Ovithe seedsarematureandshortlybeforetheinflores- positioncanonlyberealizedintheovariesof theshortcenceis maturefor dispersal)the insectlarvaehave styled flowers. Heterostyly is genetically fixed Themaleinsectsemergefirst (Valdeyron developedto maturity. flow& Lloyd,1979)andtheshort-styled "fruits" fromtheinsect-containing bybitinga holein ersdonotproduceseeds,althoughtheovariesappear thewall(pericarp). Thentheystartopeningthewalls to be inperfectstateatanthesis.Seedproduction only of the"fruits" containingthefemaleinsectsandcopu- occursinthesyconiawithlong-styled flowers,of which latewiththemwhilestillencaged.Copulation whilethe theovariesareneverusedforbreedingof thelarvae. femaleinsectsarestillencagedis to ensurethatallinsectswill laynotonly unfertilizedeggs (fromwhich Pollination Based on Breeding malesdevelop)butalsofertilizedeggs producingfein Staminate Inflorescences if a maleindividuals,whichis particularly important if it an as usual inflorescences of manytaxa, singlefemalecolonizes inflorescence, Incondensed staminate is small.Thenewlyhatchedfemaleswithpollenpock- suchas thoseof Artocarpusheterophyllus (cf. Fwgri flowerswhich & van derPijl, 1979),insectlarvaeof (presumably) etsactivelycollectpollenfromstaminate Pollination in Ficus

14

found.Theydonot smallbeetlesandfliesareregularly of thesametaxa. occurin thepistillateinflorescences Allowinginsectlarvaeto penetrateandfeed within staminateinflorescencesandpreventinglarvaefrom thepistillateinflorescenceses tobe penetrating appears thebasisforpollinationin manymoraceoustaxa(in themajorityof theArtocarpeae, probablymostrepresentativesof theCastilleae,andin somespeciesof the similaroranalogous Dorstenieae). Moreover, traitsare foundin manyothergroupsof angiosperms, suchas palms(cf. Henderson,1986)orcycads(cf. Norstoget al., 1986). Featuresrelated to the protectionof infloresencesandflowersarediscussedabove.This modeof pollinationmatchesthehabitatin whichthe indicatedtaxacanbe found:thecanopyandtheunderstoryof evergreenlowlandandmontaneforest.

FLORANEOTROPICA

Discussion Mostof themodesof pollinationindicatedabove (exceptforthatinFicus)couldhaveallowedestablishof Moraceae ment,diversification, anddistribution early in thehistoryof angiosperms,sincethesemodesare independent of biologicalvectorsor, if dependenton them,thenby a modeof pollinationwhichcanbe regardedas themostanciententomophilous one,likely to be alreadyexploitedby gymnosperms. Thiscomplieswithpaleobotanical data,suchasthosepresented by Craneet al. (1986)withregardto thefrequentoccurrenceof smallflowersin theCretaceous andthose by Romero(1993)withregardto diversityof fossils fromtheCretaceous ascribedto Moraceae. InFicusthespecies-specific relations with symbiotic regardto pollination appearto limitspeedof migration anddistancestonewestablishment, asthemaintenance of the pollinationsystemrequiressufficientlylarge populations.

Pollination in VariousTaxa with Bisexual Inflorescences characterized Forthemajority oftheDorstenieae, by DISPERSAL neitheranemophily norpollibisexualinflorescences, nationbasedon breedingby insectlarvaecanbe reTwomajortypesof dispersalcanbe recognizedin Thestaminate Moraceae: gardedaspossiblemodesof pollination. to windpollination flowersdo not showadaptations 1.Autochoryby expulsionorejectionof endocarp andtheplantsoftenoccurin evergreenforest,oftenin bodiesfromthedehiscentdrupesas describedabove. Theconstruction theunderstory. of inflorescences isoften ThisoccursinDorstenia,andsomeAfricangeneraof suchthattheydonotprovidebreedingsitesforinsects. thetribeDorstenieae,in somegeneraof theMoreae, Inmanycasesstigmascancontactpollenof adja- suchasinBleekrodea, Fatoua,andStreblus,andinthe centflowers(as in mostspeciesof Dorstenia),creat- twoAustralasian generaof theArtocarpeae, Antiaropsis for(geitonogamous) ingtheopportunity but andSparattosyce. autogamy, Astheballistically releaseddiaspores inothercases(suchasin severalspeciesof Brosimum), do nothaveelaiosomes,furthertransportion maybe in whichthestigmascannotcontactstamens,evenau- carriedoutbyrunningwater.Manyof theautochorous togamyis notfeasible.Agamospermy mightbe a com- taxaoftenoccurnearstreams. monmodeof reproduction fortaxaof thisgroup,in2. Endozoochory basedon the presenceof edible cludingthe(temporarily dioecious)Dorsteniacayapia fleshylayersprovidedby theexocarp,thefruitingpewhichcan(spontaneously) produceseedsinpistillate rianth,and/orthereceptacle. A largeassortment of speof plantskeptin greenhouses. inflorescences cializedoropportunistic frugivorous animals,includingbirds,bats,monkeys,andsquirrels,areinvolved. IntheOldWorldsomeof thecauliflorous taxaandthe Pollination in Other Taxa taxamaybe dispersedby terrestrial geocarpous mamSoroceais a peculiargenus.Mostof itsspeciesare mals.Theroleof rodentsanddungbeetleswithregard canopytrees,understory treelets,orshrubsof evergreen to survivalof seedsdispersedbymonkeys,as thoseof aremostlymoreor less severalMoraceae forest,andtheinflorescences Brosimum (Batocarpus amazonicus, laxspikesorracemesreminiscent of thosecommonin lactescens,Clarisiaracemosa,Ficusspp.,andPseudthe(anemophilous) Moreae.Thepollenis notreleased olmedialaevis),has been studiedanddescribedby activelyandneitherdo the staminateinflorescences Andresen(1999). providebreedingsites for insects.PluricellulartriIn additionto thesetwomajormodesof dispersal chomesarecommonlyabundantly presentontheperianthof thepistillateflower.Theymightplaya rolein somelesscommononeshaveto be considered: pollinationas substitutesof pollengrainsandmight 3. Exozoochory. ThismaybethecaseforTrymatoinvolvebees.Traitsquitesimilarto thoseof Sorocea coccusamazonicus,in whichtheinfructescences can arefoundin thececropiaceousgenusPourouma(cf. adheretofurorfeathers fimnhookedhairs. by persistent Berg,1990b;Berget al., 1990). Thefruitletsof Ficus,oftenwith a ? stickysurface,

INTRODUCTION TO THE FAMILYMORACEAE

mayadhereto animalseatingthesyconiaand/orto animalsextractingandeatingfruitletsfromthe syconia (e.g.,pigeons).Inadditionto thisdyszoochorous mode of dispersal,stomatochorous dispersalmayoccur,as may inFicusof which(relatively) largeinfructescences by be carriedforconsumption elsewhere,in particular batsandsquirrels. 4. Hydrochory. Inautochorous taxa,furthertransportof thediasporesby watermayoftenoccur.Accidentaltransport of the diasporescanbe expectedfor treesoccurringalongstreams.InsomeOldWorldFicus speciesthesyconiaappearto be adaptedto transportby waterby havingspongywalls.Inthetwospecies of anOldWorldrheophyticsubdivisionof Ficus (subsect.Macrostyla Coiner),thefruitletshaveexceptionallylong andpersistentstyleswithretrorsehairs facilitating adherence to thesubstrate inflowingwater. 5. Myrmecochory. Antsmayplaya rolein further transportation of Ficusfruitletsdepositedonbranches to moresuitablesites for establishment(Kaufmann etal., 1991). havelarge Themajorityof thegeneraof Moraceae seeds requiringgerminationshortlyaftermaturity. Germination normallyhappensin shadeof theforest Thelargeseedsreducethepossibilities forlongcanopy. Inmicrospermous distancedispersal. taxa,comprising thegenusFicus,somegeneraof thetribeMoreae(such asBroussonetia andMorus),andBagassaguianensis, the seedshaveextendedlongevityandoftenrequire light for germination.However,in Dorsteniasmall seeds(atleast)of forestundergrowth speciesappearto behavelikelargemoraceous seeds.Intheothermicrospermoustaxa,chancesfor long-distancedispersal appearto bemoreample.InFicusthepossibilitiesfor distantestablishment arelimitedbythepollination system,whereasin othertaxaby dioecy.

15

speciesin Madagascar thanon thecontinent,andon thecontinentwithmorespeciesin EastAfricathanin MiliciaSimis theonlygenusof thisgroup WestAfrica. confinedto Africa.Subgroupsof Ficus centeredin SouthEastAsiashowthesamepatternof distribution, againwithonegroup(sect.Galoglychia(Gasparrini) Endlicher) confinedtoAfrica. Anotherextension ofthese groups(FicusandMoreae)is to thenorthwestern part of theNeotropics: Central America, theGreaterAntilles andthenorthern partof theAndeanregion.Ulmaceae andUrticaceaeshowthesamepattern (Celtidoideae) of distribution (Berg,1983). 2. SouthAmerica-Africa forthetribesDorstenieae andCastilleae.Theneotropical groupof generaof the Artocarpeae is also associatedwith this center.The southern transatlantic linkis mostobviousin theDorstenieae,theAfricanmembersshowinga concentrationof taxaintheCameroun-Gabon region.Dorstenia is represented withaboutequalnumbersatbothsides of theAtlantic;evenoneof thesections(Lecanium) is transatlantic. Thetransatlantic distribution is lessbalancedin the Castilleae,as only two species,in the monotypicgeneraAntiarisandMesogyneEngler,occurin theOldWorld.ThefamilyCecropiaceae shows similartransatlantic connections. of this(WestGondwana) Thedistribution groupin tropicalAmericais characterized by the presenceof moreorlessclearcentersin thelowlandsof theSouth Americancontinent,as the Amazonianregion,the Guiana(Guayana) region,andeasternBrazil.Taxaof this phytogeographicelement extend to Central America, normallythroughorfromthePacificCoastal regionof Colombia,andin decreasingnumbersaway fromtheSouthAmerican continent. Thelattersuggests a relativelyrecentmigrationintoCentralAmericavia theextantPanamalandbridge. elementhasestabHowever,thisWestGondwana lished a center in northwestern Central secondary DISTRIBUTIONAND ECOLOGY America andtheGreater (southem Mexico-Guatemala) Distribution Worldwide Antilles.InthiscenterDorsteniais well represented, Themajorityof theMoraceaearetropicalandpre- and the other South American taxa by some dominantlyelementsof rainforest.Somegenera,Fi- (sub)endemic orby formsand species(Pseudolmedia) or evenwarm subspecies(Brosimum),orjust by disjunctoccurrence cus andMaclura,extendto subtropical andthe of species(e.g., Clarisiaracemosa).Representation temperate regionsof thenorthemhemisphere paleotropicalgenusStreblusto New Zealand.Two of taxaandmorphological variationsuggesttheestabgenera,BroussonetiaandMorus,canbe regardedas lishmentof thissecondary centerpriorto theestablishto subtropical elementsof warmtemperate regionsof mentof theextantPanamalandbridge (asearlyasthe andextendintothetropics. lateCretaceous theNorthemHemisphere (?);cf. Gentry,1982). can Forthefamilytwomajorcentersof distribution be distinguished: Habitat Occupation forthetribes 1.SouthEastAsia-Australasia-Africa Incontrastto theOldWorld,thetreespeciesof the MoreaeandFiceae(Ficus).Twogroupsof theArtorolein thecomposition, withthiscenter.TheMoreae Moraceaeplayanimportant carpeaearealsoassociated andtropicalAfrica,withmore structureandconstructionof lowlandrainforestin extendto Madagascar

16 manypartsof the Neotropics,in particularin the upper Amazon Basin andpartsof the Pacific Coastalregion. In these regions they usually rankamongthe top three familieswithregardto speciesrepresentation (andabundance)if thesoils arenot too poorin nutrients(cf. Boom, 1986; Gentry,1988, 1993; Balslev & Renner, 1989). Themajorityof theneotropicalMoraceaearecomponentsof humidlowlandtropicalforest. Smallnumbers of species arecomponentsof othertypes of vegetation:

FLORANEOTROPICA

sis naga),slopesof theAndesinAntioquia,Colombia (see Perebeaguianensissubsp.castilloides),andMount Roraima(see Maquiraguianensis subsp. costaricana andSoroceapubivenasubsp.oligotricha).Onthe other

hand,severalspecieswhichcanbe regardedas montanemay(regionally) occuratlowelevations, e.g.,Sorocea trophoides,Ficus crassiusculaStandley,F cuatrecasana Dugand,andF velutina Willdenow.

IngeneralMoraceaearepoorlyrepresented onnutrient-poor (oligotrophic) substrates, suchason sandy 1. Montaneor submontaneforest: several species soils,andconsequently arerarealongblackwaterrivof Ficus, Helicostylis tovarensis, the two species of ers.Thisalsoappliesto thehemiepiphytic FicusspeMorus, Pseudolmedia gentryi, R rigida p.p., and two with cies.Partof thewoodyDorstenieae areassociated species of Trophis).The only (sub)montane species relativelypoorsoils. Theassociationwithrelatively occurringoutside the Andeanregion (with extensions nutrient-rich substrates couldbe (atleastpartly)related to CentralAmericaandthe CoastalMountainRangeof to thenecessityof continuousfloweringto maintain Venezuela)is F tepuiensis C. C. Berg & Simonis, enthepollinationsysteminFicusorto thetreearchitecdemic to the GuayanaHighlands. tureinCastilleae, inwhichthelateralbranches produc2. Relatively dry types of vegetation: Brosimum ingtheflowersandfruitsarecontinuouslyshed. gaudichaudii, some Dorstenia species, some Ficus Thegroupof terrestrial woodyspeciesassociated species (including F brittoniiBoldingh andF eliadis withlowlandrainforestis relativelysmall,100- 10 Standley,both from dry parts of the Atlantic Coastal species.Theover-representation of Moraceaein lowregions in Venezuelaand Colombia, andF petiolaris landrainforestcanbe relatedto thewidedistribution Kunth, a (semi-)desert species of northernMexico), of manyspeciesandthefrequent co-occurrence of sevMaclura tinctoria subsp. mora, Sorocea bonplandii, eralspecies,evenrelatedones (or subspecies)of the andS. sprucei.

The majorityof the rain forest species are components of terrafirme forest, but some species (such as Maquira calophylla) can also occur in periodically flooded (varzea) forest. Other species, including MaquiracoriaceaandNaucleopsisinaequalis,arecommonly found in varzea forest, but may occur on terra firme. Only the shrubspecies Sorocea duckeiappears to be confined to the varzeahabitat. Maclura tinctoriacan be found in habitatsvarying fromwet to ratherdry.It shifts fromuplandhabitatsto periodicallyinundatedforestalong the AmazonRiver, being therea consistent and andratherimportantelement in earlystages of succession of varzeaforest.It is the only ecologically wide-rangingspecies amongthe neotropicalMoraceae,or one of the few, dependingon species delimitationsin Ficus. Most of the (few!) African forest tree species of Moraceaeareimportantelementsof transitionalor secondarytypes of forest. Bagassa guianensis and some species of Ficus subg. Pharmacosycea, are the few neotropicalforest tree species which match the ecology of the indicatedAfricanones, Bagassa guianensis showing close resemblancesto the ecology of Milicia excelsa (Welwitsch)C. C. Berg (cf. Berg, 1977c, 1982). Lowland species normally extend to elevations of 1500 m. However, in some cases lowland species may be found as high as 2000 m (or higher).Thatphenomenon appearsto be confined to some areassuch as the westernslopes of theAndes in Ecuador(see Naucleop-

samegenus,suchas Soroceaguilleminiana,S.pubivena subsp. hirtella and subsp. oligotricha, S. muriculata subsp.muriculata,andS. steinbachii,orPseudolmedia laevigata, P laevis, P macrophylla,P rigida subsp.

eggersii,asin someforestsof theupperAmazon Basin. SomeDorsteniaspecies,D. brasiliensis,D. cayapia (at least subsp. asaroides), and D. contrajerva, can

becomeweeds. Phytogeographic

Subdivisions

Basedon distributionandvariationpatternsand endemism, thefollowingregionswithphytogeographic significancefortheneotropicalMoraceaecanbe recognized.Pattemsas foundin Moraceaeareto a large extentshownby therelatedfamilyCecropiaceae, in the generaCoussapoaandPourouma(cf. particular Berget al., 1990). 1. Eastern Brazil, the region with rain forest extending from Bahia to Sao Paulo (and for some species to Santa Catarina). This region is inhabitedby at least 28 endemicmoraceousspecies or subspecies: 18

of thembelongingto Dorstenia(17 speciesandD. cayapia subsp.cayapia), at leastsix speciesof Ficus (F. F longifolia enormis(Miquel) Miquel,F hirsutaVellozo, Schott,F mariaeC. C. Berg, Emygdio & Carauta,and

F organensis (Miquel)Miquel,andanundescribed species), Brosimumglaucum, B. glaziovii, Pseudolmedia hirtula,andSorocea hilarii. The latterfive species are

moreor less closelyrelatedto (orderivedfrom)more

INTRODUCTION TO THE FAMILY MORACEAE

17

widespreadspecies.Theothermoraceousspeciesre- Magdalenavalley,anda few species extendfurtherinto cordedfromthisareaarewidespreadspecies,atleast CentralAmerica.Thisregionis inhabitedby31 endemic guian- moraceousspecies or subspecies:nine belongingto Fialsopresentin theAmazonBasin:Brosimum ensis, B. rubescens, Clarisia racemosa, C. ilicifolia, Helicostylis pedunculata (in Bahia), H. tomentosa, Naucleopsisoblongifolia,Pseudolmediamacrophylla, P laevigata, Soroceaguilleminiana, andnine more or

cus, five to Naucleopsis (N. capirensis, N. chiguila, N. naga, N. straminea, and N. ulei subsp.puberula), five to Sorocea (S. affinis, S. jaramilloi, S. pubivena subsp.pubivena, S. ruminata,andS. sarcocarpa), and Ficusspecies.Someof the(few)campo fourto Brosimum(B. costaricanumandB. utilesubspp. lesswidespread cerrado-savannaspecies of Moraceae,Brosimum allenii, darienense, magdalenense, and occidentale). gaudichaudii, Dorstenia brasiliensis, andD. cayapia Theothersare:Castillaelasticasubspp.costaricanaand othersthansubsp.cayapia,arelikelytohave gummifera,C. tunu, Dorstenia colombiana, D. pansubspecies fromtheeasternBraziliancenter.Thelinksof amensis, Perebea guianensis subsp. castilloides, and radiated thisregionwithAmazoniaarepartlywiththe lower Pseudolmediamanabiensis.Themoraceousfloraof this AmazonBasin (and throughthis region with the region is complemented with about 30 non-endemic Guianas),partlywiththeupperAmazonBasin. speciesor subspecies(not includingthose which canbe as (sub)montaneand sometimesoccur at low regarded 2. Southern Mexico-northwestern Central Thisgroupincludesabout15 speciesof Ficus. elevations. Antilles.Someof the America(Guatemala)-Greater this phytogeographicregiontwo subregionscan For elementsof thisregionextendto CostaRica,othersto be distinguished: position westernPanama.Thiscenter,in geographic 3a.A southernone (in Ecuadorandthe adjacentpart to thepreviousone, showsa remarkable "antipodal" with some endemicspecies (Pseudolmeof Colombia) to it.Itis inhabited intaxonomic composition similarity Sorocea jaramilloi, and S. sarcodia manabiensis, moraceousspeciesor by 41 endemic(orsubendemic) subspecies(Castilla elastica distinct and some carpa) subspecies.Nineteenof theseendemictaxabelongto as in Naucleopsis capiror forms, gummifera) subsp. ones,Ficus Dorsteniaandten(andthreesubendemic aureaNuttall,F colubrinaeStandley,andF turrialbana ensis and N. naga. 3b. A northwesternone (in Panamaand adjacent W.Burger)to Ficus.Twoof theFicusspeciesareenof Colombia and Costa Rica) with as (sub)endparts Antilles,F trigonataLinnaeus demicto the(Greater) Brosimumcostaricanum,B. utile subspp.allenii emics, Theothertaxaassoci(s.str.)andF berteroiWarburg. Castilla elastica subsp. costaricana, and darienense, atedwiththis areaare:Brosimumalicastrumsubsp. alicastrum,Castilla elastica subsp.elastica, Pseudol- Dorsteniapanamensis,Soroceaaffinis,andS. ruminata. media glabrata, P spuria, and Trophis cuspidata. Morus celtidifolia and Trophismexicana can be re-

4. The Hylea (in a broad sense). The whole region is inhabitedby about 150 species or subspeciesof thisregionis inhab- Moraceae,of which 49 belong to the tribe Castilleae, Moreover, gardedas subendemic. eitherinthe 24 (includingfourDorsteniaspecies)to theDorstenieae, itedby25 specieswitha widedistribution, lowlandsof SouthAmericaorin theAndes.Seventeen 18 to theArtocarpeae,fourto the Moreae,andabout60 ofthemarespeciesof Ficus(e.g.,F insipidaWilldenow belong to Ficus, most of them included in a treatment andF paraensis(Miquel)Miquel),theothersare:Bro- of the genus for BrazilianAmazonia and the Guianas simumguianense*,B. lactescens*,Clarisiabiflora, C. by Berg et al. (1984). racemosa*, Dorstenia contrajerva, Morus insignis, Two main subdivisions of the Hylea can be recogPoulsenia armata*,Sorocea trophoides*(?),and Tro- nizedwithregardto the distributionof Moraceae.Many phis racemosa*. In several cases markedwith an as- species aremore or less distinctly associatedwith one featuresand/orabundance of the two areas,being confinedto one of themor more terisk(*), morphological suggestthatthepresenceof thesespeciesinthisregion or less marginallyenteringthe other.Few species are Thetwo pe- (almost) confined to the Hylea region and manifestly couldbe relatedto ancientdisjunctions. Cen- presentin both of them, notjust marginallyextending centers,easternBrazilandnorthwestern ripheral Brosimum into one of these regions: Brosimumparinarioides, havea fewspeciesincommon: tralAmerica, guianense,B. lactescens,Clarisiaracemosa,Ficus am- Ficus albert-smithii Standley,F caballina Standley, ericanaAublet(s.l.,includingF guianensisDesvaux), F trigona Linnaeusf., Pseudolmedia laevis, Sorocea F citrifolia Miller(s.l.), F pertusa Linnaeusf. (s.l.), muriculata,and Trymatococcusamazonicus.Bagassa andF crocata(Miquel)Miquel(s.l.). guianensis can be addedto this category. 4a.TheGuiana(Guayana)regionin a widesense,with America of South (Ec3. Pacific Coastal region uador and Colombia) and the adjacent part of as southernborderroughly the watershedof the AmaCentral America (Panama and Costa Rica). Ele- zon Basin (butincludingthe northernpartof Roraima) mentsof this region may extendto the lower Rio andwestwardsthe Rio Orinoco,can be subdividedin:

18 4aa.An easternpart,comprisingthe StateofAmapa (Brazil), French Guiana, and eastern Suriname,and extending into easternParaor to Maranhao(Brazil). 4ab.A westernpart,comprising Guyana,northern Roraima(Brazil), and easternVenezuela. These parts are separatedby a transitional zone which could be an extension of the south-north-running zone in the Amazon Basin, partly including the "matade cipo" belt in Para(cf. Daly & Prance, 1989; Pires, 1974). The presenceof an easternanda western formof Naucleopsisguianensis accentuatesthe subdivision of the region. For the whole Guianaregion about 20 species or subspeciescan be regardedas endemicor subendemic; five ofthem (e.g.,Ficus leiophyllaC. C. Berg,F maroniensis Benoist, andPerebea rubra subsp. rubra)being more or less clearly associatedwith the easternpartof the region andfive (e.g., Ficus roraimensisC. C. Berg, F tepuiensis,andSoroceamuriculatasubsp.uaupensis) with the western part. Some of the species distinctly associatedwith the Guianaregionoccurdisjunctlyelsewhere:Bagassa guianensis in northernMato Grosso and Rond6nia (Brazil), Ficus broadwayi Urban and Helicostylispedunculatain Bahia (Brazil). About 50 more widespread species or subspecies are found in the Guianaregion. About 10 of them belong to a groupof whichthe distributionis largelynorth of the Amazon River, including some Ficus species (e.g., F malacocarpa Standley and F panurensis Standley),Brosimumpotabile, B. utile,Helianthostylis sprucei, and Trymatococcusamazonicus.This list indicates that the woody representatives of the Dorstenieae,in particularBrosimumsubg.Ferolia,Helianarelargelyassociatedwith thostylisand Trymatococcus, the Guianaregion. 4b. TheAmazonBasin (Amazonia).Approximately 90 non-Ficus moraceous species and subspecies are found here, about 50 of them are endemic to this (sub)region; 40 also occur outside this region, about 20 of them extending only to the Guianaregion. There is a distinct concentration of endemic species and subspecies in the upper part of the Amazon Basin, in particularof membersof the Castilleae and Artocarpeae.Fromwest to east the numberof species decreases and the moraceous flora reaches a borderline largely coinciding with the Rio Tapajosand the easternborderof the south-north-runningzone with types of transitionalforest,partlyincluding the "mata de cipo" forest belt (cf. Daly & Prance, 1989; Pires, 1974), in which Moraceae are poorly represented. West of this zone the moraceous flora consists of a mixtureof some trulyAmazonian,Guianan,andwidespread elements. The related family Cecropiaceae roughly shows the same pattern.

FLORANEOTROPICA Thenumberof Ficus speciesconfinedto theAmazon Basin is remarkablylow, and most of these endemic species (F aripuanensisC.C.Berg & F. Kooy,F coerulescens (Rusby) Rossberg (= F killipii Standley), F lauretanaVdzquezAvila, andF schultesii) areassociated with the upperAmazon Basin. The only typically lower AmazonianFicus species is F christianii Carauta(= F gameleira Standley), occurringin Para and Maranhao.About 20 of the Ficus species also occuroutsidetheHylea.Theothersoccurbothin theAmazon Basin and the Guianaregion. Two areasin the upperAmazon Basin are particularly rich in moraceoustaxa: 4ba. The southwestem comer, consisting of Acre (Brazil), Pando (Bolivia), and Madrede Dios (Peru): anarealargelycoincidingwiththeEastPeruPleistocene forest refuge proposed by Prance (1974, 1977). Not only aremanyspecies (andsubspecies)representedbut the family also tendsto be more abundantin the forests thanothertreefamilies (cf. Boom, 1989). It is remarkablethatthe numberof subspeciesor species (sub)endemicto thisareais verysmall,possiblyonlyNaucleopsis jamariensis. 4bb. The northwestemcomer, largelybetweenRio CaquetA (Colombia)andRio Marafion-RioAmazonasRio Solim6es(Peru-Brazil)andlargelycoincidingwith the Napo refuge (Prance, 1974, 1977). For the Ecuadorianpartof this area 53 non-Ficus species and subspecies have been recordedfor a floristic treatmentof this group (Berg, 1998a). For the whole (Napo) area thatnumberis about60, which impliesthatnearly50% of all non-Ficus woody species and subspecies recognized (129) are representedin that area. Moraceaeis often the most speciose family in uplandforestsin this region (cf. Balslev & Renner, 1989), and the species are commonly abundantas well. Ten out of these 60 species can be regardedas (sub)endemic,half of them herbaceousor (suf)frutescent.For this region nearly 30 Ficus species have been recorded,only F lauretana and a still undescribedspecies being endemic. The Napo region is distinct from otherAmazonian regions with concentrationsof moraceous species by the presenceof relatively many (10) endemics or subendemics,but also by the relatively strongrepresentation of species of shrubsand small trees, a categoryof moraceous taxa of which one gets the impression of being in generalmorevulnerableto extinctionthanthe species of medium-sizedand large trees. A third region showing a high concentration of species is the Manausarea.About 50 species and subspecies of Moraceae have been recorded. The area shows a remarkablemixture of elements associated with the Guianaregion and with Amazonia. None of the taxa is endemic.

19

TO THE FAMILYMORACEAE INTRODUCTION

FicusmolliculaPittieris found.Hylean of speciesintheAmazonBasin hemiepilithic Theconcentrations and/orjunc- elementsareextendingintogalleryforestin thisarea. forestrefuigia canberelatedto Pleistocene 6c. Twomoraceousspeciesarefoundin dryareas tionsof migrationroutes,butnotlikelyby speciation in the peripheryof the neotropicaldomain:Ficus events. petiolaris in northernMexico and Morus celtidifolia

5. Andeanregion,with extensionsto mountain- rangingfromArizonaandNew Mexicoto Honduras ouspartsof CentralAmerica,theCoastalMountain (andalmostcertainly regions introduced indrymontane Range of Venezuela,and for some species(Ficus in SouthAmerica). cuatrecasanaand Morusinsignis) to Argentina. Taxa

distinctlyassociatedwiththis regionare:Dorstenia

7. LesserAntilles.Noneof themoraceousspecies floraconareendemicto thisregionandthemoraceous sists of moreor less widespreadspecies (cf. Berg, Antilles(Dor1980),someextendingfromtheGreater

aristeguietae,Helicostylis tovarensis,Morusinsignis, Pseudolmediagentryi,P rigida subsp.araguensisand subsp. rigida, Sorocea trophoides,and severalspecies stenia contrajerva,Ficus americana,F citrifolia, and of Ficus, such as F carchiana C. C. Berg, F crassiusMaclura tinctoria),andothersfromnorthernSouth cula,F cuatrecasana,F gigantosyceDugand,F mutisii America (E guianensis, F insipida Willdeman,and Dugand, F rieberiana C. C. Berg, F tequendamae F nymphaeifolia). Dugand,F tovarensisPittier,andF velutina.Thereis

a distinctconcentration of thesespeciesinthenorthern 8. Trinidadand Tobago.The smallindigenous partof theAndes(EcuadorandColombia). moraceousfloraof theseislands,comprising13 speVenezuela cies,areelementsextendingfromnorthern 6. Neotropicalregionswithmoreor lessdryveg- ortheGuianaregion(cf. Berg&vanHeusden,1982). etation,includingcampocerrado,deciduousforest,chacoforestandscrub,thornscrub,orsavanna. Ranges of Distribution 6a. The arearichestin moraceousspeciesis the toDorstenia, species,allbelonging Mostherbaceous extensiveareawith campocerrado,chacoforestor A scrub,anddeciduousforestin CentralBrazil,eastern havesmallorevenverysmallrangesof distribution. -D. few them can behave as weeds) brasilof (which andnorthern ThefollowArgentina. Bolivia,Paraguay, withthisarea:Brosimum gaudi- iensis, D. cayapya, and D. contrajerva - have large ingtaxaareassociated or speciesof (sub)shrubs chaudii, Dorstenia brasiliensis, D. cayapia (p.p.), ranges.Non-anemophilous D. tenuis,Ficus adhatodifoliaSchott,F calyptroceras (Miquel)Miquel,F carautanaNeves & Emygdio, F obtusiuscula(Miquel) F maromaCastellano, Miquel, Macluratinctoria(subsp.mora),Sorocea bonplandii, andS. sprucei (subsp.saxicola). Soroceaspruceilinks

small trees (such as Naucleopsis humilis, Perebea

small tendtohaverelatively humilis,andP mennegae), Smallrangesareuncommon in rangesof distribution. treespeciesaswellasinthehemiepiphytic theterrestrial ones(of Ficus).Treespeciesof humidto wetlowland in spiteof (presumed) thesouthern (Bo- forestwithlimiteddistribution areawithmoreorlessdryvegetation are Brosimum costaricanum, expansion possibilities withthenorthern oneinVenezuela and livia-Argentina) Ficusroraimensis,F lauretana,HelianthoB. glaucum, Colombiaby occurrencein a dryareain Amazonian N. Peru(SanMartin),the dryforestregionin southern stylis steyermarkii,Naucleopsis chiguila, humilis, Soroceajaramilloi, S. ruminata,and S. sarcocarpa. PacificEcuadorandtheadjacentpartof Peru,andthe upperCaucavalley(Colombia).Inthedrypartof San

Martin(Peru),Sorocea sprucei (subsp. sprucei) is accompaniedby Dorstenia brasiliensis (andD. contrajerva). In westernEcuadorand Peru,Sorocea sprucei

Disjunctions

of speciesandsubInmostcasesthedistribution distrispeciesis ? continuous.Partof discontinuous and butionis dueto physicalbarriers(asmountainranges is represented bytheendemicsubsp.subumbellata is accompanied asformonbytheendemicFjacobiiV6zquezAvilaorsea)ordueto ecologicalrequirements, (andDorstenia contrajerva). tanespeciesandthoseassociatedwithrelativelydry Atlantic typesof vegetation(Brosimumgaudichaudii,Dorstenia 6b.Inthedryareaalongthenorthwestern Coast(Venezuelaand Colombia)Soroceasprucei brasiliensis,D. cayapia p.p., andSorocea sprucei). A considerablenumberof speciesoccuron both (subsp.sprucei) is accompaniedby Ficus brittoniiand F eliadis.Inthesavanna(llanos)area,extendingfrom sidesof theAndes(see list below).Thepresentdislinkof taxasuggeststwopossible"corridors" ColombiathroughVenezuelato Guyana,the same tribution andPacificlowlands:onethrough by Dorsteniabrasilien- ingtheAmazonian (sub)speciesis accompanied part linkingtheNapovalleywiththesouthern sis. Onthegranitichills(lajas)in thisareatheendemic Ecuador

20 of the Pacific Coastal region (e.g., for Maquira guianensis subsp. costaricana and Pseudolmedia rigida subsp.eggersii) anda northernone throughthe Magdalena valley (such as for Ficus casapiensis (Miquel) Miquel andNaucleopsis glabra). Disjunct occurrences that cannot readily be explainedby absenceof suitablehabitatsarefoundin several Ficus species, such as Ficus dendrocida Kunth, foundin northwesternSouthAmericaandPanamaand near Santarem (Para, Brazil), and F membranacea C. Wright,occurringin Mexico andCuba,northeastern 'Colombia,and in Ecuador.The Caribbeanform of F americanais also foundin VenezuelanGuayanaathigh elevations,andF ypsilophlebia Dugand, occurringin the MagdalenaValley, the Napo Valley, northernSan Martin(Peru), and in an arearanging from Madrede Dios (Peru)to SantaCruz(Bolivia).The latterpatternis to some extentreminiscentof thatof Maclurabrasiliensis, found in Honduras,in northernVenezuela, in an arearangingfrom Madrede Dios (Peru)to SantaCruz (Bolivia), andin easternBrazil. Trophismexicana,extendingfromMexico to Panama,is also foundin southernmostEcuador.Less pronounced(or clear)disjunctions are also found, e.g. in Bagassa guianensis, a species of the Guianas and the lower Amazon Basin also foundin northernMatoGrossoandRondonia,and in Clarisia racemosa, extendinginto CentralAmerica up to Nicaraguaand found in Oaxaca(Mexico). This in SouthAmerica. speciesalsoshowsapatchydistribution In regionswherethis species is found, it usuallyoccurs abundantly.Brosimumalicastrum subsp. bolivarense resembles C. racemosa in its patternof distribution. Lowland species occurring on both sides of the Andesare:Batocarpuscostaricensis,Brosimumalicastrum,B. guianense,B. lactescens,B. rubescens,B. utile, Clarisia biflora, C. racemosa, Dorstenia contrajerva, Ficus casapiensis, F crocata, F donnell-smithii Standley,F guianensisDesvaux,F insipidaWilldenow, F macbrideiStandley,F maximaMiller,F nymphaeifolia, F obtusifolia Kunth, F paraensis (Miquel) Miquel,F pertusa, F schippii,F yoponensis Desvaux, Helianthostylis sprucei, Helicostylis tomentosa, Maclura brasiliensis, M tinctoria, Maquira guianensis, Naucleopsis glabra, N. ulei, Perebea angustifolia, P guianensis,P xanthochyma,Poulseniaarmata, Pseudolmedia laevigata, P rigida (subsp. eggersii), Soroceapubivena, Trophiscaucana, T racemosa. Species found in the Hylea and easternBrazil are: Brosimumguianense,B. lactescens,B. rubescens,Clarisia ilicifolia, C. racemosa, Ficus broadwayi,F casteliviana Dugand, F crocata, F guianensis, F nymphaeifolia, F pertusa, F pulchella Schott (s.l. incl. F piresiana Vazquez Avila & C. C. Berg), F trigona,

FLORANEOTROPICA Helicostylis tomentosa, H. pedunculata, Maclura brasiliensis, M. tinctoria, Naucleopsis oblongifolia, Soroceaguilleminiana.

AND DIVERSIFICATION CLASSIFICATION Thefamilyasithasbeenredefined byCorner (1962) comprises6 tribes:Moreae,Artocarpeae, Dorstenieae, Castilleae,andFiceae.Thesubdivisionof thefamily hasbeenproposedby Corner(1962) andremodelled by Berg(1973, 1988, 1989b).Eachof thesetribesis describedandbrieflydiscussedin thepresentcontribution.Thetribescanbe regardedas naturalentities andmostof themhaveclearcoherenceinmorphologianddiscalcharacters andinpatterns of differentiation tribution.OnlythetribeArtocarpeae lacksthatclear coherence, andthreeratherdistinct(butnotyetformalized)subdivisions canberecognized. The familycomprises37 genera(Berg, 1998b). Mostof themaresmall,withup to 14 species.Two genera(NaucleopsisandStreblus)have20-25 species, Artocarpushas50-55 species,Dorstenia 105 species, andFicus 700-750 species.Nineteengeneraoccurin theNeotropics;14of theseareendemic,andtheothers alsooccurin theothermajorphytogeographic realms. Thepresenttaxonomicclassificationof theMoraceaeappears in evoto revealmoreclearlythepatterns lution,diversification, anddistribution thanprevious systems.However,thegenusFicus is toodiversetobe treatedasequivalent to theothergenera;onlyitsmajor subdivisionsarecomparable withtheothergenera. Thedelimitationof supraspecific taxa(generato families)aspresentlyrecognizedarebasedon a maximumof morphological coherence.Singlecharacters playeda limitedrole,butsometimestheyareimportant,suchasthedifferencein thepositionof theovule (apicalvs. basal,in facttheonlyexclusive"technical" to discriminate character) betweenMoraceae andUrticaceae,orthearchitectural modelof "Cook"to delimitatetheCastilleae.Over-evaluation of singlecharacters,oftenusedin identification keys,hascreatedmany problemsin delimitation andin classification. Forexample,thetribeMoreaeis characterized by thepresenceof stamensinflexedin thebudandbendingoutwardselastically.As someof the speciesof Maclura (theonly moraceousgenuswiththorns)do not have suchstamens,theywerekeptin anothergenus,evenin anothertribe.Dorstenia hasinflexedstamensandhas forthatreasonbeenplacedamongtaxawithinflexed butthestamensof Dorstenia aremorphologistamens, cally(andfunctionally) quitedistinctfromtheurticaceoustypeof stamen,characteristic fortheMoreae. Theaccentonmorphological coherence impliesthat also yes-or-nocharacterstateshaveplayeda limited

TO THE FAMILYMORACEAE INTRODUCTION

21

Discrepanciesbetweenresultsof traditionaland is to someextent role.Italsoimpliesthattheapproach ob- modemapproaches intuitiveandsubjective(including"impressions" may,in thecaseof the Moraceae Thisap- (andvariousotherangiosperm tainedfromfieldworkindifferentcontinents). families),notonlybe thepresent state relatedtothewayscienceis progressing. proachseemstobejustifiedconsidering Otheraspects com- to be takenintoconsideration of knowledgeof thepathwaysandpresumably aretransferof functions plexprocessesthroughwhichthepresentmorphologi- in reproductive accenton analogiesrather structures, calandecologicaldiversityandtaxonomicconstella- thanhomologies,moreopenontologicaldevelopment the common of thedominantreproductive tion developed.It also counterbalances theinfloresstructures, humaninclination to simplify. cencesratherthantheflowers,andevolutionlongbegeneramay foremodempollinationagents(witha strongimpact Keeping Maquira andPerebeaasdistinct serveasanexample.Thetechnicalmorphological char- ondifferention andspeciation)cameintoexistence. actersto separateMaquiraandPerebeaaresuchthat Severalspeciesof the samegenus,oftenclosely thesetwoentitiescouldbeeasilymergedandonlymain- relatedones,withstructurally thesameinflorescences tainedas subgenera. Featuresto tellthesetwoentities andfruitsmayoccursideby side,floweringandfruitentitiesatthegenuslevel)are ing at the sametime,withoutproducinghybrids,as, apart(asgoodandnatural incolorof driedmaterial (greenish vs.brown e.g., Pseudolmedialaevigata,P laevis,P macrophylla, differences witha lead-colored hue),theleafmargin(alwaysentire andP rigida(subsp.rigida), orSoroceaguilleminiana, andtendingto becallosevs. dentateatleastinjuvenile S. muriculata (subsp. muriculata), S. pubivena (short,inconspicuous, (subspp.hirtella andoligotricha), andS. steinbachii, stages),thetypeof indumentum orabsentvs. oftenlongandconspicuous), andtheperi- all foundin the sameone-hectareplot (Cuyabeno, dermof theleafytwigs(oftenvs. neverflakingoff). Ecuador). Thiscommontraitof (neotropical) Moraceae ThetribeFiceaeis bythefeaturesof its onlygenus incombination withinvolvement of generalized polliFicusquitedistinctandit cannotbeconfusedwithother nationagents(withtheexceptionof Ficus) andgenertribes(orgenera).However,theonlyexclusivecharlittle alistdispersalagents,widerangesof distribution, onthe actersare:presenceof waxyglands(somewhere chromosomal differentiation (withtheclearexception andanthesisof stamiplant),somekindof heterostyly, of Dorstenia) (Fedorov,1969),flexibilitywithregard nateflowerswhenthefruitsaremature,featureseviasthoserelatedto inundato ecologicalrequirements dentlyor probablyfunctionallylinkedto theunique tionandelevation,andlittleradiationoutsidetherain morphomodeof pollinationandnotstraightforward foresthabitat,do notprovideclearindicationsabout thegenusFicustends logicalones.Innumerical analyses of speciation. Littleof what modesorpathways possible to endupin a clusterof themostgeneralized generaof describedaboutspeciationin tropical Gentry (1988) thefamily. Most Inthedelimitation of speciesandsubspecies, charac- forestscanbe relatedto neotropicalMoraceae. occurred but has preallopatrically, likely speciation notonlyin tersof vegetative partsplaya dominantrole, that that distributional so patterns ago sumedly long andandrodioecious taxa,but dioecious,gynodioecious, to it no have related are longer apparent. could been alsoinmonoecious partlydue ones,inthelattercategory Incontrast to theothergroupsof Moraceae, speciatouniformity of floralcharacters (suchasinDorstenia) berelatedtopollination, becauseof its tion in can Ficus inparticular thoseof orthevariation of suchcharacters, relations. flowers(suchas inBrosimum). Genericde- commonspecies-specificplant-pollinator staminate the the of attractants in chemical composition Changes limitationalso includesto someextentcompromises rolein speciation. betweenentitiesthe specialistcan perceiveas mor- mayhaveplayedanimportant phologicallyand/orbiologicallydistinctandthosethe distinct. canrecognizeasmorphologically non-specialist USE Modemnumerical,cladistic,orchemical(includof Moraceae is limThedirecteconomicimportance methodshavehardlyplayeda ingmolecular) analytical rolein thepresentclassificationandtaxonomicdeci- ited.Sometreespecies,suchas Bagassa guianensis The havebeeninvolvedin some andBrosimumrubescens,providevaluabletimber. sions,althoughMoraceae whichsomespecieshad(Castilla of suchanalyses,unpublishedor somewith limited economicimportance &Black- elastica andC. ulei as sourcesof rubberandMaclura implications (Herreet al., 1996;Humphries (onthe tinctoria as a sourceof a yellow dye) has vanished. more,1989).As canbe expectedandpredicted revueof scientific basisof aretrospective developments) Theseedsof Brosimumalicastrum subsp.alicastrum solvesomeprob- havebeenharvestedas a starchfood source.Latexof suchanalysesarepartlyconfirmative, such formedicinal purposes, theotherhand, variousspeciesis extracted but on effects, lems,haveilluminating (andalliedspecies) andcreatenewproblems. asthatof Ficus insipidaWilldenow theyarepartlycontradictory,

22

FLORANEOTROPICA

andthatofMacworminfections, usedagainstintestinal luratinctoriato extractinflictedteeth.Latexof some poison.Bark Naucleopsisspeciesis usedasarrow/dart barkcloth. of Poulseniaarmatais usedforpreparing thoseof PseudFruitsof severalspecies,inparticular Morus olmedia,areeaten,butdonotappearinmarkets. and in SouthAmerica hasbeenincultivation celtidifolia soldinmarkets.SeveralOldWorldspecies the"fruits"

CONSERVATION STATUS Thewoodyandwidespreadspeciesof Moraceae by extinction.Some cannotbe regardedas threatened apparently rarewoodyspecieswithlimiteddistribution (such as Brosimum costaricanum, B. glaucum, Naucleopsis jamariensis, and Pseudolmedia mana-

biensis)canbe regardedas vulnerable.Manyof the herbaceousspecies,all belongingto Dorstenia,have Artocarpusaltilisand limitedrangesof distribution, of Moraceaehavebeenintroduced: andseveralof themare A. heterophyllusas food sources,Morusalba Linnaeus onlyknownfroma singlelocalityor a few neighboras a fruittreeanda sourceof silkwormfood (in Bo- ing localities.Theycanin manycasesbe regardedas livia,Dr. M. Nee, TheNewYorkBotanicalGarden, beingclose to thebrinkof extinction.A few (e.g.,D. pers.comm),andseveralFicusspeciesasomamentals. brevipetiolata)may alreadybe extinct.

TAXONOMIC TREATMENT orirreguradiallysymmetrical adnatetothereceptacle),

Moraceae

tepals(1-)2-4(-7) andfreeor Trees,shrubs, woody climbers,or herbs, terrestrial lar.Staminateflowers: dioeciousor monoecious,withlaor hemiepiphytic, oropandin spiralsordistichous, tex.Leavesalternate enlaminabasallyattached (orpeltate), posite,petiolate; tire or incised;venationpinnateor subpalmateto stipulesfully mostlybrochidodromous; subtrinervate, to lateral,freeorconnate.Inflorescences amplexicaul usuallyin axillarypairs,unisexualor bisexual,racecapitatewitha discoid mose,spicate,globose-capitate, receptacle withorwithoutaninvolucre, to cup-shaped or withanurceolatereceptacle,multi-to uniflorous, bracteate. Flowersunisexual,freeorconnate(oralso

connate,or perianthlacking;stamens1-4(-6), antipistillode tepalous,straight orinflexedbeforeanthesis; presentorabsent.Pistillateflowers:tepals(3-)4(-8), freeor connate;pistil 1;ovaryunilocular,freeor adnateto the perianth;stigmas2 or 1; ovule 1, (sub-) free Fruitanacheneordrupaceous, apicallyattached. oradnateto theperianth,oftenforminga drupaceous wholewiththefruiting perianth oralsowiththe(fleshy) receptacle;endocarpmostlycrustaceous,sometimes woodyorleathery;seedlargeandwithoutendosperm orsmallandwithendosperm; embryovarious.

Key to the tribes of the Moraceaein the Neotropics 1. Inflorescencesurceolate,enclosingthe flowersentirely,also duringanthesis;laminawith waxy glandularspots at the base of the midribor in the axils of the basal lateralveins beneath.......... .......V. Ficeae 1. Inflorescencesnot urceolatenor enclosingthe flowersentirelyduringanthesis;laminawithout waxy glandularspots. 2. Treesor shrubswith self-pruninghorizontalbranches;leaves on the verticalbranchesin spirals, those of the horizontal(and flowering)ones distichous;inflorescenceswith an involucreof imbricatebracts;uncinatehairs absent............................................................... IV. Castilleae 2. Trees, shrubs,or herbs withoutself-pruningbranches;leaves distichousor opposite,or if in spirals,then the plants herbaceous;inflorescencesnot involucrate(except in Trophiscaucana); uncinatehairs often present. 3. Inflorescencesbisexual,or if unisexual,then with peltatebractson the (sub)globosereceptacle, or the plantsherbaceous;pistillateflowersimmersedin the receptacle.......... .....III. Dorstenieae 3. Inflorescencesunisexual,the receptacleof the staminateflowerselongate,or if globose, then withoutpeltatebracts,the pistillateflowersnot immersedin the receptacle,and the plants woody. 4. Stamensinflexed in the bud, bendingoutwardssuddenlyand elastically;pistillode present;the plantswith thorns,the terminalbuds of branchescaducous,the perianth not enlargedand fleshy in fruit,or the pistillateinflorescencesracemoseand the fruitI. Moreae ing perianthorangeto red......................................................... 4. Stamensstraightin the bud, or if inflexed,then bendinggradually;pistillodeusually absent;the plantswithoutthorns,the terminalbuds of the branchespersistent,the fruiting perianthenlargedand fleshy, and if the pistillateinflorescencesracemose,then the ......................................................................II. A rtocarpeae fruitingperianth(finally) black(-ish)

Key to the neotropicalgenera of the Moraceae 1. Plants herbaceous ..................................................................... 1. Plants woody.

9. Dorstenia

TAXONOMICTREATMENT

23

2. Leaves opposite..................................................................... 4. Bagassa 2. Leaves alternate. 3. Stipules fully amplexicaul,leaving annularscars. 4. Stipulesand often also otherpartsaculeate,with prickleswith a hyaline apex.......... 7. Poulsenia 4. Stipulesand otherpartsnot aculeate. 5. Stipules connate. 6. Inflorescencesglobose to turbinateand coveredwith peltatebracts............ 10. Brosimum 6. Inflorescencesdiscoid, flabellate,cup-shaped,or if globose, then withoutpeltate bracts........................................................... 17. Castilla 5. Stipulesfree. 7. Inflorescences urceolate;1 medianor 2 lateralwaxy glandularspots at the base of the midribbeneath......................................................1 9. Ficus 7. Inflorescences discoid,globose,ellipsoidor spicate;waxy glandularspots lacking. 8. Inflorescencesnot involucrate,the staminateones spicate-clavate,the pistillate ones globose or ellipsoid...................................... (p. 143) Artocarpus 8. Inflorescencesinvolucrate,discoid or uniflorous. 9. Staminateflowersenclosedby involucralbractsuntil anthesis;pistillate inflorescencesuniflorousor with spine-likeor hardenedcushion-shaped perianthparts. 10. Staminateinflorescenceswith distinctflowers;pistillateinflorescences with "free"spine-like or hardenedcushion-shapedperianthparts ........................................................................................................ 18. Naucleopsis

10. Staminateinflorescenceswithoutdistinctflowers,but only with "free" stamensand interstaminal scales; pistillateinflorescencesuniflorous ..................................................................................................... 13. Pseudolmedia

9. Staminateflowersnot enclosedby involucralbractsuntil anthesis;pistillate inflorescences(usually)with more than 1 flower and perianthparts not spine-likeor cushion-shaped 14. Perebea .............................................. 3. Stipulesnot (but sometimesnearly!)fully amplexicaul. 11. Stipulesconnate;plantswith spines ............................................................. 3. Maclura 11. Stipulesfree; plantswithoutspines. 12. Inflorescences involucrate. 13. Stamensinflexed,bendingoutwardselasticallyat anthesis;pistillateinflorescences with one flowerwith connatetepals.............................. 2. Trophis (caucacana) 13. Stamensstraight;pistillateinflorescenceswith morethanone flower,or if only one flower,then with free tepals. 14. Lamina(and otherparts)? distinctlyhairy;leaf marginoften dent(icul)ate ........................................................................................................................ 16. Helicostylis

14. Lamina(and otherparts)(sub)glabrous;leaf marginentire......................15. Maquira 12. Inflorescencesnot involucrate. 15. Inflorescences bisexual(withthe receptaclecontaininga single embeddedpistillate flower)or staminate,the pistillodesof the staminateflowerswith long filiform stigmas................................................. 2. Helianthostylis 15. Inflorescencesbisexualor unisexual,the staminateflowers withoutstigmas. 16. Inflorescences bisexualwith a cylindricalto turbinateto subglobosereceptacle (containinga single embeddedpistillateflower)with the staminateflowers contractedat the apex of the receptacle.............................................. 11. Trymatococcus 16. Inflorescencesbisexualor unisexual,with the staminateflowers? disperse. 17. Inflorescencesglobose-to turbinate-capitate, coveredwith peltatebracts; pistillate flowers(often only 1) embeddedin the receptacle..........1..10. Brosimum 17. Inflorescencesracemose,spicate,or if globose, then withoutpeltate bractsand pistillateflowerssuperficial. 18. Leaf marginspinulose-dentate or only the leaf acumenspinulose. 19. Uncinatehairs absent;inflorescencesracemose..........................7. Sorocea 19. Uncinatehairs present;staminateinfloresecencesspicate,the 6. Clarisia pistillateones capitate..................................... 18. Leaf marginentire,or if dentate,then not spinulosenor the the leaf acumen spinulose. 20. Staminateinflorescences(slender)spikes with the flowers close togetherand the stamensstraight;pistillateinflorescencesglobosecapitateor uniflorous. 21. Leaf marginentire;inflorescencesoften on leafless branchlets (on the olderwood);pistillateinflorescences uniflorous.... 6. Clarisia

FLORANEOTROPICA

24

21. Leaf margindentate;inflorescencesaxillary;pistillateinflorescences globose-capitatewith more than 1 flower........5. Batocarpus 20. Staminateinflorescencesracemes,or if spikes, then the flowers distinct(? spaced);pistillateinflorescencesracemoseor spicate. 22. Stamensstraightor slightly inflexed, not bendingoutwards elasticallyat anthesis;perianthof pistillateflowerswith a ? distinctupperand lower part;fruitingperianthblackishat maturity,the rachisand pedicels often red...........................7. Sorocea 22. Stamensinflexed,bendingoutwardselasticallyat anthesis; perianthof pistillateflowerwithouta distinctupperand lower part;fruitingperianthmostly orangeor red at maturity. 23. Staminateflowerswith imbricatetepals;pistillateflowers with (almost)free tepals;shoot apices shed.......................1. Morus 23. Staminateflowerswith valvatetepals;pistillateflowers with connatetepals;shoot apices persistent ......................2. Trophis

I. MOREAE Trees, shrubs, climbers, (or, in Fatoua, herbs),

usually dioecious,withoutor with uncinatehairs. Leavesalternate, distichousorin spirals;laminaentire marginenorincised;venationpinnateto subpalmate, tire, dentate,crenateor serrate;stipuleslateralto Inflorescencesracemose,spicate, semiamplexicaul. or(pistillate ordiscoid-capitate globose-(sub)capitate bractspeltate bracteate (orebracteate); ones)uniflorous, orbasallyattached.Staminate flowers:tepals3, 4, or 5; stamens3, 4, or 5, (mostly)inflexedin thebudand atanthesisbendingoutwards suddenlyandelastically; present. small;pistillode anthers introrse, theconnective Pistillateflowers:tepals4, freeorconnate;ovaryfree or less commonlyadnateto the perianth;stigmas2, mostly inlength,or 1.Fruitingperianth equalorunequal enlargedandfleshy,greenishorcolored;fruita dehiscentdrupeorindehiscentandfreeoradnateto theperianth;seed smallandwith endospermor largeand testawitha thickened,vascularwithoutendosperm; izedpartbelowthehilum;embryovarious. TheMoreaeis distinctfromtheothertribesof the Moraceaein thepresenceof urticaceousstamens:inflexedinthebudandbendingoutwards elasticallyand suddenlyat anthesis.Thereareonly somespeciesof Maclurainthepaleotropical sectionCudrania (Trecul) Cornerwhichdo not havesuchstamens.Therefore, fromtheotherMoreaein theyhavebeenkeptseparate (cf.Trecul,1847;Bureau,1873; previousclassifications Bentham& Hooker, 1880; Richter,1895; Engler, 1889).However,theyarelinkedto othermembersof thegenusMaclurain so manyfeatures thattheydoubtlesslybelongto thegenusandthetribe. Theurticaceousstamensareclearlyanadaptation to anemophily: thepollengrainsthrownoutintotheair of thestamensaretransbytheexplosivestraightening portedfurtherby airmovement(asdiscussedmorein detailintheintroduction to thefamily,seeabove).Pollentransport to berareoronlysupplebywindappears

mentaryin the rest of the family.Anemophily is decisive for the type of habitatoccupiedby membersof the Moreae:mostly forest margins,scrubvegetation, and deciduous forest. Some can be found in forest undergrowth,butaretherelargelyconfinedto the marginsof (small) streams.Representativesof the Moreaeareoften shrubsor climbers.Some can become large (emergent) trees, others(in the paleotropicalgenus Fatoua) areherbaceousor suffrutescent. The inflorescences of the Moreae are relatively simplecomparedwithotherMoraceae.They aremostly racemes,spikes, or ellipsoid to globose heads. The involucratehead as found in Trophiscaucana is the most advancedtype of inflorescencein the tribe.The urticaceouspollenreleasemechanismrequiressufficientspace for eachof the staminateflowersandthatlimitsthe evolutionarycondensationof the staminate(andbisexual) inflorescences.Pistillateinflorescencesareoften more condensedthanthe staminateones, but fusion of pistillate flowers is rarein this tribe. The staminateflowers of the neotropicalrepresentativesofthe Moreaeare4-merous.Theperianthis welldevelopedandthe tepalsareimbricateor valvatein the bud.A pistillode is usually present.Togetherwith the perianththe pistillode seems to play a role in keeping the long incurved stamens in position and undertension until anthesis. The pistillateflowers also have a i well-developed perianth.The ovary is mostly free from the perianth, but is partly connate in some Trophisspecies. There are two styles, or by reduction only one. The fruitsare small to medium-sized.They areenclosed in the fleshy enlargedperianth,free or adnate, forning a pseudodrupe.If not enclosed by a fleshy perianth,the fruitsaredrupaceous.In some paleotropical species the drupesareof the dehiscenttype, ejectingor expellingthe endocarpbody.In Trophisinvolucratathe perianthdoes not even enlargeduringthe fonnation of the fruit, and the exocarp tums dark blue to black at Smallseedscontainendospermanda(relatively) maturity.

25

TAXONOMIC TREATMENT

simplyconstructed embryo;thelargeroneslackendoThetribeMoreaecanbe delimitedratherclearlyby spennandoftenhavemoreelaborate stamens.Butthegeneraof typesof embryos. thepresenceof urticaceous Thetribecomprises8 generaandapproximately 73 thistribearenotclear-cut. Fortheapproximately 73 spespecies:Bleekrodea,with2 speciesinAsiaandMada- ciesthatarerecognized atpresent, about40 generic names gascar(Berg,1977c);Broussonetia, with8 speciesinAsia havebeencreatedoverthecourseof time.ThesenumandMadagascar (Coiner,1962;Berg,1977c);Fatoua, bersindicatethetaxonomicsituation inthetribe:many with2 speciesinAsia-Australasia andMadagascar(Berg,genera(withalimitedgeographic distribution) withonly 1977c);Maclura, with 11 speciesworldwide(Berg, oneorafewspecies,andspeciesthathavebeenmoved 1986);Milicia, with2 speciesintropicalAfrica(Berg, fromonegenusto another. Corner(1962)tooktheini1982);Morus,withapproximately 12speciesworldwide; tiativetodeviseamoresatisfactory classification of the Streblus,with23 speciesinAsia-Australasia andAfrica tribebyfocusingonsimilarities ratherthandifferences. (Berg,1988);andTrophis,with9 species,inAsia,Mada- Heproposedsomeconsiderably enlargedgenera(such as Maclura,Streblus,and Trophis),with wide or even gascar,andtheNeotropics(Berg,1988). MostgeneraoftheMoreaearetropical, someoftthem pantropical distribution. By adopting Corner's concepts (asMaclura) extendingto northemtemperate inforregions andtakingintoaccountmoredetailedtaxonomic (inNorthAmericaM.pomiferaC.K. Schneider). Two mationaboutAfrican andneotropical taxa,Berg(1986, of thegenera,Broussonetia andMorus, aredistinctly 1988)redefinedthetribeandremodeledsomegenera, associatedwiththetemperate to subtropical partof the recognizingtheeightgeneralistedabove. northern Theirrangesextendto themounhemisphere. Thetribehasadistinctcenterof diversity in SEAsiatainouspartsof thetropics,orevenlowlandparts(as Australasia, andextensionsto Madasgascar, continenMorusmesozygiaA. Chevalier incontinental Africaand talAfrica,andNWtropicalAmerica.IntheNeotropics Broussonetiagrevei (Baillon)C.C. Bergin Madagas- ninespeciesof theMoreaearerepresented: two specar).In thesetwo genera,associatedwithtemperate ciesof Morus,fivespeciesof Trophis, andtwospecies climatical theshootapexis shed(abscissed) of Maclura.Thedistribution conditions, of theneotropical Moreae attheendof thegrowingseason.Onecanalwaysfind appearsto be linkedwiththenorthwestern partof the a shorttwigstumpbesidetheterninalaxillarybud;the Neotropics(Central Americaandnorthern Andes)and budfromwhichthetwigelongates.Theaxillarybuds thesetaxashowtaxonomicrelationswithtaxainAsia are ? well-developedandhave bud-scales.All (or andeasterntropicalAfrica.Thispatternalsooccursin most?)membersof thesetwo generaaredeciduous. Urticaceae, Ulmaceae,andFicus(cf. Berg,1983.) Key to the neotropicalgenera of Moreae 1. Plantsarmedwith thorns;pistillateinflorescencesglobose-capitate; stigmas2 and unequalin length or 1; tepals of both staminateand pistillateflowersand bractsmostly with yellow dye accumulated in embedded"glands" ............................................................................. 3. Maclura 1. Plantsnot armedwith thorns;pistillateinflorescencesracemose,spicate, subglobose-capitate or discoid-capitate, and involucrate;stigmas2 and equalin length;tepalsof both staminateand pistillate flowers and bractswithoutyellow dye. 2. Shoot apices deciduous;tepals imbricate;fruitingperianth? enlargedand fleshy, 4-parted....... 1. Morus 2. Shoot apices persistent;tepals valvate;fruitingperianthenlargedand 4-lobed to 4-dentateor 2. Trophis fruitingperianthnot enlarged............................................................................ Morus

1. MorusLinnaeus,Sp.P1.986. 1753;Seringe,Descr. et Cult. Mur. 1855; Bureau,in De Candolle,Prodr. 17: 273. 1873; Bentham& Hooker, Gen. PI. 3(1):

364. 1880;Leroy,Rev.Bot.Appl.29: 481. 1949. Lectotype,Britton& Brown,Ill.Fl.N. U.S.ed. 2, 1: 631. 1913. Morus nigra Linnaeus. Including: Morus Linnaeus subg. Gomphomorus Leroy,Bull. Mus. Hist. Nat. (Paris),Ser. 2, 21: 732. 1949. Type. Morusinsignis Bureau. Trees,dioeciousor sometimesmonoecious,deciduous; shoot apices shed; axillarybuds with bud scales; uncinatehairslacking.Leavesaltemateanddistichous;

lamina pinnately veined; margin crenate to serrate; stipules free, lateral.Inflorescences unisexual (or bisexual), solitaryor pairedin the axils of normalleaves orin the axilsof scaleleaves atthebaseof lateral(short-) shoots,racemoseto spicateto subcapitate,pedunculate, bracteateor ebracteate;flowers+ distant,pedicellateor sessile;tepalsfree or basallyconnate,decussate-imbricate;pistillode usually quadrangular;ovary free; stigmas 2, tongue-shaped,equally long. Fruitingperianth enlarged, ? fleshy, reddish (to blackish); fruit (sub)drupaceous; endocarp crustaceous with a placental woody plug; seed small,with endosperm,the testawith a suborbicularthickened vascularizedpartbelow the hilum; embryocurvedwith equal and flat cotyledons, not enclosing the long radicle.

26

FLORA NEOTROPICA

a comparison of themontanespecies,M.insignisand M. macroura,showingclearsimilarities,hasto precedefurtherdecisions. Theassociation of thegenuswiththenorthern-temAt theend peratezoneis expressedmorphologically. ofthelongAmerica (M. insignis) and SE Asia (M. macroura of thegrowingseasontheapicalmeristems aslowlandforestspe- shootsabortandareabscised.Besidethe "terminal" Miquel),butincontinentalAfrica lateralbudsonecanfinda shortstump,whichcanstill cies (M. mesozygia A. Chevalier). axilKunth(1817)created twonamesinthegenus,Morus be tracedto theolderwood.Fromthe"terminal" lary buds in the long-shoots elongate the next growing M and material celtidifolia collected corylifolia,basedon (accompanied by inEcuador. A thirdone,M mexicana, wasdescribed by season.Growthis thusintermittent Bentham fromMexico.Quite sheddingof leaves)andit is sympodial.Theaxillary (1841),basedonmaterial inMorusceltidifolia,butare differentwasthematerialfromColombiausedforthe budsarewell-developed in small these M. insignis; budsareprovidedwithbud descriptionof M. insignisby Bureau.Synomymsto The in scales. leaf venation M. insignisdiffersfrom thelatterspecieshavebeenaddedbyKoidzumi (1930), other in Morus species a cleartendencytowards lacking Leroy(1949),andLegame(1973),basedon material a trinervateor subpalmate pattern,thusbeingmore fromPeru,Colombia,andArgentinarespectively, pronouncedly pinnately veined.Thestaminate infloreswhereasRusby(1911) addedonetoM.celtidifolia. cencesarespicateto racemose.ThepistillateinfloresthegenusintothreesubgenLeroy(1949)subdivided cencesaremostly(sub)capitate headsin thegenus,but era:Eumorus,GomphomorusLeroy (1949), andAfroinM.insignistheyareelongateandsimilartothestamimorusA.Chevalier(1949). Subgenus Eumora comprised nateones.Incontrast to otherMorus species,thebracts allspeciesexceptfortheneotropical M insignisandits areabundant, inM insignis. peltateandsubcoriaceous synonymM trianae,whichwereplacedin subg.Gom- Morusinsignisis associatedwithhumidmontaneor phomorus,andtheAfricanM.lactea(Sim)Mildbraed subtropical conditions,beinganelementof cloudfor(= M. mesozygiaA. Chevalier),whichwasplacedin est,butM.celtidifoliais a speciesof drierhabitats. Venation subg.Afromorus. pattemsof thelamina,abunTwo Morusspecies have been introducedfrom ofthe Asia:M.alba(rathercommon)andM.australisPoiret dance,shapeandtextureof thebracts,andfeatures andembryowereusedto distinguish pericarp thesub- (rare).ThelattercanbeeasilyconfusedwithM.celtidigenera.Thesedifferencesseemtojustifyinfrageneric folia, butit canbe distinguished by theca. 1mmlong butatmostatthesectionlevel.However, styleof thepistillateflower. subdivision, Morusis, remarkably enough,theonlysmallgenus of theMoraceaethathasnotbeenrevised,and,therefore,thenumberof species,between10and13,is uncertain. Thegenusis anorthern-temperate elementwith extensionsto thetropics,athighelevationsin tropical

Key to the neotropicalspeciesof Morus 1. Inflorescences 0.5-3.5 cm long; axillary buds to 5(-10) mm long .......................................... 1. M. celtidifolia 1. Inflorescences 3-13.5 cm long; axillary buds minute ............................ .......................... 2. M. insignis

1.MorusceltidifoliaKunth,in Humboldt &Bonpland, Nov. Gen.Sp. 2: 27. 1817.Type.Ecuador.Loja: Loja (as "Loxa")or Pichincha,Puembo,(? fl), Bonpland 3350 (holotype, P).

Fig. I

M. radulina,M. vernonli,andM. vitifoliaGreene, Leafl. Bot. Observ.Crit. 2: 112-120. 1910 and 121. 1911. Based on materialfromthe U.S.A. Tree,to 10(-20) m tall or shrub,dioecious or some-

Morus corylifolia Kunth,in Humboldt& Bonpland, timesmonoecious.Leafytwigs 1-3 mm thick,glabrous

Nov. Gen.Sp. 2: 27. 1817.Type.Ecuador.Pichincha: Puembo,(st), Bonpland 3102 (holotype,P). Morus microphylla Buckley,Proc.Acad. Sci. Philad. (1862): 8. 1863. Type.U.S.A. Texas:withoutlocality,Buckleys.n. (holotype,PH,photocopyseen). Morus mexicana Bentham,P1.Hartw.71. 1841.Type. Mexico. Oaxaca:Tehuacan,1839 (a), Hartweg 514 (holotype,BM; isotypes, B, G, K, NY, P). Morus mollis Rusby, Bull. TorreyBot. Club. 38: 145. 1911; Standley,Contr.U.S. Natl. Herb.22: 204. 1922. Type:Mexico. Oaxaca:nr. Cuicatlan, 14 Jul 1910 (2 fl-fr),Rusby s.n. (holotype,NY?, not traced). Also: Morus albida, M. arbuscula, M. betulifolia, M. canina, M. confinis, M. crataegifolia, M. goldmanii, M. grisea, M. microphilyra, M. pandurata,

or sparsely to densely puberulous to hirtellous to tomentose; axillary buds to 5(-1 0) mm long, the outer scale-leaves coriaceous. Lamina narrowlyto broadly ovate to oblong, 1-20)x 0.5-9 cm, entireor 3-5-lobed (to -parted),? inequilateral,chartaceous;apex acuminate; base acute to obtuse to truncateto (sub)cordate; marginregularlyto irregularly,coarsely to finely serrate-dentate;uppersurfacesmooth and sparselyhairy or scabridulousto scabrous,sometimes? bullate;lower surfacebarbatein the axils of the lower lateralveins, otherwiseglabrousor sparselyto densely hirtellousto tomentose;venation almostplane above, ? prominent beneath;lateralveins 4-8 pairs;tertiaryvenationpartly scalariforn;petiole 0.2-4.5 cm long, 0.5-1.5 mm thick,

TAXONOMICTREATMENT

27

Fig. 1. Morus celtidifolia. 1. Leafy twig with staminateinflorescence(Buchtien3147). 2. Leafy twig with infructescences(Rosas R. 269). 3. Leafy twig with infructescences(Palmer149). 4. Leafy twig with infructescences(Palmer143). 5. Staminateflower.6. Pistillodes(McVaugh7897). 7. Staminateflower.8. Pistillode(Hartweg 514). 9. Pistillateflower. 10. Pistil. 11, lla. Fruit.12. Embryo(Pringle 6791). [By P. Pardoen,Utrecht.]

28 glabrousto denselyhirtellousto tomentose;stipules0.31 cm long, sparsely to densely tomentose, caducous. Inflorescences unisexual, sometimes bisexual, racemose to spicate to (sub)capitate,0.5-3.5 cm long, includmgthe 0.2-1.6 cm long peduncle,the peduncleand rachis sparsely to densely tomentose;flowers numerous to few, sometimes 1, distantor ? crowded, sessile or to 2(-3) mm long pedicellate, mostly ebracteate, sometimes with 1-2, triangularto ovate, puberulous bracts,to 1 mm long; perianthof staminateflower 22.5 mm long, 4-fid to -parted,sparsely to densely tomentose to puberulous;filaments 3-3.5 mm long; anthers 1-1.2 x 0.8-1 mm; pistillode quadrangularor 2-lobed, 0.5-0.7 mm long; tepals of pistillate flower pairwiseunequal,ca. 2 mm long, ? conduplicate,puberulous to tomentellousat the apex;style 0-0.2 mm long; stigmas0.5-2 mm long.Fruitingperianthto 3 mm long, red (to blackish);fruitca. 2.5-3 x 2-2.5 mm; seed ca. 1.5-2 x 1.5-2 mm. Distribution (see Fig. 72.1). Fromthe southwestem U.S.A to Honduras;in deciduous forest; to 2250 m. Also in SouthAmerica,in theAndes,butthereprobably introducedas a fruittree. Representativespecimensexamined.Severalcollections, includingtype collectionsof the speciesdescribed by Greene (1910-1911) from U.S.A.: Arizona, New Mexico, and Texasare not listed here. MEXICO. CHIAPAS: Mun. Tenejapa,Yashanal,ca. 2000 m, 17 Mar 1965 (a'), Breedlove 9393 (CAS, F,

MICH);Mun. Zinacantan,Hwy. 190, Joigel, 1900 m, 29 Aug 1976 (st), Breedlove39787 (MO);Mun.Venus5 km S of Aguatenango,1725m, 11 Apr tianoCarranza, 1981(? fl), Breedlove50802 (CAS,MO,NY);LakeJusnajav,16 km NW of Comitan,ca. 1750 m, 29 Apr 1945 (st), Sharp45445 (MO,NA); nr.Amatenangodel Valle, ca. 1900m, 14 Mar 1967(V), ShilomTon2170 (CAS,F, NY). CHIHUAHUA: Maguariche,SW of San Juanito,20 Jul 1957 (st), Knobloch96 (BM); W of Meoqui, 8 Jul 1936 (a), LeSueur 598 (F, K, MO, NY); nr. Chihuahua, 1300 m, 1-21 May 1908 (? fr), Palmer149 (F, K, MO, NA, NY, US), Apr 1886 (e'), Pringle 707 (BM, F, K,

MICH,MO,NA, NY,P, US). COAHUILA:Saltillo,29 Mar 1847 (a), Gregg 374 (MO);Mina El Popo, 2 km S of CafionEl Diablo,1600m, 29 Jul 1973(st),M. C. Johnston et al. 11929a (MO); Maguariche,SW of San Juanito, 20 Jul 1957 (st), Knobloch96 (BM);Hermanas,20 Apr 1939 (? fr), Marsh1600 (F); Saltillo, 11 Apr 1905 (? fr), Palmer 563 (F, MO, US); Canion de Indio Felipe, SierraHechiceros,Chihuahua-border, 14 Jun 1941(? fr), Stewart518 (F). DURANGO: SanRam6n,21 Apr-18May 1906 (? fr), Palmer 143 (C, F, K, MO, NA, NY, US, WIS).HIDALGO: 8 km N of Tianguistenango,1400 m, 26 Mar 1981 (? fl-fr),HernandezM. et al. 5688 (MO); Mun. Cardonal,Barrancade Tolontongo,3 May 1978 (? fr), Hiriart et al. 137 (MO); nr. Tula, ca. 2250 m, 13 Apr 1898 (? fr), Pringle 6791 (BM, F, G, K, LE, MICH,

MO,NY,P, US, Z); nr.Dublan,ca. 2200 m, 18 Apr 1904 (V), Pringle13197 (C, F, K, MICH,US), 2 Jul 1901(st), Roseet al. 5289 (US);aboveCuevaHumada,ca. 2250 m,

FLORANEOTROPICA 19 Mar 1946 (2 fl-fr), Sharp 46185 (NY). JALISCO: 16 km S of El Chanta, ca. 25 km SE of Autlan, 1450 m, 18 Mar 1965 (e), McVaugh 23054 (MICH). M?xIco: Dtto. Temascaltepec, La Sierrita, 3 Mar 1938 (ei), Hinton 3462 (K, MO, NA, NY, US), Nanchititla, 16 Mar 1933 (c), Hinton 3613 & 3627 (K, MICH, NY, US); Telpintla, 1840 m, 5 May 1933 (9 fr), Hinton 3843 (K, MO, NY, US); Mexico City, 3 May 1923 (od),Reiche s.n. (US). MICHOACAN: Nr. Morelia, Loma Santa Maria, 1900 m, 7 Aug 1918 (st), Bro. Arsene 6585 (MO, US); Quinceo, 6 km NW of Morelia, 2200 m, 18 Sep 1977 (st), Iltis et al. 168 (WIS); 6 km NW of Tuxpan, 1845 m, 3 Apr 1981 (? fl-fr), Soto Nuiiez 2934 (MO, NY). NUEVOLE6N: Galeana, 2 Oct 1958 (st), Jones 22731 & 22734 (WIS); SW of Galeana, 21 Jul 1934 (st), Mueller et al. 1138 (F); Mun. Villa Santiago, between Potrero Redondo & Laguna Sanchez, 8-15 Aug 1937 (st), Muller 2727 (MICH, NA). OAXACA: Tehuacan, 1840 (c), Hartweg 514 (B, K), 1841 (e^),Hartweg 514 (NY); Chinantla, (e), Liebmann 14212 (C); Tehuaca, (st), Liebmann 14213 & 14216 (C); 4 km NE of Ixtlan de Juarez, ca. 2200 m, 24 Apr 1982 (? fr), Lorence 4128 (MO); 18 km SE of Mishuatlan, rd. Oaxaca-Puerto Angel, km 116, 2000 m, 16 Apr 1965 (? fl-fr), Rzedowski 19662 (MICH). PUEBLA: Nr. Puebla, 2100-2200 m, 21 Mar 1907 (oi + ? fl), Bro. Arsene 910 (A, MO, US); nr. Pueblo, Mayoraygo, 20 May 1907 (? fr), Bro. Arsene 1142 (US); Queretaro, 1910-1913 ('), Bro. Arsene 10594 (US); rd. San MartinEl Verde, km 10, 2290 m, 4 Apr 1980 (? fl), Rodriguez F 24 (MO). SAN LuIs POTOSf:Charcas, 7 Aug 1934 (st), Lundell 5471 (F, US); 10 mi NE of Ciudad del Maiz, 1400-1600 m, 7 May 1949 (st), McVaugh19432 (MICH); Rascon, Jun 1911 (Q fl-fr), Purpus 5501 (BM, F, MO, NY, US); 3 km SE of Ahuacatlam,Xilitle, 10 Apr 1960 (? fr), Rzedowski 12375 (MICH). SINALOA: Mun. Badiraguato, Sierra Surutato, 3 mi N of Los Ornos, rd. to Ocurahui, ca. 2150 m, 2 Oct 1970 (st), Breedlove et al. 18374 (MICH). SONORA: 8 mi E of Imuris, nr. mouth of canyon of Rio Magdalena, 24 Mar 1934 (e), Ferris 8805 (NY, US); Rio Mayo, Curohui, ca. 1150 m, 4 Apr 1938 (? fl-fr), H. S. Gentry 3641 (F, K, MO), (e), H. S. Gentry 3657 (F, K, MO, NA); Fronteras, 25 Sep 1890 (st), Hartman 974 (MO, NY); Magdalena, 24 May 1925 (st), Kennedy 7038 (MO, US); Guadeloupe basin, 27 Aug 1893 (st), Morton 2054 & 2055 (US); 5 mi E of Esqueda, rd. to Rio de la Tierra, ca. 1400 m, 14 May 1948 (? fr), Wiggins 11764 (MICH, US). TAMAULIPAS: Sierra San Carlos, nr. Marmolejo, La Morita, 1700 m, 2 Aug 1930 (st), Bartlett 10775 (MICH, US); Sierra San Carlos, Cerro Zamora, nr. El Milagro, 25 Aug 1930 (st), Bartlett 11124 (F, MICH, US); nr. Victoria, 1 Feb-9 Apr 1907 (? fl), Palmer 158 (F, K, MO, NY, US); Mun. G6mez Farias, Sierra de Guatemala, between San Pablo & Rancho del Cielo Biol. Station, ca. 1150 m, 21 Jun 1971 (st), Sullivan 649 (NA, NY), rd. to Julilo, ca. 1300 m, 28 Jun 1971 (st), Sullivan 653 (NA, NY). VERACRUZ: Laguna del Castillo, El Castillo, 1260 m, 19 Mar 1976 (ei), MdrquezR. et al. 556 (F, NY); Toxtongo, Coscomatepec, May 1937 (? fl-fr), Matuda 1384 (K, MICH, MO, NY); valley of Orizaba, Jul 1857 (? & e), Mohr s.n. (US); 8.5 km W of Chocaman, 1760 m, 18 Nov 1981 (st), Nee 23260 (F, MO); 2.5 km N of Coatepec, 1100 m, 25 Mar 1983 (oi), Nee et

TAXONOMIC TREATMENT al. 26299 (F, MO, NY); Santa Ana Atzacan, N of Orizaba, 1320 m, 9 Apr 1967 ($ fl-fr), Rosas R. 269 (BM, MO, U). GUATEMALA. Without locality, 1846 (? fr), SavChichoy,20 Sep 1938(e), age 105 (P). CHIMALTENAGO: J. R. Johnston 1358 (F); Rio Guacalate, SE of Chimaltenengo, 1700 m, 14-23 Dec 1940 (st), Standley 79986 (F). GUATEMALA: Nr. Finca La Aurora, 1500 m, 1938-

1939 (? fr), Aguilar 316 (F). HUEHUETENANGO: Nr. Aguacatan, 1200 m, 14 Jan 1939 (st), Standley 62563 (F); nr. Aguacatan, Rio San Juan, 1600 m, 24 Dec 1940 (ei), Standley 81359 (F, US); rd. Huehuetenango-Aguacatan, km 13-14, 1950 m, 21 Dec 1940 (st), Standley 81255 (F), 2 Jan 1941 (st), Standley 91968 (F); nr. San Sebastian, 2000-2200 m, 13 Aug 1942 (st), Steyermark 50515a (F); Sierra de las Cuchumatanes,Rio Azul, below Jacaltenango, 1400 m, 4 Sep 1942 (st), Steyermark 51856 (F, US). QUEZALTENANGO: Below Santa Mariade Jesuis, 1350-1380 m, 11 Mar 1939 (st), Standley 68333 (F).

29

totype,Berg, Fl. Ecuador60: 12. 1998, P; isolectoFig. 2 types, BM, F, G, K). MorusperuvianaPlanchonex Koidzumi,Fl. Symb. Orient.-Asiat.88. 1930. Type. Peru.San Carlos, 1835 (9 fl), Mathews1634 (lectotype,designated here, K; isolectotype,BM). MorustrianaeLeroy,Bull. Mus. Hist. Nat. (Paris), Ser. 2, 21: 731. 1949. Type.Colombia.Quindio: La Palmilla,(9 fl-fr), Triana870 (holotype, P; isotypes, BM, K). Morus marmolii Legname, Lilloa 33: 334. 1973. Type.Argentina.Salta:Dtto. SantaVictoria,between Barituand Porogal,Marmolet al. 8880 (holotype, LIL-n.v.).

Tree,to 15 m tall, dioecious. Leafy'twigs 1.5-4 mm thick,white to brownishtomentoseto subvelutinousor to sparselypubescent,glabrescent;axillarybudsminute. Leavesellipticto oblong,4-25 x 2.5-14 cm,mostlybroadSACATEPEQUEZ:SE of Alotenango,1000-1260m, 9 Feb estbelowthemiddle,? inequilateral,chartaceousto sub1939 (st), Standley 65029 (F). coriaceous;apex acuminate;base obtuseto emarginate; EL SALVADOR: Parque Nacional de Montecristo, uppersurfacescabrousto scamargin (serrate-)dentate; 1845 m, 13 Mar 1988 (d'), Reyna JBL 653 (MO). often ? bullate;lower suralmost smooth, bridulous or HONDURAS. OLANCHO: MontanlaSan Estaban,N of Juticalpa, (st) 1979, Secci6n Catastro Forestal de Hon- face tomentoseto sparselypubescentor (sub)glabrous; venationalmost plane to impressedabove, prominent duras s.n. (EAP). beneath;lateralveins 4-8 pairs;tertiaryvenationlargely ECUADOR. Without locality, (st), Acosta S. 11660 (F). LOJA: Pampa Bonita, 26 May 1982 (st), Vivar C. scalariform;petiole 0.5-3(-3.5) cm long, puberulous 1547 (LOJA); Cento Loja, La Vega Grande, 1040 m, 29 to tomentose;stipules 0.5-1.5 cm long, tomentose to Mar 1985 (st), VivarC. 2345 (LOJA).PICHINCHA: Hda. sparselypubescent,caducous.Staminateinflorescences San Carlos, Cotocolao, Sodiro s.n. (Q). spicate;peduncle 0.2-0.6 cm long, puberulousto toPERU. Without locality, (? fr), Gay 2165 (P); (e), mentose; spike (l-)3-9 cm long; flowers ? crowded; Matthew750 (G). APURIMAC: Challhauaca,Aymaraes, perianthwith(almost)freetepals,ca. 2 mm long,puber2887 m, 19 Feb 1987 (st), Nuinez 7156 (MO);Abancay, ulous;filamentsca. 4 mm long; anthersca. 0.8-1 x 0.81 Sep 1920 (st), Stordys.n. (K). PIURA: Huancabamba, 1 mm; pistillode quadrangular, ca. 0.5 mm long, ca. 2000 m, Aug 1943 (d2), Sandeman 4279 (K). bractspeltate,to 1.5mm diam.,sparsely (sub)glabrous; BOLIVIA. LA PAZ: Ilimani, Cotafia, 2450 m, Nov Pistillate inflorescencesspicate;peduncle puberulous. 1911 (c), Buchtien 3147 (F, NA, NY, US). 0.6-1.5 cm long, puberulousto tomentose;spike (1-) According to the label dataof collection Bernardi 3-13.5 cm long; flowers ? crowded;perianth1-2 mm 545, the species is cultivatedandnaturalizedin Merida, long, 4-parted,the tepalsciliolate;ovaryca. 1mm long; Venezuela,at ca. 1800 m, butis not common.The small style 0.5-1 mm long; stigmas 1.5-2 mm long, pubnumberof collectionsfromSouthAmerica,lackof clear erulous; bractspeltate (or sometimes 1-2 on the peindicationsof occurrencein naturalvegetation,andthe dunclebasallyattached),to 1 mm (on the peduncleto 2 scarcityof recentcollections seem to indicatethatthe mm) diam.,sparselypuberulous.Fruitingspikeca. 0.8 presenceof Morusceltidifolia in SouthAmericais due cm diam.;fruitingperianth2.5-3 mm long, fleshy,redto introduction.It has become outmodedas a fruittree. dish (exceptfor the margins);fruitca. 2.5 x 2.5 m; seed Local names and use. Guatemala:Huehuetenango: ca. 2 x 2 mm. mora. Ecuador:Loja: palo blanco, sota. Peru:ApuriDistribution(see Fig. 72.2). FromsouthemMexico mac:moranegra;Piura:moral. throughCentralAmericaandtheAndesregionto northIn several partsof South America the species has em Argentina,also in the coastal mountain range in been in cultivation as a fruit tree, fruits being sold on Venezuela,with more or less clear gaps in the rangeof marketsin the 19thcentury. distribution;in cloud forestor moist subtropicalforest; at ca. 1300-3000 m. Representative specimens examined. MEXICO. 2. Morus insignis Bureau,in De Candolle,Prodr.17: CHIAPAS: Mun.Motozintlade Mendonza,rd. Huixtla247. 1873; Bentham.P1.Hartw.253. 1846; Leroy, Motozintla, km 45-50, 1900 m, 28 Dec 1972 (e), Bull. Mus. Hist. Nat. (Paris),Ser.2, 21: 731. 1949. Breedloveet al. 31069 (CAS). Type.Colombia.Magdalena:SierraNevadade Santa Between Finca PieGUATEMALA. EL PROGRESO: Marta, Rio Ibacha, Jan 1844 (e), Linden 1619 (lec-

monte & top of Montafia Piemonte, along Joya Pacayal,

FLORANEOTROPICA

30

AX

Fig. 2. Morusinsignis. 1. Leafy twig with pistillateinflorescences(Little15565). 2. Leafy twig with stamiet al 98674). 4, 4a. Pistillateflower with bract. nate inflorescence.3. Pistillateflowerand pistillode(Steyermark 5. Pistil. 6. Fruitingperianth.7. Fruit.8. Seed. 9. Embryo(Little 15565). [By P. Pardoen,Utrecht.]

TAXONOMICTREATMENT 2500-3000 m, 7 Feb 1942 (st), Steyermark43644 (F, MO, US). QUEZALTENANGO:Slopes of Volcan de Zunil, above Aguas Amargas, 2430-2850 m, 17 Feb 1939 (st), Standley 65384 (A, F, S); Quebrada El Pocito, S of San Martin Chiliverde, 2200 m, 27 Jan 1941 ($ fr), Standley 85036 (F); slopes of Volcan de Zunil, nr. Fuentes Georginas, 2300-2500 m, 3 Feb 1941 (od),Standley 85950 (F, US). SAN MARCOS: Barranco Eminencia, above San Rafael Pie de la Cuesta, 2100-2400 m, 15 Mar 1939 (st), Standley 68640 (F); Barranco Eminencia, between San Marcos & San Rafael Pie de la Cuesta, 2500-2700 m, 6 Feb 1941 (st), Standley 86288 (F); Rio Vega, between San Rafael & Volcan Tacana, nr. Mexico border, 2500-3000 m, 21 Feb 1940 (st), Steyermark 36300 (F); between Todos Santos & Finca El Porvenir, lower slopes of Volcan Tajumulco, 1 Mar 1940 (st), Steyermark 37046 (F). COSTA RICA. ALAJUELA:Cant6n Alfaro Ruiz, Palmares, 9 May 1938 (? fl-fr), A. Smith 519 (A, EAP, F, NY), 9 Jan 1939 (st), A. Smith 1423 (NY). CARTAGO: Cuarco, 18 Oct 1966 (st), Mora V 8 (CR, F). HEREDIA: Volcan Barva, San Jose de la Montafia, 1950 m, 17 May 1966 (? fr), Fosberg et al. 47800 (MO, NY, US); nr. Porrosati, S slope of Volcan Barva, 2000 m, 21 Apr 1989 (? fl-fr), Hammel et al. 17312 (BG, F, MO); Volcan Barba, Rio San Miguel, 1900 m, 26 Jun 1965 (ae),Jimenez M. 3237 (CR, F); between Volcan Poas & Volcan Barva, Vara Blanca, 1615 m, May 1938 (e), Skutch 3779 (K, MO, US). SAN Jost: Cerros de Escazu, Cedral, 16002300 m, 9 Oct 1991 (a), Morales 177 (BG); Las Nubes, 1500-1900 m, 20-22 Mar 1924 (? fr), Standley 38808 (F, GH, NY, US). PANAMA. CHIRIQUi: La Popa, above Boquete, 1500-2500 m, 21 Mar 1977 (a), DArcy 10863 (MO); Cerro Punta, 2000 m, 8 Sep 1971 (ae), Lao 335 (MO); 6 mi above Cerro Punta, Boquete trail, ca. 2500 m, 7 Mar 1974 (? fl), Tyson 7152 (MO, NA). COLOMBIA. ANTIOQUIA: Mun. Frontino, upper Rio Cuevas, 1780 m, 23 Sep 1987 (a), D. Sanchez et al. 1607 (HUA, MO); rd. Campamento-Las Brisas, 1570-1800 m, 8 Feb 1986 (? fr), Callejas et al. 2497 (BG, MO, NY); Mun. Campamento,Vrda. Llanadas, 6-12 km W of Campamento, 1650-1810, 7 Sep 1989 (v), Callejas et al. 8264 (BG, HUA); Mun. Jardin, rd. Jardin-Rio Sucio, km 9, 2300-2400 m, MacDougal et al. 3563 (BG). BOYACA:Rio Casanare, between "El Paleo" & Sacama, 2000-2300 m, 16 Jul 1967 (st), Jaramillo et al. 2743A (K). CUNDINAMARCA:W of Bogota, 1.5 km W of El Salto de Tequendama,rd. to El Colegio, 2400 m, 12 Apr 1972 (d'), Barclay et al. 3313 (NA, US); between Sebastopol & Las Escaleras, 2300-2400 m, 21 Mar 1942 (? fr), Cuatrecasas 13573 (F); San Antonio de Tena, Boquer6n de la Mesa, Jul 1844, (? fl), Goudot s.n. (G, P); "Prov. Bogota," Jul 1844 (a'), Goudot s.n. (K); Pacho, (e), Hartweg 1390 (BM, G, K); "Prov. Bogota," El Amacachal, 2300 m, (a), Triana 870 (P). GUAJIRA: Cuchilla de San Antonio, above Puerto Viejo, Rio Garavito, 1710 m, 19 Aug 1986 (st), Gentry et al. 55352 (BG, MO); N slope of Cerro del Espejo, Serraniade Perija, Cesar border, ca. 2560 m, 28 Apr 1987 (? fr), Gentry et al. 57176

31 (BG, MO). HUILA: Rd. Popayan-La Plata, 2000-2300 m, 31 Jul 1979 (? fl), Cabrera R. et al. 5086 (BG, MO); Finca Merenberg, E of Leticia, nr. Cauca border, 22752300 m, 7 Jul 1984 (st), Gentry et al. 47678 (BG, MO), 2 Apr 1986 (st), Gentry et al. 53930 (BG, MO). NARINo: La Planada Reserve, 7 km from Chucunes, 1800 m, 25 Jul 1986 (st), Gentry et al. 55104 (MO), 22 Dec 1987 (st), Gentry et al. 59713 (MO). QUINDIO:Mun. Calarca, Cgto. Quebrada Negra, Vrda. Vista Hermosa, Finca La Floresta, 1650-2150 m, 2 Mar 1992 (a), Agudelo et al. 895 (COL). RISARALDA:Mun. Pereira, Parque Natural Regional Ucumari, between La Pastora & Ceylan, 2400 m, 2 Dec 1989 (da), Franco et al. 2895 (COL). VALLE: Rio Cali, Quebrada Honda, La Glorieta, rd. to Miralindo, 2100-2250 m, 31 Oct 1944 (e), Cuatrecasas 18405 (F, VALLE); Rio Cali, confluence of Rio Pichidecito & Rio Pichinde, 1580-1650 m, 7 Nov 1944 (? fl-fr), Cuatrecasas 18777 (F, VALLE); Rio Pichinde, between Los Carpatos & El Olivo, 2200-2900 m, 26 Jul-6 Aug 1946 (a), Cuatrecasas 21735 (F, US). VENEZUELA.

ARAGUA: Colonia Tovar, (? fl), Moritz

FEDERAL:Cerro El Avila, 2000 1703 (BM, K). DISTRITO m, 20 Sep 1992, Meier 2807 (BG). MERIDA:Filo SaiSai, El Carrizal, 1800 m, 24 Feb 1955 (st), Bernardi 2068 (NY); 12 km WNW of Ejido, 1700 m, 30 Sep 1953 (? fl), Little 15565 (VEN); 12 km WNW of Ejido, ca. 1700 m, 19 Oct 1953 (? fl-fr), Maguire 39438 (NY); Dtto. Andres Bello, El Cinaral, 1800-1900 m, 17 Nov 1981 (a'), Marcano-Berti 306-981 (MO, U); Rio Capuri, between Cauagua & El Molino, 2530-2715 m, 11 May 1944 (st), Steyermark56473 (F, VEN). TACHIRA: Paramo de Tama, nr. Quebrada Buena Vista, 2200-2450 m, 2224 May 1967 (a'), Steyermark et al. 98674 (NY, U, US, VEN); between Villa Paez & Tama, Rio Tachira, 20002200 m, 25 May 1967 ($ fl), Steyermark et al. 98852 (MO, US, VEN); 30 km S of Alquitrana, Rio Quinimari, 2400 m, 21 Jan 1968 (a'), Steyermarket al. 101253 (VEN). TRUJILLO:Dtto. Bocon6, Montafias de Missisi, old rd. Trujillo-Bocon6, 12 km NW of Bocon6, 200-2400 m, 31 Oct 1990 (a'), Dorr et al. 7523 (BG, MO, NY). ECUADOR.

AZUAY: Guaillo,

2300 m, 30 Oct 1994

(R fl), Cornejo et al. 3479 (BG). BOLIVAR:Milagro, 2000 m, 7 Oct 1943 (st), Acosta S. 6197 (F); San Pablo de Telimbela, 1300 m, 20 Nov 1943 (st), Acosta S. 6933 (F); La Esperanza, rd. to Huamboya, 2000 m, 15 Feb 1944 (st), Acosta S. 7383 (F); rd. Chillanes-Tiquibuso, San Jose de Guayabal, 2100 m, 3 Sep 1987 (? fl-fr), Zak et al. 2705 (GH, MO, US); rd. Chillanes-Bucay, Hda. Tiquibuso, 2100 m, 10 Sep 1987 (? fl-fr), Zak et al. 2880 (MO, US); rd. Chillanes-Bucay, La Escuala, 2300-2450 m, 11 Sep 1987 (? fl-fr), Zak et al. 2933 (LL, MO, NY). CARCHI:Above Maldonado, 2500 m, 1 Aug 1989 (? fl-fr), Werif et al. 10846 (BG, QCNE). CHIMBORAZO:Las Cochas, NE of Huigro, ca. 2700 m, 21 Oct 1944 (? fr), Wiggins 11017 (US). EL ORO: Rio Palma, nr. Pampa de los Cedros, S of Cerro ChivoTurco, 2285-2315 m, 11 Aug 1943 (st), Steyermark 53781 (F); Pampade los Cedros, NE of San Pablo & Curicapo, 2285-2430 m, 12 Aug 1943 ($ fl), Steyermark

FLORANEOTROPICA

32 53817 (F). MORONA-SANTIAGO (?): QuebradaHonda,

29 Apr 1981 (st), VazquezAvila 346 (U). SANTA CRUZ:

nr. Rancho Achupallas, 2500-2700 m, 10 Oct 1943 ), Steyermark54570 (F,NY). NAPO: Nr.Baeza, 1850-

Prov. Florida, Samaipata, Chorolque, 1800 m, 28 Apr 1993 (st), Cornejo et al. 219 (USZ); Prov. Caballero, 15-25 km N of San Juan Potrero, Cerro Bravo, 20002500 m, 6 Jun 1992 (a), Killeen et al. 4109 (USZ); Prov. Manuel Maria Caballero, 50 km N of Mataral, rd. Santa Cruz-Comarapa, 2000-2100 m, 26 May 1989 (e), D. N. Smith et al. 13411 (BG, LPB, MO); Prov. Caballero, rd. Comarapa-Cochabamba, Siberia-Empalme, 2300 m, 8-9 May 1992 (? fl), I. G. Vargas et al. 1279 (USZ). TARIJA: Prov. Arce, upper Rio Cambari,trail Sidras-Tariquia, 1200 m, 13 Oct 1983 (q fr), Solomon 11201 (LPB). ARGENTINA. SALTA: Depto. Santa Victoria, rd. Porongal-Baritu, km 8, 1500 m, 19 Sep 1972 (e),

2000 m, 22 Sep 1977 (a), Berg et al. 442 (AAU, QCA,

U); rd. Baeza-Tena,1980 m, 14 Aug 1975 (e), Littleet al. 184 (MO, QAME);slopes VolcanSumaco,between El Pacto & CerroHuahuaSumaco, 1700-1800 m, 24 Oct 1989 (o), Palacios et al. 4712 (BG, MO, QAME,

QCNE);CantonEl Chaco, Rio Quijos, ca. 10 km S of Reventador,1450 m, 6-10 Oct 1990 (o), Palacios 5982 (BG, MO); CantonEl Chaco, Las Palmas, 1600-1750 m, 12 Oct 1990 (2 fl-fr), Palacios 6228 (BG, MO);nr. Baeza, ca. 2000 m, 3-4 Oct 1992 (a'), Valenciaet al. 2809 (BG).PICHINCHA: Chiriboga,nr.Rio Saloya,1800 m, 12 Jun 1989 (a), Cer6n et al. 6818 (BG, QAME, QCNE);old rd. Quito-SantoDomingode los Colorados, Chiriboga,1600-1800 m, 8 Mar 1990 (a), Cer6n et al. 8923 (BG, QCNE);5 km E of Nanegal,1550 m, 11 Feb 1991 (st), Gentryet al. 73239 (MO); rd. NanegalitoArmenia-Lomade San Jose, "LaVueltaBrava,"19002200 m, 9 Dec 1987 (e), Zaket al. 3215 (MO).TUNGURAHUA: 11 km E of Banios, 3 Jun 1968 (a' fr), Holm-Nielsen et al. 272 (AAU, C, S). PERU. Without locality, (a' + ? fr), Ruiz & Pavon s.n. (B, F, G, P). AMAZONAS: Prov. Bongara, Shillac,2300 m, 31 Aug-2 Sep 1983 (o), D. N. Smith et al. 4900 (BG, MO, NY). CAJAMARCA:ParqueNacional de Cutervo, San Andres de Cutervo, 2250 m, 3 Aug 1988 (a'), C. Diaz et al. 2967 (BG, MO). JUNMN:Tarma-Chanchamayo border, 45 km S of San Ram6n, 1800 m, 15 Oct 1982 (st), Foster 9172 (MO). LIBERTAD: Prov. Pataz, Rio Mishollo, 2400-2500 m, Aug 1914 (a), Weberbauer7040 (F, GH, US). PASCO: Prov. Oxapampa, 5 km SE of Oxapampa, 1850 m, 31 Jan 1983 (a), D. N. Smith 3149 (BG, MO, NY), 25 May 1983 (a'), D. N. Smith 4168 (F, MO); Prov. Oxapampa, Rio San Alberto, E of Oxapampa, 2600 m, 25 Jul 1984 (a), D. N. Smith 8027 (MO, NY, TEX); Prov. Oxapampa, Palmazui,ca. 2150 m, 3 Oct 1984 (a), D. N. Smith 8650 (MO, NY); Prov. Oxapampa, Quebrada Honda, 1400 m, 15 Aug 1968 (? fl), VasquezA. 179 (F, MO, U). PIURA: Prov. Huancabamba, Canchaque, between "Chorro Blanco" & "War War," 2000-2500 m, 18 Jan 1989 (a), Diaz et al. 3186 (MO). SAN MARTiN: Prov. Rioja, rd. Pedro Ruiz-Moyobamba, km 390, Venceremos, 1800 m, 27 Jul 1983 (a), D. N. Smith 4382 (MO, MOL, NY, TEX), 2100 m, 7-9 Aug 1983 (a), D. N. Smith et al. 4749 (BG, MO); Prov. Mariscal Caceres, Rio Abiseo National Park, 2650-2750 m, 25 Jul 1987 (a'), Young et al. 5005 (F, MOL). BOLIVIA. LA PAZ: Prov. Murillo, Rio Zongo, Cahua, 10 km from La Paz, 2170 m, 7 Apr. 1979 (st), Beck 1209 (BG, LPB, MO), 2000-2200 m, 20 Aug 1979 (? fl), Beck 2199 (BG, LPB); Prov. Nor Yungas, 9 km from Huancane, rd. to San Isidro, 2450 m, D. N. Smith et al. 13879 (BG, BOLV, MO); Prov. Nor Yungas, 14.3 km SW of Yolosa, rd. to Chuspipata, 2000 m, 23 Mar 1984 (st), Solomon et al. 12076 (MO); Prov. Murillo, Rio Zongo, 2600 m, 1 May 1987 (st), Solomon 16668 (BG, LPB, MO); Prov. Chapare, Rio Zongo, 2250 m,

Marmolet al. 9230 (C, K, LL, NY). Local names.Venezuela:Merida:huaguito;Tachira: rajito.Ecuador:Bolivar:guillo, sapanamarillo;Pichincha:huasca. EXCLUDEDNAMES

MorustinctoriaLinnaeus,Sp. P1.986. 1753 = Maclura tinctoria(Linnaeus)Steudel. MoruszanthoxylonLinnaeus, Syst. P1.ed. 2, 2: 1266. 1759 = Macluratinctoria(Linnaeus)Steudelsubsp. tinctoria.

Trophis 2. Trophis P. Browne, Civ. Nat. Hist. Jamaica 357. 1756, nom. conserv.; Linnaeus, Syst. Nat. ed. 10. 1289. 1759; Bureau, in De Candolle, Prodr. 17: 251. 1873; Bentham & Hooker, Gen. P1. 3(1): 365. 1880; Burger,Ann. Missouri Bot. Gard.49: 1. 1962; Corner, Gard. Bull. Straits Settlem. 19: 230. 1962;Berg,Proc.Kon.Ned. Akad.Wetensch., Ser.C, Biol. Med. Sci. 91: 345. 1988. Type. Trophis americana Linnaeus (= T racemosa (Linnaeus) Urban). Bucephalon Linnaeus, Sp. P1. 1190. 1753, nom. rejic. Type species. Bucephalon racemosum Linnaeus (= Trophis racemosa (Linnaeus) Urban). Olmedia Ruiz & Pav6n, Syst. Veg. Fl. Peruv. Chil. 1: 257. 1798; Berg, Fl. Neotropica 7: 14. 1972; Berg, Acta Bot. Neerl. 26: 73. 1977. Type. OlmediaasperaRuiz & Pav6n (= Trophiscaucana (Pittier) C. C. Berg). Malaisia Blanco, Fl. Filip. 789. 1837. Type. Malaisia tortuosaBlanco (= Trophisscandens (Loureiro) Hooker & Arnott). Dumartroya Gaudichaud, Voy. Bonite, Bot. Atlas t. 97. 1847-1848. Type. Dumartroya fagifolia Gaudichaud (= Trophis scandens (Loureiro) Hooker & Arnott). Cephalotrophis Blume, Mus. Bot. Lugduno-Batavum 2: 75. 1856. Type. Cephalotrophisjavanica Blume (= Trophisscandens (Loureiro) Hooker & Arnott).

TAXONOMICTREATMENT Maillardia Frappier ex Duchartre, in Maillard, Ann. Notes Reunion, Bot. 1: 146 (or 148). 1862 or in Maillard, Notes sur l'ile de La Reunion, Annex P. 3. 1862?. Type. Maillardia borbonica Duchartre (= Trophis borbonica (Duchartre) C. C. Berg). Calpidochiamys Diels, Bot. Jahrb. Syst. 67: 172. 1935. Type. CalpidochlamysdrupaceaDiels (= Trophis drupacea (Diels) Corner). Skutchia Pax & K. Hoffmann ex Morton, J. Wash. Acad. Sci. 27: 306. 1937, proposed as a new genus in Euphorbiaceae;Burger,Ann. Missouri Bot. Gard. 49: 6. 1962. Type. Skutchia caudata Morton (= Trophismexicana(Liebmann) Bureau).

Treesor shrubs, dioecious; uncinatehairs lacking. veined; Leavesalternateanddistichous;laminapinnately margindentateto entire;stipulesfree, lateral.Inflorescences solitary,pairedor moretogetherin the leaf axils bracteate,thebracts orjustbelowtheleaves,pedunculate, basallyattached(to subpeltate).Staminateinflorescences racemoseto spicatewitha distinctabaxialsterilestripor discoid-capitateandinvolucrate;flowersin racemoseto spicate inflorescences in longitudinalrows, distantor crowded; tepals basally connate, valvate in the bud; sometimesfilipistillodepresent,mostly quadrangular, form.Pistillate inflorescencesracemoseto spicatewith 3-ca. 35 flowers or with 1 (or 2) flower(s) and involucrate;perianthtubularor collar-shapedto campanulate; ovary partlyadnateto the perianthor free; stigmas 2, equal,filiform.Fruitenclosedandthenfreeor adnateto the enlargedfleshy, reddishto orange(or to purplish) perianth,or the fruitfree, drupaceousanddarkblue- to blackish-colored andtheperianthnot enlargednorfleshy, greenish;endocarp(thinly) crustaceous,often with a woody plug near the hilum; seed large, without endosperm;testawith a vascularizedpartbelow thehilum; embryo longitudinallyaligned; cotyledons equal and thick;radiclevery short,apical. Linnaeus(1753) initially adoptedPlumier'sname Bucephalon for this genus, but in 1759 he chose the name Trophis,thathadbeen establishedby P. Browne (1756).ThegenusTrophiswasrevisedby Bureau(1873). He recognized two species, referreda thirdone with uncertaintyto the genus, andexcludedthepaleotropical taxa which had been describedin the genus in course of time. Burger(1962) monographedthe genus again. He recognized four species and included the genus SkutchiaMorton (1937). In 1973 he describeda fifth species, T involucrata. In 1962 Cornerpublisheda revisedclassificationof the Moraceae, in which several genera, including Trophis,wereconsiderablyenlargedcomparedwithprevious concepts.In Corner'sconceptthe genusalsocomprisesseveralpaleotropicalspecies,namelythoseof the OldWorldgeneraCalpidochlamysandMaillardia.Berg (1988)remodelledthegenusagainandincludedtheAsian

33

genus Malaisia Blanco and the neotropicalgenus Olmedia.The lattergenuswas the type for the tribe Olmedieae(cf. Berg,1972),butit provedto be so differentfromothergenerainsertedin thetribethatit had to be excludedandthetribehadto be renamed(Berg, 1977a).Themainreasonto reconsideragainthe deweresomeexceptional limitation of thegenusTrophis featuresfoundin TrophisinvolucrataBurger(1973). Whenmaterialin fruitingstatebecameavailable,the speciesprovedtobequitedistinctfromtheotherTrophis species,astheperianthdidnotenlargeanddidnotbecomefleshyandorangeto redduringthedevelopment adark of thefruit,butremained unchanged, subtending blueto blackishdrupe.Similarfeaturesoccurin some andthevariagenusStreblus speciesof thepaleotropical tionin fruitsandfruitingperianthspresentin thisgeAs T involucrata nusis comparable withthatin Trophis. in Trophis, thegenusbecamedicouldbe maintained themonotypicOlmedia verseenoughto accommodate aswell.Intheenlargedgenussectionscouldbe distinrecognized thepreviously guished,largelyrepresenting smallergeneraand,in addition,a newonewascreated to accommodate T involucrata. members of Trophis aresmallormeTheneotropical areaxildiumn-sized treesorshrubs.Theinflorescences to laryoroccurjust belowtheleaves.Theyareracemose spicate,butin T caucanatheyare? obliquelydiscoidof andinvolucrate astheresultof condensation capitate of bracts.Theinflorescences therachisandenlargement similartothoseinthetribe of T caucanaareinstructure Castilleae. Thenumber of flowersin thepistillateinfloto one. variesfromnumerous rescencesof Trophis flowershavevalvatetepals,afeature by Thestaminate fromrelatedgenera. whichTrophis canbedistinguished of the Infourof thefiveneotropical speciestheperianth andbecomesenlarged, fleshy, floweris tubular pistillate andorangeto redin fruit,enclosingthefruit,whichis is disorfree.Trophis involucrata adnatetotheperianth above.Thefruiting peritinctinthisrespect,asindicated The anthof thisspeciesoftenbearsspine-like processes. specieshavebeen pollengrainsof someof the Trophis described byNiezgodaandNowaczyk(1976). IntheNeotropicsthe genusextendsfromCentral Americato AndeanSouthAmericaandto theGreater Antilles.Mostof thespeciesoccurinmoreorlessopen vegetation,such as forestmarginsandriverbanks. is anunderAccordingto thelabeldataT involuctrata storytreelet.It is somewhatdifficultto imaginehow withanemophily; cangotogether suchahabitandhabitat labeldatadonotindicatewhetherthetreeletsoccurat Six sectionscanberecograndomoralongstreamlets. thenatureof thedifnizedin thegenus.Considering as entitiescouldberegarded ferences,theinfrageneric ratherthansections.Therankof sections subgenera,

FLORANEOTROPICA

34 adoptedby Comer(1962) has beenmaintainedto avoid creationof new combinations. Threeof the sections (Calpidochlamys(Diels) Corner, Maillardia (Duchartre) Corner, and Malaisia

(Blanco) C. C. Berg) arepaleotropical,the othersneotropical(section Trophiswith threeclosely relatedspecies, andthe sections Echinocarpa and Olmedia,both monotypic).

Key to the neotropicalspeciesof Trophis 1. Laminascabrousor scabridulousabove and/orbeneath. 2. Laminainequilateral; inflorescences ..... 4. T. caucana distinctlyinvolucrate tertiaryvenationpartlyscalariform; tertiaryvenationreticulate;inflorescenceswithoutdistinctinvolucres. 2. Lamina(almost)equilateral; 3. Laminascabrousabove and often scabridulousbeneath,the marginusuallyentire;staminate flowerssessile; pistillateinflorescencesusuallywith morethan 3 flowers;perianthenlarged 3. T. racemosa in fruit, reddish,enclosingthe fruit............................................................... 3. Laminaonly scabridulousbeneath,the marginusually denticulate;staminateflowers pedicellate;pistillateinflorescenceswith 1-3 flowers;perianthnot enlargedin fruit,greenish,not 5. T. involucrata enclosingthe darkblue to blackishechinatefruit........................................................... 1. Laminasmooth above and beneath. 2. T. cuspidata 4. Laminadistinctlyhairy (hirtellousto patent-puberulous) beneath;1600-2600 m. beneath;0-1600(-2200) m. 4. Laminaglabrousor at most sparselyappressed-puberulous 5. Pistillateinflorescencesto 3.5 cm long, with (I-)3- 8(-10) flowers,the flowers sessile and usuallycrowded;staminateflowerssessile, theirperianthsdenselyhairy;fruitingperianth T. racemosa smoothor ribbed.3. 5. Pistillateinflorescencesto 14 cm long, with (2-)10- 35 flowers,the flowers often pedicellate and often ? spaced;staminateflowersoften pedicellate,theirperianthsusuallysparsely .1. T. mexicana hairy;fruitingperianthribbedand/orverrucate

2A. Trophis P. Browne sect. Trophis. Including Bucephalon Linnaeus and Skutchia Morton(see under the genus). Treesor shrubs.Staminateinflorescencesracemose or spicate;tepals 4, valvate in the bud. Pistillate inflorescences racemose or spicate with numerousto few flowers; tepals enlargedin fruit,reddish.Fruit adnate to the perianth;cotyledons equal. SectionTrophiscomprisesthreeneotropicalspecies. They areclosely relatedand sometimesdifficultto distinguish.

1. Trophis mexicana (Liebmann) Bureau, in De Candolle, Prodr.17: 253. 1873; Burger,Ann. MissouriBot. Gard.49: 13,t. 2. 1962.Soroceamexicana Liebmann,Kongel. DanskeVidensk. Selsk. Naturvidensk.Math.Afh., Ser.5,2: 335. 1851.Bucephalon mexicanum(Liebmann)0. Kuntze, Rev. Gen. 2: 624. 1891.Type.Mexico.Veracruz: Tlapacoyo,Hda. de Joro, (? fl-fr),Liebmann14260 (holotype, C; isotypes, F, 0). Fig. 3 Trophischiapensis Brandegee, Univ. Calif. Publ. Bot. 6: 178. 1915; Burger, Ann. Missouri Bot. Gard. 49: 16, t. 3. 1962. Type. Mexico. Chiapas: Cerro Boquer6n, Sep 1913 (? fr), Purpus 7091 (holotype UC, not seen; isotypes, F, MO, US).

Trophis chorizantha Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 302. 1929. Type. Honduras. Atlantida:nr. Tela, 6 Dec 1927-20 Mar 1928 (? flfr), Standley 56771 (holotype, F; isotypes, G, US). Skutchia caudata Pax & K. Hoffmann ex C. Morton, J. Wash. Acad. Sci. 27: 302. 1929. Type. Costa Rica. San Jose: nr. El General, Jan 1936 (d), Skutch 2383 (holotype, US; isotypes, A, K, MICH, MO, NY, P). Trophismatudai Lundell, Lloydia 2: 81. 1939. Type. Mexico. Chiapas: Mt. Ovando, nr. Escuintla, Dec 1937 (? fl), Matuda 2091 (holotype, MICH; isotypes, A, F, K, LL, NY, U, US). Trophisnubium Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 17. 1940. Type. Guatemala. Quezaltenango: Volcan de Zunil, 3 Aug 1934 (Y fl-fr), Skutch 925 (holotype, F; isotypes, MICH, NY, US).

Shrub or tree, to 20 m tall. Leafy twigs 1-2.5(-3) mm thick,puberulousto hirtellousto almostglabrous. Laminaelliptic to lanceolate(to almost linear),(3-)520 x (0.5-)1-7.5 cm, broadestat, above, or below the middle, almost equilateral,chartaceous to subcoriaceous; apex (caudate-)acuminate;base obtuse to acute (to rounded);marginentireor (serrate-)dentate;upper surfacesmoothand glabrousor sometimesa few hairs at the base of the midrib; lower surface glabrous or sparselypuberulouson themainveins;venationslightly

TAXONOMICTREATMENT

35

jI&~~~I

Ei.~~s

Fig. 3. Trophismexicana. 1. Leafy twig with young infructescences(Rosas R. 1213). 2. Leafy twig with staminateinflorescences(RosasR. 1016). 3. Leafy twig with staminateinflorescences.4. Leafy twig with young staminateinflorescences (Contreras6624). 5. Fruitingperianth(Burgeret al. 3367). 6. Staminateflower. 6a. Pistillode (Rosas R. 1016). 7. Embryo. 7a. Seed (Burger 3367). 8. Pistillate flower (Rosas R. 1213). [By P.

Pardoen,Utrecht.]

36

prominentto almostplaneabove,slightlyprominent beneath;lateralveins4-1 1pairs;tertiaryvenationreticulate;petiole0.3-1.5 cm long,0.5-1.5 mmthick, orglabrous;stipulesovateto triminutelypuberulous to subglabrous, angular,0.1-0.4 cmlong,puberulous

FLORANEOTROPICA

Mar 1965 (? fr), McVaugh 23393 (MICH); Sierra Madre Occidental, between La Quadrilla & San Sebastian, 1500 m, 16 Feb 1927 (e), Mexia 1701 (A, BM, F, GH, MICH, MO, NA, NY, US). OAXACA: Cafetal Nuevo Esperanza, 9 Apr 1917 (? fl-fr), Conzatti et al. 3064 (US); Dtto. Tuxtepec, nr. Chiltepec, Jul 1940-Feb 1941 (o), Marsubpersistentor caducous. Staminate inflorescences tinez-Calder6n 568 (LL, US); Dtto. Pochutla, nr. Cafetal racemose tospicate;peduncle0.2-1.5cmlong,sparsely Concordia, 1-15 Apr 1933 (? fr), Morton et al. 2478 to denselypuberulousto tomentellousto short-velu- (A, F, K, NY, US); Cafetal Calvario, Cerro Espino, 20 tinous;spikeorraceme2-12 cmlong;flowersdistant Aug 1917 (6), Reko 3519 (US); 39 km S of Valle tocrowded,sessileorpedicellate; pedicelto 3 mmlong; Nacional, 1650 m, 23 Feb 1976 (6'), Rzedowski 33938 perianthca. 2 mmlong,4-fidto 4-parted,sparselyto (NY). PUEBLA: Ajotoxco, 350 m, 29 Apr 1970 (? fr), ratherdenselypuberulous to tomentellous the VenturaA. 999 (F, MICH, NY); Mun. Atoluca, Agua de (towards apex);filaments2.5-3.5 mmlong;anthersca. 1 x 1 Obispo, 1400 m, 30 Mar 1973 (6'), Ventura A. 8100 mm;pistillodequadrangular, ca.0.5 mmlong,densely (MICH, US). TABASCO: Mun. Tacotalpa, rd. TacotalpaPaijulapa, 13 Apr 1980 (? fr), Cowan 2907 (F, MO, NY, white-tomentellous. Pistillate inflorescences mostly TEX, WIS); Mun. Huimanguillo, 8 km S of San Fransolitary,racemoseto spicate,0.2-2 cmlong,including cisco Rueda, 10 Mar 1983 (6'), Fernaindez N. 1418 (MO, thesparselyto denselypuberulous to tomentellous to NY, US); Tamulte, 13 Jan 1889 (o), Rovirosa 339 (K); short-velutinous peduncle;racemeor spike1-14 cm Mun. Huimanguillo, El Rosario, 19 Feb 1984 (e), long;flowers(2-)10-25(-ca. 35), spaced,pedicellate VenturaA. 20964 (MO, NY); Mun. Teapa, El Madrigal, (orsubsessile);pedicelto 4.5 mmlong,to 1.5cmlong 20 Feb 1984 (d6), Ventura A. 20974 (MO, NY). infruit;perianth ca.2 mmlong,4-lobed,rather sparsely VERACRUZ: Estaci6n Biol6gica Los Tuxtlas, 18 Apr puberulous to tomentellous; stigmas2-4.5 mmlong, 1972 (? fr), Calzada 763 (F, U), 22 May 1981 (? fr), sparselyto denselypuberulous. Fruitingperianth glo- Gentry et al. 32361 (BG, MO, U); Mun. Juchique Ferrer, x bose, 0.7-1 0.7-1 cm, red (to purple),apiculate, La Cima, 1700 m, 21 Jun 1972 (v), Hernandez M et al. mostly longitudinallyribbedand/or? verrucate, 1573 (F); Tlapacoyo, Hda. de Joro, Mar (? fl-fr), Liebmann s.n. (K, 0, P, U); 4 km NE of Jalapa, 1350 m, to tomentellous. sparselypuberulous 4 Mar 1973 (6), Marquez R. et al. 160 (F); Palo Dulce, Distribution(seeFig.72.3).FromcentralMexico 1 Mar 1930 (e'), Mell 685 (F, NY, US); 10 km N of to Panama,and disjunctlyin southemEcuador;in wet Sontecomapan, 21 Apr 1983 (? fr), Nee 26706 (LL, forestor less commonlyin deciduousforest;to 2200 m, NY); Mun. Hidalgotitlan, 2-3 km SE of Agustin Melgar, 5 Mar 1985 (? fl), Nee et al. 29955 (BG); Cerro de butusually600-1700 m, butin Belize andadjacentparts Chicahuaxtle, Cuauhtlapan, 1600 m, 22 Jan 1968 (6'), of Guatemalaand Hondurasonly below 600 m, andin Rosas R. 1016 (A, BM, C, F, U); Estaci6n Biol6gica Panamaand Costa Rica sometimes below 600 m. UNAM Montepio, 190 m, 20 Apr 1968 (? fl-fr), Rosas Representative specimens examined. MEXICO. R. 1213 (A, BR, U); Mun. Yecuautla, El Haya, 1300 m, CHIAPAS:Mun.La Trinitaria, LagoTsiskaw,30 mi E of 12 Feb 1971 (e'), VenturaA. 3102 (F, MICH, TEX). GUATEMALA. ALTA VERAPAZ:Sebol, 14 Apr 1964 La Trinitaria,ca. 1500 m, 14 Apr 1965 (? fr), Breedlove 9766 (CAS,F, LL,MICH);Mun.VillaCorzo,CerroTres (2 fr), Contreras 4282 & 4303 (F, LL, NY); rd. SebolCoban, Chapultepec farm, 26 May 1969 (? fl), Contreras Picos, nr. Cerro Bola, 1500 m, 9 Feb 1972 (? fr), Breedlove24074 (CAS, F, MICH,MO,NY); Mun.An- 4812 (EAP, F, LL); Rio Coban, 1200 m, 13 May 1963 gel AlbinoCorzo,FincaTuxtepec,1380m, 14 Dec 1980 (e), Molina R. et al. 12185 (EAP, F, NY); mouth of Rio (? fl), Breedlove48708 (CAS, LL, MO); Mun. Ray6n, Frio, between Tactic & Santa Cruz, 1460 m, 14 May 1963 (6'), Molina R. et al. 12267 (EAP, F, NY, US); 10 km above Ray6n Mezcalapa, 1700 m, 19 Sep 1981 (? fr), Breedlove 52681 (CAS, LL, MO); Mt. Ovando, PansamalA,ca. 1250 m, May 1887 (ci), Tuerckheim1231 (F, GH, K, MO, NY, P, US). CHIMALTENANGO: Slopes nr. Esquintla, Matuda 576 (A, LL, NY), Feb 1939 (V), Matuda 2648 (LL, NY, WIS), ca. 1500 m, 14-18 Apr of Volcan Fuego, Finca Montevideo, Rio Pantale6n, 1200-1600 m, 20 Sep 1942 (st), Steyermark 52069 1939 (? fr), Matuda 3933 (A, GH, LL, MICH, MO, NY, (A, F). ESCUINTLA: Rio Guacalate, 16 Dec 1938 (v), US), ca. 1000 m, 14-18 Nov 1939 (a'), Matuda 3978 (A, F, MICH, MO, NA, NY), ca. 1600 m (ae), Matuda Standley 60197 (A, F, MICH, NY); below Las Lajas, 4241 (A, F, LL, MO, NY, U, US); 65 km ESE of Comitan, 900-1200 m, 8-9 Feb 1939 (st), Standley 64766, 64995, Lago de Tepancuapan, 1 Jan 1953 (e), Miranda7650 65815 (F); nr. Rio Michatoya, SE of Escuintla, 250300 m, 12 Mar 1941 (st), Standley 89114 (A); Rio (US). GUERRERO:Dtto. Montes de Oca, San AntonioBuenos Aires, 10 Jun 1937 (e), Hinton10298 (GH, K, Burri6n, NE of Escuintla, 720 m, 16 Mar 1941 (st), LL, MICH, US); Mun. General Heliodoro Castillo, rd. Standley 89574 (F). IZABAL: El Estor, 24 Mar 1972 (6), El Paraiso-Puerto del Gallo, km 35, 2000-2100 m, 9 Contreras 11526 (LL, MO, US); Rio Juyama, SE of Jan 1985 (a'), Thomaset al. 3752 (NY). JALISCO:10 Cheyenne, 15 mi S of Bananera, 8 Apr 1940 (? fl-fr), km N of La Cuesta, below pass to Talpa de Allende, 31 Steyermark 39157 (F); 6 mi up Rio Dulce, Livingston,

TAXONOMIC TREATMENT

37

39451 (F,NY); Rio Trio, (MO);Schipp1261 (A, BM, F, G, GH, K, MICH,MO, 14 Apr 1940(9 fr),Steyermark 17 Dec 1941 (st), Steyermark39932 (F, US); between NY, Z); TemashRiver,ca. 50 m, 27 Feb 1935 (2 fl-fr), El Charro & El Benque, 18 Mar 1972 (e), Tun Ortiz Schipp1334 (A, BM, F, G, GH, K, MICH,MO,NY, Z). 2539 (BM, EAP, F, MICH, MO, US). PETkN: Dolores,

HONDURAS. ATLANTIDA:Montaina Lancetilla, 3

24 Apr 1961 (v), Contreras2172 (F, LL, MICH,NY);

km S of Lancetilla, 19 Feb 1962 (? fl-fr), Molina R.

Lacand6n, 7 Feb 1962 (de), Contreras 3347 (EAP, F, G,

10476 (EAP, F). COMAYAGUA: Montafia La Choca, Que-

GH, MICH,NY, US), 16 Mar 1962 (? fr), Contreras bradaEl Zope, 1500 m, 15 Apr 1957 (2 fl), MolinaR. 3535 (F, LL, MICH,NY); La Cumbre,I Mar 1967 (d), 8188 (EAP,F, NY); summitof rangeabove El Achote, Contreras6624 (LL,NY, U); San Pedro,Cadenasrd., 9 1800 m, 28 Jul 1936 (9 fr), Yunckeret al. 6171 (F, K, CerroAzul NationalPark,1540 m, 11 Feb Jan 1970 (? fl), Contreras 9483 (NY, LL, U); La NY). COPAN: Cumbre,2 Nov 1971 (v), Contreras10476 (LL, NY, 1992 (e), H. Thomas et al. 250 (EAP). EL PARAIso:Nr. U); LagunaPetexbatun,Rio Pasi6n, 1 Apr 1964 (? fl- Yuscaran,CerroPiedrasAmarillas,ca. 2000 m, 22-25 fr), Lundell18189 (EAP,F, GH, LL, MICH,MO, US); Mar 1975 (c), Nelson et al. 2520 (MO). INTIBUCA: between FincaYalpemech& Chinaja,28 Mar 1942 (? Quebrada del Pel6nde Guise,1600m, 9 Apr 1956(? fl), fl-fr), Steyermark45445 (LL, NY); Machaguila, Dolores, MolinaR. 6379 (EAP,F); Barrana Yashse, Yamaranguila, Rio Puxte, 19 Feb 1971 (? fl), Tun Ortiz 1630 (BM, 1500 m, 12 Apr 1956 (? fl), Molina R. 6529 (EAP,F, Hwy. Cito, nr. km GH, US). LA PAZ:CordilleraGuaquiro,nr. Sabanetas, EAP, F, NY, US). QUEZALTENANGO: 199, S of Quezaltenango,1200-1300 m, 22 Jan 1987 2100 m, 21 May 1964 (? fl-fr), Molina R. et al. 13875 Montafiade Celaque, 18 (o'), Croat et al. 63465 (BG, MO); Volcan Atitlan, ca. (EAP,F, NY, US). LEMPIRA: Nr. 2400 m, 24 Oct 1934 (?), Skutch 1511 (A, BM, F, NY, Nov 1974 (v), Hazlett2356 (CR, MO). MORAZAN: US); Colombo, 21 Dec 1934 (c?), Skutch 1986 (A, BM, Zamorano,Mt. Uyuca, 2000 m, 21 Feb 1952 (? fl), F, G, NY, US); Rio Samala,nr. SantaMariade Jesuis, Carlson2445 (F, MICH);CerroUyuca,2000 m, 29 May 1948(? fl), MolinaR. 909 (BM, F, GH,MO,US), 4 Sep 1500-1600 m, 25 Jan 1941 (? fr), Standley 84572 (EAP,F); S of San Martin,rd. FincaPirineos-Patzulin, 1948 (2 fl-fr),MolinaR. 1108a (BM, F, GH,MO, US); 1200-1400 m, 9 Feb 1941 (st), Standley 87012 (F); RioYeguare1800m, 4 Nov 1948 (? fr),MolinaR. 1443 VolcanSantaMaria,between SantaMariade Jesuis& (EAP, F); Cerro Uyuca, between La Labranza & Calahuache, 1300-1500 m, 5-6 Jan 1940 (st), QuebradaEl Granadillo,1800 m, 12 Dec 1948 (? fr), Steyermark 33499 & 33636 (F); Volcan Santa Clara, 1250-1650 m, 23 May 1942 (st), Steyermark46730 (F).

L. 0. Williamset al. 14815 (BM, EAP, F, GH), ca. 1900

Rio Mopa, nr. Rodeo, 14 Mar 1939 (st), Standley 68956 (F); Volcan Tujamuleo, Finca El Porvenir, Rio Cahuo, 1000-1500 m, 9-12 Mar 1940 (d), Steyermark 37426 (F) & 37576 (F, US). SANTAROSA:Cerro Redondo, 1200 m, Oct 1893 (e), E. J. Heyde et al. 6239 (A, GH, K, NY, US). SOLOLA:VolcAnAtitlan, ca. 1700-2000 m, 11 Jun 1942 (e or st), Steyermark 47333 (F), 47349 (F, NY), Volcan Atitlan, ca. 47392 (F, US). SUCHITEPkQUEZ: 2400 m, 24 Oct 1934 (? fl), Skutch 1511 (BM, NY), Finca Moca, 9 Jan 1935 (? fl-fr), Skutch 2094 (A, BM, F, NY, US); slopes of VolcAn de Zunil, between Finca Alvidas & Finca Ponchas, 2 Feb 1940 (st), Steyermark 35461 (F); Volcan Santa Clara, Finca El Naranjo, 23 May 1942 (ci, ?, or st), Steyermark 46630 & 46658 (A, F), 46687 & 46730 (F, G). BELIZE. Toledo District, Maya Mountains, 11 Mar 1987 (? fl), Davidse et al. 32292 (BG, LL, MO, NY); El Cayo District, Grano de Oro, 2 Jun 1973 (? fl-fr), Dwyer 10841 (MO, U); Stann Creek District, Stann Creek valley, Bocaurina Hill, 12 Feb 1940 (e), Gentle 3211 (A, K, LL, MICH, MO, NA, NY); Toledo District, nr. Rio Blanco, Edwards rd. beyond Columba, 15 Mar 1951 ($ fl-fr), Gentle 7234 (EAP, F, G, GH, LL, MICH, NY, US); El Cayo District, Valentin, 6 Jul 1936 (st), Lundell 6422 (F, MICH, NY); El Cayo District, Rio Frio, nr. Agustin, 27 Mar 1954 (? fl), Molina R. 238 (EAP, F); Pueblo Viejo, 9 Feb 1934 (? fl); Toledo District, between Jimmy Cut & Cabro, 9 May 1976 (Q fr), Proctor 36165

La Laguna, San Juancito, ca. 393 (EAP). TEGUCIGALPA: 1900 m, 4 Apr 1932 (e), Edwards P-43 (F). YoRo: Las

m, 20 Sep 1949 (? fr),L. 0. Williams16850 (A, BM, F, SANMARCOS: BarrancoEminencia,above San Rafael GH, MO, US). OLANCHO: AgaltaNationalPark,17 km Pie de la Cuesta, 15 Mar 1939 (st), Standley 68663 (F); NNW of Catacamas,30 May 1992 (9 fl-fr),H. Thomas

Rosas, 50 m, 13 Mar 1965 (? fl), Dickson 1422 (EAP, US); nr. Yorito, 1700-1900 m, 25 Jul 1979 (? fr), Hazlett 3156 (F). EL SALVADOR. Colina de Santa Tecla, Jul 1923 (oi), San Benito, Calder6n 1749 (GH, NY, US). AHUACHAPAN: 8 Feb 1990 (2 fl-fr), Sermefio s.n. (MO). SONSONATE: Sierra de Apaneca, Cerro El Pil6n, 23 Feb 1968 (? fl), Molina R. et al. 21613 & 21756 (EAP, F, NY); Sierra de Apaneca, Finca Colima, 17-19 Jan 1922 (9 fl-fr), Standley 20183 (GH, NY, US).

CerroAlegre, San Jose de NICARAGUA.BOACA: los Remates, 1100-1180 m, 11 Feb 1983 (oi), Moreno 20268 (MO). ESTELi: 8 km NW of Esteli, 1500-1600

m, 19 Oct 1979(? fl-fr),Grijalvaet al. 605 (MO);Cerro Quiabui,1600 m, 25 Dec 1982 (ci), Moreno 19259 (MO),

13 Aug 1976 (? fr), Neill 7759 (MO), 27 Apr 1980 (Y fl-fr), Stevens et al. 16912 (MO). GRANADA:Volcan Mombacho, 16 Jul 1975 (? fl-fr), Atwood at al. AN.185 (NY); Volcin Mombacho, 9 Jun 1981 (? fr), Moreno et

al. 9146 (MO);Plan de las Flores, N of VolcanMombacho, 1100-1200 m, 31 Mar 1982 (? fl-fr) Moreno 16023 (MO). JINOTEGA:Halfway between Matagalpa & Jinotega, 1400 m, 8 Aug 1977 (? fr), Croat 43089

(MO); rd. Matagalpa-Jinotega, 1450-1500 m, 2 Jul 1980 (9 fl-fr), Moreno 1115 (MO); NE of Cerro Kilambe,"Filasel Portal,"26 Mar 1981 (? fl), Moreno

38

FLORA NEOTROPICA

et al. 7584 (MO);FincaAventina,E of Jinotega,23 Jun MO, NY, TEX). CHIRIQUi: Nr. Fortuna Dam Camp, 1947 (st), Standley9965 (EAP,F); CordilleraCentral, 1200 m, 26 Feb 1985 (2 fl-fr), Hampshire et al. 124 (F, PMA). COCLt: Cerro Pilon, above El Valle, 9 Jan 1972 Ocotillo, nr. SantaLastenia,1550 m, 17 Jan 1965 (a), L. 0. Williamset al. 27825 (F, GH, NY, US). MADRIz:Cerro

Volcande Somoto, 1400-1600 m, 25 Sep 1980 (? fr), Moreno 2868 (MO), 3 Feb 1983 (a'), Moreno 20074 (MO). MATAGALPA:NW slopeof CerroEl Picacho,1420-

(? fl), Gentry et al. 3635 (NY); Boca del Toabre, at confluent of Rio Toabre & Rio Cocle del Norte, (? fl-

fr), Lewiset al. 5568 (CR). VERAGUAS: CerroTute,ca. 10 km NW of Santa Fe, 750-1000 m, 18 May 1975 (d'),

1520m, 25 May1983(? fr),Stevens22159(MO);Macizos Mori 6247 (MO, U), 2 Aug 1975 (? fl-fr), Mori et al. de PefiasBlancas,WNWof Hda.SanMartin,1400-1600 7542 (MO, NY, PMA). ECUADOR. ZAMORA-CHINCHIPE: Rd. Lojam, 20 Jan 1982 (a), Stevens et al. 21174 (MO); Santa Mariade Ostuma,betweenMatagalpa& Jinotega,1300- Zamora, km 45-51, 1400-1600 m, 20 Nov 1966 (? fl1500 m, 8-15 Jan 1963 (e'), L. 0. Williams et al. 23350

fr), Dodson 1423 (BM, F, GH, LL, MO, NY, US).

(EAP,F, GH, LL, NY, US); 6-10 km NE of Matagalpa, The species is variable in the leaf shape, leaf marrd. to El Tuma,1000 m, 14-16 Jan 1963 (e), L. 0. Wilgin, inflorescences,andflowers.The differencesin the liams et al. 23839 (EAP,F, G, LL, NY, US); Disparate herbarium materialat present available do not justify de Potter,SantaMariade Ostuma,1600 m, 15 Jan 1964 (? fl-fr), L. 0. Williams et al. 27620 (BM, F, GH, NY, the separationof Trophismexicana and T chiapensis, US); CerroEl Picacho, Finca SantaMariade Ostuma, as proposedby Burger (1962). 1500 m, 11 Feb 1965 (e), L. 0. Williamset al. 29170 (F, In CentralAmerica Trophismexicana can usually GH, NY, US). ZELAYA: CerroEl Hormiguero,18 Apr be distinguishedfrom T racemosa by the smooth and 1979 (9 fl-fr), Grijalva441 (MO);CerroSaslaya,1100 often almost glabrous leaf surface, or elsewhere and m, 3 May 1978 (? fl-fr),Neill 3823 (BG,MO);CerroLa otherwiseby theusuallydentateleafmargin,by themore Pimienta,900-1180 m, 16 Mar 1980 (? fl), Pipoly 6051 (MO);Caiio El Hormiguero,17 Mar 1980 (a), Pipoly or less distantandoften subsessileto pedicellatepistil6126 (MO);betweenCerroEl Inocente& CerroSaslaya, late flowers, the often sparseindumentumin the inflonr. sourceof CanloMajagua,1050-1150 m, 8 Mar 1978 rescences, and by the usually rugose or more or less clearlyribbedfruitingperianths. (d'), Stevens 6722 (BG, MO). COSTA RICA. ALAJUELA: Reserva Biol6gica The disjunctoccurrenceof the species in southemMonteverde, Rio Pefias Blancas, 24 Feb 1989 (a), Bello most Ecuadoris noteworthy,and it resembles the dis731 (BG,MO), 10 Mar1989(? fl), Bello 758 (BG,MO); junction in the range of the montane Cecropia ReservaForestalde SanRam6n,900-1200m, 9 Apr.1987 polyphlebia, in South America only known from the 11480(F);ReservaForestalde San (? fr), G6mez-Laurito Ram6n,Rio San Lorenzito,25 Feb 1987 (? fl), Herrera areain which Trophismexicanahas been collected. Local names. Mexico:Oaxaca:huanchal;Veracruz: Ch. et al. 482 (BG);N slopes of VolcanArenal,20 Apr 1973(v), Lentet al. 3367 (CR,F,MO,NY,U, US);Cant6n chischillo, ramoncillo. Guatemala: Alta Verapaz: AlfaroRuiz,Zarcero,26 Jan 1939 (e), A. Smith 1558 (F, chanaque,yax-ox. MO, NY). ALAJUELA/GUANACASTE: Slopes of Volcan Miravalles,rd.N of Bagaces,2 Mar1985(? fl), Tayloret al. 4656 (MO).GUANACASTE: Volcin Orosi,ca. 15 km 2. Trophis cuspidata Lundell, Amer. Midl. Nat. 19: SE of La Cruz,18 Apr 1985 (? fl-fr),Schatzet al. 1100 427. 1938; Lundell, Lloydia 2: 80. 1939; Burger, (MO); El Arenal, 18-19 Jan 1926 (v), Standleyet al. Ann. Missouri Bot. Gard.49: 18, t. 4. 1962. Type. 45271 (US).HEREDIA: Rio Sarapiqui, upriverfrombridge Mexico. Chiapas:Mt. Ovando, 23 Dec 1936 (o), on rd. to La Virgendel Socorro,8 May 1982 (2 fl-fr), Matuda 1051(holotype, MICH;isotypes, MO, U). Hammel12113 (EAP; MO); Colonia de la Virgendel Fig. 4 Socorro,PuertoViejo, 13 May 1975 (? fr), Poveda987 (CR, F). LIMON: Cant6nLim6n, El Progreso, 11 Apr Shrubor tree, to 15 m tall. Leafy twigs 1.5-3 mm 1984 (? fl), Herrera2588 (BG, F, MO);CantonLim6n, brownish to white-puberulous to hirtellous. thick, CerroMuchilla,Fila Matama,13 Apr 1989 ($ fr), RobLaminaelliptic to lanceolate,4.5-25(-30) x 2.5-7.5(les 2767 (BG, MO).PUNTARENAS:E of Las Cruces,5-6 11) cm, broadestatthemiddle,almostequilateral,chartkm S of San Vito, 15-16 Jan 1967 (a'), Burger et al. 4429 (BM,CR, EAP,F, MO,NY, PMA);Monteverde,1300m, aceousto subcoriaceous(to coriaceous);apex(caudate15 Dec 1979 (? fl), Haber 364 (MO), (o) Haber 365 )acuminate;base acuteto obtuse (to rounded);margin (MO), 11-14 Jun 1985 (? fl-fr), Hammelet al. 13821 (NY, entireto denticulate;uppersurfaceglabrous;lowersurTEX). SAN Jost: Bajo de la Honduras, 21 May 1974 face ratherdensely (to rathersparsely) hirtellous (to (st), Poveda 856 (CR); Parque Nacional Braulio Carrilo, on the (main)veins; venationalmostplane puberulous) 2 Jun 1984 ($ fr), Sanchez et al. 513 (F, MO); nr. El General, Jan 1936 (a'), Skutch 2386 (A, GH, K, MICH, but the midribslightly impressedabove, (very) prominentbeneath;lateralveins 6-13(-15) pairs;tertiaryveMO, P, US). nationreticulate(or in largeleaves partlyscalariform); PANAMA. BOCAS DEL TORO: Rd. to Chiriqui Grande, 30 Apr 1986 (q fl), McPherson 9147 (BG, F, petiole 0.7-2(-2.5) cm long, 1-2.5 mm thick, densely

TAXONOMICTREATMENT

39

Fig. 4. Trophiscuspidata. 1. Leafy twig with staminateinflorescences.2. Leafy twig with pistillate inflorescences (Tuerckheim1.2169). 3. Leafy twig with infructescences(Lundell 19647). 4. Pistillate flower (Matuda 2370). 5. Fruitingperianth(Dwyer 14463). [By P. Pardoen,Utrecht.]

yellowish to white-puberulousto hirtellous; stipules ovate to lanceolate, 2.5-5 mm long, puberulous, subpersistentStaminateinflorescencesspicate;peduncle 0.2-0.5 cm long,denselywhitetoyellowish-puberulous; spike 2-4 cm long, rachisdensely white to yellowishpuberulous to short-velutinous;flowers ? crowded, (sub)sessile;perianth2-2.5 mm long, 4-parted,densely white (to yellowish)-puberulousto tomentellous;filamentsca. 4-4.5 mm long; anthers0.8-1.2 x 0.8-1.2 cm; pistillode quadrangular,ca. 0.4 mm long. Pistillate inflorescencesracemose(to almostspicate);peduncle0.32.5 cm long, white to brownish-puberulousto shortvelutinous; raceme 0.5-7(-12) cm long; flowers 3-10(-12), ? spaced,pedicellate(to subsessile),pedicel to 3(-4) mm long, to 1.5 cm long in fruit;perianth2-2.5 mm long, 4-lobed, densely white to brownish-puberulous to short-velutinous;stigmas2-4 mm long, densely puberulous.Fruitingperianthsubglobose, ca. 0.8-0.9 x 0.6-0.7 cm, densely white to yellowish-puberulous.

Distribution(see Fig. 73.1). In southernMexico and Guatemala;in cloud forest;at 1650 m to ca. 2600 m. Specimens

examined.

MEXICO.

CHIAPAS: Mun. La

Independencia, rd. Las Margaritas-Campo Alegre, 2300 m, 18 Feb 1973 (d'), Breedlove 33660 (CAS, LL, MO); Mun. Villa Corzo, Cerro Tres Picos, 2100-2500 m, 23 Mar 1973 (di), Breedlove 34379 (LL, MO); Mun. Union Juarez, Volcan Tacana, above Talquian, 2200 m, 13 Dec 1976 (V fr), Breedlove 42530 (MO); Mun. Siltepec, above Siltepec, 2000-2400 m, 1 Feb 1982 (d), Breedlove et aL 58326 (CAS);Mun. Jaltenango, Reserva El Triunfo, 1900 m, 20 Apr 1989 (d), Heath et al. 79 (CAS); Mt. Tacani, 2000 m, Aug 1938 (? fl-fr), Matuda 2370 (A, EAP, F, K, LL, MICH, NA, NY, U); Cascada, nr. Siltepec, 1600 m, 11 Mar 1945 (d), Matuda 5069 (CAS, EAP, LL); Pinabeto, Motozintla, 2586 m, 9 May 1945 (? fl-fr), Matuda (1)5478 (F); Finca Prusia, rd. to Mapastepec, nr. Triunfo (S. Jultenango), 24 Feb 1951 (d'), Miranda 7028 (CAS). GUATEMALA. ALTAVERAPAZ: Coban, 1350 ni, Mar

FLORANEOTROPICA

40 II-2167 (GH, NY, US); between 1908 (e), Tuerckheim Tactic& Coban, 1700 m, Mar 1908 (? fl), Tuerckheim II-2169 (GH,NY, US). BAJAVERAPAZ:Union Barrios, 10 Aug 1975 (e'), Lundellet al. 19601 (LL, MO,NY), 15 Aug 1975 (? fl-fr), Lundellet al. 19647 (LL, MO, NY); Niiio Perdido,6 km on San Jose Espinerord., 23 Aug 1975 (2 fl-fr), Lundellet al. 19722 (MO,NY), 23 May 1977 (2 fl), Lundellet al. 20956 (LL, MO). EL PROGRESO: VolcanSantaLuisa,betweenFincaPiamonte & summit,above 2400 m, 5 Feb 1942 (st), Steyermark 43588 (F, US); between Finca Piamonte& summitof MontaniaPiamonte, 2500-3000 m, 7 Feb 1942 (6'), El Steyermark43664 (EAP, F, G). QUEZALTENANGO: Pocito, S of San MartinChili Verde,rd. to Colomba, 2200 m, Feb 1941 (st), Standley 84999 (MO); above Mujulia,between San MartinChili Verde& Colomba, 1800 m, Feb 1941 (st), Standley 85623 (F). SAN MARcos:Rafaelde Cuesta,FincaArmenia,ca. 1650m, 6-7 Jul 1977 (? fr), Dwyer 14463 (BG, MO).

Sorocea colombiana Standley, Trop. Woods 19: 39. 1929. Type. Colombia. Bolivar: Nr. San Martin de Loba, Apr-May 1916 (? fr), Curran 186 (holotype, F; isotype, NY, US). Pseudolmedia karstenii Pittier, Bol. Soc. Venez. Ci. Nat. 9: 119. 1944. Type. Venezuela. Carabobo: San Esteban, nr. Puerto Cabello, (e' + V fl), Karsten s.n. (syntypes, VEN). Shnrb or tree, to 20 m tall. Leafy twigs 1-4 nmmthick, puberulous. Lamina elliptic to lanceolate, (3-)5-20 x (1.5-)2-9 cm, mostly broadest above the middle, (almost) equilateral, chartaceous to subcoriaceous; apex acuminate; base acute to obtuse to rounded to emarginate, occasionally attenuate; margin entire or dentate, sometimes sinuate, occasionally lobate; upper surface scabrous to scabridulous or smooth, sparsely puberulous on (the lower part of) the midrib; lower surface sparsely puberulous on the main veins, sometimes scabridulous; venation slightly prominent to almost plane above, prominent beneath; lateral veins 4-9 pairs; tertiary venation reticulate; petiole 0.3-1(-1.5) cm long, 1-2 mm thick, puberulous; stipules ovate to triangular, 0.1-0.3 cm long, sparsely puberulous to glabrous,

This species is not very distinctfrom Trophismexicana, butthe presenceof indumentumat the lower surface of the lamina, the tertiaryvenation being more prominentbeneaththan in T mexicana, and its occurrence between 1700 and 2600 m, at higher elevations than normalfor T mexicana, appearto justify distin- subpersistentor caducous. Staminate inflorescences guishingseparatetaxa,althoughit is not certainwhether spicate, 0.5-10 cm long, including the 0. 1-1 cm long, minutely puberulous to short-velutinous peduncle; it should be at the species level. flowers ? crowded, sessile; perianth ca. 2 mm long, 4parted, short-velutinous to puberulous especially along 3. Trophis racemosa (Linnaeus)Urban,Symb.Antill. the margins; filaments ca. 2.5 mm long; anthers 0.84: 195. 1905; Burger,Ann. MissouriBot. Gard.49: 1.2 x 0.8-1.2 mm; pistillode quadrangular, 2-6 mm 7, t. 1. 1962. BucephalonracemosumLinnaeus,Sp. long, puberulous. Pistillate inflorescences spicate, 0.1P1. 190. 1753. Type. LINN 165.1. Fig. 5 1 cm long, including the short-velutinous to puberuTrophisamericanaLinnaeus,Syst. Nat. ed. 10. 1289. lous peduncle; spike 0.5-3.5 cm long; flowers (l-)31759.Type.P. Browne,Civ.Nat. Hist.Jamaica,t. 8(-10), usually crowded, sessile; perianth ca. 2 mm 37, fig. 1. 1756. long, 4-lobed, short-velutinous; stigmas (2-)4-5 mm TrophisramonSchlechtendal& Chamisso,Linnaea long, densely puberulous. Fruitingperianth 0.8-1.2 x

6: 357. 1831. TrophisamericanaLinnaeusvar. ramon(Schlechtendal & Chamisso)Bureau,in de Candolle,Prodr.17:253. 1873;Engler,Bot. Jahrb. Syst. 40: 543. 1908. Trophisracemosa(Linnaeus) Urbansubsp.ramon(Schlechtendal & Chamisso) W. Burger,Ann.MissouriBot. Gard.49: 10. 1962. Type. Mexico. Veracruz:Misantla,Mar 1929 (? fl-fr),Schiede& Deppes.n. (or 1118?)(holotype, B; isotypes,BM, MO). TrophisamericanaLinnaeusvar. meridionalisBureau, in De Candolle, Prodr. 17: 252. 1873. Trophis racemosa (Linnaeus) Urban subsp. meridionalis(Bureau)W. Burger,Ann. Missouri Bot Gard. 49: 8. 1962. Type. Colombia. Rio Magdalena,without precise locality, Bonpland 1512 (holotype, P; type fragment,F). Sahagunia urophylla Donnell Smith, Bot. Gaz. 40: 11. 1905. Clarisiaurophylla (Crawfordsville) (Donnell Smith) Lanjouw,Recueil Trav. Bot. N6erl. 33: 270. 1936. Type. Honduras.Puerto Sierra, 16 Jan 1903 (? fr), P. Wilson54 (holotype, US; isotype, NY).

0.6-0.9 cm, red at maturity, the surface smooth or longitudinally ribbed, densely puberulous to shortvelutinous. Distribution (see Fig. 73.2). From central Mexico to the Greater Antilles and through Central America to northern Venezuela, westem Ecuador, and northern Amazonian Peru; in (margins of) wet evergreen forest or deciduous forest, in Peru (Loreto) also in seasonally inundated (tahuampa) forest; to 1700 m. Representative

specimens examined. MEXICO.

CAMPECHE: Campo Exp. Forest, rd. Escarcega-Cande-

laria, km 5, 14-30 Dec 1965 (? fl), Chavelaset al. ES69, ES-124, ES-208 (MICH); Escarcega, 10 Apr 1963 (? fr), Marroquin168 (WIS); nr. Xpujil, 18 Mar 1973 (st), Shepherd139 (F, MICH), 12 Aug 1974 (e), Shepherd191 (WIS). CHIAPAS:5 mi SW of Pinola, ca. 1250 m, 28 Jul 1964 (Q fl), Breedlove6591 (BM, F, MICH); Mun. Tenejapa, Rio Tanate, 1100 m, 4 Oct 1965 (a'), Breedlove12884 (F, MICH); ca. 7 km N of Ocosingo,

41

TAXONOMICTREATMENT

@

:o~~~~~1

lcm

0

%11mSmm

Fig. 5. Trophis racemosa. 1. Leafy twig with staminate inflorescences (Contreras 9052). 2. Leafy twig with staminate inflorescences (Mexia 6262). 3, 4. Leafy twig with pistillate inflorescences (Hartshorn 2107). 5. Leafy twig with infructescences (Schunke 9585). 6. Leaf (Hartshorn 2107). 7. Leaf (Contreras 9052). 8. Leaf (Schunke 9585). 9. Leaves (Proctor 20588). 10. Staminate flower (Cuatrecasas 16858). 11. Pistillate flower (Cuatrecasas 14227). 12. Fruiting perianth. 13. Endocarp body. 14. Seed. 15. Embryo (Molina R. 8101). [By P. Pardnoen,Ultrecrht.]

42 24 Sep 1972 (e), Breedlove 27926 (F, NY); Esquintla, Jul 1938 (? fl), Matuda 2598 (A, F, K, MICH, NA, NY), 16 Dec 1939 (? fr), Matuda 4022 (A, MICH, MO, NY); Santa Teresa, Acapetahuy, 14 Jun 1947 (o), Matuda 16751 (EAP, F, MICH); 27 km E of Villa Flores, Rancho Lindovista, 28 Jan 1950 (v), Miranda 5969 (U, US). COLIMA: Coahuayana, Nov 1906 (e), Emrick 133 (F). GUERRERO: Cant6n de la Mano Negra, 4-8 km N of Iguala, 1100-1 150 m, 15 Feb 1970 (? fl-fr), Anderson et al. 5781, 5782, 5791 (MICH), 15 Feb 1970 (od), Anderson et al. 5795 (MICH); Montes de Osa, San Antonio, 25 Oct 1937 (a"), Hinton 11540 (F, NY); Sierra Madre del Sur, N of Rio Balsas, Temisco, 18 Nov 1937 (ei), Mexia 8844 (B, F, G, GH, K, MO, NY, U, US); Cano6n de la Mano Negra, N of Iguala, 1100 m, 15 Feb 1970 (? resp. a"),Rzedowski 27080 & 27081 (F, MICH, MO, NY); Achotla, Sep 1926 (a), Reko 4998 (US). HIDALGO: Tutotepec, El Zopilote, 1 100 m, 18 Mar 1973 (? fr), Gimate L. 907 (EAP, WIS). JALISCO: La Manzanilla, ca. 5 km E of rd. Barra de Navidad-Puerto Vallarta, 17 Jan 1986 (? fl), Ayala 535 (MO, NY); Bahia de las Bauderas, ca. 14 km SW of Puerto Vallarta,29 Mar 1959 (? fr), Carter et al. 1210 (GH, MICH); 7.5 km N of Casimiro Castillo, 7 Jan 1985 (? fl), Judziewicz et al. 5138 (WIS); 15-18 mi SW of Autlan, 9 Apr 1951 (? fr), McVaugh 11949 (G, MICH); S of Puerto Vallarta, 26 Nov 1926 (a), Mexia 1142 (A, BM, F, G, GH, MICH, MO, NY). Mitmco: Dtto. Temascaltepec,18 Jan 1933 (a), Hinton 3165 (K, MICH, NY, US), 23 Mar 1933 (? fr), Hinton 3643 (F, K, MO, NY, US). MICHOACAN: Dtto. Coalcoman, San Pedro, 4 Jun 1941 (? fr), Hinton 15901 (BM, F, MICH, NY, U, US); Dtto. Coalcoman, nr.Aquila, Jan 1942 (? fl), Hinton 16291 (K, MICH, NY, US); Mun. Artcoga, 8-10 km E of La Mira, 25 Feb 1965 (? fl), McVaugh 22564 (MICH). NAYARIT: 12-14 mi S of Las Varas, 19-20 Apr 1960 (a"),McVaugh 19167 (G, MICH). OAXACA: Dtto. Tuxtepec, nr. Chiltepec, Jul 1940-Feb 1941 (a), Martinez-Calder6n207 (A, US), (Y fl), MartinezCalder6n 263 (A, US); Mun. Jaquila, nr. San Gabriel Mixtepec, 5 km S of San Gabriel, 12 Feb 1965 (? fl-fr), McVaugh 22409 (MICH); between Pinotepa & Jamiltepec, Feb 1923 (? fr), Nelson 2350 (NY, US); Pochutla, rd. to Oaxaca, 4 Apr 1968 (? fr), Pennington et al. 9500 (A, K, NY); Dtto. Pochutla, Cafetal San Carlos, 26 Jun 1982 (a"),Torreset al. 677 (MO). QUINTANA Roo: 10 km E of Dos Aguadas, NW of Tomas Garrido, 22 Dec 1982 (? fl-fr), Cabrera et al. 4148 (MO); rd. to Uni6n, 2 km N of Estero Franco, 23 Dec 1982 (Q fl-fr), Cabrera et al. 4194 (MO); Lake Chicnancanab,between Presumida & San Francisco, 1 Oct 1982 (? fl), Darwin 2383 (F): Mun. Chetumal, 6-7 km N of Tomas Garrido, 15 Mar 1990 (? fr), Sanders et al. 9982 (MO). SANLuIs POTOSI:Rio Salto, S of El Naranjo, ($ fr), Burger 148 & 150 (MO); Tamanzunchale,26 Jul 1937 (st), Edwards 667 (F); 17 mi W of N. Morelos, 9 Jan 1948 (a), Kenoyer et al. 3770 (A, MICH); S of N. Morelos, 9 Jan 1948 (a"), Kenoyer et al. 4067 (GH); Ladora Caliza, 10 Jan 1956 (? fl-fr), Rzedowski6894 (MICH).SINALOA:Nr. Culiacan, Cerro Colorado, 5 Nov 1904 (? fl + a), Brandegee s.n. (GH); Mesa Malqueson, Cerro Colorado, 7 Dec 1939

FLORA NEOTROPICA (R fl), H. S. Gentry 5147 (GH, MICH, MO, NA, NY); Sierra Tacuichamona, ca. 1000 m, 12 Feb 1940 (? fl), H. S. Gentry 5582 (MO, NY); Mun. Rosario, Cacalotan, Oct 1932 (e'), Gonzalez-Ortega 7100 (F, K, US); Quebradadel Naracal Balboa, San Ignacio, 31 May 1919 (? fr), Narvaez-Montes 853 (US). SONORA: Dtto. Alamos, Quiricoba, 12 Nov 1933 (? fl), H. S. Gentry 736M (MICH). TABASCO: Tenosique, nr. Zapote Bobal, 12 Mar 1976 ($ fr), Calzada et al. 2227 (U); Mun. Macuspana, Santa Ana, 28 Sep 1944 (v), Gilly et al. 355 (GH, MICH), (e'), Gilly et al. 361 (GH, MICH); Ejino El Colo Chero, 17 Nov 1975 (? fl), Menendez et al. 381 (MO); Mun. Tacotalpa, Tapijulapa, 30 Sep 1986 (e'), Hammel et al. 15535 (EAP, MO); Mun. Comalcalco, Reyes Hernandez, 10 Sep 1984 (e?), VenturaA. 21239 (NY). TAMAULIPAS: Tampico de Tamaulipas, 1827 (? fr), Berlandier 172 (P); Sierra Tamaulipas, ca. 40 km NNW of Aldama, 12 Oct 1957 (? fl), Dressler 2372 (GH, MICH, MO); nr. Rancho Gomes Farias, Apr 1960 (? fr), Duke 3536 (MO, U); rd. Julillo-Rio Sabanas, N of Mante, 4 Apr 1960 (? fr), Duke 3636 (MO); nr. Rancho de Barberena, 3 Aug 1939 (? fr), LeSueur 570 (F, GH, US); nr. Tampico, 27-30 Apr 1910 (? fr), Palmer 337 (F, GH, K, MO, US). VERACRUZ: Rd. Tempoal-Panuco, 10 km, 19 Mar 1971 (? fr), Chiang 406 (F, GH, MICH); Zacaupan, (? fr or st), Liebmann s.n. or 14268 (F, GH, K, LE, P, U); 1.5 km NW of Tampico Alto, 11 Jan 1984 (? fr), Nee et al. 28716 (F); Island of Juana Ramirez, 56 km S of Tampico, 8-9 Mar 1910 (st), Palmer 454 (GH, K, MO, NY, US); nr. Zacaupan, Nov 1906 (? fl), Purpus 2316 (F, GH, MO, NY, US), Aug 1905 (6'), Purpus 5988 (A, BM, F, GH, MO, US); Mirador, Mar 1932 (e'), Purpus 14273 (A, F); Mun. Nautla, La Martinica, 25 Mar 1971 (? fr), Ventura A. 3342 (F, MICH, NY). YUCATAN:NE of Tizimin, 8 Sep 1956 (e'), Enriquez 806 (US); Progreso, Jun 1934 (9 fr), Flores 10 (F). GUATEMALA. ALTAVERAPAZ:Nr. Pancajche,5 Apr 1939 (st), Standley 70793 (F); Cubilqiiitz, Aug 1903 (6'), Tuerckheim8581 (F, G, GH, K, NY, 0, US). CHIQUIMULA: Volcn Quezaltepeque, 3-4 mi from Quezaltepeque, 8 Nov 1939 (e), Steyermark 31511 (A, F). ESCUINTLA: 2 km E of Santa Lucia, 30 Jul 1970 (oe),Harmon 3382 (NY); SE of Escuintla, nr. Rio Michatoya, 12 Mar 1941 (st), Standley 89114 (F). HUEHUETENANGO: Between Finca San Rafael & Finca Providencia, 26 Jul 1942 (Q fl), Steyermark49549 (EAP, F); Rio Trapichillo,between Paso del Boquer6n below La Libertad & Democracia, 22 Aug 1942 (6'), Steyermark 51030 (F, G), 21 Aug 1942 (st), Steyermark51185 (A, F, G). IZABAL:Rio Sarstun,Cadenas, 3 Sep 1969, Contreras 9052 (U); Cadenas, Rio Gracias a Dios, 16 Sep 1969 (6'), Contreras 9186 (U, US), nr. Quirigua, 26 Apr 1939 (st), Standley 72483 (F). PETtN: Vaxactun, 29 Mar 1931 (st), Bartlett 12354 (F, MICH, NY, US); Tikal National Park, 31 Jan 1961 (? fr), Contreras 1902 (MICH, NY); Macanhe, 30 Jan 1966 (9 fl-fr), Contreras5443 (F, GH, U); Santa Teresa,Rio Subin, 14 Apr 1933 ($ fr), Lundell 2917 (A, BM, F, GH, K, MICH, U); Sayaxche, 8 Feb 1964 (R fr), Lundell 17928 (F, GH, MICH, NY); 35 km E of Santa Elena, El RemateTikal, 9 Nov 1965 (? fr), Molina R. 15413 (BM, EAP, F,

TAXONOMICTREATMENT MO); Tikal National Park, 11 Apr 1970 (q fr), Tun Ortiz 929 (BM, F, MO, NY). QUEZALTENANGO: Colomba, 2 Dec 1934 (a), Skutch 198b (NY). RETALHULEU: Nr. Nueva Linda, between Retalhuleu & Champerico,25 Feb 1941 (st), Standley 88452 (F); nr. Retalhuleu, 17 Feb-I Mar 1941 (st), Standley 88549 & 88729 (F). SANTA ROSA: Nr. Chiquimulilla, 29 Nov-8 Dec 1940 (? fr), Standley 79291 (F, NY); S of Guazacapan, De Avellana rd., 6 Dec 1940 (q fr), Standley 79513 & 79524 (F). SUCHITEPtQUEZ: Nr. Tiquisata, 17-19 Jun 1942 (st), Standley 47670 & 47774 (F, US). ZACAPA: Loma El Pichaco, above Santa Rosalia, 1200-1600 m, 15 Jan 1942 (st), Steyermark 42744 (A, F). BELIZE. Orange Walk District, Indian Church, 20 Jul 1976 (st), Arnason et al. 17068 (MO); Mun. Cayo, Ix Chel Farm, 15 Sep 1987 (? fr), Arvigo et al. 65 BG); Rio Sibun, Bright Lookout Bank, 4 Jan 1935 (q fl-fr), Gentle 1447 (A, K, LE, MICH, MO, NY, U, US); Stann Creek District, Stann Creek railway, 19 mi, 1 Mar 1939 (q fl-fr), Gentle 2704 (A, F, K, MICH, NA, NY); Toledo District, Jacinto Creek, 24 Jan 1945 (q fr), Gentle 5161 (F, G, MICH, NY, US); Toledo District, nr. San Antonio, 8 Oct 1952 (q fl), Gentle 7781 (F, G, GH, MICH, NY); nr. Honey Camp, Sep 1928 (e'), Lundell LP-19 (COL, F, GH, NY, US); El Cayo District, Monkey Falls, 7 Aug 1936 (a), Lundell 6970 (C, GH, MICH, NY); Toledo District, Punta Gorda, "El Dorado," 6 Oct 1932 (de), Schipp 1010 (A, BM, F, G, GH, K, MICH, MO, NY, Z). HONDURAS. ATLANTIDA: Nr. Tela, 28 Dec 1927 (q fl), Standley 53841 (A, F, US), 14 Dec 1927-15 Mar 1928 (st), Standley 54766 (A, F, US). COMAYAGUA: Chichipates, Rio Yure, 22 Nov-15 Feb 1981 ((? fr), Nelson et al. 6752 (MO); La Ceiba, Rio Yure, 6-15 Feb 1981 (? fl-fr), Nelson et al. 7323 (MO); Jaitique, nr. Taulabe, 8 Dec 1974 (R fl-fr), Hazlett 2413 (MO); La Choca Mt., nr. Coyacutena, 1500 m, 14 Apr 1957 (? fr), Molina R. 8101 (EAP, F, NY). COPAN:Rio Copan, nr. Copan ruins town, 18 Nov 1969 (2 fl-fr), Molina R. et al. 24596 (BM, EAP, F, GH, NY), 19 Nov 1969 (9 flfr), Molina R. et al. 24623 (EAP, F, MO, NY). CORTES: Calan, 28 Mar 1964 (? fr), Dickson 89 (U), 4 Oct 1964 Nr. Danli, Cafetales (a'), Dickson 1327 (US). EL PARAISO: de Montafia Apauhis, 1000 m, 15 Mar 1950 (? fl-fr), Molina R. 7461 (EAP, F, NY, US). GRACIAS A DIos: Mosquito, Rio Platano, 0-4 hrs upriver from Ras, 23 May 1973 (a'), Gentry et al. 7543 (MO). ISLAS DE LA BAHIA: Roatan Island, 4.5 km E of Coxenhole, 21 Apr 1967 (2 fl-fr), Molina R. 20713 (EAP, F, NY), 24 Apr 1967 (q fl), Molina R. 20806 (EAP, F, NY, US, WIS); Flowers Bay, 13-20 Mar 1978 (? fl-fr), Nelson et al. 4504 (MO, U). OLANCHO: 6 km SE of Catacamas,30 Mar 1987 (9 fr), Ortega U. 228 (MO); between Catacamas & La Presa, N of Catacamas, 20-25 Mar 1949 (st), Standley 18683 (F). SANTABARBARA:Nr. Santa Barbara,Rio Ulua, 14 Dec 1950 (? fl), Molina R. 3810 (BM, EAP, F, GH, MO, US). TEGUCIGALPA:Mont de la Flor, Dec 1937 (st), Hagen et al. 1130 & 1177 (F, NY). YORO: Concepci6n, 19 Aug 1933 (a), Edwards 644 (A, F, K, US); nr. Progreso, 24 Jan 1928 (st), Standley 54997 (A, F, US). EL SALVADOR. Nr. San Salvador, Aug 1922 (ae),

43 Calder6n 1065 (GH, MO, NY, US), Jan 1923 (? fl-fr), Calderon 1448 (GH, MO, NY, US); Izalco, Jul 1923 (st),

Calderon 1698 (U); nr. San Salvador,(? fr), Renson 112 (NY, US); Dtto. La Libertad, ruins of San Andres, 13 Dec 1988 (? fl), Reyna 1420 (F, MO); Dtto. Ahuachapa, San Benito, 29 Dec 1992 (? fl-fr), Sandoval et al. 956 (MO); nr. San Vincente, 2-11 Mar 1922 (? fr), Standley 21404 (F, GH, MO, NY, US); nr. Sonsonate, 18-27 Mar 1922 (st), Standley 22340 (GH, US). NICARAGUA.BOACO:8.5 km NW of Camoapa, 1060 m, 24 Jan 1980 (? fr), Araquistain et al. 976 (MO); San Jose de los Remates, 2 Oct 1984 ($ fl), Moreno 24840 (MO), ('), Moreno 24855 (MO); 1-2 km SE of Santa Maria, Rio Fonseca, 12 Sep 1983 (a'), Nee et al. 27961 (MO);CerroMombachito,W of Boaco, 8 Oct 1979 ($ fl), Stevens et al. 14752 (MO). CARAZO: Estacion Biol6gica Chococentro, 19 Mar 1983 (a), Grijalva 2431 (MO), 4 Feb 1984 (a'), Grijalva et al. 3555 (MO). CHONTALES: BetweenBoaca & Ayoyapa,18 Mar 1961

(? fr), Bunting et al. 713 (EAP, F, MO); Cerro Oluma, 4 Jan 1984 (R fl), Gentry et al. 43994 (MO). ESTELf: 4 mi S of Esteli, 4 Aug 1971 (a'), Dwyer et al. 441 (MO,

US); Saltola Estanzuela,29 Mar 1983 (? fl-fr),Moreno 21136 (MO);LasLajitas,betweenEl Jobo& Estanzuela, 29 Jun 1983 (a'), Moreno 21633 (MO); SW of Pueblo Nuevo, 1 Nov 1976 (e), Neill 1208 (MO); nr. Esteli, rd.

to Estanzuela, 1000-1035 m, 13 Nov 1978 (2 fl-fr), Stevens 10787 (MO). GRANADA: Nr. Granada,17 Feb 1903 (e), Baker21 (MO). LE6N: 6 km SE of Ciudad Nagarote, 5 Mar 1983 (a'), Grijalva 2382 (MO), 27 Dec 1983 (e), Grijalva 3360 (MO). MANAGUA:Nr. Managua, Bro. Garnier 3019 & 4196 (A); CarreteraSur, km 11.5,

2 Oct 1982 (v), Grijalvaet al. 1300 (MO);betweenEl Crucero& FincaSantaJuli, 17 May 1947 (st), Standley 8375 (F); Hwy. 8, ca. 2.4 km SW of intersection with Hwy. 2, 18 Sep 1977 (), Stevens 3999 (MO). MASAYA: Miravalle, 4 km from Las Nubes, 11 Dec 1980 (9 fr), Moreno 5225 (MO); E slope of Volcan Masaya, 14 Feb 1978 (e), Neill 3192 (MO).MATAGALPA:Rd.MatagalpaSiuna, km 78, La Gloria, 13 Sep 1982 (a'), Moreno 17195 (MO); 7 km NW of Esquipulas, 8 Feb 1985 (9 fl-fr), Moreno 25423 (MO); between Matagalpa & Mayiguas, 18 Jan 1984 ($ fl-fr), Sandino 4688 (MO). NUEVA SEGOVIA:

8 km NE of Jicaro,2 Aug 1980 (o),

Moreno 1714 (MO). RIVAS:"Vergel,"15 Mar 1984 (? fl-fr), Araquistain 3823 (MO); Isla Ometepe, Volcan

Maderas,Merida, 11 Feb 1984 (? fl-fr), Robleto 206 (MO);rd. Hda.Fatima-SanJos6 de la Montafia,12 Sep 1982 (a), Sandino et al. 3598 (MO); rd. to Salinas, 19 (MO). ZELAYA: Awas Tingni, 40 km S of Waspam, 19 Mar 1971 (? fr), Little 25232 (MO); El Recreo, 21 Nov 1948 (a), Long 215

Feb 1983 (a'), Sandino 4223

(F); Rio Grande,Isabel, 28 Apr 1949 (? fl-fr), Molina R. 2497 (EAP, F, GH, US); Buenavista, 6 km NE of Siuna, 26 Feb 1983 (? fr), Moreno 20803 (MO); nr. El Recreo, Rio Mico, 28 Apr-14 May 1949 (? fl-fr), Standley 19797 (EAP, F). COSTA RICA. ALAJUELA: San Miguel Oeste, Naranjo, 1000-1400 m, 17 Oct 1986 (? fl), Herrera Ch. 11 (BG, F, MO); Upala, nr. Laguna las Camelias, 19 Nov

44

FLORANEOTROPICA

(MO); ParqueNacional Darien, Serraniade Sapo, 26 Nov 1990 (* fl-fr),H. Herreraet al. 806 (BG, EAP);S of El Real, CerroPirre,800-1070 m, 29 Jun 1988 (st), Aubreville s.n. (P), (? fr), Le6n 3221 (CR), (? fr), Little McPherson12662 (BG, MO);betweenPaya & Palo de las Letras, 10 Jun 1959 (ae), Stern et al. 210 (GH, LE, 20270 (CR). GUANACASTE: Comelco, 20 Jan 1976 (? AboveRio Guanico,5 Jan 1989 fl-fr), Frankie432A (MO); Rio Chiquitode Tilaran,2 MO,US). Los SANTOS: km NE of Rio Chiquito,1 Dec 1987 (? fr),Haberet al. (? fl-fr), McPherson 13512 (BG). PANAMA: Nr. Rio 7846 (BG, MO);Bagaces,8 Feb 1978 (? fl), Hartshorn Pescado,30 Jul 1940(? fl), Bartlettet al. 16588 (MICH, 2107 (F); SantaRosaNationalPark,Rio Guapote,4 Dec MO); BarroColoradoIsland, 12 Jun 1971 (o), Croat 15247 (GH, MO, NY, US); Rio Pita, 1-3 mi above 1977 (e), Janzen 10646 (MO); W of San Jose Pinilla, 4 Feb 1978 (oi), Liesner 4958 (MO). HEREDIA:Puerto confluencewith Rio Maestra,14 Oct 1961 (? fr), Duke Viejo, FincaLa Selva, 5 Jul 1969 (st), Frankie210a & 4736 (BM, GH, MO, US); BarroColoradoIsland, 13 210c (F, MO); 1-12 km SW of Las Horquetas,19 Apr Jun 1969 (a'), Foster 988 (GH, MICH, MO); rd. El Ll1988 (? fr), Hammelet al. 16690 (BG, MO);nr. Puerto ano-Cartird., km 4-6, 26 May 1975 (? fl-fr), Mori et Viejo, 15 Dec 1970 (? fr), Hartshorn961 (F), 19 Dec al. 6391 (MO);TrinidadR., nr. HydroelectricStation, 1987 (e), Herrera Ch. 1352 (MO); nr. Capulii, Rio

Grandede Tarcoles, 2 Apr 1924 (st), Standley 40121 (US). CARTAGO: Turrialba, 5 May 1965 (? fr),

1970 (c'), Hartshorn 1072 (DUKE, F, MO); Puerto Viejo, mouth of Rio Sarapiqui, 21 Jan 1966 (a), Jimenez M.

17 May 1914 (e), Pittier 6636 (C, GH, NY, P, US); 5 mi

W of Chepo, 29 Sep 1972 (? fr), Tyson6732 (MO). 3631 (BM, CR, F, GH, NY, US). LIM6N:5 km of rail- SANBLAS: Nr. Isla MiriaUbigandup,rd. to Digole, 19 road bridge over Rio Pacuare,20-22 Dec 1969 (? fl- Jul 1987 (e), Herrera 291 (MO). VERAGUAS: Santiago, fr), Burger et al. 6939 (CR, F, MO, NY, PMA); 4 km S 8 Mar 1961 (? fl), Dwyer 1353 (MO, NY). CUBA. CAMAGUEY: Between La Gloria& Columof Cahuita,9-14 Feb 1977 (? fl-fr),Burgeret al. 10496 (CR,EAP,F, MO,PMA);Madrede Dios, FincaLa Lola, bia, Rinc6nGrande,6 Mar 1909 (? fr), Shafer617 (NY, 18 Mar 1949 (? fl-fr), Holdridge 2525 (EAP, US); US). LA HABANA: Guatao,Feb 1917 (? fr), Bro. Le6n 7078 (NY). ISLA DE PINOS: Sierrade los Caballos, 2 Tortuguero,Caflo Harold, 10 Feb 1989 (oi), Robles 2654 (MO); Rio Reventaz6nbelow Cairo, 18-19 Feb 1926 Feb 1916 (st), Brittonet al. 15135 (NY), NuevaGerona, South5 Feb 1956 (st), Killip 45593 (US). ORIENTE: Sierra (st), Standley et al. 48591 (US). PUNTARENAS: em tip of Nicoya Peninsula,1-7 Dec 1969 (st), Burger Maestra,Lomadel Gato,Jan 1948 (? fl), Bro. Clemente et al. 6681 (CR, F); Monte Verde, 1100-1200 m, 29 5802 (GH, US); Bayate, 29 Nov 1917 (9 fl), Ekman Sep 1985 (o), Haber et al. 2935 (MO); Osa Peninsula, 9005 (F, G, MICH,NY, US); nr. PiedraGorda,23 Dec Parque Nacional Corcovado, 6 Jul 1977 (o), Liesner 1909 (? fl-fr), Shafer3272 (F, NY, US); MonteVerde, 2971 (MO,U), 6 Jul 1977 (? fl), Liesner3106 (MO,U); 1 Jul 1859(9 fl-fr), Wright589 (BM,GH, K, LER,MO, Parque Nacional Corcovado, Sirena, 27 May 1988 (ei), P), (? fl-fr), Wright592 (F, GH, K, LE, MO, NY, P). Kernan 536 (MO). SAN Jost: Santa Ana, ($ fl), Caffrey PINAR DEL Rio: SantaCruzde los Pinos,Jan 1946 (a), 56 (CR); rd. Santa Ana-Villa Col6n, km 20-21, 21 Feb Bro. A. Liogier470 (GH,US); Esperanza,13 Sep 1910 (e), Britton et al. 7337 (NY); Lomade la Gloria,Aug 1966 (? fr), Jimenez M. 3678 (CR, F, NY, US). PANAMA. BOCASDELTORO:Shepherd Island, 22 Aug 1964 (9 fl-fr), McDaniel 5168 (MO); Almirante, 29 Nov 1971, (? fl-fr), Lao et al. 441 (MO, NY); Isla Bastimentos, Bocatorito, 15 Feb 1989 (ei), Petterson et al. 6875 (MO); Changuinola valley, 10 km from Almirante by railroad, 28 Aug 1940 (a), Seibert 1581 (MO, US); between Finca St. Louis & Konkintoe, Rio

1926 (e), Bro. Le6n 12680 (A, NY); Rio San Vincente,

Vinales,17 May 1941 (9 fr), Bro.Le6net al. 20041 (A, GH); Las Martinas,19 Dec 1911 (9 fl), Shafer 11078 (NY, US); nr. Candelaria, 15 Sep 1904 (a'), Wilson 1629 (US). SANTA CLARA: Belmonte, Soledad,Cienfuegos,

26 Mar1927 (? fl), Jack5002 (A, B, K, NY, P), 17 Mar 1928 (a), Jack5842 (A, K, P, US); La Sierra,San Blas, Cricamola, 12-16 Apr 1938 (? fl-fr), Woodsonet al. 4 Dec 1928 (? fl), Jack 6777 (US); Banoa Mts., May 1924 (A, F, MO, NY). CHIRIQUi: Nr. ChiriquiLagoon, 1920 (? fr), Luna 609 (NY). 13 Nov 1940 (? fl-fr), Wedel1599 (GH, US), 13 Feb JAMAICA. Moneague, 1850 (2 fr & d), Alexander 1941 (9 fr), Wedel2075 (GH, MO, US); Peninsulade 303 (K, NY); Halls Delight, 14 Nov 1895, Harris 6075 Burica,nr. San Bartolome,28 Jul-I Aug 1940 (2 fl), (BM, F, G, NY, US); BluefieldsMts., KentuckyHill, 7 Woodson et al. 866 (MO, NY, US). COCLk: Between

Mar 1908 (a'), Harris 10211 (BM, F, K, NY, US); Upper

CerroPil6n & Vallede Ant6n,15 Aug 1967 (9 fl), Duke Clarendon,Ionis Cave Wood, 1 Mar 1910 (? fl-fr), et al. 13908 (MO); Cerro Pil6n, 9 Jan 1972 (oi), Gentry Harris10858 (F, K, NY,US); Trelawny,CockpitCounty, et al. 3653 (MO, U); Boca del Toabre, at confluence of nr. BurntHill, 14 Feb 1960 (? fr), Proctor 20588 (A, Rio Toabre & Rio Cocle del Norte, 11 Apr 1969 (? fr), BM, MO, NY, U, US); betweenRidgate& GingerHill, Lewis et al. 5568 (MO, US). COL6N:Lower Rio Guanche, St. Elizabeth, 29 May 1961 (? fr), Proctor 22304 18 Apr 1973 (2 fl-fr), Dressler 4343 (MO); between (MICH,NY, U, US); Trelawny,MangoTreeHill, 6 Nov Mosquera & Palmas Bellas, 17 Sep 1974 (? fl-fr), Mori et al. 1965 (MO, US); nr. Gamboa, 7 Jun 1914 (? fl), Pittier 6652 (G, P, US, Z). DARItN: Rio Ucurganti, 7 Jul 1967 (? fl), Bristan 1119 (F, GH, MO, UC, US); nr. Santa Fe, (? fl-fr), Duke 8392 (MO); Rio Tuira, between Rio Punusa & Rio Mangle, 1 Oct 1967 (? fl), Duke 14629

1979, Proctor 38353 (NY).

HAITI.MorneSal, Bois Neuf, 22 Jul 1924(st), Cook 13 (G, GH, US); Massif du Nord, Marmeladetowards Dondon,28 May 1927 (2 fr), Ekman8285 (US);Massif de la Salle, Papette,29 Nov 1927 (a'), Ekmnan9387 (G, K,

US); Dept.du Nord,nr. St. Michelde l'Atalaye,30 Nov

TAXONOMIC TREATMENT

45

1925 (9 fl-fr), Leonard7603 (GH,NY, US); TortueIs- San Jose del Palmar,Rio Torito,affluentof Rio Habita, land,nr.LaVallee,31 Dec 1928(st),Leonardet al. 11360 10 Mar 1980 (a), Forero et al. 6940 (MO, NY, U); (NY,US);betweenPortMargot& Correil,1903(e), Nash trail from Unguia to base of Serraniadel Darien, 18 206 (NY);Massifde la Hotte,14 kmNW of PortSalut,2 Jul 1976 (? fl), Gentryet al. 16765 (MO,NY, U); Rio Jan 1985 (e'), Zanoni et al. 33365 (MO, NY). DOMINICAN REPUBLIC. Prov. La Vega, Cotuy, 28 Jan-7 Feb 1921 (a'), Abbott 757 (US); Prov. Barahona, Barahona, Jun 1911 (6), Fuertes 1086 (BM, F, G, GH,

K, NY, P, U, US, Z); Prov. Santiago, Jacagua, nr. Santiago,5 Jan 1967 (e), Jimenez5165 (US); Rinc6n de Yuboa,6 May 1973 (? fl), Liogier 19024 (NY); San Pedrode Macoris,7 Mar 1913 (9 fl), Rose et al. 3719 (F, GH, NY, US); Prov.Macoris,Consuela,15-24 Nov 1909 (v), Taylor146 (F, NY, (e), Taylor229 (F, NY); Prov. Samana,3.5 km E of Las Terrenas,3 Nov 1981 (a') Zanoni et al. 17686 (BG, MO, NY). PUERTO RICO. Candelaria, nr. Bayam6n, Apr 1913(9 fl-fr),Brittonet al. 2853 (F,MO,NY,US); Hato Tejas,Bayam6n,24 Jun 1963 (st), Liogier9666 & 9672 (NY); Guajataca,7 Apr 1967 (st), Little 21910 (US); Utuado,Mar 1887 (9 fl), Sintenis6309 (A, BM, C, G, GH, F, K, MO, NY, P, US, Z); Rio Piedras,Ganachalez, 6 Dec 1914 (9 fl), Stevenson 2407 (NY, US); 3426 (NY); Campanillas,11 Feb 1959 (9 fl), Woodbury s.n. (NY). Sabana,4 Feb 1959 (st), Woodbury COLOMBIA. AMAZONAS:PuertoNariiio, Parque

Taparal, San Juan, 19 Aug 1962 (a), Hugh-Jones 249

(K, US); Rio San Juan,nr.Andagoya,27 Apr 1939 (? fl-fr), Killip 35392 (COL, F, US). CUNDINAMARCA: Between San Francisco & Subachoque,26 Jan 1944 (e'), Garcia-Barriga 11030 (COL). GUAJIRA: 15 km S

of Mingueo,on trailto PuebloViejo, 21 Aug 1986 (st), Gentry et al. 55384 (MO), 55456A (BG); 12 km S of Carrapia, 27 Jun 1944 (a'), Haught 4224 (COL, F, US);

Mun. Riohacha, Cgto. Tomarraz6n,2 Sep 1990 (st), Marulanda et al. 2229 (HUA). MAGDALENA: Parque

Nacional de Tayrona,between Pueblito & Calabazo, 26 Oct 1972 (a), Kirkbride 2572 (MO, NY, U); Mun.

Fundacion,ca. 5 km N of SantaRosa, 5 Aug 1971 (2 fl-fr), Romero-Castafieda 11165 (COL); Santa Marta, Sep 1900(? fl-fror d'), H. H. Smith1752 (F, MO,WIS), Jul 1901 (ei), H. H. Smith 2732 (F, MO, WIS). META: Cubarral, 16 Oct 1938 (e), Cuatrecasas et al. 3672

(COL);Sierrade la Macarena,CanioEnstrada,13 Jan 1950, Philipson et al. 2097 (COL). PUTUMAYO: Rio Putumayo,La Concepci6n,27 Nov 1940 (ae), Cuatrecasas 10840 (COL, F, US). SANTANDER: Nr. Puerto Berrio,between Rio Carare& Rio Magdalena,27 Jul

Nacional Amaca-yacui, 10 Aug 1989 (9 fl-fr), R. Vdsquez et al. 12697 (COL). ANTIOQUIA:Mun. Turbo, rd.

1935 (a'), Haught 1864 (COL, F, US). VALLE: Mun.

Curran 205 (US), (9 fl-fr), Curran 372 (US); Mun.

Williams et al. 10719 (BM, F, U, VEN); Ocumare, 8 Jul 1952 (e), Garcia 186 (VEN); Parque Nacional Henri Pittier, 1000 m, 20 Jun 1963 (a'), Montaldo 3504 (MY,

Sevilla, La Esmeralda, 1700 m, 14 May 1979 (e), Tap6ndel Darien,betweenRio Le6n& LomasAisladas, CuadrosV 653 (BG, MO); Rio Anchcaya,El Prado,4 24 Dec 1983 (9 fl-fr), Brand et al. 718 (MO); Mun. Aug 1943 (9 fl), Cuatrecasas14857 (F, GH, MO);Rio Remedios,14-17 km NW of Remedios,rd.to Zaragoza, Calima, between La Trojita & Guadalito, 19 Feb-10 15 Sep 1987 (9 fl), Callejaset al. 5188 (MO);2 km N Mar 1944 (? fr), Cuatrecasas16810 (F, GH, VALLE); of Mutata, 19 Nov 1987 (e), Callejas et al. 5655 (BG, Rio Cajambre,5-15 May 1944 (? fr), Cuatrecasas MO, NY); Rio Turbo,Turbo,2 May 1946 (o), Haught 17622 (F, GH, VALLE);Cartago,Hda. La Silvia, 214819 (COL, US); Mun. Anori, Providencia, 31 May 26 Dec 1944 (? fl), Duque-Jaramillo4067A (COL); 1973 (9 fl-fr), Soejarto 4099 (A, MO);Mun. Segovia, valley of Rio Dagua,Cisneros,21 Sep 1922 (e), Killip 24.5 km N of Remedios,20 Jul 1987 (9 fl-fr), Thomas 11479 (GH, NY), 5 May 1939 (a'), Killip 35580 (A, et al. 5498 (BG). BOLlVAR: Mun.Cartagena,Jard.Bot. BM, COL,F, US). GuillermoPinieres,Feb 1980 (? fl), CuadrosV 1097 VENEZUELA.ARAGUA: RanchoGrande,1000 m, (MO); San Martinde Loba, Apr-May 1916 (9 fl-fr), 30 Nov 1938 (2 fl-fr), Alston et al. 115 (VEN) = Ll. San Femando,betweenSantaRosa& Corocito,11 Mar 1981 (9 fl), Espina et al. 1002 (COL). CALDAS:Santa Cecilia, 21 Nov 1945 (9 fl-fr), Sneidern5169 (F, US); Salamina,1900 m, May 1943 (9 fr), Bro. Tomds1823 (US). CAQUETA: Rio Caqueta, below mouth of Rio Ortegueza,Solano, 8 km SE of Tres Esquinas,5 Mar 1945 (Y fl), Little et al. 9564 (COL,US), 8 Mar 1945 (a), Little et al. 9670 (COL, US); Morelia, 7 Oct 1941 (9 fl), Sneidern 1094 (COL), 18 Oct 1943 (ei), Sneidern

1171 (COL).CASANARE: Rio Meta,betweenCanioSan Miquel & CainoDuya, 12-13 Jan 1990 (ae), Mahecha 6384 (UDBC). CAUCA: Rio Micay, Guayabal,25 Feb 1943 (ae), Cuatrecasas 14117 (F, GH, VALLE);Rio Micay,Noanamito,26-27 Feb 19431 (9 fl-fr), Cuatrecasas 14227 (F, GH, VALLE). CHOC6: Lunar de

U); RanchoGrande,27 Jul 1979 (? fl), Nee et al. 16897 (F); Valle de Patanemo,E of PuertoCabello,Dec 1919 (e), Pittier 8659 (GH, NY, US, VEN). BARINAS:Rd.

Barinas-San Crist6bal, km 76, 14 Jul 1964 (? fl), Breteler4001 (COL,EAP,F, MO,NY, P, U, US, VEN); ReservaForestalCaparo,E of El Cant6n,10 Apr 1968 (? fl), Steyermarket al. 102044 (NY, US, VEN); between Caimital& Barrancas,26 Apr 1972 (st), Veillon 77 (US). CARABOBO: E of Puerto Cabello, 4-5 Dec 1919

(v), Pittier 8659 (US); Rio Borburata,2.5-6 km S of Borburata,3 Jan 1970 (? fl), Steyermarket al. 102409 (MO, NY, U). FALC6N:ReservaForestalTocuyo,Aug 1970 (a), Blanco 905 (NY, U, US); Mun. Jacura, Dtto.

Jurubbida,17 May 1990 (9 fl), Barbosa 6633 (MO); Acosta,CerroLa Mina, 14 Nov 1979 (? fl), MarcanoRio SanJuan,nr.Palestina,29 May 1946 (9 fl), Cuatre- Berti et al. 449-979 (MO, U); Guaidema, Tocuyo, 10 casas 21401 (F); Bahiade Solano,betweenMutis& El Jul 1923 (e), Mell s.n. (NY, VEN), Dtto. Acosa, Belen, Valle, 9 Jun 1950 (e'), Fernandez 310 (COL, US); Mun.

19 Apr 1977 (st), Ruiz Z. et al. 1147 (U). LARA: Rio

FLORANEOTROPICA

46 Guaremal,NE of Barquisimeto,28 Jul 1979 (? fr), RioClara,1100 Meijeret al. 69 (MO,NY);Dtto.Iribarren, m, 4 Aug 1968 (? fl), R. F Smith V-4171 (F); 33 km SE of Sanare,Paso de Angostura,Dtto. Jimenez,28-31 Jul 107625(MO,NY,U). M?RiDA: 1973(? fl-fr),Steyermark El Vigio, CanoAmarello,6 Feb 1955 (? fr), Bernardi 1945 (VEN);El Guaimaro,5 May 1955 (? fl), Bernardi s.n. (NY); betweenCanloZancudo& La Azulita,2 Sep 1965 (? fl-fr),Breteler4573 (COL,EAP,G, K, MO,U, US, VEN). MIRANDA: Between Chuspa& Aricagua,8 Dec 1969 (a), Steyermarket al. 102349 (MO, U); rd. to Turgua,1000-1100m, Dec 1992(? fr),Meieret al. 3188 Dtto.SanFelipe,ParqueNacionalYurubi, (BG).YARAcuY: 15 Nov 1979 (? fl), Marcano Berti et al. 482-979 (U). ZULIA:

20 km NW of El Vigia,CailoPadre,2 Dec 1966

(e), Bruijn 1330 (MO, NY, U, VEN); Machiques, be-

tween Rio Negro & Rio Tocuco, 6 Jan 1954 (? fl-fr), Little 16154 (VEN);SE of Machiques,Sierrade Perija, 25 Aug 1967 (e'), Steyermarket al. 99683 (U, US, VEN),

25 Aug 1967 (? fl), Steyermarket al. 99726 (K, MO, NY, U, VEN); Cerro Cinco de Julio, 8-10 m NW of Carichuano,4 Jun 1980 (? fl), CorpozuliaCampamento Steyermark et al. 12330 (MO, NY). ECUADOR. GUAYAS: Cord.Chong6n-Colonche,9

Jul 1995 (? fl), Cornejo et al. 4231 (BG); Reserva Ecol6gicaManglaresChurute,CerroPanchoDiablo, 3 May 1996 (? fl-fr),Cornejo et al. 5087 (BG).MANABI: Balao,Feb 1892(? fl-fr & e), Eggers 14430 (A, K, LE, US, Z). NAPO: Tena, 17 Oct 1939 (? fl), Asplund9387 (G, K, NY,P, S, US); PuertoMisahualli,3 Mar1980(st), Berget al. 1112 (MO,U); ReservaBiol6gicaJatunSacha, 8 km E of PuertoMisahualli,17-21 Nov 1988 (? fr), Cer6n et al. 5621 (AAU, MO, NY, QAME, QCNE);

CantonPompeya,La Joyade los Saches,Rio Indillama, 13 Dec 19442 (? fl-fr), Gudino et al. 2132 (QCNE); Puerto Napo, 4 Apr 1969 (e), Lugo S. 1032 (BG);

5 km NE of Coca, 18Estaci6nExp. INIAP-Payamino,

(MO);Rio Santiago,QuebradaCaterpiza,4 Dec 1979 (a'), Tunqui224 (BG, MO, U); Aramanga,5 Apr 1960 (? fr), Woytkowski 5645 (G, GH, MO, US). HUANUCO: Prov.Huaniales,QuebradaMaquizapa,rd. to Monz6n, 18 Feb 1966 (e), Schunke V 1096 (F, NY, U); Prov. Pachitea,Dtto. Honoria,BosqueNacionalde Iparia,nr. Tournavista,10 Oct 1967 ($ fr), SchunkeV 2200 (F, GH, K, NY, MO, NY); Tingo Maria,26 Oct 1938 (a'), Storket al. 9506 (F, G, K). LORETO:Nr. JenaroHerrera, 25 Nov 1988 (? fl-fr), Daly et al. 5766 (BG, NY); nr. Iquitos,Rio Itaya,20 Nov 1940 (Y fl), Asplund14619 (G, NY, P, US); Rio Napo, nr. Casaria,15 Sep 1972 (9 23 Augfl), Croat20246 (C, GH,MO,NY);Yurimaguas, 7 Sep 1929(9 fl), Killipet al. 27685 (F);Rio Putumayo, Florida,at mouthof Rio Zubineta,Mar-Apr1931 (a'), Klug 2079 (A, BM, F, GH, K, MICH, MO, NY, US); above Pongo de Manseriche,CarapisaCreek, 12 Dec 1931 (a'), Mexia 6262 (BM, F, G, GH, K, MICH,MO, NY, U, US, Z); above Pongode Manseriche,Zungarosa Creek, 19 Dec 1931 (q fl-fr), Mexia 6303 (BM, F, G, GH, K, MICH,MO,NY, U, US, Z); nr.TenientePinglo, 2117 (F, GH, NY, P, US). 4-7 Oct 1962 (9 fl), Wurdack SANMARTfN:Rio Huallaga,Juanjui,Sep 1934 (a'), Klug 3794 (BM, F, GH, K, MO,NY, US); Rio Shilcayo,N of Tarapoto,14 Apr 1986 (? fl-fr),Knappet al. 7085 (BG, MO); Prov.MariscalCaceres, TocacheNuevo, Fundo San Rafael,22 Sep 1970 (a'), SchunkeV 4430 (F, GH, K, MO,NY,US);Prov.MariscalCaceres,TocacheNuevo, Canuto,10 Oct 1970 (2 fl-fr), SchunkeV 4494 (F, MO, NY, US); Prov.Lamas,Dtto. Alsonso de Alvarado,San km 72, Juande Pacaizapa,rd. Tarapoto-Moyobamba, 1000-1500 m, 1 Jun 1977 (? fr), SchunkeV 9585 (F, MO, U, US); nr.Tarapoto,Jul 1855 (? fr), Spruce3958 (BM, K, NY), Mar 1856 ($ fl-fr),Spruce4521 (B, BM, G, K, LE, NY). UCAYALI:Prov.CoronelPortillo,Lago Yarina-Cocha, 5 Nov 1984 (? fl), Maas et al. 6208 (F).

The species is very variable,especially in the charactersof the lamina(dimensions, shape, texture,margin, andindumentum)andin the length of the inflorescences. Part of the variation is connected with distribution,which openedthe possibility to recognize varietiesor subspecies (Bureau, 1873; Burger, 1962). Most of the collections from CentralAmerica have entireandscabrousleaves and shortpistillateinflorescences (subsp. ramon (Schlechtendal& Chamiso)W. Tarquim 18 Apr 1985 (9 fl), Harling et al. 24274 (BG). Burger). In collections from the GreaterAntilles the SUCUMBtOS: Dureno,11 Mar-I Apr 1986 (? fl), Cer6n laminais smooth and glabrouswith an entiremargin; 204 (BG, MO, QAME,QCA);nr. Limoncocha,27 Sep thepistillateinflorescencesareoftenrelativelylong with 1977(R fl), Foster3877 (AAU,F, QCA);ReservaFaunis- ? distantflowers (subsp. racemosa). Most collections tica Cuyabeno,between Rio Pacuyacu& Zancudo,27 fromSouthAmericahave smooth(oftenrathernarrow) Sep 1991 (? fl), Palacios et al. 7699 (BG, QCNE), (a'), leaveswitha dentateto denticulatemarginandrelatively Palacios et al. 7720 (BG,QCNE).ZAMORA-CHINCHIPE: long pistillate inflorescences with distant flowers Rio Nangaritza,between Shaime& Mariposa,21 Nov (subsp. meridionalis(Bureau)W. Burger).In Panama 1996 (? fl), Eyndenet al. 842 (BG). PERU. AMAZONAS:Valleyof Rio Santiago,65 kmN the CentralAmerican form gradually passes into the of Pinglo, 1 Jan 1980 (2 fl), Huashikat1703 (MO, U), SouthAmericanone. In northemSouthAmerica(e.g., 18 Jan 1980 (9 fl), Huashikat 1835 (U); Quebrada in Venezuela) one can find specimens matching the Yutinca,aboveCenepa,17 Nov 1972 (? fl), Kayap107 CentralAmericanform.In the northempartof the spe26 Feb 1986 (a), Palacios et al. 1055 (BG, MO, NY,

trailCeilanQAME,QCA,QCNE).PASTAZA:RioCuraray, Rio Cononaco,6 Jun 1980 (? fl-fr), Brandbygeet al. 31654 (AAU);Rio Curaray,Lorocachi,1 Jun 1980 (9 fr),Jaramilloet al. 31743(AAU).PICHINCHA:RioToachi, rd. La Union del Toachi-Las Minas, km 3-4, 23 Feb 1981 (9 fl-fr), Berg et al. 1308 (QCA,U). Los Rios: Canton Vinces, between Mochachiana & Palenque, EsteroPeniafiel,25 Jan 1981 (? fl), Gentryet al. 30804 (AAU,U). MORONA-SANTIAGO: Rd. Gualaquiza-Nueva

TAXONOMIC TREATMENT

ciesrange(Mexico,Guatemala) onecanfindplantswith smoothleaveswitha (faintly)dentatemargin,reminiscentof theleavesof Trophismexicana.InSouthAmerica andPanama thespeciesismorevariable thanintheother thetotalvariapartsof the speciesarea.Considering tion(incomparison withthatof T mexicana)thepresent authorprefersnotto recognizeinfraspecific taxa. Localnames.Mexico:Campeche: ram6ncolorado. Guatemala: Izabal:yax-ox;Pet6n:ramoncolorado, yaxox. Belize: Cha-cox(Maya),cherry,ramon,white ramon. Honduras:Atlantida:masicaron,ramon; Olanche:barbade tamagaz,mora;Tegucigalpa: hoja tinta,ramon.Nicaragua: Masaya:ojochemacho.Costa Rica:Heredia:ojochillocolorado.Cuba:ramonde

47 Olmedia falcifolia Pittier, Contr. U.S. Natl. Herb. 13: 435. 1912. Type. Costa Rica. Puntarenas: Rio Akum (Hacum), nr. Buenos Aires, Feb 1892 (c?), Tonduz 6683 (holotype, US). Trophis aurantiaca Herzog, Repert. Spec. Nov. 7: 51. 1909; Burger, Ann. Missouri Bot. Gard. 49: 18. 1962. Type. Bolivia. Santa Cruz: Cord. de Santa Cruz, Quebrada de Cunucii [probably Rio Macufiucu], Herzog 574 (holotype, B-n.v.; isotype, Z).

Shrub or tree, to 20 m tall. Leafy twigs 1-4 mm thick, hispid to hispidulous, or strigose to strigillose, ? scabrous. Lamina oblong to lanceolate, (3-)10-25(-40) x (1-)3-7(-1 5) cm, mostly broadest above the middle, inequilateral, chartaceous to subcoriaceous; apex acumibestia.Jamaica:ramoon.DominicanRepublic:ramon. nate to caudate; base acute; margin dentate to denticuPuertoRico:ramon,ramoncillo.Colombia:Caqueta: late or subentire; upper surface scabrous to scabridulous sande;Cundinamarca: lecherocolorado.Venezuela: to almost smooth; lower surface hispid to hispidulous Aragua: marfil;Barinas:charoblanco,charoamarillo, to strigillose, ? scabrous; venation above plane, beneath charomorado;Merida: guaiimaro negro;Zulia:chatchi, prominent; lateral veins (6-)8-20(-24) pairs; tertiary guaymaromacho.Ecuador:Napo:rumicaspiyura; venation partly scalariform; petiole 0.3-1.2 cm long, Zamora-Chinchipe: pitiu(Shuar).Peru:Amazonas: strigillose, caducous; stipules 0.5-1.2 cm long, strigillose to hispidulous, caducous. Staminate inflokusuknash,pituk, uchi pituk;Loreto:insira. rescences solitary or in pairs (or as many as 6 together 2B.TrophisP.Brownesect.Olmedia(Ruiz&Pavon) on short-shoots) in the leaf axils, discoid-capitate, 0.4C. C.Berg,Proc.Kon.Ned.Akad.Wetensch., Ser. 1 cm diam., subsessile or with to 0.6 cm long peduncle, C, Biol. Med.Sci. 91: 354. 1988.Type.Olmedia involucrate; involucral bracts in 3-4 rows, ovate to lanRuiz& Pav6n(= T caucana (Pittier)C. C. Berg). ceolate, hispidulous to strigillose; flowers 10-30, sessile; perianth 4-parted, 2-3 mm long, strigillose; filaTreesorshrubs. Staminateinflorescencesdiscoid- ments ca. 3.5 mm long; anthers 1.3-1.8 x 0.5-0.9 mm, capitateandinvolucrate; tepals4, valvatein thebud. the connective narrow, sometimes appendiculate; pisPistillateinflorescenceswith1(-2)flowers,involucrate; tillode sometimes present, filiform. Pistillate infloresperianth enlargedin fruit,fleshy,orange.Fruitbasally cences solitary or in pairs (or as many as 5 together on adnateto theperianth; cotyledonsequal. short-shoots) in the leaf axils, discoid-capitate, 0.3-0.4 cm diam., subsessile or with to 0.6 cm long peduncule, SectionOlmediais monotypic. involucrate; involucral bracts in 5-7 rows, narrowly to broadly ovate, 1-3 mm long, strigillose; flowers 1 (or 4. Trophiscaucana(Pittier)C. C. Berg,Proc.Kon. 2); perianth 3-3.5 mm long, dentate, strigillose; ovary Ned. Acad. Wetensch.,C. 91: 354. 1988. Olmedia free, partly strigillose; style 1-4 mm long; stigmas 71 1 mm long, minutely puberulous. Fruiting perianth caucana Pittier, Contr. U.S. Natl. Herb. 13: 434. 1912. Type. Colombia. Valle: La Manuelita, nr. fleshy, orange to red; fruit free, subglobose, ca. 4.5-5 Palmira, 5 Dec 1904-6 Jan 1905 (o), Pittier 803 mm; inner involucral bracts, enlarged, to 10 mm long, (holotype, US). Fig. 6 those of the innermost row ? fleshy, orange inside, OlmediaasperaRuiz & Pav6n,Syst. Veg. Fl. Peruv. spreading at fruit maturity.

Chil. 1: 257. 1798;Berg,Fl. Neotrop.Monogr.7: 14. 1972.OlmediapoeppigianaKlotzsch,Linnaea 20: 525. 1847, as a synonymof Olmediaaspera Poeppig & Endlicher,Nov. Gen. 2: 31. 1838, basedon Poeppigs.n. or 1267, Peru,withoutlocality (B), non 0. poeppigianaMartius,Flora(or Bot. Zeit) 24(Beibl. 2): 93. 1841 (= Helicostylis tomentosa(Poeppig& Endlicher)Rusby).Type. Peru. Withoutlocality, (v), Ruiz & Pav6n s.n. (lectotype,Berg,Fl. Neotrop.Monogr.7: 14. 1972, MA; syntype collections, c' and/or ?, in B, Fl, HAL,K, LE, P).

Distribution (see Fig. 73.3). From Nicaragua to Amazonian Bolivia; in wet forest, often along streams and in forest margins; to 1600(-2 100) m. Representative specimens examined. NICARAGUA. ZELAYA: Cafnoel Toro, 13 May 1978 (? fl-fr), Vincelli

384 (BG, MO). COSTA RICA. ALAJUELA: San Rafaelde Guatuso, Cueva de Venado, N of Volcin Arenal, 20 Aug 1963 (a'), JimenezM. 1091 (CR, F, NY). CARTAGO: Nr.Turrialba, 23 Jun 1949 (o), Holm et al. 94 (BM, NY, U), 26 Aug 1949 (e), Le6n 1749 (US) = Skutch 4668 (CR, MO, NA,

FLORANEOTROPICA

48

B

1

0~~~~~

Fig. 6. Tophas caucana. 1. Leafy twig with staminateinflorescences(Hartmann12064). 2. Leafy twig with pistillate inflorescences (Knappet al. 2400). 3a-c. Stamens.3d. Staminateflower. 4. Staminateflower in bud (Mexia8133). Sa. Fruit.Sb, Sc. Embryo(CampE-933). 6. Pistillateinflorescence.7a. Pistil. 7b. Pistillateflower (Pittier 12041). [By P. Pardoen,Utrecht.]

TAXONOMIC TREATMENT NY, US), 4 Feb 1940 (e), Skutch 4674 (CR, MO, NA, NY, US). LIM6N: Rio Madre de Dios, nr. Siquerres, 14 Feb 1977 (a), Burger et aL. 10518 (CR, F); Bajo Telire, Rio Telire, Jul 1984 (e), G6mez 24109 (BG, MO, NY). PUNTARENAS: 5 km W of Rinc6n de Osa, 24-30 Mar 1973 (a), Burger et al. 8920 (CR, F, MO, NY, U), (? fl), Burger et al. 8989 (CR, EAP, MO, NY, U) & 8997 (CR, F, U); Rio Naranjo, nr. Londres & Villa Nueva, 10-12 Feb 1988 (? fl), Burger et al. 12272 (F); Burica Peninsula, 3 km NW of Punta de Burica, 5 Mar 1973 (? fl), Busey 771 (F, MO, NY); Quepos, 11 Feb 1988 (? fl), 11 Feb 1988 (? fl), G6mez-Laurito 11693 (F). PANAMA. BOCASDELTORO:Susruba, 10 Sep 1963 (o'), Dwyer 4364A (MO, U); Puerto Palenque, 12 Apr 1968 (e), Kirkbride et al. 554 (MO, NY); nr. Chiriqui Lagoon, 20 Oct 1940 (? fl-fr), Wedel1043 & 1094 (MO), US); Shepherd Island, nr. Chiriqui Lagoon, 22 Sep 1941 (a'), Wedel 2736 (MICH, NY, US). CHIRIQUf:Burica Peninsula, 6 km W of Puerto Armuelles, 17 Feb 1973 (a'), Liesner 11 (MO); San Fe1ix, Dec 1911 (e'), Pittier 5291 (BM, NY, US); Paso de Quebrada Gata, between Hato del Jobo & Cerro Vaca, 25-28 Dec 1911 (? fl), Pittier 5425 (BM, C, K, NY, US). COCLE: Nr. Club Campestre, 2 Jul 1967 (d), Duke 13250 (MO, US); El Valle Ant6n, 19 Mar 1939 (2 fr), Allen 1736 (EAP, F, NY), 25 Aug 1971 (? fl), Lao 289 (MO); between El Valle & La Mesa, 15 Dec 1986 (? fl & a'), McPherson 10045 (BG, MO); Penonome, Rio Guayabo, above Vado, 2 Jul 1971 (a), Webster 16814 (MICH, MO). COL6N: Rio Mendosa, 10 km N of Gamboa, 19 Oct 1975 (e'), Dressler 5196 (F, MO, NY); Mindi Hills, 28 Mar 1956 (ae), Johnston 1733 (MICH, MO); Rio Boquer6n, NW of Mina Boquer6n, 12-13 Dec 1981 (? fl-fr), Knapp et al. 2400 (MO, U) & 2451 (MO, U); Rio Guanche, 1-4 km S of Portobelo Hwy., 11 Apr 1982 (? fr), Knapp et al. 4589 (MO, U); Gatun R., 16 Aug 1914 (? fl), Pittier 6747 (BM, MO, NY, US). DARIEN:LomaPiraque,1000 m, 14 Apr 1966 (? fl), Duke 8102 (MO), Rio Balsa,

between Manene& Tusijuanda,26 Jul 1967 (? fl-fr), Duke13546 (MO,US); 18 km SE of Jaque,10 Jan 1983 (? fl), Garwood et al. 79 (BG, F, MO) & 129 (BG); Rio Paraseneco, 2-3 km SE of Pijibasal, 18 Dec 1980 (e'), Hartman 12064 (MO, U); Rio Balsas, just above Rio Coasi, 2 Jan 1981 (o), Hartman et al. 12513 (MO, U); Serrania de Pirre, 10 km S of El Real, 29 Jan 1979 (Q fl), Reveal et al. 4898 (MO). PANAMA: Serraniade Maje, trail along Rio Ipeti Grande, between Rio Charco & confluence with Rio Agua Fria, 28 Jan 1984 (a'), Churchill et al. 4452 (BG, MO); Barro Colorado Island 9 Apr 1967 (? fl), Croat 4597 (MO); Quebradala Palma, 9 Jan 1935 (a'), Dodge et al. 17365 (BM, C, EAP, Fl, K, MICH, MO, P, S, U); Serrania de Maje, 10 Feb 1977 (? fl-fr), Folsom et al. 1713 (MO, U); Cerro Galera, 15 Feb 1983 (? fr), Hamilton et al. 2976 (BG); nr. Gorgona, 10 Feb 1911 (? fl-fr), Maxon 4744 (BM, C, NY, US); Rio Charare, nr. Chepo, 29-30 Oct 1911 (e'), Pittier 4707 BM, C, F, K, NY, US). COLOMBIA. ANTIOQUIA: Mun. Turbo, rd. Currulao-Nueva Antioquia, 11 km, 3 Aug 1987 (9 flfr), Callejas et al. 4979 (BG, MO); Mun. Mutata, Aguas

49

Claras, 3.5 km N of Mutata, 20 Nov 1987 (2 fl-fr), Callejas 5705 (BG); Mun. Pueblo Rico, Vrda.Sevilla, 1450 m, 28 Sep 1989 (d'), Callejas et al. 8412 (BG);

Rio Le6n,VillaArtega,Las Caucheras,21 Oct 1961(o), Cuatrecasas et al. 26164 (COL, US); Heliconia, 28 Jun

1947(st),Daniel3978 (US);QuebradaPita,23 Apr 1946 (? fl-fr), Haught 4808 (COL, US); Uraba, Jan 1947 (C), Uribe-Uribe 1573 (COL). CAUCA: Santander de

Quichao, 1300 m, 5 Oct 1954 (st), Fernandez 2778 (COL).CHOC6:Mun.Acandi,QuebradaZardi,22 May 1989 (a), Betancur 1196 (MO); Mun. Riosucio, Peye, Rio Peye, 5 Jun 1976 (e), Forero et al. 1863 (MO, NY);

between mouthof Rio Matata& Alto de Buey, 9 Aug 1976 (? fr), Gentryet al. 17457 (MO, U); Cupica, 1011 Feb 1947 (a), Haught 5555 (COL, US), (? fl) 5564

(COL,US); QuebradaLa Platanilla,N of Rio Mecana, 10 Jan 1984 (e), Juncosa 1851 (BG, MO). HUILA: QuebradaGuache,3 km SW of Acevedo, 15 Aug 1944 (? fl-fr),Little8472 (COL,US) & 8481 (COL,US); rd. Pitalito-SanAugustin,km 9, 17 Aug 1974 (? fr), Plowman et al. 4174 (P). PUTUMAYO: Rio San Miguel, Quebrada de la Hormiga, 16 Dec 1940 (? fl-fr), Cuatrecasas 11099 (COL,US). RISARALDA: Rio Cauca, Paso de la Virginiana,900-1300 m, Aug 1884 (2 fl), Lehmann 3873 (K, US). VALLE: Rio Pichinde, Pefias Blancas,23 Apr 1978(? fl), Benalcazaret al. 220 (MO); Mun. Tulua, Mateguada, 1100 m, 23 Jan 1979 (e'), CuadrosV 597 (BG, MO);Mun.Yotoco,Lago Calima, 1500 m, 12 Oct 1980 (a), Cuadros V 1229 (BG, MO);

Mun. Sevilla, La Estella, 1250 m, 29 Dec 1980 (e'), CuadrosV 1309 (MO);Mun. Zarzal,between Caserio La Paila & Zarzal, 1000 m, 6 Apr 1986, Gentryet al. 54147 (BG);Mun.Riofrio,rd. to Sol6nici,26 May 1982 (? fl), Millan 50 (MO);Palmira,1000 m, 27 Jun 1982 (a'), Murphy et al. 627 (MO,NY, U). ECUADOR. ESMERALDAS: Timbre,4 Mar 1953 (? fl), Fagerlind et al. 2528 (S); nr. Lita, 8 Jun 1978 (e), Madison et al. 5405 (QCA, U). GUAYAS:Between Las Americanas & Daule, 30 Sep 1952 (e'), Fagerlind et al.

316 (S); ca. 13 km S of Naranjal,24 May 1980 (o), Harling et al. 19387 (BG, GB). IMBABURA: Rio Mira, by rd. ca. 110 km NNW of Ibarra,14 Oct 1982 (a), Pennington et al. 10526 (QCA, QCNE, U). Los Rios:

Cant6nVinces, Jaunecheforest, between Mocachi & Palenque,EsteroPeinafiel,8 May 1981 (st), Dodson et al. 10600 (MO); Rio PalenqueBiological Station,rd. Quevedo-SantoDomingode los Colorados,km 56, 26 Feb 1974(? fl), Gentry 10202 (MO,QCA,S). MANABt: Balao, Jan 1892 (e'), Eggers 14359 (US); rd. Portoviejo-

Pichincha,11 km E of Placido,8 May 1985(a), Harling et al. 24985 (GB); EsteroSegel, S of Olmedo,28 Sep 1942 (? fl), Haught 3487 (S, US). MORONA-SANTIAGO: Macas,Rio Upano,18 Mar1956(? fl-fr),Asplund19818 (NY, S), (a),

Asplund 19819 (NY, S); Taisha, Rio

Guambime,19 Jun 1980 (? fl), Brandbygeet al. 32071 (AAU); nr. Mendez, 8 Nov 1944 (a), Camp E-933 (NY,

US), 12 Nov 1944 (? fl-fr), CampE-951 (NY); Taisha, Rio Guagauyme,29 Jan 1962 (? fl-fr), Cazalet et al. 7560 (K, NY, US); Bomboiza, 4 Nov 1986 (a), Cer6n

et al. 452 (MO,QAME,QCA,QCNE);betweenMendez

50 & Bella Uni6n, 1 Apr 1974 (q fr), Harlinget al. 13049 (BG,GB);betweenGualaquiza& NuevaTarquiat bridge over Rio Cuchupampa,18 Apr 1985 (? fl-fr), Harling et al. 24269 (BG, GB, QCA). NAPO: BetweenTena& Napo, 7 Oct 1939 (a), Asplund 9149 (NY, S); Reserva Biol6gica Jatun Sacha, 24 Aug 1988 (d'), Cer6n et al. 4523 (NY,QCNE);8 km SE of Tena,30 Aug 1960 (e'),

Grubbet al. 1539 (K, NY); Rio Napo, 6 km below Yuralpa,15 Aug 1958 (R fr),Harling3617 (NY,S); Rio Hollin, ca. 10 km E of Archidona,30 Mar 1969 (e), LugoS. 934 (BG,GB);ParqueNacionalYasuni,nr.mouth of RioYasuni,18-28 Feb 1983(? fl), Luteynet al. 9116 (QCA); nr. Puerto Misahualli, 19 Nov 1985 ( fil), Palacios et al. 949 (BG). PASTAZA: Rio Chullana,ca. 8-15 km N of Puerto Sarayacu,16-19 Oct 1974 (a), LugoS. 4174 & 4230 (BG, GB);Rio Bobonaza,4 km E of Pacayacu, 31 Oct 1974 (a), Lugo S. 4402 (BG, GB);

3-4 km E of Puyopungo,28 Oct 1976 (? fl), Lugo S. 5022 (BG,GB);Rio Bobonaza,betweenCampChichirota CaboPozo, 20 Jul 1980 (a), Oellgaard & Destacamento et al. 34866 (AAU);Rio Pastaza,betweenDestacamento Chiriboga& ApachiEntza,24 Jul 1980 (a), Oellgaard et al. 35152 & 35179 (AAU). SUCUMBIOS:RioAguarico, ca. 76?0'W,0?20-25'S, 27 Jul 1981 (Y fl), Berg et al. 1031 (AAU, MO, US); Rio Cuyabeno,ca. 75055'W0?10'S,17 Feb 1980 (? fl), Berg et al. 1047 (AAU,NY, U); San Pablo de los Secoyas, 11 Aug 1981 (a'), Brandbyge et al. 33419 (AAU); Dureno, 11 Mar-I Apr 1986 (? fl), Cer6n262 (BG, QCA),20-31 Dec 1987 (? fl-fr),Cer6net al. 3114 (MO,NY,QAME,QCA,QCNE); Rio Pucino,tributaryof Rio Aguarico,nr. LagoAgrio, 9 Feb 1974 (? fr), Gentry9808 (QCA,US). ZAMORACHINCHIPE: Between Zamora& Zumba,Rio Jamboe, 1000-1200 m, 23 Apr 1974 (a'), Harling et al. 13878

(BG,GB);36 km NE of Zamora,17 Sep 1975(a'), Little et al. 413 (NY,US); Rio Nangaritza,1 kmupstreamfrom Shaime, 16 Feb 1994 (? fl), Werif et al. 13127 (BG). PERU. Without locality, (a'), Poeppig 1267 (B, BM, HAL, L, LE, P), (v), Poeppig 1267 (LE). AMAZONAS: Prov. Bagua, 5 km above La Peca, 2050-2160 m, 16 Apr 1984 (st), Croat 58355 (MO); Quebrada Aintami, 12 Jan 1973 (a'), Kayap 125 (MO, NY); Rio Santiago, nr. La Poza, 12 Nov 1979 (st), Tunqui 4 (MO). AYACUCHO:Prov. La Mar, 5 km W of Santa Rosa, 22 Sep 1976 (? fr), Wasshausen et al. 693 (MO, NY). Cuzco: Prov. Paucartambo,between Atalaya & Pilcopata, 31 Jul 1988 (a'), Berg et al. 1616 (AAU, BG, COL); Prov. La Convenci6n, N of Rio Apurimac & Hda. Luisiana, 31 Jul 1968 (? fl), Dudley 11494 (MO); Prov. Quispichanchis, Rio Araza, between Pande & Quince Mil, 10 Aug 1991 (a'), Nuniez 14128 (BG). HUANUCO: Tingo Maria, 1 Sep 1940 (a), Asplund 13420 (NY, P); between Venenillo & Aucayacu, 31 Jul 1948 (a'), Ferreyra 4361 (US); between La Merced & Huanuco, 2 Aug 1948 (? fl-fr), Ferreyra4373 (US); nr. Tingo Maria,rd. to Monz6n, 13 Oct 1957 (? fr), Ferreyra 12669 (US); nr. Tingo Maria, Cueva de los Lechusas, 9 Aug 1980 (9 fr), Gentry et al. 29540 (MO, U); Dtto. Churubamba,Rio Ysabel, Hda. Exito, 5 Sep 1936 (a), Mexia 8133 (GB, K, MO, NY, S, U, US); Prov. Leoncio Prado, Dtto. Rupa Rupa,

FLORANEOTROPICA Rio Monzon, 14 Nov 1971 (? fr), Schunke V 5150 (NY). JUNIN: Rio Tulumayo, 10 km S of San Ram6n, 5 Jun 1983 (? fl-fr), Gentry et al. 41522 (BG, MO); La Merced, 29 May-4 Jun 1929 (? fl), Killip et al. 23464 (F, NY, US) & 23816 (NY, US); Colonia Perene, 14-25 Jun 1929 (e'), Killip et al. 25057 (NY, US); Rio Huallaga, Juanjui, Sep 1934 (ei), Klug 3787 (BM, K, MO, NY, S, US); Prov. Satipo, Dtto. Satipo, May 1982 (? fl), Rodriguez 488 (U); Hda. Genova, 1600 m, 13 Jul 1962 (? fl-fr), Woytkowski 7374 (GH, MO). LORETO: Prov. Alto Amazonas, Rio Pastaza, Andoas, 17 Nov 1979 (st), Gentry et al. 28145 (U). LIBERTAD: Prov. Pataz, Rio Mishiollo, Ong6n, 1300 m, Aug 1914 (a'), Weberbauer 7059 (US). PASCO: Prov. Oxapampa, 12 km S of Villa Rica, 1340 m, 18 Jan 1983 (? fr), D. N. Smith et al. 3125 (BG, MO, NY). SAN MARTIN: Rio Shilcayo, N of Tarapoto, 27 May 1986 (? fr), Alcorn et al. 17 (MO); Prov. Mariscal Caceres, Dtto. Tocache Nuevo, E of Punte, 26 Jul 1974 (? fl-fr), Schunke V 7776 (MO, NY, U); Dtto. Tocache Nuevo, Quebrada de Canuto, 4 Jan 1979 (o), Schunke V 10670 (BG, MO); Tarapoto, 1855 (ei), Spruce 4034 (BM, C, K, LE, NY, P), Sep 1902 (od), Ule 6412 (MG, K, L, RB); San Roque, 1350-1500 m, 9 Jan 1930 (? fl-fr), Ll. Williams 7195 (F, US). UCAYALI: Prov. Coronel Portillo, rd. Tingo Maria-Pucallpa, 75 km NE of Tingo Maria, 25 Nov 1975 (? fr), Davidson 3679 (MO); Contamana, 28 Jul 1970 (o), McDaniel et al. 2575 (US); rd. Pucallpa-Tingo Maria, km 86, 5 Apr 1982 (? fl-fr), D. N. Smith et al. 1213 (BG, MO, NY). BOLIVIA. BENI: Prov. Ballivian, La Embocada, ca. 25 km SW of San Borja, 20 Jul 1981 (d'), Beck 6903 (BG); Prov.Yacuma,SE of San Borja, 46 km from Rio Maniqui, 14 Sep 1988 (v), Beck 16744 (BG); Prov. Ballivian, rd. Caranavi-SanBorja, Serraniadel Pil6n Lajas, 1 Nov 1989 (? fl-fr), D. N. Smith et al. 13810 (BG, MO). BENI/LA PAZ: Rio Beni, upstream from Rurrenabaque,Rio Tuichi, 13 May 1990 (v), Daly et al. 6438 (BG). COCHABAMBA: Prov. Chapare, Todos Santos, 26 Oct 1966 (o), R. F Steinbach 442 (F, MO, NY, U, US). LA PAZ: Prov. Sud Yungas, nr. Tucupi, ca. 30 km SE of Palos Blancos, nr. Rio Beni, 5 Mar 1994 (? fl-fr), Berg 1717 (BG, LPB); rd. Caranavi-Guanay,27.8 km N of Caranavi, 28 Nov 1980 (? fl-fr), Croat 5164 (MO); Prov. Sud Yungas, Rio Bopi, San Bartolome, nr. Calisaya, 1-22 Jul 1939 (ci), Krukoff 10135 (K, MICH, MO, NY, S, U, US), Krukoff 10277 (MICH, NY, U), Krukoff 10388 (K, MICH, MO, NY, S, U, US); cataracts of Rio Bopi, Sep 1921 (? fl-fr), Rusby 564 (K, NY, US); Prov. Sud Yungas, Alto Beni, Sapecho, 30 Nov 1988 (? fl-fr), E. Vargas et al. 2083 (BG); Rio Beni, Huachi (= Huanay?), 4 Sep 1921 (ad), White 1011 (MO, NY, US). SANTA CRUZ: Prov. Ichilo, 4 km SW of Buena Vista, 3 Oct 1990 (? fl), Nee 39087 (MO, NY); Prov. Ichilo, Parque Nacional Amboro, Rio Isama, 4 km NNE of summit of Cerro Ambor6, 10 Oct 1990 (ei), Nee 39208 (MO, NY); Prov. Ichilo, ParqueNacional Ambor6, Rio Surutu, 30 Aug 1985 (? fl), Solomon 14175 (BG, MO); Prov. Ichilo, 30 km SE of Buena Vista, 2 Sep 1985 (? fl), Solomon 14224 (BG, MO); Prov. Sara, Bosque del Fraile, 4 Mar 1926 (ei), J. Steinbach 7407 (BM, F, K, MO, NY, U).

51

TAXONOMIC TREATMENT

Specimens examined. COSTA RICA. ALAJUELA: 1 km E of Florencia, Rio Peje, 27 Jun 1985 (st), Hammel et al. 14082 (MO); Upala, Colonia Libertad, 2 May 1988 Santa (? fl-fr), Herrera 1934 (BG, MO). GUANACASTE: 1980 2 Jul Liberia, NW of Park, 30 km Rosa National Berg, C. C. 2C. Trophis P.Browne sect. Echinocarpa Proc.Kon.Ned. Akad.Wetensch.,Ser.C, Biol. Med. (st), Janzen 11912 (MO). HEREDIA:Rio Sarapiqui, nr. Tirimbina, E of Re, 12-15 Aug 1971 (o), Burger et al. Sci. 91: 353. 1988. Type. T involucrataW. Burger. 8081 (CR, F); 2.5 mi E of Puerto Viejo, 11 Mar 1983 (? Shrubsor treelets. Staminateinflorescences race- fr), A. Chacon 401 (NY); Cant6n Sarapiqui, Finca El Bejuco, 30 Jan 1990 (e'), Chac6n 703 (MO); Finca La mose; tepals 4, valvate in the bud. Pistillate infloresSelva, nr. Puerto Viejo, 15 Jan 1980 (a'), Denslow 79cences subcapitate,with 2-3 flowers, subinvolucrate; 125 (BG), 14 Mar 1972 (a), Frankie632 (F), 8 Feb 1981 tepalsnot enlargedin the bud,not fleshy,greenish.Fruit (? fr), Folsom et al. 8829 (MO), 10 Feb 1981 (v), free, drupaceous,darkblue to blackish,often echinate; Folsom 8902 (F), 13 Feb 1981 (? fl-fr), Folsom 8911 cotyledons slightly unequal. (NY), 26 Feb 1981 (? fr), Folsom 9130 (MICH, MO, NY); S of PuertoViejo, 2 km S of Magsasay Penal Colony, Section Echinocarpa contains only Trophis 5 Feb 1983 (st), Garwood et al. 1124 (BG, MO); Finca involucrata. La Selva, 18 Feb 1980 ($ fl-fr), Grayum 2629 (BG, F, U); Cerros Sardinal, ca. 2-2.5 km N of Chilimate de Sarapiqui, 21 Jan 1986 (a), Grayumet al. 6167 (MO); 5. Trophis involucrata W.Burger,Phytologia26: 432. Finca La Selva, nr. Puerto Viejo, 14 Mar 1980 (a'), 1973; Burger,FieldianaBot. 40: 211, t. 14. 1977. Hammel 8080 (F, MICH, MO), 18 Mar 1980 (? fr), Type.CostaRica. Heredia:FincaLa Selva, 3 km SE Hammel 8129 (F); Cant6n Sarapiqui, Estaci6n Magasay, Finca La of PuertoViejo, 19 Jan 1973 (d), Opler 1657 (ho- 18 Jul 1990 (a'), Hammelet al. 17828 (BG); Selva, 16-17 Jan 1973 (a'), Hartshorn 1091 (CR, MO), Fig. 7 lotype, F; isotypes, MO, U, US). 17 Jan 1973 (? fl), Hartshorn 1094 (CR, F, MO, PMA, 20 Dec 1973 (a'), Hartshorn 1323 (BG, F, MO, U), U), Shrubor treelet,to S m tall. Leafy twigs 1-2.5 mm 20 Mar 1974 (? fr), Hartshorn 1409 (F, MO, U); Rio thick,puberulousto hirtellous.Laminaoblongto ellipSarapiqui,Tirimbina,IstariiFarm, 26 Nov 1971 (a), Lent tic or to lanceolate,3.5-21 x 1.5-8 cm, broadestat or 2243 (BM, F, NY, PMA, U, US); Finca La Selva, 30 Mar above, or sometimesbelow the middle, almostequilat- 1956 (? fl-fr), Schubertet al. 1310 (GH). LIM6N: Rio to subcoriaceous; apex(caudate-)acumi- Sixaola, Bribri, 7-9 Sep 1978 (a'), Burgeret al. 10886 eral,chartaceous (to (F, NY); La Amistad, Quebrada Crori, Crorinia,20 Jul nate;base obtuseto subacute;marginserrate-dentate subentire);uppersurfaceglabrous;lower surfaceoften 1989 (a'), A. Chacon 243 (BG, MO); 7 km SW of Bribri, scabridulous,puberulousto hirtellousto strigilloseon 4 May 1983 (? fl), G6mez et al. 20470 (BG, MO), (a'), the veins; venationplane to slightly prominentabove, Gomez et al. 20467 (BG, MO); between Rio Chirrip6 & slightlyprominentbeneath;lateralveins (4-)7-l 0 pairs; Quebrada El Molinete, 19 Jul 1984 (a'), Grayum et al. 3551 (BG, MO); Rio Barbilla, SE of Siquirres, 11 Jan tertiaryvenationreticulate;petiole0.2-0.6 cm long,0.51987 (a'), Grayumet al. 7948 (MO); Cordillera de 2 mm thick, puberulousto hirtellous;stipules 0.1-0.2 Talamanca,Cant6n Matina, Cerro Amu, 1 Nov 1988 (a), cm long,puberulous,caducous.Staminateinflorescences Herrera Ch. et al. 2265 (BG, F, MO); CantonLim6n, Cerro solitaryin the leaf axils, racemose;peduncleca. 0.4(- Muchilla, Fila Matama, 7 Apr 1989 (? fr), Robles et al. 0.8) cm long, puberulous;raceme0.2-2.5(-5) cm long, 2693 (BG), 14 Apr 1989 (? fl), Robles 2773 (BG). SAN flowersdistant,pedicellate;pedicel 1-3 mm long;peri- Jost: ParqueNacional Braulio Carrillo, Rio Corinto, 16 anthca. 2 mm long, 4-fid to -parted,puberulous;sta- Aug 1984 (a), Sdnchezet al. 563 (BG, MO). PANAMA. BOCAS DEL TORO: Above Chiriqui mens 4; filamentsca. 3 mm long; anthersca. 0.7-0.8 x 26 Dec 1986 (? fr), McPherson et al. 10157 Grande, ca. 0.5 mm long, 0.6-0.7 mm, pistillode quadrangular, (BG, MO).

Local names. Colombia:Valle:borejore.Peru:San Martin:ishangamasha,mashonastillo.

puberulous;bractsnarrowlyovateto subulate,to 0.5 mm long, puberulous.Pistillate inflorescencessolitaryor in pairsin the leaf axils, uniflorousor subcapitatewith 23 flowers; peduncle 0.1-0.4 cm long, to 5 cm long in fruit, puberulous;flowers sessile; perianth4-fid to 4parted,ca. 1mm long,puberulous;ovaryca. 3 mm long, puberulous;stigmas 3-4 mm long, puberulous;bracts ovate, to 0.5 mm long, puberulous.Fruit ellipsoid to ovoid, 1.2-1.5 x 0.8-1.1 cm, blue-maroonto blackish, puberulous,with a few to numerousprickle-like,to 1.5 mm long protuberancesor smooth. Distribution (see Fig. 73.1). CostaRica andwestem Panama;in wet forest; at low elevations.

EXCLUDED NAMES

Trophisglabrata Liebmann,Kongel. DanskeVidensk. Selsk.Naturvidensk.MathAfh.,Ser.5, 2: 314. 1851. Type. Mexico. Veracruz:Palenque, nr. Misantla, Liebmann14174 (lectotype, Berg, Novon 6: 241. 1966, C). = Pseudolmediaglabrata (Liebmann) C. C. Berg, Novon 6: 241. 1996, to replacePseudolmediaoxyphyllariaDonnell Smith(cf. Berg, 1972). Nov. Strip.Bras.Dec.,Decas TrophishilarianaCasaretto, IX. 80. 1845. = SoroceaguilleminianaGaudichaud.

FLORANEOTROPICA

52

0~~~~~~~~~~~

Fig. 7. Trophisinvolucrata.1. Leafy twig with staminateinflorescences(Lent 2243). 2. Leafy twig with young staminateinflorescences(Hartshorn1323). 3. Leafy twig with infructescencees(Grayum2629). 4. Staminate flower and bract. 5. Stamen and pistillode (Lent 2243). 6, 6a. Pistillate flower (Frankie 632). 7. Young infructescence.8. Infructescence(Grayum2629). 9. Infructescence.10. Seeds. 11. Embryo(Hartshorn1409). [By P. Pardoen,Utrecht.]

TAXONOMICTREATMENT Trophislaurifolia Willdenow, Sp. PI. 4: 733. 1806 Styloceras laurifoliumKunth(Asteraceae). Trophis macrostachya Donnell Smith, Bot. Gaz. (Crawfordsville)40: 10. 1905. = Soroceapubivena Hemsley subsp.pubivena.

Maclura 3. Maclura Nuttall, Gen. N. Amer. P1.2: 233. 1818, nom. conserv.; Miquel, in Martius,Fl. Bras. 4(l): 153. 1853; Bureau,in De Candolle,Prodr.17: 226. 1873;Bentham& Hooker,Gen.P1.3(1): 363. 1880; Berg, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med. Sci. 89: 241. 1986.Type.Macluraaurantiaca Nuttall(= Maclurapomifera (Rafinesque)C. K. Schneider). VanieriaLoureiro,Fl. Cochinchin.564. 1790. Type. Vanieracochinchinensis Loureiro(= Maclura cochinchinensis(Loureiro)Corner). foxilon Rafinesque,Amer. Monthly Mag. & Crit. Rev. 2: 118. 1817, nom. rejic. Type. Ioxilon pomiferum Rafinesque (= Maclura pomifera (Rafinesque)C. K. Schneider). ChlorophoraGaudichaud,Voy. Uranie 509. 1830; Kaastra,Acta Bot. Neerl. 21: 657. 1972. Type. Chlorophoratinctoria(Linnaeus)B. D. Jackson (= Macluratinctoria(Linnaeus)Steudel). CudraniaTrecul,Ann. Sci. Nat. Bot., S&. 3, 8: 123. 1847, nom. conserv.Type(conserved).Cudrania javanensis Trecul (= Maclura cochinchinensis (Loureiro)Corner). Plecospermum Trecul,Ann. Sci. Nat. Bot., Ser.3, 8: 124. 1847. Type.Plecospermum spinosumTrecul (= Macluraspinosa (Willdenow)C. C. Berg). CardiogyneBureau,in De Candolle,Prodr.17: 232. 1873. Type. Cardiogyne africana Bureau (= Macluraafricana (Bureau)Corner). Treesor climbers,dioecious, often armedwith axillary thorns;scale-leaves presentat the base of lateral branches;uncinatehairspresentor absent.Leaves altemate and distichous;laminapinnatelyveined; margin subentire to dentate or lobate; stipules fused, semiamplexicaul.Inflorescences (usually) solitaryin the leaf axils or on short-shoots,pedunculate,bracteate; embeddedyellow dye-containing "glands"in the bracts and/or the (fruiting) perianths often present. Staminateinflorescencesglobose-(sub)capitateor spicate; flowers pedicellate or sessile; tepals almost free or connate; pistillode flat. Pistillate inflorescences tepalsfreeor connate,decussate, (sub)globose-capitate; thickened at the apex; ovary free; stigmas 1, or if 2, thenunequallylong, filiform.Fruitingperianth? fleshy, greenish; fruit free; endocarpcrustaceous;seed with

53 scarceendosperm;testawith a thickenedvascularized partbelow the hilum;embryocurvedor longitudinally aligned;cotyledons equal andflat;radicleratherlong. The most common neotropicalspecies of Maclura was decribedas Morus tinctoria by Linnaeus(1753). Combinationswere made with Broussonetia (Kunth, 1817), Fusticus (Rafinesque, 1838), and Maclura (Steudel,1841).In 1830 GaudichaudestablishedthegenusChlorophora,based on Morustinctoria.The combination,Chlorophoratinctoria,wasmadeby B. D. Jackson in 1893, and that name was commonly used for a long time. However, Miquel (1853) continuedto use the genus nameMaclurain his treatmentof the "Moraceae"in FloraBrasiliensis. He recognized seven neotropicalspecies, among them some based on taxa describedby Linnaeus(1759) andLiebmann(1851), M. brasiliensis, describedby Martius(1841) asBroussonetia brasiliensis, and threenew species. With the exception of M brasiliensis, Bureau(1873) included all these species, as well as a species decribed by Blume (1856) in Chlorophora tinctoria and recognized 10 infraspecifictaxain this species;moreof suchtaxawere publishedby Hassler (1919). Bureaualso transferred the AfricanMorusexcelsa Welwitschto Chlorophora. Since Bureau'srevision, two additionalspecies were decribedin Chlorophora(Fernald, 1904; Standley & L. 0. Williams, 1950). Grisebach (1879) decribed Maclura mora, which was transferredby Lillo (1910) to Chlorophoraand subsequently(1925) reducedto a variety of C. tinctoria. Macbride (1937) transferred Maclurabrasiliensisto Chlorophora.In 1972 Kaastra revisedthe genus Chlorophora.He acceptedC. tinctoria, with two subspecies,andexcluded C. brasiliensis, as a species underMaclura. In 1962 Cornerreduced Chlorophorato a section of the broadlyconstruedgenus Maclura, in which he also included the African genus Cardiogyne and the Asian genus Cudrania.Corner'sbroadconcept of the genus Maclura was reconsideredby Berg (1986). He included the Asian Plecospermum in the genus, but excluded three African taxa referredto Maclura by Corner (1962). One of them was transferredto the paleotropicalgenusBroussonetiaandtwo of themwere transferredto the Africangenus Milicia. In the present concept the genus comprises: 1. section Maclura, with only M. pomifera (Rafinesque)C. K. Schneiderin NorthAmerica; 2. section Cardiogyne (Bureau) Corner, with M. africana (Bureau)Cornerin EastAfrica and M. brasiliensis intropicalAmerica; 3. section Cudrania (Trecul) Coiner, with five species in Asia-Australasia;

54

FLORANEOTROPICA

areformednextarealternate.Theaxilarybudsof the two speciesin Asia;and oppositeandalternateleaveson the mainaxis form 5. section Chlorophora(Gaudichaud)Baillon, (sylleptically) beyond withtheleavesalternate branches withonlyM.tinctoriaintropicalAmerica. axillary budsonthesebranches theprophylls. Theprimary maydevelopintothorns,whereasaccessorybudsdeIt is remarkablethatMaclura brasiliensis is morvelopintonormalbrancheswithleaves. phologicallycloserto theAfricanspeciesM africana Thestaminate inflorescences areglobose(orellipthanto M tinctoriaandtheNorthAmericanspecies (suchas in M. in most Maclura species soid) heads M pomifera. 4. section Plecospermum(Trecul)C. C. Berg, with

MostMacluraspeciesareclimbers,a habit,however,not exhibitedby M. tinctoria.All specieshave straightorcurvedthoms,thesebeingmodifiedlateral InM.brasiliensistheyarecurved,always branchlets. present,andoftenfunctionas organsforclimbing.In M.tinctoriathethomsarestraight,usuallyoccurring on youngtrees,less so or evenabsenton abundantly adulttrees. InMacluratinctoriathereareusuallytwopairsof oppositeleavesafterthecotyledons.Theleaveswhich

brasiliensis), but they are spicate in M. tinctoria and

racemoseinM.pomifera.Thepistillateinflorescences areglobose-to ellipsoid-capitate inallspecies.Thespecies havealsoin commonthereductionof one of the two stigmas,andthe presenceof "glands"(pockets) in tepalsandinterfloral bractsin whichyelembedded These"glands" aremostapparlowdyeis accumulated. in entin thefruitingstate.Thesubstanceaccumulated thetepalsandbractsmightplayarolein theprotection in thebark. andmayalsoaccumulate againstherbivory

Key to the neotropicalspeciesof Maclura 1. Climbers;thornscurved;both staminateand pistillateinflorescencesglobose-capitate;infructes1. M. brasiliensis cences 4-6 cm diam............................................ 1. Trees or shrubs;thornsstraight;staminateinflorescencesspicate,the pistillateinflorescences 2. M. tinctoria ........................................... globose- to ellipsoid-capitate;infructescences1-2 cm diam.

3A. Maclura Nuttal sect. Cardiogyne (Bureau)Corner, Gard. Bull. Singapore 19: 237. 1962. Type. Cardiogyneafricana Bureau(= Maclura africana (Bureau)Corner). Climbers.Staminateinflorescences globose-capitate;flowers sessile or subsessile; stamensinflexed in the bud. Pistillateflowers free; tepals and interfloral bractsaccumulatingyellow dye.

IoxylonpomiferumRafinesquevar.glaberrimum0. Kuntze, Rev. Gen. P1. 3(3): 294. 1898. Type. Bolivia. SantaCruz:Rio Yapacani,Jun 1892 (? fl-fr), Kuntzes.n. (holotype,NY). Chlorophorascandens Standley& L. 0. Williams, Ceiba 1: 77. 1950. Type. Honduras.Olancho: Catacamas,18-26 Mar 1949 (? fl-fr), Standley 18431 (holotype, F; isotypes, A, EAP,US).

Climber,to ca. 25 m tall (?), scrambler,shrub, or tree, especially if climbing, then with axillarystraight or recurvedthoms, to 8 cm long, shining at the tip and bearingscale leaves and/ornonnal leaves. Leafytwigs 1-3 mm thick, almost glabrous or densely white to yellowish-hirtellous, with straight to curved hairs. 1. Maclura brasiliensis (Martius)Endlicher,Gen. P1. Lamina elliptic to oblong, (0.5-)5-10 x (0.3-)3-4.5 Suppl.4(2): 34. 1848; Miquel,in Martius,Fl. Bras. cm, mostly broadest below the middle, equilateral, 4(1): 158,t. 54. 1853;Bureau,in De Candolle,Prodr. chartaceous;apex acuminate,the acumenmucronate; 17:231. 1873;Kaastra,ActaBot.Neerl. 22: 71, t. 1. base acuteto rounded(to truncate);margin(sub)entire; 1973. Broussonetia brasiliensis Martius, Flora uppersurfaceglabrousor sparselypuberulous,partly (Regensb.)24(Beibl): 10. 1841;Miquel,in Martius, with uncinate hairs; lower surface sparsely minutely Fl. Bras. 4(1): 158. 1853. Chloroporabrasiliensis puberulous or rather densely hirtellous, partly with (Martius) Macbride,Publ. Field Mus. Nat. Hist., uncinatehairs;venationnearlyplane above, the midrib Bot. Ser. 13(2.2): 310. 1937. Type. Brazil. Rio de prominent,the otherveins slightly prominentto plane Janeiro:Campos,(o), Prince Maxim.Neuwieds.n. beneath;lateralveins 6-12 pairs;tertiaryvenationre(holotype,BR; isotype, U). Fig. 8 ticulate;petiole 0.5-2 cm long, glabrousor hirtellous,

The section comprises two species, Maclura brasiliensis and M. africana from East Africa and Madagascar.

TAXONOMIC TREATMENT

55

4

4

s}

e

Fig. 8. Maclura brasiliensis. 1. Leafy twig pistillate inflorescence (Steyermark61949). 2. Leafy twig with staminate inflorescences. 3. Leafy twig (LI. Williams 6855). 4. Leafy twigs (Standley 18431). 5. Infructescence (Foster 5616). 6. Staminate flower and bracts (LI. Williams 6855). 7, 7a. Seed. 8, 8a. Embryo (Foster 5616). [By P. Pardoen, Utrecht.]

56

FLORA NEOTROPICA

withcurvedhairs;stipulesfused,reniform tocordiform, 17 Nov 1990 (? fr), Nee 39985 (NY,USZ);Prov.Nuflo 0.1-0.3 mm long, brownish-puberulous, (sub)per- de Chavez.rd. Colonia Okinawa1-San Ram6n,2 km sistent.Staminateinflorescences globose,0.8-1.2 cm SW of Los Troncos, 2 Dec 1990 (? fl-fr), Nee et al. diam.;peduncle 0.8-2cmlong,0.5-1mmthick,minutely 40155 (BG, NY); Prov.Andres Ibaiiez, PuertoPailas, 17 Dec 1991 (? fr),Nee 42145 (US, USZ);Prov.Andres flowersto 1mmlongpedicellate; puberulous; perianth Ibafiez,10 km ESEof Don Lorenzo,20 Dec 1992 (? fl1.5-2.2mmlong,4-parted, tomentellous; filaments 2.5- fr),Nee et al. 43184 (NY);Prov.Andr6sIbainez,4 km S 4 mmlong;anthers ca.0.7-0.8x 0.7-0.8mm;pistillode of Rio GrandePlantade Gas, 3 Jan 1994 (st), Nee et al. flat,0.3-0.5mmlong,theapexuncinate; bractsoblong 44649 (NY); Prov.Ichilo, 2 km WSWof El Hondo, 15 to almostlinearorto subspathulate, oftensubcucullate, Feb 1994 (? fl-fr), Nee et al. 45025 (NY, US); Prov. toshortlyvelutinous. Pistillate puberulous inflorescences Wames,PuertoMacdonald,7 Jan 1996 (c? + separatee' intheleafaxilsoron short-shoots; peduncle2.5-3 cm fr), Nee 46733 (NY). orsparselypuberulous; In the materialfromHonduras,the laminabeneath, long,hirtellous totomentellous 4-5 mm thepetiole,the leafytwigs, andthepedunclearedensely head(sub)globose, ca. 1.5cmdiam.;perianth long,4-parted,withembedded yellowdye-containing hirtellousto tomentose,whereasthesepartsaresparsely thelobes? cucullate,intheupperpartthick- puberulous to almost glabrous elsewhere. Mature "glands," to tomentellous; ened,short-velutinous ovaryfree,ca. infructescencesarereportedto smell like rottenoranges. 1mmlong;stylelacking,thelongerstigma(10-)15Localnames.Bolivia:SantaCruz:bejucomora,mora. 25 mmlong,theshorteroneto 3(-6) mmlong;bracts few,linear(to subclavate),4-5 mmlong,to 1.5-2.5 cmlonginfruit,puberulous attheapex.Infructescences ca. 4-6 cm diam.;fruitingperianth1.5-2.5 cm long, 3B. Maclura Nuttallsect. Chlorophora (Gaudichaud) greenish,withlarge,embedded yellowdye-containing Baillon, Hist. P1.6: 193. 1875. Type. Chlorophora tinctoria (Linnaeus) B. D. Jackson (= Maclura fruitca.5-6 mmlong,oblonginoutline,flat"glands"; tinctoria(Linnaeus)Steudel). withtwolateralgrooves; tened;endocarpcrustaceous testawitha small(darkbrown)thickened partbelowthe IncludingFusticusRafinesque(see underthe genus). hilum;embryocurved,thecotelydons flat,conduplicate Trees.Staminate inflorescences spicate; stamens andslightlyfolded,? envelopingthelongradicle. inflexed in the bud. Pistillateflowers free; tepals and Distribution (see Fig. 73.4). FromHonduras bractsaccumulatingyellow dye. disjunctlyto eastemBrazil,apparently only a rather The section comprises only Maclura tinctoria. extensive? coherentoccurrencein BolivianAmaPeru(MadredeDios);inforestmarzoniaandadjacent gins, oftenalongstreams,or in (chaco)scrub;to ca. 2. Maclura tinctoria (Linnaeus) Steudel, Nomencl. ed. 2, 2: 87. 1841; Miquel, in Martius, Fl. Bras. 1000(-1350?)m. Specimens examined. HONDURAS. OLANCHE:Nr.

Jaticalpa,5-16 Mar 1949 (st), Standley17842a (F). VENEZUELA. MONAGAS:Nr. QuebradaColorado Grande,SW of Caripe,850-1350 m, 9 Apr 1945 (? fl), Steyermark61949 (F, NY, US, VEN). PERU. MADREDE Dios: ParqueNacionaldel Manu, Rio Manu,Cocha Cashu Station, 21 Oct 1980 (? fr), Foster5616 (BG, F, MOL,U), 21 Oct 1979 (? fr), Gentry et al. 27131 (BG). SAN MARTfN: JuanGuerra,nr. Tarapoto,29 Dec 1929 (a'), LI. Williams6855 (F). BOLIVIA. BENI: Prov.Yacuma,60 km E of San Bora, EstacionBiol6gica Beni, 21 Jul 1989 (st), Beck 16910 (BG); Prov.Moxos, rd. San Borja-Trinidad,km 27, 25 Aug-3 Sep 1990 (? fl-fr), D. N. Smith 14232 (BG, LPB, USZ) & 14248 (BG, BOLV,USZ); Prov. Ballivian, Estaci6n Biol6gica Beni, 50 km E of Rio Maniqui, on rd. to Trinidad, 10 Nov 1985 (? fr), Solomon14648 (BG, infructescencesmixed with materialof Brosimum SANTACRUZ:Prov.Andres lactescens). Ibaiiez,nr. San Lorenzo,13 Feb 1994 (? fr), Jardimet al. 359 (USZ); Prov.Andr6slbainez,12 km SE of Don Lorenzo,0.5 km E of EstanciaCaracore,Rio Grande,

4(1): 155, t. 51. 1853; VazquezAvila, Darwiniana 26: 318, t. 9-11. 1985. Morus tinctoria Linnaeus, Sp. P1.986. 1753.Broussonetiatinctoria(Linnaeus) Kunth, in Humboldt & Bonpland, Nov. Gen. Sp. 2: 32-33. 1817. Fusticus tinctoria (Linnaeus) Rafinesque,New Fl. Amer. 3: 45. 1838 ("1836"). Chlorophoratinctoria (Linnaeus) Gaudichaudex B. D. Jackson,Index Kew. 1: 520. 1893; Bentham & Hooker,Gen. P1.3(1): 363. 1880; Kaastra,Acta Bot. Neerl. 21: 659, t. 1. 1972. Lectotype,Kaastra, Acta Bot. Neerl. 21: 661. 1972. Sloane, Hist. Jamaica 2: t. 158, fig. 1. 1725. Fig. 9 Tree,to 30 m tall, or shrub, especially if juvenile armedwithaxillary,solitary,or sometimespairedthoms to 3.5 cm long, straightand slender, bearing minute scale-leaves at the base. Leafy twigs 0.5-4 mm thick, sparselyto densely puberulousto hirtellous or almost glabrous. Lamina elliptic to oblong, sometimes to subcordiformto suborbicularor to lanceolate, 1.5-15 (-20) x 0.8-7(-8.5) cm, broadest at or below the

TAXONOMIC TREATMENT

57

Fig. 9. 1, 3, 4, 6-lOa. Maclura tinctoria subsp. tinctoria. 1. Leafy twig with staminate inflorescences (VenturaA. 3733). 3. Leafy twig with pistillate inflorescences (Morin 267). 4. Leafy twig with spines and juvenile leaves (Steyermark et al. 10227). 6. Pistillate flower (Little 15882). 7. Fruit. 8. Endocarp body. 9, 9a. Seed. 10, 10a. Embryo (Berg 1306). 2, 5. Maclura tinctoria subsp. mora. 2. Leafy twig with staminate inflorescences (Hassler 7329a). 5. Leafy twig with pistillate inflorescences (Pires et al. 17337). [By P. Pardoen, Utrecht.]

I58

middle,oftendistinctlyinequilateral,chartaceous;apex acuminateor (sub)acute;base obtuse truncateto cordate (or to subacute); margin subentire to coarsely crenate-to serrate-dentate,or if juvenile, then sometimeslobateto lacerate;uppersurfacehirtellousto puberulous on the whole surfaceor only on the lower partof the midrib; lower surface (very) sparsely to densely puberulous,hirtellous, or tomentose on the veins; venationplaneabove,prominentorthe smallerveinsplane beneath;lateralveins 5-10(-17) pairs;tertiaryvenation reticulate;petiole 0.2-1.5(-2) cm long, sparselyto densely puberulous to hirtellous or subglabrous; stipulesfused,ovateto triangular,0.2-1 cm long, white-

FLORA NEOTROPICA

The characteristicyellow dye-containing"glands" arenot consistently present in the bractsand tepals of the pistillate flowers. They occasionally occur in the tepals of staminateflowers. The patternin the variationonly justifies recognition of two infraspecificentitiesatthe subspecieslevel. Thetwo subspeciesarelargelyallopatric,showingmarginal overlapof theirranges of distribution.

Key to the subspecies of Maclura tinctoria

1. Laminawith the smallerveins mostly plane beneath;leaf apex usually long-acuminate; caducous.Staminateinpuberulousto subsericeous, staminateinflorescences2-13 cm long; pistilflorescences:peduncle0.2-2 cmlong,denselypuberlate inflorescencesto 2 cm diam. in fruit .................................................... 2a. subsp. tinctoria ulous to tomentellous;spike 2-13 cm long, 0.2-0.4 cm diam., ? interruptedand with an abaxial sterile strip; 1. Laminawith the smallerveins prominent to subbeneath;leaf apex short-acuminate flowers crowded;tepals (3-)4, almost free, ca. 1 mm acute; staminateinflorescences3.5-4 cm long, densely puberulous;stamens(3-)4; filamentsca. long; pistillateinflorescencesto 1 cm diam. 2 mm long; anthersca. 0.6-0.8 x 0.6-0.8 mm; pistilin fruit...................... 2b. subsp.mora lode flat, ca. 1 mm long, apiculate;bractselliptic, with 2 embeddedyellow dye-containing"glands."Pistillate intheleafaxils;peduncle0.1-1(-2) cm 2a. Maclura tinctoria (Linnaeus)Steudelsubsp.tincinflorescences toria. Morus zanthoxylonLinnaeus, Syst. P1.,ed. long, puberulousto short-velutinous;head (sub)glo10,2: 1266. 1759. Fusticuszanthoxylon(Linnaeus) bose, 0.3-0.6 cm diam.;tepals almost free, 1-1.5 mm Rafinesque,New Fl. Amer.3:45. 1838 ("1836"),as long, ? cucullate and conduplicate, normally with 2 xanthoxylon.Broussonetiazanthoxylon(Linnaeus) embeddedyellow dye-containing "glands,"the upper Martius,Flora24(Beibl.): 10. 1841, asxanthoxylon. partsdensely puberulous,sometimespartlywith unciMaclura zanthoxylon (Linnaeus)Endlicher, Gen. nate hairs;ovary ca. 1 mm long; stigma 1, 3-10(-18) mm long, sometimes the second stigma developed, to P1.Suppl.4(2): 34. 1848,as xanthoxylon;Miquel,in 3 mm long;bracts1-2 mm long, cucullateto subpeltate, Martius,Fl. Bras.4(1): 156. 1853.Macluratinctoria ? swollen due to 2 embeddedyellow dye-containing (Linnaeus)Steudelvar.zanthoxylon(Linnaeus)Bu"glands,"the upperpartpuberulous.Infructescences reau, in De Candolle, Prodr. 17: 230. 1873, as globoseto ellipsoid,ca. 1-1.5 cm diam.;fruitingperianth xanthoxylon.Chlorophoratinctoria (Linnaeus)B. ca. 3 mm long;fruitlens-shaped,ca. 3.5 mm long;cotyD. Jacksonvar.zanthoxylon(Linnaeus)Chodat,Bull. ledonsplane,not envelopingthe slightlycurvedradicle; Herb.Boissier,Ser.2, 3: 349. 1903, asxanthoxylon. bractsusually strongly swollen, to 3.5 mm long. Chlorophora tinctoria (Linnaeus) B. D. Jackson subsp. zanthoxylon (Linnaeus) Hassler,Annuaire The numberof infraspecific names created,as by Conserv. Jard. Bot. Geneve 21: 113. 1919, as Bureau(1873), alreadyindicatesthatthe speciesis very andzanthoxyla.Chlorophoratinctoria xanthoxyla variable.It is particularlyso in the shapeandsize of the B. D. Jackson subsp. zanthoxylon (Linnaeus) lamina,the indumentum,andthe presence and length (Linnaeus)Hasslervar.zanthoxylonHassler,Annuaire of the thoms. The variationis to a large extent related Conserv. Jard. Bot. Geneve 21: 114. 1919, as to the stage of development;juvenile and adultstages zanthoxyla. Type. Plukenet, Phytographia3, Apshow condiderable differences. Juvenile specimens t. f. 3. 1759. pendix, 239, Fig. 9 often have lobate laminas(pinnately lobate or in particularin subsp.mora trilobate,sometimesto laciniate in subsp. tinctoria) andnumerouslong thorns.In adult specimensthe laminais normallyentireandthe thorns shortandoften few. Juvenilefeaturesmay be retained. In the reproductivepartsof the plant,variationis conspicuous in the length of the staminateinflorescences, the diameter of pistillate inflorescences, and in the length of the style.

Broussonetia plumeriSprengel,Syst. Veg., ed. 16, 3: 901. 1826. Macluraplumeri (Sprengel)D. Don ex Steudel,Nomencl.,ed. 2, 2: 87. 1841,as plumiera, with referenceto MorusplumieraBurman. Type. Plumiermsc. ex Burmann,P1.Amer.9: t. 204. 1759. Fusticusvera Rafinesque,New Fl. Amer.3: 45. 1838 ("1836"),nom. superfl.

59

TAXONOMICTREATMENT Fusticus tataiba Rafinesque, New Fl. Amer. 3: 45. 1838 ("1836"), nom. superfl. Maclura sempervirens Tenore, Cat. Hort. Bot. Neap. (1845). 87. 1845. Type. Not traced. Maclura chlorocarpa Liebmann, Kongel. Danske Vidensk. Selsk. Naturvidensk. Math. Afh., Ser. 5, 2: 314. 1851; Miquel in Martius, Fl. Bras. 4(1): 157. 1853. Maclura tinctoria (Linnaeus) Steudel var. chlorocarpa (Liebmann) Bureau in De Candolle, Prodr. 17: 229. 1873. Syntypes. Mexico. Veracruz: Rio Tecolutla, nr. Paso del Correo, Jun 1841 (a'), Liebmann 14298 (C), between Jalapa and Mirador, (? fr), Liebmann s.n. (not traced). Maclura polyneura Miquel, in Martius, Fl. Bras. 4(1): 154. 1853. Maclura tinctoria (Linnaeus) Steudel var. polyneura (Miquel) Bureau, in De Candolle, Prodr. 17: 230. 1873. Chlorophora tinctoria (Linnaeus) B. D. Jackson subsp. zanthoxylon (Linnaeus) Hassler var. zanthoxylon forma polyneura (Miquel) Hassler, Annuaire Conserv. Jard. Bot. Geneve 21: 114. 1919. Type. Brazil. Minas Gerais: nr. Sucuriu, Conceigao, (a'), Pohl s.n. (holotype, M; isotypes, L, U). Maclura affinis Miquel, in Martius,FL.Bras 4(1): 155, t 52. 1853. Maclura tinctoria (Linnaeus) Steudel var. affinis (Miquel) Bureau, in De Candolle, Prodr. 17: 230. 1873. Chlorophora tinctoria (Linnaeus) B. D. Jackson subsp. zantoxhylon (Linnaeus) Hassler var. affinis (Miquel) Hassler, Annuaire Conserv. Jard. Bot. Geneve 21: 113. 1919. Syntypes. Brazil. Rio de Janeiro: Cabo Frio, (v), Luschnath s.n. (not traced), Brazil. Pemambuco: nr. Pernambuco, (a'), Schornbaum s.n. (not traced), and Peru. Without locality, (a' + V fl), Ruiz & Pav6n s.n. (duplicates in F, G, P). Maclura subintegerrima Miquel, in Martius, Fl. Bras. 4(1): 157. 1853. Maclura tinctoria (Linnaeus) Steudel var. subintegerrima (Miquel) Bureau, in De Candolle, Prodr. 17: 228. 1873. Type. Jamaica. Without locality, (v), W Hooker s.n. (holotype, U; isotype, P). Maclura velutina Blume, Mus. Bot. 2: 82. 1856 ("1852"). Type. Puerto Rico. Without locality, (?), Wydlers.n. (lectotype, Kaastra, Acta Bot. Neerl. 21: 663. 1972, U). Maclura sieberi Blume, Mus. Bot. 2: 83. 1856 ("1852"). Type. Trinidad. Without locality, (? fl), Sieber 105 (holotype, L; isotypes, G, LE, MO). Macluratinctoria(Linnaeus)Steudelvar.subintegerrima (Miquel) Bureau subvar. quercina Bureau, in De Candolle, Prodr. 17: 229. 1873. Type. Dominican Republic. Without locality, (st), Bertero s.n. (lectotype, Kaastra,Acta Bot. Neerl. 21: 663. 1972, G; isolectotypes, M, P). Maclura tinctoria var. subcuneata Bureau, in De Candolle, Prodr. 17: 229. 1873. Type. Colombia. Cundinamarca: (Santa Fe de) Bogota, (v), Bon-

pland s.n. (lectotype, Kaastra,Acta Bot. Neerl. 21: 663. 1972, P). Maclura tinctoria (Linnaeus) Steudel var. chlorocarpa Bureau subvar. sinuata Bureau, in De

Candolle,Prodr.17: 229. 1873. Type.Venezuela. Sucre:Cumana,(st), Bonplands.n. (or 37) (holotype, P). Maclura tinctoria (Linnaeus) Steudel var. ovata

Bureau,in De Candolle, Prodr. 17: 229. 1873. Chlorophora tinctoria (Linnaeus) B. D. Jack-

son var. ovata (Bureau) Chodat, Bull. Herb. Boissier,Ser. 2, 3: 349. 1903. Type. Brazil. Rio de Janeiro:(probablythe city of) Rio de Janeiro, (? fr), Gaudichaud1068 (lectotype, Kaastra, Acta Bot. Neerl. 21: 663. 1972, P; isolectotypes, G, L, NY, P). Maclura tinctoria (Linnaeus) Steudel var. zanthoxylon

(Linnaeus)Bureausubvar.lobata Bureau,in De Candolle,Prodr.17: 230. 1873, as xanthoxylon. Type.Venezuela.Zulia:Maracaibo,(st), Plee s.n. (holotype, P). Chlorophora mollis Fernald, Contr. Gray Herb.,

N.S., 28: 52. 1904. Type.Mexico.Oaxaca:Canio Tomellin, (v), Rose & Hough 4672 (holotype, GH; isotypes,NY, US). Chlorophora tinctoria (Linnaeus) B. D. Jackson var.

acuminatissimaHuber,Bol. Mus. ParaenseHist. Nat.5: 332. 1909,nom.inval.(Art.34c Int.Code), based on Huber MG 4937 (MG), from Brazil. Para:Alemquer. Chlorophora tinctoria (Linneaus) B. D. Jackson var. acuminatissima Huber forma glabrescens Huber,

Bol. Mus.ParaenseHist.Nat. 5: 332. 1909, nom. inval. (art. 34c Int. Code), based on HuberMG 4937 (MG), from Brazil. Para:Alemquer. Chlorophora tinctoria (Linnaeus) B. D. Jackson

subsp.zanthoxylon (Linnaeus)Hasslervar.zanthoxylon forma miqueliana

Hassler, Annuaire

Conserv.Jard.Bot. Geneve21: 114. 1919. Type. Paraguay.Cordillerade Altos, Oct 1900 (? fl), Hassler 3368 (holotype,G; isotypes,A, BM, K, L, MO, NY). Chlorophoratinctoria (Linnaeus) B. D. Jackson subsp. zanthoxylon (Linnaeus) Hassler var. zanthoxylon

formatataiiba Hassler,AnnuaireConserv.Jard. Bot. Geneve21: 114. 1919. Type. Paraguay.Nr. Lago de Ypacarai, 1913 (ci), Hassler 12125 (lec-

totype, Kaastra,Acta Bot. Neerl. 21: 664. 1972, G; isolectotypes,A, BM, K, L, LIL,MICH,MO, NY, US, Z). Tree,to 30 m tall, or shrub. Lamina elliptic to oblong, sometimesto subcordiformor to lanceolate,1.515(-20) x 1-7(-8.5) cm; apex often long-acuminate; upper surface usually smooth; lower surface often subglabrous;smallerveins oftenplanebeneath;petiole 0.2-1.5(-2) cm long. Staminate inflorescences: peduncle 0.2-2 cm long; spike 2-13 cm long. Pistillate

60

inflorescences: peduncle to 1(-2) cm long; head 0.30.6 cm diam.,to 1.5 cm diam.in fruit;tepalsandbracts at the apex densely white-puberulous to shortvelutinous;stigma 6-10(-18) mm long. Distribution (see Fig. 74.1). FromMexico andthe West Indies to easternBrazil andArgentina,in northem South America eastwardspossibly to northwestem Guyanaand along the Amazon River,eastwardsto Para (Brazil); in primaryor secondary humid to dry forest, in the Amazon Basin in seasonally inundated (tahuampa or varzea) forest, in eastern Brazil in restinga, and in Argentina in subtropical forest; to 1600(-2700) m. Representative specimens examined. MEXICO. 4 km S of Constituci6n, 70 km E of Escircega, 10 Jul 1983 (9 fl), E. Cabrera et al. 5052 (MO); Tuxpefia, 16 Nov 1931 (67), Lundell 947 (MO, NY); Campo Exp. Forestal, "El Tormento," 25 Jul 1963 (st), Marr6quin 243 (TEX); Palizada, 25-28 Jul 1939 (st), Matuda 3850 (MO, NY). CHIAPAS: 13 km N of Arriaga, 18 May 1972 (? fl), Breedlove 25244 (LL, MO, NY); Mun. Venustiano Carranza, rd. Venustiano CarranzaAcala, 24 Aug 1966 (e'), Laughlin 1678 (LL); Las Garzas, 4-7 Jun 1938 (q fl), Matuda 2711 (A, LL, MO, NA, NY); Chicomuselo, 14 Jul 1941 (a'), Matuda 4506 (A, LL, MO, NY); Mun. Mazatan, Cant6n El Corralito, 1500 m, 10 Jul 1985 (? fl), E. Ventura 2074 (NY). HIDALGO: Huejutla-Macuxtepetle, rd. to Ixcatlan, (e), Moore 2930 (GH). OAXACA: Mun. Salina Cruz, Dtto. Tehuantepec, Rincon Bamba, W of Salina Cruz, 9 Jul 1986 (? fr), Martinez R. 646 (MO, TEX); Mun. Tuxtepec, 22 May 1967 (? fl), Martinez-Calder6n 1401 (LL, MICH, MO, NY); Mun. Cuicatlan, 15 km S of Dominguillo, 1500 m, 12 Apr 1976 (? fl-fr), Rzedowski 34959 (NY); Dtto. Yautepec, 5.5 km SW of Santa Maria Ecatepec, SW of La Reforma, 19 Jun 1985 (q fl), TorresC. et al. 6886 (MO). QUINTANA Roo: 20 km S of Tulum, 27 Jun 1984 (e'), E. Cabrera et al. 6527 (MO, NY); Isla de Cozumel, 8 Jun 1984 (? fl), E. Cabrera et al. 6383 (MO, NY); rd. Pantera-Margarita Maza, 5 Jul 1984 (e'), E. Cabrera et al. 6677 (TEX); Isla de Cozumel, 20 Jul 1984 (), E. Cabrera et al. 6803 (MO, NY); Xcaret, 5 km S of Playa del Carnel, 21 Sep 1985 (a'), E. Cabrera et al. 9253 (MO). SAN Luis POTOSi: Mun. Tampamolon, Tajinab, 4 Jul 1979 (e), Alcorn 3294 (TEX); Clasuapa, Tamazunchale, 21 Jul 1937 (st), Edwards 664 (MO, TEX); Mun. Cd. Valles, ca. 5 km S of Cuidad Valles, 22 May 1981 (a'), Fryxell et al. 3556A (NY, US); Paso de Botella, 24 Jun 1950 (a'), Newman 90 (US); San Dieguito, 13-16 Jun 1904 (%), Palmer 89 (F, GH, K, MO, NY, US), (? fr), Palmer 142 (F, G, K, MO, NY). SINALOA: Baromena, 24 Aug 1941 ($ fr), H. S. Gentry 6130 (F, GH, MICH, MO, NY); El Monte, Los Labrados, 15 Oct 1926 (st), Mexia 930 (MO, NY); San Ignacio, rd. Los Humayez-Los Otatez, 9 Aug 1918 (st), Narvaez M. et al. 443 (US); San Blas, 1925, (v), Ortega 5558 (US). TABASCO: Mun. Paraiso, rd. Paraiso-Barra de Tupilco, km 45, 6 Aug 1979 (? fl-fr), C. Cowan 2306 (CAS, NY). CAMPECHE:

TAMAULIPAS:Nr. Juarez, 18 Jun 1938 (a'), Happ 31 (MO);

FLORA NEOTROPICA 2 mi W of Cuidad Mante-Cuidad Victoria Hwy., on rd. to G6mez Farias, 26 Sep 1960 (st), M. C. Johnston 5733A (TEX); NW of Tampico, 45 km N of Villa Gonzalez, 9 Jul 1983 (e'), Taylor 1948 (DUKE). VERACRUZ: Laguna Verde (Punta Lim6n), 19 Jun 1972 (? fl-fr), Dorantes et al. 810 (MO, U); Mun. Actopan, La Mancha, 18 Jul 1972 (a), Dorantes et al. 1407 (LL, U); Mun. Apazapan, rd. Banios de Carrizal-Emiliano, 27 Jun 1980 (e), Hansen et al. 7484 (MO); Cosamaloapam, Playa Vicente, 24 Aug 1969 (? fr), Martinez-Calder6n 1991 (A, C, MO, NY); 2 km S of Tampico, 1-2 Jun 1910 (e'), Palmer 552 (BM, F, G, GH, K, NY, US); Remudadero, May 1923 (e'), Purpus 9030 (F, GH, MO, NY, US); Mun. Dos Rios, El Roble, 8 Jun 1979 (a), Ventura A. 16191 (MO). 17 km N of Merida, 22 May 1985, E. Cabrera YUCATAN: et al. 8590 (MO); 35 km SE of Oxkutzcab, 19 Jul 1985 (e), E. Cabrera et al. 9013 (TEX); 1 km S of Telchaquillo, 21 Jul 1985 (a'), E. Cabrera et al. 9132 (MO); Suitun, May 1916 (a' + ? fl), Gaumer et al. 23302 (BM, F, GH, K, MO, NY, US); Chichen Itza, Jun 1932 (e), Steere 1649 (MICH), (Q fl), Steere 1649 (MICH); nr. Ticul, 12 Jun 1982 (fl-fr), White 167 (MO); nr. Calcehtok, 16 Jul 1982 (e), White 215 (MO). BELIZE. OrangeWalk District, IndianChurch, 10 Oct 1977 (e), Arnason 17757 (MO); Corozal District, San Andres, Oct 1933 (a'), Gentle 831 (A, MICH, MO, NY), Jul 1933 (? fl-fr), Gentle 4738 (NY); San Andres, Corozal, 12 Jul 1933 (? fl-fr), Lundell 4738 (MO); Cayo District, Central farm, 29 Apr 1970 (a), Rose Innes 305 (MO). GUATEMALA. CHIQUIMULA: Nr. Jocatan, 9 Nov 1939 (st), Steyermark 31543 (F). EL PROGRESO: Nr. Barranquillo, 17 May 1942 (2 fl), Steyermark46436 (A, F). ESCUINTLA: Nr. San Jose, 30 Jan 1939 (st), Standley 64165 (F). HUEHUETENANGO: Paso del Boquer6n, Rio Trapichillo, 1200-1300 m, 21 Aug 1942 (st), Steyermark 51146 (F); Rio Trapichillo, between Paso del Boquer6n & Democracia, 1000-2 100 m, 22 Aug 1942 (? fr), Steyermark51028 (F, NY). IZABAL: Gualan, 15 Jun 1909 (a' fl), Deam 6283 (US), 20 Jun 1909 (? fl-fr), Deam 6378 (A, F, GH, MICH, MO, NY, US), 26 May 1919 (st), Whitford et al. 70 (US). JUTIAPA: Between Jutiapa & Las Tunas, 4 Nov 1940 (st), Standley 76250 (F). PETEN: Uaxactun, 18 Apr 1931 (st), Bartlett 12575 (F, NY); Tikal, 29 Jul 1959 (? fl), Contreras 40 (F, LL, MICH, MO, NY, US); Parque Nacional Tikal, Santa Elena, rd. to La Libertad, km 3, 8 Jun 1970 (a'), Tun Ortiz 1200 (MO, NY); nr. Uaxactuin, 16 May 1973 (a'), lin Ortiz 2559 (NY, US). RETALHULEU:Champerico, 26 Feb 1939 (st), Standley 66635 (F). SANTA ROSA:Nr. Guazacapan, 6 Dec 1940 (st), Standley 78697 (F); Capulin region, S of Los Cerritos, rd. to El Ahumado, 7 Dec 1940 (st), Standley 79606 & 79610 (F). SUCHITEPPQUEZ: 7 mi S of Tiquisate, 19 Jun 1942 (? fl), Steyermark 47816 (F). ZACAPA: 10 km W of Teculutin, 4 Aug 1970 (a'), Harmon et al. 3542 (MO); nr. Zacapa, 6-16 Oct 1940 (st), Standley 73629 (F); Rio Hondo, nr. electrical plant, 11 Oct 1940 (st), Standley 74013 (F). EL SALVADOR. LA LIBERTAD: Deininger, 5 Jun 1976 (a), Current 31 (MO). LA PAZ: Herradura, 25 Aug 1950 (a),

Rohweder

3613

(MO). LA UNI6N: Nr. La

Uni6n, 13-21 Feb 1922 (st), Standley 20795 (GH, US).

TAXONOMIC TREATMENT SANTA ANA: Nr. Santa Ana, 8 Jan 1922 (st), Standley

19688 (F, GH, MO, US). SAN MIQUEL: Laguna del Jacotal, 28 Aug 1958 (d), Allen et al. 6895 (NY, US); Laguna de Olomega, 20 Feb 1922 (st), Standley 21030 (GH, MO, NY). SAN SALVADOR:San Salvador, Jul 1922 (? fr), Calder6n 901 (GH, NY, US). SONSONATE:Finca Chilato, 26-27 Dec 1921 (st), Standley 19307 (US). Las Limas, ca. 1000 m, HONDURAS. COMAYAGUA: 3 Jul 1932 (st), Edwards P-387 (F). COPAN:Macuelino, 20 May 1919 (st), Whitfordet al. 59 (US). CORTES:Nr. La Lima, 9 Nov 1963 (e), Dickison 1204 (US), 11-20 Apr 1947 (st), Standley et al. 7557 (F). EL PARAiSO: Nr. Lipaza, 15 Oct 1951 (st), Standley 28890 (EAP). ISLAS DE LA BAHIA: Isla Santillana o de Swan, 10 Jul 1986 (9 fl-fr), Nelson et al. 9768 (MO). CHOLUTECA: Nr. Choluteca, 21 Oct-9 Nov 1949 (st), Standley 24351 (EAP, F). MORAZAN: Drainage of Rio Yeguare, Pedregal, 23 Jul 1948 (? fl-fr), Glassman 2030 (EAP, F, GH, NY); nr. Zamorano, 10 Jul 1947 (v), Molina R. 266 (EAP, F, GH, LL, MO, US); Rio Yeguare, Apr 1944 (v), ValerioR. 2122 (EAP, F). OLANCHO: Nr. Juticalpa, 5-16 Mar 1949 (st), Standley 17745 (F). SANTA BARBARA: El Mochito, 23-26 1976 (? fl), Vargas 739 (BG). VALLE: Amapala, 11 Sep 1945 (v), ValerioR. 3380 (F); San Lorenzo, 14 Sep 1945 (e), ValerioR. 3478 (EAP, F). YORO:Mun. Olanchito, San Jose, Rio Agalteca, 2 Jul 1994 (v), Davidse et al. 35518 (EAP); nr. Progreso, 24 Jan 1928 (st), Standley 55024 (US). NICARAGUA. BOACO:San Jose de los Remates, 25 km N of Teustepe, 10 Oct 1982 (? fr), Sandino 3673 (MO). CARAZO: Estaci6n Biol6gica de Chacopcentro, border Carazo/Rivas, 22-23 Jun 1984 (? fl), Soza 101 (MO). CHINANDEGA: SE of Volcan Casita, 30 Jul 1980 (? flfr), Moreno 1547 (MO); Potosi, rd. to "Los Placeres," 18 May 1983 (e'), Robleto 23 (MO); Volcan San Crist6bal, 20 km NE of Chinandega,23 Aug 1984 (? fr), Sosa et al. 168 (MO); nr. Chichigalpa, 12-18 Jul 1947 (d), Standley 11468 (F). CHONTALES: San Francisco, rd. Rama, km 93, 25 Jul 1981 (? fl-fr), Moreno 10087 (MO); 5 km N of Juigalpa, 13 Jul 1976 (a'), Neill 605 (CR, MO); 4.9 km E of PuertoDiaz, rd. to Juigalpa,5 Jun 1980 (Y fi), Stevens 17422 (MO); Hda. Corpus, W of Juigalpa, 24 Jul 1983 (c'), Stevens 22262 (MO). ESTELi: 4 mi S of Esteli, 4 Aug 1971 (d), Dwyer et al. 443 (MO); 6 km NW of Esteli, 1300-1400 m, 22 Sep 1983 (Y fr), Laguna 265 (MO); "CerroEl Picacho," S of Esteli, 24 Jul 1980 (d), Moreno 1243 (MO); Quebrada La Limosa, 2 km NW of Esteli, 25 Jul 1980 (? fl-fr), Moreno 1389 (MO); Montaniade Cucula, 25 km N of Esteli, 9 Jul 1982 (d), Martinez 7 (MO). GRANADA: 3 km E of Casa de Tejas, rd. to Charco, 5 Jun 1980 (? fl), Araquistain et al. 2844 (MO); Isla Zapatera,Hda. La Habana, 13 Aug 1982 (? fl-fr), Grijalva 783 (MO); Laguna Apoyo, "Babilonia," 30 May 1981 (? fl), Moreno et al. 8903 (MO); 12 km N of Intecna, La Playuela, 24 Jun 1981 (? fl-fr), Sandino 725 (MO). LE6N: Volcan Casita, 28 May 1983 (e'), Grijalva et al. 2534 (MO); km SE of Puerto Momotombo, 8 Jul 1983 (? fl), Grijalva et al. 2757 (MO); Isla Momotombito, 10 Jun 1979 (? fl), Stevens 13226 (BG), (a), Stevens 13227 (MO); ca. 4 km SW of La Paz Centro & 6.2 km

61 NE of Impalme Izapa, 31 May 1983 (st), Stevens et al. 22162 (MO). MANAGUA: Rd. Managua-Masaya, km 17.5, 5 Jul 1982 (? fl), Castro 2628 (MO); Peninsula de Chiltepe,SE of Hda. El Tamagas,15 Aug 1980 (a'), Moreno 1759 (MO); Volcan Chiltepe, 2 Jun 1981 (? fl), Moreno et al. 8935 (MO); between Managua & Tititapa, 26 May 1976 (? fl-fr), Neill 402 (CR, MO, NY); TamagastaPeninsula, W of Managua, Lago Julia, 25 Jun 1923 (2 fl-fr), Maxon et al. 7317 (GH, NY, US), (a), Maxon et al. 7324 (NY, US). MASAYA: Santa Teresa beach, S of Las Piedras, 28 Jun 1982 (? fl-fr), Kral 69143 (MO); ParqueNacional Volcan Masaya, NW slope, 30 Oct 1977 (st), Neill 2855 (MO). MATAGALPA:Rd. Jinotega, km 134, 15 Jun 1982 (? fl-fr), Mangas 34 (MO); Mun. Dario, El Nancital, 17 Aug 1981 (ae), Moreno 10495 (MO);QuebradaLas Palomas, nr. Chagiiitillo, 19 Jan 1979 (st), Stevens 11778 (BG, MO). NUEVA SEGOVIA: 8 km NE of Jicaro, 2 Aug 1980 (a'), Moreno 1671, (? fl), Moreno 1690 (MO). RIVAS:Rio Nagualapa, nr. Hda. Los Angeles, 23 Aug 1977 (ae), Neill 2391 (MO, U); Isla Ometepe, "La Palma," 21 Sep 1984 (st), Robleto 1209 (MO); Lago Cocibolca, Finca El Mengo, 28 Jun 1982 (? fl-fr), Sandino 3171 (MO); Quebrada Las Cafias, nr. Rio Escalante, 3 Aug 1978 (a'), Stevens 9665 (MO); confluence of Rio La Pita & Rio Escalante, 3 Aug 1978 (a'), Stevens 9716 (MO). COSTA RICA. ALAJUELA: Cuesta la Cima, Jul 1924 (a'), Brenes 3992a (NY); Rio Grande de Tarcoles, nr. Capulin, 2 Apr 1924 (st), Standley 40168 (US). GUANACASTE: Monteverde, 1300-1400 m, 8 Jun 1988 (? flfr), Bello 438 (MO); between Belvedero & Taboga, 14 Jun 1930 (? fr), Brenes 12558 (F); Palo Verde, 3 Jun 1969 (? fl-fr), Frankie 149a (MO); Filadelfia, 12-14 Jun 1984 (? fl), G6mez et al. 23042 (BG, MO), (a'), G6mez et al. 23059 (BG, MO); Santa Rosa National Park, rd. Naranjo Beach, Rio Poza Salada, 21 Jun 1977 (a"), Liesner 2277 (MO); Temisque, 7 Aug 1932 (? fr), Stork 3389 (F, GH, MICH, NY). PUNTARENAS:Nr. PalmarNorte de Osa, 10 Aug 1950 (e + ? fl), Allen 5601 (F, US); Parque Nacional Corcovado, Sirena, 22 May 1989 (? fl-fr), Kernan et al. 1106 (BG, MO); Aranjuez, 2 Apr 1963 (e), Jimenez M. 597 (CR); Atenas, 12 Jan 1942 (st), Quiros C. s.n. (CR). SAN Jost: Arenas, 1000 m, 24 Jun 1924 (? fl-fr), Brenes 3992 (F, NY, P); Poros, Hda. Lindora, 13 Jun 1965 (? fl), Hatheway 1435 & (e), Hatheway 1436 (CR, F, GH); La Caja, 1000 m, 15 Jul 1935 (? fr), Valerio 1306 & (a), Valerio 1307 (F). PANAMA. DARIEN: Nr. Garachine, Jan-Feb 1912 (st), Pittier 5695 (NY, US); S of El Real, Rio Tuira, 30 Jun 1959 (? fr), Stern et al. 754 (GH, MO, NY); 16-19 km SE of Jaque, 16 Jan 1981 (st), Garwood 1111 (MO, PMA). CHIRIQUI: Progresso, 1927 (st), Cooper et al. 237 (F, MO, US); Comarca del Baru, nr. gorge of Rio Corotu, W of PuertoArmuelles, 16 Jun 1957 (? fl), Stern et al. 136 (F, MO, NY, US); nr. Puerto Armuelles, 2831 Jul 1940 (? fr), Woodson et al. 899 (F, MO, NY). HERRERA: Divisia, 27 Aug 1976 (a'), Lao 312 (H, MO), (? fr), Lao 315 (H). Los SANTOS: Isla de Cafias, 8 Jul 1986 (a), Carrasquilla 3019 (MO, PMA), (? fl-fr) Carrasquilla 3020 (PMA); 1-2 mi W of Candelaria, 14 Jul 1967 (? fl-fr), Duke 12449 (MO, US); Las Tablas, 1

62 Jul 1961 (? fl), Dwyer 1171 (MO); Pocri, 8 Nov 1962 (? fr), Dwyer 2518 (US); Guarare, 28 Aug 1971 (? fr), Lao 315 (MO, PMA). PANAMA:Ancon, 19 Mar 1970 (st), Croat 8878 (MO); between Canita & El Llano, 1 Sep 1971 (? fr), Gentry et al. 1712 (MO, NY); Balboa Heights, 18 Sep 1917 (? fl), Killip 3000 (US); nr. Chepo, 29 Oct 1911 (st), Pittier 4710 (US); Rio Tecumen, 3 Jan 1924 (st), Standley 29381 (US). CUBA. LA HABANA:Santiago de las Vegas, 22 Jun 1905 (d'), Abarca 5143 (A, NY, US); nr. Guanabacoa, 30 Apr 1914 (a'), Ekman 553 (NY); Casa Blanca, 6 Jul 1923 (a' + V fl-fr), Ekman 16877 (US); Rincon, 9 Aug E of 1904 (a'), Wilson 587 (A, NY, US). MANTANZAS: Mantanzas, 9 Sep 1903 (? fl), Britton et al. 407 (NY). ORIENTE:Bayate, 1 Sep 1915 (? fl-fr), Ekman 6407 (F, K, MICH, NY, US); Via Azul, Yateritas, 10 Jan 1956 (st), Liogier et al. 4945 (NY); Monte Verde, Jan-Jul 1859 (a' + ? fl), Wright1439 (BM, G, GH, K, MO, NY, P). PINAR DELRio: Bay of Nariel, 19-23 Sep 1910 (juv), Britton et al. 7649 (NY); Mariel, 2 Jun 1921 (? fr), Ekman 12836 (F, G, K, MICH, US). SWAN ISLANDS. Great Swan, 15-24 Aug 1971 (9 fr), Proctor 32502 (BM, GH, LL). JAMAICA. Nr. Port Antonio, 25 Sep 1906 (? fl-fr), Britton 898 (NY); St. Andrew, Cane R. gorge, 26 Jun 1963 (e), Crosby et al. 329 (F, MO); St. Ann, Clover Hill, 28 Oct 1962 (st), Adams 11805 (MO); St. Elizabeth, nr. Malvem, 12 Sep 1954 (? fl-fr), Howard et al. 13691 (A, NY, US); Blue Mts., nr. Green Valley, 16 Jun 1916 (a), Perkins 1196 (A, F, GH, K); St. Mary, nr. Hampstead, 15 May 1954 (a), Proctor 8678 (NY); St. Andrew, Cane R., 22 May 1955 (? fl-fr), Proctor 10216 (NY, US); St. Thomas, Cambridge Hill, 24 Nov 1963 (? fl-fr), Proctor 24221 (LL, NY); St. Thomas, Yallahs R., 5 Sep 1957 (? fr), Robertson 3345 (NY); West Moreland, 20 Jul 1954 (a), Websteret al. 5056 (A, BM, MICH). CAYMAN ISLANDS. Grand Cayman, West Bay, Pool of Silence, 5 Aug 1938 (a'), Kings GC-408 (MO, NY); Grand Cayman, 9 Jun 1967 (? fl), Proctor 27982 (BM, MO, U). HAITI. Port-au-Prince, Bon-Repos, 15 Oct 1924 (9 fr), Ekman 2166 (US); Port-au-Prince, 17 May 1942 (2 fl-fr), Holdridge 1224 (BM, F, LIL, MICH, MO, NY, US), (a'), Holdridge 1225 (BM, F, LIL, MICH, MO, NY, TEX, US); nr. Fond Parisien, Etang Saumatre,3-13 May 1920 (a), Leonard 4163 (GH, NY, US); Tortue Island, La Vallee, 28 Dec 1928-9 Jan 1929 (? fl-fr), Leonard et al. 11330 (NY, US); nr. Mole St. Nicolas, 13-19 Feb 1929 (? fl-fr), Leonard et al. 13123 (GH, K, MO, US). DOMINICAN REPUBLIC: Prov. Barahona, 8 km S of Barahona, 5 Jul 1968 (? fl-fr), D'Arcy 2682 (A, MO); Jaina, 13 May 1929 (a'), Ekman 12455 (G, LL, US); Prov. Barahona,Apr 1910 ($ fl-fr), Fuertes 11 (BM, F, GH, K, LE, LIL, MO, NY, U, US, Z), (a'), Fuertes 190 (BM, C, F, GH, K, LIL, M, MO, NY, U, US, Z); N of Barahona, 14 Aug 1946 (a'), Howard et al. 8290 (B, BM, GH, NY, US); nr. San Juan de la Maguana, 5 Sep 1946 (a), Howard et al. 8810 (B, BM, GH, NY, P, US), (? fl-fr), Howard et al. 8812 (B, BM, GH, NY, P, US);

FLORA NEOTROPICA La Caleta, 27 May 1973 (a'), Liogier et al 19312 (NY); Prov. Peravia, 10 km N of Bani, 6 May 1981 (? fl-fr), Mejia et al. 13262 (BG, MO, NY); Prov. La Romana, N of Boca de Cumayasa, 11 Jun 1981 (? fl), Mejia 14762 (MO, NY); Guayacamas, 1962 (a"),Lavastre 298 (NY); Navayo, 27 Mar 1965 (a'), Lavastre 1928 (NY); Prov. Azua, 7 km N of Las Charcas, 22 Jul 1982 (? fr), Zanoni et al. 21899 (MO, NY). PUERTO RICO. Las Pijevas, 16 Mar 1913 (st), Britton et al. 2220 (NY); Rio Jueyes, 3 Apr 1930 (st), Britton et al. 9557 (NY); 7.6 km W of Carmen(st), D 'Arcy 1945 (MO); Mona Island, nr. Sardinera,26 Aug 1954 (? fl), Little et al. 16457 (BM, F, NY, US); nr. Bayamon, 28 Mar 1885 (st), Sintenis 995 (BM, GH, K, US); Cayey, 25 Sep 1885 (? fl-fr), Sintenis 2382 (BM, C, G, GH, K, MO, P, US, Z); nr. Guanica, 13 Feb 1886 (st), Sintenis 3760 (BM, GH, K, US); Mona Island, 11-12 Dec 1971 (? flfr), Woodburyet al. M-239 (NY), 1977 (v), Woodburyet al. 633 (NY); Vieques Island, nr. Martineau, 13 Jun 1969 ( fl-fr), Woodburys.n. (NY, US). ST. CROIX. Crequis, 11 May 1874 (o), Eggers s.n. (C, US); without locality, (? fl), Hansen 79 (C, NY); Cane Bay, 24 Jun 1954 (? fr), Little 16407 (NY, US); Jolly Hill, 17 Jan 1906 (st), Raunkiaer s.n. (or 2525) (C); Jerusalem Estate, 20 Jul 1896 (? fl-fr), Ricksecker 482 (F, MO, NY, US). ST. THOMAS. Without locality, (st), Riise s.n. (C). MARTINIQUE. Without locality, (? fl), T Duss 1208 (Z), (v), Plee 825 (P), (e) Plee 848 (P, U). ST. LUCIA. Soufriere, 14 Jul 1945 (a' + ? fl), Beard 526 (A, K, LIL, MO, NY, U), 14 Jul 1945 (a'), P. Beard 1177 (GH, MO, US), Aug 1886 (? fr), Duss 1404 (NY); Marquis estate, 4 Apr-12 Jun 1958 (? fl), Proctor 18177 (BM, US). ST. VINCENT. Withoutlocality, (e), Guilding 7 (BM). GRENADA. Pointe Saline, 28 Oct 1945 (? fr), P. Beard 1330 (GH, US); St. George, 25 Apr 1906 (a'), Broadway 1871 (F). BARBADOS. Without locality, (? fl-fr), Gooding 287 (BM); St. Andrew Parish, Boscobel, 2-3 Mar 1979 (juv), Howard et al. 18734 (NY); St. Michael, Jun 1901 (? fl-fr), Revalle 194 (NY). CURACAO. Klein Sint Martha, 26 Feb 1917 (st), Curran et al. 145 (NY, US). COLOMBIA. AMAZONAS: Lakes N of Leticia, 4 Jul 1969 (a'), McDaniel 11790 (MO); Rio Loretoyacu, SepNov 1944 (? fr), Schultes 6028 (F), Mar 1946 (ae), Schultes 7169 (F), (9 fl), Schultes 7171 (COL, F). ANTIOQUIA: Mun. Santa Fe de Antioquia, rd. to Anza, 1 May 1985 (9 fr), Fonnegra et al. 1664 (MO); Casabe, 3 Nov 1979 (? fl-fr), Renteria et al. 2035 (MO); Mun. Venecia, 4.2 km E of Bolomboio, 12 Mar 1987 (? flfr), Zarruchi et al. 4650 (BG), (a), Zarucchi et al. 4663 (BG, COL). ATLANTICO: Solgar y Sabanilla, 13 Jan 1962 (st), Dugand 5972 (COL); nr. Barranquilla, Jul 1934 (a'), Bro. Elias 1246 (F, US); nr. Baranquilla, Juan de Costa, 9 Jul 1963 (? fr), McKee 10458 (COL). BOLIVAR: San Martin de Loba, Apr-May 1916 (a'), Curran 7 (GH, US); Mun. San Martin de Loba, rd. to harbour, 24 Jul 1987 (? fl), Cuadros 3776 (MO); nr. Since, 7 Sep 1963

TAXONOMICTREATMENT (9 fl), Romero-Castafneda 9806 (COL); Mun. Santa Catalina, nr. Algarrobo, 13 Aug 1985 (e'), Zarucchi et al. 4122 (BG). BOYACA: Mun. Soata, Rio Chicamocha, Cani6ndel Chicamocha, Dec 1952 (9 fl-fr), Hernandez G. 6612 (COL). CALDAS: nr. La Dorada, 1-20 Feb 1946 (d' or ? fl-fr), Duque-Jaramillo 2580 (COL, NY), (9 fl), Yepes-Agredo 713 (COL). CAQUETA: Solano, 8 km SE of Tres Esquinas, Rio Caqueta, below mouth of Rio Orteguaza, 12 Mar 1945 (e), Little et al. 9778 (COL, NY, LIL, US). CUSAR: 5 km W of Manaure, rd. to La Paz, 26 Apr 1987 (st), Gentry et al. 57142 (BG, MO), 13 Jan 1988 (st), Gentry et al. 60720 (MO); Mun. Valledupar, between Antanquez & Guatapuri, 1130 m, 18 Nov 1985 (9 fr), Torres et al. 2848 (MO, NY). CUNDINAMARCA:Camboas,7 May 1940 (a'), Cuatrecasas 9415 (COL, F, US); below Fusagasuga, 1400-1500 m, 24 Jan 1944 (? fl-fr), Dugand 3545 & 3546 (COL);Apulo, 16-20 May 1946 (a'), Duque-Jaramillo 3600 (COL, LIL); Guaduas, 1000-1040 m, 5 Nov 1945 (e), GarciaBarriga 11766 (COL, US); rd. to Tibacuy, ca. 8 km from Silvania-Fusagasuga toll rd., 1500 m, 15 Dec 1982 (a), Lewis et al. 9784 (MO); W of La Mesa, ca. 1000 m, 16 Dec 1982 (9 fr), Lewis et al. 9786 (MO); Girardot, El Guayabal, 6 Aug 1939 (9 fr), Perez-Arbelaez et al. 6512 (COL, F). HUILA: Rd. Neiva-Baraya, 16 km SW of Colombia, 40 km NE of Baraya, 21 Mar 1983 (a), Croat 55324 (BG, MO); Mun. Tello, Vrda. Arenosa, Hda. Ricaurte, 8 Feb 1974 (e), L. Forero 913 (NY); Algeciras (San Juanito), 40 km S of Neiva, 1050 m, 3 Dec 1942 (st), Fosberg 19235 (NA); rd. Bogota-Neivo, km 190, 8 Jan 1974 (st), Gentry et al. 8978 (MO); 15-20 km S of Santa Ana, 1500-1700 m, 21 Feb 1944 (a'), Little 7311 (COL, MICH, US); Rio Cabrera, 18 km NE of Villavieja, 12 Jul 1950 (e), S. G. Smith 1189 (COL, GH, IAN, MO, NY, US). MAGDALENA: Upper Rio Honda, Opia, Dec 1852 (9 fr), Holton 250 (G, K, NY); Mun. Santa Marta, Parque Nacional Natural Tayrona, 9 Sep 1976 (st), Lozano C. 2752 (NY); Tueurinea, 25 Apr 1948 (9), 1063 (COL); Mun. Fundaci6n, beRomero-Castaneda tween Caserio & Rio Fundaci6n, Cgto. Santa Rosa, 3 Aug 1971 (9 fl-fr), Romero-Castanieda 11147 (MO, NY); Santa Marta, 15 Sep 1898 (d' + ? fl), H. H. Smith 425 (A, BM, F, K, MICH, MO, NY, P, TEX, U). NORTE DE SANTANDER:Nr. Cucuta, 12-23 Mar 1927 (ae), Killip et al. 21013 (A, F, GH, NY). SANTANDER: Mun. Velez, Rio Carare-Rio Op6n region, Campo Capote, Sep 1970 (? fl-fr), Cabrera 875 (COL); Rio Chicamocha, between Capitanejo & Enciso, 1200-1300 m, 18 Jul 1940 (a'), Cuatrecasas et al. 9841 (F, US); 17 km NE of Socorro, toward San Gil, 1250 m, 23 Jul 1975 (? fl-fr), Gentry et al. 15336 (MO); Rio Surata, nr. Bucaramanaga, 28 Dec 1926 (9 fl-fr), Killip et al. 16230 (A, GH, NY, US); Puerto Araujo, 18 Sep 1979 (a'), Renteria et al. 1764 (MO). TOLIMA: Armero, 6 Oct 1940 (a'), Cuatrecasas 10489 (COL, F, US), 13 May 1939 (a), Garcia-Barriga 7519 (COL, F, US); Prado, 9 Feb 1944 (a), Little 7154 (US); 10 km SE of Santa Ana, 2000-2200 m, 17 Feb 1944 (a' + ? fl-fr), Little 7261 (COL, US); between Espinal & Cuamo, 21 Jul 1971 (? fr), Pennell et al. 202 (NY); Flandes, 14 Dec 1946 (st), Schneider 213 (COL).

63 VALLE: Rio Dagua, Buenaventura, Apr (? fl-fr), Lehmann 5093 (F, K). VENEZUELA. APURE:Nr. San Fernando de Apure, 12 Apr 1951 (e), Trujillo2224 (MY). ARAGUA: San Juan de los Morros, 3 Jan 1939 (? fl-fr), Alston 6018 (BM, NY, US); Parque Nacional Henri Pittier, 29 May 1963 (? fr), Montaldo 3420 (F, MY); between El Lim6n & Guamitas ('), Pittier 15407 (VEN); Maracay, 20 Sep 1957 (? fl), Trujillo 3786 (MY). BARINAS: Ticoporo Forest Reserve, Rio Bumbun, 4 May 1964 (? fr), Breteler 3903 (EAP, F, MO, MOL, P, U, US); Mun. Pedreza, Ciudad Bolivia, 24 Apr 1953 (e), Little 15108 (VEN); Rio Caparo, between Campamento Cachicamo & Boca de Garza, E of El Cant6n, 12 Apr 1968 (st), Steyermark et al. 102227 (U, VEN). BoLfVAR: Rio Asa, above Raudal Cotua, S of Paragua, 1 Aug 1960 (R fr), Steyermark86720 (F, K, NY, US, VEN). CARABOBO: Puerto Cabello, 21 Jun 1913 (? fl), Pittier 6359 (US). COJEDES: 12 km S of Santa Lucia, 20 May 1984 (e), Ramia et al. WT3534 (NY). DISTRITOFEDERAL: CurucutoValley, 30 Aug 1927 (? fr), Pittier 12453 (A, F, G, K, L, M, MO, NY, VEN); between Caracas & Puerto Cabello, nr. Zigzag, 19 Oct 1921 (? fr), E. Pittier 74 (NY); 40 km E of La Guaira, 1-2 km S of "Los Caracas," 18 Aug 1979 (? fl-fr), Nee 17562 (F); Pariata, Sep 1938 (a), Tamayo 589 (US); Maiquetia, 25 Aug 1940 (e), Tamayo 1361 (A, F, VEN). FALC6N: Ca. 15 km NNE of Aroa, 25 Jul 1985 (? fl-fr), Colonello 952 (MO, NY); Cerro Socopo, above Socopito, 28 Jun 1979 (? fl-fr), Liesner et al. 8300 (MO, NY, U); Dtto. Silva, Cerro Chichiriviche,between Lizardo & Mallorquines, 4 Sep 1974 (d), Steyermark et al. 110724 (F, MO, NY). GUARICO: Altagracia de OritucoTamanaco, Mar 1966 (e), Aristeguieta 6035 (MO, NY, VEN); 19 km S of San Juan de Los Morros, rd. to Calabozo, 28 Jul 1974 (e), Morillo et al. 4269 (MO, NY); Rio Pascua, Zaraza, Jul 1959 (st), Ramia 1921 (VEN). LARA: Rio Guaremal, NE of Barquisimeto, nr. Yaracuy border, 29 Jul 1979 (st), Meijer et al. 72A (MO); Dtto. Palavecino, N of Cerro Grande, 2 Jun 1974, Steyermark et al. 109910 (U); Limoncito, 30 May 1964 (e), Trujillo 6443 (MY). MERIDA: Mun. Santa Cruz de Mora, Puente La Victoria, 1 Jul 1954 (9 fl-fr), Bernardi 1314 (K, NY, P, VEN); 30 km SW of Merida, 1100 m, 30 Jul 1964 (? fl-fr), Breteler 4064 (COL, MG, MO, NY, P, U, US); Chama valley, nr. Estanques, 1 Sep 1965 (e'), Breteler 4555 (COL, EAP, MO, NY, U, US); Dtto. Sucre, Mun. Estanques, Puerto del Dorado, 22 May 1980 (6'), Carrillo 28 (MO, U); Ejido, 1170 m, 7 Jun 1953 (? fl), Little 15187 (COL, VEN). MIRANDA: Dtto. Urdaneto, Cua, 2 May 1976, F Bianco 87 (VEN). PORTUGUESA: Agua Blanca, 22 Apr 1948 (st), Turner 110 (F). SUCRE: Nr. Crist6bal Col6n (Macuro), 5 Jan-22 Feb 1923 (st), Broadway s.n. (NY); valley of Cocollar, 28 Apr 1945 (st), Steyermark62352 (F, VEN); Dtto. Sucre, Paso Hondo, S of Limonal & Santa Fe, 18 Aug 1973 (st), Steyermarket al. 107825 (MO, NY). TACHIRA: San Pedro, 17 May 1917 (e), Curran et al. 1022 (A, GH, K, NY, US); S of La Mulata, nr. Colombian border, 13 Nov 1979 (6), Steyermark et al. 120264 (MO). TRUJILLO: Rd. Sabana de Mendoza-La Ceiba, km 7, 24 Apr 1977 (? fl), Bunting

FLORA NEOTROPICA

64 et al. 5038 (U); nr.Recreo,30 May 1970(a'), Benitez de Rojas 897 (MY); San Jacinto, 27 Apr 1941 (e), Tamayo 1700 (VEN). YARACUY:Rio Yaracuy, 15 Aug 1952 (? fl-fr), Schnee 1151 (F, MY). ZULIA: Dtto. Mara,7 km

23 Feb 1983 (9 fr), Berg 1306 (QCA, U); rd. Santo Domingode los Colorados-PuertoLim6n, ca. 10 Dec 1983(st),Kvist et al. 49060 (BG).ZAMORA-CHINCHIPE:

SW of CampamentoCarichuanoof Carbozulia,10 Aug 1981 (a'), Bunting et al. 10199 (NY); Dtto. Perija,between Rio Yasa& Rio Tucuco,rd. Machiques-LosAngeles de Tucuco,21 Jun 1980 (a), Davidse et al. 18398 (LL, MO, NY), (? fl), Davidse et al. 18399 (LL, MO, NY); Dtto. Mara,5 km SW of CorpozuliaCampamento Socuy,2 Jun 1980 (e), Steyermark et al. 123135 (MO); Cerro Cinco de Julio, 8.5-10 km NW of Corpozulia 4 Jun 1980 (? fl), Steyermark Campamento Carichuano, Dtto.SanFelipe, et al. 123304 (LL,MO,NY). YARACUY: rd. El Guayabo-laHoya, 0.8 km N of El Guayabo,12 Jul 1973 (2 fl-fr), Agostini et al. 1829 (U); Rio Yaracuy, 15 Aug 1952 (? fl-fr), Schnee 1151 (F).

(BG, LOJA,QCA);52 km NE of Zamora,11 Sep 1975

TRINIDAD. St Ann's, 10 Aug 1922 (e'), Broadway 5100 (BM, F, MO, Z); Woodbrook, AriapitaAvenue,14 May 1932 (?), Broadway 8046 (A, BM, MO); Pointe Gourde,29 May 1932(? fl), Broadway 8086 (A, B, BM, K, MO, Z); nr. St. Clair, 11 Sep 1918 (a'), Broadway s.n. 28 May 1929(e), Broadway (MO,NY,US);Woodbrook, s.n. (MO); without locality, 1877-1880 (a' + ? fl-fr), Fendler 718 (NY); St.Ann's,(? fl), Hart 6555 (NY); Claxton Bay, 19 Sep 1979 (? fl-fr),Ramcharan 596 (NY). TOBAGO. Rockley Vale, 21 Jul 1910 (e' + ? fl-fr), Broadway 4072 (BM, F, U, Z).

nr. Zamora, ca. 1000 m, 4 Jan 1991 (a'), Berg et al. 1663 (st), Little et al. 305 (US).

PERU. AMAZONAS: Prov. Bagua, Cajaruro,S of Bagua,14Oct 1952(o), Diazs.n. (US);Prov.Chachapoyas, Balsas, 23 Feb 1984 (st), D. N. Smith6173 (BG, MO). CAJAMARCA: San Ignacio, Caserio Calabozo, 30 Sep 1986 (a'), Diaz S. 2029 (BG); San Ignacio, 1200 m, 3 feb 1988 (a), Genry et al. 61057 (BG, NY); Colassy, 2700 m, 2 Nov 1971 (a'), Woytkowski7055 (GH, K, MO, US).

Cuzco: Prov.Convenci6n,RosarioMayo, 1100 m, 14 Oct 1968 (a'), ChavezA. 273 (MO); Prov. La Convenci6n,

Quillabamba,Salaspampa,1100 m, 26 Oct 1986 (? flfr), Nuniezet al. 6349 (BG); Prov. Convenci6n, Potrero, 1200 m, 29 Oct 1949 (? fl), Vargas C. 8523 (LIL, MO, NY); Idma, 1400-1600 m, 30 Oct 1949 (a'), Vargas C.

8535 (LIL, MO, US). HUANUCO: Rio Pachitea,ca. 20 km upstreamfrom Rio Ucayali, 14 Sep 1967 (? fl), Schunke V 2147 (F, K, MOL); Prov. Pachitea, Dtto. Honoria,Rio Pachitea, nr. Tournavista,10 Nov 1967 (a'), Schunke V 2311 (F, GH, MO, MOL, NY); Prov.

Leoncio Prado,Dtto. RupaRupa,NE of Tingo Maria, nr.Mapresa,2 Oct 1978 (? fl), SchunkeV 10623 (BG, MO). JUNIN: Rio Colorado, nr. jct. with Rio Chanchamayo,7 Feb 1983 (st), Gentryet al. 40115 (MO); GUYANA.East Coast, Demerara,Good Hope, 15 San Ram6n,8 May 1961 (9 fr), SchunkeV 4100 (MO). May 1960 (? fl), S. A. Harris EC.4 (K); East Coast,Tri- LAMBAYEQUE: Prov. Lambayeque,between Olmos & 26 Mar1985(a' + ? fr), Rios T 270 (MOL). umph 18 Sep 1952 (? fl-fr), Fanshawe 3484 (= FD Tongoriape, 7080) (K); "Coastregion,"Jun 1889 (? fl-fr), Jenman LORETO:Rio Amazonas, Yanayacu, opposite Indiana, 5067 (K); Berbice,Jan 1890 (? fr), Jenman 5869 (K). 12 Mar 1981 (? fr), Gentry et al. 32060 (F, MO, NY, ECUADOR.BOLiVAR:Valle de Limon,Acosta S. TEX, U); Reserva Nacional Pacaya-Samiria, Rio 6453 (F).ESMERALDAS: ReservaForestalde JardinTropi- Samiria, 23 Oct 1970 (9 fl-fr), Grandez et al. 2053 (BG); cal, Universidad TecnicaLuisVargasTorres,4 Feb 1991 Dtto. Sargento Lores, Quebrada de Ponguanilla, below (st), Gentry et al. 73083 (MO);Rio Guayllabamba,10 Tamshiyacu, 14 May 1980 (a'), McDaniel et al. 23674 km E of Quininde,5 Oct 1965 (st), Little et al. 21233 (F, MO, NY, US); Dtto. Mazan, Baradera Mazan, trail (NY,QAME,US). LOJA:BetweenPuenteBoquer6n& Rio Amazonas-Mazan,16 Apr 1973 (? fl-fr), Rimachi Gonzanama,1400 m, 9 Oct 1955 (9 fl), Asplund 18085 Y 195 (F, MO, NA, NY); Prov.Requena,Rio Ucayali, (B, K, LL, NY, P, S); rd. Catacocha-Macara,km 7, nr. Jenaro Herrera, 13 Mar 1984 (a'), Spichiger et al. Lucarqui,12 Apr 1980 (? fl-fr), Harling et al. 18208 1319 (MO);Prov.Maynas,PuertoAlmendras,Rio Nanay, (BG); Las Cochas, 10 km N of Zapotillo, 8 Aug 1975 3 Nov 1985 (9 fl), R. Vdsquezet al. 6891 (BG, MO, NY, (st), Samaniago et al. 39 (US); Cant6nMacara,Achima, TEX);Yurimaguas,Oct-Nov 1929 (? fr), LI. Williams 2 Jan 1989 (? fl-fr), Palacios 3296 (MO,NY, QAME). 4598 (F, GH). SANMARTiN: Pongo de Cainarachi,Rio Los Rios: Represa Daule-Peripa, Pto. Palmar, 1 Jun 1985 (st), Anonymous216 (MO); Cant6n Vinces, between Mocahi & Palenque, Estero Pefiafiel, Jauneche forest, 24 Jan 1981 (? fr), Gentry et al. 30741 (AAU, GB, MO, S, U); Pichilingue, 17-24 May 1943 (st), Little 6453 (F, K, NY), 6462 (K, NY, US), 6510 (F, K, NY), 9 Nov 1982 (9 fl), Pennington et al. 10689 (QCA, QCNE, U). MANABi:MachalillaNationalPark,EsteroMantaBlanca, S of AguaBlanca,24 Jan 1991 (a'), Gentry et al. 72721 (MO, QCNE). MORONA-SANTIAGO: Macas, 1070 m, 27 Sep 1975 (st), Little et al. 458 (US); nr. Yunganza, 20 km N of Lim6n, 19 Nov 1982 (Y fl), Pennington et al. 10786 (NY, QCA, U); Bomboiza, Misi6n Salesiana, 29 Sep 1985 (st), Pujupet RBAE.1051 (NY, QAME). PICHINCHA: Between Alluriquin & La Union del Toachi,

Cainarachi, Sep-Oct 1932 (a'), Klug 2607 (A, BM, F, G,

GH, K, MO, NY, US); Prov. Mariscal Caceres, Dtto. TocacheNuevo, Rio Huallaga, Fundo San Rafael, 18 Sep 1970 (9 fl-fr), Schunke V 4425 (F, K, MO, MOL, NY, US); Dtto. Tocache Nuevo, Quebrada de Santiago, E of PuertoPizana,29 Jul 1973, (a'), Schunke V 6510 (GH,MO,NY); Moyobamba,24 Oct 1948 (a), Scolnik 1212 (LIL, NY); Tarapoto,nr. Rio Chilcay, 14 Feb 1947 (? fl), Woytkowski35079 (F, LIL, MO). TUMBES: Cerros de Amotape, ca. 25 km SE of Cherralique, 10 Jun 1987 (st), Gentry et al. 58292 (MO); Tumbes, 14 Feb 1964 (a'), Vargas A. 34 (F, MO). UCAYALI: Prov. Coronel

Portillo, Leoncio Prado (Yarina-Cocha), 28 Mar 1981 (a'), R. Vasquez et al. 1529 (BG, MO), 14 May 1984 (a), R. Vesquez 4958 (BG, NY), 2 Apr 1988 (? fr), R.

TAXONOMICTREATMENT Vdsquezet al. 10516 (BG); Pau-Cocha, nr. Pucallpa, 5 Dec 1951 (a), Woytkowski6295 (MO). BRAZIL. ACRE: Rio Jurua, 1 km upstream from Colonia Rodriguez Alvez, 29 Sep-10 Oct 1986 (st), Campbell et al. 11001 & 10486 (BG); Mun. Sena Madureira, Rio laco, 4 Oct 1980 (9 fl), Ferreira et al. 2766 (MG, MO, NY, U, US); Mun. Senado Guiomard, BR.317, km 23, 11 Oct 1980 (st), Ferreira et al. 2822 (MG, MO, NY, U, US); Mun. Rio Branco, rd. to Quixda, km 11, 18 Oct 1980 (9 fr), Ferreira et al. 2957 (MO, NY, U) & Lowrie et al. 576 (MO, MG, NY, TEX, U, US). AMAZONAS: Itacoatiara,Igarape do Lago de Serpa, 5 Aug 1913 (a' + 9), Ducke MG 12489 (MG); Fonte B6a, 26 May 1945 (a), Fr6es 20969 (IAN, K, NY, US), (? fl-fr), Fr6es 20970 (F, IAN, K, NY, US): Mun. Carauary, Marapata, 25 May 1933 (? fl + e), Krukoff 4570 (A, BM, F, K, MICH, MO, NY, U, US); Rio Purus, between Campina & Tambaqui, 19 Jun 1971 (? fl-fr), Prance et al. 13386 (GH, K, MG, NY, U, US); JuruaMirim, Aug 1901 (e), Ule 5714 (B, G, K, MG, MO). BAHIA: Between Ajuda & P6rto Seguro, 28 May 1962 (? fr), Duarte 6739 (NY, RB, U); Mun. Caravelas, between Barrade Caravelas & Ponta da Areia, 17 Mar 1978 (9 fl-fr), Mori et al. 9631 (NY, U); Prov. Jequie, 3 km W of Jequie, 19 Nov 1978 (a'), Mori et al. 11196 (NY, U); Sao Desiderio, 14 Oct 1989 (? fr), N. Silva et al. 89 (BG, US); Rio do Meio, rd. to Aurelino Leal, 13 Jan 1971 (9 fl-fr), Santos 1308 (NY, U). CEARA: Without locality, 12 Jul 1859 (e' + 9 fl-fr), Allemdo et al. 1437 (R); Fortaleza, 12 Nov 1955 (? fl), Ducke 225 (U), Oct 1957 (e), Ducke 2625 (U); Guaramiranga, 3 Aug 1908 (st), Ducke MG 1478 (MG); Lag6a, (9 fr), Ducke MG 2058 (BM, G); Sao Antonio, Serra da Meru6ca, 8 Jan 1961 (9 fr), A. Fernandes 2077 (U). DISTRITO FEDERAL: Brasilia, C6rrego Quilombo, 18 Sep 1980 (? fl), Heringer et al. 5516 (MO, NY, US); Brasilia, Rio Sao Bartelomeu, 1 Sep 1981 (e), Heringer et al. 7358 (MO). ESPiRITO SANTO: Mun. Nova Venecia, Serra de Cima, 15 Nov 1953 (e), Duarte 3895 (RB, U); Goitacazes, Rio D6ce, 29 Mar 1934 (9 fl), Kuhlmann 89 (= RB 34261) (US). MARANHAO: Island of Sao Luiz, Jan 1940 (? flfr), Fr6es in Krukoff 11804 (A, F, K, MO, NY, U, US) & 11849 (A, F, MICH, NY, US); Chaval, 13 Apr 1949 (9 fr), Pires 1505 (NY). MATO GROSSO: Caceres, Sep 1911 (o'), Hoehne 4657 (R); 50 km N of Xavantina, Rio Vau, 9 Oct 1964 (? fl), Prance et al. 59316 (NY, US); Mun. Santa Terezinha, rd. to Santa Terezinha, 7 km E of jct. with BR.158, 13 Oct 1985 (9 fr), Thomas et al. 4364 (BG, NY, US); Mun. Vila Bela Santissima, 41 km NNW of Pontes e Lacerda, 31 Oct 1985 (9 fr), Thomas et al. 4731 (BG, NY, US). MATO GROSSO DO SUL: Mun. Rio Brilhante, rd. BR.267, 22 Oct 1970 (? fl), Hatschbach 25079 (C). MINAS GERAIS: Coraqao de Jesus, 26 Dec 1963 (9 fl), Jestis 65 (NY); P6rto Novo da Cunha, 16 Nov 1960 (9), Lanna Sobrinho 27 (K, U); Vi9osa, 17 Mar 1930 (e'), Mexia 4475 (A, BM, GH, MO, NY, US), 16 Oct 1930 (o), Mexia 5189 (A, BM, F, GH, K, MICH, MO, NY, U, US), 20 Nov 1930 (9 fl-fr), Mexia 5329 (A, BM, F, GB, K, MICH, MO, NY, TEX, U, US); Lagoa Santa, Oct 1864 (e + 9 fl-fr), Warming1939 (C, P). PA"A:

65 Mun. Altamira, Igarape Ipixuna, 3 Nov 1985 (a'), Balhe et al. 1697 (BG); Morro de Obidos, 3 Nov 1950 ($ fl), Black et al. 50-10516 (GH, IAN, NY); Altamira, 12 Aug 1971 (st), Cavalcante et al. 2896 (MG); Prainha,22 May 1903 ($ fl), Ducke MG 3648 (BM, G, MG, U); Alemquer, 2 Jan 1904 (2 fl-fr), Ducke MG 4957 (MG); Cauchauhaya, 29 Oct 1898 (? fl-fr), Huber MG 1404 (G, MG); Rio Jari, Monte Dourado, 13 Jan 1969 (R fr), N. T Silva 1625 (NY, US); Santarem, Nov 1849-Mar 1850 (d + ? fr), Spruce s.n. (or 307, 427, 651, & 1705)(B, BM, G, GH, NY, P). PARANA: Ja9arehy,12 May 1915 (st); Dusen 17034 (F, GH, K, NY); Mun. Antonina, Ponta da Pita, 22 Nov 1966 (e' fl-fr), Hatschbach 14078 (P, US); Mun. CerroAzul, Rio Turvo, 5 Oct 1977 (e'), Hatschbach 40219 (MO, NY); Mun. Antonina, Mangue Maior Santo, 25 Nov 1983 (a), Hatschbach 47136 (US); Foz do Iguassu, 1 Feb 1949 (Y fl-fr), Schwarz 7360 (LIL). PERNAMBUCO: Tapera,Dec 1930 (2 fl), Tapera,Pickel 142 (B, BM, F); Olinda, Apr 1924 (9 fl), Pickel s.n. (US). PIAui: Without locality, 1846 (2 fr), Gardner 2002 (BM, G, P). Rio DE JANEIRO: Pinheiras, (e), Dutra 202 (R); Mun. Rio de Janeiro, Corcovado, Santa Teresa, 18 Feb 1862 (? fr), Glaziou 766 (C, P); Mun. Rio de Janeiro, Floresta de Tijuca, 22 Mar 1862 (? fr), Glaziou 766a (P); Ilha de Governador, 18 Mar 1879 (e), Glaziou 14281 (B, C, F, K, P, U, US). RIO GRANDE DO SUL: Brusque, 23 Oct 1951 (a), Klein 160 (US); nr. Sao Leopoldo, 7 Dec 1948 (? fl-fr), Rambo 38586 (B, LIL, P); Butterberg, nr. Montenegro, 13 Nov 1950 (? fl), Rambo 49142 (LIL, US), (a'), Rambo 49143 (B, GB, LIL); Matador, 23 Nov 1958 (? fl), Reitz et al. 7567 (G, NY, US). ROND6NIA: Mun. P6rto Velho, Campo Novo, 20 Mar 1987 (? fr), Nee 34443 (BG, NY); BR.421, 2 km W of Mineragao Campo Novo, 18 Oct 1979 (? fr), Vieira et al. 530 (MG, MO, NY, U, US). SAO PAULO: Pirapora, 4 Dec 1924 (a'), Hoehne SP 12911 (A, US), (? fl-fr), Hoehne SP 12911a (US); Cabreuiva, 4 Oct 1933 (? fl-fr), Hoehne SP 31010 (F, NY, US); Limeira, 15 Oct 1951 (e), Kuhn 21 (NY); Campinas, Dec 1894 (? fl-fr), Novaes SP 23849 (US). SANTA CATARINA:Matador, 18 Oct 1958 (a), Reitz et al. 4071 (B, GH, K, NY); Lauro Muller, 1957 (2 fr), Reitz et al. s.n. (NY, US); Mun. Mondai, Rio Uruguai (27?07'S, 53?25'W), 16 Oct 1964 (? fl), L. B. Smith et al. 12605 (GH, MO, NY, P, R, US); Mun. Itapiranga, 3 km W of Itapiranga,17 Oct 1964 (a), L. B. Smith et al. 12659 (B, C, LE, R, TEX, US). BOLIVIA. BENI: Prov. Marban, San Rafael, 6 Oct 1979 (? fl-fr), Beck 2731 (BG); Rio Beni, Cachuela Esperanza, 22 Oct 1923 (? fr), Meyer 330 (MO, U); Prov.Vaca Diez, nr.Alto Iv6n, 25 Jun 1991 (st), Stijjhoorn 659 (NY). LA PAZ: Tipuani-Guanai, Dec 1892 (? flfr), Bang 1652 (A, BM, F, G, GH, MICH, MO, NY, US), (a'), Bang 1737 (A, BM, F, G, GH, K, M, MICH, MO, NY, US, Z); Prov. Sud Yungas, Chulumani, 1600 m, 30 Oct 1981 ($ fl), Beck et al. 4868 (BG); Prov. Nor Yungas, rd. Yalosa-Caranavi, km 8.5, 1170 m, 24 Sep 1987 (a), Beck 13590 (BG); Mapiri region, San Carlos, 21 Nov 1926 (? fll-fr), Buchtien 647 (C, K, MO, NA, NY, US, Z); Prov. Nor Yungas, N of Caravavi, 1500-1600 m, 8 Nov 1976 (? fl-fr), Davidson 4875 (NY, US). SANTACRUZ:

66 Prov.Ichilo, 2 km W of El Cairo,5 Dec 1989 (e), Nee 37960 (MO, NY); Prov.Andres Ibaniiez,3 km NW of centerof Cotoca,4 Dec 1989 (a), Nee 37941 (BG, G, MO, NY, US); Prov.Warnes,11 km SE of Montero,6 Dec 1989 (? fl), Nee 37988 (NY); Prov.AndresIba-nez, 3 kmNW of centerof Cotoca,21 Dec 1989(9 fl-fr),Nee 38214 (MO,TEX);Prov.Ichilo,E side of Rio Yapacani, atjct. withRio Surutti,30 Oct 1900(a'), Nee 39586 (MO, 29 Sep 1990 NY); Prov.N4uflode Chavez,Perseverancia, (a'), Renjel18 (MO,NY); Prov.Sara,Buenavista, 28 Sep 1924 (? fl), Steinbach 6491 (A, BM, F, K), (), Steinbach

6491bis(BM, F, K, MO);Prov.Guarayos,3-4 km SW of confluenceof Rio San Pablo& Rio Negrode Caimanes, 15 Jun 1993 (? fl-fr), I. G. Vargaset al. 2574 (USZ). PARAGUAY. CENTRAL: Cord.de Altos, 10 Nov 1902 (a'), Fiebrig 411 (A, F, G, K, L, M), 16 Mar 1903 (o), Fiebrig983 (A, F, G, K, L, M); betweenRio Apa& Rio Aquidaban, 1908-1909 (o), Fiebrig 4394 (BM, G, K,

US, Z); Alto Parana,1909-1910 (? fr), Fiebrig 6242 (G, GH,K, L, LIL,US); Cord.de Altos, Oct 1885-1895 (e'), Hassler 1366 (BM,G, K, NY,P); RioYaca,Jan1900 (a'), Hassler 6917 (A, BM, C, G, K, MICH,MO, NY, US); Lagode Ypacarai,1913 (? fr),Hassler12125a (A, BM, C, GH,K, MO,NY,US, Z);VillaRica, 15 Feb 1928 (e), Jorgensen 3835 (A, C, GH, F, LIL, MO, NY); Centro

ForestalAlto Parana, 12 km W of Puerto Presidente Stroessner,2 Feb 1984 (a), Little 40116 (MO); Villa Elisa,

11 Apr 1961 (9 fr), Pedersen5890 (MO), 16 Oct 1959 (MO);CentroForestalAlto Parana, 12 km W of PuertoPresidenteStroessner,29 Nov 1984

(a'), Pedersen 5142

(? fr), Stutz de Ortega 1955 (MO); Depto. San Pedro,

Primavera,20 Oct 1950 (? fl-fr), Woolston1026 (NY). ARGENTINA. CORRIENTES: Depto. Capital, Corrientes, 23 Oct 1970 (e), Marunak 141 (LL, P); 17 km NW of San Carlos, 1-5 Mar 1985 (? fl), Tressens et al. 3071 (MO). FORMOSA:Depto. Pilcomayo, Estancia Rancho Negro, 3 Oct 1947 (V), Morel 3746 (LIL). MISIONES:Depto. Candelaria, Pindapay, 25 Nov 1945 (9 fl-fr), Bertoni 2431 (F, LIL); Puerto Le6n, 12-21 Jul 1914 (st), Curran 683 (F, US); Posadas, 20 Feb 1944 (? fl), T Meyer 5642 (BM, LIL, U); 48 km from Eldorada

FLORANEOTROPICA

Izabal:moraor moral.Honduras:mora.Nicaragua: mora.CostaRica: Chinanadega, Esteli,andMatagalpa: moro(tree)andmora(fruit). Alajuela:mora.Panama: Cuba:fusteteor fustic.Jamaica:fustic.PuertoRico: escambr6n de madeira,fustic,mora,palode mora.St. fustic.Cura,ao:mora, Lucia:boisd'orange.Grenada: Amazonas:incira;Atlantico, paloedoesje.Colombia: Bolivar, Boyaca, and Magdalena:mora;Caldas, All dinde.Venezuela: Cundinamarca, Huila,andTolima: fustic. states:mora;Merida:palode mora.Trinidad: Ecuador:Loja:sota;Los Rios:moralfino;Moronasota. Santiago: moral,leilapo,sota;Zamora-Chinchipe: Junin: Peru:Cajamarca: moreira; Cuzco:amarillo; encira; Huanuco,Loreto:inciraor insira,insiracaspi,insira SanMartin: amarilla, limulona, moral,tsajanki (Jivaro); insiracaspi,insiraamarilla; Tumbes:sota.Brazil:Acre and Amazonas:limao-rana;Bahia:amorabranca, amoreira, moreira,moreiraamarela;Ceara:tatajuba; Maranhao: juremade espinho,tatajuba; MinasGerais: moreira, tajuba, tatajuba; Rio Parana: amoreira, moreira; de Janeiro:tatajuba; SaoPaulo:tataiba,taiuva;Santa Catarina: amoreira, tajuva.Bolivia:LaPaz:mora;Santa Cruz:moraormoragrande(subsp.mora= morachica), tataguiba, tatayieva. Paraguay: catagiva, tatajyva. Argentina:Chaco:moraamarilla, moracolorada,tata-yyba'Corrientes: sayyu,tata-y5yba-pyta; cambapunta, mora, tata-yyba; Misiones:mora,tatayiba, tata-yyba. IntheupperAmazon Basinthelatexis usedto cure toothache; JivaroIndiansapplythesapto inflictedteeth, whichthencorrodeandcanbe easilyextracted(W.H. Lewis,pers.comm.).Theyellowwoodis durableand usedforconstruction andfurniture. Thespeciesis locallycultivatedforthefruits.

2b.Macluratinctoria(Linnaeus) Steudelsubsp.mora 26: 323,t. (Grisebach) VazquezAvila,Darwiniana 11. 1985; Berg, Proc. Kon. Ned. Akad. Wetensch., on Ruta Provincial 17, 24 Oct 1978 (? fl), Renvoize Ser.C, Biol. Med. Sci. 89: 244. 1986. Maclura 3252 (MO, US); Salto Iguazii, 19 Sep 1910 (9 fl), moraGrisebach, Abh.Konigl.Ges.Wiss.G6ttingen Rodriguez460 (LIL);PuertoLeon, 26 Jun 1909 (Y fl), Phys.Cl. 24: 86. 1879.Ioxylonmora(Grisebach) Venturi 16 (LIL). 0. Kuntze,Rev.Gen.P1.3:294. 1898.Chlorophora Label annotationsindicatethatthe species is intromora(Grisebach) Lillo,in Venturi& Lillo,Contr. duced in Guyana. However, judging from the distriConoc.Arb.Argentina63. 1910. Chlorophora bution of the species, its presence in Guyana could tinctoria(Linnaeus)B. D. Jacksonsubsp.mora be natural. (Grisebach) Hassler,AnnuaireConserv.Jard.Bot.

This subspecies shows a broad ecological amplitude, ranging from humid forest to dry thickets. Most peculiaris the presenceof the subspeciesin earlystages of the succession into varzeaforest along the Amazon River (and some southerntributariesin the upperpart of theriverbasin)as canbe seen in the distributionmap.

Local names and use. Mexico:SinaloaandOaxaca: moraor moral;Yucatan:moro de la tierra.Guatemala:

Geneve 21: 114. 1919. Chlorophora tinctoria

(Linnaeus)B. D. Jacksonvar.mora(Grisebach) Lillo,Bol. Mus.Hist.Nat.6: 10. 1925.Type.Argentina.Salta:Orin, 16 Oct 1873(a), Lorentz& 479(holotype,GOETT-n.v.; Hieronymus isotypes, B, G, NY, S). Fig. 9 MacluratrilobaRojasAcosta,Bull. Acad.Int.Geogr. Bot. 24: 211. 1914. Type. Not indicated.

TAXONOMICTREATMENT Chlorophora reticulata Herzog, Meded. Rijksherb. 27: 73. 1915. Type. Bolivia. Santa Cruz: nr. Santa Cruz, Feb 1911 (st), Herzog 1526 (holotype, L; isotypes, MO, NY).

Tree,to 10m tall,orshrub.Laminaelliptic,some1-8(-l 1)x 0.8-4 cm;apexshorttimesto suborbicular, acuminate to subacute;uppersurfacehirtellous,usuto scabrous;lowersurfacedensely ally scabridulous hirtellousto tomentose;smallerveins always(very) prominentbeneath;petiole0.2-1.2(-1.5) cm long. Staminateinflorescences: peduncle0.2-0.4 cmlong; spike3.5-4 cmlong;Pistillateinflorescences:peduncle 0.1-0.4cmlong;head0.2-0.5cmdiam.,to 1cmdiam. but in fruit;tepalsandbractsattheapexsubglabrous, stigma3-7(-9) mmlong. atthemarginstomentellous; Distribution(see Fig. 74.2). FromBoliviato Argentina;in subtropicalsemideciduous,sometimes (chaco)forest;to 1600m. brieflyinundated

67 Elsa, 13 Sep 1944 (a'), Rojas 12283 (F, LIL). JujuY: Depto. SantaBarbara,PalmaSola, 23 Nov 1970 (? flfr), Cabrera et al. 21103 (MO); Depto. Ledesma, Calilegua,Tomadel Arroyo del Medio, 22 Nov 1980 (? fr), Cabreraet al. 32219 (MO);betweenJujuy& La Mendieta, 24 Oct 1948 (? fl), Killip 39584 (US); San Pedro,Oct 1940 (a), Schreiter 11444 (A, F, LIL, U); Depto. Ledesma,Sierrade Calilegua,15 Oct 1927 (a), 5396 (A, BM,F,K, LIL,F,K,MO,US, Z). SALTA: Venturi

Rio Piedras,2-13 Nov 1911 (o), Rodriguez14 (A, F, LIL,US); Depto.Oran,betweenTartagal& Yarguarenta, Oct 1949 (a), Schreiter 11443 (A, BM, F, LIL, U); Depto.Oran,ObraGrande,19 Nov 1927 (2 fr), Venturi 5551 (A, B, BM, F, GH, K, LIL,MO, US). Local names. Bolivia: Santa Cruz: mora chica, mora de la pampa, uvillo. Paraguay: tatajyba or tatA-yyba. Argentina: Formosa: mora; Salta: mora, mora colorada, mora del campo.

Representative specimens examined. BRAZIL. LI.ARTOCARPEAE MATOGROSSO:Mutinho, 26 Oct 1980 (9 fr), Pires et Trees or shrubs, mostly dioecious, with or without al. 17337 (BG, MG). uncinate hairs. Leaves alternate and distichous or opBOLIVIA. SANTACRUZ:Prov. Cordillera, Bafiados posite; lamina entire or incised; venation pinnate or del Izozog, 13 Mar 1991 (st), Navarro et al. 388 (LPB); entire, dentate, crenate, serrate, or Prov. Andres Ibafinez,12 km E of center of Santa Cruz, subpalmate; margin lateral to semiamplexicaul or fully lobate; stipules rd. to Cotoca, 20 Aug 1987 (ad), Nee 35669 (MG, MO, racemose, spicate, clavateamplexicaul. Inflorescences 28 Nov NY); Prov. Andres Ibafiez, 5.5 km NE of Cotoca, 1990 (? fl-fr), Nee 40044 (MO, NY). TARIJA:Prov. Gran to globose- or to discoid-capitate, or (pistillate ones) Chaco, rd. Villamontes-Camiri, km 29, 29 Sep 1985 (2 uniflorous, bracteate (or ebracteate), sometimes disfl), Beck et al. 11643 (BG). tinctly involucrate, the bracts often peltate. Staminate PARAGUAY. CHACO:Parque Defensores del Chaco, flowers: tepals 2-4; stamens 1-4, (almost) straight in nr. Madrej6n, 12 Jul 1985 (? fl), Brunner 1237 (G, MO); the bud, not bending outwards suddenly and elastically; nr. Concepci6n, Sep 1902 (? fl), Hassler 7329 (A, BM, anthers extrorse, latrorse, or introrse, the connective C, G, K, MO, NY), (a'), Hassler 7329a (A, BM, G, K, mostly broad; pistillode usually lacking. PistillateflowLIL, MICH, MO, NY, P); Fuerte Olimpo, 22 Oct 1946 ers: tepals 2-4(-5), mostly connate and forming a tu2 Cerro Oct Le6n, 13652 LIL); (NY, fl-fr), Rojas (2 bular, lobed or entire perianth, less commonly free or 1979 (e'), Schinini et al. 18030 (MO). NEEMBUCU: lacking; ovary free or adnate to the perianth; stigmas 2, Bernardi 9 1978 Nov (? fl-fr), Curupayty, Humaita, 18419 (MO, NY). equally long, or 1. Fruiting perianth enlarged, ? fleshy, ARGENTINA. CHACO:Las Palmas, 11 Sep 1917 greenish or colored; fruit free or adnate to the perianth (? fl), Jorgensen 2159 (GH, LIL, MO, US); Fontana, (or enclosed by fleshy bracts), sometimes a dehiscent Nov 1938 (? fl-fr), T. Meyer 3185 (LIL); Colonia drupe; seed large and (almost) without endosperm or Benitez, Nov 1941 (? fr), Schulz 1834 (LIL), 12 Nov small and with endosperm, the vascularization of the 1968 (? fr), Schulz 16475 (F, MO). CORRIENTES:Depto. testa various; embryo various. San Luis del Palmar, 18 km SE of San Luis del Palmar, The tribe comprises 12 genera and 87 species: 2 Nov 1975 (? fl), Crist6bal et al. 1378 (MO); Depto. with one species in New Guinea (cf. CorAntiaropsis, Nov 1986 15 Formosa, Formosa, Riacho San Hilario, ner, 1962); Artocarpus, with 50-55 species in Asia(? fl), Cristobal et al. 2147 (F); Depto. Capital, Corrientes, ruta 12, km 10, 23 Oct 1970 (di), Marufiak 141 Australasia (Jarrett, 1959-1960, 1975); Bagassa, with (MO); Depto. Mburucuya, Estancia Santa Teresa, 16 one species, neotropical; Batocarpus, with 3 species, Nov 1950 (e), Pedersen 847 (C, LE, MO, NY, P, U, US), neotropical; Clarisia, with 3 species, neotropical; Estancia Santa Maria, 18 Nov 1950 (9 fl-fr), Pedersen Hullettia J. D. Hooker, with 2 species in Asia (Jarrett, 856 (A, C, MO, NY, P, U, US); Depto. Capital, Corrientes, 1959-1960); Parartocarpus Baillon, with 2 species in Molina Punta, 3 Jan 1976 (? fr), Schinini et al. 12320 Asia-Australasia (Jarrett, 1959-1960); Poulsenia, with (MO); Depto. General Paz, ruta 5, Lomas de Vallejos, 24 one neotropical; Prainea J. D. two) species, (or Nov 1978 (? fr), Schinini 16043 (F, MO). FORMOSA: in Asia (Jarrett, 1959-1960); 4 with species Hooker, Puerto El Colorado, Rio Bermejo, 28 Aug 1944 (? flSorocea, with 14 species, neotropical; Sparattosyce, fr), Rojas 12127 (LIL); Depto. Pilcomayo, Isla Puerto

FLORANEOTROPICA

68 with one species in New Caledonia(cf. Corner,1962); and Treculia,with 3 species in Africaand Madagascar (Berg, 1977c). The tribeArtocarpeaelacks the homogeneityof the otherfour tribes of the Moraceae.It shows featuresof the other four tribes and is, to some extent, a crosssection through the family. The inflorescence varies from simple to complex and the flowers from normal moraceousto reduced. It is not clear whetherthe heterogeneitycan be ascribedto polyphylyor whetherthe triberepresentsisolatedfragmentsof a natural(monophyletic) group.The otherfour tribesof the Moraceae show more or less distinct centers of distributionor clear links between majorphytogeographicalregions, but not so for the Artocarpeaeas a whole. The tribe can be subdivided into three groups to which one could assign the rank of subtribes.These groups are phytogeographicallyseparatedbut do not show clear morphologicallinks. One of the groups comprises Artocarpus and the allied Malesian genera Prainea, Paratocarpus, and

Hullettia,andtheAfrican-Madagascan genus Treculia. In this group,the most complex inflorescencesamong the Artocarpeaecan be found, some reminiscent of those in Castilleaeandothersof those of Dorstenieae. The infructescencescan become very large. A second group comprises the monotypic genera AntiaropisandSparattosyce.The inflorescencesof the formerresemble those of the Castilleae and the inflorescencesof the latterresemblethoseof theFiceae.Both have dehiscent drupesexpelling the endocarpbody, a feature not occurring in the other Artocarpeae, but presentin Dorstenieaeand Moreae. The thirdgroupcomprisesthe five neotropicalgenerarevised in the presenttreatment.This grouphas the simplesttype of inflorescenceandshows in variousrespects similaritiesto the Moreae,but the generaarefar moreclear-cutthanthose of the lattertribe.This group of generais distinctlyassociatedwith the SouthAmerican continent,in contrastto the tribeMoreae,which is centered in the northwesternpart of the Neotropics: CentralAmericaandthe northernAndeanregion.

Key to the neotropicalgenera of the Artocarpeae 1. Leaves opposite............................................... 4. Bagassa 1. Leaves alternate. 2. Pricklespresenton leafy twigs, leaves, and/orstipules............................................... 8. Poulsenia 2. Prickles absent. 3. Staminateinflorescencesspicate,racemose,or subcapitate;staminateflowers? distinctand with (3-)4 stamens;pistillateinflorescencesracemoseto subumbellate,at least in fruiting 7. Sorocea state; fruitingperianthat full maturityblack................................ 3. Staminateinflorescencesspicate; staminateflowers crowded,indistinct,and with 1-3 stamens;pistillate inflorescencescapitateor uniflorous;fruitingperianthat full maturity yellow, orange, or red. 4. Inflorescencesaxillary;pistilate inflorescencesglobose-capitate ................................5. Batocarpus 4. Inflorescencesoften on the older wood, or if axillary,then the barkat the base of the trunkreddish;pistillateinflorescencesuniflorousor multiflorousand discoid-capitate ..... 6. Clarisia Bagassa 4. BagassaAublet,P1.GuianeSuppl.15. 1775;Bentham & Hooker,Gen. P1.3(1): 362. 1880;Berg,Fl. Guianas 21: 15. 1992. Type.Bagassa guianensisAublet. Laurea Gaudichaud,Voy. Bonite, Bot. Atlas t. 88. 1844. Type. Laurea tiliifolia Gaudichaud (= Bagassa guianensis Aublet).

Trees, dioecious; uncinate hairs lacking. Leaves (usually)opposite,decussate;laminatriple-nerved,entire to 3-lobed (to -parted);margin entire to crenate; stipules free, lateral.Inflorescences pairedor solitary in the leaf axils, pedunculate,bracteate.Staminateinflorescences spicate,with an abaxialsterilestrip;flowers in + distinctlongitudinalrows, close together;perianth 4-parted;stamens 2; anthersbasifixed, latrorse; pistillodepresent.Pistillateinflorescencesglobose-capi-

tate;flowersclose together;perianth4-lobed to -parted; ovary free; stigmas 2, filiform. Fruitingperianth enlarged,+ fleshy,green;fruita drupelet;endocarpthinly crustaceous;seed small;endospermpresent;testathin, not vascularized;embryowith plane and equalcotyledons and a long, curvedradicle. ThegenusBagassawasestablishedbyAublet(1775). Thedescriptionof the species,B. guianensis,was based on juvenile material(with scabrous, 3-fid leaves) and separateinfructescences. Materialin theadultstage(with entireandsmoothleaves)with staminateinflorescences wasdescribedasPipertiliifoliumby Desvaux(1825) and (in the Urticaceae)as Laurea tiliifolia by Gaudichaud (1844); the latter was combined with Bagassa by Benoist (1933). Materialwith adult leaves and pistillate inflorescences was described as B. sagotiana by

TAXONOMIC TREATMENT

69

BenthamandHooker(1880).Thematerialwithadult coriaceousto chartaceous;apex(or apices)short-acumileavesandthatwithjuvenileleaveshasbeenregarded nateto acute(whenjuvenile long-acuminate);basecoras distinctunderB. tiliifolia andB. guianensis, respec-

dateto subcordateto rounded;marginentireto crenate (whenjuvenile to serrate);upper surfacesubglabrous (when juvenile scabrousto scabridulous);lower surface densely puberulousto tomentellous (whenjuveBagassa guianensis is deciduous and forms huge nile scabrousto scabridulous);venation almost plane trees.Ithasyellowishwood,flakyandfissuredbark, above,prominentbeneath;tertiaryvenationscalariforn; andproduceslargeamountsof drinkable latex.Itis the petiole3-10(-14) cm long, puberulousto tomentellous; onlyneotropicalMoraceaewithoppositeleaves.The stipules0.5-1.5(-2) cm long, puberulousto tomentelseedis relativelysmall,a featurethatcanbe associated lous, caducous.Staminateinflorescences:peduncleIwith the ecology of the species,often occurringas 2 cm long, puberulousto tomentellous,sometimeswith treein secondary (sub)pioneer growthandin forestsin 1 or 2 ovate bracts;spike 4-12 cm long, 3-5(-10) mm tree-fallgaps. diam., sometimes irregularlyconstrictedand/orbulgAs thetreesof thespeciescanbecometallandare ing; perianthca. 1 mm long, minutelypuberulous;filadeciduous, pollenmaybereleased fromthespicatestami- ments 1-2.5 mm long; anthers0.2 x 0.2 mm;pistillode nateinflorescencesby wind.Dispersalof the seeds, subulate,0.5-1 mm long;bractsovate to oblong to lanincludingthoseinfalleninfructescences, is carriedout ceolate to subovateto subovateto (sub)spathulate,ca. bya widerangeof animals,includingmonkeys,birds, 1 mm long, minutely puberulous.Pistillate infloresdeer,rodents,andtortoises(vanRoosmalen,1985). cences:peduncle0.7-1.5 cm long, 2-4 mmthick,puberBagassais probablycloselyrelatedtoBatocarpus. ulous to tomentellous;head 1-1.5 cm diam.;perianth Intheoppositeleaves,thedifferencesbetweenjuve- ca. 2 mm long, the apex minutely puberulousoutside, nileandadultleaves,andtheshapeof theinflorescences, theupperpartdenselyminutelypuberulousinside;style B. guianensisshowsstrikingresemblances withthe 0-0. 1 mm long; stigmassubulate,ca. 1-1.5 mm long; paleotropical Broussonetia papyrifera (Linnaeus) bractslanceolateto spathulateto obovateca. 1mm long, Ventenat,a memberof thetribeMoreae. puberulous.Infructescences2.5-3.5 cm diam.; fruiting perianth1.2-1.5 cm long; fruitobovoid ca. 7-8 mm long, often ? distinctly stipitate;seed ca. 3 x 2 mm.

tively,andtreatedas suchin theFloraof Suriname by Berg(1975),althoughwithsomedoubt;thedoubtdiswhenfieldworkwascarriedout. appeared

1. BagassaguianensisAublet,P1.GuianeSuppl.15,

Distribution (see Fig. 74.3). In the Guianas and Brazil (Amapa,Para,Maranhao,Roraima,and apparFrenchGuiana.Withoutlocality,(?), Aublets.n. in the southwestemAmazonBasin:Mato ently disjunct (holotype,BM;isotype,NY). Fig. 10 GrossoandRondonia);in primaryforeston terrafirme Piper tiliifolium Desvaux, in Hamilton,Prodr.P1. andin secondarygrowth;at low elevations. Ind.Occ.4. 1825,non Schlechtendal & Chamisso, 1831.Type.FrenchGuiana.Withoutlocality,(od), Representative specimens examined. GUYANA. probablyMartin s.n. in herb. Desvaux (holo- MazaruniStation, 22 Jul 1939 (6'), Fanshawe239 (= type, P; isotype,B). FD 2975) (K, NY), 6 Mar 1941 (? fr), Fanshawe641 Laureatiliifolia Gaudichaud,Voy. Bonite, Bot. At- (= FD 3377) (K, NY); RupununiDistrict,Kumukowau las t. 88. 1844; D'Alleizette, Explicationet de- R., 14 Feb 1994 (? fl-fr), Jansen-Jacobs et al. 3767 scriptiondes planchesde l'Atlas de Voy.Bonite. (BG); KanukuMts., Moku-mokuCr., 31 Mar-16 Apr 157. 1866, descr.;Weddell,Arch.Mus. Hist.Nat. 1938 (e), A. C. Smith3543 (A, B, F, G, IAN, K, LE, Paris9: 444, t. 12D. 1856-1857;Weddell,in De MO,NY, P, U, US); KanunuMts., Wabuwak,Nov 1948 Candolle,Prodr.16(1): 235(14). 1869. Bagassa (d'), Wilson-Browne 432 (= FD 5827) (K, NY). tiliifolia(Desvaux)Benoist,Arch.Bot. Mem5(1): SURINAME.Sectie 0, 18 Nov 1915 (st), BW 1347 31. 1933. Type. Gaudichaud,Voy. Bonite, Bot. (U);Brokopondo 19 Mar1964 District,S of Kabelstation, Atlas t. 88. 1844, probablydrawnfrom Martin (st), Donselaar1097 (U); Camp8, rd. to Sarwa,24 Jun s.n., FrenchGuiana,withoutlocality,(e), in P. 1971 (st), Elburg LBB 13403 (U); Nickeri District, Bagassa sagotiana Bureauex Bentham& Hooker, KabeleboDamarea,22 km SW of AvanaveroDamsite, Gen. P1.3(1): 362. 1880. Type. FrenchGuiana. 17 Nov 1976 (st), Heydeet al. 128a (U);NassauMts.,7 Maroni,(? fr), Melinons.n. (holotype,P). Mar1949(st),Lanjouwet al. 2433 (K,NY,U); LelyMts., 26 Sep 1975 (st), Lindeman& Stofferset al. 403 (U); Tree,to 45 m tall, deciduous. Leafy twigs 2-6 mm MapaneCr., Jodensavannearea, 2-27 Apr 1953 (st), andsca- Lindeman3685 & 3712 (U); SurinameR., nr.Goddo,30 totomentellous thick,puberulous (hispidulous brouswhenjuvenile). Lamina(sub)cordiformto ovate Jan 1926 (e fr), Stahel (Exp. Wilhelminagebergte) 152 entireto 3-lobed(when (U); Powaka,nr. Carolina,Nov 1943 (st), Stahel (Sur. to ellipticorto suborbicular, juvenileto 3-parted), 6-22(-30) x 4-17(-23) cm,sub- Woodherb.)194 (K, NY, U, WIS). t. 376. 1775; Berg, Fl. Guianas21: 15. 1992. Type.

70

FLORANEOTROPICA

(D

G)~J

I

wI -

Fig. 10. Bagassa guianensis. 1. Leaf (Zarucchi et al. 2663). 2. Leaf (Oldeman B.642). 3. Leaf (Berg s.n.).

4. Staminateinflorescences(Zarucchiet al 2667). 5. Younginfructescence(OldemanB.642). 6. Staminateflower (A. C. Smith3543). 7. Stamen.8. Pistillode.9a-c. Bractsof staminateinflorescence(A. C. Smith3543). 10. Pistillate flower. 11. Pistil. 12. Fruitingperianth.13. Fruit.14. Seed. 15. Embryo.16a-c. Bractsof pistillate inflorescence (Berg et al P.F18385). [By P. Pardoen, Utrecht.]

TAXONOMICTREATMENT

71

FRENCHGUIANA.Rd. to Mana,ca. km 6, 6 Nov Local names and uses. Guyana:cow wood, kato1956 (st), BAFOG7591 (NY, P, U), 6 Mar 1957 (st), war,tuwue,yawahedan.SurinaIne:gele bagasse,jawaBAFOG 7688 (NY, P, U); rd. to Acarouany,Camp hedan, kauhoedoe or kaw-oedoe. French Guiana: Charvein,25 Mar 1957 (st), BAFOG7696 (P, U); La bacasse or bagasse, odon or odoun (Paramak).Brazil: ComteR., 0.5 kmaboveSautBief, 18 Jan1957(st),Bena Maranhao:tatajuba,tareka'y(Ka'apor);Para:amapa1302 (U); Charvein, 8 Jan 1914 (st), Benoist 517 & 518 rana, taraiko'i(Tupi),tatajuba;Roraima:tatajuba. (U, P); Saul,4 Mar1971(st), Hijmanet al. 211 (NY,U); Timber is used for nautical constructions. The Mana,Aug 1854 (? fr), Melinon 124 (P); Ile de Cayenne,GrandMatoury,2 Sep 1966 (9 fr), OldemanB.628 infructescencesareedible. (CAY,P, U); Orapu,8 km fromKaw,13 Oct 1966 (2 fr), OldemanB.642 (NY,P, U); Stationdes Nourages,27 Feb Batocarpus 1987 (st), Prevost2221 & 2501 (BG);upperCamopiR., CriqueCacao, 13 May 1987 (st), Sabatieret al. 1653 5. Batocarpus Karsten, Fl. Columb. 2: 67. 1863; (BG);Acouarany,1857 (st), Sagot 522 (BM, G, K, P), Fosberg, Proc. Biol. Soc. Wash. 55: 99. 1942; 1858 (st + ? fr), Sagot 522bis (G, P). Emygdio de Mello Filho & Emmerich,Bol. Mus. BRAZIL. AMAPA:ReservaINCRARio Falsino,22Nac. Rio de Janeiro,Nov. Ser.Bot., No. 37: 1. 1968. 26 Aug 1986 (st), Campbellet al. 14542 (BG); Rio Falsino, Type. Batocarpus orinocensis Karsten. 10 km upstreamfromconfluencewith Rio Araguari,20 Aug 1983 (? fl), Rabelo et al. 2351 (BG,NY). MARANAnonocarpusDucke,Arch.Jard.Bot. Rio de Janeiro HAO: Mun. Mon9ao, Ka'aporIndian Reserve, 9 May 3: 38. 1922. Type.AnonocarpusamazonicusDucke (= 1986 (st), Balee 2298 (NY); Mun.Moncao,Rio Turiqu, B. amazonicus(Ducke) Fosberg). P.I.Guaja,29 1987(st), Balee 3498 (NY);Rio MaracassTrees,dioecious; uncinatehairs presentor absent. ameregion,14 Jul 1932(q fr),Fr6es1783 (A, BM,F, G, Leaves alternate and distichous; lamina pinnately K, MO, U). MATTO GROSSO: Rio Aripuana,nr.Nucleo PioneiroHumboldt,10 Oct 1973 (? fl-fr),Berg et al. P. veined; marginentireor dentate;stipules free, lateral. 18385 (INPA, NY, U), 3 Oct 1873 (e), Berg et al. R 19846 Inflorescencessolitary(or paired)in the leaf axils, pe(INPA, MO, NY, U). PARA: Mun. Altamira, Igarape dunculate,bracteateor ebracteate;embeddedyellow Ipixuna,3 Nov 1985(st),Baleeet al. 1771(BG);RioXingu, dye-containing "glands" present in the perianths, AssuriniIndians,11 Jun 1986 (st), Balee 2370 (NY);nr. bracts,pericarp,and/ortheapicesof the stamens.StamiBelem, Belterra, 17 Jul 1947 (d), Black 47-1027 (COL, nate inflorescencesspicatewith an abaxialsterilestrip; IAN,INPA,NY,P, U, US, VEN);Rio Tapaj6s,BoaVista, flowers in longitudinalrows, close together;perianth 8 Aug 1932 (c'), Capucho365 (F, IAN), 1933 (? fr), with 3-5 free or basally connatetepals; stamens 1 (or Capucho510 (F, IAN, US); Belem, BosqueMunicipal, 2); anthersbasifixed, introrseto latrorse;pistillode ab27 Jun 1942 (? fl-fr),Ducke946 (F, IAN,MG,MO,NY, sent. Pistillate inflorescences globose-capitate, with R), 17 Jun 1943 (a), Ducke 1236 (IAN, K, MG, MO, severalto many flowers;perianthtubular,subentireto NY,R); Rio Brancode Obidos,31 Oct 1919(? fr),Ducke 4-lobed, the lobes decussate; ovary free; stigmas 2, RB 8296 (RB), 11 Dec 1913(v), DuckeMG15133(MG); Rio Cumina-mirim,Castanhaldos Pedos, 27 Sep 1913 short,equallylong.Fruitingperianthenlarged,? fleshy, green;fruitwith the pericarpthin and(sub)coriaceous (e), Ducke RB 18271 (RB); Braganca, 1 Aug 1926 (e), DuckeRB 19453 (G, INPA,K, P, U); Rio Jaranca,P6rto or thickandthe endocarpcrustaceous;seed large,withde Moz, 10 Oct 1955 (? fr), Fr6es32173 (IAN);Mareo out endosperm;testa conspicuously and amplyvascuda Lagod,20 Aug 1898 (? fl-fr),GoeldiMG 1587 (BM, larized;embryolongitudinallyalignedwith 2 equaland G, MG);rd. Belem-Brasilia,20 Aug 1960(? fr),Oliveira thick cotyledons and a very short,apicalradicle. 1018(IAN);Rio Jan,Rio Branca,29 Jul 1969(Q fl), N. T The genus was establishedby Karsten(1862), who Silva 2553 (IAN,NY); Rio Jari,rd. Pilao-Repartimento, described a single species, Batocarpusorinocensis.In km 45, Sep 1970 ($ fr), N. T Silva 3329 (IAN); Rio Anapu,Portel,14 Mar 1942 (st), C. W Smithin Krukoff 1922 Ducke describedthe genus Anonocarpuswith a 12519 & 12525 (G, MO,NY). RONDONIA: Mun.Costa single species, A. amazonicus,and in 1939 he addeda Marques,2 km W of Rio Cautarinho,24 Mar 1987 (? second species, A.peruvianus. Fosberg (1942) united fr), Nee 34473 (BG, NY); Minera9ao Taboica, at the two genera andreducedA. peruvianus to the synMassangana,35 km WSW of Ariquemes,10 Oct 1979 onymy of B. orinocensis. A third species, B. costa(a'), Zarucchi et al. 2667 (GH, MO, NY, RB, U, US). ricensis, was described by Standley and L. 0. WillRORAIMA: SEMAEcol. Reserve,Ilhade Maraca,21 Sep iams (1952). The genus has been revised by Emygdio 1987 (st), Milliken et al. 580 (BG). de Mello Filho andEmmerich(1968), who recognized The species is uniformthroughoutits rangeof dis- a fourth species, B. maranhensis. tribution.Leaves ofjuvenile specimensdiffer considThe barkat the base of the trunkmay be reddish,as erably from those of adulttrees in shape, texture,and in Clarisia biflora and C. racemosa. The unicellular indumentum. hairs can be uncinate in Batocarpus amazonicus and

72

FLORA NEOTROPICA

variation inthetextureofthe (sub)globose-capitate largefruitsshowapeculiar B. costaricensis.Pluricellular, inB. amazonifromthinand(sub)coriaceous trichomesoccurin thethreespecies,althoughusually pericarp: in inthe cusandB. orinocensisto ratherthicklycrustaceous veinletsterminating InB. costaricensis, sparsely. is notconfinedto hydathodes attheend;such B. costaricensis. Thevascularization havegland-like leafmargin athickened partof thetesta,asoftenoccursinMoraceae. sometimesoccurinB. amazonicus. structures inthestructure of theinflorescences Thesimilarities inflorescences arespiThelongandslenderstaminate whenen- andflowerssuggestthatBatocarpusis closelyrelated cate(orprobably so).Thelobesoftheperianth inflorescences andflowers fis- to Bagassa.Thestaminate pattern of undulate largedinfruitformaremarkable thehumanbrainsurface,particularly areverysimilarto thoseof Clarisia.Thethreespecies sures,resembling in Batocarpus amazonicus and B. costaricensis. The recognizedin thepresentrevisionarecloselyrelated.

Key to the speciesof Batocarpus 1. Laminawith the marginentire;tertiaryvenationreticulate;hairs on the leafy twigs not uncinate .3. ...........................................................................................................................................................

B. orinocensis

1. Laminawith the marginusually? distinctlydentate,tertiaryvenationpartlyscalariform;some of the hairson the leafy twigs uncinate. 1. B. costaricensis 2. Laminaglabrousabove....................................................................... 2. Laminahairyabove, at least on the midrib......................................................................2. B. amazonicus

1. Batocarpus costaricensis Standley & L. 0. Williams, Ceiba 3: 25. 1952; Emygdio de Mello Filho & Emmerich,Bol. Mus. Nac. Rio de Janeiro,Nov. Ser. Bot., No. 37: 8, t. 3. 1968; Burger,Fieldiana Bot. 40: 111. 1977. Type. Costa Rica. Puntarenas: PalmarNorte de Osa, 27 Feb 1951 (? fr), Allen 5971 (holotype, EAP; isotypes, BM, F, GH, US). Fig. 11 Tree, to 30 m tall. Leafy twigs 1.5-4 mm thick, sparselyto densely brownish-puberulousto hirtellous, some of the hairsuncinate.Laminaelliptic to oblong, 7-25 x 3-13 cm, broadest at or above the middle, ? inequilateral, subcoriaceous; apex acuminate to subcaudate;base acuteto obtuseto subcordate;margin (often ? irregularly)dentate;upper surfaceglabrous; lower surfacesparselywhitish-puberulousto hirtellous to brownish-strigoseon the veins, some of the hairs uncinate;venationplane above, even the smallerveins prominentbeneath;lateralveins 8-14 pairs;tertiaryvenationpartlyscalariform;petiole 0.3-0.7 cm long, 1-2 mm thick, puberulousto hirtellous, some of the hairs uncinate; stipules 0.3-1 cm long, appressed-

the outerones conduplicate,upperpartof the perianth thickened,withnumerous,small,embeddedyellow dyecontaining "glands,"the apex brown puberulous to hirtellousoutside,the upperpartwith dense stiff brown hairs inside, the lower part with sparse brown hairs outside; ovary glabrous;style ca. 1 mm long; stigmas subulate, 3-4 mm long; bracts lacking (?). Infructescences 4-5.5 cm diam.,greenishat maturity;fruiting perianthca. 1.5-2 cm long, tuberculateat the apex; pericarpwithoutaccumulationof yellow dye; endocarp crustaceous;seed ca. 1.2-1.5 x 1 cm; hilum large. Distribution. (see Fig. 74.3). From Costa Rica to AmazonianBolivia, apparentlywith discontinuousdistribution;in non-inundatedwet forest;atlow elevations. Specimensexamined.COSTA RICA. PUNTARENAS: Nr.PalmarNortede Osa,22 Feb 1951 (? fl), Allen5948 (EAP,F, GH, US); ParqueNacionalCorcovado,Sirena Field Station, 13 Aug 1984 (? fr), Gentryet al. 48530 (MO);ca. 7 km SE of Quepos,trailPlayaManuelAntonio-PuertoEscondido,22 Aug 1985 (? fr), Grayumet al. 5923 (MO);Cant6nOsa, ca. 5 km W of Rinc6n, 17

Oct 1990 (st), Hammel et al. 17692 (MO); Cant6n Osa, Fila Retina, Quebrada Benjamin, 28 Jun 1991 ($ fl-fr), caducousorsubpersistent. puberulous, Staminatein- Hammel et al. 18241 (BG); Cant6n Osa, nr. Guerra, 6 florescences: peduncle 0.3-0.4 cm long, 1-1.5 mm Aug 1991 (? fr), J. Marin 98 (MO); Parque Nacional thick,brown-puberulous;spike 7-15 cm long, ca. 2-3 Corcovado, Pavo Forest, 16 Jun 1988 (? fl), Kernan et mm diam.;tepalsfree,2-4, (sub)spathulate,ca. 0.5 mm al. 588 (BG, F, MO); Parque Nacional Corcovado, lower long, often i cucullate,with 2 (or 3) embeddedyellow Claro Ridge, 19 Jun 1988 (e'), Kernan et al. 596 (MO); dye-containing"glands,"ciliolate;filaments0.1-0.2 mm ParqueNacional Corcovado, Ollas Trail to Skyway Trail, 21 Jun 1988 (d'), Kernan et al. 608 (BG, F, MO); Parque long; anthers ca. 0.5 x 0.3-0.4 mm, the connective Nacional Corcovado, Sirena, 5 Jul 1977 (? fl-fr), Liesner narrow;bracts lacking (?). Pistillate inflorescences: 2909 (MO). SAN Jost: CerroBijagualito-Tarcoles,4 peduncle 0.5-1 cm long, ca. 2 mm thick, puberulous; Aug 1975 (st), Poveda 1092 (CR, F, MO); Parque head 1.5-2 cm diam.;perianthca. 5 mm long, subentire Nacional de Manuel Antonio, Quepos, 4 Aug 1975 (st), to 4-lobed, the lobes decussate, the inner ones plane, Poveda et al. 1713 (CR).

TAXONOMICTREATMENT

73

Fig. 11. Batocarpus costaricensis. 1. Leafytwig with pistillateinflorescence(Allen 5948). 2. Leafy twig with staminateinflorescences(Mutis 1139). 3. Leaf. 4. linfructescence(Allen 5971). 5. Stamen. 6a-c. Tepalsof staminate flower.7. Bract(?) and bristlesof staminateinflorescence(Mutis 1139). 8. Pistillateflower.9. Fruitingperianth. 10, 10a. Seed. 11. Embryo(Bristan 1459). [By P. Pardoen,Utrecht.]

74 PANAMA. CHIRIQUi: Burica Peninsula, nr. Quebrada Tuco, 21 Feb 1973 (st), Croat 22093A (MO); Burica Peninsula, nr. Punta de Piedra, 1 Mar 1973 (st), Croat 22459B (MO).DARIEN: Rlo Uruceca, Nov 1967 (? fr),

Bristan1459 (MO,PMA,U); Rio Balsa,betweenManene & Tusijuanda,26 Jul 1967 (? fr), Duke 13583 (MO, NY, US); Rio Tuira, between Rio Punusa & Rio Mangle, 1 Oct 1967 (? fr), Duke 14601 (MO, US). COLOMBIA. Without locality, (e^),Mutis 588, 589, 1139, 2958 (US). ANTIOQUIA: Rio Le6n, 26 Jan 1962 (st), Cain 63 (COL); Mun. Turbo, 11 km from Currulao, ENE of Turbo, 3 Aug 1987 (? fr), Callejas et al. 4982 (BG, COL). CALDAS: La Dorada, 23 Apr 1963 (? fr), Espinal T 1205 (COL, PSO), (? fl-fr), Espinal T 1211 (COL). CHOC6: Mun. Riosucio, Zona de Uraba, Cerro del Cuchillo, rd. Cuchillo-La Cumbre Noroeste, 29 Mar 1988 (? fr), Cdrdenas 1572 (JUAM, MO); Mun. Riosucio, between Subida & Tilupo, ParqueNaturalNacional Los Katios, 5 Jul 1976 (? fl-fr), Le6n 252 (MO); Parque NaturalNacional Los Katios, Alto del Lim6n, 9 May 1983 (? fl-fr), Zuluaga R. 757 (COL). ECUADOR. NAPO: Reserva Biol6gica Jatun Sacha, 8 km E of Puerto Misahualli, 30 Dec 1987 (st), Gentry et al. 60207 (BG, MO), 8 Aug 1987 (st), Palacios 1748 & 1798 (BG, MO, QAME), 21 Oct 1989 (st), Palacios 4680 (QAME, QCNE) & 4695 (BG, QCNE); Aiiangu, Parque Nacional Yasuni, 30 May-21 Jun 1982 (st), SEF 8864 (QCA, U), 8962 (QCA, QCNE, U), 9107 & 9133 (QCA, U); nr. outlet of Rio Aiiangu, 30 Jun-9 Jul 1982 (st), SEF 10134 & 358 (QCA, U). PERU. HuANUco: Prov. Puerto Inca, Dtto. Yuyapichis, 2 Nov 1989 (? fr), Kroell S. 772 (BG). LORETO: Prov. Maynas, Rio Amazonas, above mouth of Rio Napo, QuebradaYanomono,9 Nov 1979 (st), Gentryet al. 27772 (MO, U); Rio Amazonas, between mouth of Rio Napo & Indiana, 15 Jul 1983 (st), Gentry et al. 43081 (BG, MO). MADRE DE Dios: Prov. Tambopata,ComunidadNativa de Infierno, Hermosa Chica, 16-18 Nov 1988 (st), Alexiades et al. 109 (BG, MO) 29 Mar 1989 (st), Alexiades et al. 544 (BG, MO); ManuiNational Park, Cocha Cashu Biological Station, 5 Aug 1983 (st), Gentry 43370 (MO); Prov. Tambopata, 15 km ENE of Puerto Maldonado, 12 Dec 1989 (? fr), Gentry et al. 68612 (MO); Prov. Tambopata, Comunidad Nativa de Infiemo, Hermosa Chica, 22 Dec 1990 (? fr), Pesha 71 (BG, US). PASCO: Valley of Rio Palcazu,Iscozacin, 8 Aug 1981 (st), Foster 4573 (F); drainage of Rio Palcazu, new rd., km 51-60, NW of Villa Rica, 4 Mar 1982 (st), Gentry et al. 36006 (BG). UCAYALI: QuebradaShesha, ca. 65 km NE of Pucallpa, 25 Jun 1987 (st), Gentry et al. 58501 (BG, MO). BOLIVIA. PANDO: Prov. Manuripi, Conquista, rd. Puerto Rico-Sena, km 18, 7 Oct 1991 (st), Beck et al. 20176 (BG, LPB).

FLORANEOTROPICA Filho & Emmerich,Bol. Mus. Nac. Rio de Janeiro, Nov. Ser. Bot., No. 37: 6, t. 4 & 5. 1968. Anonocarpus amazonicusDucke, Arch. Jard.Bot. Rio de Janiero3: 39. 1922;Arq. Serv.Florest. 1(1): 1, t. 1 & 2. 1939; Type. Brazil. Para:Obidos, Paranabelow Obidos, 3 Jan 1916 (o), Ducke MG 15922 = RB 13077 (lectotype, Emygdio de Mello Filho & Emmerich, Bol. Mus. Nac. Rio de Janeiro,Nov. Ser. Bot., No. 37: 6. 1968, R; isolectotypes, B, G, K, MG, NY, U, US). Fig. 12 BatocarpusmaranhensisEmygdio de Mello Filho & Emmerich,Bol. Mus. Nac. Rio de Janeiro, Nov. Ser. Bot., No. 37: 9, t. 6. 1968. Type.Brazil. Maranhao: Viana, 1936 (o), Carvalho1 (holotype, R).

Tree, to 30 m tall. Leafy twigs 1.5-6 mm thick, sparselyto densely andminutelypuberulous,some of the hairs uncinate. Lamina oblong to elliptic, 5-26 x 2.5-13 cm, broadestabove the middle, ? inequilateral, subcoriaceous;apex acuminate;base obtuseto rounded to subcordate;marginsubentireto repandto irregularly crenate;uppersurfacesparsely(especially on the midrib)to ratherdensely puberulousto (subtomentellous); lower surfacesparsely,on the main veins mostly more densely puberulous,some of the hairsuncinate;venation slightly prominentabove, the main veins or also the smallerveins prominentbeneath;lateralveins 8-14 pairs;tertiaryvenationpartlyscalariform;petiole 0.31.5 cm long, ca. 1-2.5 mm thick, densely puberulous, some of the hairs uncinate, the epidermis sometimes flaking off, stipules 0.2-0.6 cm long, puberulous(to hirtellous), caducous. Staminate inflorescences: peduncle 0.6-1 cm long, ca. 1 mm thick, densely, white to pale brown-puberulous;spike 8-22 cm long, ca. 24 mmthick;tepalsfree,3-5, 0.5-1.5 mm long, minutely puberulous;stamens 1 or 2; filaments ca. 1 mm long; anthersca. 0.4-1 x 0.25-0.5 mm, the connective glandularat the apex;bractsnarrowlyto broadlyspathulate to subpeltate,0.5-1.5 mm long, with 2 embeddedyellow dye-containing "glands,"puberulous. Pistillate inflorescences: peduncle 0.7-1.5 cm long, ca. 2 mm thick, densely puberulous;head globose to ellipsoid, ca. 1-2 cm diam.;perianthca. 5 mm long, subentireto 4-lobed, densely puberulousto subvelutinousoutside andinside, the upperpartwith embbeded,yellow dyecontaining"glands";ovarypuberulous;style ca. 1 mm long; stigmassubulate,ca. 3 mm long; bractsoblong to subrectangular(to subpeltate),2-4 mm long, with 1 or 2 embeddedyellow dye-containing"glands,"the apex Local names. CostaRica:ojochemacho.Colombia: puberulous. Fruitingperianth1-1.3 cm long; pericarp Choco:gusanero.Ecuador:Napo:frutode pande monte. Peru:Madrede Dios: chimicua.Bolivia: Pando:chicle. subcoriaceous,with accumulationof yellow dye; seed ca. 6-8 x 5-7 mm, flattened;testa adnateto the pericarp;hilum large(?). 2. Batocarpus amazonicus (Ducke) Fosberg, Proc. Distribution (see Fig. 75.3). In the Amazon BaBiol. Soc. Wash. 55: 99. 1942; Emygdio de Mello sin, whenceit extendsto Maranhao(Brazil)andFrench

TAXONOMIC TREATMENT

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.IY.

75

@?

Fig. 12. Batocarpus amazonicus. 1. Leafy twig with infructescence. 2. Leaf (Reynel 702). 3. Leafy twig with

youngstaminateinflorescences.4. Staminateinflorescence(Lao et al 1664). 5. Leafy twig with pistillateinflorescence (Steinbach 7207). 6a, 6b. Staminateflowers. 7. Stamen.8. Bracts of staminateinflorescence (Lao et al 1664). 9. Pistillate flower. 10. Bractsof pistillate inflorescence(Foster 5691). 11. Fruit. 12. Seed. 13. Embryo (DuckeMG 16924). [By P. Pardoen,Utrecht.]

76 Guiana;in non-inundatedor, in Peru, also in seasonally inundated(tahuampa)forest;at low elevations. Specimens examined. FRENCH GUIANA. Arataye R., Sauts Parare, 28 Oct 1985 (st), Barrier 4830 (CAY) 21 Nov 1980 (? fl-fr), Sabatier 28 (U); piste de St. Elie, 5 Oct 1991 (st), Sabatier et al. 3900 (BG). ECUADOR. NAPO: 3 km S of Tena, 12 Dec 1985 (d), Zaruma 379 (BG, QAME, QCA, QCNE). SucuMBIOS:Cant6n Aguarico, Reserva Faunistica Cuyabeno, Rio Aguarico, Zancudo, 27 Sep 1991 (de), Palacios et al. 7715 (BG, QCNE). PERU. Cuzco: Prov. La Convenci6n, Alto Urubamba, Manguyari,3 Feb 1989 (? fr), Niuiiezet al. 10148 (AG). HuANUCO:Tingo Maria, 8 Aug 1940 (d), Asplund 12915 (G, R), (? fl), Asplund 12916 (G, R); Prov. Puerto Inca, Dtto. Yuyapichis, Dantas, 2 Nov 1989 (? fr), Kroell S. 772 (BG); Tingo Maria, 9 Dec 1981 (c?), Plowman et

FLORANEOTROPICA Center,14 Aug 1973 (st), Berg et al. P18524 (MO,NY, U). PARA:Belem, BosqueMunicipal,27 Jan 1942 (de), Ducke947 (F,IAN,MG,MO,NY,R, US);Obidos,Cacaval Imperial,8 Sep 1910(q fr),DuckeMG 11006(MG,US), 21 Jan 1918 (9 fl-fr)DuckeRB 13077 (B, G, GH,K, R, U), 21 Jan 1918 (Y fl), Ducke MG 16924 = RB 13077

(G, GH, K, MG, R, U, US); Belem, Bosque Rodrigues Alves, 5 Aug 1950 (d), A. Silva 464 (COL, F, IAN). BOLIVIA. BENI: Prov. Vaca Diez, Tumi Chucua, 27

km SW of Riberalta,30 Sep 1991 (q fl-fr),Beck20065 (BG, LPB);Prov.Yacuma,EstacionBiol6gicadel Beni, Rio Maniqui,El Chacal,18Aug 1991 (st),Palacios 7566 (LPB);Rio Maniqui,nr.SanBora, 25 Sep 1976(q fl-fr), Meneces344 (MO);Prov.Ballivian,Bajio, 15 Sep 1993 (st), Michel et al. 1159 (BG, LPB). COCHABAMBA: Prov.

Carrasco,Estaci6nValle del SajtaUMSS,(st), Galarza s.n. (USZ).PANDO: Prov.Nicolas Suarez,nr.Cobija,23 Oct 1978 (st), Meneces 793 (BOLV). LA PAZ: Rio Bopi

al. 11270 (F, U). LORETO: Prov.Maynas,Rio Tahuayo, valley,Rio Inque,25 Sep 1921(9 fr),Rusby751 (F,NY). 12 Dec 1961 (q fr), Aristegui V 15 (U); Varadera de SANTACRUZ:Prov. Nuflo de Chavez, Ascenci6n de Mazan, 27 Sep 1972 (de), Croat 20772 (NY); Prov. Guarayos, 30 Aug 1985(st),Beck12254(BG,LPB);Santa Maynas, Rio Amazonas, between mouth of Rio Napo & Cruz,JardinBotanico,9 Oct 1977(? fl), Hartshornet al. Indiana,28 Aug 1983 (st), Gentryet al. 43770 (BG,MO); 2004 (BG); nr. SantaCruz,Nov 1907 (st), Herzog 682 Prov. Maynas, Dtto. Iquitos, rd. to Quisto Cocha, 28 May (Z); Prov.Nuflo de Chavez, San Ram6n,22 Feb 1991 1973 (? fl), McDaniel17355 (NA);Prov.Uyacali,Dtto. (st), Quevedoet al. 361 (LPB, MO, USZ); Prov. Sara, Contamana, rd. Contamana-Aguas Calientes, km 10, 27 Japacani,Aug 1925 (? fl), Steinbach7207 (A, BM, G, Oct 1982 (? fr), Reynel R. 782 (BG, U); Prov. Loreto, GH,MO,NY,U); Prov.Ichilo,ParqueNacionalAmbor6, Tipischca, Santa Cruz, Rio Pacaya, 17 Jul 1985 (st), R. 8 kmSWof El Carmen,15-22 Nov 1991(st),I. G. Vargas Vasquezet al. 6680 (BG); Prov. Maynas, Maniti, Recreo, et al. 1208 (LPB, USZ); Prov. Guarayos,4 km SE of 15 May 1988 (? fl-fr), R. Vasquez10667 (BG, GH, MO); confluenceof Rio Negrode Caimanes& Rio San Pablo,

Prov. Maynas, Indiana, Yanamono, 3 Dec 1988 (? fl-fr), R. Vasquezet al. 11309 (BG); lower Rio Huallaga, Puerto Arturo,21 Nov 1929 (? fr), Lt. Williams5334 (A, F, US). MADRE DE DIos: ParqueNacional del Manu, Rio Manu, Cocha Cashu Station, 1 Nov 1980 (? fl), Foster 5691 (F); Tambopata, Rio Tambopata, 19 Feb 1984 (st), Gentry et al. 45617 (BG, MO); Tambopata Nature Reserve, jct. of Rio La Torre & Rio Tambopata, 24 May 1987 (st), Gentryet al. 57563 (BG, MO).SAN MARTIN:Prov. Mariscal Caceres, Dtto. Tocache Nuevo, rd. to Shunte, 26 Feb 1970 (? fl-fr), Schunke V 3826 (COL, F, GH, K, MO, NY, US, WIS). UCAYALI: Rd. Pucallpa-Lima, ca. km 99, 6 May 1975 (o), Lao M. et al. in Hartshorn

1664 (BG, F, MO, U). BRAZIL. ACRE:upper Rio Moa, Fazenda Arizona, 24-30 Sep 1984 (st), Campbell et al. 8054 (BG); Mun. Cruzeiro do Sul, Rio Tejo, 13 Mar 1992 (? fl-fr), Daly et al. 7383 (BG); Mun. Sena Madureira, Rio Macaua, Coloca9do Cachorra Macha, 2 Apr 1994 (? fr), Daly et al. 8140 (NY); mun Senador Guiomard, rd. Rio BrancoBrasileia, km 68, 10 Oct 1991 (9 fr), Ferreira et al. 10261 (INPA). AMAPA:Reserva INCRA Rio Falsino, 22-26 Oct

1986 (st), Campbellet al. 14316 (BG). AMAZONAS: Ca. 15 km SE of Sao Gabriel da Cachoeira, 28 Nov 1987 (9 fr), Kawasaki 341 (BG, INPA); Mun. Humaita, nr. Tres Casas, 14 Sep-1 I Oct 1934 (9 fr), Krukoff 6422 (A, B, BM, F, G, K, LE, MO, NY, U); Rio Piria, Sep 1959 (q flfr), N. T Silva 556 (IAN, NY, US). MARANHAO: Mun. Mon9ao, Ka'apor Indian Reserve, 9 May 1986 (st), Bal,e 2277 (NY). MATOGROSSO:Rio Aripuana, Humboldt

18 Jun 1993 (e), I. G. Vargas et al. 2604 (USZ).

Local names.Brazil:Acre:mururemole;Maranhao: murure. Peru: Huanuco: leche caspi, mashonaste; Loreto: mashonaste; San Martin: sacha ishpingo; Ucajali:mashonaste,mashonastenegro.Bolivia:Beni: murure,quechoamarillo;Cochabamba:murure;Pando: quecho amarillo;SantaCruz:murure,mururehuasa.

3. Batocarpus orinocensis Karsten, Fl. Colomb. 2: 67. t. 134. 1862; Emygdio de Mello Filho & Emmerich, Bol. Mus. Nac. Rio de Janeiro,Nov. Ser. Bot., No. 37: 4, t. 1 & 2. 1968. Type. Colombia. Meta:Llanode San Martin,Apiai, (v), Karsten s.n. (holotype, LE). Fig. 13 Anonocarpus peruvianus Ducke, Arq. Serv. Florest.

Rio de Janeiro1(1): 1. 1939. Type.Peru.Loreto: Rio Huallaga,nr.Yurimaguas, 13 Feb 1924 (? fr), Kuhlmann1399 (holotype,RB; isotypes,B, K). Tree, to 35(-40?) m tall. Leafy twigs 1-5 mm thick,

sparsely to densely, puberulous. Lamina oblong (to elliptic),7-21.5 x 2-8.5 cm, mostly broadestabovethe middle, slightly inequilateral,(sub)coriaceous; apex acuminate(to subcaudate);base obtuse to rounded; margin entire;upper surface glabrous;lower surface on the midrib;venation sparselyappressed-puberulous

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I

77

4l$1;

Fig. 13. Batocarpus orinocensis. 1. Leafy twig with staminate inflorescence (Kayap 159). 2. Leafy twig with pistillate inflorescence (Gentry et al. 29733). 3, 3a. Staminate flower. 4. Stamen. Sa-e. Bracts of staminate inflorescence (Asplund 12915). 6, 6a. Pistillate flower (Schunke V 7453). 7. Fruiting perianth. S. Seed. 9. Embryo. 10a-c. Bracts of pistillate inflorescence (Tunqui 668). [By P. Pardoen, Utrecht.]

78

FLORA NEOTROPICA

(almost)planeabove,? prominent lateralveins beneath; 8-16 pairs;tertiary venationreticulate; petiole0.7-1.2 cmlong,1-2.5mmthick,puberulous; stipules0.4-0.8 cm long, puberulousto subsericeous,caducousor subpersistent.Staminateinflorescences:peduncle0.3-

Reserva Floristica El Chuncho, Estaci6n Exp. INIAPPayamino, 5 km NW of Coca, 12-14 Oct 1987 (e), Neill et al. 7901 (BG, MO, QAME); Reserva Biologica Jatun Sacha, 8 km E of Puerto Misahualli, 12 Aug 1987 (R fr), Palacios 1837 (BG, MO, QCA, QCNE); Canton Francisco de Orellana, rd. to Los Zorros, Pozo Petrolero Jaguar I, 23 Oct 1988 (a'), Palacios 3227 (BG, MO); Afiangu, Parque Nacional Yasuni, 30 May-21 Jun 1982

0.4cmlong,1-1.5mmthick,brown-puberulous, sometimeswithI or2 triangular bracts,to 1mmlong;spike 7-15(-30) cmlong,ca.2-2.5 mmthick;perianth 0.6- (st), SEF 8542 (QCA, U). PASTAZA: Rd. Coca-Auca, 0.8mmlong,2-4(-5)-parted,glabrous; filaments0.6- 115 km S of Coca, 4 km S of Rio Tigiiino, 22-28 Feb 1 mmlong;anthers0.5 x 0.3 mm;interfloralbracts 1989 (? fr), Zak 3974 (BG, QCNE); Cant6n Pastaza, topeltate, 0.8-1mmlong, Pozo Petrolero Ramirez, ca. 20 km S of Curaray,21-28 tonarrowly broadly spathulate theupperpartthickened,withembeddedyellowdye- Feb 1990 ($ fl-fr), Zak et al. 5170 (BG, QCNE). containing"glands,"puberulous. Pistillate inflores- ZAMORA-CHINCHIPE:Rio Nangaritza,Pachicutza,3 Dec cences:peduncleca. 1 cm long;headsubgloboseto 1990 (e), Palacios et al. 6480 (BG, MO). PERU. AMAZONAS: Nr. Yuwi Entsa, 13 Mar 1973 (R ellipsoid,ca. 1-1.5 cmdiam.;perianthca. 3 mmlong, fr), Ancuash 100 (MO, U); Rio Santiago valley, 65 km theupperpartwithoutyellowdye-containsubentire, N of Pinglo, nr. Caterpiza, 24 Mar 1980 (2 fr), Huashikat ing "glands,"white-subvelutinous,the lower part 2355 (MO); Quebrada Aintami, 12 Jan 1973 (? fl-fr), sparselypuberulous; ovaryglabrous; styleca.2-3 mm Kayap 113 (MO, U); Rio Cenepa, Huampami, 18 Jan long;stigmaslingulate,1.5-2mmlong,conspicuously 1973 (? fr), Kayap 159 (MO, NY, U); Soledad, Jun 1925 to subpeltate, papillate; interfloral bractsspathulate 0.5- (e'), Tessmann 5243 (NY); Rio Santiago, Quebrada 1mmlong,with1or2, embedded yellowdye-contain- Caterpiza, 29 Jan 1980 (? fr), Tunqui 668 (BG, MO). ing"glands," puberulous attheapex.Infructescences HUANUCO:TingoMaria,8 Aug 1940(a), Asplund12915 3.5-4.5cmdiam.;fruitingperianth ca.2 cmlong;peri- (R), (? fl), Asplund 12916 (K, R, US); Prov. Puerto Inca, carpsubcoriaceous, withoutaccumulation of yellow Dtto. Yuyapichis, 2 Nov 1989 (st), Kroell S. 728 (BG); Tingo Maria, 24 Oct 1962 (c), Schunke V 6191 (F, US), dye;seedca. 1.5 x 1-1.2 cm;hilumsmall.

29 Aug 1940 (9 fr), Skutch4960 (F, K). JUNMN:Prov.

Distribution (see Fig. 75.2). In the upperAmazon

Basin,fromColombia(Putumayo) to Brazil(Acre),in alsoin Meta;innon-inundated Colombia forest;to 1200 m (ormore?). Representativespecimensexamined.COLOMBIA. AMAZONAS: Araracuara,Rio Yari, 18 Apr 1986 (? fr), Galeanoet al. 709 (COL). META:Villavivencio,Llano de San Martin, (e' fl-fr), Triana 835 (K, NY, P). PUTUMAYO: BetweenMocoa& PuertoLim6n,Jun 1935

(2 fl), Garcia-Barriga3675 (AAU, COL). ECUADOR. MORONA-SANTIAGO: Nr. CentroTuutin Entsa,nr.Taisha,10 Sep 1985 (st), AnananchRBAE.178 (BG); Rio Yuquipa,10 km SE of Macas, 18 Mar 1986 (st), Baker 6774 (BG); CentroShuarYukutais,5 Nov 1988 (a), Bennett et al. 3502 (BG, QCNE); 6 km NE of Gualaquiza, 16 Sep 1975 (a'), Little et al. 389 (US); Cord.

de Cutucu,6-10 km E of Logrofio, 1200-1500 m, 78 Oct 1975 (ae), Littleet al. 660 (LOJA,US); Bomboiza, nr. Misi6n SalesianaShuar,5 Nov 1986 ($ fr), Neill et al. 7442 (AAU,BG); 16.7 km NE of Taisha,14-19 Aug 1976(st), OrtegaU. 30 (US). NAPo:10 kmNW of Coca, 25 Feb 1980 (st), Berg et al. 1079 (AAU, MO, NY, QCA, US), (? fl-fr), Berg et al. 1080 (BG, QCA, U); rd. CocaAuca, ca. km 30-35, 26 Feb 1980 (st), Berg et al. s.n.

(BG);ReservaBiol6gicaJatunSacha,8 km E of Puerto Misahualli, 23-27 Jun 1987 ($ fr), Cer6n 1667 (BG), 8 Nov 1987 (a), Ceron 2651 (BG, MO, QAME, QCA, QCNE, US); Parque Nacional Yasuni, Pozo Petrolero Daimi 2, 26 May-8 Jun 1988 (a'), Ceron et al. 3849 (AAU, BG, MO, QAME, QCNE); Huashito, 20 km N of

Satipo, Dtto. Puerto Ocapa, 11 Jun 1981 (st), Reynel R. 318 (U). LORETO: Varadera de Mazan-, 27 Aug 1972 (e), Croat 20772 (MO,NY); Prov. Alto Amazonas, Rio Pastaza, Andoas, 15 Aug 1980 (a), Gentry et al. 29733 (BG, F, MO); Prov. Maynas, Dtto. Las Amazonas, Rio Napo, nr. Sucusari, 3 Mar 1991 (? fl-fr), Pipoly et al. 14223 (BG); Dtto. Pebas, Brillo Nuevo, Rio Yaguasyacu, 4 Nov 1981 (?), Treacy et al. 337 (BG, F); Prov. Loreto, Rio Maranion,Nauta, 9 Nov 1982 (? fl-fr), R. Vdsquez et al. 3426 (NY); Prov. Maynas, Rio Itaya, San Antonio, 13 Dec 1982 (? fl-fr), R. Vdsquezet al. 3571 (MO); Prov. Maynas, rd. Iquitos-Nauta, km 44, 14 Mar 1989 (? fr), R. Vdsquezet al. 11927 (BG). PASCO: Prov. Oxapampa, La Esperanza, Rio Pichis, 1 hr. below Puerto Bermuidez, 17 Jun 1983 (st), Gentry et al. 42134 (MO); Prov. Oxapampa, Dtto. Iscozacin, 15 Oct 1986 (? fr), Pariona et al. 1021 (BG, F, MO). SAN MARTiN: Prov. Mariscal Caceres, Tocache Nuevo, rd. to Shunte, Rio Tocache, 16 Jul 1974 (? fl), Schunke V 7453 (MO, NY, U); Prov. Mariscal Caceres, Tocache Nuevo, Quebrada de Cachiyuca, 22 May 1975 (a), Schunke V 8531 (F, MO, NY, U). BRAZIL. ACRE: Upper Rio Moa, nr. Fazenda Arizona, 10-16 Oct 1985 (st), Campbell et al. 6174 (BG); Reserva INCRA Santa Luzia, BR.364, km 40, 5-19 Oct 1984 (st), Campbell et al. 6828 (BG); Rio Jurua',1 km upstream from Col6nia Rodriguez Alvez, 20-25 Oct 1986 (st), Campbell et al. 12887 (BG).

Local names and uses. Ecuador: Morona Santiago:pikiu, pitiu or pitiuk (Shuar),pituca;Napo: Coca, 3-21 Nov 1989 (a), Gudifio103 (BG, QCNE); sachapaparagua(Quichua);Zamora-Chinchipe:pitiu

TAXONOMIC TREATMENT

79

ThethirdspecieswasdeAllemao(1857)respectively. in thegenusExcoecaria scribedintheEuphorbiaceae, by Sprengel(1821),as E. ilicifolia.Thesamespecies was also describedin the generaAcanthinophyllum Allemao(1858)andAliteriaBenoist(1929),bothcreatedto accommodate thespecies.Ina seriesof studies and onMoraceae, Lanjouw (1936)includedSahagunia thetwogeneracreatedbyAllemaoin Clarisiaandrecognizedeightspecies.Inarevisionof thegenus,Burger Clarisia in whichhe rec(1962)reinstated Acanthinophyllum, 6. ClarisiaRuiz& Pav6n,Fl. Peruv.Chil.128.1794, ognizedtwospecies,leavingin Clarisiaonlytwospenom.conserv.,nonClarisiaAbat,1792,nom.rejic.; cies,namelythosedescribedbyRuiz&Pavon. Lanjouw,RecueilTrav.Bot.Neerl.33:254. 1936; Clarisia biflora and C. racemosa can become tall Burger,Ann. MissouriBot. Gard.49: 20. 1962. trees,whereasC.ilicifoliais a shruborsmallunderstory Type(conserved). ClarisiaracemosaRuiz& Pavon. tree.Thetwoformerspecieshaveconspicuously warty, Sahagunia Liebmann, Kongel. Danske Vidensk. lenticellate bark.Thebarkonthebaseof thetrunkand Selsk. Naturvidensk.Math.Afh., Ser. 5, 2: 316. thatontherootsareorangetored,duetoaccummulation 1851. Type. SahaguniamexicanaLiebmann(= of dye.Uncinate hairsarealwayspresentin C.ilicifolia, Clarisia biflora Ruiz & Pav6n). sometimespresentin C. biflora,andhaveneverbeen (Shuar).Peru:Amazonas:pitu,uyapitu(Huambisa); Huanuco:mashonastenegro; Junin:mashonaste; Loreto: pandilla, pittiu (Jivaro); San Martin: mashonastillo; Pasco:chimicua. JivaroIndiansusethe latexmilksapforstomach(fruits, acheandintestinal diseases.Theinfructescences seeds)areedible.

SoaresiaAllemao,RevistaBrazil.1: 210. 1857,nom. rejic., non C. H. Schultz-Bip., 1863, nom. conserv. (Asteraceae). Type. Soaresia nitida Allemao(= ClarisiaracemosaRuiz & Pav6n). Acanthinophyllum Allemao, RevistaBrazil. 1: 368. 1858; Burger,Ann. MissouriBot. Gard.49: 27. 1962. Type. Acanthinophyllum strepitans Allemao(= Clarisiailicifolia(Sprengel)Lanjouw & Rossberg). AliteriaBenoist,Bull. Mus. Hist.Nat. (Paris),Ser.2, 1: 163. 1929. Type. Aliteria sagotii Benoist (= Clarisiailicifolia(Sprengel)Lanjouw& Rossberg).

foundin C. racemosa.The leaf marginof C. ilicifoliais

evenlobate,but oftenspinulose-dentate, andsometimes it is entireinthetwootherspecies. Theinflorescencesin Clarisiabifloraareusually bomein theaxilsof theleaves,butin theotherspecies normallyonleaflessshootseitherwithlongintemodes in (C. racemosa)orwith short internodesin (C. occurin ilicifolia).Ontheseshoots,theinflorescences pairsandaresubtendedby scale-likebracts(= fused stipules).As the inflorescencesof C. bifloraandC. theleaflessshootwith racemosaareusuallyuniflorous, solitary, fornsa racemose the inflorescences, paired or Treesor shrubs, dioecious;uncinatehairspresent as a raceme, in often indicated structure, descriptions and lamina orabsent.Leaves alternate distichous; pinorlo- althoughit is a compoundinflorescence. natelyveined;marginentire,(spinulose-)dentate arespicate,short,ofThestaminate inflorescences bate;stipulesfree,lateral.Inflorescences pairedorsoliin Clarisia or sometimes even subcapitate ten subclavate taryintheaxilsof theleavesoronleafless(short-)shoots, in are The flowers arranged longitudinal rows, ilicifolia. to subpeltate, withor bractsbasallyattached bracteate; the with of bracts. At floral altemated rows anthesis, withoutembeddedyellow dye-containing "glands." is ? obscure,especiallyin C. bifloraandC. Staminateinflorescencesspicate,with a distinctabaxial structure in whichthe tepalsareusuallyfree.The racemosa sterilestrip;flowerscrowded,in longitudinalrows; of is mostlyreducedto one,butflownumber stamens tepals(2-)3-6, freeorconnate;stamens1-3; anthers arenotuncommon, andevenflowers with two stamens basifixed,latrorse; pistillodeabsent.Pistillateinfloreswith be sometimes found.The ers three stamens can oruniflorous; subentire cencescapitate perianth tubular, have been described all Clarisia of species pollen grains oralmostfree; or4-lobed;ovaryadnateto theperianth and (1976). by Niezgoda Nowaczyk or stigmas2, equallylong, filiformto band-shaped in arediscoid-capitate Thepistillateinflorescences red,orange, tongue-shaped. Fruitingperianthenlarged, Clarisiailicifolia. Those of C. bifloraandC. racemosa paleyellow,or greenish;fruitadnateto theperianth; areusuallyuniflorous,buta secondfloweris someorcoriaceous; seedlarge, endocarp woody,crustacous, Thesinglefloweris subtended bya small timespresent. withoutendosperm; testaamplyvascularized; embryo involucreconsistingof 3-7 bracts.Theovaryis adnate longitudinally aligned,withequalandthickcotyledons to the perianthin C. biflora and C. racemosa. It is alanda veryshortapicalradicle. mostfreein C.ilicifoliaatanthesis,butthefruitis advariesconsiderably; Theendocarp ThegenusClarisiawasestablished byRuiz&Pavon nateto theperianth. in C. (1794)who describedtwo species,C. biflora andC. it is coriaceousin C. biflora,thinlycrustaceous racemosa. Bothtaxaweredescribed again,eachin an- ilicifolia, andwoody in C. racemosa. The vascularizaothergenus,SahaguniaLiebmann (1851)andSoaresia tion is not confinedto a thickerpartof the testa.In

FLORA NEOTROPICA

80

Clarisiabiflora,theperianth,the connective,thebracts, andthe cotyledonshave very smallembedded"glands" containinga yellow to orangedye. Clarisia biflora and C. racemosa can be regarded as ratherclosely related species. The differences between these two species and C. ilicifolia occur, e.g., in

thesize of theplant,in thespinuloseleafmarginand pistillateinflorescences. apex,andin themultiflorous the occurrenceof similardifferencesin Considering at the othergenera,noneof themjustifiesseparation atthesecgenericlevel(cf. Burger,1962).Separation tionlevelis proposedherewith.

Key to the speciesof Clarisia 1. Margin and/or apex of the lamina spinulose, the margin often dentate; midrib prominent above....... 3. C. ilicifolia 1. Margin and apex of the lamina not spinulose, the margin entire; midrib slightly impressed above. 2. Inflorescences on leafless branchlets on the older wood; uncinate hairs absent ........I............2. C. racemosa 2. Inflorescences in the leaf axils or on leafless branchletsjust below the leaves; uncinate hairs usually 1. C. biflora present ...................................................................

6A. Clarisia Ruiz & Pavon sect. Clarisia. Including Sahagunia Liebmann and Soaresia Allemao (see underthe genus). Trees. Lamina entire. Tepals of the staminate flower (usually) free. Pistillate inflorescences (usually) uniflorous. Bracts with embedded yellow dyecontaining "glands." 1. Clarisia biflora Ruiz & Pavon,Syst. Fl. Peruv.Chil. 255. 1798; Burger,Ann. Missouri Bot. Gard.49: 21, t. 5. 1962.Type.Peru.Huanuco:Pozuzu-Macora, Ruiz& Pavons.n. (holotype, MA-n.v., photograph in F seen; isotypes?, F, G). Fig. 14 Sahagunia mexicana Liebmann,Kongel. Danske Vidensk. Selsk. Naturvidensk.Math.Afh., Ser. 5, 2: 316. 1851. Clarisia mexicana(Liebmann) Lanjouw,Recueil Trav.Bot. Neerl. 33: 270, t. 13B. 1936; Woodson& Schery,Ann. Missouri Bot. Gard.47: 124, t. 40. 1960. Clarisiabiflora Ruiz & Pav6n subsp. mexicana(Liebmann)W. Burger,Ann. MissouriBot. Gard.49: 22. 1962. Barranca de SantaMaria, Type.Mexico.Veracruz: Tlaltetla, E of Mirador,(d%),Liebmann14259 (holotype,C; isotypes, F, GH). SahaguniacolombianaRusby,Descriptionsof 300 new species of SouthAmericanPlants11. 1920. Clarisia colombiana(Rusby) Lanjouw,Recueil Trav.Bot. Neerl. 33: 266. 1936. Type. Colombia. Magdalena:SantaMarta,Don Amo rd., 13 Jan 1899 (f), H. H. Smith424 (holotype, NY; isotypes, MICH,MO, US). Clarisiamattogrossensis Lanjouw,RecueilTrav.Bot. Neerl. 33: 268. 1936.Type.Brazil.MatoGrosso: Rio Serragem,nr. Diamantino, Jul 1927 (o), Collenette131 (holotype,K; isotype, NY). ClarisiapanamensisWoodson,Ann. MissouriBot. Gard.47: 123. 1960. Type.Panama.Cocle:N of El Valle de Ant6n, 27 Sep 1946 (? fl-fr), Allen 3741 (holotype,MO;isotypes,BM, F, G, GH, P, U, US).

Tree,to 35 m tall;barkconspicuouslylenticellate, thatof thebaseof thetrunkandthe(larger)rootsreddish.Leafjy twigs2-4.5 mmthick,sparselyto densely puberulous, sometimeswith(retrorse) uncinatehairs. 4.5-25 x 1.5Laminaellipticto oblong(tolanceolate), 11 cm, broadestat or above the middle, often ? inequilateral, subcoriaceous to coriaceous; apexacuminate;baseacuteto rounded;marginentire;uppersurwith faceglabrous;lowersurfacesparselypuberulous straight and/oruncinate hairs;midribslightlyimpressed, the otherveins slightlyprominentto planeabove,? prominentbeneath;lateralveins 8-15 pairs;tertiary venationreticulate orpartlyscalarifonn; petiole0.5-2 cm long, 1-2.5 mm thick,sparselypuberulous,the epidermis oftenflakingoff;stipuleslanceolate, 0.3-0.8 cm long,puberulous,caducous.Staminateinflorescencesin the leaf axils or on leaflessbranchlets just belowtheleaves;peduncle0.1-0.8cmlong,puberulous to almostglabrous;spike2-12 cmlong;perianth ca. 1 mmlong;tepals3-5, variouslyshaped,oftenspathulate,sparselyandminutelypuberulous;stamens12(-3); filamentsca. 1-1.5 mmlong;anthersca. 0.30.5 x 0.3-0.4mm,theupperpartof theconnective with a yellowglandular apex;bractsbasallyattached, reniform to broadlyspathulate,ca. 0.5-0.8 mm long, sparselyandminutelypuberulous, withsmall,embedded yellow dye-containing"glands."Pistillate inflores-

cencesin the leaf axils or on leaflessbranchlets just belowtheleaves,uniflorous; peduncle0.2-0.8cmlong, withstraight puberulous and/oruncinate hairs;perianth 3-4 mmlong, 2-4-lobed to subentire,glabrousexandbasalparts,withsmall,emceptfortheuppermost beddedyellowdye-containing "glands"; ovaryadnate to theperianth; stigmasfiliformto band-shaped and512mmlongortongue-shaped, 2-4 mmlong,smooth orsometimes bracts3-7, reniconspicuously papillate; formto elliptic,puberulous, withsmall,embedded yellow dye-containing"glands."Fruitingperianthglobose

to ellipsoid(oroblongoid),1.8-2 x 1.8-2 cmor3.5-4

TAXONOMICTREATMENT

81

JIII

Fig. 14. Clarisia biflora. 1. Leafy twig with pistillate inflorescences (Skutch et al. 2003). 2. Leaf (Schunke V 6230). 3. Leaf (Dodson 5235). 4. Leaf (Gentry et al. 32312). 5. Leaf. 6. Branchlet with staminate inflorescences (Knapp et al. 5473). 7, 7a. Stamens. 8, ga. Bracts of staminate inflorescence. 9a-c. Tepals of staminate flower (Cuatrecasas 21318). 10, 10a. Stamens with glandular apex (Klug 3698). 11, lla. Pistillate flowers (Skutch 2003). 12. Fruiting perianth and fruit. 13. Seed. 14. Embryo (Gentry et al. 32312). [By P. Pardoen, Utrecht.]

82 x ca. 1.5 cm, yellowish(-green)or red at maturity,glabrous;endocarpcoriaceous. Distribution (see Fig. 75.1). FromsouthernMexico through CentralAmerica and the Andean region to northwestemVenezuela,the Pacific Coastallowlands of Colombia and Ecuador,andthe upperAmazon Basin; in wet non-inundatedforest;to 1750 m, sometimes (such as in Colombia and Peru) to ca. 2100 m. Representative specimens examined. MEXICO. Esquintla, 30 Dec 1939 (d), Matuda 4023 (A, MO, NY); Esperanza, nr. Escuintla, 9 Aug 1941 (? fr), Matuda (1)5470 (CAS, F, LL), 2 Jan 1948 (? fl), Matuda (1)7439 (F, K); 30 km NE of Ocozocuantla, 30 Mar 1950 (9 fl-fr), Miranda 6339 (CAS, US); 4-5 km S of Palenque, 27 Jun 1970 (? fr), Thorne et al. 40536 (CAS, RSA) & 40540 (CAS). OAXACA: Mun. Santa Maria Chimalapa, 7 km SW of Santa Maria, 24 Jul 1984 ($ fr), Hernandez G. 241 (F, MO). VERACRUZ: Mun. San Andres Tuxtla, 30 km N of Catemaco, 12 Apr 1984 ($ fl), Cedillo T et al. 2710 (MO); between Playa Escondida & UNAM Field Station Los Tuxtlas, 25 May 1981 (? fr), Gentry et al. 32312 (U, MO); Estacion Biol6gica Tropical Los Tuxtlas, 18 Apr 1983 (? fl), Ibarra M. 529 (MO, NY); Cazadero de Santa Maria, Consoquitla, Mar 1842 (d), Liebmann 14273 (F); Zacaupan, Barranca de Tenampa, Jun 1905 (? fr), Purpus 5989 (F, GH, NY, US), Mar 1912 (e'), Purpus 6492 (F, GH, MO, NY, US). GUATEMALA. QUEZALTENANGO: Colomba, 22 Dec 1934 (? fl), Skutch 2003 (A, BM, F, G, NY, U, US), (6'), Skutch 2004 (A, BM, F, G, NY, U, US) ATLANTIDA: Nr. Tela, 6 Dec 1927HONDURAS. 20 Mar 1928 (st), Standley 54677 (A, US) & 55377 (A). COMAYAGUA: Pito Solo, Lake Yojoa, 22 Aug 1932 (st), Edwards 415 (A, F) & 460 (A, EAP, F, K). NICARAGUA. CHONTALES: Cerro Oluma, 4 Jan 1984 (st), Gentry et al. 43926 (MO). MATAGALPA: Rd. Tuma-Waslala, ca. 1 km E of El Trebol, 25 Nov 1981 (? fl), Stevens et al 20940 (MO). ZELAYA: Rd. Nueva Guinea-La Fonseca, 9 Sep 1982 (st), Araquistain 3179 (MO); 30 km SW of Waspan, 24 Mar 1971 (st), Little 25287 (F, MO, US); 5 km E of San Antonio, 7 Sep 1983 (2 fr), Nee et al. 27848 (MO). COSTA RICA. AJALUALA: Rio Segundo, 4 Sep 1943 (? fl-fr), Orozco C. 1674 (CR). CARTAGO: Nr. Turrialba, 29 Jan 1953 (? fr), J. Leon 4203 (NA), Feb 1902 (?), Nov 1893 (? fr), Tonduz8338 (BR, US). GUANACASTE: Canton Santa Cruz, rd. Santa Cecilia-La Esperanza, 29 Dec 1989 (? fr), Gereau et al. 3461 (MO); Cant6n Hojancha, Rio Ora, rd. to El Carmen,29 Dec 1988 (? fl), Flores et al. 15 (BG), 1 May 1989 (? fr), Flores 41 (BG, MO); Tilaran, Rio Chiquito, 6 Jun 1987 (2 fr), Haber et al. 7429 (F, MO); Tilaran, Rio Chiquito, La Chiripa, and Cerro Frio, 9 Sep 1987 (? fl-fr), Haber et al. 7481 (BG, MO). HEREDIA: Finca La Selva, Rio Puerto Viejo, 27 Aug 1980 (d), Hammel 9598 (F, MO); Puerto Viejo, Rio Sarapiqui, 21 Dec 1974 (9 fr), Hartshorn 1477 (F, MO, U). PUNTARENAS: Golfo Dulce area, nr. Jalaca Farm, 25 Mar 1949 (2 fr), Allen 5207 (EAP, MICH, MO, US); N CHIArAs:

FLORANEOTROPICA of Palmar,N of Osa, 22 Feb 1951 (9 fl), Allen 5955 (EAP, F, GH, US); between PalmarSur & Puerto Cortes, 21 Aug 1965 (?), Jimenez M. 3458 (F); Parque Nacional Corcovado, Sirena, 4 Apr 1988 (9 fr), Kernan 374 (BG, MO), 30 Jan 1989 (9 fl), Kernanet al. 918A (BG, MO). SAN Jost: Villa Col6n, 28 Feb (9 fr), McCaffrey 158 (CR), 16 Feb 19.60 (? fl), Fournier822 (CR), 28 Apr 1966 (2 fr), Fournieret al. 979 (CR). PANAMA. CHIRIQUi: Without locality, Jul-Aug 1927 (st), Cooperet al. 232 (NY); Burica Peninsula, nr. Punta de Piedra, 1 Mar 1973 (st), Croat 22469A (MO). COL6N: Rio Guanche, rd. to Portobelo, 24 Mar 1975 (d'), Moriet al. 5182 (MO,NY, PMA)*. PANAMA: Cerro Jefe, Altas de Pacora, 18 Dec 1980 (2 fr), Antonio 3206 (BG, MO)*; Serrania de Maje, S of Ipeti, 2 Apr 1982 (st), Knapp et al. 4561 (MO); rd. El Llano-Carti, km 8, 9 Jun 1982(e), Knappet al. 5473 (MO,U)*. SAN BLAS: Rio Acla, Aila Tiwar, 11 Mar 1979 (st), Sugden et al. 548 (MO). COLOMBIA. ANTIOQUIA: Mun. Mutata',Vrda. Caucheros, 31 Mar 1983 (a'), Bernal et al. 526 (NY); Mun. Mutata, Vrda. Caucheras, 6 Jun 1987 (ei), CastanoRosalba 12 (HUA)*, 6 Jun 1987 (? fl), CastanoRosalba et al. 44 (HUA)*; Mun. Mutata, Finca La Palmera, 3 May 1987 (oi), Fonnegra et al. 2120 (BG); nr. Villa Arteaga, 6 Dec 1948 (d), Lopezet al. 34 (US); Mun. Venecia, Bolombolo, 9 Mar 1985 (9 fr), Renteria et al. 3633 (JUAM);Mun. Anon', Buenos Aires, ca. 4 km from Providencia, 12 Dec 1972 (d), Soejarto et al. 3633 (AAU, F, GH, MEDEL, MO, NY, VALLE). CHOC6: Mun. Bahia Solano, between Bahia Solano & Rio Juna, 27 Aug 1989 (? fl-fr), Espina et al. 3082 (HUA)*. CUNDINAMARCA:BetweenSanFrancisco& Subachoque,

1900-2100 m, 26 Jan 1944 (e), Garcia-Barriga11030 (COL, US). GUAJIRA: 15 km S of Mingueo, trail to Pueblo Viejo, 21 Aug 1986 (st), Gentry et al. 55421 (MO). HUILA: Rio Villalobos, nr. Rio Suazita, 1400 m, Jan 1943 (ei), Schultes et al. 5189 (GH). MAGDALENA: Sierra Nevada de Santa Marta, (? fl), Karsten s.n. (LE); nr. San Javier, 1300 m, 22 Jan 1959 (9 fl-fr), Romero-

Castaneda 7030 (COL, F, MO, NY). QUINDIO: Mun. Filandia, Vrda. Bambuco, la Curva, Quebrada La Armenia, 16 Jan 1990 (2 fl-fr), Velez et al. 988 (COL); Mun. Filandia,rd. El Roble-Circasia, 1780 m, 24 Jan 1990 ($ fl-fr), Velezet al. 1022 (COL). PUTUMAYO: Mocoa, 3-7 Dec 1942 (? fr), Schulteset al. 2087 (F). RISARALDA: Mun. Pereira, Finca El Pital, 1500 m, 25 Jul 1980 (9 fl), Idrobo et al. 10022 (COL). VALLE:Rio Anchicaya, El Retiro, 13 Oct 1943 (? fr), Cuatrecasas 15278 (F, US)*; Rio Sanquinini, La Laguna, 1250-1400 m, 10-20 Dec 1943 (? fl-fr), Cuatrecasas 15487 (COL, F), (oi), Cuatrecasas 15493 (F); Rio Calima, nr. La Brea, 22-25 May 1946 (d%),Cuatrecasas21318 (F); Cartago, Hda. La Silvia, 21-26 Dec 1946 (9 fl), Duque-Jaramillo 4115A (COL); Santa Helena, above Topacio, 1930 m, 10 Dec 1985 (st), Gentry et al. 53038 (MO). AMAZONAS: Upper Rio Orinoco, VENEZUELA. Ugueta, 29 Sep 1951 (st), Croizat 762 (NY). BARINAS: Barinitas, 25 Jun 1956 (st), Bernardi 3317 (NY);Reserva Forestal Caparo, N of Rio Uribante & Rio Apure, S of

83

TAXONOMIC TREATMENT

CaiioAnaru,9 Jan 1972(st), Marcano-Berti 2909 (MER, U), 13 Jan 1972 (st), Marcano-Berti2924 (U); Dtto. Pedraza,Rio Zapa,15 Aug 1975(d), Rodrigues173 (F); Dtto.Obispos,7 kimS of Barrancas, 4 Dec 1962(st),RuizTeranet al. 1242 (U);DKo.Pedreza,betweenRio Socopo & Rio Michay,Anaro,3 Mar1960 (st), Veillon17.V (F). MARIDA: El Vigia, CainoAmarillo, 4 Feb 1955 (a'), Bernardi 1886 (NY, VEN); Santa Elena, 27 Jan 1954 (c?),Little16025(VEN).ZUUA: Nr.Campamento Guasare, 10 May 1983($ fr),Berget al. 1504 (U); Dtto.Mara,10 kmSWof Campamento Socoyde Carbozulia, 24 Aug 1982 (st), Bunting10313 (U); QuebradaPeraya,affluentof Rio Tonuku,SW of Machiques,29 Aug 1967 (9 fl), Steyermarket al. 99865A (U, VEN). ECUADOR. CARCHI: Between Las Juntas & El Coraz6n, 1500-1600 m, 18 Aug 1994 (d), Palacios 12411 (QCNE).ESMERALDAS: BetweenSantoDomingo & Quininde,8 Apr 1943 (9 fr), Little 6204 (F, US); 10 km SE of Esmeraldas,29 Sep 1965 (st), Little et al. 21170 (NY,QAME,US); Busingade Rio Verde,25 km S of mouthof Rio Verde,30 Sep 1965 (st), Littleet al. 21183 (COL, F, MO, NY, QAME, US); Cant6n Eloy Alfaro, Reserva Ecologica Cotachi-Cayapas,Charco Vicente, 6-9 Sep 1993 (? fr), Palacios et al. 11336 (BG)*. Los Rtos: Rio PalenqueBiological Station,rd. Quevedo-SantoDomingode los Colorados,km 56, 11 Jun1973(? fl), Dodson5235 (CR,F,QCA,U);Jauneche forest, Cant6nVinces, between Mocachi & Palenque, EsteroPeniafiel,15 Aug 1978 (st), Dodson et al. 7098 (MO);nr. Montalvo,27 Jul 1962 (st), Jativa et al. 223 (NY); Pichilingue,20 May 1943 (st), Little6531 (F, K, US). MANABI: Canton Jipijapa, Parque Nacional Machalilla,2-5 kmW of Guale,9 Aug 1992(9 fr), Yanez et al. 1306 (QCA). MORONA-SANTIAGO: Puerto Morona,NE of Rio Morona,30 Sep-2 Oct 1975 (? fl),

Jul 1984 (st), Dodson et al. 14740 (QAME). SUCUMBfos: Dureno, 5 Aug 1986 (st), Cer6n 348 (BG, MO); Limoncocha, 27 Sep 1977 (2 fl-fr), Foster 3875 (F, K, MO, S); Canton El Chaco, Las Palmas, 1600-1750 m, 12 Oct 1990 (? fl-fr), Palacios 6229 (MO, QCNE). Pachicutza, 70 km NE of Zamora, ZAMORA-CHINCHIPE: 1000 m, 14 Sep 1975 (d), Littleet al. 341 (LOJA,QAME, US); Canton Nangaritza, Pachicutza, 5 Dec 1990 (9 fl-

fr), Palacios et al. 6562 (MO, QCNE). PERU. Cuzco: Urubamba valley, Santa Rosa, 1200 m, 9 Jul 1915 (ci), Cook et al. 1719 (F, US); Prov. La Convencion, nr. Hda. Potrero & Quillabamba, 1200 m, 24 Mar 1989 (st), Nuniezet al. 10305 (BG). JUNMN: Tarma/Chanchamayo border, Rio Tulumayo drainage, Rondoyacu, 45 km S of San Ram6n, 1800 m, 15 Oct

1982 (st), Foster et al. 9176, 9178 (MO), 9180 (BG,

Prov.Pataz, Rio Mishiollo, below Ongon, MO). LIBERTAD: 1200 m, Aug 1914 (2 fl-fr), Weberbauer 7057 (F, GH, MOL, NY, US). LORETO:Prov. Loreto, Airico, 11 Dec 1984 (q fr), R. Visquez 6073 (MO, NY); Prov. Maynas, Yanamono, 10 Oct 1986 (9 fr), R. VYsquez8302 (BG, MO); Prov. Maynas, Santa Maria de Nanay, Quebrada Yarina, 19 May 1989 (q fl-fr), R. Vasquez et al. 12172 (BG); Rio Amazonas, Cabello Cocha, 5 Aug 1929 (st), Ll. Williams2086 (F, US). MADREDE DIos: Tambopata, Comunidad Nativa Infierno, Hermosa Chica, 15 Mar 1989 (st), Alexiades et al. 467 (BG); Prov. Tahuamame, Dtto. Iberia, rd. Iberia-Ifnapari,km 32, 17 Nov 1981 (q fr), Diaz S. 1722-4 (MO, MOL, U); Prov. Manui,Parque Nacional del Manu',Rio Manu',Cocha Cashu Station, 29 Jul 1984 (ci), Foster 9711 (MO); TambopataNature Reserve, jct. of Rio La Torre & Rio Tambopata,2 Jun 1987 (st), Gentry et al. 58063 (BG, MO); rd. Inapari-Iberia, km 17, Chilina, 27 Feb 1982 (st), Ruiz C. 12 (BG). SAN MARTtN:Zepelacio, nr. Moyobamba, Jun 1934 (o'), Klug Little et al. 571 (LOJA, MO, NY, QAME, US); Rio 3698 (A, BM, F, GH, K, MO, NY, US); Prov. Lamas, E of Santiago,Santiago,16-17 Oct 1975 (st), Littleet al. 754 San Juan de Pacayzapa, 10 May 1973 (o), Schunke V (LOJA, NY, QAME, US); Bomboiza, 17 km SE of 6230 (F, MO, NY, U); Tarapoto,Nov 1856 (? fl), Spruce Gualaquiza,Jul-Oct 1985 (? fr), Zaruma290 (BG, 4923 (K); Prov. Rioja, rd. Pedro Ruiz-Moyobamba, km QAME,QCA, QCNE);rd. Bomboiza-Gualaquiza,km 390, Venceremos,2100 m, 7-9 Aug 1983 (st), D. N. Smith 8 from Mision, 2-3 Nov 1986 (9 fl-fr), Zaruma361 et al. 4722 (MO); San Roque, 1350-1500 m, 8 Jan 1930 (AAU, BG, MO, NY, QAME,QCA,QCNE).NAPO:20 (2 fr), Li. Williams 7205 (F). UCAYALI:Rd. Pucallpakm E of Baeza, 1500 m, 22 Jul 1986 (a"), Gentryet al. Lima, km 99, arboretum, 28 Jan 1980 (st), Begazo 87 55007 (BG); EstacionExp. INIAP-Payamino,5 km N (MO); El Sacramento, 99 km S of Pucallpa, 23 Jul 1957 of Coca, 9 Sep 1986 (? fl), Neill et al. 7342 (BG, NY, (st), Ellenberg 2511 (U); rd. Pucallpa-Lima, km 99, 12 QCA), (a), Neill et al. 7343 (BG, MO, NY, QCA), 29 May 1975 ($ fl), Hartshorn 1691 (MO, MOL, U), (e) Nov 1986 (2 fr), Neill 7529 (NY); rd. Hollin-Loreto- Hartshorn 1692 (MO, MOL, U). Coca,km40, betweenRio Guamani& Rio Pucuno,1200 BRAZIL. ACRE:Nr. Tarauaca,14 Sep 1966 (q fl-fr), m, 11 Dec 1987 (? fl-fr), Neill et al. 8117 (BG, MO, Pranceet al. 7270 (F, GH, MG, NY, P, U, US). AMAZONAS: QAME);ReservaBiol6gicaJatunSacha,8 kmE of Puerto Nr. mouth of Rio Embira, Rio Tarauaca,26 Jun 1933 (? Misahualli, 2 Oct 1986 (a"fl-fr), Palacios 1425 (BG, MO, fr), Krukoff5008 (MICH, MO, U). ROND6NIA: Mun. NY, QAME,QCA, QCNE);Cant6nQuijos, Chaco,rd. Presidente Medici, rd. Cuiabai-P6rtoVelho, km 300, 28 to Cayambe-Coca,Rio San JuanChico, 1750 m, 12 Jan Jun 1984 (c), Ferreiraet al. 4841 (BG, NY). BOLIVIA. BENI: Prov. Moxos, 130 km S of San 1992 (? fl-fr), Palacios et al. 9570 (BG). PASTAZA: Cant6nPastaza,Pozo PetroleroDanta2, 50 km SSE of Ignacio, Rio Chirizi, 12 Aug 1991 (ae), Del Aguila et al. Curaray,1-19 Oct 1990 (? fl-fr),Rubioet al. 787 (BG, 81 (LPB); Prov. Ballivian, rd. Yucumo-Rurrenabaque, QCNE). PICHINCHA: Rd. Santo Domingo de los km 35, Mar-Jul 1990 (st), D. N. Smith et al. 14381 (BG, Colorados-Quininde,km 170-175, 4 Sep 1949 (st), MO); Prov. Moxos, rd. San Bora-Trinidad, km 27, 25

AcostaS. 13733 (F); Centinela,15 km E of Patricia,31

Aug-3 Sep 1990 (q fl), D. N. Smithet al. 14205 (BG),

84 (d?),D. N. Smithet al. 14206 (BG). COCHABAMBA:Prov.

Carasco,CantonIvirgarzama,CampamentoII, 27 Oct 1987 (? fr), Beck 13684 (BG). LA PAZ: Prov. Abel Alto Madidi,acrossof mouthof Rio Entatagua, Iturralde, 22 May 1990 (st), Gentryet al. 70341 (MO);Prov.Sud Yungas,Rio Bopi, San Bartolome,nr. Calisaya, 1-22 Jul 1939 (? f), Krukoff10410 (A, F, G, K, MICH,MO, NY, U, US); Prov. Larecaja,Tuiri, nr. Mapiri, 12-30 Sep 1939 (9 fl), Krukoff10728 (A, F, G, K, MO,NY,U, US), (e'), Krukoff10813 (A, F, G, MICH,MO, NY, U, US); Prov. Sud Yungas,Alto Beni, Serraniade Marimonos,ColoniaSan Martin,25 May 1991 (st), Seidel et al. 5308 (BG). PANDO: Prov. Manuripi,12 km W of 5 Oct 1991(st), Becket al. 20143 Conquista-Trampolin, (BG,LPB);Prov.NicolasSuarez,Mukden,Jun-Dec1979 (st), Izawa 7A (MO);Prov.Nicolas Suarez,rd. CobijaPuertoRoca,km 53, 2 Jun 1977(? fl), Meneces656 (IS); 74 km SW of Cobija,21 Aug 1988 (? fr),Penningtonet al. 158 (BG). SANTA CRUZ: Prov.Ichilo, ReservaForestal Chore,Rio Ibabo,20 Aug 1990 (? fl), Neill et al. 9447 (MO, QCNE); Prov. Ichilo, Parque Nacional Ambor6,ca. 5 km SE of Rio Surutu,Rio Pitasama,31 Aug 1985 (? fl-fr), Solomon 14191 (BG, MO). Clarisia biflora is rathervariablein shape,dimensions, andtextureof the leaves. Some collections from Panama, Colombia (Antioquia), and Ecuador (Esmeraldas),indicatedwith (*) in the list of specimens examined, such as Castano-Rosalba 12 and 44, Cuatrecasas15278, andAllen5955 (thetype collection of C.panamensis),possiblyrepresenta distinctinfraspecific taxon,differingfromthemajorityof the collections by the large ellipsoid-oblongoid fruitingperianthand 2-4 x ca. 1.5 cm long andtongue-shapedstigmaswith long papillae.These features,however, cannotbe correlateddifferencesin vegetative characters,otherthan therelativelylargelaminas.ThecollectionMatuda15470 from Mexico also has a large fruiting perianth(2.3 x 1.2 cm), but it has finely papillate,ca. 5 mm long stigmasandis ? intermediate betweenthematerialwithlarge fruiting perianths (from Colombia, Ecuador, and Panama)andthatwiththe usualsize of fruitingperianth. Thefruitingperianthis mostlyyellowish to greenish or rarelyred in SouthAmerica,but in CentralAmerica (Mexico to CostaRica) bothred andyellowish fruiting perianths occur, probably equally represented. The fruits are dispersed by canopy bats (Romo, 1996), at least in AmazonianPeru. The collection by Croizat from the upper Rio Orinoco (Venezuela) is not indicated on the distributionmap,partlybecausethe localitycouldnot be traced.

FLORA NEOTROPICA

Merida:maitin. Ecuador:Esmeraldas:savaleta; Los Rios: palo cerano, tillo serrano; Pastaza: wica (Huaorani); Zamora-Chinchipe:chicalminia. Peru: Madrede Dios: mashonaste,mashonasteblanco; San Martin:cousajo;Ucayali: sachavaca micuna.Bolivia: Beni:chicleorpalochicle;Cochabamba: murare;Pando: chicle; SantaCruz:isirillo.

2. Clarisia racemosa Ruiz & Pavon, Syst. Fl. Peruv. Chil. 255. 1798; Ducke, Arch. Jard. Bot. Rio de Janeiro3: 40. 1922; Lanjouw, Recueil Trav.Bot. Neerl. 33: 265. 1936; Burger,Ann. Missouri Bot. Gard.49: 24, t. 6. 1962.Type.Peru.Huanuco:Chinchao-Pozuzo,Ruiz&Pavons.n. (holotype,MA, seen as photographin F; isotype?,F). Fig. 15 SoaresianitidaAllemao,RevistaBrazil. 1: 210, cum tab. 1857; Lanjouw,Trav.Bot. Neerl. 33: t. 11. 1936. Sorocea nitida (Allemao) Warburg,Bull. Soc. Bot. France49(Mem.3): 644. 1913. Clarisia nitida(Allemao)Macbride,Publ.FieldMus.Nat. Hist., Bot. Ser. 11: 15. 1931. Type. Allemao, RevistaBrazil. 1: tab. 1857.

Tree,to 40 m tall; barkconspicuously lenticellate, that of the base of the trunkand of the (larger) roots reddish. Leafy twigs 0.5-3.5 mm thick, minutely puberulous to subglabrous. Lamina elliptic to lanceolate, 4-14 x 2-6.5 cm, often broadest below the middle,sometimesslightly inequilateral,coriaceousto subcoriaceous;apex acuminateto subacute;base acute to obtuse to truncate(to emarginate);margin entire; upper surface glabrous; lower surface sparsely puberulous;venation? prominentto plane above, but the midribmostly ? impressed,the venation ? prominentbeneath;lateralveins 9-18 pairs;tertiaryvenation reticulate; petiole 0.5-1.5 cm long, 1-2 mm thick, sparsely and minutely puberulous; stipules broadly ovate to subcordiform, 0.1-0.3 cm long, minutely puberulous, caducous. Staminate inflorescences on (usually) leafless branchlets, sometimes on shortshoots (spurs),often on the older wood; peduncle0.10.8 cm long, minutelypuberulous;spike 1-I 0(-1 8) cm long; tepals 3-7, free or sometimes connate,variously shaped, 0.5-0.8 mm long, minutely puberulous;stamens 1-2(-3), filaments 0.3-1 mm long, anthersca. 0.3-0.4 x 0.3-0.4 mm;bractsvariously shaped,partly suborbicularand subpeltate,ca. 0.5-0.6 x 0.5-0.8 cm, often ? fimbriate, minutely puberulous, and with minute, embedded yellow dye-containing "glands." Local names. Honduras: Atlantida: masicaran. Pistillateinflorescenceson (usually)leaflessbranchlets, Costa Rica: lechilla, ojoche; Puntarenas: ojoche. sometimeson short-shoots(spurs),usuallyon the older Panama:Chiriqui:nisperoblanco. Colombia:Cundin- wood; peduncle 0.4 cm long, to 1.5 cm long in fruit, amarca:lechero colorado; Huila: caudillo colorado. minutelypuberulous;flowers 1, or sometimes 2; periVenezuela:Barinas:charomorado,charonegro,maitin; anth3-4 mm long, subentireto 4-lobed, puberulousin

TAXONOMICTREATMENT

85

'~~

0

4

0

Fig. 15. Clarisia racemosa. 1. Leafy twig. 2. Branchlet with staminate inflorescences (ex Hort. Mus. Goeldi 120).

3. Branchletwith pistillateinflorescencesand leaf (DuckeRB 13051). 4. Branchletswith infructescences(Bruijn 1553). 5. Leaf (Prance 22889). 6. Leaf (Ducke RB 13051). 7. Leaf (Blanco 509). 8. Stamen. 9a-c. Tepals of stami-

nate flower.10, 10a. Bractsof staminateinflorescence(ex Hort.Mus.Goeldi120). 1l. Pistillateinflorescence.1la. Pistillateflowerand bracts(DuckeRB 13051). 12. Fruitingperianth.13. Seed. 14. Embryo(Prance22889). [By P. Pardoen,Utrecht.]

FLORANEOTROPICA

86

theuppermost part,andwithminute,embeddedyellowdye-containing "glands"; ovaryadnateto theperianth;style1-3 mmlong;stigmasfilifonn,4-7 mmlong; bracts 3-7, suborbicularto broadlyovate, often and 0.8-1.2mmlong,minutelypuberulous, subpeltate, with minute, embedded yellow dye-containing "glands." Fruiting perianth subgloboseto ellipsoid, 1.5-3(-4) x 1-1.7(-2.4) cm, glabrous,orangeto red;

endocarpwoody. Distribution (see Fig. 76.2). FromsouthernMexico to Surinameand easternBrazil, with a ? clearly discontinuousrangeof distribution,butoften locally common; in wet, non-inundatedforest;to 1500 m. Representative specimens examined. MEXICO. OAXACA: Pochutla, Cerro del Machete, 1000 m, Feb

1941 (9 fr), Reko 6291 (F). NICARAGUA. CHONTALES: CerroOluma,top of Cord. Amerisque, 4 Jan 1984 (st), Gentry et al. 43957 (MO). COSTA RICA. ALAJUELA: Upala, Bijagua, El Pil6n, rd. to Chimurria, 20 Jul 1988 (? fr), Herrera 2136 (F, MO). CARTAGO: Turrialba, 26 Oct 1949 (9 fl-fr), J. Leon 1885 (EAP); Turrialba, Punta Caj6n, 21 Mar 1972 (st), Poveda 57 (CR); Rio Reventaz6n, CATIE, 22 Sep 1984 (st), Zamora V 759 (F). PUNTARENAS: Parque Nacional Corcovado, Sirena Trail, 4 Jan 1989 (? fl), Kernan et al. 866 (BG, F, MO), 6 Jan 1989 (9 fl), Kernan et al. 869 (MO), (d') Kernan et al. 871 (BG, F, MO), 22 Mar 1989 (9 fr), Kernan et al. 994 (BG,MO). SAN Jost: 4 km SW of Ciudad Col6n, 16 Feb 1990 (st), Grayumet at. 9671 (BG). PANAMA. CHIRIQUi: Burica Peninsula, W of San

Bartolo, 24 Feb 1973 (st), Croat 22155A (MO). SAN BLAS:Nusagandi, El Llano-Carti rd., ca, km 20. 30 Apr 1992 (st), Paredes 718 (SCZ). COLOMBIA. AMAZONAS:Rio Caqueta, nr. Isla Norrocoy, 22 Sep 1986 (st), Brandet al. 1442 (BG); Rio Loretoyacu, 20 Jan 1973 (st), GlenboskiC.242 (COL); Rio Caqueta, CanioSolarte, 25 Aug 1986 (st), P Palacios et al. 1949 (BG). ANTIOQUIA:Mun. San Luis, Rio Claro, 2 May 1984 (? fr), Cogollo1663 (MO), 3 Jun 1984 (st), Cogollo 1789 (MO). ANTIoQUIA/BOLiVAR: 38 km W of Barrancabermeja,24 Feb 1967 (st), Bruijn 1488 & 1498 (COL, MO, NY, U, VEN), 2-3 Mar 1967 (? fr), Bruijn 1553 (COL, MO, NY, U, VEN). BOLiVAR: 150 km N of Barrancabermeja, Mico-Ahumado Camp, 18 Aug 1966

(st), Bruijn1056 (COL,MO, NY, U, VEN). CAQUETA: Araracuara, 27 Aug 1991 (st), Vester et al. 399 (BG). CHOC6: Mun. Riosucio, Zona de UbarA, Cerros del Cuchillo, rd. Nova-La Cumbre, 23 Mar 1988 (9 fr), Cdrdenas1461 (JUAM, MO). META: San Martin, Finca La Becerra, 23 May 1968 (st), Devia U. s.n. (US); Macarena, Serrania de Chamusa, 21 Apr 1988 (9 fl-fr), Stevenson 71 (MO). VALLE: Rio Digua, Pedra de Moler, 19-28 Aug 1943 (c? & 9 fr), Cuatrecasas 15148 (F); Rio Digua, El Engaiho,27 Mar 1947 (st), Cuatrecasas 24000 (F). VICHADA: Las Gaviotas, CanioAriba, 19 Feb 1973 (6'), Cabrera2662 (COL).

VENEZUELA. AMAZONAS: Upper Rio Orinoco, Isla Rat6n, 31 May 1940 (st), LI. Williams 13191 (VEN). BoLfVAR:Sierra Imataca, El Palmar, Rio Grande, 16 Feb 1959 (q fr), Bernardi 7069 (F, G, NY); 48 km NE of Caserio Los Rosos, 17 km E of Upata, 29 Apr-4 Jun 1965 (st), Blanco 101 (MO, VEN); NE of Upata, El Paralso Camp, 19 Mar 1966 (st), Breteler 5054 (WAG); Dtto. Piar, El Palmar, Rio Grande, La Isabel, Aug 1961 (st), Conejos 85 (VEN); Rio Canaracuni, 13-16 Apr 1988 (e'), Stergios 11746 (BG). BOLiVAR/DELTA AMACURO: Rio Grande o Toro, E of Upata, 8 Apr 1967 (st), Bruijn 1658 & 1659 (MO, U, VEN). DELTAAMACURO:E of Rio Grande, ENE of El Palmar, Nov 1965 (d), Blanco 509 (NY, U, US, VEN), 29 Jun 1964 (st), Marcano-Berti 277 (MO, NY, US, VEN). SURINAME. Kabourie, (tree 597), 6 Nov 1920 (st), BW 4821 (MO, U), 4 Nov 1920 (st), BW 4854 (F, NY, U), 2 Oct 1920 (st), BW 4968 (F, NY, U); Coppename R., Voltzberg, Feb 1978, Troon (LBB) 16297 (U). Nr. Coronel C. Concha, ECUADOR. ESMERALDAS: 23 Dec 1977 (st), Berg et al. 435 (U); nr. San Lorenzo, (st), Houssin 4 (P, S, US); Cant6n Eloy Alfaro, Reserva Ec6logica Cotocachi, Cayapas, Charco Vicente, Rio San Miguel, 6-9 Sep 1993 (? fr), Palacios et al. 11336 (QCNE); 3 km E of Quininde, 12 Apr 1943 (? fr), Little 6240 (K, US); 30 km S of mouth of Rio Verde, Businga, 1 Oct 1965 (st), Little et al. 21188 (NY, QAME, US). Los Rios: Rio Palenque Biological Station, 16 Feb 1974 (st), Gentry 9964 (MO, U); Cant6n Vinces, between Mocachi & Palenque, Jauneche forest, 4 Oct 1979 (st), Dodson et al. 8785 (MO); Jauneche forest, Estero Peniafiel,24 Jan 1981 (? fr), Gentry et al. 30724 (AAU, Cord. de Cutuca, 5-10 NY, S, U). MORONA-SANTIAGO: km E of Logroino, 1200-1500 m, 7-9 Oct 1975 (6), Little et at. 652 (US); Rio Cuyes, rd. Bomboiza-Gualaquiza, 1 Nov 1986 (e), Palacios 1453 (BG, MO, QAME, QCNE). NAPO:Parque Nacional Yasuni, Pozo Petrolero Amo 2, 9-19 Jan 1988 (st), Neill et al. 8307 (BG, QAME, QCNE); Reserva Biol6gica Jatun Sacha, 8 km E of Puerto Misahualli,4 Sep 1989 (e), Neill et al. 9032 (MO, QAME, QCNE, US); 4 km N of Coca, 15 Sep 1986 (? fr), Palacios et al. 1291 (BG, MO, NY, QAME, QCA, QCNE); ca. 30 km NNW of Coca, Rio Huashito, 27 Oct 1982 (? fl), Pennington 10643 (QCA, QCNE, U). PASTAZA:40 km N of Montalvo, 1-17 May 1990 (? fl-fr), Espinoza 148 (QCNE); rd. Coca-Auca, 10 km S of Napo border, nr. Rio Tigiiino, 26-31 Jan 1989 (st), Neill et al. 8859 (BG); rd. Coca-Auca, 110 km S of Coca, 10 km from Rio Tigiiino, 7 Jan 1989 (? fr), Palacios et at. 3388 (MO, NY, QAME, QCNE); 115 Km S of Coca, 7-9 Jan 1989 (? fr), Hurtado 1316 (MO, QCNE). PICHINCHA:Rd. Santo Domingo de los Colorado-Quininde, 2 Sep 1949 (st), Acosta S. 13681 (F); rd. Santo Domingo de los Colorados-Puerto Lim6n, km 23, 22 Jul 1982, Kvist et al. 40713 (AAU). SUCUMBiOS: Rio Cuyabeno, ca. 0?lO'S, 75?55'W, 20 Feb 1980 (st), Berg et al. 1068 (AAU, NY, QCA, U); Dureno, 14 Sep 1986 (? fr), Ceron 385 (MO, NY, QAME, QCNE); Lumbaqui, 14 May 1987 (st), Pennington 12258 (BG, QAME, QCA); Reserva Faunistica Cuyabeno, nr. Laguna Grande, 1 Apr

TAXONOMICTREATMENT 1988-1 Apr 1990 (st), Valencia et al. 67656 (QCA). ZAMORA-CHINCHIPE: 52 km NE of Zamora, 12 Sep 1975 (st), Little et al. 314 (LOJA, QAME, US), 19 Sep 1975 (st), Little 434 (US); Canton Nangaritza, Shaimi, 27 Oct 1991 (9 fr), Palacios et al. 8705 (QCNE). PERU. Cuzco: Prov. La Convenci6n, Alto Urubamba, Manguyari, 3 Feb 1989 (st), Nnfiez et al. 10193 (BG). HuANuco: Nr. Tingo Maria, 10 Aug 1940 (e'), Asplund 12949 (G); Prov. Pachitea, Dtto. Inca, Dantas, 1984 (st), Reynel R. 1047 (U); Prov. Leoncio Prado, Dtto. Rupa Rupa, Bella, 9 Jul 1964 (6'), E. Vdsques A. 34 (F, NY, P, WIS). JUN1N:Prov. Satipo, PichanakiRodal Kimiriki, 15 Jun 1981 (st), Diaz 1698 (MO); Prov. Chanchamayo, Rio Colorado, rd. San Ram6n-Puente Paucartambo, km 23, 6 Oct 1982 (? fr), D. N. Smith et al. 2492 (MO, NY); Prov. Tarma, La Merced, (9 & o), Weberbauer 1912 (F); Rio Negro, 20 Aug 1960 (? flfr), Woytkowski 5865 (GH, MO, US). LORETO:Prov. Requena, Rio Ucayali, 04?55'S, 73?45'W, Aug-Sep 1976 (st), Bernardi 16278 (G, U); Prov. Maynas, Rio Amazonas, above mouth of Rio Napo, Quebrada Yanomono, 5 Nov 1979 (st), Gentry et al. 27476 (MO, U); Rio Amazonas, ca. 2 km W of Indiana, 14 Feb 1987 (st), Gentry et al. 55871 (MO); Prov. Alto Amazonas, Dtto. Yurimaguas, rd. Cainarachi-Yurimaguas, 16 Aug 1968 (st), Soria S. 22 (U); Prov. Maynas, Puerto Almendras, 11 Apr 1988 (6'), R. Vcsquez et al. 10544 (MO); Prov. Maynas, rd. Iquitos-Nauta, km 44, 6 Apr 1989 (9 fr), R. Vasquez12004 (MO); La Victoria, 5 Sep 1929 (st), Ll. Williams 3079 (F, US). MADREDE MIOS: Tambopata, Cumunidad Nativa de Infierno, Hermosa Chica, 29 Mar 1989 (st), Alexiades et al. 542 (BG), 14 May 1989 (st), Alexiades et al. 736 (BG); Mani National Park, Rio Manui, Cocha Cashu Camp, 4-19 Aug 1983 (st), Gentry 43358 (BG, MO); Tambopata, 19-25 Feb 1984 (st), Gentry et al. 45692 (MO); Tambopata Nature Reserve, jct. of Rio La Torre & Rio Tambopata, 23 Jul 1985 (st), Gentry et al. 51170 (BG, MO). PASCO: Rd. Villarica-Cedropampa, 1350 m, 4 Feb 1981 (? fr), Diaz 1638 (MO); Prov. Oxapampa, rd. Puente Paucartambo-Oxapampa, km 8, 2 Jul 1984 (e), D. N. Smith 7567 (MO, NY). SAN MARTiN: Prov. Mariscal Caceres, Dtto. Tocache Nuevo, rd. to Pushurumbo, 1 Mar 1978 (e), Schunke V 9956 (MO, NY, U). UCAYALI: Rd. Pucallpa-Tingo Maria, km 86, 20 Jan 1980 (? fr), Begazo 91 (MO); SE of Pucallpa, nr. jct. of Rio Pachita & Rio Yuyapichis, Panguana, 15 Oct 1985 (st), Morawetz et al. VJO-151085 (BG), 18 Oct 1985 (? fr), Morawetz et al. 14-1810985 (BG); Prov. Coronel Portillo, Atalaya, 2 Mar 1981 (st), Reynel R. 135 (U); rd. Pucallpa-San Alejandro, km 82, 29 Feb 1972 (st), Simpson et al. 60 (F, MO); Yarina Cocha, Oct 1925 (e), Tessmann 5433 (G, NY). BRAZIL. ACRE: Fazenda Sossego, 27 Sep-7 Oct 1985 (st), Campbell et al. 9322 (BG); Mun. Serra Madureira, rd. to Bonsucesso, km 7, 1 Oct 1980 (? fr), Ferreira et al. 2648 (MO, NY, US); mouth of Rio Macaua, tributary of Rio Yaco, 9 Aug 1933 (9 fl-fr), Krukoff 5380 (A, BM, F, G, K, LE, MICH, MO, NY, U, US); rd. Rio Branco-Brasileia, km 42, 16 Oct 1980 (fr),

87 Nelson 759 (BG, MG); Seringal Auristella, Jul 1911 (od), Ule 9317 (K, MG). AMAPA: Reserva INCRA, Rio Falsino, 15-20 Aug 1983 (st), Campbell et al. 9861 (BG). AMAZONAS: Serra Araca, 11 km E of Rio Jauari, 8 Mar 1984 (st), Amaral et al. IG2-2-73 (BG); nr. Manaus, 26 Oct 1935 (a'), Ducke 59 (A, B, F, K, MO, NY, US), 26 Oct 1935 (e'), Ducke 231 (NY); Rio Solimoes, Igarape do Maruja, 5 Jun 1941 (st), Fr6es 132 (F, G, GH, K, NY); Rio Negro, Mun. Sao Gabriel, Yuco, 5 Dec 1941 (st), Fr6es 410 (F, G, MO, NY); mun Sao Gabriel, Rio Castanha, Busseh Cussi, 27 Apr 1942 (st), Fr6es 852 (G, NY); Ponta Alegre, 6 Jun 1904 (st), Huber MG 4367 (MG); nr. mouth of Rio Embira, tributary of Rio Tacauaca, 4 Jul 1933 (? fr), Krukoff 5173 (A, BM, F, G, K, LE, MICH, MO, NY, U, US); Mun. HumaitA,nr. Tres Casas, 14 Sep-11 Oct 1934 (2 fl), Krukoff 6327 (A, B, BM, IAN, K, LE, MICH, MO, U, US), nr. Livramento, 12 Oct-6 Nov 1934 (? fl), Krukoff 6628 (A, BM, F, G, K, LE, MICH, NY, U, US); Mun. Sao Paulo de Oliven9a, nr. Palmares (? fl), Krukoff 8312 (A, BM, F, G, K, LE, MICH, MO, NY, P, U, US); Mun. Sao Paulo de Oliven9a, Belem Creek, 26 Oct-1I Dec 1936 (9 fl), Krukoff 8666 (LE, NY, U); rd. Manaus-P6rto Velho, BR.319, km 380, 2 km S of Rio Jutai, 13 Oct 1974 (? fr), Prance et al. 22889 (NY, U); Rio Negro, Igarape das Araras, SIDERAMA forest, 9 Nov 1963 (R fr), Rodrigues 5519 (F). BAHIA: Without locality, 19 Oct 1946 (st), Bondar s.n. (F); Rio Grongogy, 1 Oct-30 Nov 1915 (st), Curran 96 (US); between Ajuda & P6rto Seguro, 22 Aug 1961 (st), Duarte 5969 (NY, U). ESPiRITO SANTO:Linhares, Reserva Florestal de CVRD, 14 Sep 1982 (e), Folli 395 (BG), 10 Mar 1972 (9 fr), Lino 37/72 (BG), 1 Feb 1985 (st), Peixoto et al. 3128 (BG, MO). MARANHAO: Carutapera,Rio Gurupi, nr. SerraTiracamba,5 Nov 1982 (st), Balee 16 (MG). MATO GROSSO: Rio Machado, nr. Tabajara, Nov-Oct 1931 (2 fr), Krukoff 1450 (A, BM, F, G, K, MO, NY, P, U, US); Aripuana, Nucleo Juina, 16 Jan 1979 (2 fr), M. G. Silva 4269 (MG, NY); Mun. Alta Floresta, 17.5 m N of Alta Paraiso, 30 Sep 1985 (a), Thomas et al. 4123 (MO, NY, US). MINAS GERAIS: Mun. Caratinga, Fazenda Montes Clara, 11 Jan 1980 (? fr), Mishimura 8 (FEEMA, U). PARA: Rd. CachoeiraPalhao, Rio Curuia,(? fr), Cavalcante et al. 1536 (MG); Rio Tapaj6s, Boa Vista, 11 Apr 1933 (? fr), Capucho 568 (F); Faro, 23 Jan 1920 (? fl), Ducke RB 13051 (G, K, U, US); Rio Tapaj6s, nr. Cachoeira da Montanha, 18 Dec 1919 (e?), Ducke RB 13052 (G, K, MO, U, US); Rio Tapajos, Sao Luiz, 4 Dec 1919 (? fl-fr), Ducke RB 13052 (K, U); Rio Trombetas, Morro da Agua Fria, 25 Oct 1919, Ducke RB 13054 (U); Rio Xingu, Volta Grande, 14 Dec 1916 (e), Ducke MG 16606 (G, MG, US); Gurupa, 14 Aug 1918 (st), Ducke MG 17204 (MG); Rio Tocantins, Rio Itacaiunas, Serra Buritirama, Aug 1970 (st), Pires et al. 12813 (IAN); rd. Belem-Brasilia, km 100, Aug 1960 (? fl), N. T. Silva 600 (IAN); Rio Trombetas, 26 Apr 1941 (st), C. W Smith 601 (FG, G, NY, US). RIO DE JANEIRO:Mun. Rio de Janeiro, between Morro da Urca & Pao de Acucar, 19 Mar 1980 (st), Berg et al. s.n. (U); Mun. Rio de Janeiro, Corcovado, 7 Feb 1863 (? fr), Glaziou 859 (BR, F, K, NY, P, US);

88 Mun. Rio de Janeiro, Cosmo Velho, 19 Mar 1876 (q fr), Glaziou 8289 (F, K, P, US); Mun. Rio de Janeiro, Corcovado, 24 Jul 1893 (d + ?), Glaziou 20489 (A, B, BR, C, K, LE, NY, P); Mun. Rio de Janeiro, Mundo Novo, Aug 1920 (c), Kuhlmann RB 13055 (U) & 19 Sep 1920 (? fl), Kuhlmann RB 13055 (U, US). ROND6NIA: Mun. P6rto Velho, Usina Hidroelectrica Samuel, 12-24 Sep 1988 (st), Thomas et al. 6444 &

6520 (BG). RORAIMA: SEMA Ecol. Reserve, Ilha de Maraca, 29 Jul 1987 (? fr), Milliken et al. 526 (BG); Maita Mts., trail Surucucu-Uaica', 13 Feb 1971 (? fr), Prance et al. 10488 (F, GH, K, MG, NY, P). BOLIVIA. BENI: Prov. Ballivian, rd. YacumaRurrenabaque,km 35, 20-25 Jun 1989 (st), D. N. Smith et al. 13568 (MO); Prov. Carrasco, rd. Santa Cruz-Villa Tunari, km 240 , 12-14 Jul 1989 (? fr), D. N. Smith 13727 (BG, MO); Prov. Ballivian, rd. Caranavi-San Bora, Serraniadel Pil6n Lajas, 2 Nov 1989 (9 fr), D. N. Smith et al. 13866 (BG, MO); Rurrenabaque, 14 Oct 1921 ($ fr), 0. E. White 892 (NY), Rio Beni, Huachi, 17 Sep 1921 (st), 0. E. White 1061 (GH, K, NY, US). COCHABAMBA: Prov. Carrasco, Estacion Valle del Sajta UMSS, 4 Apr 1988 (st), Fedlmeier 2 (BG). LA PAZ: Prov. Abel Iturralde,nr. mouth of Rio Enlatagua, 21 Aug 1990 (st), Gentry et al. 70228 (MO); Prov. Sud Yungas, Rio Bopi, San Bartolome, 1-22 Jul 1939 (o), Krukoff 10533 (A, F, G, K, MICH, MO, NY, U, US); Prov. Sud Yungas, Rio Bopi, Asunta, nr. Evenay, 27-31 Jul 1939 (st), Krukoff 10679 (F,NY); Prov. Larecaja, Tuiri, nr. Mapiri, 12-30 Sep 1939 (st), Krukoff 10744 (A, F, G, K, MICH, MO, NY, U, US); Prov. Sud Yungas, Sapecho, 8 Oct 1991

(? fl), Seidelet al. 5458 (BG,LPB).PANDO: Prov.Madre de Dios, rd. Sena-Rio Beni, km 10, 11 Oct 1991 (? fl), Beck et al. 20375 (BG, LPB); Prov. Manuripi, 36 km S of Porvenir, 25 Oct 1977 (9 fr), Hartshorn et al. 2073 (MG, NY); Prov. Nicolas Suarez, rd. Cobija-Puerto Rico, km 53, San Francisco, 31 May 1977 (2 fl), Meneces 643 (IS, MO); Prov. Nicolas Suairez, rd. Porvenir-Puerto Rico, km 124, 11 Aug 1979 (9 fr),

FLORANEOTROPICA Local names and use. Mexico: Oaxaca:juandiego amarillo.Colombia:Bolivar:aji;Caqueti:pelacara.Colombia:Amazonas:guariuba.Venezuela:Amazonas: billo; Bolivar:cajiman;DeltaAmacuro:cajim6n.Ecuador:Esmeraldas:bobo, moral,moralbobo,moralbasto, moralcomida de mono; Los Rios: moral bobo; Napo: moral; Pastaza:chisruya, pitiiik (Jivaro). Peru: Huanuco: tulpay; Junin: ciruela, mashonaste, naranjillo, tulpar;Loreto:Guariuba,mashonaste,sunquirimsuwe (Jivaro);Pasco: tulpay blanco; Ucayali: mashonaste amarillo.Brazil:Acre: guariubaamarela;Amazonas: guariuba,guariubade caranazal;Bahia:orticica;Espirito Santo: oiticica; Maranhao:guar; Mato Grosso: guariuba;Para:janita; Rio de Janeiro:siticica; Roraima: houyoufi(Uaica).Bolivia:Beni:huehue;Cochabamba: urupi; La Paz: mascajo rojo; Pando: murure;Santa Cruz:murur6. The latexis put in boiling wateranddrunkby Jivaro Indiansto treatdiarrhea.

6B. Clarisia Ruiz & Pavon sect. Acanthinophyllum (Allemao)C. C. Berg,stat.nov.,basedonAcanthinophyllumAllemao,RevistaBrazil.1:368. 1858.Type. AcanthinophyllumstrepitansAllemao (= Clarisia ilicifolia (Sprengel)Lanjouw& Rossberg). IncludingAliteria Benoist (see underthe genus) Shrubsor treelets. Lamina mostly spinulose-dentate. Tepalsof the staminateflower connate.Pistillate inflorescences capitatewith several to many flowers; ovary almost free from the perianth.Bracts without yellow dye-containing"glands."

Meneces879 (NY). SANTA CRUZ: Prov.Ichilo,between 3. Clarisia ilicifolia (Sprengel)Lanjouw& Rossberg, Recueil Trav. Bot. Neerl. 33: 717. 1936. ExcoePalacios & Ayacucho, 27 May 1976 (6'), Meneces 424 caria ilicifoliaSprengel,Neue Entdeckungen2: 117. (NY); Prov. Ichilo, 1 km SW of center of Buena Vista, 7 Dec 1990 (9 fr), Nee et al. 40212 (BG, NY); Prov. Ichilo, 1821. Acanthinophyllumilicifolia (Sprengel) W. Reserva Forestal Chore, Rio Ibabo, 16-18 Aug 1990 (e'), Burger,Ann. MissouriBot. Gard.49: 27, t. 7. 1962. Neill et al. 9384 (MO); Parque Nacional Ambor6, 8 km Type.Brazil.Withoutlocality,Anonymouss.n. (hoSW of El Carmen, 15-22 Nov 1991 (st), I. G. Vargas et lotype, B). Fig. 16 al. 1119 (LPB). Pseudosorocea poeppigii Baillon, Adansonia 11: In the central part of the Amazon Basin (as near 297. 1857. Type. Brazil.Amazonas:withoutlocality, (d%),Poeppig s.n. or 2582 (holotype, P; Manaus)two forms can be recognized:one with relaisotype, LE) tively small, narrow,andthin laminaswith a relatively Acanthinophyllum strepitans Allemao, Revista Braacumen da long ("guariuba folha fina") and the other zil. 1: 369, cum tab. 1858. Sahagunia strepitans with larger,broader,and thicker laminaswith a rela(Allemao), Bentham& Hooker, Gen. P1. 3(1): tively shortacumen("guariubada folha larga").How377. 1880. Clarisia strepitans (Allemao) Lanjouw, ever,intermediatesoccur. Recueil Trav.Bot. Neerl. 33: 274. 1936. Type. The isolated occurrenceof the species in Oaxaca Allemao,RevistaBrazil. 1: tab. 1858. (Mexico) is remarkableandmightbe relatedto the secSahagunia racemifera Huber, Bol. Mus. Paraense ondarycenterdiscussedin theintroductionto thefamily. Hist. Nat. 5: 334. 1909. Type. Brazil. Para:nr. The fruits are eaten by birds and spidermonkeys. Obidos, (e), Ducke MG 2885 (holotype, MG).

TAXONOMIC TREATMENT

@3

89

_

i4

I

Fig. 16. Clarisia ilicifolia. 1. Leafy twig with staminate inflorescences (Lanna 1813). 2. Branchlet with staminate inflorescences (Mori et al. 8789). 3. Leaf (M. G. Silva et al 3435). 4. Leaf (Prance et al. 10622). 5. Leaf (Donselaar 1221). 6. Leaf (Pires et al. 12879). 7. Branchlet with pistillate inflorescences (Kuhlmann 6458). 8. Branch with infructescences (Prance et al. 10622). 9. Branchlet with staminate inflorescences. 10, 10a. Staminate flowers. 11. Bract of staminate inflorescence (N. T. Silva 1171). 12. Pistillate flower (Kuhlmann 6458). 13. Pistillate flower. 14. Fruiting perianth. 15. Seed. 16. Embryo (Wessels Boer 1080). [By P. Pardoen, Utrecht.]

90

FLORA NEOTROPICA

Aliteriasagotii Benoist,Bull. Mus.Hist.Nat. (Paris), Ser. 2, 1: 163. 1929. Type.FrenchGuiana.Acarouany,1857 (? fr), Sagot 530 (lectotype,Berg, Fl. Suriname5(1): 185. 1975, P). SoroceastenophyllaStandley,Trop.Woods43: 18. 1935.Type.Brazil.Para:Rio Tapaj6s,B6a Vista, (?fl-fr), Capucho447 (holotype,F; isotypes,G, GH, IAN, NY). ClarisiaspruceanaLanjouw,RecueilTrav.Bot.Neerl. 33: 272. 1936. Acanthinophyllumspruceana (Lanjouw)W. Burger,Ann. MissouriBot. Gard. 49: 30. 1962.Type.Venezuela. Amazonas: RioCasiquiare,betweenLago Pacibaand Rio Pacimoni, Jan 1853(o), Spruce3279 (holotype,K; isotypes, BM, BR, G, P). Clarisiastrepitans(Allemao)Lanjouwvar.guianensis Lanjouw,RecueilTrav.Bot. Neerl.33: 274. 1936. Clarisiailicifolia(Sprengel)Lanjouw& Rossberg var.guianensis(Lanjouw)Lanjouw& Rossberg, RecueilTrav.Bot.Nerl. 33: 718. 1936.Type.Suriname.SurinameR., nr.Goddo,28 Jan 1926(? flfr), Stahel 120 (holotype,U). Clarisiastrepitans(Allemao)Lanjouwvar.paraensis Lanjouw,RecueilTrav.Bot. Neerl.33: 275. 1936. Clarisiailicifolia(Sprengel)Lanjouw& Rossberg var.paraensis (Lanjouw)Lanjouw& Rossberg, Recueil Trav.Bot. NMerl.33: 718. 1936. Type. Brazil. Parai:without locality, Jan 1905 (? fl), HuberMG 6965 (holotype,U; isotypes,BM, P). Clarisiastrepitans(Allemao)Lanjouwvar.micrantha Lanjouw, Recueil Trav. Bot. NMerl.33: 275. 1936. Clarisia ilicifolia (Sprengel)Lanjouw& Rossbergvar.micrantha(Lanjouw)Lanjouw& Rossberg, Recueil Trav. Bot. Neerl. 33: 718. 1936. Type.Brazil.Rio de Janeiro:Mun.Rio de Janeiro,between Mendanhaand Realengo, 29 Aug 1880 (e), Glaziou 12172 (holotype, K; isotypes,B, BR, F, G, GH, LE, P). Clarisia strepitans(Allemao) Lanjouwvar. cuyunensis Lanjouw,RecueilTrav.Bot. Neerl.33: 275. 1936. Clarisia ilicifolia (Sprengel)Lanjouw& Rossbergvar.cuyunensis(Lanjouw)Lanjouw& Rossberg, Recueil Trav. Bot. Neerl. 33: 718. 1936. Type. Guyana. Cuyuni R., nr. Camasia rd., Feb 1931 (a), Davis (FD) 1029 (holotype, U; isotype, K).

venationreticulate; petiole0.3-1.5 cmlong,1-2.5mm thick,oftenpuberulouswithuncinatehairs;stipules subulate,0.1-0.3 cm long, puberulous,caducous. Staminate inflorescences usually on short-shoots

to 0.5cmlong,(rather) (spurs); peduncle denselypuberulouswithuncinatehairs;spike0.2-2(-6.5) cmlong; perianthca. 0.5 mm long, (2-)3-4-lobed to -parted, minutelypuberulous;stamens1(-3); filamentsca. 0.5-1.5 mmlong;anthersca. 0.2-0.3 x 0.2-0.3 mm; bractsvariouslyshaped,often subpeltate,minutely puberulous, sometimeswithuncinatehairs.Pistillate inflorescences usually on short-shoots (spurs); pe-

duncleto 0.3 cmlong,to 2 cmlongin fruit,sparselyto denselypuberulouswithuncinatehairs;headca. 0.5 cmdiam.;flowers6-20; perianthca. 1.5-2 mmlong, subentire,almostglabrousto ratherdenselypuberulous,sometimespartlywithuncinatehairs;ovary(almost) free; style 0.5-1 mm long; stigmastongueshaped,1-2 mm long;bractsbasal(subtendingthe marginalflowers), sometimesamongthe flowers, broadlyto narrowlyovate,to 1 mm long, minutely oftensomeof thebractsthickenedandto puberulous, 2 mmlongin fruit.Fruitingperianth (subgloboseto) ellipsoidto oblongoid,1.2-2 x 0.8-1.3 cm, (yellow to)orange(tored);fruitadnateto thelowerpartoronly to thebasalpartof theperianth. Distribution (see Fig. 75.1). In the AmazonBasin,

theGuianas,andeastemBrazil;in theundergrowth of non-inundated forest;atlow elevations.

Representative specimens examined. COLOMBIA. GUAVIARA: Rio Vaupes, nr. Miraflores, 6 Feb 1944 (? fr), Gutierrezet al. 759 (COL, GH). VAUP1S:Confluence of Rio Macaya & Rio Ajaju, Sierra de Chiribiquete, 1516 May 1943 (? fr), Schultes 5417 (COL, F, MO). VICHADA: Las Gaviotas, Monte del Cafio Ariba, 17 Jul 1972 (V fr), Cabrera 2400 (COL). VENEZUELA. AMAZONAS: Cacuri, 24 Apr 1976 (o), Colchester 2382 (K); Depto. Atures, 23 km SE of Puerto Ayacucho, 18 Jan 1989 (? fl-fr), Cuello 336 (MO), (e), Cuello 337 (MO); Depto. Atabapo, Rio Cunucunuma, Mar 1990 (? fr), A. Ferndndez 7492 (BG, NY); Rio Ventuari, nr. Salto Tencua, 26 Apr 1971 (? fr), Foldats 145A (NY, VEN); Sierra Parima, Rio Matacuni, nr. Simarawochi, 23 May 1973 (? fr), Steyermark 107428 Shrubor tree, sometimes to 20 m tall. Leafy twigs (F, NY, U); Depto. Atures, 25-30 km SE of Puerto 1-4 mm thick,puberulous,usuallywith uncinatehairs; Ayacucho, Cafio Culebra, 12 May 1980 (a'), Steyermark peridermoftenflakingoff. Laminaellipticto (narrowly) et al. 122349 (MO, NY, U); nr. Puerto Ayacucho, belanceolate, (3-)8-33 x (I-)2.5-12 cm, broadestat or tween airport& Rio Cataniapo,26 Jul 1967 (Vfr), Wessels sometimes below the middle, almost equilateral, Boer 1879 (NY, U, VEN). BOLiVAR: Mun. Aut6nomo (sub)coriaceous;apex(caudate-)acuminate, the acumen Sucre, lower Rio Caura, 18 km SW of Maripa, 1 Apr 1990 spinulose; base acute to obtuse; marginentire, spinu- (? fl), Aymard et al. 8586 (BG, NY); rd. Calcara del lose-dentate, sometimes sinuate or lobate;uppersur- Orinoco-San Juan de Manapiare, km 202, Rio Suapure, Mar 1975 (a'), Delascio et al. 2823 (VEN); Mun. Raul face glabrous;lowersurfaceglabrousor sparselypuberLeoni, Rio Paragua-Rio Paramichi, Mar 1988 (9 fl-fr), ulous with uncinatehairs;midribdistinctlyprominent Delgado 64 (BG, MO, NY); Mun. Raul Leoni, Rio and other veins slightly prominentto plane above, ? Paragua, 13 km N of Estaci6n Karun de EDELCA, May prominentbeneath;lateralveins (5-)8-20 pairs;tertiary 1988 (st), A. Fernandez 4614 (BG); Rio Canaracuni, 16-

TAXONOMIC TREATMENT 26 Apr 1988 (d), Stergios 11831 (BG); Rio Caura,Raudal Guanagujania,9-26 May 1988 (? fl-fr), Stergios et al. 12605 (BG, MO, NY). GUYANA. Mazaruni Station, 20 Jun 1940 (? fr), Fanshawe 502 = FD 3238 (NY), 23 Jun 1942 (e), Fanshawe 736 = FD 3472 (NY); Essequibo R., Moraballi Cr., 20 May 1943 (? fl-fr), Fanshawe 1303 (= FD 4039) (NY); Upper Demerara-Berbice region, Berbice R., N of Kwakwani, 16 Apr 1993 (? fl-fr), Hoffman et al. 3959 (BG). SURINAME. Emmaketen, 7 Nov 1959 (st), Daniels et al. 1079 (U); Brokopondo District, 2 km S of Gansee, 21 Apr 1964 (st), Donselaar 1221 (U); upper Saramacca R., nr. Jan Basi Gado, 24 Feb 1951 (? fr), Florsch0tz et al. 1633 (U); 8 mi from Afobakka, 24 Feb 1965 (st), Lems 104 (NY); Mapane Creek area, Jodensavanne, 30 Sep 1953 (a), Lindeman 4778 (MO, NY, U), 9 Dec 1953 (? fl-fr), Lindeman 5178 (F, MO, NY, U, US); Marowijne District, S of Moengo Tapoe, 14 Jan 1954 (st), Lindeman 6161 (U); Coppename R., nr. Bitagro, 9 Aug 1954 (st), Lindeman 6458 (U), Jodensavanne, 10 Dec 1954 (st), Lindeman 6776 (NY, U); Oelemari R., 3?6'N, 54?33'W, 24 Mar 1963 (? fr), Wessels Boer 1080 (NY, U). FRENCH GUIANA. Arataye R., Sauts Parare, 26 Aug 1983 (st), Barrier 4047 (BG); Mt. Yaroapi, Saut Coueki, 29 Apr 1970 (9 fl-fr), Granville T618 (P); Mt. Grand Matoury, 13 Nov 1974 (a), Granville 2310 (NY); upper Inipi R., N of Emerillon Creek, 25 Sep 1980 (st), Granville 3977 (U); nr. Saul, 7 Feb 1972 (? fl-fr), Granville B.4283 (P), 21 Oct 1972 (o'), Granville B.4622 (U); basin of Maroni R., Gobaya Soula, 6 Jan 1989 (st), Granville et al. 2120 (US); Trois Sauts, Zidockville, 23 Jan 1975 (st), Grenand 675 (U); Mt. Mahury, 16 Feb 1965 (? fr), Halle 1092 (NY, U, US); nr. Cayenne, (a), Martin s.n. (BM, K); Saul, 17 Dec 1976 (e), Mori et al. 8789 (U);Yaloupi R., Saut Quaimicouare, 13 Mar 1970 (? fr), Oldeman 342 (U); Camopi R., Saut Quasseye, 13 Dec 1967 (2 fl-fr), Oldeman 2676 (NY, U); Yaroupi R., Saut des Polissoirs, 23 Apr 1970 (? flfr), Oldeman B.3064 (NY); upper Oyapoque R., Saut Couyari, 9 Apr 1970 (? fl-fr), Oldeman B.3173 (P); Acouarany, 1857 (e'), Sagot 530 (BM, P). PERU. LORETO: Prov. Maynas, Dtto. Iquitos, nr. Santa Clotilde, 19 Dec 1975 (? fr), Rimachi Y 2027 (NA); Prov. Loreto, nr. Nauta, 19 Jan 1996 (st), Ruokolainen et al. 8456 (TUR). BRAZIL. ACRE:Mun. Rio Branco, rd. Rio BrancoP6rto Acre, km 33, 12 Oct 1980 (e), Ferreira et al. 2865 (MG, MO, U, US), (9 fl-fr), Lowrie et al. 487 (MG, MO, NY, U). AMAZONAS: Mun. Tefe, Vila Nogueira, 10 Nov 1986 (9 fl-fr), Daly et al. 4310 (BG, MO, NY); Manaus, Igarapedo Passarinho,23 Sep 1955 (a), Francisco INPA 2025 (MG, MO); nr. Calama, Nov-Dec 1931 (? fr), Krukoff 1309 (B, BM, MICH, MO, NY, K, P, U); Mun. Borba, nr. Bella Vista, 4-6 Sep 1934 (st), Krukoff 5991 (A, K, NY, U); Mun. Manicore, nr. Santa Fe, 8-11 Sep 1934 (? fl-fr), Krukoff6049 (A, B, BM, F, G, LE, MICH, MO, NY, U, US); Mun. Manicore, nr. Concei9do, 8-11 Sep 1934 (e), Krukoff6033 (A, K, NY, U); Mun. Humaita, nr. Livramento, 12 Oct-6 Nov 1934 (? fr), Krukoff6849 (A, BM, F, G, LE, MICH, MO, NY, U, US); Rio Madeira, Democracia, 31 Aug 1923 (a'), KuhlmannRB 19853 (U);

91

rd. Manaus-Itacoatiara,km 75-79, 16 Oct 1963 (st), Oliveira2681 (IAN, U); Maues,30 Nov 1946 (? fl-fr), Pires 118 (COL,IAN, NY, US, VEN);SerraAraca,base of westernmassif,11 Jul 1985(2 fr),Pranceet al. 29467 (MO,NY). BAHIA: Rio Mucuri,FazendaCanaao,9 Oct 1946 (st), Bondars.n. (F, GH, NY, US); Mun.Aurelino Leal, rd. AurelinoLeal-Lagodo Banco, km 10-11, 16 Feb 1994 (? fl), Kallunkiet al. 562 (SPF);rd. PradoItamaraju, 65 kinNW of Prado,18 Sep 1978(d'), Moriet al. 10659 (NY,U); Mun.Itanagra,rd.Itanagra-Subauma, 8kmWofItanagra,26May 1981(2 fr),Morietal. 14117 (NY);Almadina,Serrada Pancahinha,3 Aug 1971 (d), Pinheiro 1534 (U); Ubaitaba,4 Sep 1970 (d), Santos 1086 (U). ESPiRITOSANTO:35 kanN of Conceigao da Barra, 11 Nov 1968 (R fr), Almeidaet al. 223 (U); Linhares, ReservaFlorestada CVRD,27 Aug 1988 (a), Folli 784

(BG), 4 Jan 1989 (a' fr), Folli 828 (BG); Mun.Castelo, Caxixa,17 Sep 1985 (e), Hatschbach et al. 49887 (MO); nr. Linhares, 2 Oct 1971 (a), Santos 2039 (U). MARANHAO: Mun.Mon9ao,Rio Turia9u,P. I. Guaja,29

Jun 1987 (st), Balee 3501 (BG);FazendaBacaba,3 km NW of Lago do Junco, 4 Oct 1980 (a'), Daly et al. 483

km 50, 24 Oct (NY,U, US); rd. SantaLuzia-A9ailAndia, Boa 1980(9 fl-fr),Daly 732 (NY,U); Rio Maracassume, Esperanca, 5 Sep 1932 (a), Fr6es 1869 (A, BM, F, G,

MICH,MO, NY), Boiciquara,Viana, 16 Nov 1959 (? fr),Fr6es34905(IAN);rd.BR.222,km293, 17 Dec 1978 (? fr),Rosa et al. 3035 (F, NY, U). MATOGROSSO:Rd. km 755, FazendaCachimbo,19 Nov Santarem-Cuiaba, 1976 (? fr), Cordeiro1079 (NY,U), 2 Dec 1976 (? fr), Cordeiro 1290 (NY); Rio Machado, source of Rio Dec 1931 (? fr),Krukoff1669 (A, BM, G, K, Jatuarana, MICH,MO,NY, P, U); Mun.Alta Floresta,12 km N of Alta Paraiso,30 Sep 1985 (? fl-fr), Thomaset al. 4109 (BG, MO, NY, US). MINAS GERAIS: Mun. Coronel Fabriciano,Rio Piracicaba,28 Aug 1957 (a'), Fr6es 33426

(IAN), 12 km E of PonteNova, 7 Dec 1958(? fr),Irwin 2254 (NY,US); Mun.Governador Valadares,Campode Sementes,25 Nov 1941 (? fl-fr),Magalhaes843 (IAN, US); Vigosa, 8 Oct 1930 (a), Mexia 5158 (A, BM, F, G,

GB, GH,K, MICH,MO,NY, U, US, Z); Mun.Valadares, Campo de Semuites, 21 Aug 1942 (a'), Oliveira 1072 (IAN, US); Simonesia, 16 Aug 1969 (a), Lanna Sobrinho

1813 (NY,U, US); confluenceof Rio Doce & Rio Turvo, ReservaFlorestalde Rio D6ce, 30 Aug 1973 (a'), Sucreet al. 10162 (F). PARA:Belem, 26 Jan 1943 (? fl-fr), Arkm cher 8197 (IAN, NY, P, US); rd. Altamira-Itaituba, 23, 14 Aug 1978 (? fl), Bahia 36 (MO,U); Rio Xingu, AssuriniIndians,10-12 Jun 1986 (st), Balee 2322 (BG); Belterra,17 Oct 1947 (? fr), Black 47-1715 (IAN, NY, VEN); 18 S of Tucurui, 5 Nov 1981 (a), Daly et al. 1215

(MO, NY), (? fr), Daly et al. 1223 (MO, NY); Rio Itacaiunas,ca. 80 km S of Maraba,3 Dec 1981 (? fr), Daly et al. 1644 (BG, MO,NY); Serrade Carajas,8-12 Dec 1981(? fr),Daly et al. 1805 (BG,MO,NY); Belem, 10 Jul 1945 (da), Ducke 1726 (A, F, IAN, MG, NY, U, US); Alemquer, 26 Dec 1903 (a), Ducke MG 4899 (BM,

2 Sep 1923(d), Piquiatuba, MG,US); Serrade Santar6m, DuckeRB 18325(K, U); Obidos,Mamauru,13 Sep 1926 (a), Ducke RB 19473 (G, K, U, USA); Rio Tocantins, 24 Sep 1948 (a), Froes 23516 (IAN, U); Monte Alegre, 8 Sep 1953 (a'), Fr6es 30546 (MO); Rio Tapaj6s, Ford-

FLORANEOTROPICA

92 landia, Sep 1931 (a'), Krukoff1054 (A, BM, F, K, MICH, P, U); Rio Moju, Fabrica,Dec 1953 (a), Pires et al. 4559 (IAN, US); rd. BR.22, km 96, nr. Cachoeira,29 Oct 1965 (a'), Prance et al. 1780 (F, GH, IAN, NY, P, U, US); rd. BR.230, 15 km W of Ruropolis, Presidente Medici, 29 Nov 1977 ($ fr), Prance et al. 25884 (MO,U) & 25888 (MO, NY, U); Rio Xingu, Gleba Bacaja,just below mouth of Rio Bacaja, 22 Nov 1960 (q fl-fr),Prance et al. 26381 (NY); Santarem, Estrada do Pilhao, km 35, Igarape Curupira,3 Sep 1969 (a), M. Silva et al. 2492 (COL, F, GH, MG, NY, U, US); Rio Jari, Monte Dourado, 8 Oct 1968 (ae), N. T Silva 1144 (IAN, NY, U, US); Rio Jari, Pilao, 24 Jan 1969 (q fr), N. T Silva 1689 (IAN, NY); Serrade Carajas,8 km SE of Rio Itacaiunas, 17 Jun 1982 (), Sperlinget al. 6400 (MO,NY,US). RIODE JANEIRO: Mun. Rio de Janeiro,Corcovado, 28 Sep 1921 (a), Ducke et al. RB 6153 (K, U, US);Mun. Rio de Janeiro,Mata do Pai Ricardo, 8 Oct 1926 (e' + q), Kuhlmann 673 (U); Goitacases, 12 Nov 1943, Kuhlmann 6458 (U); Mun. Rio de Janeiro, Trapicheiro, 14 Nov 1925 (q fr), Kuhlmann

RB 19292(K, U, US). RONDONIA: Mun.CostaMarques, Chapadados Perecis, 15 Jun 1984 (a'), Ferreiraet al. 4555 (BG); Porto Velho, 10 Sep 1971 (? fl-fr), Maguire et al. 56675 (F, GH, K, MO, U, US); rd. Cuiaba-P6rto Velho, nr. Santa Barbara, 17 Aug 1968 (e), Prance et al. 7180 (F, GH, MG, NY, P, U, US); Forte Principe da Beira, Igarapeda Viuva, 5 Jan 1962 (9 fr), Rodrigues et al. 4234 (F); Mun. Porto Velho, Usina Hidroelectrica Samuel, 1224 Sep 1988 (st), Thomas et al. 6245 (BG);Mineragao Campo Nova, 120 km WSW of Ariquemes, 18 Oct 1979 (? fr), Vieiraet al. 512 (MO, NY, US); nr.Vilhena, 31 Oct

1979 (? fr), Vieiraet al. 813 (MG,NY, U). RORAIMA: SEMA Ecol. Reserve, Ilha de Maraca, 8 Apr 1987 (a'), Milliken 57 (BG),25 Apr 1987 (q fl), Milliken 95 (BG); nr. Maita, between Sucurucu & Uaica, 15 Feb 1971 (? fr), Prance et al. 10535 (F, GH, K, MG, NY, P, U), 20 Feb 1971 (R fr), Prance et al. 10622 (MG, MO, NY, U); Rio Mucajai, Porto Mucajai, 19 Mar 1971 (? fr), Prance

et al. 11115 (F, GH, K, NY, U). BOLIVIA. BENI: Rio Sena, 4 Oct 1923 (? fl-fr), KuhlmannRB 19831 (K);Rio Beni, Cachuela Esperanza, 1924 (st), Meyer 9 (U, Z), Nov-Dec 1923 (? fr), Meyer 105 (MO, U, US, Z), 10 Aug 1921 (a' + ? fl), Meyer 112 (U, US, Z), Sep 1921 (a'), Meyer 172 (MO, NY, U, US, Z), 1923 (? fl-fr), Meyer 221 (MO, U, Z); Prov. Vaca Diez, rd. Riberalta-Guayamerin, 17 Sep 1981 (e),

Solomon6289 (GB, LPB,NY). PANDO: Prov.Manupiri, Conquista, rd. Puerto Rico-Sena, km 18, 7 Oct 1991 (9 fl-fr), Beck et al. 20211 (LPB); prov.Madre de Dios, rd. Sena-Rio Beni, km 10, Beck et at. 20354 (LPB); Prov. Madre de Dios, Rio Madre de Dios, 19 km WSW of Ribralta, 2 km E of Candelaria, 18 Aug 1985 (st), Nee 31391 (BG, MO, NY), (a'), Nee 31397 (BG, LPB, MO, NY); prov Abufia, Rio Orton, 8 Jul 1992 (st), L. Vargas et al. 727 (LPB).

ally long (2-6.5 cm) staminateinflorescences.Justas in C.biflora,thefruitingperianthvariesfromoblongoidto globose.Accordingto the labelof Meyer112 (Bolivia), the staminateinflorescencesareveryfragrant.Thefruits areeatenby monkeysandbirds(vanRoosmalen,1985). Local names. Suriname:letterhout,manletterhout, man-redhout, oeman-redhout. Brazil: Amazonas: hihounahi (Yanomani),janete; Bahia: amorabranca; EspiritoSanto:almorabranca;Mato Grosso:moracca chocolate; Minas Gerais: f6lha da serra; Roraima: hualina (Uaica), xamamore (Mucajai), xamaroenai (Mucajai). NAME EXCLUDED Clarisia mollis Standley, Ann. Missouri Bot. Gard. 30: 85. 1943 = Soroceapubivena Hemsley subsp. pubivena.

Sorocea 7. Sorocea Saint-Hilaire,Mem. Mus. Hist.Nat. 7: 473. 1821. Gaudichaud,Voy.Bonite, Bot. Atlast. 71-74. 1844; Miquel, in Martius,Fl. Bras. 4(1): 111, 213. 1853; Burger,Lanjouw& Wessels Boer,Acta Bot. Neerl. 11:428. 1962;Marques,Carauta,Gongalves Costa & Sucre,Anais Acad. Brasil. Ci. 48(2): 286. 1976; Berg & Akkermans,Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med. Sci. 88: 381. 1985. Lectotype,Burger,Lanjouw& Wessels Boer,Acta Bot. Neerl. 11: 430. 1962. Sorocea bonplandii (Baillon) W. Burger,Lanjouw & Wessels Boer. BalanostreblusKurz,J. Asiat. Soc. Bengal., Pt. 2, Nat. Hist., 42: 247. 1873. Type. Balanostreblus ilicifolius

Kurz (= Sorocea

guilleminiana

Gaudichaud). PseudosoroceaBaillon,Hist. P1.6: 210. 1875,Adansonia 11: 296. 1875. Lectotype,designatedhere. Pseudosorocea bonplandii Baillon (= Sorocea

bonplandii (Baillon) W. Burger, Lanjouw & WesselsBoer). Trophisomia RojasAcosta, Bull. Acad. Inst. Geogr. Bot. 24: 211. 1914. Lectotype,IndexNominum Genericorum 3: 1808. 1979. Trophisomia edulis Rojas (= Sorocea sprucei (Baillon) Macbride

subsp.saxicola (Hassler)C. C. Berg). ParaclarisiaDucke,Arq.Serv.Florest.1(1):2. 1939. Type. Paraclarisia amazonica Ducke (= Sorocea

duckeiW. Burger).

Treesor shrubs,dioecious; twigs usually with conspicuous lenticels; uncinate hairs lacking. Leaves alThe leaves are very variablein shape, dimensions, temate and distichous;laminapinnately veined; marandmargin.The leaves may resemblethose of Sorocea gin (spinulose-)dentateto entire;stipules free, lateral. guilleminianaandrelatedspecies with a spinulose-den- Inflorescencesin pairsor solitary (or several together tate margin and acumen. Clarisia ilicifolia can be distinon short-shoots)in the axils of the leaves or below the guished from those species by the midribprominent leaves,racemoseto spicateto subcapitate(oruniflorous), above. The type collection of C. spruceanahas unusu- bracteate;bractssmall,basallyattachedto peltate,cordi-

TAXONOMIC TREATMENT

staminate flowerswith4 form,ovate,orsuborbiculate; tepals;tepalsdecussate-imbricate, mostlybasallyconnate;stamens(3-)4, free,straightin thebud,atanthesis oftenbentinwardsor outwards;anthersbasi-to theconnectivemostly dorsifixed, extrorse(tolatrorse), broad,often(gland-like) apiculate; pistillodenormally absent;pistillateflowerswith a tubular,4-lobedto subentireperianth,its upperandlowerpartusually? different (indiameter, surface, indumentum); lowerpart of theovaryadnateto thelowerpartof theperianth, the ovarygraduallynarrowedinto the style; stigmas2, equal,short,mostlytongue-shaped; the rachis,the pedicel,or sometimesalsothepedunclemoreor less camoseandred(ororange)in fruit;fruitingperianth enlarged,fleshy,red,pink,ororange,(always?)tumingblack(ish)atfullmaturity, mostlyona(nelongated) pedicel.Fruitlarge,mostlygloboseto ellipsoid,adnate to the perianth;endocarpthin, subcoriaceousto subcrustaceous; seedlarge,withoutendosperm; testa thin,witha thickened,(sub)orbicular partbelowthe hilum;embryo(always?)green,longitudinally aligned; cotyledonsveryunequal,thesmalleroneminuteand plane,enclosedby thelarger,thick,curved,conduplicate,andlobedcotyledon;radicleshort,apical.

93

species areevergreen;only S. duckeiandS. spruceiare deciduous. The lenticels are mostly conspicuous and prominent.The latex is white and usually copious. Leaves: The lamina is slightly to pronouncedly inequilateral,mostly coriaceous, sometimes subcoriacious to chartaceous.The leaf marginis mostly dentate or denticulate. In Sorocea guilleminiana and S. bonplandii, the teeth, as well as the leaf acumen, are spinulose.In S. hilarii andS. sprucei,the teethandleaf acumenaresometimesvery shortlyspinulose. In most species,thetertiaryvenationis scalariform,butin some, includingS. affinis and S. muriculata,it is reticulate. Indumentum: Two types of trichomesare found: 1. Unicellularhairs on all partsof the plant, but often (glandular) hairs sparse.2. Globose-capitate, pluricellular on young partsof the plant,often frequenton the lower leaf surface.In some species (e.g., Soroceabriquetii,S. guilleminiana, and S. ruminata),they are abundantly presenton the lowerpartof the perianthof thepistillate flower.Therethey form or exude a substratefor a fungus with a dense white mycelium, which can be found as whiteblotcheson the enlargedfruitingperianth.The fungussometimesproduces"fruit-bodies." Inflorescences: The inflorescencesarebome solitary or in pairs in the leaf axils, or several (to 4 or 6) HISTORY togetheron axillaryshort-shoots.In most species (such ThegenusSoroceawasestablished bySaint-Hilaire as in Sorocea hilarii), the inflorescences also appear (1821)in a studyof theveryunequalcotyledonsinthe on the older wood, occasionally on the trunk.The ingenus.In 1844,Gaudichaud publishedplatesof some florescencesarebasicallyracemosewith the flowers ? Thefirsttreatment with less distinctlyadaxialon therachis.The staminateflowspecies,withoutdescriptions. of species,sevenin total,wasby Miquel ers aresessile in most species, but arepedicellate in S. descriptions (1853)in MartiusFloraBrasiliensis.Someadditional affinis, S. bonplandii,S. guilleminiana, andS. hilarii. specieswerepublishedby variousauthorsupto 1962, In S. affinis, the flower can be either pedicellate or whenarevisionof thegenuswaspublishedbyBurger, sessile. Pistillate flowers are often sessile at anthesis, Lanjouw,andWesselsBoer.Thisrevisioncomprised but afterwardsa pedicel is developed.Pedicels present 22 species,of whichsevenwerenew.In 1966,Cuatre- at anthesisusually elongate afterwards.The bractsare to thepresent small and basally attachedto peltate. They cover the casasaddedthreespecies.A precursor revisionwaspublished byBergandAkkermans (1985): very young flowers, especially in staminateinflores16 specieswererecognized,oneof themnew;andin cences. The bractsaresubcoriaceousto coriaceous,but Conclu- submembranaceousin S. duckei and S. sprucei. fourspecies,twosubspecieswererecognized. Staminate Flower: The perianthof the staminate sionsbasedon morerecentcollectionshavereduced thenumberof speciesto 14,includingtwospeciesand flower consists of four decussate-imbricate tepals, one new subspecies,whichhavebeenmorerecently mostly basally or up to halfway connate. Sessile flowdescribed(Bergin Berg& Franco,1996). ersoften have a very broadbase. The flowersnormally Somegenerarecognizedas distinctfromSorocea, have four isomorphic stamens,but the numberof staBalanostreblus Kurz(1873),Pseudosorocea Baillon mens is often reduced to three. In species where this (1875b),andParaclarisiaDucke(1939),wereincluded reductionoccurs,one can also find anisomorphyin the in thegenusby Burgeret al. (1962),Paraclarisia be- androecium.The pair of stamens opposite the outer ingreducedto a subgenus.TrophisomiaRojasAcosta tepals can have distinctly shorterfilaments and sometimesalso smalleranthers.Moreover,aberrationsin the (1914)provedto be a synonymof Sorocea as well. length of the filaments and the dimensions of the anthers are not uncommon. In Sorocea duckei and S. MORPHOLOGY Habit: MostSorocea speciesareshrubsor small sprucei, the filaments are very short and the anthers trees,butseveralspeciesaretrees,whichcanreacha subsessile.Althoughin most species, the filamentsare of theSorocea ratherthickandshorterthanthe perianth,they areslenheightof 20 oreven25 m.Themajority

94

FLORA NEOTROPICA

der and longer than the perianthin S. trophoides.At anthesis,the stamensarestraight,bent inwardsor outwards.Incurvedstamensare,althoughnot very clearly, associatedwith a broadlysessile flower.Whenthe stamens are straight, the antheris often perpendicular, pointingoutwards.The anthersareextrorse(to slightly latrorse).The connective is mostly broadand is often extendedinto a gland-likeapiculum.The pollen grains of several Sorocea species have been described by Niezgoda andNowaczyk (1976). Fruit: White blotches on the fruitingperianthare caused by fungi. The embryo is (always?) green and has very unequalcotyledons,a phenomenondescribed by Saint-Hilaire,who establishedin the samepaperthe genus Sorocea (1821).

with a disjunctdistributionin SouthAmerica,arespecies of (rather)dry forestor shrubvegetation.Sorocea duckeiis an element of varzeavegetation in the Amazon Basin. Three elements can be distinguished: a group of species in CentralAmerica and the Pacific Coastalregions of Colombia and Ecuador:a groupof species in the Amazon Basin, in particularthe upper Amazon Basin: and a small group of species south of the Amazon Basin. Sorocea pubivena and S. guilleminianaarethe most widespreadspecies, occurring in more thanone phytogeographicalregion. The other species have smallerranges of distribution,and some, such as S. jaramilloi, S. ruminata, and S. sarcocarpa, have very smallranges,occupying only a part of the phytogeographicregions they are associatedwith.

AND DISPERSAL POLLINATION

Nothing is known about pollination in Sorocea. None of the features of flowers and inflorescences suggest the occurrenceof wind pollination.Moreover, most species arecomponentsof the understoryof evergreen forest. On the other hand, there are not clear adaptationsto insectpollination,certainlynot to insect pollinationbasedon breeding.The commonoccurrence of pluricellulartrichomes on the perianthof pistillate flowers suggests thatthese trichomesmightplay a role in pollination, as substitutesfor pollen grains. These trichomes are apparentlynutritiousas they provide a substratefor fungi. According to some label data, the fruits (fruiting perianths)are eaten by birds. According to Kubitzki and Ziburski (1994), the fruits (fruitingperianths)of Sorocea duckei float due to buoyancyprovidedby air enclosed by the folded (large) cotyledon.

SYSTEMATICRELATIONSHIPS AND SUBDIVISIONOF THE GENUS

The genus Sorocea is in variousrespects quite distinct.It is the only genus amongthe non-anemophilous ones having predominantlyracemose inflorescences, which may have implications for pollination (see above). It has clearlydefined and ? separatestaminate flowers, in contrastto the genera that areprobablyits closest relatives, such as Clarisia and Batocarpus. Furthermore,the perianthat least in the fruitingstateis intoa lowerpartadnateto theovary clearlydifferentiated and an upperpartfree from the ovary. The genus has been divided into two subgeneraby Burgeret al. (1962: 468): Sorocea and Paraclarisia, the lattercomprising only S. duckei andS. sprucei. In subg. Sorocea,one can recognizea groupwith a spinulose-dentateleaf marginand/ora spinulose acumenof the lamina,comprisingS. bonplandii,S. guilleminiana, DISTRIBUTIONAND ECOLOGY andS. hilarii, as somewhatdistinctfrom the restof the The genus rangesfrom southernMexico to Argen- subgenus. These three species are morphologically tina. It is absent from the West Indies (except for veryclose andoftennot easy to distinguish.In the larger Cura9ao)andpoorlyrepresentedin the Guianas.Most group of species, the marginof the laminais entireor of the species are elements of lowland rainforest, be- varies from entireto dentate,but neitherthe teeth nor ing understorytreeletsor shrubsor canopytrees.Only the acumenof the laminaarespinulose. The species of Soroceatrophoidesis a (sub)montanespecies,although this group areclosely relatedand it is often difficult to it is also found at low elevations.Sorocea bonplandii, distinguish them. The exception is the clear-cut S. in the southernpartof the genus range,andS. sprucei, jaramilloi.

Key to the species of Sorocea 1. Lamina scabrous above and/or beneath. 2. Pistillate inflorescences with 1-3 flowers; anthers not apiculate; leaf base mostly (sub)acute; Amazonia .............................. 14. S. duckei 2. Pistillate inflorescences with 4-12 flowers; anthers apiculate; leaf base mostly rounded to 13. S. sprucei emarginate; Venezuela to Argentina.................................... 1. Lamina smooth above and beneath. 3. Leaf margin and/or acumen spinulose. 4. Stigmas (0.5-)1-2.2 mm long, deflexed; tepals of the staminate flower horizontally spreading to deflexed at anthesis; anthers suborbicular in outline, slightly smaller than the tepals; SE Brazil, Argentina, and Paraguay)........................... 2. S. bonplandii

TAXONOMIC TREATMENT

95

4. Stigmas 0.5-1 mm long, horizontally spreading; tepals of the staminate flower erect at anthesis; anthers ovate in outline, or if suborbicular, then distinctly smaller than the tepals. 5. Anthers in outline ovate (distinctly longer than broad); pistillate inflorescences with peduncles 0.5-16 cm long; lamina usually distinctly inequilateral; E Brazil ................. 3. S. hilarii 5. Anthers in outline suborbicular (nearly as long as broad); pistillate inflorescences with peduncles 0.5-1.5 cm long; lamina usually almost equilateral; E Brazil and Amazonia ..................................................................................................................................... .1. S. guilleminiana 3. Neither margin nor acumen spinulose. 6. Lamina subhirsute, hirtellous, or (sub)tomentose beneath. 7. Inflorescences subcapitate (or the staminate ones shortly spicate, to 1.5 cm long); bracts 13. S. sprucei membranaceous; Venezuela to Argentina............................................................ 7. Inflorescences racemose to spicate, the staminate ones at least 1.5 cm long; bracts subcoriaceous .................................................................. 4. S. pubivena 6. Lamina appressed- or patent-puberulous or glabrous beneath. 8. Inflorescences (sub)capitate (or very shortly spicate); bracts membranaceous; leaf scars prominent ............................................................ 13. S. sprucei 8. Inflorescences distinctly spicate or racemose; bracts subcoriaceous; leaf scars plane. 9. Lamina minutely puberulous on the midrib above. 10. Filaments longer than the perianth; pedicels of the pistillate flowers 1.5-2 cm long in fruit; Central America to Peru ...................................................... 8. S. trophoides 10. Filaments shorter than the perianth; pedicels of the pistillate flowers to 1.5 cm long in fruit. 11. Petiole longer than 1.5 cm. 12. Lateral veins 8-10 pairs; staminate inflorescences 16-33 cm long; fruiting perianth 2-2.5 cm long; W Ecuador.................................. 12. S. jaramilloi 12. Lateral veins (6-)10-20(-25) pairs; staminate inflorescences usually less than 15 cm long; fruiting perianth 1-1.7 cm long; Central America to Amazonia ..................................................... 4. S. pubivena 11. Petiole to 1.5 cm long. 13. Hairs on the midrib beneath of different lengths (the longer ones ? appressed, the shorter ones often patent); Amazonia .......................... 5. S. steinbachii 13. Hairs on the midrib beneath of similar length (patent or appressed) or lacking. 14. Margin of the lamina denticulate to dentate. 15. Staminate flowers pedicellate or narrowly sessile; fruiting perianth 0.6-0.8 cm diam.; Central America to Colombia (Choc6) ........ 9. S. affinis 15. Staminate flowers ? broadly sessile; fruiting perianth 0.8-1.7 cm diam.; Central America to Amazonia ..................................... 4. S. pubivena 14. Margin of the lamina entire. 16. Staminate flowers pedicellate or subsessile; fruiting perianth densely hairy, or if subglabrous, then its surface smooth and the perianth 0.6-0.8 cm diam.; Central America to Colombia 9. S. affinis (Choc6) ............................................ 16. Staminate flowers sessile; fruiting perianth subglabrous and its surface ruminate or the fruiting perianth 0.5-1.7 cm diam. 17. Petiole 0.3-0.8 cm long; staminate inflorescences 0.8-4.5 cm long; fruiting perianth 0.7-0.9 cm diam., its surface ruminate 11. S. ruminata and subglabrous; Panama......................................... 17. Petiole (0.5-)0.8-2(-2.5) cm long; staminate inflorescences 1.5-20 cm long; fruiting perianth 0.8-1.7 cm diam., its surface smooth, hairy or subglabrous; Central America to 4. S. pubivena Amazonia .............................................................. 9. Lamina completely glabrous above. 18. Filaments longer than the perianth; perianth of the pistillate flower subglabrous, in fruit ca. 1 cm diam. and pedicel to 2 cm long; midrib in the acumen of the lamina usually impressed; at (700-)900-2200 m; Central America to Peru ... 8. S. trophoides 18. Filaments shorter than or as long as the perianth; perianth of pistillate flower puberulous, at least with globose-capitate hairs at its lower part, or if subglabrous, then the fruiting perianth ellipsoid or at most 0.8 cm diam. and on pedicels to 1 cm long, or, if the perianth is similar to that of S. trophoides, then the midrib in acumen of the lamina plane; elevations to 900 m.

FLORANEOTROPICA

96

19. Laminadistinctlyinequilateral;staminateflowers pedicellate;anthersovate withpeduncles0.5-16 cm long, mostlypenin outline;pistillateinflorescences 3. S. hilarii dulous;E Brazil.................................................. 19. Laminausually slightly inequilateral;staminateflowerssessile, or if pedicellate, then the antherssuborbicularin outline;pistillateinflorescencesusually with pedunclesto 0.5 cm (in fruitto 1.5 cm) long. 20. Lateralveins usually 10-20 pairsand the laminamostly (sub)entire;leafy twigs 2.5-6 mm thick; petiole (I-)1.5-2.5 mm thick; fruitingperianth 1-1.7 cm diam.;Amazonia,Guyana,SE and N Venezuela,and N and SE Colombia ...............................................

4. S. pubivena

20. Lateralveins usually7-12 pairs and the laminadentateor denticulate, or if entire,then the leafy twigs 1-3.5 mm thick and the petiole 0.51.5 mm thick;fruitingperianthat most 1 cm diam. 21. Materialwith staminateflowers. 22. Flowerspedicellateor subsessile;CentralAmericato Colombia 9. S. affinis

(Choc6) .............................................

22. Flowers? broadlysessile. 23. Perianthof staminateflowers (sparsely)puberulousoutside. 24. Anthers apiculate. 25. Lamina(usually) with several to many parallel tertiaryveins; NW Ecuador.........................10. S. sarcocarpa 25. Laminawithoutor with a few paralleltertiaryveins; CentralAmericato Colombia(Choc6).....................9. S. affinis 24. Anthersnot apiculate;N Amazonia,E Venezuela,and Suriname .........................................

6. S. muriculata

23. Perianthof staminateflowersciliolate or glabrous. 26. Stamensincurved;N Amazonia, S Venezuela, and Suriname ..................................

S. muriculata

26. Stamensrecurvedor straightand the anthersperpendicular to the filament;AmazonianPeru,Brazil,and Bolivia.. 7. S. briquetii 21. Materialwith pistillateflowers. 27. Flowers (shortly)pedicellate. 28. Upperpartof the perianthdistinctlybroaderthanthe lowerpart. 29. Laminausually with severalto numerousparalleltertiary 10. S. sarcocarpa ................................... veins; NW Ecuador 29. Laminawithoutor with a few paralleltertiaryveins. 30. Lowerpart of the perianthdensely puberulous; CentralAmericato Colombia(Choc6).....................9. S. affinis 30. Lowerpart of perianthsparselypuberulousto muriculate; Amazonia, S Venezuela, and Suriname... 6. S. muriculata

28. Upperpartof the perianthnearlyas broadas the lower part; N Amazonia,S Venezuela,Suriname,and FrenchGuiana . 6 . S. muriculata .........................................................................................

27. Flowerssessile. 31. Pistillateinflorescenceswith 3-12(-20) flowers;perianthwith or withoutsparseglobose-capitate hairs;fruitingperianthsubglobose, or if oblongoid(to ellipsoid), then 1-1.2 x 0.5-0.6 cm; Amazonia,S Venezuela,and Suriname .................6. S. muriculata 3 1. Pistillateinflorescenceswith 10-40 flowers;perianth,at least the lower part,with dense globose-capitatehairs;fruitingperianth ellipsoid, 0.8-1.3 x 0.6-1 cm; AmazonianPeru, Brazil, and Bolivia ....................................... 7. S. briquetii 7A. Sorocea Saint-Hilairesubg. Sorocea. IncludingBalanostreblusKurzandPseudosorocea Baillon (see underthe genus). Treesor shrubs,evergreen;internodesusuallylong; scarsof the leaves plane;axillarybuds inconspicuous.

Inflorescencesdistinctlyracemoseor spicate;bractssubcoriaceousto coriaceous;stamenswithdistinctfilaments. This subgenuscomprises 12 species, occurringin Centraland SouthAmerica.

TAXONOMICTREATMENT

97

Fig. 17. Sorocea guilleminiana. 1. Leafy twig with infructescence.2. Leaf (Prance et al. 7989). 3. Leafy twig withpistillateinflorescence(Prance et al. 19247). 4. Leaf. 5. Leafytwig with pistillateinflorescences(Krukoff 8849). 6. Leafy twig with pistillate inflorescences(Berg et al. P19825). 7. Leaf (Prance et al. 19247). 8. Leaf. 9. Leafy twig with staminateinflorescence (Prance et al. 2344). 10. Leafy twig with staminateinflorescences (Berg et al. P18527). 11. Staminateflower and bract. 12. Stamen(Maguire et al. 56625). 13. Staminateflower. 14. Stamens(Prance et al. 2344). 15, 15a. Pistillate flower (Philcox et al. 3059). [By P. Pardoen,Utrecht.]

FLORANEOTROPICA

98 1. SoroceaguilieminianaGaudichaud, Voy.Bonite,Bot. Atlas t. 74. 1844;Trecul,Ann. Sci. Nat. Bot., Ser 3, 8: 145, t. 6, fig. 185-188. 1847;Miquel,in Martius, Fl. Bras.4(1): 213, t. 34. 1853; Burger,Lanjouw& WesselsBoer,Acta Bot. Neerl. 11: 462, t. 11. 1962; Marques,Carauta,Gon9alvesCosta& Sucre,Anais Acad. Brasil. Ci. 48(2): 287, t. 16, 17, 19. 1976. Type. Brazil. Rio de Janeiro:Mun. Rio de Janeiro, Corcovado,1839 or 1840 ($ fr), Guillemin131 (holotype, P; isotypes, F, G, U). Fig. 17 Sorocea houlletianaGaudichaud,Voy. Bonite, Bot. Atlas t. 74. 1844; Miquel, in Martius,Fl. Bras. 4(1): 214. 1853. Type. Brazil. Rio de Janeiro: Mun.Rio de Janeiro,(e), Gaudichaud1089 (holotype, P; isotypes,A, B, F, G, U, US). Trophishilariana Casaretto,Nov. Stirp.Bras. Dec. 80. 1845. Soroceahilariana(Casaretto)Bureau, in De Candolle,Prodr.17: 254. 1873. Syntypes. Brasil. Bahia: Sao Amaro,Casarettos.n. (TOn.v.);Rio de Janeiro,Restingade Taypu,Casaretto Mun.Riode Janeiro, s.n. (TO-n.v.);Riode Janeiro: Corcovado,Riedel s.n. (TO-n.v.). Sorocea klotzschianaBaillon, Adansonia1: 212, t. 6. 1860; Burger,Lanjouw& WesselsBoer,Acta Bot. Neerl. 11: 460, t. 10. 1962. Type. Brazil. Amazonas:Rio Negro, mouth of Rio Padauari, Dec 1984 (a' fl), Spruce 3794 (holotype, P; isotypes,BM, BR, G, GH, K, LE, MG, NY). Balanostreblus ilicifoliusKurz,J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist., 42: 248, t. 19. 1873. Type.Not traced,the plate couldbe designatedas the type. SoroceagrandifoliaS. Moore,Trans.Linn.Soc. London Bot., Ser.2, 4: 476. 1895.Type.Brazil.Mato Grosso:SantaCruz,1891-1892(d'& V fl), Moore 556 (holotype,BM; isotypes,B, K, NY, P). Sorocea castaneifolia Huber,Bol. Mus. Paraense Hist. Nat. 5: 333. 1909;Chodat& Vischer,Bull. Soc. Bot. Geneve, Ser. 2, 11: 230. 1920. Type. Brazil.Para:Obidos,11 Jan 1905 (v), DuckeMG 6961 (lectotype, Burger, Lanjouw & Wessels Boer,Acta Bot. Neerl. 11: 460. 1962, MG). SoroceamacrogynaLanjouw& WesselsBoer,Acta Bot.Neerl.11:459, t. 10. 1962.Type.Brazil.Amazonas:Mun.Humaita,TresCasas,14 Sep-i1 Oct 1934 (? fl-fr), Krukoff6143 (holotype,U; isotypes,A, B, BM,F,G, K, LE,MICH,MO,NY,US). Sorocea jureiana RomaniucNeto, Albertoa 4(8): 97, cumtab.1996.Type.Brazil.SaoPaulo:Peruibe, EstagaoEcol6gicada Jureia,(e'), Mamadeet al. 451 (holotype, SP-n.v.). Tree,to 20(-30) m tall,orshrub.Leafytwigs 1-4 mm thick, sparselypuberulous.Lamina oblong to elliptic or to lanceolate, 4-20(-32) x 2-8(-15) cm, usually broadestat the middle, almost equilateral,(sub)coriaceous;apexacuminate,theacumenspinulose;baseacute to obtuse (to subcordate);margin(usually) spinulose dent(icul)ate; upper surface glabrous; lower surface sparsely puberulous on the main veins or sometimes

hirtellous,the globose-capitate,pluricellularhairs mostlyabundant; venationnearlyplane,butthemidrib ? impressed above,prominent beneath; lateral veins1018pairs;tertiaryvenationmostlypartlyscalarifonn; petiole0.3-1.2(-1.5)cmlong,1-2.5mmthick,puberulous;stipules0.4-0.8 cm long, puberulousto glaStaminateinfloresbrous,caducousorsubpersistent. the0.1-1.2 1.5-8cmlong,including cencespendulous, cm long,puberulouspeduncle;flowersdistant,with 2-3 mmlong,4-parted pedicelsto4 mmlong;perianth to 4-fid,the lobesmembranaceous towardsthe margin, erectat anthesis,sparselypuberulous,ciliolate; orstraight, stamens(3-)4, free,atathesisbentoutwards filaments1butthentheanthersusually? transverse; 1.5mmlong;anthers(0.5-)0.8-1 x (0.5-)0.8-1 mm, broad. distinctlysmallerthanthetepals,theconnective Pistillateinflorescencespatent(topendulous?),2-12 cm

long;peduncle0.5-1.5 cmlong,to 2 cmlongin fruit, redin fruit;flowers5puberulous; rachispuberulous, 30(-ca.50),oftenreflexed,especiallyin fruitingstate; pedicellateorsometimessubsessile;pedicelto 0.5 cm long,to 1(-1.2)cmlongin fruit,minutelypuberulous to muriculate; perianth ca.2 mmlong,faintly4-lobed, theupperpartovoidtohemispherical, sparselyandminutelypuberulous, sparselyhispidulousoralmostglabrous,thelowerpartusuallyratherdenselypuberulous to hispidulousandusuallywithdense,globose-capitate,pluricellular hairs;stylesometimesexceedingthe to subulate,ca.0.5-1 perianth; stigmastongue-shaped mmlong,coarselypapillate. Fruitingperianth ellipsoid to globoseto ovoid,ca. 0.6-1 x 0.5-1 cm,red,finally black, apex rounded(to acute), surfacesmoothto muriculate orsometimesverrucate. Distribution(seeFig.76.4).IntheAmazonBasin, eastemBrazil,andcentralBrazil;in evergreen,wetor relativelydry,non-inundated forest;atlowelevations. Representativespecimensexamined.COLOMBIA. AMAZONAS: BetweenLeticia& Marco,20 Aug 1946 (? fl), Black et al. 46-41 (IAN); Puerto Nariiio, Parque NacionalAmacayacu,3 Aug 1989 (6'), Vasquezet al. 12539 (F, MO). CAQUETA: Tres Esquinas, Aug-Sep 1973 (st), Machecha1110 (UDBC). ECUADOR. SUCUMBiOS: ReservaFaunisticaCuyabeno,nr.LagunaGrande,12 Mar1990(? fl-fr), Valencia et al. 361 (QCA,QCNE). PERU. Cuzco: Prov.La Convencion,Rio Manguriari(= Manuyari),2 Feb 1991 (st), Ntuez et al. 12677 (BG). HUANUCO:Prov. Pachitea,2-8 km E of Puerto Inca, 12 sep 1982(st), Foster8716 (MO);Prov.Pachitea, ca. 24 km SSE to ca. 20 km SE of PuertoInca, 9 Jan 1988 (? fr), Wallnoefer11-090188 (BG). JUNIN: Prov. Chanchamayo,Hda. La Genoa,Finca Italia, 1140 m, 4 Mar1991(? fr), Gentryet al. 73320(BG);Prov.Sapito, Dtto. Pangoa,PuertoOcopa,5 Jun 1981 (st), ReynelR. 292 (MOL,U); Prov. Satipo, Dtto. Satipo, Zapallar,3 May 1982 (st), ReynelR. 478 (MOL,U); Prov.Tarma,

TAXONOMIC TREATMENT valley of Rio Chanchamayo, La Merced, 1000 m, (9 flfr), Weberbauer 1853 (G). MADRE DE DIos: Parque Nacional Manu, Rio Cumerali, 20 Oct 1986 (d'), Foster et al. 11942 (BG, MOL); Parque Nacional del Manu', Rio Manul,Cocha Cashu Station, 17 Oct 1979 (st), Gentry et al. 26819 (U); 26 km S of Puerto Maldonado, Rio Tambopata, 2 Nov 1979 (9 fl-fr), Hartshorn 2401 (BG, F, MO, MOL, NY, U); Rio Madre de Dios, Lago Sandobal, 10 km NE of Puerto Maldonado, Jun 1900 (st), Ntniez 12279 (MO); Prov. Tambopata, Tambopata Reserve, 18 Aug 1990 (si), Reynel R. et al. 5216 (BG, MO); Tambopata Wildlife Reserve, 30 km S of Puerto Maldonado, 1 Dec 1984 (? fl-fr), Younget al. 323 (BG, MO, NY). PASCO: Prov. Oxapampa, Dtto. Puerto Bermudez, Cahuapanas, 4 Nov 1980 (? fl-fr), Reynel R. 25 (MOL, U). UCAYALI:Prov. Coronel Portillo, Carretera "Federico Basadre," km 99, 2 Dec 1978 (9 fr), Diaz 673 (MO, U); jct. of Rio Yuyapichis & Rio Pachitea, SE of Pucallpa, 21 Oct 1985 (st), Morawetz et al. X3-211085 (BG, MO). BRAZIL. ACRE:Mun. Rio Branco, Reserva Florestal da Universidade Federal do Acre, 15 Oct 1980 (? fl), Ferreira et al. 2907 (MO, NY); Sao Luiz, Varadouro, 26 Oct 1923 (? fl), J. G. Kuhlmann RB 19828 (MO, U); nr. Tarauaca, 23 Sep 1968 (? fr), Prance et al. 7489 (INPA, NY, U, US). AMAZONAS:Mun. Manaus, Rio Taruma,23 Oct 1941 (e), Ducke 810 (F, IAN, MG, MO, NY, US); Rio Madeira, Calama, Nov-Dec 1931 (? fr), Krukoff 1304 (A, B, BM, G, K, MICH, MO, NY, P, U); Mun. Humaita, nr. Tres Casas, 14 Sep-lI Oct 1934 (? fl-fr), Krukoff 6223 (A, BM, F, G, K, LE, MICH, MO, NY, U, US); Rio Livramento, nr. Livramento, 12 Oct-6 Nov 1934 (? fl-fr), Krukoff 7039 (A, BM, F, G, K, LE, MICH, MO, NY, U, US); Sao Paulo de Oliven9a, 26 Oct11 Dec 1936 (? fl), Krukoff 8849 (A, B, BM, F, G, K, LE, MICH, MO, NY, P, U, US); Tefe (= Ega), (9 fl-fr), Poeppig 2683 (B, P); 2-3 km SW of Labrea, 28 Oct 1968 (Y fr), Prance et al. 7989 (F, GH, INPA, NY, P, U, US). BAHIA:Mun. Ilheus, Solobrinho, 18 Dec 1968 (? fl-fr), Almeida et al. 290 (U); between Ajuda & P6rto Seguro, 8 Jun 1962 (e'), Duarte 6635 (MO, NY, U, US); Itacare, nr. mouth of Rio de Contas, 28 Jan 1977 (a), Harley et al. 18355 (U); Mun. Una, rd. BA-265, 25 km from Una, 26 Dec 1976 (? fl-fr), Mori et al. 9309 (NY, U); Mun. Itabuna, 3 km NW of Jucari, 9 Mar 1978 (9 fl), Mori et al. 9587 (U); Vit6ria, (e), Sello 1185 (B, US). DISTRITOFEDERAL.C6rrego Quilombo, 6 Oct 1980 (e), Heringer et al. 5557 (MO); C6rrego do Gama, 21 Oct 1988 (a), Nogueira Silva 33 (MO); Reserva Ecol6gica do IBGE, 14 Sep 1984 (a), Pereira 1184 (BG, SP). ESPIRITOSANTO:Mun. Nova Venecia, Serra dos Aymores, 16 Nov 1953 (Q fr), Duarte 3779 (MO, RB, U, US); Linhares, Reserva Florestal da CVRD, 23 Nov 1989 ( fl), Farias 343 (BG); Mun. Concei9ao do Castelo, Venda Nova, 17 Oct 1985 (a), Hatschbach et al. 49883 (NY); Linhares, 29 Mar 1971 (9 fl-fr), Santos 1503 (U); Linhares, Reserva Florestal da CVRD, 31 Oct 1978 (a), I. A. Silva 26 (BG). GoIAs: Serra dos Pirineus, 60 km N of Corumba de Goias, 23 Jan 1968 (9 fr), Irwin et al. 19091 (LD, MO, NY, US); Serra do Caiapo, 13 km N of

99 Jatai, 21 Oct 1964 (a'), Prance et al. 59532 (B, MO, NY, U, US). MARANHAO:Maraca9ume, 10 Feb 1976 (st),

Ribeiroet al. 1294 (IAN);betweenEstreito& Maraba, 10 Feb 1976 (st), Ribeiro et al. 1296 (IAN). MATO GROSSO:Rio Aripuana, nr. Humboldt Centre, 14 Oct 1973 (c), Berg et al. P18527 (INPA, MO, NY, U), 22 Oct 1973 (? fl-fr), Berg et al. P19825 (INPA, MO, NY, U); Rio Machado, nr. Tapajara, Nov-Dec 1937 (Y fr), Krukoff 1349 (A, B, BM, G, K, MICH, MO, NY, P, U); Cuiaba, 6 Jan 1892 (? fr), Malme 1312 (B, G, K, NY, P); 1 km E of rd. Xavantina-Cachimbo, 15 Nov 1967 (? fl-fr), Philcox et al. 3059 (IAN, K, MO, NY, P); Serra de Roncador, 50 km N of Xavantina, Garapu airstrip, 30 Sep 1964 (a), Prance et al. 59186 (MICH, NY, U, US); Mun. Alta Floresta, rd. Alta Floresta-Rio Apiaca, 44.5 km E of Rio Apiaca, 29 Sep 1985 (e), Thomas et al. 4104 (BG, MO, NY, US); Mun. Luciara, 43 km S of P6rto Algere do Norte, 17 Oct 1985 (? fl), Thomas et al. 4479 (MO, NY, US). MINAS GERAIS: Mun. Muriae, rd. BR.116, 12 Dec 1984 (e'), Hatschbach et al. 48790 (BG, F); Mun. Itinutaba, Correio, 11 Nov 1954 (d' + ? fl), Macedo 3817 (BM, IAN, MO, RB, SP, US); Rio Branco, Retiro de Antonio Avelino, 14 Nov 1930 (o), Mexia 5303 (A, BM, F, G, GB, GH, K, MICH, MO, NY, U, US, Z); Vi9osa, rd. to Sao Miguel, 22 Oct 1930 (? fr), Mexia 5379 (A, BM, F, G, GB, GH, K, MICH, MO, NY, U, US, Z); Belo Horizonte, 22 Oct 1942 (? fl), Oliveira 1147 (IAN, US); Lagoa Santa, (? fl-fr), Warming 1949 (C). PARk: Obidos, 20 Nov 1907 (? fl), Ducke MG 8845 (BM, F, G, MG); Faro, 21 Jan 1920 (? fr),

DuckeRB 13042(B, K, MO,U, US); Serrado Cachimbo, 12 Dec 1956 (? fl), Pires et al. 6150 (IAN, MO, NY); Serra de Carajas, nr. AMZA headquarters, 17 Oct 1977 (d), A. Silva et al. 54 (MO, NY, RB, U); ParqueNacional de Tapajos, Ilha Grande, 28 Nov 1976 (? fr), M. G. Silva et al. 4000 (MG, U); Rio Jari, rd. Monte DouradoBraco, 9 Jan 1969 (? fl-fr), N. T Silva 1591 (IAN, NY, U, US). PARAIBA:Arei, Escuela de Agronomia do Nordeste, 28 Apr 1953 (e + ? fl-fr), Coelho de Moraes 693A (IAN, MO, US), 9 May 1953 (o), Coelho de Moraes 1073 (NY). PERNAMBUCO:Without locality, (? fl), Guillemin s.n. (F); Recife, Dois Irmaos, 5 May 1949 (a), Andrade-Lima 49-202 (IAN), 8 May 1949 (? fl), Andrade-Lima 49-204 (IAN); Gurjau, 19 Mar 1953 (o), Andrade-Lima 53-1227 (IAN); Recife, Dois Irmaos, 27 Feb 1962 (a'), Mattos et al. 9818 & 9819 (MICH, US); Tapera, 15 Mar 1934 (? fl-fr & e'), Pickel 3580 (GH, NY, US). RIO DE JANEIRO:Corcovado, 30 Sep 1921

(? fl-fr), Ducke et al. RB 114 (B, K, U, US); without locality, probably near the city of Rio de Janeiro, (? fl), Gaudichaud 1089 (P); Mun. Rio de Janeiro, Corcovado, 1878 (a), Miers 3910 (BM, K, P, U, US); Mun. Rio de Janeiro, Tijuca, Oct 1870 (? fl), Glaziou 4919 (B, C, K, P); Larangeiras, nr. Cantagalo, 19 Mar 1887 (? fl), Glaziou 16356 (B, G, K, LE, P); Mun. Cabo Frio, Arraial do Cabo, 2 Feb 1985 (a'), Plowman 13921 (NY); Mun. Rio de Janeiro, Feb 1838 (? fr), Saint-Hilaire 131 (P). ROND6NIA: Bananeiras, along Madeira-Mamore railway, 13 Sep 1968 (a), Maguire et al. 56625 (F, K, NY, U, US); Rio das Gar9as, 27 Nov 1949 (R fr), N. T Silva

100

386 (IAN, MO);MineragdoCampoNovo, 100 km S of Ariquemes,14 Oct 1979 (? fl-fr), Zarucchiet al. 2698 (MO, NY, U, US); Mun. POrtoVelho, Vila CanecoMineragao,Jacunda,22-27 Oct 1979 (? fl-fr), Vieiraet al. 322 (MG, MO). SAO PAULO:Conceiqao de Itahaim, (st), Loefgrenet al. 1669 (C). BOLIVIA. BENI: Prov.VacaDiez, nr.Alto Iv6n, 2 Nov 1983 (? fl-fr),Boom4024 (NY, U, US); Rio Beni, CachuelaEsperanza,19 Oct 1923 (e), Meyer87 (NY, U, Z); Prov. Vaca Diez, Tumi Chucua, 30 km S of Riberalta,29 Sep 1981 (ei), Solomon6516 (LPB, MO, NY, U). LA PAZ: Prov.Abel Iturralde,Alto Madidi,25 May 1990 (st), Gentryet al. 70550 (LPB, MO); Prov. Inquisivi,mouth Rio Aguilani, trail to Khora,19 Nov 1991 (e), M. Lewis40611 (BG),mouthof Rio Aguilani, 26 Nov 1991 (? fl-fr),M. Lewis 40685A(BG). PANDO: Prov.Manupiri,rd. PuertoRico-Rio Madrede Dios, km 33, 10 Oct 1991 (? fl), Beck et al. 20300 (BG, LPB); Prov.Abuna,Rio Orton,9 Jul 1992 (st), L. Vargaset al. 786 (USZ). SANTACRUZ: Prov. Velasco, Serraniade Huanchaca,ParqueNacional Noel KempffM., 6 Nov 1991(e), Foster13719 (LPB,USZ);Cuestade Suspiros, Mar 1911 (V fr), Herzog 1630b (G); Prov. Nuflo de Chavez, 14 km NW of San Javier,30 Nov 1990 (? flfr),Nee 40107 (BG, LPB);Prov.Ichilo,ParqueNacional Ambor6, Rio Macuinucu,6 Dec 1991 (9 fl-fr), Nee 41974 (BG, LPB). This widespread species is rathervariable in the dimensions, shape, margin,texture,and indumentum of the leaves, and in the shape and dimensions of the fruitingperianth.Specimens with the lower leaf surface hirtellous,subhirsute,or hispidulousarefound in Rio de Janeiro,MinasGerais,andin southernAmazonia In southernAmazonia, leaves with such indumentum areoften relatively large and with the lateralveins impressed above. Except for the spinulose leaf margin, these specimensremindone of large-leavedspecimens of Soroceahirtellasubsp.hirtella.Inbothspecies,large leaves are correlatedwith relatively large fruits. The materialfromPernambucoandParaibahas rathersmall leaves, often with sharplyspinulosemargins;they approachS. hilarii in its leaf characters. Whensterile,SoroceaguilleminianaandS. bonplandii cannotbe distinguished;only if globose-capitatepluricellulartrichomesareabundantlypresenton the lower leaf surface can one be rathersure that the specimen belongsto S. guilleminiana.SterilespecimensofS. hilarii can be distinguishedfromthose of S. guilleminiana by the distinctlyinequilaterallaminaandthenon-spinulose (or very shortlyspinulose)marginandacumenof the lamina. Material of Sorocea guilleminiana and Clarisia ilicifolia is often confused. The latterspecies can be readily recognized by the midribof the laminabeing prominentabove, andby the presenceof uncinatehairs on various parts.

FLORA NEOTROPICA

Theperianthof the pistillateflower is often covered by a white mycelium, at anthesisthe lower partand in fruitthe greaterpart. Local names. Peru: Junin: yanchama; Pasco: yanchama.Brazil: Bahia: bainha de espada, trempa; EspiritoSanto:bainhade espada;MinasGerais:paude serra; Para: lingua de onca. Bolivia: Beni: xoqueshequere(Chacobo). 2. Sorocea bonplandil (Baillon)W.Burger,Lanjouw& WesselsBoer,Acta Bot. Neerl. I1: 465, t. 12, 1962; Marques,Carauta,GoncalvesCosta& Sucre,Anais Acad. Brasil. Ci. 48(2): 289, t. 3-6. 1976. PseudosoroceabonplandiiBaillon,Adansonia 11:296. 1875. Type.Argentina.Corrientes:withoutlocality,(? flfr), Bonpland s.n. (holotype, P). Fig. 18 Sorocea ilicifolia Miquel forma laxiflora Hassler, AnnuaireConserv.Jard. Bot. Geneve 21: 121. 1919;Chodat& Vischer,Bull. Soc. Bot. Geneve, S&. 2, 11: 257. 1920. Type. Paraguay. Nr. Sapucay,Aug 1913 (d), Hassler 12205 (holotype, G; isotype, GH). SoroceasylvicolaChodat,in Chodat& Vischer,Bull. Soc. Bot. Geneve, Ser. 2, 11: 230, 256, t. 239 and 240. 1920. Type. Paraguay.Yagauro,Jun 1877 (? fl-fr), Balansa 1971 (lectotype, designated here, G). SoroceasylvicolaChodatvar.caaguazuensisChodat, in Chodat & Vischer, Bull. Soc. Bot. Geneve, Ser. 2, 11: 257. 1920. Type. Paraguay.Rio Yhu, Caagauzu,Sep 1905 (? fl), Hassler 9413a (holotype,G; isotypes,A, B, BM, G, K, MO,NY,P). Shrubor tree,to 10m tall.Leafytwigs 1-4mm thick, puberulous.Laminaoblong to lanceolateor to elliptic, 4-12(-24) x 1-4.5(-6) cm, usually broadest at the middle,equilateralto slightly inequilateral,coriaceous; apex acuminate,the acumenspinulose;base (acuteto) obtuse to rounded;margin spinulose-dentate;upper surface glabrous; lower surface sparsely puberulous on the main veins, the globose-capitate, pluricellular hairs(very) sparse;venation almost plane or the midrib slightly impressedabove, ? prominentbeneath;lateralveins 8-4(-18) pairs;tertiaryvenationreticulateor partly scalariform;petiole 0.3-1 cm long, 1-2.5 mm thick,puberulous;stipules 0.3-1 cm long, puberulous to glabrous,caducous or subpersistent.Staminateinflorescences patentto pendulous, 0.5-2 cm long; peduncle0.1-0.3 cm long, puberulous;rachispuberulous; flowers more or less crowded;pedicel to 1.5(-3) mm long;tepalsalmostfree,ca. 1.2mm long, entirelymembranaceous,minutelypuberulous,ciliolate, spreading horizontallyto reflexedat anthesis;stamens4, (almost) free,bentoutwardsatanthesis;filaments0.8-1 mm long; anthersca. 1 x 0.8-1 mm, slightlysmallerthanthetepals,

TAXONOMICTREATMENT

101

Fig. 18. Sorocea bonplandii.1. Leafy twig with staminateinflorescences(Spegazzini15454). 2. Leafy twig with staminateinflorescence(Schwan 4513). 3. Leafy twig with pistillateinflorescence(Pedersen472). 4. Leafy twig with young infructescences(Renvoize3241). 5. Leafytwig with infructescence(Renvoize3184). 6. Stamens andtepals.7. Bractof staminateinflorescence(Fiebrig951). 8, 8a. Pistillateflowerand bract.9. Stigmas(Glaziou 7851). 10. Fruitingperianth(Klein s.n.). [By P. Pardoen,Utrecht.]

102

the connective broad.Pistillate inflorescencespatent (pendulous?in fruit),1-5 cm long, to 6 cm long in fruit; peduncle0.5-2 cm long, to 3 cm long in fruit,puberulous; rachispuberulous,red in fruit;flowers 3-15 (often reflexedin fruitingstate);subsessileorpedicellate; pedicel to 0.3 cm long, puberulous, to 1.8 cm long, camose, red and smooth or ? muriculatein fruit;perianthca. 2 mm long, 4-lobed,puberulous,theupperpart conicalto narrowlyovoid;styleoftenexceedingtheperianth;stigmasnarrowlytongue-shaped(0.5-)1-2.2 mm long, reflexed, finely papillate.Fruitingperianth globose to ovoid to ellipsoid, 0.6-1.2 x 0.6-0.9 cm, apiculate,red,finallyblack,the surfacesmooth,muric(ul)ate, verruculate,or ? wrinkled,minutely puberulous. Distribution (see Fig. 76.3). In southem Brazil, Paraguay,and northemArgentina;in moist forest, often gallery forest; to 1200 m. Representative specimens examined. BRAZIL. MINAs GERAIS:Bom Jardim de Minas, Serra da Mira, 29 Jan 1977 (? fr), Carauta2292 (RB); Caxambui,20 Sep 1956 (d), Heringer5349 (RB);Mun.Cristina,rd. to Casmo de Minas,22 Oct 1989(? fl), Mello-Silva et al. 80

(SPF);Mun.Tiradentes, Serrade Sao Jose, 1200m, 3 Oct 1987 (e), Peron351 (RB);Uberlandia, Parquede Sabia, 27 Oct 1992(2 fl), Alveset al. 12 (SP);withoutlocality, 1845 (e), Widgren s.n. (K, LE, 0). PARANA:Parque

Nacionalde Igua9i,Rio Sao Joao, 14 May 1959(? fl-fr), Duarte et al. 1780 (RB); Serra do Mar, 4 Nov 1915 (?), Dusen17283 (F, K, MICH,MO,NY,P); Mun.Piraquara, Campininha, 9 Oct 1944 (e), Hatschbach 1521 (US); Mun.Aropoti,Fazendade Tigre,10 Sep 1960(o'), Hatschbach 7242 (US), (? fl), Hatschbach 7243 (US); Mun. PontaGrossa,Rod. do Cafe-Itaiacoca,27 Sep 1964 (e'), Hatschbach 11601 (F); Mun. Capina Grandedo Sul, Sitio do Belizario, 17 Aug 1966 (6), Hatschbach 14629

(P,US);Mun.Sao Josedos Pinhaes,Purgatorio, 1982(a'), Hatschbach45300 (F, GB, MBM,WIS);Tamanadre,24 Sep 1914 (? fl), Jonsson 969a (F, GH, K, MICH,MO, NY,P). RIODEJANEIRO:(ParqueNacional da) Serra dos Orgaos, 10 Oct 1942 ($ fl), Barros 1034 (RB, U, US), ca. 1000 m, Mar 1891 (? fl), Gardner 5859 (BM, K); Mun.Petr6polis,Itamaraty, PontaLuzia,13 Sep 1874(Q

FLORA NEOTROPICA 824 (B, NY, US); Ibarama,20 Sep 1956 (a'), Reitz et al. 3733 (B, NY, US); Piloes, Palhoca, 28 Sep 1956 (a), Reitz et al. 3798 (B, G, GH, K, NY, U, US, WIS); Sao Fransisco do Sul, Tres Barras, Garuva, 5 Oct 1957 (a'), Reitz et al. 5036 (B, US); Serrado Matador, 29 Dec 1958 (? fl-fr), Reitz 6088 (B, F, G, K, Z); Blumenau, Morro Spitzkopf, 21 Aug 1959 (? fl), Reitz et al. 8977 (NY, US, SP). SiO PAULO:Serra de Canteira, (a'), Andrade RB 6519 (U); rd. Jacapiranga-Cananeia, km 3-5 9 Sep

1976 (? fl), Davis et al. 60827 (SP); Mun. Botucatu, Campus of UNESP, 13 Nov 1978 (2 fl-fr), Filho 17 (F); Mun. Teodoro Sampaio, Reserva Florestal Morro do Diabo, 28 Oct 1986 (? fl-fr), Pastora et al. 157 (MBM, RB); Sao Paulo, 21 Aug 1941 (a), Pickel 5357 (US), Mun. Vinhedo, nr. Esta9ao de Tratamento de Agua de Valinhos, 23 Sep 1977 (a), Taroda et al. 6620 (F, RB). PARAGUAY. Depto. Itapua, El Tirol, 11 Oct 1984 (Q fl-fr), Brunner 867 (MO);Depto. Paraguari, Parque Nacional Ybycui, 13 Sep 1980 (? fl), Fernandez C. et al. 3576 (NY); Depto. Alto Parana,29 km from Puerto Presidente Stroessner, 9 Dec 1982 (a), Fernandez C. et al. 7361 (MO, NY); upper Rio Parana, 1909-1910 (st), Fiebrig 5386 (GH), 5386a (B, G, US), 6404 (B, GH); Depto. Paraguari,Ybycui National Park, Arroyo Minas, 1 Oct 1985 (a), Gentry et al. 51851A (BG, MO), (? flfr), Gentry et al. 51858 (BG, MO); Depto. Itapua, Pirap6, 4 Aug 1984 (a'), Hahn et al. 2731 (MO, RB); Rio Yhu, Sep 1905 (a'), Hassler 9413 (A, B, BM, G, K, NY, P); nr. Sapucay, Jul 1913 (a), Hassler 11883 (B, BM, C, F, G, GH, K, MICH, MO, NY, US, Z), (? fr), Hassler 11883a (B, BM, C, F, G, GH, K, MICH, MO, NY, US, Z); Pirapo, Colonia Elisa, 5 Jun 1893 (? + a), Lindman A.1719 (B, Z); Depto. Guaira, Tororo, Cerro Polilla, 13 Nov 1988 (9 fr), Ortiz 820 (MO); Santiago, Estancia La Soledad, 19 Nov 1956 (? fl-fr), Pedersen 4353 (A, C, US); Depto. Amambay, Cerro Cora, 15 Aug 1980 (? fl-fr), Schinini et al. 20158 (MO); Colonia Primavera, 5 Aug 1956 (a'), Woolston705 (C, NY,SP,U); Depto. Caazapa, Dtto. Yuty, Tres de Mayo, 10 Sep 1987 (? fl), Zardini et al. 2982 (MO); Depto. San Pedro, Colonia 8, 12 km SE of Chore, 30 Sep 1987 (? fl-fr),Zardini et al. 3156 (MO); Depto. Guaira, Cord. de Ybytyruzui, Cerro Per6, 13 Jan 1989 (a'), Zardini et al. 9750 (BG).

ARGENTINA.CORRIENTES: Depto. San Tome,23

km NW of Gobernador Varasoro, 2 Dec 1970 (? fl-fr), fl + 6), Glaziou 7851 (B, C, F, G, LE, P, US), 7 Jul 1879 Krapovickas et al. 16884 (F,P,WIS); Depto. Mburucuya, Estancia Santa Teresa, 13 Nov 1949 (? fl), Pedersen (c'), Glaziou 11565 (B, C, F, K, LE, NY, P); Itataia, Maromba, 25 Oct 1928 (6), Porto 1836 (B, RB), 23 Oct 3978 (A, C, G, P, US), 21 Aug 1954 (a'), Pedersen 2780 (A, C, G, P, US, WIS); Depto. General Paz, Fernandez 1931 (Q fl), Porto 1918 (= RB 13047) (B, MO), Porto 2100 (=RB 25928) (B, RB, U); Serrados Orgaos,Cas- Guazu, 11 Sep 1956 (9 fl), Pedersen 3978 (A, C, P, US). cata do Imbui, Oct 1952 (9 fl), Rizzini RB 86692 = MISIONES:Depto. San Ignacio, Acaragua, 31 Aug 1946 Markgraf 10063 (MO, RB, U). RIO GRANDEDO SUL: (a'), Bertoni 2885 (F, MO, WIS); Depto. Candalaria, Loreto, 4 Aug 1931 (? fl), Gruener 273 (F, GH, NY); Sao Leopoldo, Sep 1941 (d), Leite 415 (A), (? fl-fr), Leite 416 (A, SP); nr.P6rtoAlegre, 10 Oct 1945 (R fr), Salto Iguazu, Dec 1910 (? fl-fr), Lillo 7277 (A, G, U, US); Depto. Monte Carlo, Puerto Piray, 19 Oct 1978 (? Rambo 27131 (MO, NY), 3 Oct 1949 (e?), Rambo 43725 (C, K, US), 2 Sep 1950 (? fl), Rambo48800 (MO);Sao fr), Renvoize 3184 (MO,U); Depto. Iguazui,Salto Iguazu, Leopoldo, Oct 1942 (?), Reitz 524 (B, NY); Cerro 21 Aug 1910 (? fl-fr & a), Rodriguez 366 (A, P, U, US); Depto. Candelaria, Loreto, 20 Apr 1947 (a), Schwartz Botucarai, Candalaria, Sep 1986 (ci), Sobral et al. 5169 (MBM, SP), (? fl), Sobral et al. 5173 (MBM,SP).SANTA 4513 (MO, U). Local names. Brazil:Parana:bainhade espada;Rio CATARINA: Mata da Limeira, 1 Sep 1950 (? fl-fr & d), Klein21 (US); Itajai,Cunhas,29 Oct 1954 ($ fr), Klein de Janeiro:fZolha de serra;Rio Grandedo Sul: cincho,

TAXONOMICTREATMENT soroca, sorocaba;Parana:cancorosa,serraleiro,suchu; SantaCatarina: carapicica, carapicicadef-ohlamitida, cego olho; Sao Paulo: caxim-mirim, ganchincho-canxim, soroco; Paraguay:niandyra, niandypami.Argentina: Cornentes:mariamolle; Misiones:nandpa,naudipa.

3. Sorocea hilarii Gaudichaud,Voy.Bonite, Bot. Atlas t. 71. 1844; Trecul,Ann. Sci. Nat. Bot., Ser. 3, 8: 145, t. 6, 1847; Burger,Lanjouw& Wessels Boer, Acta Bot. Neerl. 11: 457, t. 9. 1962; Marques, Carauta,Gon9alves Costa & Sucre, Anais Acad. Brasil. 48(2): 291, t. 1, 14, 16, 18. 1976. Sorocea ilicifoliaMiquelvar.hilarii(Gaudichaud)Miquel,in Martius,Fl. Bras.4(1): 213. 1853.Type.Brazil.Rio de Janeiro,1834 (? fl-fr) Gaudichaud1088 (holotype, P; isotypes, B, F, G, U). Fig. 19 SorocearacemosaGaudichaud, Voy.Bonite,Bot.Atlas t. 72. 1844;Trecul,Ann. Sci. Nat. Bot., Ser.3, 8: 145.t. 6. 1847;Miquel,in Martius,Fl. Bras.4(1): 213. 1853;Burger,Lanjouw& WesselsBoer,Acta Bot. Neerl. 11: 455, t. 8 & 9. 1962; Marques, Carauta,GonsalvesCosta & Sucre,AnaisAcad. Brasil. Ci. 48(2): 291, t. 2, 7, 13. 1976. Type. Brazil. Rio de Janeiro, (? fl-fr), Gaudichaud 1090bis(holotype,P; isotypes,F, S, US). SoroceamacrophyllaGaudichaud,Voy.Bonite,Bot. Atlas t. 73. 1844. Type. Brasil. Rio de Janeiro, (e), Gaudichaud1090 (holotype, P). Sorocea uriamenMiquel,in Martius,Fl. Bras.4(1): 113. 1853. Type.Brazil.Bahia:withoutlocality, (Y fl), Martius1818 (holotype,M, photographs ex M in F, GH, NY; isotype,B). Shrubor tree,to 5 m, occasionallyto 10m tall.Leafy twigs 1-4 mm thick, sparselyto ratherdenselypuberulous or glabrous;lenticels mostly ratherconspicuous. Laminaoblong to elliptic (or to lanceolate),2-12(-27) x 1-5(-IO) cm, mostly broadestabove the middle, ? inequilateral,coriaceous;apex acuminate,the acumen not or very shortly spinulose; base obtuse to rounded (to subcordate);margin in the upper part(very short spinulose-)dentateto subentire;uppersurfaceglabrous; lower surface sparsely puberulouson the main veins to glabrous,globose-capitate,pluricellularhairsrather abundant;venationalmostplane or the midribslightly impressedabove, ? prominentbeneath;lateralveins 712(-18) pairs,tertiaryvenationmostly reticulate;petiole 0.2-0.8(-1.2) cm long, 0.5-1.5(-2.5) mm thick, puberulous;stipules 0.2-0.8 cm long, puberulousto glabrous, caducous or subpersistent.Staminateinflorescences 2-20 cm long, includingthe 0.2-7 cm long, sparsely puberulous peduncle; flowers distant, with pedicels to 4 mm long, sometimes (sub)sessile; perianth4-fid to 4-parted,2-3 mm long, only the margins of the lobes membranaceous,sparselypuberulousand ciliolate, or to subglabrous,erect at anthesis;stamens

103 4, straightat anthesis; filaments 0.8-1 mm long; anthers1.2-1.5 x 0.8-1 mm, distinctlylongerthanbroad, the connectiverathernarrow.Pistillate inflorescences pendulous(to patent?),2.5-25(-45) cm long;peduncle 0.5-16 cm long, (sparsely)puberulous;rachispuberulous, ? camose, red to purplein fruit;flowers (3-)l 0ca. 35, subsessileto pedicellate;pedicel to 0.2 cm long, sparselypuberulousto hispidulous, to 1.5 cm long, ? swollen and red in fruit; perianthca. 2 mm long, 4lobed,theupperpart(broadly)ovoid, sparsely,minutely puberulous,smooth,muriculateorverruculate,thelower part mostly densely, minutely puberulous; stigmas tongue-shaped,ca. 0.5 mm long, often dentate,finely papillate. Fruiting perianth ovoid to ellipsoid (to oblongoid), the apex rounded (to truncate)or obtuse (to subapiculate),the surfacesmooth,sparselyto puberulous to almostglabrous. Distribution (see Fig. 76.1). In eastemBrazil,from Bahia to Sao Paulo;mostly in coastal vegetation, forest, or restinga;at low elevations. Representativespecimensexamined.BRAZIL.BAMun.Itamaraju, rd. Itamarajui-Eunapolis,km 5, 3 Nov 1983 (e), Carvalo et al. 2034 (BG, NY); Rio Gongogy, 4 Oct-30 Nov 1915 (? fl-fr), Curran248 (US); HIA:

Mun. Itacare, nr. mouth of Rio de Contas, 31 Mar 1974 (d), Harley 17533 (CEPEC);nr. Almada,(d), Martius 2065 (F, GH, NY); Mun. Mara6,rd. Ubaitaba-Itacare, ca. km 15, 5 Feb 1979 (R fl-fr), Moriet al. 11344 (NY); Ilheus, 11 Mar 1969 (c), Santos 386 (U). ESPiRITO SANTO: Mun. Itagua9u,Jatiboca,27 May 1946 (Y fr), Brade 18441 (U); Mun. Rio Bananal, rd. Rio BananalNova Brasil, km 10, 21 Apr 1995 (? fl-fr), Kallunkiet al. 708 (SPF); Guarapari,rd. BR.101-Praia Setibana, km 6, 23 Feb 1988 (? fl), Piraniet a. 2423 (BG);SantaTereza, 4 Feb 1985 (a), Peixotoet al. 3436 (BG, MO);without locality, (Y fr), Saint-Hilaire92 (cat. B2-no. 278: P, U, US), (e'), Saint-Hilaire94 (cat. B2-no. 403: P). RIO DE JANEIRO: Corcovado, 17 Aug 1973 (e), Berg 211 (MO, U), (? fl-fr), Duarte793 (MO); JurugubaBay, 1841 (d),

Gardner5629 (BM, K); Mun.Rio de Janeiro,Gavea,4 Mar 1861 (? fl-fr), Glaziou73 (BR, P); Itaipu, 27 May 1969 (? fl-fr), Plowman 2831 (= Sucre 5131) (GH); nr. Rio de Janeiro, 1836 (? fl), Riedel s.n. (LE); Reserva de Cabo Frio, 1 Nov 1966 (? fl-fr), Sucre 1153 (U); Bahia de Sepetiba, Ilha Furtada,24 Dec 1967 (? fl), Sucre 2005 (MO, U); Itapuca, 1 Jul 1968 (Qfl), Sucre et al. 3399 = Bragaet al. 1001 (F, MO); Restinga do Grumari, 14 Aug 1968(V),Sucre3518 (MO).SAOPAULO: Mun. Ubatuba, Pontal da Lagoinha, rd. Sao Paulo-Unatuba, km 237, 31 Aug 1980 (? fl-fr), Forero7673 (SP); Piassaguera, 14 Mar 1923 (), GehrtSP 8238 (US).

Sorocea racemosa and S. hilarii, as recognizedby Burgeret al. (1962), areunitedin the presenttreatment becauseimportantdifferentiating characterscouldnotbe of (coastal) found.Thespeciesoccursin theundergrowth forestandin moreopen littoralvegetation(restinga).In the formerhabitat,the leaves arerelativelylargeandof-

FLORA NEOTROPICA

104

,3~

~

!!j

?

!I

f'I

4

Fig. 19. Sorocea hilarii. 1. Leafy twig with staminate inflorescences (Santos 386). 2. Leafy twig with staminate inflorescences (Berg 211). 3. Leafy twig with young infructescences (Berg 210). 4. Leafy twig with young infructescences (Sucre 2005). 5. Twig with infructescence (Pinheiro 2031). 6. Staminate flower. 7. Stamens (Nadeaud s.n.). 8. Staminate flower. 9, 9a. Stamens (N. T Silva 58429). 10a, b. Pistillate flower. 10c. Stigmas (Moraes 693A). 11. Fruiting perianth (Mori et al. 12095). [By P. Pardoen, Utrecht.]

TAXONOMIC TREATMENT

105

tenentire,andtheinflorescences arependulous andlax andhavelongpeduncles. Intheothertypeof habitat, the leavesaresmallerandoftendistinctlydentate,andthe infloresences areoftenshorter, morecompact, andoften patentwithshortpeduncles.Whether thesedifferences aremerelyvariations orarebasedoninfraspecific morphologicalandecologicaldifferentiation couldnotbe established onthebasisof thepresentmaterial anddata. ThecollectionKallunkiet al. 708 hasextremelylong pistillate inflorescences, upto45 cmlong,withtheflowersupto 5 cmfromeachother. Thespecieshasbeenin cultivation in theBotanical Garden of St.Petersburg sincethemiddleofthe19thcentury.Staminate material hasbeendistributed underthe names: SoroceagracilifloraRegel&K6rnickeorS. hilarii var.gracilifloraRegel&K6rnicke. Thisnameis alsoused forpistillate material collectedbyRiedelin 1836.

ciliolate; stamens (3-)4, bent inwards or outwards, straightor bentinwardsat anthesis,mostly isomorphic; filaments0.2-1.5 mm long; anthersca. 0.5-0.8 x 0.50.8 mm, the connectivebroad,sometimesshbrt-apiculate.Pistillateinflorescencespatent(to pendulous?),26.5 cm long, to 9.5 cm long in fruit; peduncle 0.1-0.5(-l) cm long, to 1 cm long in fruit,puberulous; rachis puberulous, red(dish) in fruit; flowers 3-60 (-100), sessile, sometimeswithpedicelsto 0.3 cm long, subsessile or mostly pedicellate in fruit;pedicel to 1.5 (-2) cm long, tuming red, sparsely puberulous;perianth 2-4 mm long, 4-lobed (to subentire),the upper partnarrowlyto broadly ovoid, glabrous or sparsely puberulous,the lower partdistinctor hardlyso, mostly densely andminutelypuberulous;style sometimesexceeding the perianth;stigmas tongue-shaped,0. 5-1.5 mm long, rathercoarsely to minutely papillate.Fruiting perianth subglobose to ovoid to oblongoid or to Localnames.Brazil: Bahia: deleite,uriamen. jaquitia oblate, 1-1.7 x 0.8-1.7 cm, mostly apiculate,black,the surfacesmooth,sparselyto denselyandminutelypuberulous (to hirtellous) or subglabrous. 4. SoroceapubivenaHemsley,Biol.Centr.Amer.Bot. 3: 150. 1883;Burger,Ann. MissouriBot. Gard. In contrastto therelatedspeciesSoroceasteinbachii 47: 122. 1960;Burger,Lanjouw& WesselsBoer, andS. muriculata,this species shows quite a variation Acta Bot. Neerl. 11: 449, t. 6. 1962; Burger, in the androecium. FieldianaBot.40: 209. 1977.Type."Guatemala," Threesubspecies can be recognized. Friedrichsthals.n. (holotype,K). Figs.20, 21 Key to the subspecies of Sorocea pubivena Shrubortree,to 15(-25) mtall.Leafytwigs(1-)2.56 mmthick,whiteto brownish-subhirsute tohirtellous 1. Laminawith appressedhairson the veins beneathor subglabrous;northernSouth or patentto appressed-puberulous to subglabrous. . 4b. subsp. oligotricha Americato E Panama Lamina oblongto ellipticorto lanceolate,(6.5-)121. Laminawith patenthairs beneath. 30(-50) x (2.5-)4-13(-16) cm, broadestat or above 2. Pistillateinflorescenceswith ca. 15-60 themiddle,equilateral to slightlyinequilateral, coria(-100) flowers;CentralAmerica........ ceousto chartaceous; apexacuminate; baseacuteto .4a. subsp.pubivena ............................................ obtuseto rounded;margin(sub)entireor dentateto 2. Pistillateinflorescenceswith 3-15(-25) onthe denticulate; uppersurfaceminutelypuberulous flowers; upper Amazon Basin ........... .4c. subsp.hirtella ................................................ midribor glabrous;lowersurfaceratherdenselyto

sparselyhirtellous(tosubhirsute), denselyto sparsely patent-or sparselyappressed-puberulous (withhairs 4a. Sorocea pubivena Hemsley subsp. pubivena. of about the same length) on the main veins to Fig. 20 venationalmostplane,butmostlythe subglabrous; Bot. Gaz. Donnell Smith, Trophis macrostachya midribandoften also the lateralveins i impressed 10. 1905. Type.CostaRica. 40: (Crawfordsville) above,? prominentorthe smallerveinsalmostplane Nov 1898 Cartago:Rio de las Vueltas,Tucurrique, lateralveins(6-)10-20(-25)pairs,sometimes beneath; (V fl-fr), Tonduz 8124 (lectotype, Burger, connectedby a slightlyarcuatesubmarginal vein;terLanjouw& Wessels Boer, Acta Bot. Neerl. 11: tiaryvenation(usually) partlyscalariform; petiole(0.5-) 449. 1962, US). 0.8-2(-2.5) cm long, (l-)l.5-2.5(-3) mm thick,puber-

Clarisiamollis Standley,Ann. MissouriBot. Gard.

ulousto hirtellousorsubglabrous; stipules0.3-1.2 cm long, puberulousto hirtellous,caducous.Staminate (topatent?),1.5-20cmlong, inflorescencespendulous includingthe0.1-0.5 cm long,puberulous peduncle; flowersspacedto rathercrowded,broadlyto rather narrowlysessile;perianth1-2.5 mmlong,deeply4or only partedorthetepalsfree,sparselypuberulous

30: 85. 1943. Type. Panama. Bocas del Toro: Nr. Chiriqui Lagoon, 8 Oct 1940 (e), Wedel1090 (holotype, F; isotype, GH). Sorocea cufodontisii W. Burger, Acta Bot. Neerl. 11: 447. 1962, as S. cufodontii;Burger, Fieldiana Bot. 40: 208. 1977. Type. Costa Rica. Puntarenas: PuertoJimenez, 4 Apr 1930 (d), Cufodontis200 (holotype, F).

106

FLORANEOTROPICA

;'.

-.

El.:.

.

Fig. 20. Sorocea pubivena subsp. pubivena. 1. Leafy twig with staminate inflorescences (Holm et al. 389). 2. Leafy twig (McDaniel 5098). 3. Twig with infructescences (Burger et al. 10488). 4. Young infructescence. 5. Leaf (Burger et al. 10049). 6. Staminate flower. 7. Stamens (Holm et al. 389). 8, 8a. Pistillate flower (Lent 1416). 9, 9a. Fruiting perianth (Burger et al. 4290). [By P. Pardoen, Utrecht.]

TAXONOMIC TREATMENT

Leafy twigs puberulousto hirtellous.Lamina subcoriaceousto chartaceous;marginentireor (especially in specimenswith the laminahirtellousbeneath)? distinctly dentateto denticulate;upper surfaceminutely puberulouson the midrib;lower surfacehirtellousor densely to sparsely minutely patent-puberulous.Pistillate inflorescencesto 20 cm long in fruit;flowers ca. 15-60(-l 00). Fruitingperianthellipsoidto (sub)ovoid to subglobose, 1-1.4 x 0.8-1.2 cm, rathersparselyto densely puberulous.

107

1892 (d), Tonduz6751 (BR, CR, G, US). SAN JOSE: Tarrazu,11 Jul 1987 (? fr), G6mez-Laurito11601 (F, MO); ca. 25 km NE of Bijagualde Turrubares,Carara Reserve,26 Jun 1985 (? fl-fr),Grayumet al. 5485 (BG, MO);1.5 km E of SantaRosade Puriscal,14 May 1987 (? fl-fr), Grayumet al. 8348 (BG, F, MO). PANAMA. BOCASDEL TORO:Almirante,1922 (? fl-fr), Cooper et al. 28 (US); Changuinola valley, 1927

(st), Cooperet al. 107 (F, GH, NY, US); Duwebdulup Peak, N of Rio Terebe,across from QuebradaHuron, 13 Apr 1968(R fl-fr),Kirkbrideet al. 569 (MO);4 mi N of Almirante,18 Aug 1964 (? fl-fr), McDaniel 5098 (MO);CerroPila de Arroz,rd. to ChiriquiGrande,10 mi from continental divide, 10 Mar 1986 (? fl-fr),

Distribution (see Fig. 77.1). Westem Panamato Nicaragua(Rio San Juan);in wet forest;at low elevations. It is not likely that the species occurs in Guate- McPherson 8776 (BG, MO); rd. to Chiriqui Grande, N mala,as is indicatedon thetypespecimenof S.pubivena. of Fortuna Dam, 29 Jun 1987 (? fl), McPherson 11120 (BG, MO). CHIRIQUi: Cerro Galera Chorcha,

Gualaca, Representativespecimens studied. NICARAGUA. 1 Aug 1947 (? fr), Allen5059 (BM, BR, EAP, G, MO); Rio SAN JUAN: Rio Indio, Siempreviva,30 Nov 1982 (? fl-fr), Araquistain3317 (MO); between Rio Santa 23 km N of Puento de Burica, 5 Feb 1973 (st), Busey 748A (MO); Progreso, Jul-Aug 1927 (2 fr), Cooper et Cruz& CanioSantaCrucita,La Palma,30 Nov-2 Dec 1094 (st), Stevens 23444 (MO). COSTA RICA. ALAJUELA: E of San Rafael,W of La Marina, 19 May 1968 (d'), Burger et al. 5082 (CR, F,

NY). CARTAGO: BetweenTuis & Bajo Pacuare,15 Nov 1975 (e), Burgeret al. 10046 (F, U), 10049 (CR, F, U); Rio Chitaria,12 Oct 1967(? fr),Lent1419 (F,GH,NY); Tuis,nr.Turrialba, Jul 1897 (? fl-fr),Pittier11266 (BR, CR, F, MICH,US); 2 km W of Turrialba, 6 Sep 1990 (9 fl), Solomon 19197 (BG); Rio Yurquin,Mar 1894 ( fl), Tonduz11357 (CR, US). GUANACASTE: Tileran, Quiros27668 (CR). HEREDIA: 8 km S of PuertoViejo, 7 Jan 1967(? fl-fr),Burgeret al. 4290 (BM,F, MO,U); FincaLa Selva,PuertoViejo de Sarapiqui,15 Aug 1979 (? fl-fr), Grayum2365 (DUKE), (e), Grayum2375 (DUKE),23 Jun 1980 (? fr),Hammel9106 (F), 15 Mar 1985(Y fl-fr),Schatzet al. 1055 (WIS).LIM6N:Between Cahuita& Suretka,23-25 Apr 1982 (? fr), Barringeret al. 2660 (F); 4 km S of Cahuita,9-14 Feb 1977 (? fr), Burgeret al. 10488 (CR, EAP,F, MO);Los Diamantes, Rio Clara,1.6 km E of Guapiles,11 Jul 1949 (e), Holm et al. 389 (A, BM, F, G, K, MO, P, U, US); Rio Blanca, Santa Clara, Jul 1899 (a), Pittier 13453 (CR, G, US); ParqueNacional Tortuguero,5 Nov 1987 (e), Robles

1262 (BG),22 Nov 1987 (? fl),Robles1288 (BG,MO), 7 Feb 1989 (? fr), Robles2620 (BG, F, MO);nr.mouth of Rio Reventazon,23 Oct 1951 (e'), Shanket al. 4384 (BM, EAP, F, GH, US); CerroCoronel, E of Laguna Danto, 16-23 Jan 1986 (st), Stevens 23754 (MO). PUNTARENAS: BetweenPuertoJimenez & Rio Tigre, 3 Apr 1930, Brenes 18950 (CR); 4 mi W of Rinc6n de Osa, 4-7 Jan 1968 (6e), Burger et al. 5500 (BM, CR, F,

MO, PMA,U); betweenQuebradaBonita& CararaReserve, 25 Jul 1985 (? fl-fr), Grayumet al. 5709 (BG, MO);ParqueNacionalCorcovado,LowerOllas, 11 May 1988 (e), Kernan 406 (BG, MO); Rincon de Osa, 10 Feb 1974 (st), Liesner 1984 (MO); Peninsulade Osa, CorcovadoNationalPark,5 Jul 1977(? fr),Liesner2939 (MO,U); 4 mi W of Rinconde Osa, 6 Aug 1967 (? fr), Raven21558 (BM, CR, F, MO, U); N of Palmar,Mar

al. 174 (F, NY); 11 mi S of Puerto Armuelles, 22 Feb 1973 (st), Croat 22139 (MO); W of San Bartolo Limite,

24 Feb 1973(st), Croat22182A& 22195A(MO);Burica Peninsula, nr. Brazo Seco, 3 Mar 1973 (st), Croat 22557B (MO); 7 mi on rd. Puerto Armuelles-San Bartolo Limite, 19 May 1976 (st), Croat 35043 (MO, U).

4b. Sorocea pubivena Hemsley subsp. oligotricha (Akkennans& C. C. Berg) C. C. Berg, Novon 6: 243. 1996. Sorocea hirtella Mildbraed subsp. oligotricha Akkermans& C. C. Berg, Proc. Kon. Ned. Akad. Wetensch.,Ser. C, Biol. Med. Sci. 88: 383, t. 1. 1985. Type.Brazil.Amazonas:Mun. Sao Paulode Oliven9a,nr.Palmares,11 Sep-26 Oct 1936 (? fl-fr),Krukoff8157(holotype,U; isotypes,BM, F, G, GH, K, LE, MICH,MO, NY, US). Fig. 21 Sorocea faustiana Cuatrecasas,Ciencia (Mexico) 24(5/6): 185, t. 1. 1966. Type. Colombia. Valle: Rio Anchicaya, El Prado, 4 Aug 1943 (? fl-fr), Cuatrecasas 14836 (holotype, F).

Leafy twigs appressed- (to patent-)puberulous. Lamina subcoriaceousto coriaceous; margin mostly entire(occasionally distinctly dentate);uppersurface minutelypuberulouson the midribor glabrous;lower surfaceon the mainveins sparselypatent-to appressedpuberulouswith hairsof aboutequal length.Pistillate inflorescencesto 1O(-12)cm long in fruit;flowers ca. 10-20(-30). Fruitingperianth usually (sub)globose, mostly distinctlypedicellate,subglabrous. Distribution (see Fig. 77. 1). In the upperAmazon Basin, extendingto Guyana,easternandnorthernVenezuela, northernand western Colombia, and eastern Panama;in wet forest;to 1250 m, but in the Pakaraima Mountains(Guyana)and Pico da Neblina to 2200 m.

108

FLORA NEOTROPICA

Fig. 21. 1-3. Sorocea pubivena subsp. hirtella. 1. Part of leaf (Prance et al. 24016). 2. Pistillate flower (Kayap 611). 3. Infructescence (Prance et al. 24016). 4-13. Sorocea pubivena subsp. oligotricha. 4. Leafy twig. 5. Young infructescence (N. T. Silva 60618). 6. Pistillate flower (Pires 343). 7. Pistillate inflorescence (Fr6es 22148). 8. Infructescence (Maguire 45945A). 9. Pistillate flower. 10. Fruiting perianth (Schultes 6171). 11. Staminate inflorescence (Pires 248). 12. Staminate flower (Croat 20357). 13. Staminate flower (Pires 338). [By T. Schipper, Utrecht.]

TAXONOMIC TREATMENT

109

Representative specimens examined. PANAMA. fr), Fanshawe 1144 (= FD 3880) (K); Cuyuni-Mazaruni COL6N:Ca. 2-3 mi up to the Rio Guanche,19 Jan 1973 region, PakaraimaMts., Mt. Ayanganna, 1100-1200 m, (e), Kennedy et al. 2124 (F, MO);SantaRitaRidge,nr. 7 Nov 1992 (6'), Hoffman et al. 3262 (BG), (9 fl), end of rd., 25 May 1987 (? fl), McPherson 10970 (BG,

MO).SANBLAS:Rio Play6n,Chico, 11 Jun 1994 (? flfr), H. Herrera1664 (BG);El Llano-Cartird., km 19.1, 11 Mar 1986 (st), Nevers et al. 7358 (BG, MO, PMA). COLOMBIA. AMAZONAS: Rio Caqueta, Villa Azul, 213 Nov 1989 (st), Andel et al. 500 (BG); Leticia, SepNov 1944 (? fr), Schutltes 6171 (F, K, US); Rio Loretoyacu, Sep 1946 (? fr), Schultes et al. 8325 (COL, GH, K, US); Puerto Narinio,Rio Loretoyacu, 16 Jun 1973 (? fl), Soejarto et al. 4178 (HUA, NY); PuertoNariiio, Parque Nacional Amacayacu, 3 Aug 1989 (2 fl), R. Vasquez et al. 12520 (COL, MO). ANTIOQUIA: ca. 38 km W of Barrancabermeja, 1 Mar 1967 (d), Bruijn 1541 (MO, NY, U); Mun. Carepa, 45 km S of Turbo, 31 Jul 1987 (? flfr), Callejas et al. 4854 (BG, HUA, MO); Mun. San Luis, Rio Claro, 2 May 1972 (? fl-fr), Cogollo 1672 (HUA, MO); Rio Claro, rd. Refugio-Antioquia, 10 Oct 1982 (? fl-fr), Renteria et al. 2823 (MO); Mun. Anori, Rio Anori, 4 km from Providencia, Buenos Aires, between Dos Bocas and Anori, 26 Apr-3 May 1973 (e), Soejarto et al. 3931 (MO), and 5 km from Providencia, (? fl-fr), Mun. Soejarto et al. 4092 (F, GH, HUA, MO). BOLIVAR: Morales, Cgto. Norosi, rd. to Tiquisio Nuevo, 9-14 Apr 1985 (a'), Cuadros V 2108 (MO); Quebrada la Culebra, La Raya, Caiio Caribona, between jct. of Rio Cauca and Rio Magdalena, 4 May 1987 (st), Gentry et al. 57259 (MO). BOYACA: El Humbo region, 130 km N of Bogota, ca. 1000 m, 31 Mar 1933 (ae),Lawrance 730 (A, F, G, K, MO, US). CAQUETA: Mun. Vicente de Caguan, rd. San Vicente-Neiva, 27 Nov 1990 (o), Betancur et al. 2207 (HUA). CHOC6: Mun. Riosucio, Cerros de Uraba, rd. La Eugenia (Nova)-La Cumbre Sureste, 18 Apr 1988 (e'), Cardenas 1727 (BG, MO); Mun. Riosucio, Cerros del Cuchillo, Quebrada El Cedro, 20 May 1988 (? fl-fr), Cardenas 2008 (MO). C6RDOBA: Mun. Ayapel, Finca Canime, 4 Apr 1990 (e'), Marulanda 2067 (HUA). META: La Macarena, Rio Guayabero, El Refugio, 1 Mar 1976 (? fl), Idrobo 8629 (COL). NARINO: Mun. La Espriella, Estacion CONIF, Rio Mira, 9 Jun 1986 (? fr), Le6n et al. 1329 (BG, U). VALLE: Rio Anchicaya, La Planta, 17 Sep 1943 (Y fl-fr), Cuatrecasas 15204 (F, VALLE).VICHADA: Territorio Faunistica del Tuparro, 1 km N of El Tap6n, 13 May 1979 ( fl), Vincelli 1139 (COL). VENEZUELA. AMAZONAS: Between Rio Mawarinuma and Rio Baria, 7 May 1984 (a'), Gentry et al. 47244 (BG, MO), Thomas et al. 3390 (BG, NY). ARAGUA: Rd. Guamites-Rancho Grande, Parque Nacional, 1 Jul 1947 (a'), Pittier 15477 (US, VEN). BOLIVAR:Between Campamento Los Vaquiros and El Cruzero, 14 Jul 1960 (? fl-fr), Steyermark 86332 (NY, U, US). CARABOBO: Ca. 15 km S of Puerto Cabello, 6 km S of San Estaban, 14 Apr 1982 (a), Liesner et al. 13635 (MO, NY, U). GUYANA. Cuyuni R., Oct 1904 (a), Bartlett 8383 (B, K, NY, U); Mazaruni station, 6 Jan 1940 ($ fr), Fanshawe 337 (= FD 3073) (K, U); Lower Essequibo R., Tiger Cr., 9 Dec 1940 (? fl), Fanshawe 338 ( FD 3074) (K, U); Potaro R., Eagle Mt., 26 Jan 1943 (R fl-

Hoffman et al. 3268 (BG); Macouria R., Nov 1886 (d), Jenman 2480 (K); Kurupung, Tacoba, 25 Nov 1922, Lang & Persauds.n. (F); southernPakaraimaMts., Kamana Falls, 2200 m, 25 Aug 1961 (? fl-fr), Maguire et al. 45945A (U); Kako R., 23 Sep 1960 (? fl-fr), Tillett et al. 45495 (MO, NY). ECUADOR. ESMERALDAS:Canton San Lorenzo, Reserva Indigena Awa, Ricaurte, centro Pambilar,21 Jan 1993 (e), Aulestia et al. 1072 (QCNE); Eloy Alfaro, Charco Vicente, Rio Miguel de Cayapas, 20 Feb 1993 (st), Mendez et al. 43 (QCNE); Cant6n San Lorenzo, Reserva Indigena Awa, Ricaurte, 15-29 Apr 1991 (? fr), Rubio et al. 1352 (BG, QCNE). MORONA-SANTIAGO: Centro ShuarYukutais, 12 Mar 1990 (? fl-fr), Bennett et al. 4092 (BG) and 4121 (BG, NY). NAPo: rd. HollinLoreto, km 31, Challua, 1200 m, 18-21 1989 (st), Hurtado et al. 2287 (BG); ParqueNacional Yasuni, Pozo Petrolero Amo 2, 9-19 Jan 1988 (e), Neill et al. 8223 (MO, QAME, QCNE); Reserva Biol6gica Jatun Sacha, 8 km E of Puerto Misahualli, 9 Jan 1990 (c'), Palacios 4782 (BG, MO, QCNE); Parque Nacional Yasuni, Ahiangu,30 May-21 Jun 1982 (Y fl-fr), SEF 8719 (MO, NY, U). PASTAZA: Rd. Coca-Auca, 11 km S of Coca, 10 km from Rio Tigiiino, 7 Jan 1989 (e'), Palacios et al. 3352 (MO); Cant6n Puyo, Santa Cecilia, Villano, 1 May 1992 (? fr), Palacios 10101 (BG, QCNE); Pozo Petrolero Ramirez, ca. 20 km S of Curaray,21-28 Feb 1990 (? flfr), Zak et al. 5101 (BG). SUCUMBiOS: Reserva Faunistica Cuyabeno, nr. Laguna Grande, 12 Mar 1990 (? fl-fr), Valencia et al. 357 (QCA). TUNGURAHUA: Rio Negro, ca. 1250 m, 2 Mar 1972 (6), Harling 11117 (GB). PERU. LORETO: Varadero de Mazan, Rio Amazonas-Rio Napo, 22 Aug 1972 (? fl), Croat 19532A (MO); Rio Napo, Negro Ulco, 17 Sep 1972 (st), Croat 20357 (GH, MO); Quebrada Yanamono, above mouth of Rio Napo, 11-14 Nov 1979 (st), Gentry et al. 27815 (MO, U); Rio Amazonas, Explorama Inn, 1 km S of Indiana, 16 Jun 1986 (o), Gentry et al. 54582 (BG, MO); Prov. Requena, Rio Amazonas, Sinchucuy, 8 Aug 1986 (e), R. Vdsquezet al. 7762 (BG, MO); Rio Amazonas, Caballo Cocha, 5 Aug 1929 (? fr), Ll. Williams 2092 (F); Rio Amazonas, La Victoria, 30 Aug 1929 (st), Ll. Williams2965 (F). MADRE DE DIos: Prov. Manu, Parque Nacional Manu, Rio Manu, Rio Sotileja, 19 Oct 1986 (e), Foster 11730 (F). SAN MARTIN: Prov. Lamas, rd. Tarapoto-Yurimaguas,km 72, Santa Rosa de Davidcillo, 28 Sep 1986 (? fl-fr), Knapp et al. 8459 (BG). BRAZIL. ACRE: Rio Jurua, nr. Colonia Rodriguez Alvez, 28 Sep-10 Oct 1986 ($ fl-fr), Campbell et al. 10849 (BG); nr. Sena Madureira, 27 Sep 1980 (? fr), Ferreira et al. 2573 (BG, MO); Mun. Mancio Lima, Rio Moa, Fazenda Arizona, 19 Sep 1984 (9 fl-fr), Ferreira et al. 5313 (BG); rd. Abuna-Rio Branco, km 242-246, nr. Campinas, 18 Jul 1968 (6), Forero et al. P6361 (U); 2-4 km W of Cruzeiro do Sul, 22 Oct 1966 (? fl-fr), Rio Prance et al. 2733 (INPA, NY, U, US). AMAZONAS: Negro, Foz do Caiary, 17 Apr 1947 (v), Fr6es 22148

110 (IAN, U); Rio Negro, Tapuruquara,8 Apr 1947 (d), Pires 248 (IAN, MO, US); Rio Negro, Jupati, 17 Apr 1947 (o'), Pires 343 (IAN, MO, NY, US); Rio Negro, Uaupes, 1 May 1947 ( fil), Pires 518 (IAN, MO, NY, US); between Igarape Anta and Pico da Neblina, 1350-2000 m, 14 Dec 1965 (? fl), N. T Silva et al. 60618 (NY, U, US); Rio Uaupes, Panure, Oct 1852 (st), Spruce 2658 (B, BM, BR, P). ROND6NIA: 1 km S of Riberao, 27 Jul 1968 (9 fr), Prance et al. 6572 (U).

This subspeciescan be distinguishedfromSorocea steinbachiiby the equallengthof the hairson the lower leaf surface,mainlyoccurringon themidrib,andby the greaternumberof lateralveins. It can be distinguished InGuyana alsofromS. muriculataby thelattercharacter. andthe adjacentpartof Brazil,the midribof the lamina is glabrous above, whereas in the other parts of the species range it is minutely puberulous.The glabrous midrib, the relatively small fruitingperianth,and the possible disjunctoccurrenceof this form could be argumentsto recognize a distinctinfraspecifictaxon. The collections from high elevations, 1350-2200 m (N. T Silva et al. 60618 andMaguireet al. 45945a) hardly differ from the lowland collections. Subspies oligotrichaandsubsp.hirtellaoftenco-occurandflower simultaneously,apparentlywithoutproducingintermediateforms,as has beenpersonallyobservedin Reserva FaunistaCuyabenoin Ecuador.

FLORANEOTROPICA ulous), at least ? densely patentpuberulous(to hispidulous) on the main veins; lateralveins often connected with a slightly arcuatesubmarginalvein. Pistillate inflorescences to 9.5 cm long in fruit;flowers 3-15(-25). Fruitingperianth usually (sub)globose, sometimesto ovoid, to oblongoid,or to oblate,sometimessubsessile, sparsely(to densely)puberulousto hirtellousor to subhispidulous. Distribution (see Fig. 77.7). In the upperAmazon Basin; in non-inundatedwet forest; to 1250 m. Representative specimens examined. COLOMBIA. AMAZONAS: Rio Cotuhe, 8 Nov 1946 (st), Black et al.

46-354 (US); Rio Caquetu,nr. Isla de las Palmas, 28 Aug 1986 (st), P Palacios et al. 2014 (BG); Rio Loretoyacu, nr. Leticia, 12 Sep 1963 (9 fl-fr), Soejarto et al. 794 (GH). VENEZUELA. AMAZONAS: Depto. Atures, 44-45 km SE of PuertoAyacucho, 9 May 1980 (q fl-fr), Steyermark et al. 122182 (MO, U). ECUADOR. NAPO: ParqueNacional Yasuni, Aiiangu, 1-30 Apr 1985 (? fl-fr), Korning et al. 58632 (QCA); Rio Napo, 9 km downstream Misahualli, 10 Mar 1985 (a'), Neill et al. 6040 (BG, MO, QAME, QCNE); Reserva Floristica El Chuncho, 5 km N of Coca, 9 Oct 1987 (st), W Palacios 2140 (BG, MO, QAME). PASTAZA: Lorocachi, 23-29 May (9 fl-fr), Brandbyge et al. 30650 and 31238 (AAU). SUCUMBIOS: Reserva Faunistica Cuyabeno, nr. Laguna Grande, 11 Mar 1990 (? fr), Balslev et al. 97004 (AAU, BG, QCA). PERU. AMAZONAS: Rio Cenepa, QuebradaKayamas, Local names. Colombia:Amazonas:puruhi(Yuc.), 14 May 1973 (9 fr), Ancuash 377 (MO, U); Rio Cenepa, takatakaramio(Mui.), tujiguarce(Mui.), viecrinajae Aintami, 19 Jan 1973 (o'), Kayap 184 (GH, MO); Que(Mir.). Venezuela: Bolivar: charo macho. Ecuador: brada Huampami, 15 Feb 1973 (9 fl-fr), Kayap 362 (MO, Napo: vegotahuemo(Huaorani). NY); QuebradaChigku Shinun, 11 Apr 1973 ($ fr), Kayap 611 (MO, U). HUANUCO:Prov. Pachitea, 2-5 km E of Puerto Inca, 11 Sep 1982 (a), Foster 8691 (MO, U); 4c. Sorocea pubivena Hemsley subsp.hirtella (Mild- Prov. Pachitea, 26 km S of Puerto Inca, 27 Sep 1988 bread) C. C. Berg, Novon 6: 243. 1996. Sorocea (e'), Wallnoefer15-27988 (BG).LORETO:Rio Corrientes, Notizbl.Bot.Gart.Berlin-Dahlem between Tnte. L6pez and Puerto Avanzado, 4 Apr 1977 hirtella Mildbraed, 10: 183. 1927; Macbride, Publ. Field Mus. Nat. (9 fl-fr), Gentry et al. 19034 (MO, U); Prov. Alto Hist., Bot. Ser. 13(2.2): 311. 1937; Berg & Akker- Amazonas, rd. Yurimaguas-Tarapoto,15 km SW of Yurimans, Proc. Kon. Ned. Akad. Wetensch., Ser. C, maguas, 10 Oct 1985 (? fl-fr), Gentry et al. 52209 (BG, nr. Iquitos, Jan 1930 (d), Klug 739 Biol. Med.Sci. 88:383, t. 1. 1985.Type.Peru.Loreto: MO); Mishuyacu, (F, US); Iquitos, 27 Feb 1924 (do), Kuhlmann 1517 (G, upperRio Marafion,Pongo de Manseriche,11 Sep U, US); Rio Javari, 2 hrs. above Rio Javarizinho, 24 Oct 1924 (o), Tessmann4016 (holotype, B; isotypes, 1976 (9 fl-fr), Prance et al. 24084 (INPA, NY, U); Prov. Fig. 21 Alto Amazonas, Shushuyacu, 14 Sep 1981 (o), R. Vdsquez F, GH, NY, US). et al. 2498 (BG); Prov. Maynas, Puerto Almendras, 20 Sorocea opima Macbride,Publ. Field Mus. Nat. Hist., Apr 1983 (9 fr), R. Vdsquez et al. 3984 (BG, MO); Prov. Bot. Ser. 11: 64. 1931; Burger,Lanjouw& Wessels Maynas, Yanamono, 23 Nov 1983 ($ fr), R. Vdsquezet Boer, Acta Bot. Neerl. 11: 445, t. 4. 1962. Type. al. 4695 (BG); Prov. Maynas, rd. Iquitos-Nauta, km 42, Peru.Loreto:Caballo-Cocha,Aug 1929 (q fr), LI. 18 Aug 1987 (o), R. Vasquezet al. 9374 (BG,MO); Prov. Williams 2337 (holotype, F; isotype, US). Requena, Sapuena, Jenaro Herrera, 13 Sep 1987 (o), R. Vasquezet al. 9560 (BG, MO); Previsto, 17 Oct 1962 (a), Leafy twigs brown to whitish-subhirsuteto hirtel- Woytkowski7628 (GH, MO, US). MADRE DE DIos: Prov. lous. Lamina chartaceousto subcoriaceous;margin Tambopata, Zona Reservada de Tambopata, Laguna

entireto obscurelydent(icul)ate;uppersurfaceminutely puberulouson themidrib;lowersurface(rather)densely hirtellous (to subhirsute)to puberulous(or to hispid-

Cocococha, 11 June 1989 (st), Phillips et al. 323 (BG); Prov. Tambopata, Zona Reservada de Tambopata, 18 Aug 1990 (st), Reynel R. et al. 5217 (BG). PASCO: Shirin-

TAXONOMIC TREATMENT

111

gamazu,ca. 20 km S of Iscozacin,Rio Palcazuvalley,6 slightlyimpressedabove,sometimesalsothelateralveins Jul 1988 (st), Gentryet al. 63332 (BG); Prov. Oxa- slightlyimpressed,prominentbeneath;lateralveins 9pampa,Palcazuvalley,Rio San Jose in the Rio Chuchu- 14 pairs; tertiaryvenation mostly partly scalariform; rrasdrainage,13 May 1983 (? fl-fr),D. N. Smith3998 petiole0.5-1.5(-3.5) cm long, 1.5-2 mm thick,sparsely (BG, MO);Prov.Oxapampa,GranPajonal,nr. Chequiandminutelypuberulousto glabrous;stipules0.3-0.8 tavo, 1250 m, 23 Sep 1983 (? fl-fr), D. N. Smith5164 cm long, puberulousto glabrous,caducous.Staminate (BG,MO,MOL).SAN MARTiN: Prov.Lamas,SantaRosa patentto pendulous,(0.5-)2-6(-10) cm inflorescences de Davidcillo,E of km 72 on rd. Tarapoto-Yurimaguas, 28 Sep 1986 (? fl-fr), Knappet al. 8459 (MO);Prov. long, including the 0.1-0.4 cm long, puberulouspeMariscalCaceres,Dtto. TocacheNuevo, Rio Tocache, duncle;flowersrathercrowdedto distant,(rather)broadly 12 Apr 1970 (Q fl-fr),SchunkeV 3895 (COL,F, G, GH, sessile;perianthca. 2 mm long, 4-parted,ciliolate;staK, MO,MOL,NY, US, WIS);Dtto.TocacheNuevo,Rio mens 4, isomorphicor anisomorphic,straight(with the Huallaga, Puerto Pizana, 8 Jan 1971 (e), Schunke V anthers? transverse)or bent outwardsat anthesis;fila4632 (COL,F, G, GH, K, MO, MOL,NY, US), 17 Apr ments0.5-1.2 mm long; anthers0.54.7 x 0.6-0.8 mm, 1971 (9 fr), SchunkeV 4827 (COL,F, G, GH, K, MO, the connectivebroad.Pistillate inflorescencespatentto MOL,NY, P, US); Rio Huallaga,QuebradaHuicte,27 pendulous,1-3 cm long, to 9 cm long in fruit;peduncle May 1964 ( fr), Schunke V 6506 (F, MO, US).0.1-0.5(-2) cm long, to 0.8 cm long in fruit, puberuUCAYALI: Jct. Rio Pachiteaand Rio Yuyapichis,SE of Pucallpa,4 Oct 1985 (? fl-fr),Morawetzet al. 14-41085 lous; rachisminutelypuberulousto glabrous,camose (BG), 26 Sep 1985 (e'), Morawetz et al. 18-26985 (BG) (?) and red (?) in fruit; flowers (5-)15-40, sessile to BRAZIL. ACRE: Mun.MancioLima,Serrada Moa, shortlypedicellate;pedicel 0. 1-0.3(-0.5) cm long, or 30 Sep 1984(? fl), Ferreiraet al. 5088 (BG,NY);mouth occasionallyto 2 cm longin fruit,to 3 mmthick,sparsely of Rio Embira,30 June 1932 (o), Krukoff5084 (A, LE, puberulous;perianth2-2.5 mm long, 4-lobed,theupper km partbroadlyovoid, glabrous,the lowerpartshort,denseMO, U, US). AMAZONAS: Rd. Manaus-Itacoatiara, 26, ReservaFlorestalDucke,Igarapedo Tinga,23 Nov ly, minutely puberulousto glabrous;stigmas tongue1955 (? fl), Brito et al. 13 (INPA); Sao Gabriel da shaped,0.4-0.6 mm long, finely papillate.FruitingpeCachoeira,25 Nov 1987 (? fl-fr), Kawasaki281 (BG); rianthsubglobose(to broadlyovoid orobovoid),0.5-0.8 Sao Paulo de Oliven9a, 26 Oct-Il Dec 1936 (? fl), x 0.5-0.8 cm, tuming fromred to black, subapiculate, Krukoff8648 (BM, F, G, K, MICH,MO,NY, P, U, US); Rio Negro, Ilha Tamandua(Ilha Maraj6),19 Oct 1987 the surface smooth, subglabrous,sometimes densely (? fl-fr),Maaset al. 6787 (BG,MO);Rio Javari,Esterao puberulous or with rather dense, globose-capitate, do Ecuador,22 Oct 1976 (? fr), Prance et al. 24016 pluricellularhairs. km 160, 10 Dec 1965 (? (NY); rd. Manaus-Itacoatiara, Distribution. (see Fig. 77.2). In the upperAmazon fl-fr), Rodrigues et al. 7348 (INPA, U).

Local names. Colombia: Amazonas: dicrinajai (Mir.). Peru:Amazonas: tsakaya; San Martin:sacha chimicua;Pasco:jopi.

5. Sorocea steinbachii C. C. Berg, Proc. Kon. Ned. Akad.Wetensch.,Ser.C, Biol. Med. Sci. 88: 383, t.

2. 1985. Type.Bolivia. SantaCruz:Prov.Sara, Buenavista,27 Nov 1925 (? fr),Steinbach7356bis (holotype, U; isotypes B, BM, F, G, GH, K, MO, Fig. 22 NY). Tree,to 30 m tall,orshrub. Leafy twigs 1-3.5 mm Laminaoblong thick,sparselyto denselypuberulous. to elliptic (or to lanceolate),(2.5-)5-15(-32) x (1.5-) 2.5-7(-13) cm, mostly broadestabove the middle, al-

base mostequilateral, subcoriaceous; apexacuminate; todistinctly acutetoobtusetorounded; margin obscurely to subentire; uppersurfaceon themidrib dent(icul)ate torather glapuberulous, denselyandminutely sparsely lowersurfaceonthemainveinsverysparsely brescent; hairs withmostlyappressed torather denselypuberulous of different lengths;venationalmostplaneorthemidrib

Basin, fromColombiato Bolivia; in non-inundatedor seasonally inundated(tahuampa)forest; to 1200 m. Representative specimens examined. COLOMBIA. Rio Caqueta,30 Sep 1990 (e), AMAZONAS: Araracuara,

Alvarezet al. 120 (NY);PuertoNarifno,ParqueNacional Amacayacu,3 Aug 1989 (ae),R. Jksquezet al. 12525 (MO),10Aug 1989(? fl), R. V6squezet al. 12697 (BG). RioCaqueta, CAQUETA:Solano,8 kmSEof TresEsquinas, belowmouthof Rio Orteguaza,7 Mar1945(a'), Littleet Rio al. 9651 (COL,GH,NY). META: Mun.La Macarena, mouthof Rio Losada,11 Aug 1988(? fl-fr), Guayabero, Callejas et al. 7087 (MO); Mun. La Macarena,Rio S of LaMacarena,12Aug 1988(st), Callejas Guayabero, et al. 7126 (MO);MacarenaSerraniade Chamusa,Rio Duda, 20 Jun 1988 ($ fr), Stevenson69 (MO). PUTUMAYO: Mocoa, Jul 1989 (st), Mahecha et al. 5682a (UDBC).VICHADA: ParqueNacionalNaturalEl Tuparro, ca. 6 kmN of El Tap6n,21 Mar1985(e), Zarucchiet al. 3800 (BG, MO),($ fl), Zarucchiet al. 3817 (BG, MO). ECUADOR. MORONA-SANTIAGO: Taisha, 6 Feb 1962(e), Cazaletet al. 7684(US).NAPO:ParqueNacional Yasuni,confluenceof Rio Tivacunoand Rio Tiputini,1 Apr 1988(a'), Coello217 (BG, QCNE),(9 fl-fr),Coello 221 (BG, QCNE);ReservaBiol6gicaJatunSacha,8 km E of PuertoMisahualli,12 Nov 1987 (? fr), Cer6n2697

112

FLORANEOTROPICA

I

,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

(Fosteret aL.5187). 2. Pistillateflower.3. Fruiting Fig. 22. Soroceasteinbachii.1. Leafytwig withinfructescence perianth.4. Seed. 5. Embryo(Lieraset at P.16931). 6. Staminateinflorescence(Revilla845). 7. Staminateflower and stamens(Croat 20112). [By H. Ryp)kema,Utrecht.]

TAXONOMICTREATMENT (BG,MO,QCA,QCNE);rd. Hollin-Loreto,km 17, 1100

113

m, 14-22 Feb 1989 (e), Hurtado et al. 1702 (BG), 14-

Croat et al. 62457 (BG, MO); nr. Sena Madureira,2 Oct 1980 (? fr), Ferreira et al. 2679 (MO, NY); Rio Yaco,

16 Sep 1989 (Y fl), Hurtadoet al. 2109 (BG); 7 km N of Coca, Estaci6nExp. INIAP-Payamino,6 Apr 1985

mouthof Rio Macaua,4 Sep 1933(? fl-fr),Krukoff5785 (A, BM, F, G, K, MICH,NY, U, US); 2-4 km W of

(? fl-fr), Neill et al. 6241 (BG, MO, QAME, QCA,

Cruzeirodo Sol, 24 Oct 1966 (9 fl-fr), Prance et al. 2769

QCNE);20 km W of Coca, 22-23 Apr 1985 (Y fl-fr),

(MG,MICH,MO,NY, U, US); Taraca,24 Sep 1968 (9

Neill et al. 6391 (BG, MO, QAME, QCA, QCNE).

fl-fr), Prance et al. 7531 (F, GH, MG, NY, U, US); 4 km

PASTAZA: trailCeilan-RioCononaco,N

of Rio Curaray, E of SenaMadureira, 27 Sep 1968(? fl-fr),Pranceet al. 7619(C, MG,MICH,MO,NY,U, US);nr.Serrada Moa, Pozo PetroleroMasaramu,40 km NNE of Montalvo, 24 Apr 1971 (9 fl-fr),Pranceet al. 12417 (GH,K, MG, 26-30 Apr 1990 (? fl-fr), Gudino248 (BG, MO); Rio MO,NY, P, U); SantaLucia,Transamazonian Hwy.,40 km E of Cruzeirodo Sul, 14 Oct 1987 (9 fr), Pruskiet Curaray,ca. 10 km E of Curaray,22 Mar 1980 (e), Harlinget al. 17636 (BG, GB);Pozo PetroleroRamirez, al. 3462 (K). AMAZONAS:San Antonio de Ia, 7 Sep 20 km S of Curaray, 21-28 Feb 1990 (? fl-fr), Zak et 1906 (? fl-fr),Ducke MG 7655 (B, G, U); Rio Tarauca, al. 5020 (BG, QCNE). SUCUMBiOS: Dureno, 27 Dec nr.mouthof Rio Embira,21 Jun 1933 (st), Krukoff4954 6 Jun 1980 (? fl-fr), Brandbyge et al. 31631 (AAU);

1988 (st), Gentry et al. 64242 (BG); Cant6n Gonzalo

(A, BM, F, G, K, LE, MICH, MO, NY, U, US); Rio Javari,

Pizarro,Rio Tigre, 900-1 100 m, 18-21 Feb 1987 (d'),

Miraflor,30 Jul 1973 (? fl-fr), Lleras et al. P16931 (MO,NY, U); Tefe (= Ega), (9 fl-fr), Poeppig2563 (B, F, G, P); nr. Boca do Acre, 16 Sep 1966 (a), Prance et

Palacios et al. 1613 (BG, MO, NY, QAME, QCA,

QCNE);Cant6nGonzalo Pizarro,Santa Rosa Luz de Belen, 26 Mar 1992 (9 fl), Quelal et al. 419 (QCNE); ReservaFaunisticaCuyabeno,nr. LagunaGrande,15 Mar 1990 (? fl-fr), Valenciaet al. 362 (QCA). PERU. HUANUCO: Dtto. Monz6n, 1000 m, 11 Sep 1964 (Y fl), Schunke V 6601 (F, K, MO, US); Prov. Pachitea, ca. 26 km S of Puerto Inca, 22 Oct 1988 (Y flfr), Wallnoefer 13-221088 (BG). LORETO: Rio Amazonas, SE of Iquitos, 17 Aug 1972 (st), Croat 19245 (GH, MO, NY); nr. mouth of Rio Napo, 14 Sep 1972 (ce), Croat 20122 (C, GH, MICH, MO, NY); Prov. Alto

Amazonas,Andoas, Rio Pastaza, 15 Aug 1980 (Y fl), Gentry et al. 29732 (U); Reserva Nacional PacayaSamiria, Cocha Atum, 10 May 1985 (? fr), Grdndez et al. 389 (BG); Rio Marainon,nr. mouth of Rio Tigre, 19 Aug 1929 (? fl), Killip et al. 27531 (F, NY, US); Prov. Maynas, Dtto. Pebas, Rio Ampiyacu, 19 Jul 1976 (a), Revilla 845 (MO, NY, U); Puerto Margarita,19 Apr 1964 (Y fr), Schunke V 6415 (F, MO, US); lower Rio Itaya, Soledad, 29 Jan 1925 (c), Tessmann 5251 (B, F, NY, US); Prov. Loreto, Atun Cocha (Rio Samiria), 10 May 1985 (? fr), R. Vasquezet al. 6486 (BG, MO). MADRE DE Dios: Prov. Tambopata, Rio Heath, 8 km above confluence with Rio Madre de Dios, 22 Nov 1995 (? fl-fr), Alexiades 1248 (BG, NY); Parque Nacional del Manu, Rio Manu, Cocha Cashu Station, 21 Nov 1976 (? fr), Foster et al. 5220 (F), 21 Nov 1980 (2 fr), Foster et al. 5815 (F); Parque Nacional Manu, Rio Cumerjali, 20 Oct 1986 (? fl), Foster et al. 11939 (BG); Tambopata,"Cuzco Amazonico," 20 Feb 1989 (st), Gentry et al. 65957 (MO). PASCO: Puerto BermAdez,14-17 Jul 1929 (? fr), Killip et al. 26485 (NY, US); Prov. Oxapampa, Rio Pichis, nr. Paujil, 10 km downriver from Puerto Bermudez, 23-24 Sep 1982 (e), Foster 8853 (BG, MO), (? fl-fr), Foster 8892 (MO). SANMARTiN: Prov. Mariscal Ckceres, Tocache Nuevo, Quebrada de Tananta, 17 Dec 1970 (a), Schunke V 4573 (COL, F, MO, NY, US); Prov. Rioja, Dtto. Yorongos, Nuevo Tabaloso, 1020 m, 29 Oct 1996 (? fl-fr), I. Sanchez V et al. 8332 (BG). UCAYALI: Prov. Coronel Portillo, Leoncio Prado (Yarina-Cocha), 14 May 1984 (? fr), R. Vasquez 4994 (BG, MO). BRAZIL. ACRE: Rio Moa, 21 Aug 1986 (? fl-fr),

al. 2317 (F, GH, MG, NY, P, U, US); Rio Jurua, Jurua-

Mirim,Sep 1901(9), Ule 5871 (B, G, K), (d), Ule 5871a (B). ROND6NIA:Nr. P6rtoVelho, 14 Aug 1987 (? fl-fr), Dionizia et al. 156 (MO, NY). BOLIVIA. BENI: Prov.Yacuma,65 km SE of Bora, 65 km SW of SanIgnacio,27 Oct 1989(st), Foster et al. 13382 (F). COCHABAMBA:Prov. Carasco, Cant6n

Ivirgarzama,Estaci6nValle del Sajta UMSS, Campamento II, 27 Oct 1987 (9 fl-fr), Beck 13719 (BG). LA PAZ: Mapiriregion, San Carlos,28 Nov 1926 (9 fl-fr), Buchtien 1658 (B, US); Prov. Larejaca,rd. CaranaviGuanay,km 27.8, 28 Nov 1980 (st), Croat51634 (MO); Prov.Larecaja,Rio Mapiri,Tuiri,nr.Mapiri,12-30 Sep 1939 (c?), Krukoff10819 (F, G, MICH,NY, U, US) and 10932 (A, F, G, K, MICH,MO, NY, U, US); Prov.Sud Yungas,Alto Beni, rd. Palos Blancos-Yucumo,km 4, 24 Dec 1987 (9 fr), Seidel et al. 2181 (BG, LPB). PANDO:Prov.Nicolas Suarez, Rio Tahuamanu,Bella Flor,7 Oct 1989(9 fl-fr), Beck et al. 19220 (BG). SANTA CRUZ: Prov. Ichilo, Parque Nacional Ambor6, Rio Saguayo, 19 Dec 1988 (? fr), Nee 37261 (MO, NY); Prov.Ichilo, 30 km SE of Buena Vista, Rio Surutu,2 Sep 1985 (d), Solomon 14226 (BG, MO); Prov. Sara, BuenaVista,7 Oct 1916 (d), Steinbach 2939 (B), 2 Oct 1924 (d'), Steinbach 6548 (A, B, BM, F, G, K), 2 or 6 Oct 1925 (c?), Steinbach 7272 (B) and 7272a (B, BM, F, GH, K, MO, NY, U, Z). Materialof Soroceasteinbachiihasbeen treatedunder S. hirtella by Burger et al. (1962) due to a misidentificationof the type collection of S. hirtella, whichis presentlyreducedto a subspeciesofS. pubivena. Soroceasteinbachiishows morphologicalaffinities to both S. muriculata and S. pubivena. In the lamina characters,S. steinbachiidiffers fromS. muriculatain the minutely puberulousmidribabove. It differs from S. pubivena subsp. oligotricha in the indumentumof the lower leaf surface, especially on the midrib. In S. steinbachii,this indumentumconsists of minutepatent hairsanddistinctlylonger,mostly + appressed,some-

114

FLORA NEOTROPICA

times ? patenthairs.In S. pubivena, these hairsareal- sparselypuberulouswith equally long hairs;venation most equally long, either patent or appressed.More- planeor mostly the midrib? impressedandsometimes over,the numberof lateralveins is mostlymorethan12 the lateralveins as well above, prominentbeneath;latpairsin S.pubivena,butusuallyupto 12pairsin S. stein- eralveins (6-)8-12(-14) pairs;tertiaryvenation(usubachii. The pistillate inflorescence of S. steinbachii ally) reticulate;petiole 0.2-0.8 cm long, 0.5-1.5(-2) mostly has more flowers than those of the two other mmthick,mostlypuberulous;stipules0.2-0.4 cm long, species.The fruitsaresmallerthanthoseof S.pubivena. sparselypuberulousto glabrous,caducous.Staminate The stamens are bent outwardsin S. steinbachii, but inflorescencespendulous,1-3.5(-4.5) cm long, includinwardsin S. muriculata.Sorocea steinbachii cannot ing the 0.1-0.3 cm long puberulouspeduncle;flowers be distinguishedfromS. pubivena in charactersof the distant,broadlysessile;perianth1-1.5 mm long, deeply staminateflowers. 4-parted,puberulousor only ciliolate; stamens4, bent Specimensfromthe southernpartof the speciesarea inwards at anthesis; filaments 0.3-1.2 mm long; anoften have oblong to lanceolateleaves, broadestabove thers0.4-0.6 x 0.4-0.6 mm, the connectivebroad.Pisthe middle and mostly distinctlydentate.In the north- tillate inflorescencespatent,0.4-1.5 cm long, to 4 cm ern partof the species area,the leaves areusually ob- long in fruit;peduncle 0.1-0.3 cm long, puberulous; long to elliptic, broadestat the middle and subentire. rachispuberulous,camose andred in fruit;flowers(3-) Some specimens(e.g., Pruskiet al. 3462) have excep- 5-12(-20), (sub)sessile to shortlypedicellate;pedicel tionallylong pedicels,to 2 cm long in the fruitingstate, to 0.6(-1.2) cm long, camose andred in fruit;perianth similarto those of Sorocea trophoides. 1.5-2 mm long, 4-lobed to subentire,variousin shape Sorocea steinbachii has been cultivatedin the bo- andindumentum(see the subspecies);stigmastonguetanical gardenof Museu Goeldi in Belem (Para,Bra- shaped,ca. 0.5 mm long, finely papillate. zil) from materialcollected along Rio Ucayali (Peru) Sorocea muriculatais a variablespecies, especially by Kuhlmann.Fromthese staminatespecimensin cul- withregardto thefruitingstate.Remarkabledifferences tivation,severalherbariumcollectionshavebeenmade: occur in the shape of the fruiting perianth,fruit, and Ducke 929 (F, K, IAN, MG, MO, NY, US), Ducke seed. These differencescan be correlatedwith a minor MG 17044 (A, BM, F, G, US), Huber MG 6977(BM, differencein the staminateflower andallow the recogG, MG). nition of two subspecies. Gradationbetween the two Local names. Ecuador:Napo: ikicaspi (Quichua), subspecies occurs. oboye (Huaorani), tementayi (Huaorani). Peru: Key to the subspecies of Sorocea muriculata Hudnuco:mashonastillo;Loreto:turuuru(Jivaro). 1. Perianthof the staminateflower glabrous outside;upperpartof the perianthof the pistillateflowerdistinctlybroaderthanthe 6. Sorocea muriculata Miquel, in Martius,Fl. Bras. lowerpart;AmazonBasinto eastemVene4(1): 113, t. 34. 1853; Huber,Bol. Mus. Paraense zuela and Suriname .................4a. subsp.muriculata Hist.Nat. 5: 333. 1909;Macbride,Publ.Field Mus. 1. Perianthof the staminateflowerminutely Nat. Hist., Bot. Ser. 13(2.2): 312. 1937; Burger, puberulousoutside;upperpartof the periLanjouw& WesselsBoer,Acta Bot. Neerl. 11:436, anthof the pistillateflower about as broad t. 1. 1962; Berg & Akkermans,Proc. Kon. Ned. as the lower part;easternVenezuela,the Akad.Wetensch.,Ser.C, Biol. Med. Sci. 88: 387, t. adjacentpart of AmazonianBrazil, Suri3. 1985. Type. Brazil.Amazonas:nr.Tefe (= Ega), name, and FrenchGuiana..... 6b. subsp. uaupensis (V fl-fr), Martiuss.n. (lectotype, Burger,Lanjouw & Wessels Boer, Acta Bot. Neerl. 11: 436. 1962, M). Fig. 23 6a. Sorocea muriculata Miquel subsp. muriculata. Fig. 23 Shrubortree,to 5(-12) m tall.Leafytwigs0.5-3(-4) Sorocea amazonica Miquel, in Martius,Fl. Bras. mm thick, minutely and sparsely to rather densely 4(1): 111, t. 34. 1853;Burger,Lanjouw& Wessels puberulous.Lamina elliptic to oblong, sometimes to Boer,Acta Bot. Neerl. 11: 440, t. 1. 1962. Type. (narrowly)lanceolate,(2-)5.5-16.5(-22) x (I-)2-6.5 Brazil. Amazonas:nr. Tefe (= Ega), (? fl-fr), (-7.5) cm, broadestat or above the middle, usually ? Martiuss.n. (holotype,M; isotypes,BR, U). inequilateral,subcoriaceousto chartaceous;apex (cauSoroceadentataHuber,Bol. Mus.ParaenseHist.Nat. base acuteto obtuse(to rounded);mardate-)acuminate; 5: 333. 1909. Type.Brazil.Para:Rio Trombetas, gin obscurely to distinctly dent(icul)ateto subentire; 29 Sep 1907 (V), DuckeMG8957 (holotype, uppersurfaceglabrous;lowersurfaceon the mainveins MG;isotypes,BM, G, US).

TAXONOMICTREATMENT

115

Fig. 23. 1-7. Sorocea muriculatasubsp.muriculata.1. Leaf. 2. Infructescence.3. Pistillateflowerand fruiting perianth (Berg et al. P18517). 4. Pistillate flower and fruiting perianth (Davidse et al 16972). 5. Fruiting perianth (Fr6es 21448). 6. Fruiting perianth (Schultes et al 1.6972). 7. Staminate flower (Cardona 1344). 8-12. S. muriculata

subsp.uaupensis.8. Leafytwig with pistillateinflorescences(Aristeguietaet al 7361). 9. Leaf. 10. Infructescence. 11. Fruitingperianth(Alencar554). 12. Staminateflower (N. T Silva et al. 60808). [By T. Schipper,Utrecht.]

116

Perianthof the staminateflower glabrousoutside,

FLORA NEOTROPICA stream from Puerto Misahualli, 10 Mar 1985 (9 fl), Neill

thusonlyciliolateorcompletely glabrous. Pistillateflow- et al. 6059 (BG, MO, NY); 7 km N of Coca, 6 Apr 1985 ers3-12(-20) perinflorescence; upperpartof theperi- (? fl-fr), Palacios et al. 262 (AAU, BG, MO, NY, QAME, anthmostly(broadly) thanthe QCA, QCNE); Reserva Biol6gica Jatun Sacha, 8 km E broader ovoid,distinctly of Puerto Misahualli, 6 May 1990 (9 fl-fr), Palacios et lowerpartandminutelypuberulous to muriculate, the al. 4951 (BG, QCNE), 14 May 1990 (9 fr), Palacios lowerpartmuriculate to pusticulate. Fruitingperianth 5116 (BG, QCNE).PASTAZA: Lorocachi,26 May 1980 subglobose, ca.0.5-0.8x 0.5-0.8cm,red,tuning(blue-) (? fl-fr), Brandbyge et al. 31039 (AAU); Rio Curaray, black,thesurfacemuri(cul)ate, orwith? 10 km E of Curaray, 22 Mar 1980 (e?), Harling et al. pust(ic)ulate irregularly shapedprotuberances; pediceloftenstrongly 17651 (GB); Cant6n Pastaza, Pozo Petrolero Ramirez, swollenand/or? irregularly 21-28 Feb 1990 (? fl-fr), Zak et al. 5099 (BG, QCNE). shapedinfruit. SUCUMBIOS: Reserva Faunistica Cuyabeno, nr. Laguna Grande, 30 Mar 1992 (? fl), Jaramillo et al. 14698 (GB, NY), 12 Mar 1990 (9 fl-fr), Valencia et al. 311 (QCA). PERU. AMAZONAS:Rio Santiago, nr. Caterpiza,ca. 65 km N of Pinglo, 20 Nov 1979 (ei), Huashikat 1346 (MO, U), 4-12 Feb 1980 (? fr), Huashikat 1960 (MO, U); Representativespecimensexamined.COLOMBIA. Quebrada Chikishinuk, 31 Jan 1973 (? fl-fr), Kayap 283 AMAZONAS: nr. Leticia,24 Aug 1946 (e), Black 46-69 (GH, MO); Rio Santiago,nr. Caterpiza,29 Jan 1980 (? fr), (MO);ParqueNacionalAmayacu,nr. Mocagua,20 Jan Tunqui671 (MO, U). HuANuco: Prov. Pachitea,ca. 26 km 11988 (st), Gentryet al. 60814 (MO);Araracuara, Rio S of PuertoInca, 18 Oct 1988 (? fl), Wallnoefer13-181088 Caqueta, 25 Apr 1990 (? fr),Londonoet al. 1679 (NY); (BG); Prov. Pachitea, 21-24 km SE of Puerto Inca, 8 Oct

Distribution (see Fig. 77.3). In the upperand middleAmazonBasin,extendingto easternVenezuela andto Surinane;innon-inundated forestorsometimes in occasionallyinundated forest;to 1250m.

TrapecioAmaz6nico,Rio Loretoyacu,Oct 1946 (2 flfr), Schultes et al. 8547 (F, GH, IAN, K, U, US); Puerto Narinlo,ParqueNacionalAmacayacu,3 Aug 1989 (? fl), R. Vasquez et al. 12541(MO),4 Aug 1989(e'), R. Vsquez et al. 12566 (BG).GUAINIA: Rio Guainia,Rio Naquieni, base of CerroMonachi,Jun 1948 (? fr), Schulteset al. 10128 (GH,IAN, NY). VAUPtS:Mitul, 12 Nov 1939 (? fl-fr),Perez-Arbelaezet al. 6740 (COL);upperRio Vaupes, nr.Miraflores,4 Feb 1944 (e'), Gutierrezet al. 749 (GH, NY). META: Mun.Restrepo,upperRio Guacavia,(? flfr), Mahechaet al. UDBC 11179 (UDBC). VICHADA: TerritorioFaunisticadel Tuparro,1 km N of El Tapon, 13 May 1979 (? fl-fr), Vincelli1187 (COL). VENEZUELA.AMAZONAS: Depto.Atabapo,middle partof CainoCamane,Cucuritalde Caname,30 Apr-I May 1979 (? fl),Davidseet al. 16972 (MO,NY,U, US); nr. SanCarlosde Rio Negro, 15-20 May 1979 (? fl-fr), Liesner7430 (MO,U); SierraParima,Rio Matacuni,67 km W of Brazilianborder,18 Apr-23 May 193 (q fl), Steyermark107063 (MO, NY, US); ESE of Puerto Ayacucho,10-30 km on rd. to Gavilan,11 Jun 1977 (? fl-fr),Steyermarket al. 113900 (MO, U, US, VEN); San Carlosde Rio Negro, ca. 20 km S of confluenceof Rio NegroandBrazoCasiquiare, Mar1983(? fl-fr),Uht 457

1988 (e), Wallnoefer16-81088 (BG). LORETO: Rio

Corrientes,upstreamof Tnte. L6pez Oil Camp, 3 Apr 1979 (? fl), Gentry et al. 18956 (MO, U); Prov. Maynas, Yanomono, 19 Feb 1981 (e?), Gentry et al. 31470 (MO, NY, U); Mishuyacu, nr. Iquitos, Oct-Nov 1929 (e), Klug 153 (F, NY, US); Prov. Maynas, Casario Santo Tomas, Rio Momon, 29 Oct 1987 (a'), Ruiz et al. 1090 (BG); Dtto. Pebas, Brillo Nuevo, 11 May 1981 (? fl-fr), Treacyet al. 372 (F, WIS); Prov. Maynas, Rio Nanay, Mishana 25 Oct 1980 ('), R. Vdsquez et al. 609 (BG, MO); Prov. Alto Amazonas, Shucushuyacu (Rio Huallaga), 14 Sep 1981 (d'), R. Vasquezet al. 2499 (BG,MO, NY); Prov. Requena, Dtto. Saquena,Rio Ucayali, nr. JenaroHerrera,7 Feb 1979 (a'), Rimachi Y 4229 (MO). SAN MARTiN: Rio Huallaga, Chazuta,Mar 1935 (? fl), Klug 3984 (BM, F, GH, K, MO, NY, US). UCAYALI: Prov. Coronel Portillo, rd. PucallpaTingo Maria, Bosque Nacional Alexander von Humboldt, 11 Sep 1980 (o), Oliveira 31 (MO); jct. of Rio Pachitea and Rio Yuyapichis, SE of Pucallpa, 3 Oct 1985 (e'), Morawetz et al. 11-31085 (BG). BRAZIL. ACRE: Rio Yaco, nr. mouth of Rio Macaua, 3 Aug 1933 (e), Krukoff 5250 (A, BM, F, G, K, MICH, MO, NY, NY, U, US), 8 Aug 1933 (? fl), Krukoff 5352 (A, BM, F, G, K, MICH, MO, NY, U, US); Rio Abuna, (MO). BOLIvAR: Mun. Foraneo Aripao, Rio Caura, 1 km Seringal Orion, 23 Oct 1923 (9 fl-fr), Kuhlmann 708 = belowAraguania,I May 1988 (? fl),Aymardet al. 6743 RB 19821 (MO, U); 4 km W of Sena Madeira, 27 Sep (BG, MO,NY); middlepartof Rio Paragua,13 May-13 1968 (9 fl-fr), Prance et al. 7616 (F, GH, INPA, MG, Jun 1987 (? fl), Stergios 10389 (MO). NY, P, U, US); Rio Jurua, Jurua-Mirim, Sep 1901 (e), SURINAME.BakhuisMts., CoppenameR., 11 Feb Ule 5869 (A, B, F, G, K), (9 fl-fr),Ule 5870 (A, B, F, G, 1965(? fl-fr),Florschuitz et al. 2805(NY,U, US);Sipaliwini K). AMAZONAS: Rio Javary, Esperanca, 5 Oct 1942 (9 savanna,SipaliwiniR., 22 Jan 1970 (a'), Oldenburger et fl-fr), Ducke 1040 (MG, MO, NY, US); Sao Antonio do al. 1127 (U, MO),Feb 1970(a'), Oldenburger et al. 1402 Ica, 16 Sep 1906 (c), Ducke MG 7670 (BM, G, U); (U); upperSurinameR., nr.Goddo,20 Jan 1926(? fl-fr), Manaus, 27 Dec 1932 (a), Ducke RB 25256 (MO, U); Stahel(Exp. Wilhelminagebergte) 143 (U). Rio Negro, Sao Gabriel, P6rto de Curucuhy, 27 Apr ECUADOR. MORONA-SANTIAGO: Taisha, Rio 1947 (e), Fr6es 22163 (IAN, U); Mun. Borba, nr. Bella Guaguayme, 27 Jan 1962 (? fr), Cazalet et al. 7540 (K, Vista, 4-6 Sep 1934 (? fl), Krukoff 5977 (A, B, BM, F, NY, US). NAPO: Rd. Coca-Auca, km 50-60, 28 Feb G, GH, K, LE, MICH, MO, NY, U, US), (e), Krukoff 1980(st), Berg et al. 1097 (BG);Rio Napo,9 km down- 5982 (A, B, BM, F, G, GH, K, LE, MICH, MO, NY, U,

117

TAXONOMICTREATMENT US); Mun.Humaita,nr.Livramento,12 Oct-6 Nov 1934 (? fl-fr),Krukoff6988 (A, B, BM, F, G, K, MICH,MO, NY,U, US); Sao Paulode Oliven9a,11 Sep-26 Oct 1936 (? fr), Krukoff8520 (A, B, BM, F, G, LE, MICH,MO, NY, P, U, US); Rio Madeira,Democracia,31 Aug 1923 (e), Kuhlmann256 (MO, U); mouthof Rio Branco,5 Jan 1924 (? fl-fr), Kuhlmann 1103 (MO, U); Rio Javary,

PalmeirasArmyPost, 2 Aug 1973 (? fl-fr),Lleraset al. P.17067 (MO, U); Rio Negro, Ilha Tamandua,19 Oct 1987 (? fl-fr), Maas et al. 6788 (BG); Rio Negro, nr. Manaus, Dec 1850-Mar 1851 (Y fl-fr), Spruce s.n. (B, BM, C, G, K, LD). MATO GROSSO: Rio Aripuana, nr.

kusuknash(Huambisa);Loreto:mashonastillo;Pasco: comawantzaui.Bolivia: Beni: xanajoxo (Chacobo).

6b. Sorocea muriculata Miquel subsp. uaupensis (Baillon) C. C. Berg, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med. Sci. 88: 387, t. 3. 1985. Pseudosorocea uaupensis Baillon, Adansonia 11: 297. 1875. Sorocea uaupensis (Baillon) Macbride,Candollea 5: 349. 1934. Type. Brazil. Amazonas:Rio Uaupes,nr.Panure,Nov 1852 (a), Spruce2715 (holotype,P; isotypes,B, BM, F (fragFig. 23 ment), G, K, LE, US).

HumboldtCenter,13 Oct 1973 (e), Berg et al. P18468 (INPA,MO, U), 14 Oct 1973 (? fl), Berg et al. P 18549 (INPA,MO, NY, U); Rio Machado,nr. Tabajura,NovSorocea guayanensis W. Burger,Acta Bot. Neerl. Dec 1931(? fr),Krukoff1314 (A, B, BM, G, K, MICH, 11: 439, t. 2. 1962. Type.Venezuela.Amazonas: MO,NY, P, U); Rio Aripuana,nr. HumboldtCenter,10 San Carlosde Rio Negro,28 Feb 1942 (? fr),Ll. Oct 1973 (de),Pranceet al. 18349 (INPA,MO,NY, U); Williams 14532 (holotype,F; isotypes,A, G, IAN, Mun.Alta Floresta,17.5 km N of Alta Paraiso,30 Sep NY, US, VEN). 1985 (d), Thomaset al. 4128 (BG, MO). PARA: Rio 14 Dec 1906(v), DuckeMG 7959 (BM, Cumina-Mirim, Perianth of the staminate flower minutely F, G, GH, MG);Rio Tapaj6s,5 Oct 1922 (e), DuckeRB puberulousoutside. Pistillateflowers 3-7(-9) per in18318 (U); Rio Tapaj6s, Vila Nova, Cachoeira Chacoro, florescence; upper part of the perianthcylindrical to 18 Dec 1951 (? fl-fr), Pires 3539 (IAN, US); rd. Cuiabaovoid, almostas broadas the lowerpart,both narrowly Santarem, km 1056, 14 Nov 1977 (? fl-fr), Prance et Fruiting perianth cylindrical, parts puberulous. al. 25401 (MO,NY, U), km 1135, nr. IgarapeNatal, 16 oblongoidor sometimesellipsoid, 1-1.2 x 0.5-0.6 cm, Nov 1977 (? fl-fr), Prance et al. 25508 (MO, RB, U). ROND6NIA: P6rtoVelho,Estradado Dique,11 Aug 1987 red, turningblack (or darkred?), the apex sometimes (? fl-fr), Dionizia et al. 109 (NY); Mun. P6rtoVelho, curved, the surface smooth and puberulous;pedicel rd. Vila Caneco-Minera9aoJacunda,BR-164, km 106, usuallyratherslender, 1-2 mm thick, but occasionally 26 Sep 1979 (? fl-fr), Vieira et al. 291 (MO, U); to 4 mm thick,cylindricalandregularin outlinein fruit. MineraqaoJaboca, 10 Oct 1979 (? fl-fr), Vieiraet al. Distribution (see Fig. 77.2). In southeasternVen375 (MO,U); Estradados Padres,14 km fromJi-Parana, 21 Oct 1979 (? fl), Vieira et al. 571 (MO, RB, U). ezuelaandadjacentBrazil,also in SurinameandFrench RORAIMA: Rio Branco,Paranado Marara,4 Mar 1977 Guiana;in non-inundatedforest;to ca. 1200 m. (e), M. R. Santos 131 (NY). BOLIVIA.BENI: Prov.VacaDiez, Alta Iv6n, 8

Dec

1983 (? fr), Boom 4109 (NY); 17 km S of Riberalta, 23 Nov 1983 (? fl-fr), Boom 4030 (NY, U, US), 18 Nov 1989 (? fl-fr), Daly et al. 6243 (BG, USZ); Rio Beni,

Jun 1921(? fl), Meijer102 (NY,U, CachuelaEsperanza, Z); confluenceof Rio Beni and Rio Madrede Dios, Aug 1886 (e), Rusby 2217 (F, GH, K, MICH, NY, US); Prov.

km 20.8, 25 Sep VacaDiez, rd. Riberalta-Guayaramerin, 1981 (d), Solomon6426 (GB, MO, NY). PANDO: Prov. 5 Oct Manupiri,12 km W of rd. Conquista-Trampolin, 1991 (Y fl-fr), Beck et al. 20154 (LPB); Prov. Manupiri,

Conquista,rd. PuertoRico-RioMadrede Dios, km 18, 8 Oct 1991 (? fl), Becket al. 20254A(BG, LPB). The collection Steyermarket al. 11390 shows features intermediate between subsp. muriculata and subsp.uaupensis.The sterilecollection FD 7629 from Guyana (summit of Mt. Faiafun) might belong to subsp. muriculata.

Representativespecimensexamined.VENEZUELA. AMAZONAS:upper Rio Orinoco, between Ocamo and

et al. 7361 Mavaca,10-23 Jan1970(? fl-fr),Aristeguieta (MO, NY, VEN); Santa Barbaradel Orinoco, 27 May 1975 (? fl-fr), Berry752 (MO); Sierra Parima, Rio Siapa, 1123 Mar 1946 (a), Cardona1344 (F, NY, US); Depto. Atabapo, upper part of Cafio Caname, 3 May 1979 (? flfr), Davidse et al. 17136 (MO); Depto. Atabapo, Rio Putaro, Raudal Xikita-pora, Feb 1990 (? fl), Fernandez 7150 (NY); trail S from Cerro de La Neblina Camp 5,

1200-1300m, 12 Apr 1984(st), Gentryet al. 46552 (BG, MO);SanCarlosde RioNegro,ca. 20 km S of confluence of Rio NegroandBrazoCasiquiare,6 Apr 1979(? fl-fr), Liesner6275 (MO);Depto.Atabapo,base of Duida,nr. Culebra, on Rio Cunucunuma, 23 Feb 1985 (st), Liesner 17840 (MO); Depto. Atabapo, slope of Huachamacari, 5-6 Mar 1985 (a'), Liesner 18285 (MO); Depto. Atures,

20 km SE of PuertoAyacucho,21 Jun 1984(? fr),Miller

1593 (BG, MO, NY); Depto. Atures, 44-45 km SE of Puerto Ayacucho, 9 May 1980 (9 fl), Steyermark et al. Local names. Venezuela:Amazonas:palo de brujo. 122172 (MO, NY); Depto. Atabapo, upper Rio Orinoco, Ecuador:Napo:ayamoyegui(Huaorani),tapaguemohue Ocamo (Santa Maria de los Guaicas), 15 Feb 1981 (e), (Huaorani),tapoguene(Huaorani),tentemoeyi(Huao- Steyermarket al. 124341 (MO, NY); Mt. Duida, Aug 1928rani).Peru:Amazonas:kusuknashikumpari,shitapach Apr 1929 (a), Tate 947 (US); Isla de Rat6n, Jun 1940 (?

118

FLORANEOTROPICA

Jun fl-fr), LI. Williams13443 (VEN, US); Rio Sanariapo, 1940 (9 fr), LI. Williams 13488 (F, K, US, VEN); Rio Orinoco,SanAntonio,28 Apr 1942 (? fl-fr), Ll. Williams 15079 (US);uppermouthof Rio Casiquiare, 5 May 1942 (? fr), LI. Williams 15184 (F, G, MICH,US, VEN);Rio Yatua,PiedraArauicauca,15 Jul 1959 (? fl-fr), Wurdack et al. 43455 (NY,U, US). BOLIVAR:Dtto. Heres,90 km S of La Paragua,12 May 1987 (? fl), Aymard C. 5777 (MO,NY); Rio Parguaza,CainoBejuco, 15 Apr 1946 (? fl), Badillo 1424 (VEN),(e^)Badillo 1431 (VEN);Mun. Raul Leoni, 58 km SE of Pijiguaos, Jul 1989 (? fl), Delgado 375 (BG,MO);Mun.Sucre,RioAriza,May1990 (? fl), Delgado 964 (MO);Mun.Raul Leoni,upperRio Paragua,25 km N of Macizo Ichuin,Apr 1988 (a), A. Fernandez 4301 (MO);CanloPablo,ca. 6 km ESEof Las Pavas(Salto Para),8 May 1982 (9 fl-fr),Liesner et al. 13934 (MO, NY); middle Rio Paragua,nr. "Minasde Manaima,"13 Jun 1987 (9 fl-fr), Stergios 10225 (BG, NY,MO);Rio Canaracuni, 13-26Apr1988(? fl), Stergios 11734 (BG, MO, NY), (a), Stergios 11840 (BG, MO); Rio Caura,ca. 2 km belowCafioGuacamaya,13-26 Apr 1988 (9 fl), Stergios 12069 (BG, NY); Mun. Sucre, Boca

de Nichare,May 1988 (d'), Velazco 203 (BG, MO). SURINAME.NatureReserveVoltzberg,23 Feb 1977 (? fl-fr), Lindeman & Mennega et al. 125 (U); Oelemari R., nr. airport,24 Mar 1963 (9 fl), Wessels Boer 1069 (NY, U, US). FRENCHGUIANA.Paul Isnardregion,nr. Citron, 12 Feb 1983 (9 fl-fr), Feuillet 715 (CAY). 11 Jul 1979 BRAZIL. AMAZONAS: SerraCuricuriari, (9 fr), Alenqar 554 (MO,NY, U, US); Rio Negro,Serra do Sao Gabriel, 1 May 1947 (v), Pires 555 (NY, US); SerraCuricuriari, 10 Jul 1979(9 fr),Poole 1939 (NY,U, US); Mun. Uaupes, Rio Negro, 29 Oct 1962 (9 fl), Rodrigues et al. 4743 (INPA,U); Rio Negro, Cucui, 6 May 1973 (9 fl-fr), M. F Silva et al. 1275 (RB); Rio Maturaca,MissaoSalesiana,1 Jan 1966 (e), N. T Silva 60808 (NY,U, US). RORAIMA: Rio Uraricoera, Cachoeira Tocuxema,9 Mar 1979 (e), Pires et al. 16894 (NY). The three collections from Surinameand French Guianadeviatesomewhatfromthe othercollectionsof subsp. uaupensis. Additional collections may clarify the statusof these threecollections. Local names. Venezuela: Amazonas: fnepo (Yanomani);Bolivar:cafecillo.

pasa trail, 29 Oct 1901 (? fl-fr), R. S. Williams 412 (holotype,NY; isotype, US). Tree,to 15 m tall or shrub.Leafytwigs 1-4(-5) mm thick, sparsely to densely and minutely puberulous. Laminaoblongto lanceolate,4.5-17(-42) x 1.5-6(-15) cm, broadestat or abovethe middle,almostequilateral, (sub)coriaceous;apex acuminate,the acumenacuteto obtuse;baseacuteto obtuseto rounded;margindistinctly to obscurelydent(icul)ate(to subentire);uppersurface often shining, glabrous;lower surfacesparselypuberulouson themainveins,theglobose-capitate, pluricellular hairsabundantto sparse;venation almost plane or the midribslightly impressedabove, ? prominentbeneath; lateralveins 8-18 pairs;tertiaryvenationmostly partly scalariform;petiole 0.4-1.2 cm long, 1-2.5 mm thick, sparselypuberulousor glabrous;stipules 0.3-0.6 cm long, sparselypuberulousto glabrous,caducous.Staminate inflorescencespatent, 1-6 cm long, includingthe 0.1-0.2 cm longpuberulouspeduncle;flowerscrowded, narrowlyto ratherbroadlysessile; perianthca. 1.5 mm long, deeply 4-parted, sparsely puberulous or only ciliolate;stamens4, sometimesanisomporphic,straight with the anthers? transverseor bentoutwardsat anthesis; filaments0.5-1.5 mm long; anthers0.5-1 x 0.4-0.8 mm, the connectiverathernarrow,not or very shortly apiculate.Pistillateinflorescencessometimeson theolder wood, patent,0.5-2.5 cm long, to 7 cm long in fruit; peduncle0.1-0.5 cm long, to 1 cm long in fruit,puberulous; rachispuberulous;flowers I 0-ca. 40, sessile or subsessile,pedicellatein fruit;pedicelto 1(-1.2) cm long, minutelypuberulousto muriculate;perianth1.5-2 mm long, subentire,the upperpartbroadlyovoid to almost discoid, glabrous,the lower partwith dense, globosecapitate,pluricellularhairs;stigmas narrowlytongueshaped, ca. 1-1.5 mm long, finely papillate.Fruiting perianthellipsoid(to ovoid),0.8-1.3 x 0.6-1 cm,chromeyellowto orangetoredorpurple,(finallyblack?),theapex truncatewith a collar-like rim, or obtuse, the surface smoothwithdense,globose-capitate,pluricellarhairs. Distribution (see Fig. 77.4). In Amazonian Peru and Bolivia, extending to Brazil (Acre); in non-inundatedforest;at low elevations. Representative specimens examined. PERU.

7. Sorocea briquetiiMacbride,Candollea4: 311. 1931; Macbride, Publ. Field Mus. Nat. Hist., Bot. Ser. 13(2.2): 311. 1937; Burger, Lanjouw & Wessels Boer, Acta Bot. Neerl. 11: 441, t. 2. 1962. Type. Peru. San Martin:nr.Tarapoto,Jun 1855 (? fl-fr), Spruce 4220 (holotype, G; isotypes, B, BM, C, F, GH, K, LD, LE, P, S). Fig. 24

HUANUCO:Prov.PuertoInca, Dtto.Yuyapichis,Unidad Modelo DANTAS, 1 Dec 1988 (a), Kroell S. 187 (BG), 4 Dec 1988 (9 fl-fr),Kroell S. 203 (BG); nr. confluence of Rio Cayumba with Rio Huallaga, 9 Oct 1936 (e), Mexia 8260 (BM, F, GB, GH, K, MO, NY, U, US); Rio

Monzon,Tazo Grande,21 Sep 1965 (? fl), SchunkeV 871 (F, MOL, NY, US), 22 Sep 1965 (e), Schunke V 884 (F, MOL, NY, US); Prov. Pachitea,ca. 26 km S of

Puerto Inca, 3 Nov 1988 (a' fl), Wallnoefer 11-031188 (BG). JUNMN: Chanchamayo, 5-10 km S of San Ram6n, t. 2. 1962. Type.Bolivia.La Paz:Ixiamas-Tumu- 1000-1500 m, 5 Sep 1982 (st), Foster 8514 (U); Prov.

Sorocea pileata W. Burger,Acta Bot. Neerl. 11: 443,

TAXONOMICTREATMENT

119

Fig. 24. Sorocea briquetii. 1. Leafy twig with young staminateinflorescences(Mexia 8260). 2. Leafy twig 7088).4. Leafytwig with infructescence (Klug4161). 3. Leafytwig with youngpistillateinflorescences(Woytkowski (SchunkeV 3365). 5. Staminateflower.6. Stamens(Mexia8260). 7. Pistillateflower.8. Fruitingperianth.9. Seed. 10, lOa. Embryo.11. Radicle and small cotyledon (SchunkeV 3365). [By P. Pardoen,Utrecht.]

120

Satiopo,Dtto. Pangoa,PuertoOcopa, 13 May 1981 (st), Reynel R. 251 (MOL,U). LORETO: Prov.Uacayali,Dtto.

VargasGuerra,Orellana,27 Nov 1982 (st), ReynelR. 932 (MOL);Rio Yaguasyacu,Brillo Nuevo, ca. 150 km ENEof Iquitos,13 Nov 1981 (st), Treacyet al. 534 (F); Prov.Ucayali,Canchahuayo, Rio Ucayali,25 Nov 1985 (e), R. Vasquez et al. 6914 (BG, MO, NY). MADRE DE

FLORA NEOTROPICA

Prov.Oxapampa,GranPajonal,nr. Chequitaro,1250 m, 23 Sep 1983, with juvenile inflorescences, could not be placed with certaintyin this species as the midrib is minutelypuberulousabove. The presence of a white mycelium on the perianth of the pistillateflower is very common in this species. Fruits(still unripe)have been reportedto be eatenby Cebusalbifrons(Foster 5225).

DIos: TambopataCommunidadNativa de Infierno, HermosaChica,22 Nov 1988 (st), Alexiadeset al. 121 (BG, MO) and 200 (BG); Prov.Tahuamanui, Dtto. IbeLocal names and use. Peru: Junin: sintaque; km 32, 1 Dec 1981 (e'), Diaz S. ria, rd. Iberia-Iniapari, 1730-95 (BG, MO, MOL);ParqueNacionaldel Manu', Loreto: yanchama;Madre de Dios: capinuri blanca, chatojioji, etona-tewe (Ese-ejhal), manchinguilla, Cocha Cashu Station, 23 Sep 1976 (a), Foster et al. 5060 (BG), 18 Nov 1976 (? fl-fr), Foster et al. 5210 (F); tamamuri;Ucayali:yanchama.Brazil:Acre:jaca brava. Tambopata NatureReserve,jct. of Rio La Torreand Rio In Peru(Madrede Dios) a decoction of the latex is 24 May 1987 (st), Gentryet al. 57537 (BG, used against fever. Tambopata, MO). SANMARTiN: UpperRio Huallaga,Juanjui,Dec 1935 (? fl), Klug 4161 (A, F, GH, MO, NY, U, US); Prov.MariscalCaceres,3 km fromTocacheNuevo, 21 8. Sorocea trophoides W. Burger,Acta Bot. Neerl. Aug 1969 (? fr),SchunkeV 3365 (F, GH,K, MO,MOL, 11:450, t. 6. 1962; Burger,FieldianaBot. 40: 210. NY, US), 4 Jul 1974 (? fr), SchunkeV 7154 (MO,NY); 1977; Berg & Akkermans,Proc. Kon. Ned. Akad. upperRio Huallaga,28 Dec 1929 (9 fl-fr),LI. Williams Wetensch., Ser. C, Biol. Med. Sci. 88: 389. 1985. 6803 (F, G); Juanjui,9 Mar 1962 (9 fl-fr), Woytkowski Type. Costa Rica. Alajuela: CantonAlfaro Ruiz, 7088 (GH, MO). UCAYALI: El Sacramento,89 km S of Pucallpa,22 Jul 1957 (st), Ellenberg2463 (U). Zarcero, 17 Jul 1937 (? fl), A. Smith 141 (holoBRAZIL. ACRE: UpperRio Moa, FazendaArizona, type, MO; isotypes, F, U). Fig. 25 10-16 Oct 1985 (st), Campbellet al. 6522 (BG);Mun. Sorocea rhodorachis Cuatrecasas, Ciencia (Mexico) Brasileia,rd. to Assis Brasil,km 16, 2 Nov 1980 (? fr), 24(5/6): 188. 1966. Sorocea trophoides W. Burger Ferreiraet al. 3093 (MO,NY,US);rd.Rio Branco-P6rto subsp. rhodorachis (Cuatrecasas)C. C. Berg, Proc. Acre, km 39, 13 Oct 1980 (6'), Lowrieet al. 510 (NY); Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med. rd. Rio Branco-Quixada,km 11, 18 Oct 1980 (9 fl-fr), Sci. 88: 389. 1985. Type. Colombia. Valle: Rio Nelson 730 (BG, MO);Mun. SenaMadureira, Rio laco, Fazenda Nova Olinda, 29 Oct 1993 (e), M. Silveira et al. 688 (NY). BOLIVIA. BENI: Rurrenabaque, 23 Nov 1921 (9

Dagua, Piedra de Moler, 19 Aug 1943 (fl-fr), Cuatrecasas 14891 (holotype, F). Sorocea martineziana Cuatrecasas, Ciencia (Mexico) 24(5/6): 187. 1966. Type. Colombia. Valle: Rio Sanquinini, La Laguna, 16 Dec 1943 (? fl-fr), Cuatrecasas 15572 (holotype, F; isotype, VALLE).

fl-fr), M. Cdrdenas1757 (F, NY, US) Prov.Ballivian, rd. Yumuco-Rurrenabaque, km 35, Mar-Jul 1990 (st), D. N. Smith et al. 14387 (MO). LA PAZ: Prov. Abel Iturralde,Alto Madidi,25-27 May 1990 (st), Gentryet Tree,to 15 m tall. Leafiytwigs 1-3(-4) mm thick, al. 70559 (MO); Prov. Franz Tamayo, Serrania de minutely puberulous.Lamina elliptic to oblong, 4.5Chepite, CampoSismico de Texaco, 15 km W of Rio 12(-35) x 2-6(-9.5) cm, broadest at the middle, ? Tuichi,3-8 Apr 1992 (6'), Killeen 3726 (LPB). SANTA inequilateralto equilateral,subcoriaceousto chartacous; CRUZ: Prov. Ichilo, Parque Nacional Ambor6, Rio apexacuminateto caudate,the acumennarrow,acuteto Macuiiucu,5 Dec 1991 (9 fr), Nee 41935 (LPB). obtuse;base acuteto obtuse (to rounded);margin(serSoroceabriquetiiresemblesS. trophoidesin theshape rate-)dentateor (sub)entire;uppersurfaceof the midandmarginof the laminaandin the long pedicel of the rib minutely puberulous or glabrous; lower surface pistillateflower in fruitingstate. It is likely thatS. bri- (sub)glabrous;venationalmostplane, but the midribi quetii is more closely relatedto S. trophoidesthanto impressedabove, the venation ? prominentbeneath; the groupofAmazonianspecies constitutedby S.pubi- lateralveins 6-10(-14) pairs;tertiaryvenationreticuvena, S. muriculata,and S. steinbachii. lateorpartlyscalariform; petiole0.5-1.5(-2.5) cm long, Two forms occur within the species. A form with 1-2(-3) mm thick, sparselypuberulous;stipules 0.3relatively large leaves and large fruiting perianths 0.8 mm long, puberulousto glabrous,caducous.Stami(longerthan 1 cm) is representedby the collectionsfrom nate inflorescences pendulous (?), (1 .5-)2.5-6.5 cm Madrede Dios (Peru)andBolivia. Anotherform with long, including the 0.2-0.3 mm long puberulouspesmallerleaves and fruitingperianthsis representedby duncle; flowers crowded, sessile or (in Mexico and the other Peruviancollections cited. Intermediatesin Guatemala)with pedicels to 2 mm long; perianthca. leaf size occurin Bolivia andBrazil(Acre).The collec- 1.5-2 mm long, 4-parted, ciliolate, stamens (3 or) 4, tion D. N. Smith 5172 (BG, MO) from Peru, Pasco, usually (S-shaped)curvedwith the antherstransverse;

TAXONOMICTREATMENT

121

;

'J

(Burger Fig. 25. Soroceatrophoides.1. Leafytwig with staminateinflorescences(Pittier3045). 2. Infructescence et al. 9585). 3. Leafy twig with infructescences(Garcia et al. 431). 4. Leafy twig with staminateinflorescences (Uribe-Uribe1221). 5. Staminateflower(A. Smith4227). 6. Staminateflowerand bract.7. Youngstamens(Klug 3566). 8, 8a. Pistillateflower (Cuatrecasas14891). 9, 9a. Pistillateflower. 10. Fruitingperianth(Raven21775). [By P. Pardoen,Utrecht.]

122

filaments2.5-3 mmlong,mostlydistinctlylongerthan thetepals;anthersca.0.8-1 x 0.7-0.8mm,theconnective rathernarrow,apiculate.Pistillate inflorescences

2-4 cmlong,to 10cmlongin patent(topendulous?), rachispuberfruit;peduncle0.3-1cmlong,puberulous; ulous,brightyellowto redin fruit;flowersca. 10-40, infruit;pedicel1.5-2cmlong, (sub)sessile, pedicellate red(?),almostglabrousinfruit;perianth pileate,ca.2.5 mm long,4-lobed,almostglabrous;stigmastongueshaped,ca. 1mmlong,finelypapillate.Fruitingperianth(sub)globose, 0.8-1 x 0.8-1 cm,red,finallyblack, theapextruncatewitha low collar-likerimto obtuse, thesurface? ridged,almostglabrous. Distribution(see Fig. 77.6). In southernMexico inCostaRicaandwestandadjacent partof Guatemala, ernPanama,andfromColombiato Peru(SanMartin); in wetmontaneorlowlandforest;in thenorthempart atlowelevations,in CostaRicaand of thedistribution Panamausuallyat 1000-1800m, in SouthAmerica mostlyat 1000-2000m, butin somepartsof Colombiacommonatlow elevations(seebelow). Representative specimens examined. MEXICO. TABASCO:Mun. Teapa, rd. Teapa-Tacotalpa, km 13, 26 Mun. Jun 1980 (e), Cowan 3082 (NY, CAS). VERACRUZ: Minatitlan, Rio Uxpanapa, nr. Oaxaca border, 27 Sep 1980 (c'), Wendtet al. 2764 (BG, CAS, MO, NY). GUATEMALA. IZABAL: Rio Tameja, 24 Dec 1941 (2 fl-fr), Steyermark 41801 (EAP, F, US). COSTA RICA. ALAJUELA: San Pedro de San Ram6n, ca. 1050 m, 24 Jul 1925 (? fl-fr), Brenes 4332 (F); Reserva Forestal de San Ram6n, 800-1000 m, 30 Nov 1986 (9 fl-fr), Herrera Ch. 202 (BG, MO), 30 Apr 1987 (e), Herrera Ch. et al. 561 (BG, F, MO); Upala, Bijagua, El Pil6n, 13 Nov 1987 (? fl-fr), Herrera Ch. 1261 (BG); Monteverde Reserve, PeniasBlancas, 800 m, 7 Oct 1986 (? fl-fr), Haber et al. 5667 (BG, MO); Cant6nAlfaro Ruiz, (region of) Zarcero, 1900 m, 22 Jun 1938, A. Smith 802 (A, F, NY), 1650 m, 10 May 1937 (d%),A. Smith 2720 (F, U), ca. 1750 m, 6 Jul 1937 (od), A. Smith 4227 (F, MICH, NY); Cant6n San Carlos, Villa Quesada, 750 m, 14 Apr 1939 (ei), A. Smith 1951 (F, NY, US). CARTAGO: El Mufleco, 1200 m, 9 Jul 1982 (9 fr), G6mez-Laurito8670 (F); Casa de Teja Ridge, above Rio Gato, 800 m, 15 Sep 1973 (V), Lent 3611 (F); nr. Finca Las C6ncavas, 7-8 Dec 1925 (st), Standley 41434 (F, US); El Mineco, Rio Navarro, 6 Mar 1926 (st), Standley et al. 51156 (US); CerroLa Carpintera,N of Tres Rios, E of San Jose, 16001800 m, 1 Jun 1984 ('), Taylor 3865 (DUKE). GUANACASTE: Parque Nacional Guanacaste, 1200 m, 2 Jun 1990 (od),Apu 27 (MO); SW of Volcan Rinear,Volcan La Vieja and Volcan Santa Maria, 1400 m, 30 Jul 1971 (st), Burgeret al. 7794 (CR, F); nr. Monteverde, 13001450 m, 20 Oct-2 Nov 1975 (2 fr), Burger et al. 9585 (CR, EAP, U). HEREDIA: San Jose de la Montafia, 1600 m, 8 Aug 1965 (? fl-fr), Hatheway 1489 (CR, F, US); nr. Cariblanco, Rio Sarapiqui, 800 m, 19 Apr 1949 (ei), L. 0. Williams16450 (BM, EAP, F). PUNTARENAS:Between

FLORA NEOTROPICA Lindora and Rio Guacimal, ca. 2 km WSW of Monteverde, 840-900 m, 26 Aug 1982 (9 fl-fr), Alverson 1879 (MO, WIS); E of Monteverde, 1300-1450 m, 29 Oct-2 Nov 1975 (9 fr), Burger et al. 9585 (F,MO, WIS); Monteverde, 1400 m, 10 Jun 1985 (), Grayum et al. 5397 (BG, MO); Cerro Chivo, 1600-1800 m, 17 Jul. 1983 (d'), Hazlett 5243 (F); 6 km S of San Vito de Java,Wilson's finca, ca. 1650 m, 14 Aug 1967 (? fr), Raven 21775 (BM, F, MO, U). SANJost: El Cedral de Santa Maria de Dota, 2000 m, 8 Mar 1987 (d'), Burger et al. 12090 (F); Cerros de Turrubares, 1550 m, 31 Mar 1990, Jimenez M. 825 (F, MO); El Cedral de Santa Maria de Dota, 12 Jun 1975 (? and d), Poveda s.n. (CR); nr. Santa Maria de Dota, 1500-1800 m, 26 Dec 1925-3 Jan 1926 (st), Standley et al. 43305 (F, US). PANAMA. CHIRIQUi: Las Lagunasarea, W of El Hato del Volcan, 1400 m, 23 Aug 1982 (? fr), Hamilton et al. 943 (MO, PMA); La Fortuna hydroelectric project, 20 Mar 1978 (9 fl-fr), Hammel 2051 (MO, U); Cerro Pelota, 1800-2000 m, 10 Oct 1981 (? fr), Knapp 1496 (MO, PMA, U); 8-10 km N of Los Planos de Hornito, 1250-1300 m, 9 May 1982 (e), Knapp 4990 (MO, U), 1150 m, 18 Jun 1982 (? fl-fr), Knapp et al. 5616 (MO, U); nr. Fortuna Dam, 1250 m, 28 Apr 1986 (? fl), McPherson 9097 (MO); nr. border with Costa Rica, rd. from Rio Sereno, 1400-1800 m, 12 May 1991 (? fl), McPherson 15300 (BG, EAP); nr. El Boquete, 10001200 m, 5 Mar 1911 (e), Pittier 3045 (BM, GH, NY, US). VERAGUAS: Above Santa Fe, Cerro Tute, 12001400 m, 28 Sep 1972 (9 fr), Gentry 6276 (MO). rd. El COLOMBIA. ANTIOQUIA: Mun.Sabanalarga, Oro-El Placer, 8 km from Sabanalarga, 1100 m, 8 Apr 1986 (9 fl-fr), Callejas et al. 2267 (BG, HUA, NY); Mun. San Luis, Rio Claro, 8 Jul 1983 (? fl-fr),Cogollo et al. 508 (HUA, MO), 16 Aug 1983 (6'), Cogollo 684 (MO),28 Jan 1984 (? fl-fr), Cogollo 1253 (MO), 30 Mar 1984 (d), Cogollo 1471 (MO), 29 Apr 1984 (? fl-fr), Cogollo et al. 1599 (MO), 2 Sep 1984 (? fl-fr), Cogolo 1917 (MO). CUNDINAMARCA:San Francisco,Vrda.Torriba,1550 m,

12 Jan 1944 ($ fl-fr), Garcia-Barriga10988 (COL, F, US). Serra de la Macarena, Vrda. El Tablazo, Morro Bello, 1000-1200 m, 28 May 1973 (? fr), Garcia et al. 431 (NY, U). QUINDfO: Mun. Calcara,Cgto. Quebrada Negra, 1 Oct 1987 (st), Arbeldez et al. 2289 (COL). META:

RISARALDA:Rd. LaVirginia-Cerritos, km 6, 6 May 1989

(e'), Silverstone-Sopkinet al. 5165 (MO), 16 Oct 1989 (? fl), Silverstone-Sopkin 5660 (MO); rd. Cerritos-Pereira, km 4, 27 Nov 1989 (? fl), Silverstone-Sopkin5716 (MO); rd. La Virginia-Cerritos, km 6, 1 Dec 1989 (? fl-fr), Silverstone-Sopkinet al. 5808 (MO) TOLIMA: "Gda. de la Plata,"1844(? fl-fr),Goudots.n. (K). VALLE: Mun. San Pedro, Cgto. Buenos Aires, La Siria, via Buenos Aires-Las Esmeraldas,1800 m, 14 Apr 1988 (o), Devia A. et al. 2103 (BG, TULV,US); Mun. Darien, 1700 m, 27 Jun 1988 (9 fl-fr), Devia A. 2158 (BG, TULV,US); Rio Frio, Finca El Porvenir, 1200 m, 5 Apr 1986 (st), Gentry et al. 54056 (MO); nr. Cali, La Cumbre, 1800-2100 m, 14-19 May 1922 (? fr), Pennell et al. 5778 (GH, NY, US), 1600 m, 24 Apr 1946 (a'), Uribe-Uribe 1221 (COL, US). ECUADOR. AZUAY:Nr. Molletura, 1200-1500 m, 11-14 Jan 1990 (e), Ortiz 167 (QCNE). BOLiVAR:San

TAXONOMICTREATMENT Pablode Telimbela,1000 m, 20 Nov 1943 (e), Acosta S. 6941 (F). COTOPAXI:Rd. Quevedo-Latacunga,km 55, Tenefuerte,24 Jul 1984 (st), Dodson et al. 14382 (MO, QAME,QCNE).IMBABURA:Cotacachi,2000 m, 14 Apr 1992 (d), Palacios et al. 10040 (BG, QCNE). LOJA:Rd. Alamor-Cazaderos, just W of El Limo, 1400 m, 20 Feb 1985 (? fr), Harling et al. 22313 (BG, GB, QCA);rd. Chaguarpamba-Buenavista, km 2.8, 1100 m, 6 Feb 1995 (? fl-fr), Jiggins et al. 175 (LOJA).Los Rios: CerroSamama,Hda. Clementina,550 m, 4 Apr 1996 (? fr), Cornejo et al. 4926 (BG). MORONASANTIAGO:7-8 km N of Gualaquiza,1450 m, 16 Apr 1985 (Q fl), Harling et al. 24190 (BG, GB, QCA). PICHINCHA:Chiriboga,ReservaForestalLa Favorita, 1600-1800 m, Ceron et al. 8926 (QCNE); Maquipucuna,5 km E of Nanegal, 1550 m, 10 Feb 1991 (st), Gentryet al. 73121 (MO, QCNE);ReservaFloristicaEcol6gica Rio Guajalito,ca. 2.5 km NE of km 59 on old rd. Quito-SantoDomingode los Colorados,1800-

123

Neerl. 11:453. 1962;Burger,FieldianaBot. 40: 207. 1977.Type.Panama. Panami:nr.RioGrande, ParaisoStation,19Sep 1862(? fl),Hayes684(holotype,K;isotypes,BR,G,NY,P). Fig.26 Shrub or tree, to 10(-20) m tall. Leafy twigs 1-3

mmthick,minutelypuberulous. Laminaoblongto elliptic,(5-)7-23.5 x (1.5-)2-8.5 cm,broadestaboveor inthemiddle,equilateral to slightlyinequilateral, chartaceousto subcoriaceous (to coriaceous);apexacuminate;baseacuteto subobtuse; margindistinctlyto obscurelydent(icul)ate(to subentire);uppersurface glabrousor sparselyandminutelypuberulous on the midrib;lowersurfacesparselyandminutelypuberulouson themainveins;venationaboveplane,butthe midribimpressedabove,the venationprominent beneath;lateralveins7-12 pairs;tertiary venationreticulate(orpartly scalariform); petiole0.3-1.2cmlong,0.52200 m, 24 Apr 1992 (e), Jaramillo 14763 (GB, QCA, stipules0.2-0.5 NY). ZAMORA-CHINCHIPE: ParqueNacionalPodocarpus, 1.5mmthick,minutelypuberulous; cm long, puberulous to glabrous, caducous. Staminate GuarderiaBombuscana,1050 m, 11 Jul 1992 (V fl-fr), pendulous,1-10 cmlong,includingthe Freire-Fierro2280 (QCA);rd. Zamora-Gualaquiza, 4 inflorescences km from Los Encuentros,850 m, 26 Jun 1996 (? fl.), 0.1-0.5 cmlongpuberulous peduncle;flowersdistant Klitgaardet al. 278 (LOJA);rd. Zumba-Valladolid, be- torathercrowded,withpedicelsto 1(-4) mmlong,(in tween Palanumaand Palanda,1150-1200, 13 May 1985 someinflorescences sometimessubsessileornarrowly (? fr), Stein et al. 2777 (BG). sessile);perianth1.5-2 mmlong,4-fid to deeply4PERU. SAN MARTfN: Zepelacio, nr. Moyobamba, parted,sparselypuberulous andciliolate,orglabrous; 1200-1600m, Mar1934,(e'), Klug3566 (A, F, G, GH,K, stamens (3-)4, straight to bent inwards,isomorphic or MO,NY,US);Prov.Lamas,Dtto.Alonsode Alvarado, San filaments 0.5-1.5mmlong;anthers 0.JuanPacaizapa,rd.Tarapoto-Moyobamba, km 72, 1000- anisomorphic; Pistillate 1050 m, 6 Jun 1977 (? fl-fr),SchunkeV 9630 (GH,U). 0.8 x 0.4-0.7 mm,theconnectiveapiculate. inflorescences patent,0.5-4 cm long,to 5 cm longin The materialfrom CentralAmerica has usually a fruit;peduncle0.1-0.4 cmlong,to 4 cmlongin fruit, dent(icul)ate leafmarginandthemidribis minutelypuberrachispuberulous, infruit);flowpuberulous; (reddish? ulous andplane in the acumenabove. In materialfrom ers 5-25(-40), pedicellate(or subsessile);pedicels SouthAmerica,the leaf marginis entire,the midribis 0.05-0.2 cmlong,to 0.6 cm longin fruit,denselyand usuallyglabrousabove,andthemidribusuallyimpressed minutely puberulous; perianth ca.2 mmlong,subentire, in theacumenabove.Thetwo morphologicaltypeshave theupperpart(depressed)subgloboseto hemispheribeen treatedas distinctsubspecies(Berg& Akkermans, cal,glabrous,thelowerpartdenselypuberulous; stig1985), butconsideringthe totalvariationin the species, mas0.5-1.2 mmlong,coarselypapillate.Fruitingpepartlyfoundinmaterialrecentlyavailableforstudy,aswell rianthellipsoidtobroadly ovoidtosubglobose, ca.0.7-1 as the degreeof variationfoundin otherspecies,the dif- x 0.6-0.8 cm,orangeto red(nottumingblack?),the ferencesindicateddo notmeritdistinctionatthe subspeapex obtuseto umbonateto apiculate,the surface cific level. ThecollectionsfromGuatemalaandMexico smooth,forthegreater partrather denselyandminutely occurfaroutsidethemainareaof thespecies.Theydiffer puberulous (orsubglabrous).

somewhatin the shapeofthe laminafromthecollections fromCostaRicaandPanama,andthe staminateflowers aremorepronouncedly pedicellate. Moreover,allarefound atlow elevations,50-200 m. Thesecollectionscouldrepresentadistinctinfraspecific taxon.Inthecollectionsmade by Cogollo in Antioquia(Colombia)at 300-400 m (!), thelaminais relativelynarrow(oblongto lanceolate)and the inflorescencesareshort(1-2 cm long). 9. Sorocea affinis Hemsley,Biol. Centr.Amer.Bot. 3: 150. 1883;Hemsley,Biol. Centr.Amer.Bot. 5: t. 79. 1886; Burger,Ann. Missouri Bot. Gard.47: 121. 1960; Burger,Lanjouw& Wessels Boer,Acta Bot.

Distribution (see Fig. 77.5). From Nicaragua

(Zelaya)throughCostaRica(AlajuelaandPuntarenas) andPanamato Colombia(ChocoandAntioquia);in wetforest;to ca. 1200m. Representativespecimens studied. NICARAGUA. ZELAYA:RioAleman,Montafias de Esquipulasy Aleman,

27-29 Nov 1951 (? fl-fr), Shanket al. 4827 (EAP, F).

COSTARICA. ALAJUELA: Upala, Dos Dios, 5 km of Brasilia,30 Oct 1987(? fl-fr),HerreraCh.1043(BG); 2 km S of Canalete,12 Nov 1975 (2), Utleyet al. 3325 (F, MO). GUANACASTE: ParqueNacional Rinc6nde la Vieja,Hda.SantaMaria,10 Oct 1987 (? fl-fr),Herrera Ch.et al. 841 (BG). PUNTARENAS:BetweenRio Esquinas

124

FLORANEOTROPICA

'I

Fig. 26. Soroceaaffinis. 1. Leafytwig with staminateinflorescences(Woodsonet al. 1383). 2. Leafytwig with pistillateinflorescences.3. Leafy twig (Maas et al. 1746). 4. Leaf (Moriet al 3084). 5a-c. Staminateflowers.6. Stamens(Seibert464). 7. Staminateflowerand bract.8. Stamen(Cooper554). 9a, b. Pistillate flower. 10, 10a. Fruitingperianth(Dukeet al. 14067). [By P. Pardoen,Utrecht.]

TAXONOMIC TREATMENT and Palmar Sur de Osa, Tinoco Station, 16 Mar 1950 (a'), Allen 5492 (CR, US); Osa Peninsula, 5 km W of Rincon, 24-30 Mar 1973 (? fl), Burger et al. 8899 (BM, CR, EAP, F, NY, U). PANAMA.BOCAS DEL ToRo: 7.5 km fromChanguinola, 3 Aug 1976 (o), Croat 38099 (MO); above Almirante, 28 Nov 1971 (? fl-fr), Gentry2678 (MO); N of Rio Teribe, Duwebdulup Peak, 13 Apr 1968 (? fr), Kirkbride et al. 569 (MO, NY); Escudo de Veraguas Island, 8 Aug 1987 (e), McPherson 11442 (BG, MO) (? fl-fr), McPherson 11423 (BG); Chiriqui Lagoon, 6 Sep 1940 (? fl), Wedel 598A, 28 Sep 1940 (? fl-fr), Wedel 961 (GH, MO), 20 Nov 1940 (a), Wedel 1715 (GH, MO, US); nr. Almirante, Jan-Mar 1928 (e'), Cooper 554 (F, K, NY). CHIRIQUI: Burica Peninsula, Rabo de Puerco, 8 km W of PuertoArmuelles, 18 Feb 1973 (a'), Croat 21960 (CR, F, GH, MO); Quebrada Aleman, 8 mi N of Los Planes de Homito IRHE FortunaHydroelectricProject, 13 Mar 1982 (a'), Knapp et al. 4130 (MO); 7 km N of El Cope, 10 Sep 1977 (? fl), Folsom 5244 (U). COCLI: Rd. to Coclesito Logging Camp, 12 mi from Llano Grande,9 Dec 1983 (2 fl-fr), Churchill et al. 3992 (BG), 16 Dec 1983 (d), Churchill et al. 4117 (BG); El Valle de Ant6n, 23-27 Jul 1935 (a'), Seibert 464 (A, K, MO, NY); 15-20 km NE of La Pintada,towardsToabre, 15 Feb 1981 (? fl-fr), Sytsma et al. 3609 (U). COL6N: 1-3 mi W of Portobello, 7 Sep 1971 (? fl-fr), Gentry 1732 (MO, SCZ) and Gentry 1756 (F, NY, SCZ); Santa Rita Ridge, W of TransisthmianHwy., 16 Dec 1972 (? fl-fr), Gentry 6554 (MO); Gamboa, 21 Aug 1971 (? fl), Kennedy 1130 (F, MICH, MO), (o) 1133 (MO); Mosquere, 17 Sep 1974 (? fl-fr), Maas et al. 1746 (CR, F, MO, U); QuebradaAncha, 4 km E of Buena Vista, 21 Nov 1973 (Q fl-fr, Nee 7778 (GH, MO); Loma de la Gloria, nr. Fato, Jan-Aug 1911 (d), Pittier 3856 (BM, C, GH, NY, US); Rio Trinidad,Jul 1911 (? fl), Pittier 4034 (BM, GH, NY, US); nr. Palenque7-8 Aug 1911 (a), Pittier 4123 (BM, MO, NY). DARIMN:Nr. Santa Fe, Rio Sebana, 30 Sep 1961 (fr), Duke 4156 (GH, MO, US); Rio Pirre,20 mi W of El Real, 14 Jul 1962 (a'), Duke 5159 (GH, MO, PMA, US); Rio Paya, 31 Aug 1967 (? fr), Duke et al. 14001 (MO, NY) and 14067 (F, MO); between Punta Guayabo Grande and Punta Guayabo Chiquita, 25 Jan 1957 (? flfr), Sternet al. 180 (A, MO, NY, US); nr.El Real, Rio Tiura, 1 Jul 1959 (a), Stern et al. 770 (GH, LE, MO, US), ($ fl), BarroColoStern et al. 771 (GH, LE, MO, US). PANAMA: rado Island, 5 Sep 1929 (? fl-fr), Bangham 558 (A, F, US), 3 Aug 1971 (? fl), Croat 16545 (GH, MO, NY), 27 Oct 1967 (? fl-fr), Dressler 3136 (MO, PMA, SCZ, US); Rio Pita, 1-2 mi above confluence with Rio Maestro, 14 May 1961 (? fl-fr), Duke 4744 (GH, MO); nr. El Llano, 14-19 Oct 1962 (e), Duke 5836 (GH, MO, US); S of Ipeti, 1 Apr 1982 (2 fr), Knapp et al. 4539 (MO, PMA, U); Cerro Campana, 45 km W of Panama City, 8 Aug 1975 (? fl), Mori et al. 7693 (MO, U); Rio Mosombi Grande, 1-5 km NW of Summit Gardens, 12 Sep 1973 (? fl-fr), Nee 6908 (CR, LD, MO, PMA, WIS); nr. Arraijan,21 Jul 1938 (a), Woodson et al. 1383 (A, F, MO, NY, NA). SAN BLAS: Chucunaque, 2-10 mi above Cuna/Darien boundary, 21 Aug 1966 (Y fl-fr), Duke 8575 (MO); between Cangandi

125 and Mandinga Indigena, 25 Oct 1967 ('), Duke 14754 (MO, NY); between Rio Diablo and Rio Acuati,nr.Nargana, val3 Nov 1967 (? fl-fr), Duke 14871 (MO). VERAGUAS: ley of Rio Dos Bocas, between Alto Piedraand Calovebora, 29 Aug 1974 (2 fl), Croat 17435 (MO); ca. 4 mi above Santa Fe, 13 Dec 1971 (? fr), Gentry 3078 (F, MO); Alto Piedra, Santa Fe, 1200 m, 27 Sep 1972 (9 fr), Lao 517 (MO); 5 mi NW of Santa Fe, 700-1200 m, 18-19 Mar 1973 (q fr), Liesner 969 (F, GH, MO, US); valley of Rio Dos Bocas, NW of Santa Fe, 14 Nov 1974 (? fr), Mori et al. 3084 (MO, U); W of Santa Fe, CerroTute, 15 Dec 1981 (Q fr), Knapp et al. 2472 (PMA, U). COLOMBIA. ANTIOQUIA:Mun. Turbo, rd. Tap6n del Darien, between Rio Le6n and Loma Aisladas, 28 Aug 1983 (e), Brand et al. 465 (MO), 26 May 1984 (? fl-fr), Brand 1186 (MO), 10 Jan 1985 (R fl-fr), Brand et al. 1282 (HUA, MO); Mun. Mutata, Rio Mutata, Finca Puentiadero-Finca La Palma, 5 May 1987 (? fl-fr), Fonnegra et al. 2188 (BG, MO). CHOC6: Rio Chintado, N of La Nueva, 6 Feb 1967 (a'), Duke 9875 (A, MO); between Teresita and Rio Salada, 4 Jun 1967 (e), Duke 12214 (US); Mun. Acandi, Cgto. de Unguia, rd. DarienQuebrada del Lim6n, 16 Jun 1976 (? fl-fr), Forero 427 (MO), 27 Jun 1976 (e'), Forero 429 (MO); Mun. Acandi, Cgto. de Unguia, Cuna de Arquia, 10 Jul 1976 ($ fl), Forero 615 (MO, TULV); Rio Atrato, Tagachi, 8 Apr 1982 (? fl), Forero et al. 8979 (COL, MO); Mun. Riosucio, Parque Nacional Los Katios, Cararicas, 16 Jul 1976 (Q fl), Le6n 383 (MO); Mun. Acandi, mouth of Rio Cuti, 25 Jul 1957 (a), Romero-Castafneda6427 (COL, GH, MO).

Soroceaaffinisis variablein severalcharacters.The laminais mostly chartaceousto subcoriaceouswith a dentatemargin,but it can be entireandcoriaceousand then quite similarto thatof S. ruminata.The midribis eitherglabrousor sparselypuberulousabove.Thestaminate flowers are mostly pedicellate, but they can be subsessileornarrowlysessile in the northernpartof the species range (see, e.g., Allen 5492). The collection Seibert464 showstransitionsfrompedicellateto sessile flowers. The collections with pistillate inflorescences made by Brand et al. in Colombia (Antioquia)have a subglabrousfruitingperianthand are,therefore,difficult to distinguishfrommaterialof S. trophoidesfrom the same region. Sorocea affinis appearsto be closely relatedto the geographicallyseparatedS. sarcocarpa.

10. Sorocea sarcocarpaLanjouw& WesselsBoer,Acta Bot. Neerl. 11: 452, t. 7. 1962. Type. Ecuador. Esmeraldas:4 km W of Borbon,26 Apr 1943 (V flfr),Little 6375 (holotype,F; isotype, K). Fig. 27 Tree, to 20 m tall. Leafy twigs 1-3.5 mm thick,

sparselypuberulous.Lamina(sub)obovate,7-23 x 3.510.5 cm, slightlyinequilateral,subcoriaceousto chartaceous; apex acuminateto subcaudate;base acute to

126

FLORA NEOTROPICA

.,(s

' (9;

()

i~(S)

(43t'

Fig. 27. Sorocea sarcocarpa. 1. Leafy twig with staminate inflorescences (Dodson et al. 7492). 2. Leafy twig with young infructescence. 3. Leafy twig (Gentry 9911). 4. Staminate flower. 5. Stamen (Eggers 14410). 6a. Pistillate flower and bract. 6b, c. Pistillate flower. 7. Fruiting perianth (Little 6192). [By P. Pardoen, Utrecht.]

127

TAXONOMIC TREATMENT

subobtuse;margin(mostly)distinctly(serrate-)dentate; uppersurfaceglabrous;lower surfacesparselyandminutely appressed-puberulouson the main veins; venation plane, but the midriband sometimes also the lateral veins slightly impressed above, the venation prominentbeneath; lateral veins 7-13 pairs; tertiary venationmostly partlyscalariform;petiole 0.2-1.5 cm long, 1-2 mm thick,ratherdenselyto sparsely,minutely puberulous;stipules0.2-0.5 cm long, appressed-puberulous, caducous.Staminateinflorescencespendulous, 2-8.5 cm long, including the 0.1-1 cm long puberulous peduncle; flowers distant to rather crowded, broadlysessile; perianthca. 1- 1.5 mm long, (deeply) 4-parted, subglabrous;stamens4, bent inwards,filaments ca. 0.8-1.2 cm long; anthersca. 0.6-0.7 x 0.60.8 mm, the connectivebroad,shortlyapiculate.Pistillate inflorescencespatent(or pendulous?in fruit), 1-5 cm long, to 9 cm long in fruit;peduncle 0.2-0.5 cm long, to 1.5 cm long in fruit, puberulous;flowers 515(-40) (in fruitingstateoften ? reflexed),pedicellate (or subsessile);pedicelto 0.3 cm long, 0.5-1.5 cm long in fruit; perianth ca. 2 mm long, faintly 4-lobed to subentire,theupperpartovoid to subglobose,glabrous, the lower partvery sparselyandminutelypuberulous; stigmas 0.5(-1) mm long, coarsely papillate.Fruiting perianthovoid to globose,ca. 0.8-1 x 0.6-0.9 cm, soon black,theapexroundedor apiculate,the surfacesmooth, (sub)glabrous. Distribution (see Fig. 77.5). Confinedto thePacific Coastalregionof Ecuador;in wetforest;atlow elevations. Specimens examined. ECUADOR. BOLiVAR:Cordillera Occidental,Valle del Lim6n, 16 Oct 1943 (st), AcostaS. 6428 (F). EL ORO: 10 km W of Pinias, 8 Oct 1979 (? fl), Dodson et al. 8984 (MO, QCNE,U). ESMERALDAS: Timbre,22-27 Feb 1953 (9 fl), Fagerlind et al. 2445 and 2503 (GH, NY, S); W of San Mateo, Reserva Forestal de Jardin Botanico, Universidad

LuisVargasTores,4 Feb 1991 (e), Gentryet al. T&enica 73073(MO);Borb6n,8 Sep 1965(&'),Littleet al. 21017 (F, NY,US); ca. 20 km S of Esmeraldas,13 Apr 1967(? fl-fr), Sparre 15490 (S). GUAYAS:Balao, Mar 1897 (e),

Eggers14319 (F, K, 0), (e and i), Eggers14410 (A, B, K, LE, US), May 1892 (9 fl-fr), Eggers 14640 (A, F,

6-7 May 1959 (? fl-fr),Harling5000 US); Manglaralto, (S); 20 km S of Esmeraldas,13Apr1967(? fl-fr),Sparre 15490 (MO).IMBABURA: Lita, 28 Apr 1949 (a), Acosta S. 12292 (F). Los Rios: Rio PalenqueBiological Station, 6 Mar 1974 (? fr), Dodson 5450 (F, MO, QCA, U, US), 21 Jan 1977 (e'), Dodson 6628 (AAU, MO, QCA); Cant6n Vinces, rd. Quevedo-Palenque via Mocachi, Jauneche forest, 4 Feb 1979 (a), Dodson et al. 7492 (U), 23 Mar 1980 (9 fr), Dodson et al. 9804 (MO, U); Clementina, 30 Mar 1953 ($ fl-fr), Fagerlind et al. 2665 (S); Rio Palenque Biological Station, rd. Quevedo-Santo

Domingode los Colorados,km 56, 13 Feb 1974 (? flfr), Gentry9911 (GB, GH, MO, S); Jaunecheforest, CantonVinces,betweenMocachiand Palenque,Estero Pefiafiel, 24 Jan 1981 (C), Gentry et al. 30683 (AAU,

GB, MO,NY,S, U); Pichilingue,21 May 1943(st),Little 6478 (US);Pichilingue,8 Nov 1982(? fl-fr),Pennington et al. 10679 (QCA,QCNE,U); betweenBabahoyoand Montalve, 18 Feb 1967 (2 fl-fr), Sparre 14496 (LD, MO, S). MANABi: Rd. Portoviejo-Pichincha,12 km E of SanPlacido,2 May 1985(R fl-fr),Harlinget al. 24741 (BG, GB). PICHINCHA:Rio Blanco,justbelowconfluence withRio Toachi,16-18 Mar1959(? fl-fr),Harling4499 (S); Finlandia, 16 km E of Santo Domingo de los Colorados,29 Jun 1976 (? fl), Gentryet al. 12158 (MO, QCA, U); rd. Santo Domingo de los ColoradosQuininde,km 43-68, 7 Apr 1943 (? fr), Little6192 (F, K, NY, US); betweenQuinindeand Santo Domingode los Colorados,2 Sep 1949 (st), Acosta S. 13676 (F). Local name. Ecuador:Bolivar:petaquilla.

11. Sorocea ruminata C. C. Berg, Novon 6: 244, t. 10. 1996. Type. Panama.Darien:ParqueNacional Darien,betweenCampamentoCasaViejaandCerro Sapo, 22 May 1991 (? fl-fr), Herrera et al. 975 (holotype, MO; isotypes, BG, BM, CR, MEXU, PMA). Fig. 28 Shrubor tree, to 12(-20) m tall. Leafy twigs 1-2.5 mm thick, sparselyandminutely puberulous.Lamina oblong to elliptic (to lanceolate),3.5-17 x 1.5-6.5 cm, broadestat or above the middle, ? inequilateral,coriaceous; apex acuminate;base acuteto rounded;margin entire;uppersurfaceminutely puberulouson midrib; lower surfacesparselyandminutelypuberulouson the (base of the) midrib;venation (almost) plane above, prominentbeneath; lateral veins 8-12 pairs; tertiary venationreticulate(or tendingto scalariform);petiole 0.3-0.8 cm long, 1-2 mmthick,(minutely)puberulous; stipules 0.3-0.5 mm long, (minutely) puberulous,caducous. Staminateinflorescences patent (?), 0.8-4.5 cm long, includingthe 0.1-0.2 cm long puberulouspeduncle;flowersrathercrowdedto ? dispersed,narrowly (orbroadly)sessile;perianth1-1.5 mm long, 4-parted, ciliolate, sparsely and minutely puberulous outside; stamens(3 or) 4, straight,isomorphic;filaments 0.51.2 mm long; anthersca. 0.5-0.7 x 0.4-0.7 mm, the connective broad, apiculate.Pistillate inflorescences patent(?), 0.5-3 cm long, to 5 cm long in fruit,including the 0.1-0.8 cm long, puberulouspeduncle;rachis sparsely and minutely puberulous; flowers 3-12, (sub)sessile, pedicellate in fruit;pedicel to 1 cm long, (very) sparsely hispidulous to minutely puberulous; perianthca. 2 mm long, glabrous,the upperparthemi-

128

FLORANEOTROPICA

Oi~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~l

Fig. 28. Sorocearuminata.1. Leafytwig with pistillateinflorescences(Herreraet al. 975). 2. Staminateflower. Stamenand bract(Gentry6554). 3. Fruitingpistillateflower(Herreraet al 975). [By HendriekeBerg, Bergen.]

TAXONOMICTREATMENT sphericalto broadlyovoid; stigmastongue-shaped,ca. 1 mm long, coarsely papillate. Fruiting perianth subglobose, ca. 0.74.9 x 0.7-0.9 cm, with the apical part? discoid and the lower partwith a ruminate,almost glabroussurface. Distribution (see Fig. 78.1). In easternPanamaand the adjacentpartof Colombia(Choco);in wet forest;at low elevations. Specimens studied. PANAMA. COCLt: Rd. to Celesito, 12 mi from Llano Grande, 16 Dec 1983 (d), Churchill et al. 4117 (MO). COL6N: Santa Rita Ridge rd., between TransisthmianHwy. and Agua Clara, 11 Dec 1973 (st), Berg et al. 302 (BG) and 309 (BG); Santa Rita Ridge, E of TransisthmianHwy., 16 Dec 1972 (e'), Gentry 6554 (NY); Santa Rita Ridge, E of Panama-Col6n Hwy., 13 May 1986 (? fl-fr), McPherson 9166 (BG, MO). DARIJN: Cerro Pirre, 4 Aug 1967 (? fr), Bristan 1231 (MO, US); Manene, mouth of Rio Cuasi, 28 Apr 1968 (9 fl), Kirkbride et al. 1393 (MO, NY); Cerro Pirre, above RENARE Camp, 28 Jul 1988 (? fl-fr), McPherson 12640 (BG, BM); Cerro Pirre, above Rio Tuquesa, Lower Tuquesa Mining Camp Charco Chiva, 5 Jul 1975 (Y fl-fr), Mori 6984 (BG, MO, U); Rio Tuquesa, Middle Tuquesa Mining Camp Charco Peje, 8 Jul 1975 (2 fr), Mori 7033 (MO, PMA); nr. Cana, 23 Jun 1959 (? fr), Stern et al. 662 (MO, US). PANAMA:ca. 20 km NE of Chepo, 12 Dec 1973 (st), Berg et al. 336 (BG); El Liano-Carti rd., km 8.7, 3 Sep 1977 (st), Berg et al. 403 (BG) and 405 (BG); Cerro Campana, 22 Jun 1972 (d'), Croat 17191 (MO); Barro Colorado Island, 1960 (e), Ebinger 161 (MO); El Llano-Carti rd., km 12-16, 5 May 1973 (? fl-fr), Kennedy et al. 3158 (MO, NY); foothills of Serrania de Maje, Rio Pirati, 16 May 1982 (st), Knapp et al. 5142 (MO); El Llano-Carti rd., 8.2 mi from Pan-American Hwy., 6 Jan 1982 (? fr), Knapp 5902 (MO, PMA, U); Cerro Jefe, rd. Alto Pacora-Cerro Brewster, km 4.5-5, 19 Jun 1988 (? fl-fr), McPherson 12597 (BG, MO). SAN BLAS: Cangandi, 19 May 1985 (? fl-fr), Nevers et al. 5772 (BG, MO, NY), 27 Mar 1986 (st), Nevers et al. 7455 and 7527 (BG, MO). COLOMBIA. CHOC6:Riosucio, Urabaregion, Cerros del Cuchillo, 19 Apr 1988 (de), Cardenas 1727 (JUAM), 20 Mar 1988 (Y fl-fr), Cardenas 2008 (JUAM), 24 Jun 1988 (? fl-fr), Cdrdenas 2121 (JUAM), trail Alto Curiche-Camp Curiche, E of Boca Curiche, 20 May 1967 (? fl-fr), Duke et al. 11301 (NY, US); upper Rio Baud6, Resguardo Indigena Embera, Quebrada de Condoto, 3 Jun 1985 (st), La Rotta et al. 680 (COL).

This species differsfromSorocea affinisin the consistentlyentireleaf margin,in the flowersbeing sessile at anthesis, and in the ruminate,subglabrousfruiting perianth.The inflorescencesaremorecompactandthe laminais more coriaceousthanthose of S. affinis.The fruitingperianthis often coveredby mycelium. Local name. Colombia: Choc6: bibobetubu (Embera).

129

12.SoroceajaramilloiC.C. Berg,Novon6: 241,t. 9. Pichincha: rd.Quito-Puerto 1996.Type.Ecuador. Quito,km 113, 10 kmN of rd.,ReservaForestal ENDESA,22 Aug 1984 (? fr), Jaramillo 7055 (holotype,QCA;isotypes,AAU,GB,MO,QCA). Fig. 29 Tree,to 20 m tall.Leafy twigs 2-5 mmthick,minutely puberulous,the older partsconspicuously lenticellate. Laminaelliptic,(5-)10-35 x (3-)7-17 cm, apex? abruptly acumi? inequilateral, (sub)coriaceous; nate;baseroundedto obtuseatthebroadside,acuteto obtuseatthenarrowside;marginentire;uppersurface ? on themainveins;lowersurminutelypuberulous, faceminutelypuberulouson the (main)veins;venabeneath; lateralveins8tionplaneabove,? prominent 10 pairs,the basalpairoftenrunningparallelto the reticulate orpartlyscalariform; margin; tertiary venation petiole (1.5-)2.5-3.5(-5) cm long, 1.5-2.5 mm thick,

stipules0.3-0.8 cmlong,puberminutelypuberulous; ulous, caducous. Staminateinflorescences in the leaf

axils,pendulous,ca. 16-33 cm long, including0.51.8 cm long, sparselypuberulouspeduncle;rachis flowers? spaced,broadlysessile; sparselypuberulous; perianth2.5-3 mmlong,4-parted,sparselyciliolate; stamens4, isomorphicor anisomorphic, straightor atanthesis; filaments 0.2-0.6mmlong; slightlyincurved anthers0.8-1.2 x 0.7-12 mm,the connectivebroad, not apiculate.Pistillate inflorescencesin the leaf axils

or belowthe leaves,pendulous,ca. 6-17 cm long,to 40 cmlongin fruit;peduncle0.5-3 cmlong,minutely rachisminutelyandyellowish-puberulous; puberulous; flowersca. (I0-)20-60; pedicel0.1-0.2 cm long,to 21.5cmlonginfruit,yellowish-puberulous; perianth 2.5 mmlong,minutelypuberulous,the upperpart? ruminate; stigmasovate,ca.0.5mmlong.Fruitingperianth ellipsoidto globose,2-2.5 mmlong,black. Distribution(seeFig.78.1).InthePacificCoastal regionsof EcuadorandColombia(Valle);in wetforest;atlowelevations. Specimens examined. COLOMBIA.VALLE: Ca. 16 km NW of Buenaventura,28 May 1987 (st), FaberLangendoen751 (BG). ECUADOR.CARCHI: N of Lita,N of Rio Miraand E of Rio Baboso, 7 Aug 1994 (st), Boyle et al. 3469 (BG);Cant6nTulcan,ReservaInigenaAwa,Chical,2327 May 1992 (2 fl-fr), Tipazet al. 1041 (BG, QCNE), (ei), Tipazet al. 1044 (BG, QCNE); Cant6n Tulcan, ReservaInigenaAwa, TobarDonoso, 19-28 Jun 1992 (? fl-fr), 27pazet al. 1240 (QCNE);GualpiAlto, 18-25 May 1985 (ei), Thomsen 58829 (QCA, QCNE). ESMERALDAS:Bilsa BiologicalStation,MarcheMts., 35 km W of Quininde,24 Jan 1995 (a'), J. L. Clark417

FLORANEOTROPICA

130

2mm

Fig. 29. Soroceajaramilloi. 1. Leafy twig. 2. Pistillateinflorescence(Jaramillo7055). 3. Pistillate inflorescence (Jaramillo6791). 4. Pistillateflower and bract(Jaramillo7055). [By HendriekeBerg, Bergen.]

(BG, QCNE), 28 Mar-l Apr 1995 (V fl-fr), J. L. Clark et al. 524 (BG, QCNE); Cant6n Eloy Alfaro, Reserva Ecol6gica Cotacachi-Cajapas, Charco Vicente, Rio San Miguel, 20-27 Mar 1993 (st), Mendez et al. 116 (QCNE),

6-9 Sep 1993(2 fr),Palacioset al. 11356 (BG, QCNE); Alto Tambo,13 May 1992 (? fl-fr), Quelal et al. 521 Cord. Chong6n-Colonche,Jul (BG, QCNE).GUAYAS: 1995 (? fl-fr), Bonifaz et al. 3096 (BG). Los Rfos/

TAXONOMIC TREATMENT

131

subspinulose-denticulate; uppersurfacesmoothand glabrousor puberulouson the midribor scabrousto al. 10278 (MO),23 May 1983 (st), Dodsonet al. 13806 scabridulous; lowersurfacesparselyto denselypuber(MO). PICHINCHA: Rd. Quito-Puerto Quito, km 113, ulous to hispidulous to subhirtellous or to 10 km N of rd., Reserva Forestal ENDESA, Rio Silanchi, midribandsometimesalsothelateral subtomentose; 10 Jul 1984 (? fl-fr), Jaramillo6791 (AAU, GB, MO, smaller veinsslightlyimpressedabove,the veins and NY, QCA). smallerveinsplaneori prominent beneath; lateral veins This species is characterizedby the largeand? dis- (5-)8-12(-14) pairs;tertiaryvenationreticulate(or tinctly inequilateralleaves and the pistillate inflores- partlyscalariform); petiole(0.1-)0.3-1.2cmlong,0.5cences which can become 40 cm long in fruit. 2 mm thick,puberulous;stipules0.2-0.9 cm long, PICHINCHA: El Centinela, Montaniasde Ila, rd. Patricia

Pilar-24 de Mayo, km 12, 6 Apr 1980 (d), Dodson et

Local name. Ecuador:Guayas:miguelillo.

puberulous,caducous.Staminateinflorescencesshortly

spicate(to 1(-1.5) cm long)or subcapitate; peduncle 0.2-0.5cmlong,puberulous; flowerscrowded,sessile or withpedicelsto 1 mmlong;tepals4, free,ca. 1.5 7B. Sorocea Saint-Hilairesubg.Paraclarisia (Ducke) mmlong,membranaceous, minutelypuberulous; staW. Burger, Lanjouw & Wessels Boer, Acta Bot. mens4; filaments0-0.1 mmlong;anthersca.0.9-1.1 Neerl. 11: 468. 1962. Paraclarisia Ducke, Arq. x 0.8-1 mm,theconnectivebroad,apiculate. Pistillate Serv. Florest. 1(l): 2. 1939. Type. Paraclarisia inflorescences veryshortlyspicateto subcapitate, in amazonica Ducke (= Sorocea duckeiW.Burger). fruitingstatetoracemoseorsometimessubumbellate; IncludingTrophisomiaRojasAcosta (see underthe peduncle0.2-1.5 cm long,sometimesto 2.5 cm long infruit,puberulous; flowers4-12, sessileorsubsessile, genus). infruit;pedicelmostlyto0.5cmlong,someShrubsor trees, deciduous;intemodes often short; pedicellate scarsof the leaves prominent;axillarybuds conspicu- timesto 1.5cm longorto 3.5 cm long,reddish;perious, globose. Inflorescencessubcapitate(or shortlyspi- anthca. 1.5-2mmlong,4-lobed,sparselyandminutely to band-shaped stigmastongue-shaped, to cate); bracts submembranaceous;stamens with very puberulous; 1-2 mm subfiliform to or sometimes subulate, long, shortfilaments. twisted,coarselyorsometimesfinelypapillate.FruitThis subgenus comprises two species from South ingperianth ellipsoidtoovoid,sometimes (sub)globose America,occurringin habitatsdifferentfrom the pre- oroblongoid,(0.6-)l-I.3(-I.5) x (0.6-)0.8-0.9(-1.2) dominantone in subg. Sorocea. sometimestruncate cm,black,theapex(sub)apiculate, withalowcollar-like rim,thesurfacesmooth(orslightly wrinkled?), sparselyandminutelypuberulous. 13. Sorocea sprucei (Baillon) Macbride,Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 16. 193 1; Macbride, Publ. Field Mus. Nat. Hist., Bot. Ser. 13(2.2): 313. 1937; Burger,Lanjouw& Wessels Boer,Acta Bot. Neerl. 11: 468, t. 13. 1962; Berg & Akkermans, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med. Sci. 88: 391, t. 4. 1985.Pseudosoroceasprucei Baillon, Adansonia 11: 296. 1875. Type.Peru.San Martin:nr. Tarapoto,Jun 1855 (c), Spruce 4483 (holotype, P; isotypes, B, BM, C, F, G, K, LD, LE, NY). Fig. 30

Thespeciesis veryvariable.Threesubspeciescan berecognized atpresent.A fourthdistinctinfraspecific taxonfromthedrypartsof theupperCaucavalley(Colombia,Valle)mightberecognizedasdiscussedbelow, andpossiblya fifthoneto accommodate thesomewhat deviatingcollectionsfromBolivia(Beni),Fosteretal. 12469 andMichelet al. 1040 and 1042), withrelatively largechartaceouslaminas,? scabrousbeneath.

Key to the subspecies of Sorocea sprucei

1. Smallerveins of the laminaplaneto slightly prominentbeneath;Curacao,northern Shrub or tree, to 8 m tall, sometimes lianescent. Venezuela,and Colombiato Bolivia ........... Leafytwigs 1-3 mm thick,puberulousto hirtellous;in13a. subsp.sprucei .................................................. ternodes often short;lenticels often conspicuous and of 1. the lamina Smaller veins prominent dark (brown or blackish) colored. Lamina elliptic to beneath. lanceolate(to subobovate),(l-)6-16 x (0.5-)2.5-8 cm, 2. Apex of the laminausuallyroundedto ? to mostly broadest above the middle, inequilateral obtuse; Bolivia to N Argentina.......... equilateral,chartaceousto coriaceous;apex acuminate 13b. subsp. saxicola .......................................... to subacuteto obtuseto rounded,the acumenapiculate; 2. Apex of the laminausually acuminate; base acuteto roundedto cordulate;margin(sub)entire SW Ecuadorand NW Peru ................. ................................... 13c. subsp. subumbellata to denticulate to (crenate-)dentate, occasionally

132

FLORANEOTROPICA

'21t

i(53

qs

ql@3q

i

0t 893

4

Fig. 30. 1, 3-6, 8, 9. Soroceaspruceisubsp.sprucei.1. Leafytwig with pistillateinflorescence(Breteler4333). 3. Leafytwig with infructescence(Cuatrecasas15340). 4. Pistillateinflorescence.5. Pistillateflower (Steyermark 101957). 6. Fruitingperianth(Arnoldo1104). 8. Twigs with staminateinflorescences(Marcano-Berti38-3-77). 9. Staminateflowerand stamen(Fr6es23120). 2, 7. S. spruceisubsp.subumbellata. 2. Leafytwig with younginfructescence(Plowman5471). 7. Fruitingperianth(Plowman5471). 10, 11. S. spruceisubsp.saxicola. 10. Leafy twig with infructescence(Pedersen1208a). 11. Staminateflowerand stamens(Hassler12222). [By T. Schipper,Utrecht.]

TAXONOMICTREATMENT 13a. Sorocea sprucei (Baillon) Macbride subsp. sprucei Fig. 30 GuatteriacuspidataRusby,Mem.N.Y. Bot. Gard.7: 245. 1927.Type.Bolivia.Beni:Nr.LakeRogagua, 1921 (v), M. Cardenas1706 (holotype,NY). OlmediavirgataPittier,Bol. Soc. Venez.Ci. Nat. 8: 305. 1942; Berg, Fl. Neotropica7: 145. 1972. Type.Venezuela.Bolivar:BajoCaura,2 May 1939 (ei), Ll. Williams 12040 (holotype,VEN;isotypes, F, G, S, US). SoroceaarnoldoiLanjouw& WesselsBoer,ActaBot. Neerl. 11:472, t. 13. 1962.Type.Cura9ao.Christoffelberg,10 Aug 1952 (? fl-fr), Bro. Arnoldo 2106 (holotype,U; isotypes, F, NY, P, US). Laminato 16 x 7.5 cm; apex acuminate,occasionally subacute;base cordulateto subacute;lower surface hirtellous to tomentose to puberulous to hispidulous; the smaller veins slightly prominentto almost plane beneath. Staminate inflorescences subcapitateto shortly spicate,to 1(-1.5) cm long. Distribution (see Fig. 78.2). In northemVenezuela andCura9ao,northemColombia,southemGuyana andthe adjacentpartof Roraima(Brazil), centralColombia, Peru(San Martin,Tarapotoarea),andBolivia (Beni); usually in relativelydrytypes of forest,suchas subxerophyticforest, savannaforest, semi-evergreen riverine forest, but also in seasonally flooded forest (Roraima);to 1300 m. Representativespecimens examined. CURACAO. 2 Nov 1946(o), Bro.Arnoldo6 (U);hills Christoffelberg, nr. Knip, nr. Pose Kajoeda,Sep 1948 (? fl-fr), Bro.Arnoldo 1088 (MO, U), Sep 1948 (2 fr), Bro. Arnoldo1104 (U), May 1951 (st), Bro.Arnoldo1632 (US); Christoffelberg, 10 Aug 1952 (ei), Bro.Arnoldo2106A(NY, U); hills nr. Knip, 30 Nov 1963 (c), Stoffers5112 (U). COLOMBIA:ATLANTICO: Manati,4 Jan 1934 (st), Dugand 590 (F); nr. Usiacuri, Sep 1938 (st), Dugand 1213 (COL, US); confluence of Rio Honda and Rio Le6n, rd. Barranquilla-Puerto Colombia,25 Apr 1965 (st), Dugand6557 (US), 25 Apr 1965 (st), Dugand6904 Mun.Morales,Cgto. Norosi,Tiquisio(US). BOLIVAR: nuevo,9-14 Apr 1985 (ei), CuadrosV 2120 (MO,US); nr. San JuanNepomuceno,70 km SW of Cartagena,7 May 1987 (st), Gentryet al. 57452 (MO), 11 Jan 1988 (st), Gentryet al. 60567 (MO). CASANARE: Orocue, Canio Guirripa, Jul 1990 (st), Mahecha6906 (UDBC). CUNDINAMARCA: QuebradaCarmargo,N of Apulo, 5 May 1944 (? fl-fr), Killip et al. 38232 (US). GUAJIRA: Mun. Fonseca, Cgto. Distracci6n, 31 Aug 1990 (st), Marulanda et al. 2122 (HUA).HUILA: Mun. La Plata, Hda. Limona, 1020 m, 8 Oct 1984 (c), Lozanoet al. 4796 (F, NY). MAGDALENA:Rio Cesare,E of LosVenados,23 Apr 1963 (st), Dugand6321 (COL, US); Tucurinca,Dec 1947 (st), Romero-Castafieda 646 (COL); Pivijay, 27 Apr

1948 (? fl), Romero-Castanieda1107 (COL, F, US); Santa Marta, 1898-1899 (c), H. H. Smith2411 (A, F,

133 GH, MICH, MO, NY, P, U, WIS). NORTEDE SANTANDER: Rio Zulia, between San Cayatanoand Cucuta, 24 Jul 1940 (st), Cuatrecasas et al. 10160 (COL, F, US). RISARALDA: Mun. Marsella, Puerto Caldas, 1000 m, 1980 (st), Idrobo et al. 10158 (COL). QUINDfO: Mun. La Tebaida, rd. Tebaida-El Alambro, 1100 m, 28 1987 (st), Arbeldez S. et al. 2041 (MO). SANTANDER: Capote, Jun 1968 (st), Saa 405 (UDBC). SUCRE: Colosa, nr. Estaci6n de Primatas, 17 Nov 1981 (st), Gentry et al. 34825 and 34829A (COL, MO). TOLIMA:Mariquita, 2 Apr 1976 (st), Mahecha UDBC 10871 (UDBC). VALLE: Mun. Tulua, Mateguadua Botanical Garden, 1070 m, 22 Oct 1980 (e), Cuadros V 1222 (MO,TULV); Rio Saquinini, 1200 m, 8 Dec 1943 (? fl-fr), Cuatrecasas 15340 (F); Palmira, 13 Mar 1937 (e), D. Jaramillo 236 (VALLE); nr. Buga, 1000 m, 5 May 1982 (? fr), Murphy352 (COL, MO, NY, TULV); Mun. Carrito, Hda. San Gerardo, 12 Dec 1987 (? fl), Ramos et al. 850 (CUVC); Pan-American Hwy., between La Paila and Zarzal, 20 Jun 1987 (? fl-fr), Silverstone-Sopkin et al. 3209 (MO); Hda. San Gerardo, nr. border of Mun. Guacari, nr. Guabas, 10 Mar 1988 (? fl-fr), Silverstone-Sopkin et al. 3659 (MO); 3 km S of Buga, 16 Jun 1988 (? fr), Silverstone-Sopkin et al. 4122 (MO). VENEZUELA. ANZOATEGUI: Caiio Mamo, 40 km W of San Fe1ix, Jul 1988 (? fl-fr), Colonello 1228 (MO). APURE:San Fernando,Rio Arichuna,Fundo Las Maropas, 3 Apr 1969 (st), Aristeguieta et al. 7057 (MO,NY, VEN); Dtto. Pedro Camejo, Rio Capanaparo, 14 km SW of Uranon, 6-7 May 1977 (st), Davidse et al. 12820 (MO, U); Dtto. Pedro Camejo, Rio Capanaparo, 11 km E of Paso de San Pablo, 8-9 May 1977 (st), Davidse et al. 12841 (MO). BARINAS:Rio Michan, Reserva Forestal de Ticoporo, 4 May 1966 (st), Marcano Berti 860 (MER, MO, U); NW of Barrancas, 30 Mar 1977 (e), Marcano Berti et al. 38-3-77 (MER, MO, U); Pedreza, 65 km SE of Cuidad Bolivia, 19 Apr 1953 (a"),Little 15088 (VEN); Reserva Forestal Caparo, E of El Cant6n, 9 Apr 1968 (? fl), Steyermark et al. 101957 (F, MO, U, VEN), 11 Apr 1968 (a), Steyermarket al. 102135 (K,U, VEN). BOLIVAR: Reserva Forestal El Cauro, Jun 1970 (? fl-fr), Blanco 854 (MO, U); Dtto. Cedefno,rd. Maniapure-Caicara,km 6, 4 Oct 1985 (st), Boom et al. 6243 (NY), 23 May 1986 (? fl), Boom et al. 6696 (MO, NY); between Guaniamo and Perro de Agua, 9 Jun 1984 (dc fl-fr),L6pez-Palacios et al. 4522 (NY); Parque La Llovizna, Isla Zapateral, Puerto Ordaz, 5 Aug 1989 (? fr), Rosales et al. 624 (BG). DIssRITo FEDERAL: Caracas, JardinBotanico, May 1958 (a'), Aristeguieta 3084 (VEN), Jun 1958 (? fl-fr), Aristeguieta 3214 (VEN), 12 Apr 1979 (da),Berry 3485 (MO, NY); Cotiza, nr. Caracas, May 1918 (? fl), Pittier 7870 (F, GH, MICH, MO, US, VEN); Caracas,Colinas de Bello Monte, 1300 m, 9 Jun 1975 (9 fl-fr), Ruiz Z. et al. 75-0051 (MO, VEN). FALC6N: Peninsula Paraguani, Cerro Santa Ana, 176 Dec 1964 (9 fl-fr), Breteler 4333 (G, MO, NY, U, US, VEN); Dtto. Colina, Rio Macoruca, between Las Barrancasand El Turaguai de Macoruca, 12 Feb 1977 (st), Steyernark et al. 113749 (U, VEN); Sierra San Luis, 2 km W of Carrizalira, 1170 m, 24 Mar 1970

134

FLORANEOTROPICA

(de), Wingfield5730 (U); Paraguana,CerroSantaAna, V 1220, Cuatrecasas 15340, D. Jaramillo 236, and 8037 (U). GuALco: Ramos et al. 850. They havepistillateinflorescences Moruy,2 Nov 1980(? fl), Winglield Dtto.Miranda,Mun.Calabozo,Rio Orituco,28 Aug 1975 to 0.5cmlongwith1-5 flowers,globosefruitingperi(st), Castillo146 (BG,VEN);Rio Guarico,S of Calabozo, anths0.6-0.7 cmdiam.,andpedicelsrelativelyshort, 14 Feb 1979 (st), Gentryet al. 24734 (MO). MIRIDA: to 0.5 cmlong.Theleavesarerelativelysmall,smooth Mun.Lagunillas,Dtto. Sucre,rd. Estanquez-Paramo Las the above,andsubtomentose to subhirtellous beneath; Coloradas,6 May 1978 (? fr), MarcanoBertiet al. 1-5margin is are usually entire. The leafy twigs densely 78 (MER,U). MIRANDA: W of SantaFe, Quebrada Barute, 3 Aug 1975 (st), Steyermarket al. 111807 (MO,VEN). hairy.It mayrepresenta distinctinfraspecifictaxon fromtheupperCaucavalley.Somecollectionsfrom MONAGAS: Caicara,25 May 1952 (d), F D. Smith221 (US, VEN). PORTUGUESA: Dtto. Guanare,groundsof Bolivia(Beni)areplacedundersubsp.sprucei asthey UNELLEZ,3 Jul 1984 ($ fl-fr),Aymard2623 (NY);32- are more similarto this subspeciesthanto subsp. 35 km fromAcarigua,Feb 1969 (st), Aristeguietaet al. saxicola. Theymightrepresent a distinctinfraspecific 6919 (VEN); 30 km W of Guanare,Rio Tucupido, 11 taxonaswell. Mar 1982 (e), Liesneret al. 12438 (MO, NY, U); Rio Localnames.Colombia: Atlantico: copey,gigant6n, Maria,Boca de Monte,38 km N of Hwy.at Las Marias, 3 Nov 1982(st),Steyermark et al. 127141(MO).YARACuY: guaimarolechoso;Magdalena: guaimaro,guaimaro Chivacoa,Filade Eljalma,20 Jun 1970(? fr),R. F Smith lechoso,mamonde leche.Venezuela: Apure:charo; V-6260andV-6261(VEN).ZULIA:Dtto.Colon,nr.Casigua Barinas: charo,charonegro;Bolivar: charo,lechero,weEl Cubo, 7-14 km N of Casigua, 29 Apr 1979 (? fr), yo (Panare);DistritoFederal:lechero;Gutrico,Monagas, Buntinget al. 7379 (NY, U); Dtto. Mara,Sierrade la andPortuguesa: Zulia: charo; guaimaro, gudimaro negro. Corbata,3-4 km NNE of ParcelamientoCachari,5 Jun 1980(? fl-fr),Steyermark et al. 123361(MO,NY,VEN); Dtto. Mara, 3 km WSW of CorpozuliaCampamento 13b. Sorocea sprucei (Baillon) Macbridesubsp. Carichuano,30 May 1980 (? fl-fr), Steyermarket al. C.C.Berg,Proc.Kon.Ned.Akad. saxicola(Hassler) 122949(MO,NY);Dtto.Mara,Rio Socuyo,betweenCerro Ser. Biol. Med. Sci. 88: 391. 1985. Wetensch., C, La Culebraand CerroCaracara,27 May 1980 (? fl-fr), Soroceasaxicola Hassler,Bull.Herb.Boissier,Ser. et al. 12295 (MO,NY,VEN). Steyermark 2, 7: 11. 1906; Hassler,Annuaire Conserv. Jard. GUYANA.South Rupununi,Oct 1969 (st), Davis Bot. Geneve21: 111. 1919;Chodat& Vischer,Bull. 1696 (A, GH, NY); Upper Takutu-UpperEssequibo region,65 km NE of Warimure Ranch,4 Nov 1991 (st), Soc. Bot. Geneve,Ser.2, 11:233. 1920;Burger, E. M. Harriset al. 1133 (BG);RupununiDistrict,footLanjouw& WesselsBoer,Acta Bot. Neerl. 11:470. hills NW of KanukuMts., Moco-Moco, 26 Oct 1979 t. 13. 1962. Type.Paraguay.Concepcion,Sep 1901 (st), Maas et al. 3826 (U); without locality, (e + (? fl), Hassler 7338 (holotype,G; isotypes, B, BM, Schomburgk1239 (B, K). C, K, MICH, MO, NY, P,S). Fig. 30 PERU.SAN MARTIN:valleyof Rio Huallaga,29 km Soroceasaxicola Hasslervar.dentataHassler,Bull. S of Tarapoto,nr.El Abra,5 Feb 1984 (st), Gentryet al Herb.Boissier, Ser. 2, 7: 12. 1906. Type. Para45008 (BG, MO); Dtto. Juanjui,rd. Tarapoto-Juanjui, guay. San Bernardino,(c), Hassler 1071 (holo23 Nov 1982 (st), Rios T 2 (U). type, G; isotypes,BM, NY, P). BRAZIL. RORAIMA: Mun. Boa Vista, SEMAEcol. Soroceasaxicola HasslerformasubrepandaHassler, Reserve,Ilha de Maraca,16 May 1987 ($ fl), Lima et Bull. Herb.Boissier, Ser. 2, 7: 12. 1906. Type. al. 750 (BG), 19 May 1987 (? fl-fr), Lima 787 (BG, Paraguay.CordilleraCentral,Cerrosde Tobaty, INPA),27 Mar-l Oct 1987(st), Millikenet al. 103 (BG); Sep 1900 (c), Hassler 6455 (holotype, G; Rio Surumu,(e), Ule 8115 (B). isotypes,B, BM, F, GH, K, MO, NY, P, US). BOLIVIA. BENI: LakeRogagua,8 Nov 1921 (Q flTrophisomiaedulis Rojas Acosta, Bull. Acad. Int. fr), M. Cardenas1697 (NY); Prov.Yacuma,E of San Geogr. Bot. 24: 211. 1914. Type. Argentina. Bora, Bosquede Chimanes,10 km N of timbermill, 23 Chaco: nr. Rio Parana, nr. Colonia Benitez, Nov 1988 (? fr),Fosteret al. 12469 (BG, F, LPB);Prov. Niederlein3359 (not traced). 13 Sep 1993 (d'), Ballivian,rd. Rurrenabaque-Yucumo, Michel et al. 1040 (BG, LPB), (st), Michelet al. 1042 Lamina to 10 x 5.5 cm; apex mostly rounded to (BG, LPB);Prov.Cercado,rd. Trinidad-SanBora, km orshortlyacuminate; subacute base 40, 18 Jun 1995 (st), Moraeset al. 2237 (LPB);Prov. obtuse,sometimes Moxos, rd. San Bora-Trinidad,Bosque de Produccion cordulateto rounded;lowersurfacehirtellousto tomenbeneath.StaminateinChimanes,4-5 Sep 1990 (st), D. N. Smith14310 (LPB, tose;allveins(very)prominent MO) and 14331 (LPB, MO). florescences capitate.

The materialfromnorthernColombiaandVenezuDistribution(seeFig.78.2).IntheupperRioParaela, Roraima(Brazil),andCuracaois ratheruniform.A guayBasin,fromBrazilthroughParaguay to northem more or less distinct form occurs in Valle (Colombia) Argentina, andinnortheastem Bolivia;in savannaforat900-1200 m. Itis represented by thecollectionsCuadras est;atlow elevations.

TAXONOMICTREATMENT

135

Representative specimensexamined.BRAZIL.MATO

EstanciaLasTresMarias,15 U, US);Depto.Empedrado,

GROSSO:Cuiaba,Sep 1827 (? fl-fr),Riedel 1184 (LE);

Sep 1971 (? fl), Pedersen 9839 (A, MO); Depto. Capital,

Caceres,(st), Rondon4481 (R), (v), Rondon4504 (R); Perich6n,Rio Parana,3 Oct 1976 (? fl), Schininiet al. Mun.Pocone,7-12 km SW of Rio Cassange,rd.to Porto 13639 (MO, WIS), (d6), Schinini et al. 13640 (MO); Jofre, 25 Oct 1985 (? fl-fr), Thomaset al. 4577 (BG, Depto.Salada,LagunaSofo, 28 Nov 1944 (d), Schwarz 35 (A, NY); Depto. San Cosme, QuertoToledo, 19 Jul INPA, MO, NY, SPF). MATOGROSSODO SUL: Mun. 7 Oct 1953(? fr),Pereira 1944 (st), Ybarrola671 (F, NY). FORMOSA: Depto. Corumba,FazendaMarilandia, N of Clorinda,13 Sep 1946 et al. 262 (RB); FazendaCongonha(P7-RADAM),22 Pilcomayo,RiachoPortenio, Oct 1980 (R fr), Pires et al. 17254 (U). (d'), Morel 1291 (NY). BOLIVIA.BENI:Prov.Marban,San Rafael,28 Sep This subspecies is rather uniform. Some of the col1979(9 fr), Beck2501B (MO);Prov.Ballivian,Espiritu, from Bolivia show features ? transitional to lections nr.RioYacuma,28 Sep 1979(Y fl), Beck2501 (MO)and They match the indumentum of the subsp. sprucei. 2501A(BG, MO), 29 Sep 1979 (st), Beck2587 (BG), 1 Oct 1979 (st), Beck2602 (BG, MO), 15 Oct 1980 (? fl- leaves of subsp. saxicola, but the apex of the lamina is fr), Beck et al. 5078 (BG); Prov. Yacuma,Estacion often acuminate to subacute as typical of subsp. sprucei. Biol6gica Beni, 16 Nov 1985 (o), Moraes 655 (BG); Prov.

Moreover, some Bolivian collections are more similar

Ballivian,EstanciaEl Porvenir,50 km E of Rio Manique, to subsp. sprucei in the dimensions, the apex, and the rd. to Trinidad, 14 Nov 1985 (9 fl), Solomon 14721 venation of the lamina than to subsp. saxicola and are, 27 Oct 1991(? fr), Gen- therefore, listed under the former taxon. The differences Curuyuqui, (BG).SANTA CRUZ: tryet al. 75311(USZ);Prov.AngelSandoval,SanMattias, simi7 Sep 1994 (a), 7 Sep 1994 (ci), 7 Sep 1994 (a), Guillen

2229 (USZ);Prov.AndresIbanfez,12 km E of centerof SantaCruz,rd. to Cotoca, 24 Aug 1987 (? fl-fr), Nee 35729 (MG, MO, NY), 28 Aug 1987 ('), Nee 35813 (GB,

between subsp. sprucei and subsp. saxicola show larities to those of the two subspecies of Maclura tinctoria.

Local names. Bolivia: Beni: sapito. Paraguay:

MO,NY), 26 Jul 1991(st), Nee 41815 (BG);Prov.Sara, neandipa mei. Argentina: Chaco: nand pa Portachuelo,26 Sep 1929 (a), Steinbach6477 (B, BM, F, G, GH); Prov. Sara, La Perdix, 8 Aug 1927 (d), Steinbach7917 (A, B, BM, F, G, GH,K, MO,NY,U, Z). 13c. Sorocea sprucei (Baillon) Macbride subsp. Nr.Ascuncion,Sep-Nov PARAGUAY. Rio Paraguay, subumbellata C. C. Berg, Novon 6: 245. 1996. 1876 (? fl-fr), Balansa 1969 (B, G, K, LD, P); Cerro Ecuador. Guayas: rd. Guayaquil-Nobol, km Type. 24 Sep 1881(? fl-fr),Balansa3192 Pelado,nr.Paraguari, 14,21 Jan 1985(? fl-fr),Harlingetal. 21069 (ho(G, P); EsteroCamba,Misiones-Neebucu,8 Nov 1978 Fig. 30 lotype, GB; isotype, BG). deAltos, (Y fl-fr),Bernardi18374(F,MO,NY);Cordillera Sep 1892(? fl), Fiebrig182(A, B, F,G, K, P);AltoChaco,

Shrub or tree, to 8 m tall, sometimes lianescent. Lamina oblong to elliptic to (sub)obovate, 3-11 x 1.51325a(B, G, K, Z);Depto.PresidenteHayes,VillaHayes, Estancia La Golandrina, 9 Sep 1982 (ci), Hahn 707 (G, 5.5 cm; apex acuminate to subacute; base rounded (to MO, RB); nr. Concepci6n,Rio Ypane, Sep 1901 (a"), obtuse); margin distinctly (to rather faintly) dentate; Hassler7338a (B, BM, G, K, MO,NY); CordilleraCen- venation impressed above, prominent beneath. Pistiltral, nr. Lago de Ypacarai, Aug 1913 (c) Hassler 12222 late inflorescencessubumbellatein fruit;peduncle0.321?S, 1906 (a"),Fiebrig 1325 (B, G, K, Z), (?), Fiebrig

and (? fl) Hassler 12222a (A, B, BM, C, F, G, GH, K, MO,NY, US, Z); Depto.Capital,betweenIta Enramada and PuertoPaula,Rio Paraguay,9 Sep 1978 (? fl-fr), CerroMbatori, Schinini15358 (MO);Depto. Paraguari, 9 Aug 1988(st),Soziaet al. 2315 (MO);LasGolandrinas, Cerro Aug 1982 (a"), Wright83 (MO);Depto.Paraguari, Mbatovi,2 Jul 1988 (st), Zardini5493 (BG). 12 NovARGENTINA. CHACO:nr. Barrangueras, 15 Dec 1913 (st), Curran27 (US); Las Palmas,9 Sep 1917 (? fl-fr),Jorgensen2156 and (a")2157 (GH,MO, US);nr.Rio Pilcomayo,(2 fl), Rojas480 (G);Resistencia, 26 Sep 1944 ('),

Rojas 11649 (A, NY); Depto. Puerto

de Mayo,ColoniaBenitez,Sep 1953 (2 fl), Schulz8541 Rio Empedrado, (? fl), Empedrado, (US). CORRIENTES: Krapovickaset al. 19924 (F); Depto. Mburucuya,Estancia

SantaTeresa,20 Nov 1949(? fl-fr),Pedersen443 (C, K, P, U, US); Depto.Mbumcuya,EstanciaSantaMaria,30 Sep 1951 (a), Pedersen 1208 (C, G, MO, NY, P, U, US),

22 Dec 1951(? fl-fr),Pedersen1208a(C, G, MO,NY,P,

1 cm long, to 3.5 cm long in fruit; flowers 3-12(-18); pedicel 1-1.8 cm, to 3.5 cm long in fruit, (brownish)red in fruit; immature fruiting perianth (brownish-)red. Distribution (see Fig. 78.2). In southwestern Ecuador and the adjacent part of Peru (Tumbes); in dry forest; at low elevations.

Specimensexamined.ECUADOR.EL ORO: ca. 60 km SE of Arenillas,Rd. to Loja, 13 Nov 1982 (? fl-fr), Pennington et al. 10720 (QCA, QCNE, U); Cant6n Arenillas, rd. Arenillas-Piedras, 16 Feb 1976 (? fr), Plowman 5471 (GH, S, U). GUAYAS:Rd. GuayaquilSalinas, km 24, 18 Mar 1980 (? fr), Dodson et al. 9601 (F, MO, U); rd. Guayaquil-Daule, km 21, Capeira, 18 Sep 1981 (st), Dodson et al. 11346 (F, MO), 23 Sep 1981 (st), Dodson et al. 11454 (MO), 11 Feb 1982 (? fr), Dodson et al. 12346 (F, MO). LOJA: QuebradaEl Chirimoyo, Bosque Petrificado Puyango, 13 Apr 1996

136

(st), Cornejoet al. 5021 (BG); PuenteChico, 12 km N of Alamor,9 Aug 1975 (st), Samaniegoet al. 50 (LOJA, QAME,US). PERU. TUMBES:Prov.Tumbes,Pampasde Hospital, El Caucho,22 Jan 1989 (Q fl), C. Diaz et al. 3224 (BG, MO). Local names. Ecuador: Guayas: pinto de cerro; Loja:guacora.

FLORA NEOTROPICA

512 (COL, P, U); Rio Loretoyacu, Sep 1946 (? fr), Schultes et al. 8273 (COL, GH, US); Rio Caqueta, nr. La Pedrera, Apr 1944 (d fr), Schultes 5858j (F). Manco PERU.LORETO:Prov.Maynas,Yarina-Cocha, Cocha, 13 Dec 1987 (e'), Ayala 5899 (MO); Prov.

Requena,nr. JenaroHerrera,CafioSupay,16 Feb 1988 (? fr), Daly et al. 5657 (BG, MO, NY); Prov.Requena, Rio Pacaya, CainoYarina, 22 Mar 1977 (? fl-fr), Encarnaci6n E-1073 (MO, US); Prov. Requena, nr.

Jenaro Herrera,CainoIricahua, 17 Feb 1982 (? fl), Encarnaci6n 1277 and 1279 (MO, NY, US), 24 Aug 1982

14. Sorocea duckei W. Burger,Acta Bot. Neerl. 11: 473, t. 13. 1962, as new name for Paraclarisia amazonica. Paraclarisia amazonica Ducke, Arq. Serv.Florest. 1(1): 2, t. 3. 1939. Type.Brazil.ParA: Santarem,Genipatuba,2 Feb 1933 (9 fr), Ducke RB 23995 (lectotype, Burger,Lanjouw& Wessels Boer, Acta Bot. Neerl. 11: 473. 1962, RB-n.v.; Fig. 31 isolectotypes, B, G, K, U, US). Shrubor tree,to 7 m tall.Leafytwigs 1-3 mmthick, hispidulous to puberulous; lenticels inconspicuous. Lamina oblong, (2-)5-12 x (1-)2-5 cm, broadestat or above the middle, (almost) equilateral,chartaceous to (brittle)subcoriaceous;apex acuteto subacuminate to obtuse; base acute to obtuse; margin entire, sometimes obscurelydenticulate;uppersurfacescabrousto scabridulous;lowersurfaceon the mainveins scabrous, on the smallerveins tomentoseto hirtellous;venation slightly impressedto plane above, prominentbeneath; lateralveins (6-)9-12 pairs;tertiaryvenationpartlyscalariformto reticulate;petiole0.2-0.8 cm long, 1-1.5 mm thick, scabrous;stipules0.1-0.4 cm long, puberulous, caducous. Staminateinflorescences subcapitate,0.40.6 cm diam.;peduncle 0.1-0.3 cm long, puberulous, bracteate;flowerssessile;tepals4, free,ca. 1.5mmlong, membranaceous,minutelypuberulous;stamens4; filaments0.1-0.3 mm long; anthers0.6-0.7 x 0.5-0.7 mm, the connectivebroad,not apiculate.Pistillate inflorescences with 1-2(-3) flowers; peduncle 0.1-0.3 cm long, puberulous, bracteate; flowers sessile, but subsessile or with pedicels to 0.2 cm long in fruit;perianthca.2-2.5 mm long,4-lobedto subentire,theupper partsparselyand minutelypuberulous;stigmas subulate, 2-3 mm long, finely papillate.Fruiting perianth ellipsoid to subglobose (to obovoid), 1.2-1.5 x 1-1.3 cm, ? apiculate,purpleto black,the surfacesmooth(or slightly wrinkled?),subglabrous. Distribution (see Fig. 78.2). In the upperAmazon Basin, mainly along the Amazon River, also near Santarem(ParI);in periodicallyinundatedmarginsof rivers and lakes.

(? fl-fr),Encarnaci6n26494 (MO, US); Prov.Maynas, Dtto.Pebas,Rio Ampiyacu,13 May 1976(? fr), Revilla 596 (MO,NY, U); Prov.Maynas,PuertoAlmendras,10 Jul 1990 (st), J. Ruiz C. 1519 (BG); nr.JenaroHerrera, QuebradaSupay,16 Apr 1989 (? fl-fr),R. Vcsquezet al. nr.Pucallpa, 12032 (BG).UCAYALI: LagoYarina-Cocha, 20 Jan 1987 (? fl-fr),Stein et al. 3931 (BG, MO). SaoPaulode Olivenga, 28 May BRAZIL.AMAZONAS: 1946 (? fr), Ducke 557 (MO, NY, US); Rio Solim6es, Tonantins,21 Feb 1944 (e'), Ducke 1530 (A, IAN, K, MG,NY, US), (? fl) Ducke1531 (A, IAN, K, MG,NY, US), (2 fl-fr);Sao Paulode Oliven9a,Camatia,10 Apr 1944 (? fl-fr)Ducke1532 (A, NY, US), 29 Jan 1937 (? fl-fr), Sao Paulo de Olivenqa,Ducke RB 35613 (G, U, US); FuroManaquiri,21 Feb 1941 (? fr), Fr6eset al. 2 (G, NY, US); Tonantins,17 Aug 1941 (st), Fr6es 12209/ 122 (F);nr.Manaus,Paranado Xiborema,FazendaSanto Antonio,May 1953 (? fr), Fr6es29629 (GH,IAN, NY, U, US);Mun.Humaita,2 km N of Humaita,1 May 1955 (st), Gottsbergers.n. (INPA);Lago Arapari,Paranado Autaz-Mirim,12 May 1966 (? fl-fr), Mello 18 (INPA, U), 17 May 1966 (q fl-fr), Mello 40 (INPA);Lago do Inemazinho,Cambixe,6 May 1962 (? fr), Mello INPA 12732 (INPA,U); Rio Purus,Lago Marrah, Seringal Sao Clemente, 11 May 1975 (st), Prance et al. 23430 (NY, U, US); Lago do Janauari,2-5 Jun 1961 (? fr), Rodrigues et al. 2695 (NY) and 2723 (INPA, NY); Lago

do Rei, IgarapeGrande,5 Jun 1964 (? fl-fr),Rodrigues et al. 5892 (INPA);Mun.Careiro,Lago do Castanho,9 Jul 1972 (? fr),M. Silva et al. 309 (INPA,U); mouthof Rio Purus,Anori,2 Apr 1967 (2 fl), M. Silva 754 (MG, NY), 5 Apr 1967(? fr), M. Silva et al. 787 (MG,NY); Lago Curu9a,10 km below Itacoatiara,18 Jun 1977 (q fr), N. Smith 128 (INPA); Manaquiri,Jun 1851 (st), Spruce 1635 (P). PARA: Santarem,2 Feb 1933 (? fr), Ducke RB 23994 (B, U). RONDONIA:Rio Madeira, Calama,Apr 1980 (? fl-fr), Goulding 14a and 103a (MG);Rio Machado,Jan 1981 (? fl-fr), Goulding1137 (MG).RORAIMA:Rio Amajary,23 Mar 1948 (v), Fr6es 23120 (GH, IAN, NY, VEN, U, US).

Local names. Colombia:Amazonas:uvo de monte. Brazil: Amazonas: Caimbe, caimbe, caimbe miuido, caraipe,caraip6do igap6; Para:janita

NAMES Specimens examined. COLOMBIA.AMAZONAS: EXCLUDED Rio Loretoyacu,9 Oct 1961 (? fr), Idrobo4693 (COL); Sarcodiscus amazonicus Martius - see Miquel in Martius,Fl. Bras. 4(1): 111. 1853, nomen. 2 km N of Leticia, 27 Jan 1969 (? fl-fr), Sastre et al.

137

TAXONOMICTREATMENT

I~~~

l

dl

*

Fig. 31. Soroceaduckei.1. Leafytwig withstaminateinfloresceces(DuckeRB 23995).2. Leafytwig. 3. Leafytwig with infructescences.4. Leafy twig (Mello INPA12732). 5. Staminateflower.6. Stamen(Ducke1530). 7a. Pistillate flowerand bracts.7b. Pistillateflower.8. Fruitingperianth.9. Seed. 10. Embryo,large cotyledon. 11. Small cotyledonand radicle. 12. Radicle (Mello INPA12732). [By P. Pardoen,Utrecht.]

FLORA NEOTROPICA

138

connate;perianthtubular,4-dentateto -lobed,in fruit fleshyandyellowish;ovaryfree,stigmas2, equal,filiSorocea ilicifolia Miquel, in Martius,Fl. Bras. 4(1): form.Fruit (almost)free;exocarpmembranaceous; seedlarge,withoutendosperm; crustaceous; 114. 1853,see Burger,Lanjouw& WesselsBoer, endocarp partsbelowthe vascularized with 2 lateral thin, testa ActaBot. Neerl. 11:429. 1962.Typenot extant, hilum;embryowiththe cotyledonsequalandthick; identitydoubtful. radicleshort,apical. Miquel, in

SoroceacolombianaStandley,Trop.Woods19:39. 1929. = Trophisracemosa (Linnaeus) Urban.

Sorocea ilicifolia Miquel var.grandifolia

Martius,Fl. Bras.4(1): 213. 1853,as the typical

byMiquel(1854) ofPoulseniawasdescribed Material asOlmedia?armata.Manyspecieshavebeenincluded Kongel.DanskeVidensk. in Olmedia,whichinthecourseof timehavebeentransLiebmann, Soroceamexicana Math.Afd., Ser.5, 2: ferredto othergenera(Berg,1972, 1977a).In 1916 Selsk. Skr.,Naturvidensk. 335. 1851 = Trophismexicana(Liebmann)Bureau. Pittierestablished thegenericnameInophloeumforthis of Poulsenia thepublication Bull. Soc. Bot. species,butheoverlooked Soroceanitida(Allemao)Warburg, France49(Mem.3):644. 1913,basedonSoaresia aculeatabyEggers(1898)forthesamespecies.Standley arnata in 1933,buthadearlier nitidaAllemao = ClarisiaracemosaRuiz & Pavon. madethecombinationP the as Coussapoarekoi. species decribed (1919) Trop.Woods43:18.1935. Standley, Soroceastenophylla of Poulsenia alreadyinhistory The nomenclatural = Clarisiailicifolia(Sprengel)Lanjouw& Rossberg. features, dicatesthatthegenushascommonmoraceaous buthasotherfeatures asinflowersandinflorescences, Poulsenia somewhatunusual,as in its vegetativeparts.Thege8. PoulseniaEggers,Bot. Centralbl.73: 49. 1898; nusis characterized of prickleswith by theoccurrence Standley,Trop. Woods 33: 4. 1933; Burger, a hyalineapexonmostpartsof theplant.Theprickles FieldianaBot. 40: 199. 1977. Type.Poulsenia resemblethoseoccurringin Urticaceae(like Urera), aculeata Eggers (= Poulsenia armata (Miquel) butarefirmer. passintorigidhairs. Theymaygradually Standley). Onyoungplantpartsonecanfindbrown(moniliform) InophloeumPittier,J. Wash.Acad. Sci. 6: 113. 1916. pluricellular thosefoundinmany resembling trichomes, Type. Inophloeumarmatum(Miquel) Pittier ( inCecropiaceae). Ficus species(andmorepronouncedly Poulsenia armata (Miquel) Standley). Itis notclearwhetherParmatais mostlyandthroughmonoecious(withstamiTrees,monoeciousor dioecious,aculeate.Leaves outits rangeof distribution or simultaneously and inflorescences, nate pistillate laminapinnatelyveined;maralternate anddistichous; or dioecious. often present) subsequently stipulesfree,fullyamplexicaul. gin(sub)entire; InfloThe genus probablyconsists of two species, rescencespairedor solitaryin theaxilsof theleaves; andinvolucre- Poulsenia armata, rangingfromMexicothroughthe apexof thepedunclemostlybroadened like;staminateinflorescencescapitate(sub)globose, Andeanregionto Bolivia,andanotherspecies,found withfew interfloralbracts;perianth4-lobedto -fid; inFrenchGuiana,butnotyetdescribed,asonlysterile Poulseniacannoteasilyberelated is available. tepalsdecussate;stamens4, straightbeforeanthesis; material pistillodelacking;pistillateinflorescencescapitate, to one of the otherneotropicalArtocarpeaenor to generaof thetribe. (sub)discoid,withbasalbracts;flowers1-15, basally paleotropical varietyof S. ilicifolia.

Key to the speciesof Poulsenia 1. Lamina broadest below or in the middle; widespread, Mexico to Venezuela and to Bolivia ......... 1. P. armata 2. P. sp. 1. Lamina broadest above the middle; French Guiana........................................................................

1. Poulsenia armata (Miquel) Standley,Trop.Woods Fig. 32 33: 4. 1933. Olmedia? armata Miquel, in Seemann, Bot. Voy. Herald 196. 1854. Type. Panama. Darien: Without locality, (2 fl), Seemann s.n. (holotype, K). Inophloeum armatum (Miquel) Pittier, J. Wash. Acad. Sci. 6: 114. 1916.

Poulsenia aculeata Eggers, Bot. Centralbl. 73: 50, cum tab. 1898. Type. Ecuador. Manabi: El Recreo, 1892-1897 (d' + $ fl), Eggers 15651 (holotype, B; isotypes, BG, BM, C, F, GH, H, K, LE, MO, NT, 0, P, S, US). Coussapoa rekoi Standley, Contr. U.S. Natl. Herb. 20: 211. 1919. Type. Mexico. Oaxaca: Cafetal Concordia, 15 Nov 1917 (e'), Reko 3590 (holotype, US).

TAXONOMICTREATMENT

139

Fig. 32. Poulsenia armata. 1. Leafy twig (Steyermark92080). 2. Leafy twig with staminateinflorescences (Eggers 15651). 3. Staminateinflorescence.4. Staminateflower. 5. Stamens(Cazalet et al. 5058). 6. Pistillate inflorescence.7. Pistillateflower.8. Pistil (Steyermark61158). 9. Pistillateflower(Cuatrecasas17669). 10. Fruit. 11. Seed. 12, 12a. Embryo(Steyermark 61158). 13. Prickle(Miller1742). [By E. M. Hupkensvan der Elst and P. Pardoen,Utrecht.]

140

FLORA NEOTROPICA

Tree,to 30 m tall,oftenwithbuttresses, (sometimes Matuda3645 (A, F, K, MICH,NA, NY); 2 km from climbers?), on youngpartsaculeatewithprickleswith Estaci6nEl Suspiro,7 Sep 1967 (? fl), Penningtonet al. a hyalineapex.Leafytwigs2-10 mmthick,sparselyto 9148 (GH,K, NY); 5 km S of Palenque,11 May 1982(? denselypuberulous, withappressed, brownhairs,gla- fl-fr), Sousa et al. 12381 (MO). OAXACA: 3-4 km E of SantaMariaChimalapa, 27 Jul 1984 (? fl-fr),Herndndez brescent,sometimesalsohispidulous,oroccasionally G. 266 (MO);CerroConcordia,17 Apr 1933 (? fl-fr), alsowithlong,straightto uncinaterigidhairs,passing Mortonet al. 2719 (K, US);rd.to Pochutla,22 Nov 1967 intoprickles.Lamina(suborbicular) to ellipticto ob- (st), Pennington et al. 9329 (NY); Mun. Matias Romero, long (to lanceolate),5-55 x 3-25 cm,broadestat or 7.2 km W of Esmeralda,8 Mar 1982 (? fl), Wendtet al. belowthemiddle(injuvenilematerial somtimesabove 3617 (MO).TABASCO: 30 km S of Chontalpa,May 1963 themiddle),often? inequilateral, subcoriaceous tochar- (st),Le6n Cazaressn. (GH). VERACRUZ:SaltodeEyipantla, taceous;apexacuminate to subacute; baseacutetocor- SanAndresTuxtla,(st), Caizada 44 (F,GH,MICH,MO); 5 Jun 1981 (? fr), date;marginentireto repand,sometimesaculeate; up- PlayaEscondida,N of Sontecomapan, lowersurfacepuberulous Gentryet al. 32636 (MO);SanAndresTuxtla,Estaci6n persurface(sub)glabrous; withsparseto dense,appressed, brownhairs,glabres- BiologiaLos Tuxtlas,2 Jun 1983 (9 fr), IbarraM. 646 (MO),22 May 1970 (? fl-fr), Martinez-Calder6n 3007 cent, sometimesscabridulous, oftenaculeateon the (A, F, MICH,MO);7 km NNE of Sontecomapan, 1 Nov midribandthe lateralveins;venationalmostplane 1981 (st), Nee 22577 (F); 1 km S of AgustinMelgar,3 above,themidribandlateralveinsprominent beneath, Mar1984(? fl), Nee et al. 29813 (F,MO);rd.Catemacotheotherveinsalmostplane;lateralveins5-16 pairs; Montpio, 3 Sep 1967 (st), Pennington et al. 9114 (K, tertiaryvenationpartlyto largelyscalariform; petiole NY); 8 km NW of Catemaco,23 Jul 1985 (v), RoblesG 0.5-7 cmlong,coriaceous,oftenaculeate,puberulous 121 (F);Hidalgotitlan, CedilloJuntaat Rio Soloxuchil,I withdense,appressed, brownhairs;stipules(0.5-)l- Apr 1974 (?), Vdzquez276 (F); Fortuna,Rio Coatzawith(dense)brownpluricellular coalcos,Mar 1937 (st), Ll. Williams8426 (F, MICH). 8 cmlong,coriaceous, GUATEMALA. IZABAL:Escoba,W of PuertoBarhairs,usuallyaculeate,caducous.Staminateinfloresrios, 3 May 1939 (st), Standley72867 (F). PETtN: 8 mi cencesglobose,oblongoid-orobovoid-capitate, 0.5-3 w of Rio SanRam6n,NW of Chinaja,29 Mar 1942(st), x 0.5-2 cm;peduncle0.5-2 cmlong,puberulous; flowSteyermark 45505 (A, F, MICH);Rio Machaquila,N of ersfreeorbasallyconnate;perianth2-5 mmlong,the El Cambio,25 Apr 1942 (st), Steyermark45955 (F). upperpartpuberulouswith brownhairs,often also BELIZE. ToledoDistrict,nr. SapoteCamp,ca. 6.5 theoutertepals? cucullate; hispidulous, stamensoften mi W of MedinaBank,23-27 1976( fr),Proctor35958 filaments3-6 mmlong;anthers anisomorphic; 0.7-1.6 (MO);Rio Grande,MissouriRancho,23 Mar 1933 (ae), x 0.7-1.2 mm,theconnectivenarroworbroad,some- Schipp 1154 (A, BM, F, G, GH, K, MICH,MO, NY) timesapiculate; bractsoblongto linear,to 1.5mmlong. and 1155 (MO, NY). COSTA RICA. ALAJUELA: Reserva Biol6gica Pistillate inflorescencessubsessileorpeduncleto 1 cm Monteverde,Rio PeniasBlancas,8 May 1989 (d'+? fl), Bello 875 (BG, MO);Rio Peiias Blancasvalley, NE of withappressed puberulous to patent,brownhairs,ofSan Carlos, 29 Jun 1985 (st), Hammel et al. 14059 tenalsohispidulous, occasionally denselyhirtellous or (MO); 1 km S of Fortuna,10 May 1988 (ae), Hammel aculeateat the apex;ovary2-3 mmlong;styleca. 5 16838 (MO). CARTAGO: Turrialba,(st), Oersted 14345 mm long, puberulous;stigmas(0.2-)3-7 mm long; (F); Turrialba,PuenteCaj6n, 13 Apr 1972 (st), Poveda bractsa few to numerous(40-50), ovateto subulate, 29 (CR).LIM6N: Alta Soki,Amubri,Talamanca,20 Apr often? swollen,indumentum similarto thatof thepe- 1976 (st), Ginzbergs.n. (CR);Cord.de Talamanca,Rio rianth.Infiuctescences 1-4 cmdiam.;fruitingperianth Madrede Dios, 2 Sep 1988 (9 fl), Grayumet al. 8668 1.5-2 cmlong,withyellowishwarts;fruitca. 1.3-1.8 (BG, F, MO);nr.Matina,Mar 1980 (st), Lent4148 (F); x 1-1.5(-2) cm; bractsoften cushion-shapedand api- Rio Coen, Sep 1947 (st), Shank24 (F); Cant6nPococi, Cariari,4 Apr 1967 (2 fl-fr),Soels KSW-4=y-37(CR,F, culateoralmostspine-like,to 1 cmlong. NY). PUNTARENAS: Osa Peninsula,5 km W of Rinc6n, Distribution (see Fig. 78.3). FromsouthemMexico 24-30 Mar 1973 (st), Burgeret al. 9013 (CR, F); Osa to northeastem Guatemalaand southem Belize and Peninsula,CorcovadoNational Park,nr. Sirena Field from Costa Rica throughthe Andeanregion to north- Station,13Aug 1984(st), Gentryet al. 48499 (BG,MO); em Venezuela,to AmazonianBolivia, andto Acre(Bra- ParqueNacionalCorcovado,Sirena,LowerOllas,4 Apr zil); in wet forest; to ca. 1500 m, in Colombia to ca. 1988 (a'), Kernan 375 (BG, MO), 3 Feb 1989 (a), Kernan et al. 945 (BG, MO). 2000 m. PANAMA. BOCAS DEL TORO: Chinquinolavalley, Representative specimens examined. MEXICO. 1927 (st), Cooper et al. 88 (A, F, US) and 132 (US); CHIAPAS: Mun. Palenque, rd. Palenque-Ocusingo, km Almiranteregion, Daytona farm, Jan-Mar 1928 (st), 6-12, 10 May 1973 (9 fl-fr), Breedlove34978 (CAS, Cooper455 (F, K, NY); Isla Col6n, 16 Mar 1993 (st), Mo); nr. Palenque, 11 May 1982 (o'), Davidse et al. 20356 Fosteret al. 14538 (SCZ, see note);Almirante,29 Nov (MO, U); Javalinero, Palenque, 6-9 Jul 1939 (? fr), 1971 (st), Lao et al. 448 (U). COL6N: Pipeline rd., 10 long; flowers (I-)3-15;

perianth ca. 6 mm long,

TAXONOMICTREATMENT km NW of Gaboa,14 Dec 1973 (? fl), Berg et al. 356 (BG, U); QuebradaAncha, 4 km E of BuenaVista, 4 Nov 1973(? fl), Nee 7785(MO);4 kam NWof Salamanca, 12 km NE of Buenos Aires, 30 Dec 1973 (? fl), Nee 9087 (MO);rd. to Portobello,2 km N of PuertoPilon, 27 Mar 1974 (st), Nee et al. 10919 (MO);Rio Fatovalley, nr. Dos Bocas, 16 Aug 1911 (2 fl-fr), Pittier 4202 (K, US). PANAMA: BarroColoradoIsland, 1931 (e), Aviles 22 (F, MO), 18 May 1932 (R fl), Carpenter75

141 92080 (NY,in VEN: 57579); Rio Chichiriviche, between Portachudo and Penita, 1 May 1967 (st), Steyermark et

Hwy.,23 Jul al. 98263 (VEN).MtRIDA: Pan-American 1975 (st), Bernardi2353 (F, MER, NY); SantaElena,

27 Jan 1954 (2 fl-fr), Little 16211 (MER). SucRE: Dtto. Sucre, QuebradaEl Tigre, S of Fila La Baqueta, 21 Nov 1981 (9 fl and e"), Davidse et al. 19227 (MO, U, VEN). ZULIA: Dtto. Perija, 3.5 km SW of Los Angeles de Tukuko, 24 Jun 1980 (o'), Davidse et al. 18553 (MO, (F), 12 Feb 1969 (d), Croat 7914 (F, MO, NY); nr. El VEN); Quebrada Perayra, Rio Tokuku, Misi6n de Los Llano, 11 Nov 1973 (? fl), Nee 7951 (MO);Alhajuela, Angeles de Tokuku, SE of Machiques, 29 Aug 1967 (st), 25 May 1911 (st), Pittier3731 (BM, GH,NY, US). SAN Steyermark 99873 (U, VEN). BLAS: Play6nChico,4 Oct 1972 (? fl), Gentry6378 (F, ECUADOR. AZUAY: Nr. Manta Real, Rio Patul, 22

MO); El Llano-Cartird., km 19.1, 12 Mar 1985 (st), Neverset al. 5130 (MO);Cangandi,19 May 1985 (st), Nevers et al. 5748 (MO).

COLOMBIA.Withoutlocality, (? fl), Triana852 (P). ANTIOQUIA: Mun. Mutata, Rio Mutata, Finca Puentiadero-Finca La Palma, 5 May 1987 (? fl), Fonnegra et al. 2166 (BG). CHOC6: Rio Pichima,

Waunana,22 Nov 1976 (st), Forero718 (MO);Rio San Juan, nr. Tado, 15 Aug 1976 (o), Gentry et al. 17765

(MO,U); ElValle,27 Jun1958(st),Pintoet al. 107 (COL, P); Rio Sucio,Saltodel Rio Truando,2 Nov 1956(? fr), Romero-Castanteda6176 (COL).CUNDINAMARCA:Tena, (st), Anonymous s.n. (US). GUAJIRA: Rd. Mingueo-San Antonio, 16 Nov 1985 (st), Cuadros V 2415 (MO). MAGDALENA: BetweenMincaand San Lorenzo,10 Sep 1965 (q fl-fr), Espinal et al. 1784 (COL);Serraniade SantaMarta,SE of Minca,ca. 2000 m, 3 Jul 1984 (st), Gentryet a. 47608 (BG, MO);SierraNevadade Santa Marta, N slope, Feb 1948 (st), Romero-Castanieda 722

Oct 1994 (st), Cornejoet al. 3464 (BG).BOLIVAR: Hda. Changuil, Sector Nuevo Mundo, Oct 1995 (? fl-fr), Bonifaz et al. 3352 (BG);Bucay, 87 km E of Eloy Alfaro, 4 Jul 1943 (st), Little6735 (F, K, QAME, US). CARCHIf: San Marcos valley, ca. 24 Nov 1983 (st), Kvist et al. 48985 (BG). ESMERALDAS:Rio San Miguel, 28 Mar-6 Apr 1959 (a"), Harling 4676 (S); San Lorenzo, 20 Apr 1943 (st), Little 6297 (F, K, QAME,US);Businga, Rio Verde, 2 Oct 1965 (9 fl), Little et al. 21203 (F,MO, US); San Lorenzo, Finca La Chiquita, 8-15 Jul 1988 (st), Palacios 2594 (BG, MO); Cant6nSan Lorenzo, Ricaurte, 15-29 Apr (st), Rubio et al. 1399 (MO). Los Rios: Rio Paleque Biological Station, rd. Quevedo-Santo Domingo de los Colorados, km 56, 13 Sep 1973 (e), Dodson et al. 5386 (F, QCA),20 Aug 1976 (o), Dodson 6183 (MO); Jauneche forest, Cant6n Vinces, between Mocachi and Palenque, Estero Pefiafiel, 145 Aug 1978 (st), Dodson et al. 7075 (MO); Pichilingue, Exp. Station, 2 Aug 1962 (st), Jdvita et al. 301 (MO, S); Pichilingue, 21 May 1943 (? fr), Little et al. 6482 (A, F, K, NY, US), 23 May 1943 (? and a"),

(COL).META: Nr. Villavicencio,Llanode San Martin, Jan1856(st), Trianas.n. (BM).NARINo: Tumaca,Salisbi, Littleet al. 6499 (A, F, K, NY,US). MANABt: Machallila 8 Jun 1951 (d), Romero-Castanieda2656 (COL, F); Rio

Rosario,3 km fromRio Caunapi,5 Mar(st), RomeroMar1853(st), Triana Castaneda5586 (COL);Barbacoas, 1861 (COL, US). PUTUMAYO: Rio Putumayo,Puerto Porvenir,17 Nov 1940 (st), Cuatrecasas10618 (COL, F, US); Rio San Miguel o Sucumbios,SantaRosa, 7-8 Apr 1942 (st), Schultes 3617 (GH). QUINDtO: Mun. Circasia,Vrda. Los Pinos, 1765 m, 28 Dec 1989 (st), Velez et al. 908 (COL)and 914 (COL).TOLIMA: Mun. SantaIsabel,Vrda.La Pava,2030 m, 4 Aug 1980 (st), between Idrobo et al. 16583 (U). VALLE: Rio Chabirru, PuertoMerizaldaand Meregildo, 2 Mar 1943 (? fr), Cuatrecasas14342 (F); Rio Cajambre,5-15 May 1944 (? fl-fr), Cuatrecasas 17669 (F); Rio Cali, Pichinde, 1700 m, Jul 1946 (? fr), Duque-Jaramillo 3895 (COL); Tulua, Cgto. Monteloro, 1700-2300 m, 9 Nov 1972 (st), Jimenes Saa 1839 (U).

VENEZUELA. ANZOATEGUI: QuebradaNegra, tributaryof Rio Zumbador,NE of Bergantin,24 Feb 1945 (e + ? fr), Steyermark 61158 (F, G, RB, VEN). ARAGUA: ParqueNacionalHenriPittier,RanchoGrande,

16 Nov 1975 (st), Huber200 (VEN), 15 Jul 1963 (st), Merida 16 (VEN).DISTRITO FEDERAL: CerroNaiguata,

Las Delicias, (st), Bunting2095 (MY);CerroNaiguata, Lomasde las Delicias, betweenQuebradade Basenilla andQuebrada Guayoyo,15-19 Nov 1963(st),Steyermark

National Park, San Sebastiin, 8-9 km SE of Agua Blanca, 20 Jan 1991 (st), Gentryet al. 72439 (MO). NAPO:Rd. Puerto Napo-Puerto Misahualli, km 5-25, (st), Dodson et al. 14972 (MO, QCA); Parque Nacional Yasuni, rd. Pompeya Sur-Iro, km 38.7, 20 Oct 1994 (st), Jaramillo et al. 17280 (QCA);Cant6nArchidona,rd. Hollin-Loreto, km 31, Challua Yacu, 1200 m, 20-25 Mar 1989 (st), Palacios 4026 (MO); Rio Napo, 8 km downstreamPuerto Mishualli, 25-30 May 1985 (st), Neill et al. 6517 (BG, MO, QAME, QCA); Aniangu, mouth of Rio Afiangu, 30 Jun-9 Jul 1982 (st), SEF 10042 (QCA, QCNE, U). PICHINCHA: Rd. Santo Domingo de los ColoradosQuininde, km 170-175, 12 Sep 1949 (st), Acosta S. 13964 (F); Santo Domingo de los Colorados, 20 Sep 1930 (st), Benoist 3047 (P, U); 20 km W of Santo Domingo de los Colorados, 18 Oct 1961 (6'), Cazalet et al. 5058 (K, NY, US); Santo Domingo de los Colorados, 29 Oct 1960 (a" + 9 fl-fr), Pennington 6350 (K, NY). SucuMBfos: Rio Aguarico, Dureno, 5 Aug 1986 (st), Cer6n 349 (MO), 6 Mar 1937 (st), Pinkley 237 (COL, S). Rio Cenepa, nr. Huampami,5 km PERU. AMAZONAS: E of Chavez Valdivia, 1 1 Jul 1978 (st), Ancuash 1018 (MO, U). AYACUCHO:Tambo, Oserato, 17 Sep 1963 (st), Weiss178 (F). HUANUCO:Prov. Pachitea, Sungaro, W of Puerto Inca, 14 Sep 1982 (st), Foster 8796 (MO); Prov. Leoncio Prado, UTCF, 24 Nov 1979 (st), Lao M. 2a (U).

142

FLORANEOTROPICA

LORETO:Prov. Maynas, Tarapoto,ca. 20 km W of Iquitos, 21 Feb 1996 (st), Ruokolainen et al. 9467 (TUR). MADRE DE DIos: Prov. Tambopata, Comunidad Nariva de Infierno, Hermosa Chica, 19 Nov 1988 (st), Alexiades 118 and 155 (BG, MO); Rio Manui,Cerro de Pantiacolla, Rio Palotoa, 10-15 km W of Shintuya, 16-17 Dec 1985 (st), Fosteret aL 10944 (BG) and 11025 (F); Rio Manu, Cocha Cashu Station, 17 Oct 1979 (st), Gentryet al. 26877 (MO), 2 Aug 1983 (st), Gentry 43311 (BG); Prov. Tambopata, Cuzco Amazonico Tourist Lodge, 27 May 1989 (st), Nuiinezet al. 10528 (MO). PASCO: Prov. Oxapampa, Villa Rica, 1640 m, 19 Jan 1983 (st), D. N. Smith et al. 3131 (BG, MO). SAN MARTIN:Prov. Lamas, rd. Tarapoto-Yurimaguas, km 66, 27 Apr 1986 (st), Knapp et al. 7245 (BG, MO); Prov. Lamas, Alonso de Alvarado, E of San Juan de Pacayzapa, 11 May 1973 (2 fl and de), Schunke V 6232 (MO, U); Prov. Mariscal Caceras, NE of Tocache Nuevo, Rio de la Plata, 7 May 1975 (? fl-fr), SchunkeV 8411 (MO,U). UCAYALI: Prov. Coronel Portillo, San Alejandro, 6 Sep 1973 (st), Bellido s.n. (NY, US); Prov. Coronel Portillo, CarreteraMarginal, 22 km S of km 86 on Pucallpa-Tingo Maria rd., 11 Feb 1981 (st), Gentry 31239 (MO); nr. Brazilian border, QuebradaSapallal, base of Cerra Las Cachoeiras, 19 Jun 1987 (st), Gentryet al. 58459 (MO);SE of Pucallpa, nr. jct. of Rio Pachitea and Rio Yuyapichis, 10 Oct 1985 (st),

ca. 15 km (SE) up Rio Pitasama, from Rio Surutui,29 Aug 1985 (st), Solomon 14155 (BG, MO); Prov. Sara, Rio Surutui,Buenavista, 8 May 1981 (st), VazquezAvila 386 (U).

BRAZIL. ACRE: Mun. Cruzeiro do Sul, Reserva Extrativistado Alto Rio Jurua, Seringa Sao Joao, 19 Mar 1992 (st), Daly 7519 (NY). BOLIVIA. BENI: Prov. Yacuma, SE of San Borja, nr. Rio Cuverene, 13 Sep 1988 (Q fl), Beck et al. 16702 (BG); Prov. Ballivian, Serrania del Pil6n Lajas, 21 km from Yucumo, 1035 m, 17 May 1989 (9 fl-fr), D. N. Smith et al. 13169 (BOLV, MO); Prov. Ballivian, rd. Yucumo-Rurrenabaque, km 35, 20-25 Jun 1989 (? fl), D.N. Smith et al. 13567 (BG, MO); Prov. Ballivian, rd. Caranavi-San Borja, Serrania del Pilon Lajas, 1 Nov 1989 (? fl-fr), D. N. Smith et al. 13831 (BOLV, MO); Prov. Ballivian, Serrania del Pilon Lajas, rd. CaranaviSan Borja, 2 km from Yucumo, 18 Feb 1990 (Q fl), D. N. Smith 13981 (BOLV, MO); Prov. Moxos, rd. San Borja-Trinidad, km 27, 25 Aug-3 Sep 1990 (st), D. N. Smith et al. 14175 (BG). LA PAZ: Prov. Sud Yungas, Santa Ana de los Mosetenes, 21 Mar 1990 (st), Hinojosa 1102 (LPB); Prov. Sud Yungas, Rio Bopi, San Bartolome, 1-22 Jul 1939 (? fl-fr), Krukoff10282 (A, F, G, K, MICH, MO, NY, U, US); Prov. San Yungas, Rio Bopi, Asunta, nr. Evenay, 27-31 Jul 1939 (? fl-fr), Krukoff 10667 (F, NY). PANDO:Prov. Manuripi, rd. PorvenirPuerto Heath, km 26, 16 Aug 1974 (? fl-fr), Meneces 903 (BOLV). SANTACRUZ:Prov. Ichilo, ParqueNacional Amboro, 2.5 km ENE of summit of Cerro Ambor6, 10 Oct 1990 (st), Nee 39158 (MO); Prov. Ichilo, Parque Nacional Ambor6, Rio Isama ("Rio Pitasama"), 12 Oct 1990 (st), Nee 39262 (MO); Prov. Ichilo, Parque Nacional Amboro, 23 km S of Buena Vista, Rio Chonta, 18 Nov 1988 (9 fl), Nee 36848 (MO); Prov. Ichilo, Reserva Forestal Rio Ibabo, 16-18 Aug 1990 (? fl), Neill et al. 9368 (MO); Prov. Ichilo, ParqueNacional Ambor6,

Localnamesanduse.Mexico:Chiapas: chileamate, mazitoto(Zoque); Oaxaca: camero,chirimoya; Tabasco: amateblanco;Veracruz: abasbabi,chagane,comede pescado, huichilnma, CostaRica: guachilamo, guarilama. Cartago: mastate; Limon:kal-dtsi(Bribri), tetsi mastate, Panama: BocasdelToro:mastate; SanBlas: (Cabecra). ikor-wala(Kuna),omeganiko(Kuna);Colon:cucua; Darien:cocua or cucua. Colombia:Choco:cocua, damagua; Magdalena: cucua;Narifno: damaque, punte; Valle:corbon. Venezuela: Anzoategui: guaritota; Merida: habillo.Ecuador: higueron Carchi: mahana (Coaiquer); Esmeraldas: damagua,damajagua blanca;Los Rios: majagua; Manabi: majagua; Napo:matopalo; Sucumbios: k'arap'acha(Kofan). Peru:Amazonas:kamush; Ayacucho: HuanucoandLoreto:yanchama; Kiriniroki; MadredeDios:yanchama, maja-seji ormaja-sehi (Eseejhal),maja-nei(Ese-ejhal);SanMartinandUcayali: yanchama.Brazil:Acre:chanchama.Bolivia:Beni: corocho;SantaCruz:tachioreortachori. Theinnerbarkprovidesbarkcloththroughbeating andwashing.

In thenorthern partof the speciesrange(Mexico, andBelize)thestipulescanbe aslongas 8 Guatemala, cm;elsewhere thestipulescanbeto6 cmlongonquickly inPanama growingshoots,although andColombia they usuallyareno longerthan3 cm.Thepistillateinfloresinthenorthern cenceshaverelatively longpeduncles part of thespeciesrange,buttheyareto (sub)sessileinVenInthesouthern ezuela,Colombia,andEcuador. partof thespeciesarea(e.g.,Ecuador andBolivia)thenumber of pistillateflowersperinflorescence is oftensmall,2it increasesto 15. 3, butnorthwards Theperianth of thefemaleflowersin Cuatrecasas 14342and17669is somewhat deviating, asthetepalsare andpairwise theouter distinctly decussate, differentiated, ones? cucullate, theinneronesplanewitha thinapex, resembling theperianth ofthestaminate flowers.Thestigmasareextremely short,only0.1-0.2mmlong,inthese twocollections. ThecollectionFoster 14538fromPanama deviatesinhavingoblongtosubobovate resemlaminas, fromFrenchGuiana, as regarded Morawetzet al. V2-101085(BG);MonteReal, Rio San blingthoseof material to a distinct belonging species (see below). Alejandro, 22 Jul 1958 (st), Woytkowski5114 (MO).

2. Poulseniasp. Tree,to 25 mtall.Leafytwigs2-5 mmthick,sparsely andminutelypuberulous andwithstraightto slighty curvedprickles. Laminaobovatetooblanceolate, 6-25 x 2-10 cm;apexacuminate tocaudate; baseacute;margin bothsurfaces withprickles onthemain aculeate; glabrous,

143

TAXONOMIC TREATMENT

ArtocarpusJ. R. & G. Foster,Char.Gen.P1.ed. 1, 110,t. 51. 1775,nom.cons.;Jarrett, J.ArnoldArb. 40: 1-29, 113-155,298-368. 1959;41: 73-109, 113-140.1960;Jarrett, Blumea22:409-410.1975. Type.Artocarpus communis J. R. & G. Foster Specimensexamined.FRENCHGUIANA.Region (= Artocarpusaltilis (S. Parkinson) Fosberg).

veins beneath;lateralveins 4-8 pairs;tertiaryvenation scalariform;petiole 0.7-1.5 cm long; stipules 1.5-3 cm long, coriaceous,with sparsebrownpluricellularhairs, aculeate,caducous.Inflorescencesunknown.

de PaulIsnard,along trailCitron-Mt.Decou Decou, 10 Sep 1983(st), Cremers8207 (BG);regionde PaulIsnard, Trees,monoecious. Leaves alternate andin spirals; base of Mt. Lucifer,2 km N of Citron,9 Nov 1982 (st), laminapinnately veined;stipulesfree,frillyamplexicaul. Granville5237 (BG);regionPaulIsnard,Citron,15 Feb Inflorescencesunisexual,solitaryintheleafaxilsoron 1995 (st), Loubry 2022 (BG); region Paul Isnard, ontheolderwood.Staminateinflorescences branchlets BarthlermyCr.,20 Feb 1995 (st), Loubry2097 (BG). stamen1.Pistillate infloresspicate;perianth tubular; Thesterilematerialexaminedbelongswithoutdoubt cences globose-to ellipsoid-capitate; flowersconnate; to Poulseniaandlikely representsan additionalspecies perianth tubular; stigma1.Infructescenceslarge,with of this genus. theouterlayerfirmanda softinnerlayercontaining the

CultivatedSpecies of Artocarpus

fruits;seedlarge,withoutendosperm; cotyledons thick andequal.

A genuswith50-55 speciesof theAsianmainland, Two speciesof Artocarpus arecommonlycultivated andPacificOceanislands.Twospecies,treated in many partsof the Neotropics and often regardedas Malesia, thetropics. indigenous.Therefore,they arebriefly treatedhere. below,havebeenintroduced throughout

Key to the speciesof Artocarpusin cultivationin the Neotropics 1. Lamina pinnately incised .................... 1. Lamina entire ....................

1. A. altilis 2. A. heterophyllus

1. Artocarpus altilis (S. Parkinson)Fosberg,J. Wash. Tree,to 10(-15)cmtall.Leafytwigs 3-7 mm thick, Acad.Sci. 31: 95. 1941.Sitodiumaltile S. Parkinson, glabrous. Laminaellipticto obovateto oblong,6-25 x J. Voy. South Seas 45. 1773. marbaseacuteto rounded; 3-11 cm;apexacuminate;

lateralveins5ginentire;bothsurfaces(sub)glabrous; 10pairs;tertiary petiole1-2.5cm venationscalariform; to long;stipules1.5-8cmlong,appressed-puberulous Tree,to 35 m tall.Leafy twigs ca. 10-20mmthick, glabrous.Inflorescences mostlyon branchlets on the Laminaelliptictoobovate,ca.20-50 x 15- olderwood(mostlytrunkandmainbranches). puberulous. Stamibase(sub)acute; nate inflorescencesclavate,2.5-10 x 0.7-2.5 cm;peapexacuminate; 30(40) cm,coriaceous; to duncle1.5-5 cmlong.Pistillate inflorescences ellipmarginpinnatelyincised;bothsurfacespuberulous hirtellous,on theveinsto hirsute;lateralveins 12-16 soidto obovoidto oblongoid,5-10 x 3-6 cm,covered petiole3-6 cmlong; withconicalprocesses; venation scalariform; pairs;tertiasy peduncle5-10 cmlong,broadto ap- enedattheapex;stigma1-2 mmlong.Infructescences stipules8-25 cm long,hirsuteto subsericeous pressed-puberulous.Inflorescences on terminal 30-60(-l 00) x 25-40(-50) cm. toclavStaminateinflorescencescylindrical branches. Introduced fromIndia,as a fruittreein cultivation ate,8-30 x 1-4 cm;peduncle2.5-9 cmlong.Pistillate thetropics;boththeformwithfinnfruiting throughout 4-9 inflorescencesellipsoidto globoseto subobovoid, andthatwithsoftonesarein cultivation. x 3-6 cm,coveredwithconicalprocesses (orintheseed- perianths 4-7 cmlong;stigma lessform)rathersmooth;peduncle 1-2 mmlong.Infructescences15-30cmdiam. III. DORSTENIEAE ArtocarpuscommunisJ.R. & G. Forster,Char.Gen. P1.ed. 1. 102. 1775.

Trees, shrubs (of variousgrowthhabit),or (in Introduced fromPacificislands,Moluccas,orNew thetrop- Dorstenia) herbs, commonlymonoecious,less comthroughout asa fruittreeincultivation Guinea; ordioecious,mostlywithunciics;boththeseededandseedlessfornsareincultivation. monlyandrodioecious natehairs.Leaves alternate,in spiralsor distichous; laminaentireorincised,sometimespeltate;venation or(inpeltatelaminas)to radiate; 2. Artocarpus heterophyllus Lam., Encycl. 3: 209. pinnate,subpalmate to lateral. orsemiamplexicaul 1789. stipulesfullyamplexicaul

FLORANEOTROPICA

144

inoutlineorsecbisexualandorbicular Inflorescences ondarilyunisexualorshapedotherwise,discoidto turor(pistillate ones) multiflorous binateortocup-shaped, neverdistinctly (orebracteate), bracteate to uniflorous, bractsmostlypeltate.Staminate interfloral involucrate, flowers: tepals(l-)2-4, or the perianthvestigialor absent;stamens1-4, straightin thebud,orif inflexed, pistillode gradually; so orstraightening thenremaining mostlyabsent.Pistillateflowersmostlyconnatewith inthereceptacle, otherflowersand/orlargelyembedded rarelyfree;tepals2-4; ovaryfreeoradnateto theperianth;stigmas2, equallylong,butin Dorsteniaoften unequallylongoronly 1.Fruita dehiscentdrupe(let) wholewithotherflowersand! orforminga drupaceous or orthereceptacle; seedlargeandwithoutendosperm embryo inDorstenia(p.p.)smallandwithendosperm; withthickandoftenunequalcotyledons,or if small, thenflatandwithequalcotyledons.

withone speciesin Africa(Berg, 1977c);Trilepisium 1977c); Trymatococcuswith two species in the andUtsetelaPellegrin,withonespeciesin Neotropics; U. Africa(Berg,1977c),includingthenewlydescribed

The tribecomprises8 generaand 129 species: withonespeciesinAfrica(Berg,1977c); Bosqueiopsis, with15speciesintheNeotropics; Dorstenia, Brosimum, with105speciesof which47 arefoundintheNeotropics, onespecies,inAsia(India,SriLanka),andtheothersin Socotra,andYemen; Africa,Madagascar, continental with two speciesin the Neotropics; Helianthostylis, Baillon,withtwospeciesinAfrica(Berg, Scyphosyce

distinct theGuianaShieldregion,andinthatassociation Brosimum of theMoraceae. fromgeneraor subgenera doesnotshowclearaffinitiesto oneof subg.Brosimum regions. themajorphytogeographic bisexualand arepredominantly Theinflorescences Incontrast totheothertribes, oftencomplexinstructure. variationin poltheDorstenieaeshowa considerable lenfeatures(Hoen& Punt,1989).

neglecta Jongkind(1995; Berg, 1998b). The genus Dorstenia also includes Craterogyne,

some by Lanjouw(1935)to accommodate established describedunderTrymatoAfricanspeciesincorrectly is atpresent(Berg,1978c; Lanjouw coccus.Craterogyne Berg& Hijman,1999)in thesynonymyof Dorstenia sect. NothodorsteniaEngler (1898).

Therelationsbetweenthe moraceousfloraof the andthatofAfricaareclosestinthistribe.Asa Neotropics of theserelations itislargelya SouthAmericonsequence (Berg,1998b).However, canelementintheNeotropics corDorsteniahasa distinctcenteron thenorthwestem nerof theNeotropics. Brosimum subg.Ferolia(7 spp.), Helianthostylis,and Trymatococcusareassociatedwith

Key to the neotropicalgeneraof Dorstenieae 1. Plants herbaceous(or suffrutescent) 9. Dorstenia ............................... 1. Plants woody. 2. Plantsdioecious,or if monoecious,then the staminateflowerswith or withouta vestigial perianth......................... 10. Brosimum 2. Plants monoeciousor androdioecious;the staminateflowers with well-developedperianth. 3. Inflorescencesbisexualand cylindricalor turbinate,with the flowersat the upperend of 11. Trymatococcus the receptacle;pistillode minute...................... 3. Inflorescencesbisexual and globose or staminate;pistillode (usually)well-developed,with long stigmas...................... 12. Helianthostylis

Dorstenia 9. DorsteniaLinnaeus,Sp. Pl. 121. 1753;Plumier,Nov. P1.Amer.Genera29, t. 8. 1703; Houstoun,Philos. Transact.37: 195. 1731;Sprengel,Syst.Veg.3: 777. 1826; Nees von Esenbeck,Plant.Med. t. 98. 1828; Endlicher,Cat.Hort.Vindob. 1:241. 1842; Fischer & Meyer, Index Sem. Hort. Petrop. 11: 55. 1846; Miquel,in Martius,Fl.Bras.4(1): 159. 1853;Lernaire, Ill. Hort. 10: t. 362. 1863; Bureau,in De Candolle, Prodr.17: 258. 1873; Bentham& Hooker,Gen. P1. 3(1): 366. 1880; Engler,Monogr.Afr. Pfl. 1(Moraceae):5. 1898;Britton,Bull. TorreyBot. Club51: 7. 1924; Barker & Dardeau, Fl. Haiti 89. 1930; Rossberg, Notizbl. Bot. Gart. Berlin-Dahlem 12: 168. 1934; Sauget& Liogier, Fl. Cuba2: 54. 1951; Carauta,Valente& Sucre,Rodriguesia27(39): 225.

1974; Carauta,Bradea2(21): 149. 1976; Carauta, Rodriguesia29(44):53. 1978;Berg& vanLeeuwen, Proc.Kon.Ned. Akad.Wetensch.,Ser.C, Biol. Med. Sci. 85: 29. 1982;Carauta&Valente,Atas Soc. Brasil 1(20): 111. 1983; Berg, Proc. Kon. Ned. Akad. Wetensch.,Ser. C, Biol. Med. Sci. 89: 113. 1986. Lectotype,Britton& Wilson,Bot. PortoRico 5: 242. 1924. Dorstenia contrajervaLinnaeus. Sychinium Desvaux, MWm.Soc. Linn. Paris 4: 216. 1826. Type. Sychinium ramosum Desvaux (= Dorstenia ramosa (Desvaux) Carauta,Valente& Sucre). Dorstenia Linnaeus sect. Sychinium (Desvaux) Fischer

& Meyer,IndexSem. Hort.Petrop.11: 57. 1846; Carauta, Bradea 2(21): 151. 1976 (as Sychinia); Carauta, Rodriguesa 29(44): 88. 1978. Dorstenia Linnaeus sect. Lecanium Fischer & Meyer, Index Sem. Hort. Petrop. 11: 62. 1846; Carauta,

TAXONOMIC TREATMENT

Bradea2(21): 151. 1976 (as Lecania); Carauta, 29(44):65. 1978. Lectotype,Carauta, Rodriguesia Bradea2(21): 151. Dorstenia turnerifolia Fischer & Meyer. Dorstenia Linnaeussect. Emygdioa Carauta,Bradea 2(21): 151. 1976; Carauta,Rodriguesia29(44): 105. 1978.Type.Dorstenia brasiliensis Lamarck.

145

tropicalAmerican specieswereadded,suchasD. drakenaby Linnaeus(1759), D. brasiliensisandD. cordi-

folia by Lamarck(1786),andD. tubicinaby Ruiz& Pavon(1798).Inthebeginningof the 19thcenturythe genusattracted horticultural andbotanical interest based on the remarkable featuresof the inflorescencesof materialcollectedin easternBrazil(e.g., Fischer& Plants herbaceous orsuffrutescent, monoecious, to Meyer,1846;Hooker,1840,1841).Ithasbeenin cul1(-2) m tall,unbranched orsparselybranched; stems tivationin greenhouses, andmanyspecieswereillusmainlysupraterranean to entirelysubterranean; intern- tratedinbotanicalmagazinesin Europe. odeselongate,short,orbothshortandelongate;unciInthefirstcomprehesive treatment of thegenusfor natehairsoftenpresent.Leaves alternate andin spirals theNeotropics by Miquel(1853),16specieswererec(oralmost),oftensubrosulate; laminabasallyattached ognized,themajoritySouthAmerican. Fora whileinorpeltate,entireorpinnatelyto palmatelyorpedately terestmovedto theGreaterAntilleswheremanyspelobedto parted;venationpinnateto subpalmate or(in cieswerediscovered(Britton,1908,1924;Grisebach, peltateleaves)almostradiate,brochidodromous to re- 1860,1866;Urban&Ekman,1929).Rossberg(1934) ticulate; marginusuallydentate to crenate; stipulessub- provideda surveyof thespeciesof theGreater Antilles. foliaceous to subulate, plurinervate to uninervate, (sub)- Theinterestin theBrazilianspecieswasrenewedby persistent,mostlyfirmlycoriaceous.Inflorescences studiescarriedoutby Carautaandhis collaborators bisexual(butinD. cayapiausuallyunisexual), usually resultingin numerouspublications(Carauta,1972, solitaryin the leafaxils,pedunculate; receptacledis- 1974a,b,1976;Carauta et al., 1973a,b,1974a,b,1975, coidto turbinate to cup-shaped, orbicular, elliptic,qua- 1976a,b;Valente &Carauta, 1974,1975,1977;Valente drangular, (irregularly) stellate,lingulate,orbifurcate, etal., 1977),andmanynewspeciesweredescribed. A green,(partly)yellow,(partly)purplish, or(partly)red- study of the taxa in the northwesternpart of the entireto (irregularly) dish-brown, lobedto cren(ul)ate, Neotropics,asa precursory studyforthepresentreviusuallywith 1-5 rowsof smallto minutemarginalor sion,wastakenupby BergandvanLeeuwen(1982) submarginal bracts,sometimeson shortto ratherlong andwasfollowedby a precursory studyof thespecies marginalorsubmarginal oroccasionally of thesoutheastern appendages, partof theNeotropics(Berg,1985), withoutbractsandwithonly filiform(sub)marginal thelattercarried outincollaboration withCarauta (Berg interfloral appendages; bractslackingoroccasionally &Carauta,1985). a (narrow)fringeoftenpresentbetween rudimentary; Althoughthe AfricanDorstenia flora is more thefloweringfaceandtheedgeof thereceptacle; flow- speciosethantheneotropical one,it wasfora longtime ersconnate; hairsoftenpresentontheperianths, among knownonlyby a speciesdescribed by Forsskil(1775) theflowers,andon thefringe,minuteandgloboseto asKosariafoetida,basedon materialfromYemen. In orto conical,hyalineorpurplish.Stami- theendof the 19thandthebeginningof the20thcenclub-shaped nateflowers amongthepistillateonesorconcentrated tury,numerous newspeciesweredescribed,manyof in or confinedto theperipheryof thefloweringface, themby Englerbetween1894and1914. pedicellate (pedicelsconnateandadnateto theperianth Parallelto the studyon neotropicalrepresentatives of thesessilepistillateflowers);tepals2 or3(-4), (al- of the genus,revisionalstudiesof the paleotropical most)free;stamens2 or3, inflexedinthebud;pistillode specieshavebeen carriedout by Berg (1978c) and occasionally present.Pistillateflowerssessile;perianth Hijman,resultingin severalpapersandtreatments for tubular,onlytheupperpartfree,entire,2- or3-lobed; Africanfloras(Berget al. 1977,1984, 1985;Hijman, ovaryfree;stigmas2, shortlyfiliform,oftenunequalin 1980,1989,1990,1991).Thissituation created aunique length.Fruit a dehiscentdrupelet,exocarpwhiteand opportunity theneotropical andpaleotropical tocompare turgid-fleshy,at maturityejectingthe crustaceous, Dorsteniaflorason the basis of currenttaxonomic mostlytuberculate endocarp body;seedsmall;testathin, work.Thisresultedin a proposalfor the subdivision witha small,slightlythickened,vascularized partbe- of thegenus(Berg& Hijman,1997). low the hilum, endospermpresent;embryosmall, curved;cotyledonsflatandequalin size;radiclerela- MORPHOLOGY tivelylong. inhabitand Thegenusshowsconsiderable variation lifeformsandadiversityinreproductive structures (inHISTORY matchedonlyby a fewotherangiosperflorescences), The genuswas establishedby the descriptionof mousgenera. Dorstenia contrajerva and its variety houstonii from herbaceous Habit:Thegenusis a predominantly Mexico(Linnaeus1753).In the 18thcenturya few one in a family of woody plants. The woody habit is

146

onlyfoundin 11Africanspeciesintwosectionsof the genus.Incontrastto theNeotropics,varioustypesof succulencehavedevelopedin someof theAfricansectionsof thegenus. All neotropical speciesare(basically)herbaceous,

FLORANEOTROPICA

culencein supraterranean orsubterranean stemparts,a in dryhabitats(cf. featureconnectedwithoccurrence Engler,1898;Berg, 1977b,1978c;Berg& Hijman, 1977,1999;Hijman,1989).

Leaf:Thegenusshowsconsiderablevariationin theleaves.Thecaulescentspeciesusuallyhaveelliptic andlowerpartthe(leafy) tooblongleaves,mostlywithshortpetioles. tescent,as therhizomatous Thelamina stemcanbecomemoreor less woody.Thesespecies is entire,often lobed to partedonly in Dorstenia andmanyothershave(rather) slender,erect(toascend- choconiana. Theothercaulescentspecieswithlobed ing)stemswithlongintemodesandusuallyrathershort to partedleaves,D. ramosa andD. grazielae, have rhizomatousparts.They aremoreor less distinctly relativelylongpetiolesandarerelatedto a groupof short subcaulescent phanerophytic. As a ruletheleaveshaverelatively specieswithlongpetiolesandoftenincised laminas. In the group of subcaulescentto petiolesinthesecaulescenttaxa. Another groupof specieshasstemswithatleastsome (sub)acaulescent species,one canroughlyrecognize of theinternodes short.Thestemsareoftenforthegreater twogroups.Onegroup,including,e.g.,D. contrajerva Theleavesaremoreorless andD. arifolia, hasrelativelylargeleaves,thesebeing partorentirelysubterranean. rosulateandusuallyhaverelatively distinctly longpeti- oftenlobedto parted.Theothergroup,including,e.g.) oles. Subterranean partsof thestemwithshortintern- D. brasiliensisandthegroupofAntilleanspecies,have odescanbe moreor less tuberous.Thesespeciesare relativelysmallandentireleaves.withtheexceptionof to geophytic.Thetwocategoriesof D. cayapia p.p.,in whichthelaminamaybepalmately hemicryptophytic InDorstenia habitformsarenotwellseparated. ramosa, lobedto parted. oneof thesubspecies haslongintemodes, theothersubInthegroupof caulescent andinthegroupof largeleaved,subcaulescent species,thevenationis usually specieshasonlyshortinternodes. Inthemajorityof thespecimensof D. contrajerva, moreor less distinctlybrochidodromous, thuswith areshort,butoccasion- loop-connected atleastsomeof theinternodes lateralveins.Inthe subcaulescent to ally all intemodesarelong.Youngspecimensof D. acaulescent, small-leaved species,thevenationpasses withtheleavesrosulate; intothecraspedodromous bahiensisaresubacaulescent type(withlateralveins,ofolderspecimens haveelongateinternodes andtheleaves tenafterbranching atthemargin).Inlamiterminating habit naswithbrochidodromous distinctlyspaced.Twospecieswitha procumbent venation,partof thetertiary andlong,moreor less repent(andmoreor less rhi- venationcanbe scalariform (withmanyorsomeof terparallel,perpendicular zomatous)stems,D.flagellifera andD. belizensis,have tiaryveinsrunning to thelateral mostorall internodeslong,respectively. veins).Inthegroupof Antilleanspecies,peltateleaves Thesuffrutescent habitcanbe regarded as a primi- arecommon.Basallyattachedandpeltateleavescan tivestateandtheshortening of theintemodes asderived, occurin thesamespecies.Indistinctlypeltateleaves, withthe possible exceptionof Dorsteniabelizensisand thevenationpassesfrompinnateinto(alnost)radiate. D.flagellifera, inwhichthelongintemodes andtheproc- Thevenationis mostlypinnate,butsubpalmate insome umbenthabitcouldberegarded asderived.Thus,gen- of thespecimens ofD. cayapia andD. contrajerva.The eral trendsseem to be from a phanerophyticto a marginof thelaminais usuallycrenateto dentate. to a geophyticlife formandfroma hemicryptophytic Thestipulesarecoriaceous andpersistent in allspecaulescent tosubcaulescent toanalmostacaulescenthabit.cies.InDorsteniaelata,thestipulesaremostlydistinctly In theneotropical Dorsteniaspecies,monocaulyis a foliaceousandplurinervate. Largestipules,eitherindisdistincttendency. the tinctlyplurinervate Sometimes, e.g.,inD.turnerifolia, oruninervate, arefoundin several (supraterranean partsof the)stemsarebranched. species,e.g.,D. urceolataandD. arifolia. InthemajorMostspeciesoccurin humidandshadedplacesin ityof thespecies,thestipulesaresmall,mostlytrianguforest undergrowth.Only a few subcaulescentto larorsubulate, anduninervate. Insomeof thecaulescent acaulescentandgeophyticspecies,Dorsteniabrasil- species,thesmallstipulescanbecomepatenttodeflexed. iensisandD. cayapia,have(partly)"escaped" from The(sub)acaulescent formwithlong-petiolate leaves thishabitatto moreopenanddryvegetation,suchas andformswithlobedto partedlaminasarerarerin the savannaandcerrado;bothcanbehaveas weeds. Old WorldDorstenia flora, representedby, e.g., Theneotropical DorsteniafloradiffersfromtheOld D. barnimianaSchweinfurth andD. letestuiPellegrin. Worldonein lackingtrulywoodyfrutescent to almost Broadstipulesarefoundonlyin woodyspeciesandin arborescent forms,in lackingformswithdistinctlyand oneof theherbaceous species,D.picta Bureau(cf.Berg, oftenmuch-branched stems,andinlackingdistinctsuc- 1978c;Berg& Hijman,1977). althougha few species, like Dorstenia appendiculata, D. colombiana, and D. turnerifolia, can be suffruc-

TAXONOMIC TREATMENT

147

Inflorescence:Theinflorescences areusuallysoli- paleotropical ones,partlydueto thefactthatthemajortaryandborneintheaxilsof theleaves,in somespecies ity of thepaleotropical specieshavereceptacleswith onmoreorlessdistinct short-shoots withreduced appendagesandlack stipules. well-developed(sub)marginal Thereceptaclevariesin shape;it canbe orbicular, bracts.Moreover,the fringeof thereceptaclecanbe elliptic,obovate,moreorlessdistinctly (quadr)angular,verybroadinAfricanspecies(see Fig. 33D).Theinstellate,orfurcate.Themostextremeshapeof recep- florescences arealwaysbisexual,exceptinD. cayapia. tacle,thefurcateoneof Dorstenia ramosa, canbe re- The smallstaminateandpistillateflowersareintergardedas derivedfromthequadrangular one,suchas mingled,althoughthepistillateonesareoftenmoreor in D. contrajerva(Fig.33C). less concentrated in the centralpartof theflowering Thereceptacle is centrallyto eccentrically attached face.Ina few species(e.g.,D. turnerifolia), thestamito the peduncle.Its margincan be entire,dentate, nateflowersare(almost)confinedtotheperipheral part crenate,moreor less deeplylobed,or appendiculate. of thefloweringface.Anthesisof thepistillateflowers Thereceptacle and/orthestructure of themarginof the mostlypreceedsthatof thestaminate onesin thesame receptacle is oftenshapedaccording to thepositionand inflorescence.D. cayapiais the only speciesof the thesizeof thebracts;e.g,thelargerbractsarefoundon genusin whichunisexualinflorescences areusual.The thecornersof quadrangular receptacles oronthelobes staminate inflorescences appearfirston theplant,and of stellatereceptacles. whenthey have witheredthe pistillateones appear Inalmostallneotropical species,thereceptacle bears (Hoen,unpubl.thesisUtrecht).However,bisexualinsmallbracts,mostlyarranged inseveralrows.Theyare florescences sometimesoccur,buttheyshowcleardiffoundonthemargin,(just)belowthemargin,andon ferencesin theproportions of staminateandpistillate thefringe(thenarrowareabetweentheedgeof there- flowers. ceptacleandtheflower-bearing partof theflowering Justas in.the inflorescenceof therelatedAfrican surfaceof thereceptacle; seeFig.33D).Thebractsdif- genusScyphosyce (cf.Berg,1977a),thepistillateflowfer in dimensionsandshape.Especiallyin inflores- ersaresessileandthestaminate Inthe onespedicellate. cenceswitha singlerowof bracts,thelargerandthe saucer-shaped to cup-shaped receptacle,theperianths smallerbractsarealternating. Thebractsaremostly of thepistillateflowersareconnateand/oradnatetothe flowers(andin some appressed,butin severalspecies,especiallythemar- (fused)pedicelsof thestaminate ginalbractsarepatentandradiating,whichoftenac- cases also withthe stalksof interfloralbracts),thus centuates thestellateshapeof thereceptacle. Thebracts forminga layerof tissuewithcavities,containingthe onthefringeareoftenminute. (free)pistils(see Fig. 33A). In somespecies,teethandappendages Theinflorescenceorpartsof theinflorescenceare mayarise onorjustbelowthemargin,atplaceswherethebracts, green,yellowish,red-brown, orpurplish(toblackish). especiallythelargerbracts,aresituated.InDorstenia In somespecies,thepeduncleandall partsof thereappendiculata, the larger bracts are born by ceptaclearecolored,inotherspeciesonlythepeduncle, semiorbicular to linear,to 1.2 cm long appendages, onlythemargin,oronlythefloweringface.Sometimes colored.Thecolorsand whicharisebeloworalsoatthemargin.InD.grazielae, onlythestigmasare(purplish) themarginalbracts(oralsothosebelowthemargin) thepartsthatarecoloredoftenvarywithinthespecies. aresometimesbornonto 2.5 cmlongfiliformappendFlowers:Theflowersareratheruniformin thegeages.Filiformappendageson (orbelow)themargin of thepistillatefloweris tubularand arefoundin all specimensof D. ramosa. As in many nus.Theperianth paleotropical species,bractsareusuallynotformedin the apex is sometimesminutely3-lobed (e.g., in D. ramosa, and,thus,themarginof thereceptacleis Dorstenia turnerifolia).The pistil is free. The stigmas thanbracteate(see Fig. 33B). aresmall,filifonn,andequalorunequalin length. ratherappendiculate flowersof neotropicalspecieshave West Thestaminate bractsareonlyfoundinthe(primitive) Interfloral Africanspecies,D. djettiiGuillaumet (cf.Berg,1978c); sometimesthree,mostlytwo,small,almostfreetepals Thestatheyarepeltateandtheupperpartcaducous(seeFig. andthreeortwo,smallstamens,respectively. bractssometimes mensaresmall.Thefilamentis oftenswollenatthebase 33A).However,vestigalinterfloral occurin D. turnerifolia fromBrazil.The flowering and inflexed before anthesis,becominggradually (sur)faceof thereceptaclemaybe turneddownwards, straightat anthesis,andoften elongating.A minute to side- pistillodecansometimesbe found. sideways,or it canchangefromdownwards Theminutehairson the perianthsandalso those waysorfromsidewaysto upwardsin thecourseof the of theinflorescence orduringanthesis. amongtheflowersareoftengloboseto clavate.They development andmaycause Dorstenia speciesshowmuchless arehyalineandpurplishorred-brown Theneotropical variationin the shape of the receptaclethan the glisteningof (partsof) thefloweringface.

148

FLORANEOTROPICA

A~~~~~ *nf'fl. ~

.'es)

it 'a

SI

B_D ApSSf

o

DIbantImMes

.

S

e

*

CVb

,

.

.

it

e...

.

;;X A,mdas

t7e. i ? np.ndqe

'

w fthat

s

Fig. 33. Schematic drawings. A. The basic structure of the inflorescence of Dorstenia. B-D. The differentiation

of the inflorescenceof Dorstenia.B. Developmentof appendages.C. Differentiationof the shape of the inflorescence. D. Differentiation of the position and the presence of bracts, the development of appendages and the expan-

sion of fringeof the receptacle.

TAXONOMICTREATMENT

149

of theovulesvary.Insomeofthespecies,the sistsof moreorlessswollentissueinsteadof twodistinct Features in inothers,the integument layers.Theembryosacshowsdifferences nucellusis surrounded bytwointeguments; andpolarnuclei. antipodes, of synergids, sideof thecoverof thenucellustowards thefuniclecon- thenumber Surveyof dataon ovulesandembryosacsof neotropicalspeciesof sect.Lecanium: D. turnerifolia Integuments partlyfuseda incl.D. argentata Integuments 2a D. hirta Integuments partlyfuseda D. panamensis Integuments 2b D. elata Integuments 2a D. urceolata Integuments partlyfusedc Surveyof dataon ovulesandembryosacsof neotropicalspeciesof sect.Dorstenia: Integuments 2; polarnuclei2b Integuments 2; polarnuclei2b Integuments 2; polarnuclei2; synergidspresent; antipodes10/20b D. drakena Integuments 2; polarnuclei2; synergidsabsent;

D. bahiensis D. ramosa D. contrajerva

antipodes5/lOd

Surveyof dataon ovulesandembryosacsof neotropicalspeciesof sect.Emygdioa: Integuments 2; synergidsabsentorpresent; antipodesOb 2b D. tenuis Integuments D. cayapia subsp.asaroides

aSchleuss, 1958. bHoen, 1983 (unpubl.thesis), based on materialin the BotanicalGardenat Utrecht.

cDe Granville, 1971. dModilewsky, 1908.

bodyis surfaceof theendocarp ejected.Thetuberculate (cf.Overbeck,1924; relatedto theejectingmechanism woodyAfrican Schleuss,1958).Inthemacrospermous species,the endocarpbodyis smoothandis simply species,thesurface squeezedout.Ina fewneotropical is (almost)smooth,asinD. cayapiawith Pollen:A studybyHoenandPunt(1989)onpollen oftheendocarp largeendocarp bodies.Thesmallfruitconof Dorsteniaandothergeneraof thetribeDorstenieae itsrelatively Nine typesof tainsa seedwitha thintesta,witha slightlythickened revealsa considerabledifferentiation. partbelowthesmall Dorsteniapollencouldbe recognized,somerareones andwitha (poorly)vascularized is presentandtheembryohassmall, andsomemorecommon.Onthebasisof 18trendsin hilum.Endospenn in flat,andequalcotyledonsandarelativelylongradicle. thepollentypescouldbebe arranged differentiation, Africanspecies,theseedsdo Inthemacrospermous of pollenmatchesonly twogroups.Theclassification is large,withthick endosperm andtheembryo notcontain partlythegroupingof speciesasproposedhere. andoftenunequalcotyledonsanda smallradicle.Inthe epiphyticspeciesD.astyanactisAkeAssi, drupe(let), WestAfrican Fruit:ThefruitofDorstenia isadehiscent of andis expelledbyelongation stipitateandwitha whiteexocarp.Thefruitis initially thedrupeis indehiscent 1971). baseof thefruit(cf.deGranville, of thefruitandelongation thestipitate enclosed,butby enlargement of thestipeitbreaksthroughtheuppersurface(= flowTheupperpartof thetur- CYTOLOGY eringface)of thereceptacle. Thechromosomenumbersrangefrom2n = 24 to engid-fleshyexocarpsplitsandkeepsthecrustaceous docarpbodybetweenthe two halves,whichcan be 2n= 72 (Krause,1931;Le Coq,1963,1964;Federov, thetops 1969;Hoen,unpubl.thesis1983),withx = 12 and13 releasedwhenthevascularbundleconnecting of twohalvesoverthetopof theendocarp bodybreaks. as basicnumbersforAfricanspeciesandx = 14, 15, Dorstenia and16forAmerican species,andincludediploids,tripInthesmallfruits,asfoundinallneotropical andis loids,tetraploids, hexaploids,andaneuppentaploids, bodyis usuallytuberculate species,theendocarp also InAfricanspecies,bothtypesof integuments occur;polarnucleicanbe twoorfour;synergidscan be occasionallyabsent,andthenumberof antipodes variesfrom0 to 25.

150

FLORANEOTROPICA

inpositionof theflowersandpossibilities loids,as listedbelow.Themostcommonbasicnum- bydifferences pollination. However,somespecies bersinthefamilyarex = 13and14(cf.Federov,1969). for"autogamous" seedsevenafterthestigmashavebeenremoved produce of inflorescences (Hoen,unpubl. thesis),andthepistillate Chromosome numbers:sect.Lecanium: (thefunctionally dioecious)Dorsteniacayapiaproduce D. turnerifolia 32a seedswithoutthepresenceof pollenvectors.Thesedata incl. D. argentata 28b theoccurrence ofsometypeofagamospermy. Both suggest D. hirta 28a,c autogamy andagamospermy couldexplainvariation pat32a D. panamensis terns,asahighdegreeof localmorphological uniformity D. elata 26,c 32a,d evenover andstriking differences betweenpopulations, D. urceolata 32d,, shortdistances. De Granville (1971)hasobservednor32a var.variegata malfertilization inD. contrajervaandD. embergeriG. Chromosome numbers:sect.Dorstenia: Mangenot, butdidnotfindseedproduction afterremoval 32a D. bahiensis of thestigmas,andsuggestedthatpollination is needed 32a D. arifolia Hoen(unpubl. forreproduction. thesis,1983)foundthat 32c D. ramosa ingreenhouse differences conditions, probably largely light 30a,c,d D. contrajerva levels,didinfluence ("spontaneous") seedproduction. 30a D. drakena Vegetativereproduction,which occursin some branchlets Africanspecies(byarrested (semi)succulent Chromosomenumbers:sect. Emygdioa: thatmaybecomesmalltubersandgetdetached)is not 32a D. cayapia subsp. cayapia foundin theneotropical species. 32a D. cayapia subsp. asaroides tobe a commonphenomenon in Autochory appears 32a D. tenuis Dorstenia species.As the speciesoften growalong aHoen, 1983 (unpubl.thesis), based on material stream(let)s, further transport of ejectedendocarp bodin the BotanicalGardenat Utrecht. iesmayoccurbyrunning water.Thegenusdoesnothave b Le Coq, 1964. obviousmeansof long-distance-dispersal. the Moreover, cKrause, 1930. seedsappearto be viablefor only a shorttime.The dKrause,1931. Sudanianannualspecies,D. annua Friis& Vollesen eMangenot& Mangenot, 1962. withregard (1982),ispresumedly exceptional tothelongevityof theseeds,normallyviablefora shorttime. Chromosomenumbersfound for Africantaxa are: 2n = 24, 26, 28, 36, 40, 42, 48, 52, ?64, 72.

DISTRIBUTION

POLLINATION ANDDISPERSAL REPRODUCTION: Nothingis knownaboutthepollinationof Dorstenia. Severalfeatures,such as shapeof the inflorescence,ap-

causedbythecharpendages, colors,glistening surfaces hairsontheperianths andoftenelsewhereon acteristic theuppersurfaceof thereceptacle,and,in severalcases,

canbeassociated withmyophily; thereareno fragrance, recordsconfirmingit. Visits of beetles have been observedin some species in Camerounand Gabon(pers. comm.Hijman).Onemay wonderwhetherthedifferentiationandvariationin the inflorescencehas any adaptive significanceat all (cf. Berg 1977a). As the staminateandpistillateflowers occurmixed andcrowded,so thatstigmascan get in touchwith pollen, one may assumethatgeitonogamousself-pollination will often happen.Thereis a tendencyfor anthesis of pistillate flowers to preceedthatof staminateflowers. In several species, there is a distinct time lapse betweenanthesisof pistillateandstaminateflowers,but in others, there is no or minimal lapse. Severalspecies kept in greenhousesproduceseeds othersnot.Thisdifferencecannotbe explained profuisely,

Thedistribution of theneotropical Dorstenia flora showssomepecularities. TheNeotropics canbedivided into two (almostexclusive)domains,of whichthe boundaries aredefinedbythedistributions ofD. brasiliensis andD.contrajerva(seemaps,Fig.80.6& 81.8). The areasof these two specieshardlyoverlap.The southeastern domainhasitscenterineasternBrazil,the northwestern domainhasitscenterin northern Central AmericaandtheGreater Antilles. TheeasternBrazilian centercontains20 species,includingthemostprimitiveneotropical Dorstenia species.Thenorthern Central American andAntillean centercontains19species:12in theGreaterAntilles and5 on thecontinent.IntheAndeanpartof theD. contraisolatedspejerva domaintherearefourgeographically cies, clearlyrelatedto eachotherandto two central Americanspecies.The threeneotropicalsectionsof in bothdomains.Thereare Dorstenia arerepresented in themorphological no essentialdifferences diversity of thetwogeographically separated groupsof species. Itis striking thatDorsteniais almostabsentfromthe Guianaregion.Thewidespread D. brasiliensis is the

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FischerandMeyer(1846)designeda subdivision onlyspeciesfoundinthisregion.IntheAmazonBasin, thegenusis represented onlyby themoreorlesswide- basedon thegrowthhabitandcharacters (mainlythe spreadgeophyticspeciesD. brasiliensisandD. cayapia shape)of thereceptacle: subsp.asaroides,inplaceswithmoreorlessopenveg1.Sychinium,withonly Dorsteniaceratosanthes etation,andby thetwophanerophytic forestspecies, (= D. ramosa); D. peruviana and D. umbricola,close to the Andes. 2. Lecanium,comprisingthecaulescentneotropiThereareonlytwowidespread species,Dorstenia calspecies; brasiliensisandD. contrajerva.Dorsteniacayapiaand 3. Dorstenia,comprisingthe subacaulescent neoD. drakena,relatedtoD. brasiliensisandD. contrajerva tropical species;and respectively, havemoderately extensiveareas.Theother 4. Kosaria,comprising thecaulescentandacaulesspecieshaverelativelysmallto verysmallareas.Sevcentpaleotropicalspecies, all with a radiate eralspeciesareknownonlyfroma singlelocalityora receptacle. few neighboringlocalities. The greatnumberof Thissubdivisionwas adoptedby Walpers(1848(sub)endemic speciesis striking, beinginclearcontrast 1849)and,recently, largelysobyCarauta (1976,1978a). to thewoodyspeciesof thefamily. Carauta sect.Lecaniaandsect.Sychinia, recognized Themajorityof thespeciesarecomponents of the undergrowth of (rain)forests.Theyoftenoccuron anddividedFischer& Meyer's"section"Dorstenia slopesnearstream(let)s, ofteninplaceswithrocks.The intosect.Dorstenia(withD. contrajervaandseveral Antillean species and Dorstenia belizensis appearto relatedspecies)andsect.Emygdioa(withD. brasilsubdivibe(almost)confinedtocrevicesof steeplimestone rocks iensisandseveralrelatedspecies).Carauta's sion was on the habit based life form growth and and and cliffs. Dorstenia lindeniana and D. contrajerva of theinflorescenceandtheleaf. mayoccurinthesamehabitat.OnlyD. brasiliensis and on characters It is surprisingthatMiquel(1853) neglectedthe mostsubspecies of D. cayapiaarecomponents of open, basedoncharacters of theinfloD. drakenaand previoussubdivisions andcerradovegetation; savanna(-like) rescenceby combiningspecieswitha furcaterecepD. excentricacanoccurin (rather)dryforest. tacle(DorsteniaramosaandD. ceratosanthes)and to ellipticreceptacle(D.arispecieswithanorbicular CONSERVATION Inthetreatfolia) intoa singlespeciesD. multiformis. Smallor very smallrangesof distribution, some mentofDorsteniaforFloraBrasiliensis (1853),Miquel speciesbeingknownonly froma singlelocality(as groupedspeciesaccordingto thegrowthhabitinthree Dorstenia belizensis andD. conceptionis), andrather groups: strictecologicalrequirements, mostlysuitable placesin 1.frutescent to suffrutescent plantswithstemswith primaryforest,meanthata goodnumberof Dorstenia long intemodes; species are seriouslythreatenedby extinction(cf. 2. suffrutescentplants with stems with short Carauta & de Castro,1982).Oneof thesespecies,D. and intemodes; brevipetiolata,may be extinctalready. 3. subacaulescent plants. SUBDIVISIONOF THE GENUS

In 1863Lemaireproposedto subdivideDorstenia Thefirstsubdivisionof thegenuswasproposedby into: Sprengel(1826).Atthattime,thespeciesknownwere 1.Furcatae,comprisingthespecieswitha furcate neotropical,exceptforDorsteniaradiata(Forsskal) receptacle; Lamarck(= Kosaria radiata ForsskAl)from Yemen. 2. Quadratae, comprisingthe neotropicalspecies wasbasedonthegrowthhabit: Sprengel'ssubdivision withquadrangular toorbicular and receptacle; 1.Scapigerae, theneotropical comprising species; 3. Radiatae,comprisingthepaleotropical species and witharadiatereceptacle. 2. Caulescens, with Dorstenia radiata.

Endlicher'ssubdivision(1842) was basedon the shapeof theinflorescence: 1. Dorstenia,comprisingall species except for one;and 2. Sychinium, with only Dorstenia ramosa (= SychiniumramosumDesvaux, 1826), a species

witha furcatereceptacle.

When Bureaumade his importantrevision of Dorstenia(1873),the richDorsteniafloraof Africa becamebetterknown.Bureau,however,didnotproposea formalsubdivision,butonly groupedthespeAfterseparating theNew ciesbasedonkeycharacters. WorldandOldWorldspecies,he keyedoutgroupsof oracaulescent; base species,basedon:stemscaulescent of thestem(forthecaulescent OldWorldtaxa)rhizoma-

152

tous,tubiferous, ortuberous; inflorescences orbicular orellipticalto (!) furcate;laminaspeltateorbasallyatortritached;andstipulesfoliaceousandplurinervate angularandpungentto subulate. Ina treatment of theAfricanrepresentatives of the Dorsteniaintothree genus,Engler(1898)subdivided (African) sections: (1) Nothodorstenia, (2) Eudor-

FLORA NEOTROPICA

Bazzemia,resemblesin habitmembersof theneotropicalsectionEmygdioa. TwoAfricanspeciesbelongto thelargelyneotropical sectionLecanium. Thethreeneotropical sectionscannotbesharplydelimitedbecauseof thelackof verycleardiscontinuities in morphological (andecological)variation. 1.SectionLecanium comprises distinctly caulescent (suffrutescentto herbaceous)species, with spaced, mostlyshortlypetiolateleaves.All membersof this of theundergrowth grouparecomponents of rainforest.Thespeciesof thisgroupoccurin easternBrazil, theAndesregion,andCentral America. Thegroupcom-

stenia,and(3) Kosaria.This subdivisionwas based of bractson thereon thepresenceandarrangement ceptacle,thenumberof stigmata,andpartlyoncharactersof thestemandgrowthhabit. Itis clearthatthesubdivisions weremainly proposed basedonthegrowthhabit(andthelifeform)andchar- prises species like Dorstenia appendiculata and D. whichcanberegarded as themostprimiTheweaknessof thebasis turnerifolia, actersof theinflorescence. is clear,astheylackconsistency in tive ones amongthe neotropicalspecies,due to the forthesubdivisions stem,thepresenceof threestamens,and theuseof characters andarelargelybasedon material suffrutescent the occasional occurrence of interfloral bracts. fromonlyoneof themajorareasof distribution. This section the also comprises species whichshow A classification of Dorstenia,basedonfloristicand distinct with African relations Dorstenia species: the studies for both New and Old revisional World,is showssimilarities to thefrutescentAfproposedby BergandHijman(1999). In the whole appendiculata ricanspecies,D. dorsteniodes(Engler)Hijman& C. genus,ninesectionscanbe recognized: C. Berg;D. turnerifoliashowssimilaritiesto African 1.sectionNothodorstenia Engler,withfivewoody D. subdentata Hijman& C. C. Berg; andD. urceolata speciesin Africa(mainlyWestAfrica); resemblesAfricanD.picta.Thus,thetransatlantic af2. sectionXylodorstenia Hijman,withsix woody finitiesareclosestbetweeneastern Brazilian speciesand speciesinAfrica(mostof themin WestAfrica); speciesoccuringin Cameroun andGabon. 3. section Lecanium, with 22 non-succulent WithinsectionLecanium onecanrecognizea group caulescent herbaceousspecies, 20 in the of specieswithsmall,triangular to subulate, uninervate Neotropics,andtwoin (West)Africa; stipulesanda groupof specieswithrelativelylarge, 4. section LomatophoraHijman,with 26 non- (broadlyto narrowly) ovate(inDorsteniaelataalmost caulescent succulent herbaceous speciesinAfrica foliaceous) stipules, often with (distinctto faint) (mostof theminWestandCentral Africa); plurinervate venation.Thegroupwithsmallstipules 5. section Dorstenia,with eight non-succulent, comprisesthecaulescentspeciesof CentralAmerica partlyacaulescentherbaceousspecies in the andtheAndeanregion(thespeciesnumbered 10to 17) Neotropics; andseveralof theeasternBrazilianspecies(thespecies 6. sectionKosaria(Forsskal)Fischer& Meyer, numbered1 to 9); theymostlyhavehirsuteto hirtelwith 16 (semi)succulentcaulescentspecies, lousindumentum on theupperleafsurface.Thegroup includinganannualoneandanepiphyticone, withlargestipulescomprisestheeasternBrazilianspein Africa(mainlyEastAfrica),withextensions cies, D. elata, D. hildegardis, and D. urceolata. This to Madagascar, Yemen,SaudiArabia,Socotra, groupshow distinctrelationswith the (partly)subSriLanka,andIndia; caulescentspeciesincludedin sectionDorstenia. 7. sectionBazzemia withonenon-succulent Hijman, 2. SectionDorsteniacomprisessubcaulescent (to acaulescent speciesinAfrica(Mozambique); specieswithlarge,oftenlobedtoparted leaves 8. section Emygdioa, 18 non-succulent,partly caulescent) withlongpetioles.Somespeciespairsofthisgroupshow acaulescentspeciesintheNeotropics;and remarkable similarities andparallel variation. Whenster9. section Acauloma Hijman, three succulent fromD. acaulescenttubiferousspeciesin EastAfrica ile,Dorsteniaarifoliacanhardlybe separated ramosa ssp. ramosa, nor D. contrajerva from D. andSocotra. drakena.Dorsteniaarifolia andD. drakenahave inflo-

Inmostcases,thedelimitations of theseinfrageneric rescenceswithanellipticreceptacle. InD. contrajerva, subdivisionsarenot veryclear-cut.Six sectionsare thereceptacle is basicallyquadrangular; thefurcatereandincludetwosectionsof woody ceptacleof D. ramosacanbe regarded entirely paleotropical as derivedfrom to suffrutescent taxa,two sectionswith succulentor a quadrangular one.Especiallydepauperate specimens semisucculent taxa,andtwowithherbaceous non-suc- of D. drakenaand D. contrajervashow similaritiesto culent taxa. One of these, the monotypicsection

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153

representatives of thefollowingsection.Thegroupdoes peculiarvariationpatterns)unsolved.Thestillscarce numbersand notshowresemblances to anyof theAfricanspecies. dataaboutthevariationin chromosome the(probable)occurrenceof agamospermy pointsto 3. SectionEmygdioa comprisesspecieswithrelathe necessityof biosystematicstudies,which may tively smallleavesandlong petiolesandstemswith solve,at leastpartly,the problemsandthe questions mostly all or many internodesshort. Dorstenia indicatedby thepresentstudy,in particular thoserebelizensis andD. flagellifera areexceptional because garding Dorsteniadrakena. InAfrica,manymorecomof thepresenceof only longinternodes.Severalspeplex situationsexist(cf. Berg&Hijman,1999). cies of thissectionhavea geophyticlife formandare adaptedto moreor less dryandopenvegetation(D. Local Names and Use brasiliensisandD. cayapia p.p.).WithinthisthirdsecIn Mexico andCentralAmerica,thecommonvertion,the 13 speciesof theGreater Antilles(including nacularnameforDorsteniacontrajervaandD. drakena D. belizensis fromBelize)constitutea moreor less is contra hierba or contrahierba (or variantscontra distinctgroupof species,beingcloselyrelated,partly In theseregions,thisnameis yerba or contrayerba). in arathercomplexway.Exceptfortheirhabitat preferused also for otherDorsteniaspecies, (sometimes?) ence,theyarecharacterized byoftenhavingverysmall such as D. lindeniana. Dorstenia drakena is also leavesandby peltatelaminas.Oneof themembersof known bythenamesbarbodilloorbarboriain Jalisco thisgroup,D.fawcettii, showsremarkable similarities and Sonora,Mexico,respectively.In Panama(San to D. tenuis,occuringin thesouthemmost partof geoBlas),D. contrajervais knownby the Kunaname graphicrangeof thegenus.TheMexicanD. excentrica nanduli and in PuertoRicoa as tusilla. showsclose resemblances to D. brasiliensis. In secInBrazilthemostcommonnameforallspecies,but tionEmygdioa,therareD. conceptionisis morphologiinparticular fortheacaulescent orgeophytic species(e.g., callyratherisolated.Thissectiondoesnot showdisDorstenia brasiliensis and D. cayapia), is caiapia or tinct relations with the AfricanDorstenia flora. variantssuchas caapia, capia',carapa, carapia,caripa, However,one of theAfricanspecies,D. zambesiaca somevariantis alsousedinadjacent is in habitsimilarto the carpales,caxapia; Hijman(fromMocambique) countries andParaguay). Othervernacular (Argentina onein sectionEmygdioa. prevalent Systematics

namesusedin Brazilareapii, chupa-chupa,contra-erva, figueira terrestre,figueirilha,figueirilho, liga-liga, liga osso, tarope,teiu'(or teiu-apu),and tiu'(or tiu'-a9u).

Thesubterranean partsof theplantareusedto preThepresentstudyis almostentirelybasedonmorappliedformanymedicinal purposes, a rather phologicalcharacters, unsatisfactory approach, pareconcoctions as it leavesmanytaxonomicproblems(relatedto the e.g.,againstsnakebites(cf. Peckolt& Peckolt,1890). Keys to the neotropicalspecies of Dorstenia Becausethespeciesof Dorsteniainhabitalmostexclusivelytwodomainsin theNeotropics,twokeyshave whichco-occurin beenconstructed. Thereareonlytwowidespread species,D. brasiliensisandD. contrajerva, andin Trinidad andtheadjacentpartof Venezuela. Peru,nearTarapoto, Key to the species of the southeasternpart of the Neotropics Argentina, Paraguay, Guianas, Thiskeycoversthedomainof Dorsteniabrasiliensis, including Brazil,Uruguay, andTrinidad, alsopartlyBolivia,Peru,andVenezuela. 1. Intemodeselongate,to 1 cm long or longer,the leaves ? spacedalong the stem. 2. Stipulesat least 0.5 cm long. 9. D. setosa 3. Stipules subulate................................................... 3. Stipuleselliptic, ovate, or (broadly)triangular. 4. Petiole (8-)15-60 cm long; base of the laminacordate. 5. Receptacle furcate............................ ...................... 26a. D. ramosa subsp. dolichocaula 5. Receptacleorbicularto elliptic. 6. Receptacleeccentricallyattached;internodesrathershort,occasionallyto 1.2 cm ..........................25. D. arifolia long, 5-15 mm thick;laminaoften pinnatilobateto -partite 6. Receptaclecentrallyattached;internodes(normally)to 1.5 cm long, 4-10 mm thick; laminaentire, rarelypinnatelylobed............................................. 22. D. grazielae 4. Petiole 2.5-12 cm long; base of the laminacordateto subacuteor to subattenuate.

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154

7. Laminawith (11-)15-22 pairs of lateralveins; stipules 1-2 cm long, distinctly internodesto 8 cm long; receptacle(usually)elliptic, at least 1.5 cm plurinervate; 18. D. elata broad ...................................................... 7. Laminawith 6-13 pairsof lateralveins; stipules0.5-1.2 cm long, faintlyplurinervate to uninervate;internodesto 2.5 cm (or to 4 cm) long; receptacleorbicularor angular,if elliptic then at most 1 cm broad. 8. Bractsca. 10-12, in a single submarginalrow, spathulate,(0.5-)2-3 mm long, 24. D. bonijesu

radiating ...................................................

or to lanceorows, ovate to semiorbicular 8. Bractsnumerous,in 1-2 (sub)marginal late, to 2 mm long, appressed. ................................................... 19. D. urceolata 9. Receptacle cup-shaped,red-brown 9. Receptaclediscoid to broadlyturbinate,green or partlypurplish. petiole 3.5-8(-10) cm long; 10. Base of the laminaacute to (sub)attenuate; receptaclediscoid, ca. 1 cm diam .............................................20. D. hildegardis 10. Base of the laminacordateto subacute;petiole 10-12 cm long; receptacle discoid to broadlyturbinate,1-4 cm diam.............................21. D. bahiensis 2. Stipulesto 0.5 cm long. appressed;bractsspathulate,(0.5-)2-3 mm long, radiating...23. D. albertii 11. Stipulesovate, plurinervate, 11. Stipulessubulateor (narrowly)triangular,? patentor deflexed;bractsovate to semiorbicular, to 2 mm long, usuallyappressed. 9. D. setosa 12. Petiole 4-14 cm long ............................................................ 12. Petiole 0.3-3(-4) cm long. 4. D. brevipetiolata 13. Petiole 0.3-0.5 cm long; internodesto 5 cm long........................................ 13. Petioleat least 0.5 cm long; internodesto 2 cm long. angularto stellateand 3-4 cm 14. Base of the laminaattenuate;receptacleirregularly 5. D. contensis ................................................ diam.;penduncleto 3 cm long ... then the 14. Base of the laminaacute, obtuse,rounded,cordate,or if subattenuate, receptacleorbicularor to 2 cm diam. or the penduncleto 1 cm long. 15. Stamens3 in all or manyflowers;staminateflowersdistinctlyconcentrated on the peripheryof the floweringface. 16. Receptacleirregularlystellate,with ca. 5-6 semiorbicularto linear,submarginalappendagesto 1.2 cm long........................................1. D. appendiculata 16. Receptacleorbicularto slightly angular,with at most numerousand ...................................................2. D. turnerifolia very short marginalappendages 15. Stamens2; staminateand pistillateflowers (usually)intermixed. 17. Receptacleorbicularto slightly angular,green; marginalbractsabout equal in size, 1-3 cm diam.;peduncle1.5-6 cm long ............................3. D. hirta often partlypurplish, 17. Receptacleangularto substellate(to suborbicular), often 0.3-1.5 cm diam.;marginalbractsdistinctlyunequalin size; peduncle 0.5-1.5(-2)

cm long.

18. Base of the laminacordate. 19. Peduncle 1.5-4 cm long; receptacle(I-)1.5-2.5 cm diam.; 9. D. setosa petiole 1.5-4 cm long, hirsute....................................... 19. Peduncle0.7-1.5(-2) cm long; receptacle0.3-1(-1.3) cm diam.; petiole 0.5-1(-2.5) cm long, hirtellous............................. 8. D. milaneziana

18. Base of the laminaacute, obtuse,or rounded. 20. Laminalanceolateto oblong (to subovate),to 10 x 3 cm; petiole 0.5-1.5(-2.5) cm long; Rio de Janeiro,Minas Gerais,Sao Paulo .......................................

6. D. bowmaniana

20. Laminaoblongto elliptic(to ovate),to 16 x 7 cm; petiole (0.5-) 1-4 cm long; Sao Paulo, Parana,SantaCatarina ...................7. D. carautae 1. Intemodesshort,at least some of them, the leaves (sub)rosulate. 26. D. ramosa 21. Receptacle furcate....................................................................... to irregularlylobed. or quadrangular 21. Receptacleelliptic to (sub)orbicular 22. Receptacleeccentricallyattached;laminausually 10-30 cm long, often pinnatelyincised .25. ....................................................................................................................................................

D. arifolia

22. Receptaclecentrally(or slightly eccentrically)attached;laminausually to 15 cm long, entire or palmatelyincised. 23. Lamina3-parted(sometimes3-lobed to subentire);receptacle2.5-3 cm diam.; inflor24. D. bonijesu escences bisexual........... 23. Laminaentire, or if 3-7-lobed to -parted,then the receptacle 0.5-2 cm diam. and the inflorescencesunisexual.

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24. Stems and leaves glabrous......................................................... 21. D. bahiensis 24. Stems and/orleaves hairy. 25. Inflorescencesusually unisexual;stipules obtuse to rounded;endocarpbody smooth;laminaovate to cordifornto reniform,entireor palmatelylobed to 30. D. cayapia parted................................................... 25. Inflorescencesalways bisexual;stipules acute to acuminate;endocarpbody mostly tuberculate;laminaelliptic to oblong to (sub)ovateor cordifonn,entire. 26. Laminacordiformwith a wide sinus; lateralveins 4-6 pairs;receptacledisc31. D. tenuis ................................................ oid, often (sub)stellate 26. Laminaelliptic to ovate to oblongto subobovate,or if the base (sub)cordate, thenthe sinusnarrow;lateralveins at least6 pairs;receptacleoften? cupshaped, (sub)orbicular. 27. Lateralveins of lamina6-7 pairs,? distinctlyloop-connected,or if not so, then not furcate;stipulesconspicuous ..................................29. D. conceptionis 27. Lateralveins of lamina6-7 pairs,usuallyterminatingin the marginand furcate,if faintlyloop-connected,then at least 10 pairsof lateralveins; stipules inconspicuous.............. ............................... 32. D. brasiliensis

Key to the species of the northwesternpartof the Neotropics This key covers the domainof D. contrajerva, includingthe Antilles, CentralAmerica,Andeanregion, and northemVenezuela. 1. Internodeselongate,to 1 cm long or longer. 2. Laminapeltate...................................................................... 38. D. belizensis 2. Laminabasally attached. 3. Petiole 3-20 cm long; laminabroadlyovate to cordiform 27. D. contrajerva .......................................... 3. Petiole at most 5 cm long; laminaelliptic to lanceolate,to subobovateor to subovate. 4. Laminaat the base attenuateto decurrentwith narrowwings along the upperpartof the petiole............................................................. 10. D. choconiana 4. Laminanot attenuateto decurrentat the base. 5. Laminascabrousabove with long bristle-likehairs,the apex almostroundedto 17. D. lindeniana ....................................................... broadly (sub)acute 5. Laminasmoothabove,or if scabrous,then with minuteconicalhairs,the apex acuminateto acute. 6. Stipulesbroadlytriangular to ovate or to lanceolate,(usually)patentto deflexed. 7. Petioleca. 1 cm long; stipulesto 0.7 cm long; Panama ....................16. D. panamensis 7. Petiole0.7-4(-9.5) cm long; stipulesto 0.25 cm long; Colombiaor Mexico. 8. Laminamorethan 5 cm broad,the base cordateto rounded;Colombia ........................................................................................................... 15. D. colombiana

8. Laminaless than5 cm broad,the baseacuteto obtuse;Mexico.... 11. D. uxpanapana 6. Stipules narrowlytriangularto subulate,appressed. 9. Base of the lamina(sub)acuteto obtuse;receptacleoutside sparselyand/or minutely puberulous. 10. Uppersurfaceof the laminascabridulous;receptaclesuborbicularand centrallyattached;Peru, Ecuador.............................................. 12. D. umbricola 10. Uppersurfaceof the laminasmoothor nearlyso and glabrous;receptacle ? angularin outlineand eccentricallyattached;Mexico.... 11. D. uxpanapana 9. Base of the laminacordateto truncate,or if subobtuse,then the receptacle outside ratherdensely hirtellousto puberulous. 11. Tertiaryvenationof the laminaprominentand conspicuousbeneath; 14. D. aristeguietae ............................................. receptaclelobed; Venezuela 11. Tertiaryvenationof the laminaalmost plane (and ratherinconspicuous) beneath;receptaclesubentire;Peru, Bolivia...................................13. D. peruviana 1. Internodesshort,at least some of them. 12. Receptacleeccentricallyattached,at least 0.5 cm diam. to 4 cm broad,entire;Mexico..... 33. D. excentrica 13. Laminaoblongto elliptic'(to ovateor suborbicular), to elliptic or to broadlyovate, usuallymore than 4 cm 13. Laminacordiformto subsagittiform broad,mostly lobed to parted. Mexico to Costa Rica......................28. D. drakena 14. Receptacleelliptic to obovateor suborbicular; to irregularlylobed; Mexico to Peruand Venezuela,West 14. Receptaclequadrangular Indies.............................27. D. contrajerva

156

FLORANEOTROPICA

12. Receptacle(almost)centrallyattached,or if eccentrically,then the receptacledeeply cup-shaped and less than 0.5 cm diam.;GreaterAntilles. 15. Laminapeltate. 16. Receptaclediscoid;bracts(almost)in a single (sub)marginal row, all or most of them radiatingand/oron short appendages. 17. Laminawith only shorthairs above;Jamaica.................................................. 36. D. jamaicensis 17. Laminawith both shortand long hairsabove;Cuba............................................. 35. D. petraea 16. Receptaclecup-shapedto discoid;bractsin (2- )3-4 rows, appressed. 18. Stems (partlycreeping)with many long internodes;laminausuallydistinctlylonger than broad; Haiti......................................................

37. D. flagellifera

18. Stems with most internodesshort;laminamostly not distinctlylongerthan broad. 19. Receptacledeeply cup-shaped,with bractson and below the margin,not on the 46. D. rocana fringe; Cuba................................................... 19. Receptacleshallowlycup-shapedto discoid, with bractsalso on the fringe. 20. Largermarginalbractssetulose;stigmasof equal length, 0.4-0.5 mm long; Cuba, Hispaniola................................................ 42. D. erythrandra 20. Largermarginalbractsnot setulose;stigmasof unequallength, 0.1-0.2 mm long. 21. Receptaclecup-shapedto turbinate;laminawith rathersoft shorthairs above; Cuba, DominicanRepublic................................................ 41. D. peltata 21. Receptaclediscoid;laminawith stiff hairsabove;Cuba. 22. Stems 3-5 mm thick;lamina0.5-5 cm long, above with ratherlong stiff hairs;perianth(almost)glabrous...................................40. D. nummularia 22. Stems 4-8 mm thick;lamina(3-)5-10 cm long, above with short stiff hairs;perianthwith dense, minute(sub)globosehairs............39. D. roigii 15. Laminabasally attached. 23. Receptaclediscoid,the bracts(almost)in a single (sub)marginal row, all or most of them radiatingand/oron short appendages(or sometimesalso with minuteappressedbracts on the fringe). 24. Laminaovate to (sub)cordiform, to 7 cm long, the apex mostly subacuminateto obtuse;endocarpbody distinctlytuberculate;Jamaica,Haiti..............................34. D. fawcettii 24. Laminacordiformto discoid,to 2.2 cm long, the apex mostly rounded;endocarp 45. D. tuberosa body (almost) smooth;Cuba............................................................ 23. Receptaclecup-shapedto discoid, the bractsin (2-)3-4 rows, appressed. 25. Receptacledeeply cup-shaped,with bractson and below the margin,not on the 46. D. rocana fringe; Cuba...................................................... 25. Receptacleshallowlycup-shapedto discoid, with bractsalso on the fringe. 26. Plantson all partswith only minutehairs;Haiti......................................... 43. D. caimitensis 26. Plantson severalparts,at least on the laminaabove and/orthe marginalbracts, with long hairs. 27. Receptaclewith entiremargin;marginalbractsdistinct,the largerones setu42. D. erythrandra lose; Cuba,Hispaniola ................................................ 27. Receptaclewith crenulatemargin;marginalbractsobscureand not setulose; Cuba ................................................

SECTION LECANIUM 9A. DorsteniaLinnaeus sect. LecaniumFischer & Meyer, Index Sem. Hort. Petrop. 11: 62. 1846. Dorstenia sect. Lecania Carauta,Bradea 2(21): 151. 1976. Lectotype,Carauta,Bradea2(21): 151. 1976. Dorstenia turnerifoliaFischer & Meyer. Dorstenia Linnaeussect. Dorstenia series CaulescentesEndlicher,Gen. P1.Suppl.4(2): 35. 1848. Lectotype, designatedhere. Dorstenia nervosa Desvaux(= DorsteniaurceolataSchott). Plants herbaceousto suffrutescent,caulescent;stem mostly unbranched;internodeslong. Leaves in spirals or tendingto distichous;laminamostly oblong to ellip-

44. D. crenulata

tic or (sub)ovate,entire(to pinnatilobateto -fid); venation (basically)brochidodromous; petiole mostly short, sometimes long; stipules broad (and plurinervateto uninervate)to subulate.Inflorescencesactinomorphic (to slightly zygomorphic),mostly orbicularto elliptic, sometimessubquadrangular, substellate,or irregularly lobed,bracteate(bractssometimeson appendages),the fringe narrow;pistillate flowers numerous.Endocarp body mostly tetrahedraland tuberculate,sometimes subgloboseandsmooth;seed small,endospermpresent; embryowith flat and equal cotyledons. This section comprises 22 species. Only two of them, Dorsteniapicta andD. subdentata,areAfrican;

TAXONOMICTREATMENT

theyoccurin the Guineo-Congolese region.Twelve speciesoccurin easternBrazilandeightspeciesare scattered fromBoliviato Mexicoandthecoastalmountainsof Venezuela.All speciesarecomponents of the undergrowth of rainforest. Becauseof thecharacters of theinflorescence and the occurrenceof suffrutescent forms(in Dorstenia appendiculataandD. turnerifolia)thissectionlinksup withtheAfricansectionNothodorstenia,oneof thetwo sectionsof woodyspecies,andmay be regardedas derivedfromthelatter. Withinthe section,two groupsof speciescanbe recognized: 1. a groupwith ratherthick stems,with broad, often plurinervatestipules, and with rather thick, somewhatfleshy leaves, mostly with inconspicuousindumentum,representedby, e.g., Dorstenia elata andD. urceolata; 2. a groupwithratherslenderstems,sometimes moreor less woody at the base, with small, often subulatestipules,and with ratherthin leaves, often with hirtellousto hirsuteindumentum,representedby, e.g., Dorstenia turnerifoliaandD. choconiana.

157 in ca. 3 rows, on the margin,on the apices of the appendages,anda few to severalon the fringe,oblong to broadlyovate,to 0.7 mm long,withminuteconicalhairs; staminateflowers among the pistillateones, but ? distinctlyconcentrated in theperipheryofthe floweringface; tepals 3 or 2; stamens 3 or 2; filaments as long as or shorterthanthe perianth;stigmas not seen. Endocarp bodyca. 4.5 x 3.5 mm, slightly tuberculate. Distribution (see Fig. 79.5). In eastem Brazil(Bahia, Espirito Santo, and Minas Gerais); in moist and shadedplaces; at low elevations. Specimens examined. BRAZIL. BAHIA: Mun. km 7.3, 25 Jan 1996, Uru9uca,rd. SerraGrande-Itacare, Amorimet al. 1941 (NY); Mun. Itacare,6 km SW of Itacare,S of mouthof Rio de Contas,Jan 1977,Harley et al. 18345 (K, SPF, U); Mun. Maraui,rd. MarauUbaituba,km 26, Mattos Silva et al. 475 (RB); Mun. FazendaBarra Ilheus,at km 10 fromrd.I1h6us-Olivenga, do Manquinho,5 Feb 1982, Mattos Silva et al 1428 (GUA);Mun.Ilheus,6 km fromOliven9a,12 Feb 1994, Piraniet al. 2952 (SPF,NY); Mun. Itacare,rd. ItacareUbaituba,km 6, 12 Apr 1980, Plowman et al. 10085 (U), Santoset al. 3558 (GUA, RB); Mun. Uruguca,rd. SerraGrande-Itacare,km 7.4, 9 Feb 1994, Thomaset al. 10334 (NY). ESPiRITO SANTO: Mun. Cachoeirodo Itapemirim,FazendaSanto Antonio da PedraBranca, 27 May 1949, Brade 19913 (RB, U). MINAs GERAIS: Carangola,FariaLima,Novaes SP 19606 (SP).

1. DorsteniaappendiculataMiquel,in Martius,Fl. Dorstenia appendiculata is a distinct species and Bras.4(1): 162. 1853;Bureau,in De Candolle, Prodr.17:269. 1873;Carauta, Rodriguesia29(44): in severalfeatures,e.g., in habit,lifeform,andshapeof 79, t. 10. 1978.Type.Brazil.Bahia:RiodeContas, thereceptacle,is reminiscentof membersof anAfrican SerraGrande,Nov 1818,Martius2141 (holotype, group of (suf)frutescent species, especially D. M;isotype,U). Fig. 34 dorstenioides (Engler) Hijman & C. C. Berg (cf.

Hijman,1980).The speciesis characterizedby therelaSubshrub(orherb),to 1.5mtall;stem2-4 mmthick, tively large (and broad) appendagesdepartingbelow with? retrorse to 2 the marginof the receptacle. hirtellous uncinate hairs;internodes cm long.Leaves in spirals,buttendingto distichous;

to subobovate, 6-25 x 1.5laminaoblongto lanceolate 6 cm,chartaceous; apex(sub)acuminate (toacute);base 2. Dorstenia turnerifolia Fischer & Meyer, Index acuteto attenuate (? decurrent); marginfaintlyandirSem.Hort.Petrop.11:63. 1846;Miquel,in Martius, crenate-dentate to subentire; regularly uppersurfacealFl. Bras. 4(1): 164. 1853; Bureau,in De Candolle, lowersurface mostglabrous, oftenscabridulous; sparsely Prodr.17: 268. 1873; Carauta,Rodriguesia29(44): puberulouswith ? retrorseuncinatehairs, often 82, t. 13. 1978. Type.Brazil.Rio de Janeiro:Rio de lateralveins10-18pairs,loop-connected; Janeiro,Corcovado,Riedel & Luschnath1011 (hoscabridulous; venation tertiary reticulate; petiole0.3-1.4cmlong,hirlotype, LE). tellousto hispidulouswith straightto curvedhairs; DorsteniaargentataJ. D. Hooker,Bot. Mag.t. 5795. stipulessubulate,0.1-0.2 cm long,coriaceous,? de1869; Bureau,in De Candolle, Prodr.17: 267. onminute flexed,uninervate, puberulous. Inflorescences feui1873;A. Dalliere,Les plantesornamentales withreduced 0.3short-shoots Ilage panacheet colore 1: t. 23. 1873; Carauta, stipules,green;peduncle Rodriguesia29(44): 80, t. 11. 1978. Type. Dewithretrorse uncinate 0.8cmlong,sparselypuberulous scribedfrommaterialcultivatedin Hort.Saunders ircm 0.8-3 hairs;receptacle centrallyattached, diam., and Hort. Kew, collected in Brazil, nr. Rio de regularlystellatein outline,with5-12, to 1.2cmlong, materialin K; progenystill in Janeiro;herbarium almostlineartotriangular (tosemiorbicular) appendages cultivationin severalEuropeanbotanicalgardens. belowthemargin,theoutsidesparsely departing puberSubshrub(or herb),to 50(-140) cm tall, sometimes ulouswithuncinatehairs,themarginentireorfaintly bracts branched;rhizomeshort;stem 2-3 mm thick, puberuthefringeto 1mmbroad; lobatetoappendiculate,

FLORA NEOTROPICA

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TAXONOMICTREATMENT

159

lousto hirtellous, partlyormainlywithuncinatehairs; tins, property of Roberto A. Kautsky, 11 Jun 1985, intemodes to 1.5(-2)cmlong.Leavesinspiralsortend- Kautsky883 (GUA);Castelo,FomoGrande,6 Dec 1956, ingtodistichous; lamina(narrowly) lanceolate tooblong, Pereira 2117 (HB). RIO DE JANEIRO:Mun. Marica, (5-) 7-17 x (l-)1.5X-4 cm,thinlychartaceous, sometimes Itaipuacui,Pico Alto Moirao, 14 Apr 1982, Andreataet al. 438 (GUA);Mun.Campos,Morrode Coco, Cruzde variegated; apexacuteto acuminate; base(sub)acute to Serra,base of Morrodo Bau, 13 Dec 1982, Carautaet attenuate; margin rather coarselyandirregularly crenate- al. 4421 (GUA); Mun. Rio de Janeiro, Jacarepagua, dentateto subentire; uppersurfacepuberulous tohirtel- Florestados Tres Rios, 7 Jan 1952, Duarte 4164 (RB, lous,mainlywithuncinatehairs,scabrous; lowersur- U); Mun.Rio de Janeiro,Florestada Tijuca,9 Mar1864, facepuberulous tohirtellous withstraight anduncinate Glaziou777 (BR,P); Mun.SantaMariaMagdalena,Alto hairs,? scabrous; lateral veins(7-) l0-17, ? faintlyloop- Imb6,Rio Imbe,ca. 1000 m, 18 Feb 1981,Martinelliet connected; tertiary venationreticulate, inconspicuous; al. 7589 (GUA,RB); Mun.Petropolis,ReservaPatydo petiole0.5-2 cmlong,sometimes purplish, puberulous Alferes, 1 Nov 1970, Sucre 7340 (RB). to hirtellous, mainlywithuncinatehairs;stipulestrianDorstenia turnerifoliais a rathervariablespecies. gular,to 0.2 cm long,uninervate, soondeflexed,gla- It almostcertainlyincludesD. argentata,basedon culbrousorsparselypuberulous. Inflorescences solitaryor tivatedmaterial,andis differentfromthe "normal"form often2-3 togetherin theleafaxils,withthepeduncle of the species, mainly in the conspicuouslyvariegated palegreen,thereceptacle darkgreenoutsideandtheflow- laminaandthe purplishmarginof the receptacle.The eringfacebrightgreenorthemarginpurplish; peduncle morphologicaldifferentiatingcharacterswouldjustify 0.5-1.5(-2.5)cmlong,hirtellous tohispidulous, mainly distinctionof a variety.However,the cultivatedmatewithstraight hairs;receptacle centrally attached, discoid, rial on whichD. argentatawas based apparentlyhas a suborbicular to? angular orfaintlylobedinoutline, (0.S-) chromosomenumberdifferentfrom thatof cultivated 1-2.5cmdiam.,theflowering faceplaneorslightlycon- materialof the "normal"D. turnerifolia,withoutvariecave,the outside(sparsely)puberulous to hirtellous, gatedleavesandentirelygreenreceptacles,2n = 32 and mainlywithuncinatehairs,themarginentireto crenu- 2n = 28, respectively (cf. Krause, 1931). lateordent(icul)ate, thefringedistinct,1-1.5mmbroad, bractsin2-3 rowsonthe sparsely minutelypuberulous; marginandonthefringe,appressed orsomeof themar- 3. Dorstenia hirta Desvaux, Mem. Soc. Linn. Paris oron to 0.5 mmlongappendages, 4: 218. 1826, as var. y; Bureau, in De Candolle, ginalonesradiating ovatetotriangular, to2 mmlong,themarginal Prodr.17: 267. 1873; Carauta,Atas Soc. Biol. Rio onesdistinctlydifferentin length,shorterandlongerones ? de Janeiro16:7. 1972;Carauta,Rodriguesia29(44): 77, t. 9. 1978.Type.Brazil.Withoutlocality,Anonyor the largermarginal clearlyaltemating, puberulous mous s.n. (holotype, P). Fig. 35 bractssetulose;minuteinterfloralbractssometimes flowersintheperiphery present;staminate of theflowDorsteniaerectaVellozo,Fl. Flumin.53. 1829("1825"), Ic. 1: t. 142. 1831 ("1827"); Miquel, in Martius, eringface,sometimesa few amongthepistillateones; Fl. Bras. 4(1): 161. 1853; Bureau, in De Candolle, tepals(2 or)3; stamens(2 or)3; filaments(to3 times) Prodr. 17: 267. 1873; Valente, Carauta & Braga, of thepistillate flower longerthantheperianth; perianth Albertoa4(25): 329, t. 1. 1998. Dorsteniavarrowith3 minutelobesattheapex;stigmas(almost)equal niifoliaFischer& Meyer,IndexSem.Hort.Petrop. in length,0.2-0.8mmlong,white.Endocarp body1.511: 63. 1846, as a synonym of D. erecta Vellozo; 2 x 1.5-1.7mm,distinctlytuberculate. Miquel, in Martius, Fl. Bras. 4(1): 163. 1853. Distribution(seeFig.79.6).IneastemBrazil(from in shadeorhalf-shade; Bahiato RiodeJaneiro); atlow elevations. Representative specimens examined. BRAZIL. Mun. Almadina,rd. Almadina-Ibutipa,km 7, 19-20 Feb 1998, Jardimet al. 1706 (NY); Mun. Jussari, rd. Jussari-Palmeira,ca. km 7.5, 10 Apr 1998, Jardim et al. 1737 (NY); Itamaraju,Campo Alegre, rd. to Piraja, 14 May 1971, Santos1638 (CEPEC,U); Mun.Aurelino BAHIA:

Leal, ca. 11.2 km E of Aurelino Leal, 3 May 1992, Thomas9106 (US). ESPiRiTO SANTO: Mun. Santa Teresa, Valsugana Velha, 18 Feb 1986, BoudetFernandes1849 (GUA, US); Mun. Itagua9u, Jatibocas, Altamira, 15 May 1946, Brade 18242 (NY, RB, U); Mun. Domingos Mar-

Dorstenia erecta Vellozo var. varroni(folia (Fischer

& Meyer) Bureau, in De Candolle, Prodr. 17: 267. 1873. Type. Fl. Flumin. Ic. 1: t. 142. 1831. Dorstenia hispida Hooker, Ic. PI. Ser. 1, 3: t. 220. 1840; Miquel, in Martius, Fl. Bras. 4(1): 163.

1853; Bureau,in De Candolle, Prodr.17: 267. 1873. Dorstenia erecta Vellozo var. hispida (Hooker) Bureau, in De Candolle, Prodr. 17: 267. 1873. Type. Brazil. Rio de Janeiro: Rio Comprido, Sep 1836, Gardner 2116 (or 116?) (holotype, K; isotype, BM). Dorstenia minor Fischer & Meyer, Index Sem. Hort. Petrop. 11: 63. 1846. Dorstenia hispida Hooker forma minor (Fischer & Meyer) Miquel, in Martius, Fl. Bras. 4(1): 163. 1853.

FLORA NEOTROPICA

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Dorstenia erecta Vellozovar.minor (Fischer& Meyer) Herb, to 80 cm tall; rhizome short and stem for the Bureau,in De Candolle, Prodr.17: 267. 1873. greaterpart supraterranean,2-5 mm thick, hirsuteto

Type.Describedfrommaterialcultivatedin Hort. hirtellouswith straightand uncinatehairs, sometimes Petropolis,driedspecimenin herb.Fischer(LE). puberulous;intemodes to 2 cm long. Leaves almost

161

TAXONOMICTREATMENT distichous;laminaoblong to (narrowly)lanceolateto subobovate or to elliptic, (3-)7-25 x (1.5-)2.5-8.5 (-10) cm, (thinly) chartaceous,sometimesvariegated; apex acuminate;base subacuteto obtuse, sometimes roundedor subcordate;marginfaintly to distinctly ? irregularlycrenate-dentateto subentire;uppersurface strigoseto hirsuteto hirtellous,sometimessparselyso, mostly ? scabrous;lower surfacewith minuteuncinate hairs and on the main veins usually with much longer appressedto patent hairs, mostly ? scabrous;lateral veins 7-16 pairs,faintlyloop-connected;tertiaryvenation scalariform(to reticulate);petiole 0.5-2(-3) cm long, hirsute,sometimespuberulous;stipulesnarrowly triangular,to 0.25 cm long, uninervate, usually deflexed, ciliolate. Inflorescences sometimes in pairs, green, the flowering face pointing upwards;peduncle 1.5-7 cm long, sparsely to densely hirsute, also with minute and ? retrorseuncinatehairs;receptaclecentrally attached,discoid, (sub)orbicularto slightly angularin outline,1-3 cm diam.,mostly? lobed,theflowering face plane to ? convex, the outside puberulous, mainly with ? retrorseuncinate hairs, the margin ? faintly and irregularlylobed, the fringe very narrow, sometimes to 3 mm broad;bractsin 2-3 rows on the margin,ovateto ellipticto oblong, 0.5-1 mm long, glabrous or sparsely puberulous,the bracts of the outer row not distinctly different in size; staminateflowers amongthe pistillateones or ? distinctlyconcentratedin the peripheryof the flowering face; tepals 2; stamens 2; filamentslongerthanthe perianth;stigmas(almost) equalin length, 1.5-2 mm long, white.Endocarpbody 2.5-3 x 2-2.5 mm, tuberculate. Distribution (see Fig. 79.7). In eastemBrazil(from Bahiato Parana);in shadeor half shade,oftenon sandy soil; at low elevations. Representative specimens examined. BRAZIL. BAHIA: Ipaiu,rd.to Jequi6,27 Oct 1970,Santos1205 (RB);

Rio Branco, rd. to Prata, 27 Jan 1971, Santos 1437 (CEPEC, U). MINAS GERAIS: Without locality, Langsdorffs.n. (US). ESPIRITO SANTO: Mun.Domingos Martins, Campinho, rd. to Chapeu, 15 Sep 1975, Carauta 1810 (RB); Mun. Itaguagu, Jatibocas, 15 May 1946, Brade et al. 18242 (RB); Mun. Cachoeiro do Itapemirim, Vargem Alta, 5 May 1949, Brade 19756 (RB). PARANA: Guaraque9aba, Serra Negra, 10 Dec 1970, Hatschbach 25780 (MBM). RIO DE JANEIRO: Mun. Nova Igua9u, Serra da Bandeira, 17 Oct 1977, Araujo et al. 1875 (GUA); Mun. Rio de Janeiro, Serra Carioca, 20 Apr 1970, Carauta 1085 (MBM, RB, U); Mun. Petr6polis, Serra da Estrela, 9 Mar 1978, Ferreira et al. 295 (RB); Mun. Maj6, Paraiso, Centro de Primatologia, 19 Oct 1984, Lima et al. 2271 (GUA, RB). SAO PAULO: Caraguatatuba,26 Oct 1974, Carauta 1745 (GUA, MICH); Ubatuba, 5 Nov 1962, Fontella et al. 82 (SP, US), 8 Jan 1985, Gentry et al. 49309 (BG); Sao

21 Jan1955,Joly SP Vincente,Praiade Paranapoan, 14343(B, SP,US);Salesopolis,Boracea,28 Jan1949, Kuhlmannet al. 1736 (SP). Dorstenia hirta is a rathervariablespeciesandin hardlydistinguishable sometimes vegetative characters fromD. turnerifolia.Thesetwospeciesare,however, Thetwo collecverydistinctin thefloralcharacters. Sao Paulo,Carauta 1745 tionsfromCaraguatatuba, sparselyhairyleavesand andLanna 139,havenarrow, resemble D. brevipetiolata.Theseandothercollections withsimilarfeatureshaverecentlybeenplacedin D. erecta erectabyValente etal.(1998).TheyreinstatedD. largelybecauseof anatomicaldifferencesfoundbetweenthisformandthemorecommonandmoredistinctlyhairyformof D. hirta. Specimensat LEfrom cultivatedmaterialare annotatedas D. erecta var. maculata Regel,whichwasprobablynotpublished.

4. DorsteniabrevipetiolataC. C. Berg,Proc.Kon. Ned.Akad.Wetensch.,Ser.C, Biol. Med.Sci. 88: Riode 264,t. 2. 1985.Type.Brazil.RiodeJaneiro: Apr1839,Guillemincat.no. Janeiro,Corcovado, Fig. 36 752(holotype,P). Herb, to 40 cm(ormore?)tall;stem4-8 mmthick, withstraight andminutely anduncipuberulous sparsely natehairs;intemodesto 5 cm long.Leaves in spirals, tendingto distichous;laminanarrowlylanceolateto (tosubcosuboblong,13-19 x 2.5-5 cm,chartaceous basesubobtuse riaceous); apexacute(tosubacuminate); crenate-dentate; margin? irregularly to cord(ul)ate; on the uppersurfacesparselyappressed-puberulous lowersurfacesparsely(onthemain midrib,glabrescent; with? retrorsestraightand veinsdensely)puberulous uncinatehairs;lateralveins 13-20 pairs,loop-convenationoftenpartlyscalariform; petinected;tertiary ole 0.34.5 cmlong,minutelypuberulous, mainlywith to retrorseuncinatehairs;stipulesbroadlytriangular ovate, 1-1.5 mm long, (often broaderthan long), deflexed,sparselyciliolate.Inflorescences: uninervate, withretpeduncle6-8 cmlong,minutelypuberulous disrorseuncinatehairs;receptacle centrallyattached, toellipticinoutline,2.5-4 cm coid,plane,suborbicular theoutsideminutelypuberulous in (largest)diameter, the withretrorseuncinatehairs,themarginsubentire, fringealmostlacking;bractsin 2-3 rowson themargin, ovateto oblong,to 1 mmlong,sparselypuberulousto glabrous;staminateflowersamongthepistillateones;tepals2; stamens2;filamentslongerthanthe stigmas(almost)equalin length,0.6-0.8mm perianth; long.Endocarp body ca. 3 x 2 mm,tuberculate. Distribution (see Fig. 79.5). In eastem Brazil(Rio

de Janeiro),knownonlyfromthetype.

162

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FLORA NEOTROPICA

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Thisspeciesis reminiscent of bothDorsteniaelata 5. Dorstenia contensis Carauta& C. C. Berg, Proc. andD. hirta(cf. Carauta,1978a).It differsfromthe Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med. formerin its veryshortpetioleandstipulesandfrom Sci. 88: 261, t. 1. 1985. Type. Brazil. Bahia: Rio thelatterin its shortindumentum, das Contasvalley, Ipiau,rd. to Itaiba,7 Nov 1970, longerinternodes, shorterpetiole,longerpeduncle,andthegreaternuniSantos 1265 (holotype, CEPEC;isotypes, RB, U). berof lateralveins. Fig. 37

TAXONOMICTREATMENT

163

Brazil.MinasGerais:LagoaSanta,2 Jan-17Feb Herb,to40 cmtall,rhizomatous; stem3-5 mmthick, 1864, Warming1940 (or 1940/3) (holotype,P; varyingfromhirsute(orhirtellous) tohispid(orhispiduisotypes,C, P). lous)withstraightanduncinatehairs,or almostglabrous;internodes to 2 cmlong;latexwhiteto yellowHerb,to 25 cmtall;stempartlyrepentto rhizomaish. Leaves in spirals,tendingto distichous;lamina tous,1-3 mmthick,puberulous, mainlywithretrorse oblongto lanceolate,14-27 x 3.5-9 cm,chartaceous; uncinate hairs;internodes to 1.5cmlong.Leavesinspiapexacuteto subacuminate; baseattenuate; margin? rals,tendingto distichous; laminalanceolate to oblong irregularly andfaintlycrenate-dentate to subentire; up- to subovate,(1-)3.5-10(-13.5)x (0.5-)1-3 cm,mempersurfaceglabrous;lowersurfacepuberulous to hir- branaceous to chartaceous, sometimes variegated; apex tellousorsubglabrous; lateralveins14-20pairs,loop- acute to subacuminate;base acute to obtuse (or connected; tertiary venationoftenpartly? scalarifonn; rounded); marginrepandto ? irregularly andcoarsely petiole1-3 cmlong,varyingfromhirsuteto hirtellous crenate-dentate; uppersurfacesparselyminutely pubertohispidto hispidulous withstraight anduncinate hairs ulous,glabrescent; lowersurfacesparsely, ontheveins oralmostglabrous; stipulesnarrowly triangular tosubu- densely,puberulous, mainlywith? retrorseuncinate late,2-5 mmlong,rigidlycoriaceous(subspinosein hairs;lateralveins7-13 pairs,loop-connected, somedry material),uninervate,soon patentto deflexed, timesfaintlyso;tertiaryvenationreticulateortending hispidulousto almostglabrous.Inflorescenceson to scalariform; petiole0.3-1.5(-2.5)cmlong,puberuminuteshort-shoots withreducedstipules,green;pe- Ious,mainlywithretrorse,uncinatehairsand/orwith duncle2-8 cmlong,puberulous withretrorse uncinate curvedto straighthairs,often (especiallyin young hairs;receptaclecentrallyto somewhateccentrically leaves)partlypointingtowardsthe lamina;stipules discoid(slightlyconcave?),irregularly attached, 0.1-0.3cmlong,patenttodeflexed, angu- narrow-triangular, larto stellatein outline,3-5.5 cm diam.,theoutside sparselyand minutelypuberulous.Inflorescences withretrorse, puberulous uncinatehairs,themargini green;peduncle0.4-1(-2.5)cmlong,slender, minutely withstraightpatenthairsor withretrorse clearlydentate,the fringe1-1.5 mmbroad,sparsely puberulous andminutelypuberulous; bractsin ca. 3 rowson the uncinatehairs;receptacle centrallyattached, discoidto substellateto ? angularto suborbicular marginandon thefringe,(semiorbicular to) ovate,to subturbinate, ca.0.7mmlong,ciliolate;staminate flowersamongthe andthenlobedtoentireinoutline,0.3-1(-1.3)cmdiam., pistillateones;tepals2; stamens2; filamentsaslongas theoutsidepuberulous,mainlywith? retrorseunciorshorter thantheperianth; ribsleadingto stigmasequalinlength,ca. natehairs,sometimeswith? prominent the largerbracts,the fringeverynarrowor to 1 mm 0.2-0.3 mmlong.Endocarpbodynotseen. broad;bractsin 2-3(-4) rows,on themarginandon Distribution(seeFig.81.4).IneasternBrazil(Bathefringe,ovateto oblong,to 0.5 mmlong,glabrous hia);in shadedplaces;atlowelevations. orsparselypuberulous; staminateflowersamongthe Specimensexamined.BRAZIL. BAHIA: BetweenUru- pistillateones;tepals2; stamens2; filamentsshorter gucaand Ubaitaba,22 Apr 1970, Santos 760 (CEPEC); thantheperianth; stigmasequalin length,ca.0.2 mm Gand6,BR.l01, km 3, 8 Jul 1971,Santos 1743 (CEPEC). long.Endocarp bodyca.2 x 2 mm,tuberculate, brown. Thisspeciesis characterized leafbase byitsattenuate Distribution (see Fig. 79.8). In easternBrazil andthelargereceptacle, angulateto stellateinoutline. (MinasGerais,RiodeJaneiro, andSaoPaulo);inmoist It shows affinities to Dorstenia bowmaniana. andshadedplaces;to 900 m.

6. Dorstenia bowmaniana J. G. Baker, in W. W. Saunders,Refug. Bot. 5: t. 303. 1871; Bureau,in De Candolle, Prodr. 17: 268. 1873, as D. bowmanniana; Carautaet al., Rodriguesia29(44): 81, t. 12. 1978. Type. Described from materialcultivated in Hort. Saunders,collected by Bowman in Brazil,nearRio de Janeiro,herbariummaterialcollected in 1870, Anonymouss.n. (holotype,K).

Specimens examined. BRAZIL. RIO DE JANEIRO: Mun. Rio de Janeiro,Mata do ParqueLage, 16 May 1971, Carauta 1377 (GUA);Mun. Resende, Itatiaia, Lote 21, 22-28 Nov 1938, Markgrafet al. 3626 (GUA, RB); Mun.

Rio de Janeiro,Matado Rumo,16 Oct 1969,Sucre6089 (RB); Mun.Petr6polis,ReservaPaty de Alfares, 1 Nov 1970,Sucre7340(GUA).Sio PAULO: Morrodas Pedras, Morrode lguape,Oct 1917, Brade 7874 (RB). Large-leavedspecimensof Dorsteniabowmaniana

ofD. turnerifolia, but resemble small-leaved specimens Dorstenialagoensis Bureau,in De Candolle,Prodr. thetwospeciescanbe easilytoldapartby thearrangeflowersandthenumberof sta17: 268. 1873; Carauta,Rodriguesia29(44): 83, mentof thestaminate thestaminate flowers,mostly t. 14. 1978;Carauta,Valente& Barth,Arq.Jard. mens:in D. turnerifolia, Bot. Rio de Janeiro23: 106, t. 3. 1979. Type. with3 stamens, arefoundintheperiphery oftheflower-

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

165

ing face,andin D. bowmaniana, thestaminateflow- Hatschbach14137 (MBM);Paranagua,Rio Cambara, ers,with2 stamens,areintermixed withthepistillate 28 May 1968, Hatschbach 15254 (MBM, MICH); flowers. Dorstenia bowmaniana also resembles D.

QuartoRamos,Rio do Carvo, 12 Jan 1967,Hatschbach

milaneziana;thelattercanbe distinguished fromthe 15700 (MBM);Guaraque9aba,FazendaAbebreira,15 formerbythe(sub)cordate leafbaseandthelonghairs Nov 1969, Hatschbach22461 (MBM);Morretes,Serra Marumbi,Picadodo Olimpio,19 Jan 1971, Hatschbach on theuppersurfaceof thelamina. 1 7. DorsteniacarautaeC. C. Berg,Proc.Kon.Ned. Akad.Wetensch., Ser.C,Biol.Med.Sci.89: 136,t. 9. 1986.Type.Brazil.SaoPaulo:Eldorado, nr.Gruta da Tapagem,22 Jan 1971, Carauta1277 (holotype,RB;isotypes,AC,COL,E, F,GH,GUA,K, NCU, SING,SP,U, US, UT,ZA). Fig. 38 Herb,toca.50cmtall;stempartlyrepent torhizomatous,1-4 mmthick,(rather) denselypuberulous tohirtellouswithstraight,curved,and/oruncinatehairs;intemodesto 3.5 cm long.Leavesin spirals,tendingto distichous;laminaoblongto elliptic,4-16 x 2-7 cm, chartaceousto membranaceous; apex subacuteto (sub)acuminate; baseacuteto obtuseto rounded; margin ? coarselycrenate-dentate to subentire; uppersurfacescabrous withminuteconicalhairsorglabrous; lower withminutestraight surface touncinate puberulous hairs, mainlyontheveins,orto hirtellous; lateralveins6-1 1 pairs,rather distinctly loop-connected; tertiary venation reticulate topartlyscalariform; petiole(0.7-)1-4cmlong; withminutestraight, oruncinate hairs pubenulous curved, to almost (orto hirtellous); stipulesnarrowly triangular subulate,0.1-0.4 cmlong,coriaceous, uninervate, appressedor? patentto deflexed,sparselyandminutely puberulous. Inflorescences green;peduncle0.7-2 cm long,minutely puberulous; receptacle centrally attached, in discoidto broadlyturbinate, angularto (sub)stellate outline,(0.7-)1-2cmdiam.,theoutsideminutely puberulous,thefringedistinct,ca. 1mmbroad;bractsin 2-4 orsomeandonthefringe,appressed rows,onthemargin ovateto suborbicular, to timesthelargeronesradiating, 0.7mmlong,ciliolate;staminate flowersamongthepisaboutaslongas tillateones;tepals2;stamens 2;filaments theperianth; stigmas(almost) equalinlength,ca.0.1-0.2 mm long. Endocarpbodyca. 2 x 1.5 mm, tuberculate.

Distribution(seeFig.79.12).IneastemBrazil(Sao in shadedplaces;atlow elPauloto SantaCatarina); evations. Specimens examined. BRAZIL. PARANA:Morretes, 23 Jan 1914, Dusen 14383 (F, K, MICH, NY, P); P6rto de Cima, 12 Dec 1914, Dusen 14617 (MO); Morretes, Estacao Marumbi, Rio Taquaral, 12 Dec 1948, Hatschbach 1147 (MBM, US); Guaratuba,Rio Boguaqu, 23 Oct 1957, Hatschbach5185 (MBM); Guaratuba, Pedra de Araraquara, 9 Mar 1961, Hatschbach7887 (HB, HBR, MBM); Morretes, 16 Dec 1964, Hatschbach 12038 (MBM); Adrian6polis, Barra Rio Pardo, 5 Apr 1976,

26006 (MBM);Morretes,Rio dos Padres, Dec 1972, Hatschbach30893 (MBM);Rio Iriranga,2 Mar 1967, Hatschbachs.n. (MBM); CerroAzul, BarraRio Bom Sucessa, 24 Jan 1974, Hatschbach 33745 (MBM); Paranagua,14 Dec 1948, Tessmanns.n. (MBM, MO). SANTA CATARINA: Itajai,Morroda Fazenda,Carautaet al. 4464 (GUA);Ilhade SantaCatarina,MorroCostada Lagoa, 21 Dec 1966, Klein 6985 (Herb. Fac. Farm, Parana);Arar,Terao,Reitz 139 (RB); Itajai,Morroda Fazenda,4 Mar 1954,Reitzet al. 1720 (HBR,US); Sao Franciscodo Sul, Garuva,Tres Barras, 19 Dec 1957, Reitzet al. 5759 (HBR,K);Pil6es,Palhoca,17 Feb 1957, Reitzet al. 5922 (HBR,MBM,K); Rio do Sul, Serrado Matador, 17 Oct 1958, Reitz et al. 7327 (HBR, K); Florian6polis, MorroItacoburi,12 Mar1952,L. B. Smith et al. 6152 (US);Piloes,Palhoca,4 Apr 1957,L. B. Smith et al. 12355 (US). SAOPAULO: Mun.Cananeia,Ilhado Cardoso,Maruja,24 Oct 1989, Cordeiroet al. 613 (SP); Mun. Cananeia,Ilha do Cardoso,Rio Pedro, 17 May 1988, Sugiyamaet al. 746 (SP).

Thematerialreferredto Dorsteniacarautaehasbeen included in D. lagoensis (Carautaet al., 1978). In the presentrevisionD. lagoensis is regardedas a synonym of D. bowmanianawith a scattereddistributionin the States of Minas Geraisand Rio de Janeiro.Dorstenia bowmanianadiffers from D. carautae in the smaller leaves and inflorescences and in featuresof the indumentum, such as the length of the hairs and the frequency of uncinatehairs. Dorstenia carautae andD. bowmanianaappearto be closelyrelated,butsufficiently distinctto justify recognitionat the species level.

8. Dorstenia milaneziana Carauta,Valente& Sucre, Bradea 2(48): 473, cum. tab. 1975; Carauta, Rodriguesia29(44): 86, t. 18, 1978. Type. Brazil. Espirito Santo: Cachoeiro do Itapemirim,25 Apr 1972, Sucre8961 (holotype,RB; isotypes, F, GUA, HB, MICH, R, SP). Fig. 39 Dorsteniagracilis Carauta,Valente& Araujo,Bull. TorreyBot. Club 103: 172, t. 2. 1976; Carauta, Rodriguesia29(44): 87, t. 19. 1978. Type. Brazil. EspiritoSanto: Linhares,Reserva Florestal de Linhares, 14 Jun 1969, Sucre 8628 (holotype, RB; isotype, F). Herb, to 20 cm tall; stem partlyrepentto rhizomatous, slender, 1-2.5 mm thick, hirtellouswith straight anduncinatehairs;internodesto 1(- 1.5) cm long.Leaves in spirals;laminaoblongto lanceolateorto elliptic,often

FLORA NEOTROPICA

166

'.

..

A

.

.17k

Fig. 38. Dorstenia carautae. 1. Habit(Lofgren sn.). 2. Leafy partof stem with inflorescence(Dus, [By E. M. Hupkens van der Elst, Utrecht.]

14383).

167

TAXONOMIC TREATMENT

Fig. 39.

Dorstenia milaneziana.

1. Habit.

2. Leaf

(Santos

subovate or subobovate, 1-10(-1 2) x 0.5-4(-5) cm, chartaceousto membranaceous,often variegated;apex acuteto subobtuseto subacuminateor to rounded;base (sub)cordate;marginrepandto ? coarselycrenate-dentate;uppersurfacehirtellousto strigoseto hispidulous, often distinctlypunctate;lower surfacepuberulous,on the main veins to hirtellouswith straightanduncinate hairs, or partly hispidulous; lateral veins 3-10 pairs, loop-connected (sometimes faintly so); tertiaryvenation reticulate,sometimes partlysubscalariform;petiole 0.5-1(-2.5) cm long, hirtellous with straightand

1255).

[By

van"': der Elst, Utrecht.]~~~~~~~~~~~~~~~~~~~~~~

,.-'' E. M. Hupkens

stipulesnarrowly uncinatehairs,orpartlyhispidulous; unitriangular to subulate, 0.2-0.5cmlong,coriaceous, nervate,patentto + deflexed,sparselypuberulousor glabrous. Inflorescences green; peduncle 0.7-1.5

with to (brownish-)hirtellous (-2) cmlong,puberulous centrallyattached, patentor? retrorse hairs;receptacle to + angular to discoid,suborbicular broadlyturbinate in outline,0.3-1(-1.3) cm diam.,theoutsidepuberulousto hirtellouswith? retrorsehairs,themarginsubthefringedistinct, entiretodentateto subappendiculate, bractsin ca.4 to 1.5mmbroad,minutelypuberulous;

168 rows, most of them on the marginand on the fringe, a few below the margin,most of themnarrowlyovate, to 0.5 mm long andciliolate, a few marginalandsubmarginal ones distinctly larger,to 1.3 mm long and often setulose(orthemarginalteethsetulose);staminateflowers in the periphery of the flowering face, or a few among the pistillate flowers; tepals 2; stamens2; filaments almost as long as the perianth;stigmas slightly unequalin length,ca. 0.3 mm long. Endocarpbodyca. 2 x 2 mm, distinctly tuberculate,pale brown. Distribution (see Fig. 79.12). In eastern Brazil (BahiaandEspiritoSanto);in moist andshadedplaces; at low elevations. Specimensexamined. BRAZIL. BAHIA:Sao JoseBuerarema,rd. to Una, km 19, 1 Apr 1980, Berg et al. 1144 (CEPEC,U); SantaCruzde Cabralia,7 Apr 1972, Pineiro 1812 (CEPEC);Anapolis, Itabela,Guaratinga, 4 Jul 1970, Santos 892 (CEPEC,RB, U); Itiau, rd. to Jequie,2 Nov 1970, Santos 1255 (CEPEC,U); rd. Rio Branco-Una,km 40, FazendaDendhavea,4 Sep 1974, Santos2768 (CEPEC);Mun.Ilheus,2 kmNNEof Banco da Vitoria,28 Sep 1994,Thomaset al. 10708(NY);Mun. Ilh6us, 3 km N of Rodovaria,Mata da Esperan9a,17 SANTO: Mar1996, Thomaset al. 11078 (NY). ESPilUTO Mun. Linhares,Goitacazes, Rio Doce, 19 Nov 1943, Kuhlmann6490 (K, RB, U); Mun. Linhares,Reserva Florestal,Cia. Vale do D6ce, 29 Sep 1978, Martinelli 5000 (RB); Mun. Aracruz,21 Feb 1994, Pirani et al. 3068 (SPF,NY); SantaTeresa,valley of Rio Canaa, 1 Feb 1969, Sucre et al. 4566 (RB). The species is rathervariablein the dimensions of the leaves and the inflorescences and approaches Dorstenia setosa.

FLORANEOTROPICA a reddishmargin;peduncle1.5-4 cm long, sparselyhirtellousto hispidulousto hirsute;receptaclecentrallyattached,discoid,almostplane,orbicularin outline,sometimesslightly lobate,(l-) 1.5-2.5 cm diam.,the outside puberulouswithretrorseuncinatehairs,partlyhispidto hispidulous,especially on the appendages,the margin dentateby to 1.5 mm long appendages,the fringeca. 1 (-2) mm broad,minutelypuberulous;bractsalmostin a singlerowon themarginalappendages,appressed,ovate, to ca. 0.7 mm long, minutelypuberulousor the larger ones setuloseto setose, sometimesalso with ca. 4 rows of minutebractson the fringe;staminateflowersamong thepistillateones;tepals2; stamens2; filamentsas long as or shorterthantheperianth;stigmas(almost)equalin length, 0.4-0.5 mm long. Endocarp body ca. 1.5 x 1 mm,tuberculate. Distribution (see Fig. 79.9). In eastem Brazil(Bahia); in shadedplaces; at low elevations. Specimensexamined.BRAZIL.BAHIA: Mun.Buerarema,rd. Buerarema-Vila Brasil,km 14, 9 Feb 1982, Carvalhoet al. 1159 (GUA);Mun. Itacare,rd. BR101, 18 km fromItacare,FazendaDas Almas, 13 Feb 1994, Kallunkiet al. 532 (SPF,NY); Almadina,Matada Serra Pancadinho, 19 Mar 1971, Raimundo S.P. 1093 (CEPEC,RB, U); Sao Domingo,Feb 1822, Riedel 759 (LE);Gongoi,Barragemdo Funil, 16 Nov 1971,Santos 2154 (CEPEC,GUA);Rio Branco,rd. to Pratas,27 Jan 1971, Santos 1439 (CEPEC,GUA, U). Dorstenia setosa is a distinctive species, differing fromits relatives,like D. milanezianaandotherspecies withhirsuteto setoseindumentumandnarrowandsmall stipules,mainlyin the greaterdimensionsof mostparts.

10. Dorstenia choconiana S. Watson, Proc. Amer. Acad. 22: 477. 1887: Burger, Fl. Costaricensis, Fieldiana Bot. 40: 137, t. 17. 1977. Type. Guatemala. Alta Verapaz:"Black River,"branchof Rio Chocon,20 Mar 1885, Watson296 (holotype,GH). Fig. 41 Dorstenia choconiana S. Watson var. integrifolia Donnell Smith, Bot. Gaz. (Crawfordsville)13: Herb, to 50 cm tall; stem 2-5 mm thick, hirsuteto 76. 1888. Type. Costa Rica. Limon: Llanura(s) setose;intemodesto 2 cm long. Leavesin spirals,tendde SantaClara,La Concepci6n,Feb 1896,Donnel ingto distichous;laminaellipticto oblong,(to lanceolate), Smith6777 (holotype,GH; isotypes, BM, F, K). (6.5-)9-21 x (3-)4-8.5 cm, thinly chartaceous;apex Dorsteniacordato-acuminata Cufodontis,Arch.Bot. Sist. Fitogeogr. & Genet. 10: 27. 1934. Type. (sub)acuminate;base cordate; margin ? irregularly Guatemala.BajaVerapaz:nr. Purulha,Apr 1907, crenate-dentateto coarsely crenateto sublobateor to Tuerckheim (II-)1709(lectotype,designatedhere, subentire;uppersurfacestrigose;lowersurfacesparsely G; isolectotypes,F, G, GH, NY, US, Z). hirsute(to setose) on the main veins; lateralveins (9-) 13-16 pairs,loop-connected; tertiaryvenationoftenpartly Herb,to ca. 50 cm tall;stem2-5 mm thick,sparsely scalariform;petiole (1.5-)4-14 cm long, hirsute(to se- to ratherdensely puberulousto hirtellouswith straight tose); stipules subulate,0.3-1.1 cm long, uninervate, and uncinate, often ? retrorse hairs or subglabrous; often ? patent,hispidulousto hirsute.Inflorescenceson intemodes 1-5 cm long. Leaves in spirals;laminaobminuteshort-shootswithreducedstipules,greenor with long to elliptic, to subovateor subobovate,sometimes 9. Dorstenia setosa Moricand,P1.Nouv. Amer. 103, t. 64. 1841; Miquel, in Martius,Fl. Bras.4(1): 170. 1853;Bureau,in De Candolle,Prodr.17:269. 1873; Carauta,Rodriguesia29(44): 76, t. 8. 1978. Type. Brazil. Bahia: Withoutlocality,Blanchets.n. (holotype, G). Fig. 40

TAXONOMICTREATMENT

169

Fig. 40. Dorsteniasetosa. Leafy partof stem with inflorescences(Santos 1439). [By E. M. Hupkensvan der Elst, Utrecht.]

170

FLORA NEOTROPICA

Fig. 41. Dorstenia choconiana. 1. Habit (Tuerckheim111 709). 2. Leaf (Lent 691). (By E. M. Hupkens van der Elst, Utrecht.]

TAXONOMICTREATMENT

171

4558 (F, MICH);nr. La Ceiba, 16 Jul to lanceolate,entireorpinnatilobed to -parted,5-26 x Jul 1934, Yuncker 2.5-8.5 cm,orif thelaminapinnatifid to -parted,then 1938, Yuncker et al. 8493 (F, GH, MICH, MO, NY). to 17.5cmbroad,chartaceous; apexsubacute to acumi- YORO:Cord.Nombrede Dios, hills S of San Jose de nate;base? distinctlyattenuate orroundedto subcor- Texiguat, 17 May 1991, Davidse et al. 34481 (MO). NICARAGUA. Rio SANJUAN:La Palma,between Rio dateanddecurrentalongthe petiolewith ? narrow Santa Cruz and Cafio Santa Crucita,21 Mar 1985, Moreno wings;marginsubentire torepandto lobate;uppersur- 25505 (MO),30 Nov-2 Dec 1984,Stevens23412 (MO). face(sub)glabrous, smooth;lowersurfacepuberulous ZELAYA:El Escobillo, rd. Colonia Serrano-Yolaina, km. tohirtellousorsubstrigillose with? retrorse straight to 2, 29 Jul 1982, Sandino 3319 (MO);Cailo Costa Riquita, uncinate hairs,moredenselyonthemainveins,smooth ca. 1.8 km SW of Colonia Naciones Unidas, 6-7 Nov to scabridulous;5-10(-12) pairs of lateralveins, 1977, Stevens 4974 (BG, MO). Reserva UCR, Fila Volcan (faintly)loop-connected; tertiary venationscalariform COSTA RICA. ALAJUELA: to reticulate; petiole1.5-5 cmlong,sparselyto rather Muerte, between headwaters of Rio San Lorenzo and denselypuberulous tohirtellouslikethestem;stipules Bajo Jamaical, 1000-1200 m, 15-17 Apr 1982, Barringer (narrowly) triangular to 1.2cmlong,to 3.5mmbroad, et al. 2605 (F); Cataratas (Los Angeles) de San Ram6n, 18 Feb 1935, Brenes 18920 (CR), 17-19 Apr 1935, Brenes puberulous toglabrous. Inflorescences onminuteshort- 20526 (F, NY); Reserva Forestal San Ramon, Rio Lorenshootswithreducedstipules,greenor themarginof cito, 19-20 Feb 1985, Gomez-Laurito 10256 (F); Upala, thereceptaclepurple-brown; peduncle1-5 cm long, Dos Rios, 8 Apr 1988, Herrera Ch. 1741 (BG); Alfaro graduallypassingintothe? broadlyturbinate recep- Ruiz, La Pena de Zarcero, 30 Jul 1938, A. Smith 1004 (F, tacle;receptaclecentrallyattached,(sub)orbicular in GH, NY); Cant6n San Carlos, Pueblo Nuevo, 1100 m, 15 outline,(0.5-)1-3.5 cm diam.,the outsidewithminute

Apr 1939, A. Smith 1917 (F). CARTAGO: Nr. Turrialba,12

conicalandlongerretrorseuncinatehairs,themargin Jul 1965,Jimenez M 3280 (CR), Feb 1966, Kern 822 (F, entire,thefringe0.5-3.5mmbroad,withsparseminute NY), 1 Aug 1965, Lent 691 (F, MICH, NY); Quebrada swollenhairs;bractsin ca. 3 rows,on andbelowthe Rojas, nr. Muneco, 1240 m, 15 Sep 1969, Lent 1791 (BM, F, U); Rio Gato, 15 Sep 1973, Lent 3632 (CR, marginandon the fringe,(broadly)ovateto semi- Pejivalle, Jan 1940, Skutch 4582 (CR, F, GH); (Mt.) F); Talato 1.5mmlong,appressed, orbicular, sparselypuberu- manca, hills of Sikurbete, Apr 1895, Tonduz9388 (BR), lous;staminate flowersamongthepistillateones;tepals Feb 1895, Tonduz9389 (BR, CR); (Mt.) Talamanca,forest 2; stamens2; filamentslongerthantheperianth; stig- of Tsaki, Apr 1895,Tonduz9524 (BR, CR, F, Z). GUANAmasequalorunequal inlength,0.3-0.8mmlong.Endo- CASTE:ParqueNacional SantaRosa, 24 Jul 1983, Barringer carp body ca. 2.5 x 2 mm, slightly tuberculate.

Distribution(see Fig.79.13).FromGuatemala to inmoistforest,oftenalongstreams; Panama; to 1800m. Representative specimensexamined.GUATEMALA. ALTAVERAPAz: Between SebolandCoban, 25May1964,

Contreras4802 (F, NY); betweenSepacuiteand Secanquin, 1100m, 18 May 1905,Pittier325 (US)andPittier 326 (US);mountainsE of Tactic,rd. to Tomalhu', 15001600 m, 9 Apr 1939,Standley71411 (F);betweenSanta CruzandTactic, 1375 m, 14 Apr 1941, Standley92292 (F);mtns.E of Tactic,1500-1650m, 9 Apr 1939, Wilbur 14930 (MICH);Cobain,Rio Chiui,1350 m, Feb 1908, Tuerckheim 2133 (US). BAJA VERAPAZ: Pansamala,ca. 1200 m, Jun 1885, Tuerckheim 751 (B, GH, K, MICH, NY, P, US). IZABAL: Escobas,W of PuertoBarrios,3 May 1939, Standley 72959 (F, NY) and 73022 (F); Montafladel Mico, betweenDarmouthand Morales,7 Apr 1940, Steyermark39057 (F, US); Rio Juyuma, 22 km SW of Banarera,8 Apr 1940, Steyermark39122 and 39123 (F); Rio Bonita, 21 Dec 1941, Steyermark 41663 (F). HONDURAS. ATLANTIDA: Mt. Nombre de Dios,

between Saladito and San Francisco, 26 Apr 1967, MolinaR. 20847 (F,NY); nr.SanAleje,22-27 Apr1947, Standley7595 (F); nr. Tela, 6 Dec 1927-20 Mar 1928, Standley52647, 53114, and 55601 (F, US), 30 Jul 1962, Websteret al. 12701 (F, MO, U); Lancetillavalley, 12

et al. 4006A (F);ParqueRincon de La Vieja, Liberia, 9501100 m, 28 Nov 1987, Herrera Ch. 1448 (BG); Tileran, Jan 1943, Quiros C. s.n. (CR). HEREDIA:Rio Sarapiqui, Colonia Virgen del Socorro, 3 Feb 1984, G6mez-Laurito 9861 (F). LIM6N:Between Bri Bri on Rio Sixaola and the coastal plain, nr. Rio Catarata,10-13 Feb 1977, Burger et al. 10483 (CR, F); rd. Cerro Chimun-RioSegundo, 1200 m, 1 May 1985, G6mezet al. 23600 (BG); between Cerros Jacr6n and Bitarkara,25-29 Mar 1983, Hazlett 5147 (F); nr. Guapiles, 9 Jul 1949, Holm et al. 346 (P). PUNTARENAS:Palmar Norte, 27 Feb 1951, Allen 5973 (GH);5 km W of Rinc6n de Osa, 24-30 Mar 1973, Burger et al. 8849 (CR, F); Monteverde, 1350-1400 m, 13 Oct 1984, Haber 668 (BG); Montes de Oro, 1000 m, May 1961, Jimenez L. s.n. (CR); San Vito de Java, 22 Oct 1964, Jimenez M 2453 (CR, F); ca. 4 km SW of Rinc6n de Osa, Kennedy 1630 (MO); Rincon de Osa, nr. airfield, 6-7 Feb 1974, Liesner 1800 (MO, U); Burica, 16 Apr 1978, Ocampo S. s.n. (CR); nr. Golfito, Mar 1896, Pittier 9898 (BR, CR). SANJose: Between Canaan and Chimirol, Rio Chirripo del Pacific6, 1000 m, 30 Dec 1969, Burger et al. 7131 (CR); El General, 1050 m, Jan 1936, Skutch 2526 (MICH, NY, S), 1100 m, Dec 1936, 3006 (NY, S). PANAMA.CHIRIQui:Rd. Questa Piedra-Cordillera, QuebradaGrande, 3 Jun 1980, Folsom et al. 8042 (MO).

Specimenswith entireand with pinnately lobed to partedleaves can be found throughoutthe rangeof the species, even in the same population.

172

FLORANEOTROPICA

TwocollectionsfromPanama (Cocle:ElCope,Car- Brecha93, 12 Jul 1980, Wendtetal.2527(CHAPA, MEXU); ibbeansideof divide,4 Feb1983,Hamiltonetal. 2753 Mun. Minatitlan,hills S of Poblado 11, ca. 27 km E of La (BG), and Col6n: 9 km W of Llano Grande,S of Laguna,3 Jun 1981, Wendtet al. 3395 (CHAPA,MEXU). Cascajal,11Oct1978,Hammeletal. 5058(MO,PMA, This species shows strongsimilaritiesto bothDorsimilartoDorstenia U))areinmostfeatures choconiana, stenia choconiana andD. umbricola. butthereceptacleis discoidandi irregularly lobed,as in D.panamensis.Itis notclearwhetherthesecollectionsrepresent a distincttaxonorcanberegardedasa 12. Dorstenia umbricola A. C. Smith, Bull. Torrey formof D. choconianawithunusualinflorescences. Bot. Club 58: 87. 1931; Carauta,Rodriguesia Thefactthatthesecollectionsoccur(far)outsidethe 29(44): 84, t. 15. 1978. Type.Peru.Loreto:Rio mainrangeof individualswithnormalinflorescences Itaya,Soledad,20-22 Sep 1929,Killip et al. 29639 adistinct taxon. thattheyrepresent couldbeanindication (holotype, US; isotypes, F, NY). Fig. 43 stem2.5-4 mm Herb,toca.30cmtall,rhizomatous; to hirtelthick,sparselyto ratherdenselypuberulous louswithretrorsestraightanduncinatehairs;intemodes 1-2 cmlong.Leavesin spirals;laminaoblong(to lanceolate orto elliptic),3-12 x 1.5-5 cm,membranaceousto chartaceous; apexacute,sometimesalmost rounded; basesubacute to subobtuse; marginsubentire torepandor? coarselycrenate;uppersurfacescabridontheveinspubercoralloidrhi- ulous;lowersurfacescabridulous, Herb,to 40 cmtall,froma horizontal with uncinatehairs;latulous to hirtellous and straight to hispidulous zome;stem1.5-4mmthick,puberulous eral veins 8-12 pairs, (faintly) loop-connected; tertiary withretrorse to patent,partly tohirtellous to subhispid, withuncinate,hairs;intemodes0.8-2 cmlong.Leaves venation reticulate; petiole 0.5-1.5(-2.5) cm long, in spirals;laminaoblongorellipticto obovate,6-14.5 puberulousto hirtellous with retrorse,predominantly x 2-5 cm, chartaceous; apexacuteto subacuminate; uncinatehairs;stipulestriangular,often swollen at the to faintlycrenate; base,orto subulate,to 0.25 cm long, uninervate,puberubaseacutetoobtuse;marginsubentire lowersurface lous. Inflorescences on minute short-shoots with reuppersurfacescabridulous, subglabrous; onmidrib; lateral veins7- ducedstipules,green;peduncle1.5-3.5 cm long, gradusparsely minutely pubemlous to sparselypuberulous venationreticulate; allypassingintothereceptacle, 12,faintlyloop-connected; tertiary predominantly uncinatehairs; petiole 0.7-2.5 cm long, ca. 1 mm thick,minutely hirtellouswithretrorse, (almost)centrally attached, broadlyturbinate, to lanceolate,0.1-0.15 receptacle puberulous; stipulestriangular in cm 0.8-1.7 (sub)orbicular outline, diam.,theoutcmlong,uninervate, oroften,fromthefirst appressed oronlylater,patentorreflexed,puberulous. Inflores- side (rather)sparsely appressed-puberulousto strigilwithuncinatehairs,the margin cencesonminuteshort-shoots, greento yellowish,the lose, predominantly floweringfacepurple;peduncle0.5-1.7cmlong,pass- entire,thefringeca. 1 mmbroad,rathersparselyand intothereceptacle, inggradually minutelypuberulous; minutely puberulous;bracts in ca. 3 rows on and be? eccentrically ? low themarginandon thefringe,appressed, broadly receptacle attached, broadlyturbinate, angularin outline,ca. 0.2-1 cmdiameter,theoutside ovateto semiorbicular,to 1mm long, sparselyminutely themarginlobed,lobesnarrow, puberulous;staminateflowersmainlyperipheral,a few minutelypuberulous, ? alternating longer(ca.1mm)andshorter (ca.0.5mm) amongthepistillateflowers;tepals2; stamens2; filalobes,thefringeverynarrow;bractsin 2 rows,a row ments shorterthan the perianth;stigmas equal or unonthereceptacle lobesanda rowinthehollowsbetween equal in length, ca. 0.5 mm long, often purplish.Enlobes,submarginal, ovatetodeltoid,ca.0.5-1 mmlong, docarp body ca. 2.5 x 2 mm, tuberculate. flowersamongpistilminutelypuberulous; staminate Distribution(seeFig.79.14).InPeru(Loreto)and lateones;tepals2; stamens2; filamentsaboutaslong Ecuador (Morona-Santiago, Napo, andPastaza);in rain astheperianth; stigmasnotseen.Endocarp bodyca.2.5 forestundergrowth; atlow elevations. x 2 mm,slightlytuberculate. 11. DorsteniauxpanapanaC. C. Berg& T.Wendt, Proc.Kon.Ned. Akad.Wetensch.,Ser.C, Biol. Med.Sci. 89: 129. 1986.Type.Mexico.Veracruz: Mun.Minatitlan,14.7kmE of LaLaguna,8.2 km et al. 4879 (holoN of Brecha,6 Jul 1985, Wendt type, MEXU;isotypes,CAS, CHAPA,MO,NY, TEX,U). Fig. 42

Distribution (see Fig. 79.13). In Mexico onrockoutcropsandsteepslopesinhigh(Veracruz); rainforest;atlowelevations. precipitation Specimensexamined.MEXICO.VERACRUZ: Mun. Minatitlan, 14.7 km E of La Laguna then 8.2 km N on

Specimens

examined.

ECUADOR.

MORONA-

SANTIAGO:Nr. San Luis, ca. 10 km S of Macas, 14 Nov

1995, Stahl et al. 1903 (AAU). NAPO:Aiiangu, Parque Nacional Yasuni, 1-30 Oct 1983, Korning et al. 47180 (AAU); Rio Napo, Afiangu, 16-27 Apr 1985, Lawesson et al. 39466 (AAU); Reserva Floristica El Chuncho, 5

TAXONOMIC TREATMENT

Fig 42 Dorstenia uxpanapana Habit (Wendt et al. 339)

173

[By E M Hupkens van der Elst Utrecht.

FLORA NEOTROPICA

174

C,',

Fig43

;,

Dorstena umbcola Habit (Pa

) [y

H

n

Br

Bg

f 'S ,S~~~~~~~~~~~~~~~~~~~~~ 0 fXr~ ~~~~~~~~~r

f:::

........

s 2

,

00-0 010~~~~~~~~~~~~~~~~~~~~~~~~~~~ iD0 FE aywL?ULf-Su-

...............

-

S .

.......


S .*.

;

i

-k0'tr

0;90

*-

;0S100'S'

Fig. 43. Dorstenia umbricola. Habit (Palacios 2012). [By Hendrieke Berg, Bergen.]

*;<J~~~~~~~~~~~~

TAXONOMICTREATMENT km NW of Coca, 2 Oct 1987, Palacios 2012 (QCNE). PASTAZA:Canton Pastaza, nr. Villano, 24 Jul 1992,

Palacios 10285 (BG, QCNE). PERU. LORETO:Prov. Maynas, Rio Amazonas, 2 km W of Indiana, 15 Feb 1987, Gentry et al. 55967 (MO); Prov. Maynas, Rio Amazonas, nr. Indiana, 14 Feb 1989, Gentry et al. 65779 (BG, MO, USM); Quebrada Yanamono, 4 Nov 1994, Maas et al. 8316 (U).

The inflorescencesof Dorsteniaumbricolaarevery similarto those of D. choconiana.The relativelybroad

175 staminateflowers mainly peripheralor also numerous among the pistillate flowers; tepals 2; stamens2; filamentslongerthantheperianth;stigmasalmostequalor unequal in length, 0.4-0.5 mm long. Endocarp body ca. 3 x 2-2.5 mm, tuberculate. Distribution (see Fig. 79.14). In Peru(Cuzco,Huanuco, Madrede Dios, andPuno) andBolivia (La Paz); in moist forest;to 1400 m. Specimens examined. PERU. Cuzco: Prov.Quis-

12949 (toelliptic)laminaresemblesthatof D.peruviana,but pichanis, nr. Quince Mil, 18 Feb 1991, Nuniez

stem3-5 mmthick, Herb,to 80cmtall,rhizomatous; sparsely,? retrorselyand minutelypuberulousto denselyhirtellousto subhirsutewith uncinateand straighthairs;intemodes1-4.5 cm long. Leaves in

Prov. Pachitea,Codo de Pozuzo, 22 (BG). HUANUCO: Oct 1982,Foster9381 (BG,F, MO);Prov.LeoncioPrado, Dtto. HerilioValdizan,old roadto La Divisoria,16 Apr DEDios: Prov.Tambo1976,PlowmanS904(U). MADRE NativoInfiemo,betweenChicaHermosa pata,Comunidad and Lago Cocococha, 21 Nov 1988, Diaz et al. 3148 (MO);Prov.Manui,Atalaya,nr. Hda.Amazonia,12 Dec 1983, Foster et al. 7443 (BG, MO); ParqueNacional del Manui,Rio Manu',CochaCashuStation,6 Nov 1986, Fosteret al. 12176 (BG); ParqueNacional Manui,Rio Manu,PakitsaStation,18 Dec 1988, Fosteret al. 12621 (F);ParqueNacionaldel Manu,Rio Manui,CochaCashu Station, 29 Jul 1986, Nuinez 5543 (BG). PUNO:Rio Tavara,17 May 1992 (st), Gentryet al. 76763 (MO). BOLIVIA. LA PAZ:Prov.Abel Iturralde,Campamento Alto Madidi, 29 May 1990, Beck et al. 18325 (BG, LPB); Prov. Sud Yungas,Alto Beni, Concesi6n Sapecho, 23 Oct 1990, Seidel et al. 4391 (BG, LPB); Prov.SudYungas,Alto Beni, ColoniaTarapaca,27 Oct

spirals; lamina oblong to subobovate to elliptic to

1993, Seidel et al. 7401 (BG, LPB).

it has fewer lateralveins, 8-12 vs. 10-18 pairs.

13.DorsteniaperuvianaC.C.Berg,Proc.Kon.Ned. Akad.Wetensch.,Ser.C,Biol. Med.Sci. 85:35, t. 3. 1982. Type. Peru. Huanuco:trail from Rio 15Oct 1936, Huallagato RiachuelaChontalagua, Mexia 8303(holotype,U;isotypes,BM,F,G, GH, Fig. 44 K, MO,NA, NY, S, US). Dorstenia boliviana C. C. Berg, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med. Sci. 85: 33, t. 2. 1982. Type. Bolivia. La Paz: Charopampa, nr. Mapiri, Nov 1907, Buchtien 1948 (holotype, US).

subpandurateor to lanceolate,6-22 x 2-9.5 cm, The species shows similarities to Dorstenia basesubcorto acuminate; chartaceous; apexsubacute choconianafromwhichit mainlydiffersin the absence dateto cordateor sometimesalmostobtuse,often? of an attenuatebase of the lamina and in the gradual torepandor(coarsely) broadeningof the peduncle into the receptacle. The marginsubentire inequilateral; crenate;uppersurfacescabridulousor smoothand species appearsto be relatedto D. umbricola. on theveins lowersurfacescabridulous, subglabrous; The collections from Huanuco differ from those sparselyandminutelypuberulouswith straightand fromMadrede Dios andBolivia in the sparseandshort uncinatehairs;lateralveins 10-18 pairs,loop-con- indumentumon variousplantparts.Thisdifferencewas to reticulate; usedto nected;tertiaryvenationsubscalariform distinguishDorsteniaperuvianafromD. bolivitohirtellous ana petiole2-5 cmlong,minutelypuberulous (Berg & van Leeuwen, 1982). Due to more recent uncinateto straighthairs;stipules(nar- collecting,the differencesbetweenthe northemandthe with? retrorse to subulate, to0.8cmlong,uninervate, more distinctly hairy southem form became obscure rowly)triangular onminuteshort- andmade uniting of these two entities necessary. Inflorescences puberulous. (minutely) shootswithreducedstipules,yellowishgreenwitha darkerrim;peduncle1-7 cmlong,sparselyto densely to hirtellous,mainlywithuncinatehairs; 14. DorsteniaaristeguietaeCuatrecasas, puberulous Bol. Soc. to centrally attached, gradually eccentrically receptacle Venez. Cienc. Nat. 15: 108. 1954: Carauta, to passingintothepeduncleornot,broadlyturbinate 29(44):85, t. 16. 1978.Type.VenezuRodriguesia to ellipticinoutline,ca. 1.5almostdiscoid,orbicular ela.Aragua:ParqueNacionalHenriPittier,Rancho 5 cmdiam.,theoutsideminutelyappressed-puberulous Grande,Aug 1953, Aristeguieta 1980 (holotype, to hirtellous,themargindenticulateto subentire,the Fig. 45 F-n.v.;isotypes,US,VEN). fringeca. 0.5 mmbroad;bractsin ca. 3 rowson and to ca.50cmtall;stem2-7 mmthick, belowthemarginandonthefringe,ovatetotriangular Herb,probably orsome- hirsuteto hirtellouswith(almost)straightoruncinate to 0.7mmlong,appressed orto semiorbicular, times ? radiating,sparselyappressed-puberulous;hairs;internodes(0.2-)0.5-2 cm long.Leavesin spi-

176

FLORA NEOTROPICA

Fig. 44. Dorstenia peruviana. 1. Habit. 2. Leafy stem with inflorescences (Mexia 8303). [By E. M. Hupkens van der Elst, Utrecht.]

rals;laminaoblong to lanceolateor subobovate,5-15 x 1.2-7.5 cm, membranaceousto chartaceous;apex subacuminateto acute;base (narrowly)truncateto cordate;margincoarselycrenate(to subentire);uppersurface scabrouswith minuteconical hairs;lower surface scabrous,on the veins puberulousto hirtellous to hirsute;venationplaneto slightlyimpressedabove,mostly prominent(and conspicuous)beneath;lateralveins 814 pairs,loop-connected;tertiaryvenationsubscalariform to reticulate;petiole 1-4 cm long, puberulousto hirtellous or subhirsute;stipules to 0.8 cm long, (narrowly) triangularto subulate,uninervate,puberulous.

Inflorescences on minute short-shoots with reduced

stipules, green;peduncle 2.5-6 cm long, hirtellous(or puberulous),mainlywith straighthairs;receptaclecentrallyto somewhateccentricallyattached,discoid, suborbicularto ? irregularlylobed in outline, ca. 1-2 cm diam.,theoutsidepuberulous,mainlywithuncinatehairs to hirtellouswith straightand uncinatehairs,the margin : lobed,the fringeca. 0.5-1 mm broadwith minute conical hairs; bracts in ca. 3 rows on and below the margin and on the fringe, smaller ones appressed, to 1.5mm long,puberubroadlyovate(to semiorbicular), lous;staminateflowers amongthe pistillateones; tepals

TAXONOMIC TREATMENT

<;:

va dr Elst Utrecht.]

177

X~~~

178

FLORANEOTROPICA

rowsonandbelowthemarginandonthefringe,broadly ovateto semiorbicular, to 1.5mmlong,appressed, subglabrous; staminate flowersamongthepistillateflowers;tepals2; stamens2; filamentslongerthantheperianth;stigmasalmostequalin length,0.5-0.6mmlong. Distribution (see Fig. 79.10). InVenezuela,known Endocarp body ca. 2.5 x 1.5-2 mm,? tuberculate.

2; stamens 2; filaments as long as or slightly longer thanthe perianth;stigmas equal or unequalin length, ca. 0.2-0.3 mm long. Endocarp body ca. 1.5 x 1 mm, distinctlytuberculate. froma few localitiesin the CoastalMountainRange;in shady and moist places; to ca. 1000 m.

Distribution(seeFig.79.10).InColombia(Magdalena andValle);in forest;at900-1400 m. Specimensexamined.VENEZUELA.Withoutlocal-

ity, 1865,Moritzs.n. (K).ARAGUA: ParqueNacionalHenri Specimensexamined. COLOMBIA.MAGDALENA: Pittier, RanchoGrande,Nov 1965, Aristeguieta5946 SierraNevada de Santa Marta,nr. QuebradaIndiana, (VEN). DISTRITo FEDERAL:Quebrada of Rio de La 1350 m, 29 Aug 1972, Kirkbride1962 (COL);upper Guaria,Apr 1854, Birschels.n. (K); nr. Caracas,Ernst Rio Buritaca,Altode Mira,rd.betweenCascadadel Cailo 1710 (BM); Tomade Mala de Platano,ca. 1000 m, 31 Negro and Finca Merado, 900-1 100 m, 15 Jul 1989, Mar 1972, Morillo et al. 2128 and 2131 (VEN); Madriiainet al. 281 (MO). CerroNaiguata,QuebradaFrontina,900-1 100 m, This speciesshows close affinitiesto Dorstenia 2 Nov 1963, Steyermark91841 (F, NY, US, VEN); panamensis. betweenLasDeliciasandLaTomade Quebradade Mata de Platano,Oct 1966, Steyermark97521 (NY, U, US, VEN); above NaiguatA,Rio Camuri Grande,20-22 16. Dorstenia panamensis C. C. Berg,Proc.Kon.Ned. May 1992, Meier et al. 2299 (BG). Akad.Wetensch.,Ser.C, Biol. Med. Sci. 85: 32, t. 1. Dorstenia aristeguietaeis ratherpolymorphic.The 1982. Type. Panama.Col6n: Rio Boqueron,5 Sep two collections madeby Steyermarkareespeciallydis1977, Berg, Maas & Dressler 415 (holotype, U; tinct,e.g., in the greaterdimensionsof severalpartsand isotypes,BG, K, MO, NY, PMA, RB). Fig. 47 in shorterand less dense indumentum.These differHerb,to ca.25 cmtall;stem3-4 mmthick,(rather) ences resemble those of D. peruviana.

tohirsutewithuncinate,straightand denselyhirtellous curvedhairs;intemodes0.5-2 cm long. Leaves in spi-

15.DorsteniacolombianaCuatrecasas, RevistaAcad. Colomb. Ci. Exact. 9: 341. 1956; Carauta, Rodriguesia 29(44):85, t. 17. 1978.Type.Colombia.Valle:valleyof RioSanquinini, LaLaguna,14

rals;laminaoblong,5.5-11 x 2.5-4.5 cm,membranaceousto chartaceous; to subacute; base apexacuminate (narrowly)roundedto cordate;marginrepandto ? dis-

tinctlycrenate; uppersurfacesmooth,glabrous;lower Dec 1943, Cuatrecasas 15508 (holotype, F; surfacescabrous, to hirtellousontheveins; puberulous isotype,US). Fig. 46 midribslightlyimpressed above,thevenation prominent beneath;lateralveins 10-15, loop-connected; tertiary Herb,probablyto 50 cmtall;stem2-4 mmthick, venation subscalariform to reticulate; petioleca. 1 cm withretrorsestraight sparselyappressed-puberulous tohirsute;stipulesbroadlytriangular long,hirtellous to andcurvedhairs;internodes1-5 cm long.Leavesin ovate,to0.7cmlong,faintlyplurinervate, acuteto sublaminaelliptictooblong,9-15 x 4.5-6 cm,charspirals; acuminate,on the older partof the stem often patentto basesubcordate taceous;apexacuminate; to rounded; deflexed,sparselypuberulous.Inflorescencesgreen;pemarginratherfaintlyandcoarselycrenate;uppersur-

duncle0.5-3.5 cm long, puberulouswith retrorseunciface smoothor ? scabrous,with sparseminuteconical natehairs;receptaclecentrallyattached,discoidto broadly onthemain hairs;lowersurface? scabrous, puberulous ? irregularly turbinate, lobedto suborbicular inoutline,

veins;lateralveins8-10 pairs,loop-connected; tertiary toreticulate; venationsubscalariform petiole1-4(-9.5) cm long, puberulousto substrigillosewith retrorse to straightorcurvedhairs;stipulesbroadlytriangular, 0.5cmlong,oftenwitha ? auriculate base,onthelower partof thestempatentto reflexed,subglabrous. Inflorescencesgreen;peduncle1-5 cmlong,puberulous to subhirtellouswith ? retrorsehairs;receptaclecentrally attached,discoid, (sub)orbicularin outline, 0.5-2 cm diam., the outside sparsely puberulous, partly with uncinatehairs,themarginlobed,the fringe0.5-1.5 mm

broad,onlywithminuteglobosehairs;bractsin ca. 3

0.5-1.5cmdiam.,theoutsidepuberulous withretrorse uncinate hairs,thefringe0.5-1.25mmbroad,withminute globose hairsandappresseduncinatehairs;bractsin ca. 2 rows on and below the marginor also on the fringe, to 2 mm long, appressed,broadlyovateto semiorbicular, staminateflowersamongthepistillateflowsubglabrous; ers;tepals2; stamens2; filamentsshorterthanthe perianth;stigmasequalorunequalin length,0.2-0.3 mmlong, white.Endocarpbodyca. 1.5 x 1-2 mm, ? tuberculate. Distribution (see Fig. 79.10). In Panama(Col6n);

in forest;atlow elevations.

179

TAXONOMIC TREATMENT

-

fgz

f:.'~~~~~~~~L

Fig. 46. Dorstenia colombiana. Leafy partof stemwith inflorescences(Cuatrecasas15508). [By E. M. Hupkens

van der Elst, Utrecht.]

Specimens examined. PANAMA. COL6N: Rio 17. DorstenialindenianaBureau,in De Candolle,Prod. Guanche,5 Jul 1979,Antonio1240 (MO);Rio Guanche, 17: 269. 1873. Type. Mexico. Tabasco:nr. Teapa, 21 Aug 1974,Dressler4690 (F); Rio Boquer6n,6-8 km Sep-Oct 1839, Linden64 (holotype,P; isotypes, G, upstreamfrom Peluca HydrographicSation. 29 Aug Fig. 48 K, LE). 1974, Dressler 4698 (PMA);betweenCerroBrujaand Rio Escandalose,20 May 1978, Hammel3221 (U). Herb,to 50 cm tall;stem 2-5 mm thick,hirtellousto The inflorescencesofthe collectionsHamiltonet al. hirsute with predominantly straight or a mixture of 2753 from Cocle and Hammelet aL 5085 from Col6n straightand uncinate patent or often ? retrorsehairs; intemodes 1-3 cm long. Leaves in spirals;laminaob(also mentioned in the discussion of Dorstenia choconiana) have inflorescences like those of D. long to obovate to elliptic or oblong, 2.5-14 x 1.5-6.5 panamensis,buttheleavesmatchthoseof D. choconiana. cm, chartaceous,usually variegated;apex roundedto

FLORANEOTROPICA

180

1cm.

Fig. 47. Dorsteniapanamensis.Leafy partof stem with inflorescences(Berg et al. 415). [By E. M. Hupkens van der Elst, Utrecht.]

broadlysubacute;basecordate;marginrepandto coarsely crenate;uppersurfacescabrouswithminuteconicalhairs and sparsebristle-likehairs;lower surface? scabrous, puberulousto hirtellouswith straightanduncinatehairs, on the veins denserand almosthirsute;lateralveins 811 pairs, faintly loop-connected or terminatingat the margin;tertiaryvenation subscalariformto reticulate; petiole1.5-5 cm long,withindumentumlikeon thestem; stipulesnarrowlytriangular,sometimeswith a swollen base,to 0.6 cm long, to 0.2 cm broad,usuallywith a few bristle-likehairs,sometimeswith? patentInflorescences on minuteshort-shootswith reducedstipules,green or the receptaclewith a purplemarginandfloweringface; peduncle2-4 cm long, puberulousto hirtellousor sometimes hirsutewith straightto uncinate,often ? retrorse hairs;receptaclecentrallyattached,broadlyturbinateto almost discoid, (sub)orbicularin outline, 0.5-1.2 cm diam.,the outside sparselypuberulous(to hirsute)predominantlywith uncinate,? retrorsehairs,the margin (sub)entireto dent(icul)ate,the fringe ca. 1 mm broad, sparsely,minutelypuberulous;bractsin ca. 3 rows, on and below the margin and on the fringe, appressedor

theouterones often? radiatingand/oron theteethofthe margin,ovate to oblong, 0.3-0.8 mm long, puberulous or the largerones setulose;staminateflowers amongthe pistillateones;tepals2; stamens2; filamentsshorterthan theperianth;stigmasalmostequalin length,0.2-0.3 mm long. Endocarpbodyca. 3 x 2 mm, slightlytuberculate. Distribution (see Fig. 79.1 1). In Mexico, Guatemala, andBelize; in shade,often in rocky places, often limestone, along streams;to 1100 m. Specimensexamined.MEXICO. CHIAPAs: 6-12 kmn S of Palenque,23 Nov 1972,Breedloveet al. 29773(CAS); Mun.Ocosingo,70 km SW of Palenque,11 Nov 1980, Breedlove47409 (CAS), 4 Dec 1980, Breedlove48296 (CAS),14 Jan 1981,Breedlove49587 (CAS);Mun.Palenque, nr.AguaAzul, 11 Nov 1981,Breedloveet aL 55402 (CAS);25 kmS of Palenque,15 Jan1982,Breedloveet al. 57334 (CAS);15 IkmS of Palenque,13 Dec 1986,Breedlove et al. 66322 (CAS);Mun.Ocosingo, S of Frontera 4 Dec 1984,MartinezS. 8966 Rio Usumacinta, Echeverria, (F); 2 km N of Naja,24 Sep 1988,Stevens25840 (BG). GUATEMALA. ALTA VERAPAZ: Chahal,Sebol rd., 30 Sep 1968, Contreras7771 (LL, U); nr. Cubilquitz,

181

TAXONOMIC TREATMENT

:

4~~~~~~~c

Fig. 48. Dorstenia lindeniana. 1. Habit with inflorescence in fruit. 2. Inflorescence (Tuerckheim I.5808). [By E. M. Hupkens van der Elst, Utrecht.]

3-6 Mar 1942, Steyermark 44542 (F), 44732 (F, NY), Aug 1907, TuerkheimII-43 (BR, F, MO, NY, 0, US, Z), Mar 1900, (in Donnell Smith) 7667 (F, GH, NU, US); nr. Chirriacte, 9 Apr 1941, Standley 91635 (F); between Campur and Socoy6, 9 Apr 1941, Standley 91728 (F). IZABAL:Puerto Mendez, 20 Aug 1966, Contreras 6024 (LL); Lago Izabal, 26 Apr 1966, Jones et al. 3130 (F, NY); Mun. Livingston, rd. Laguna Salvador-Calix, 30 Jul 1988, Martinez S. et al. 23137 (BG); Mun. Puerto Barrios, rd. Puerto Barrios-Punta de Palma, Rio Las Escobas, 10 Sep 1988, Martinez S. et al. 23679 (MO); nr. Escobas, W of Puerto Barrios,2 Jun 1922, Standley 24850 (GH, NY, US), 3 May 1939, Standley 72975 (F), 13 Apr 1940, Steyermark 39289 (F); Montanias del Mico, 7-8 km W of Santo Tomas de Castilla, 19 Aug 1988, Stevens 25552 (BG); Montanas del Mico, ca. 10 km W. of Izabal,

2 Apr 1940, Steyermark38607 (F); Rio Frio, 17 Dec 1941, Steyermark39972 (F, US). PETAN: Dolores, 4 Sep 1961, Contreras 2842 (LL): rd. Sebol-Coban, km 4, 16 Apr 1964, Contreras 4338 (LL); Sebol, Rio Sebol, Jul 1964, Contreras5303 (LL); Los Arcos, Cadenas rd., 9 Dec 1969, Contreras 9308 (BM, LL, U); San Luis, 11 Jul 1959, Lundell 16376 (LL); Laguna Petexbat6n, 11 Apr 1964, Lundell 18192 (LL); Rio Pasi6n, San Juan Acul, 17 Mar 1964, Lundell 18193 (LL); La Cumbre, 30 Jun 1975, Lundell et al. 19482 (LL), 12 Jul 1976, Lundell et al. 20023 (LL); Rio Roman, W. of Chinaja, 30 Mar 1942, Steyermark45516 (F, NY); Cerro Ceibal, between mouth of Rio San Martin and mouth of Rio Santa Monica, 29 Apr 1942, Steyermark 46070 (F, LL, NY). BELIZE. TOLEDO: Southern Maya Mtns., Bladen Nature Reserva, 13 May 1996, Davidse et al. 35875 (BG,

182

FLORANEOTROPICA

MO),20 May 1996,Holstet al. 5455 (BG);PuebloViejo, purple, or also the perianthsof the pistillate flowers 9 Feb 1934, Schipp 1276 (F, GH, MICH, MO, NY, Z). purplish;peduncle 2-25 cm long, sparsely to rather Dorstenia lindeniana is distinct from the other denselypuberulouswithretrorsehairs;receptacleeccenmembers of the CentralAmerican-Andean group of trically(to almostcentrally)attached,subflabellate,concave, elliptic to suborbicularin outline, 1.5-5 cm wide species by the scabrousupper surfaceof the lamina. (orlong), occasionallylanceolate-lingulate,the outside minutelyappressed-puberulous with retrorsehairs,the 18. Dorstenia elata Hooker, Ic. P1., Ser. 1, 3: t. 220. marginentireor faintly lobed, the fringe narrowto al1840; Miquel, in Martius,Fl. Bras.4(1): 160, t. 55. most lacking;bractsin ca. 2(-3) rows on the margin, 1853;Bureau,in De Candolle,Prodr.17:270. 1873; ovateto subovate,0.5-2.5 mm long, ciliolate;staminate Carauta,Rodriguesia29(44): 68, t. 1. 1978. Type. flowers amongthe pistillateones; tepals 2; stamens2; Brazil.Rio de Janeiro:Serrados Orgaos,Feb 1837, filamentslongerthantheperianth;stigmas(almost)equal Gardner 621 (holotype, K; isotypes, BR, FI, G, in length, ca. 1-1.2 mm long, white; on the perianths GH, K, NY, P, US). Fig. 49 andamongthe flowersminuteconical hairs.Endocarp bodyca. 4 x 3.5 mm, slightly tuberculate. Dorstenia longifolia Moricand, P1. Nouv. Amer. 90, t. 58. 1840; Miquel, in Martius, Fl. Bras. 4(1): 164. 1853; Bureau, in De Candolle, Prodr. 17: 270. 1873; Carauta, Rodriguesia 29(44): 70, t. 2. 1978. Type. Brazil. Bahia: without locality, 1834, Blanchet 1960 (holotype, G; isotype, BM). Dorstenia plumeritfolia Fischer & Meyer, Index Sem. Hort. Petrop. 11: 65. 1846; Miquel, in Martius, Fl. Bras. 4(1): 161. 1853; Bureau, in De Candolle, Prodr. 17: 270. 1973. Type. Brazil. Rio de Janeiro: Serrade Macae, May 1832, Riedel & Luschnath 308 (holotype, LE; isotype, US). Dorstenia sucrei Carauta, Bot. Mus. Bot. Curitiba 17: 2. 1974; Carauta, Rodriguesia 29(44): 74, t. 6. 1978. Type. Brazil. EspiritoSanto:Mun. Itagua9u, Limoeira, Santa Maria, 29 Jun 1946, Brade et al. 18278 (holotype, RB; isotypes, F, GUA, U).

Distribution (see Fig. 81.4). In eastemBrazil(from Bahiato Rio de Janeiro);in moist andshadedplaces;at low elevations.

Representative specimens examined. BRAZIL. BAMun. Itamarajui,BR.101, km 4, Ribeiro de Fora, 7 Apr 1972, Pineiro 1825 (CEPEC, GUA, K, U); valley of Rio Mucuri, BR.101, 2 Apr 1971, Santos 1539 (U, WAG); Mun. Itamarajfi, Campo Alegre, Piraja, 14 May 1971, Santos 1626 (CEPEC, RB). EsPIRITO SANTO: Mun. Nova Venecia, Serra dos Aimores, 15 Nov 1953, Duarte 3623 (RB); Mun. Santa Teresa, Esta9ao Biol6gica da Caixa d'Agua, 8 Aug 1985, Boudet Fernandes 1379 (US); Mun. Concei9ao do Castelo, Alto Baranal, 18 Oct 1985, Hatschbach 49956 (BG); Mun. Domingos Martins, BR.262, km 11, nr. Concei9ao do Castelo, 20 Oct 1979, Kautsky 606 (GUA, RB); Mun. Linhares, Goitacazes, Rio Herb, to 1 m or to 2 m tall; stem 2-8 mm thick, D6ce, 31 Mar 1934, Kuhlmann 127 (K, MBM, RB, U), sparselyto densely, ? retrorselystrigillose, strigose, 19 Nov 1943, Kuhlmann6500 (RB, U, US); VargemAlta, subsericeous, subpilose, hirtellous to subhirsuteand/ Sao Jose de Frunteira,12 Dec 1956, Pereira 2285 (GUA, or (partly)puberulous;intemodesto 8 cm long. Leaves RB). MINAs GERMS: Mariana,Jul 1824, Riedel 359 (LE, almost distichous; lamina elliptic to lanceolate to US). RiO DE JANEIRO: Mun. Maje, Dtto. Guapimirim, Centro de Primatologia, 27 Aug 1984, Andreata et al. subobovate or tending to pandurate,11-35 x 3.5-14 cm, chartaceousto subcoriaceous;apex acuminateto 624 (RB); Mun. Santa Maria Magdalena, Serra de Rifa, obtuse to roundedor emarginate;base obtuse to trun- Parque Nacional do Desengano, 25 Nov 1977, Carauta cateto cordate;marginrepandto coarselyto finely,regu- 2800 (RB); Mun. Campos, Morro do Coco, Cruz da Serra, Morro Bau, 13 Dec 1982, Carauta et al. 4419 (GUA); larly or irregularly,crenate-dentateto crenate;upper Mun. Nova Friburgo, Alto Macae do Nova Friburgo, 2 surfaceglabrous;lower surfacemainly or only on the Oct 1892, Glaziou 20495 (B, BR, K, P); Mun. Macae, veins sparsely to densely hirtellous to hirsute to Bicuda Pequena, nr. Cachoeiros, 14 Jul 1982, Gurkens.n. retrorselypilose to appressed-puberulous, often ? sca- (GUA); Mun. Teres6polis, Parque Nacional da Serra dos brous; at least the midrib impressedabove, the vena- Orgaos, Rio Soberbo, 28 Nov 1979, Lima 1165 (GUA, tion prominentbeneath;lateralveins 15-22 pairs,loop- RB); Mun. Silva Jardim, Reserva Biologica de Pogo das connected;tertiaryvenationreticulateto (sub)scalariform; Anta, 5 Nov 1982, Martinelli et al. 8898 (RB); Mun. petiole 0.5-8 cm long, retrorselystrig(ill)ose to sub- Casimiro de Abreu, Corrego da Luz, Pogo Pai Joao, 11 sericeousto hirtellousor to patentpuberulous;stipules Oct 1982, Pedroso et al. 612 (GtJA).

ovate to triangular,(0.7-)1-2 cm long, coriaceous, conspicuouslyplurinervatein broadstipules,less so in narrowones, retrorselypilose to (sparsely)puberulous to hirtellous,especially on the marginandthe midrib, the apex subobtuse to acute. Inflorescences green or the receptacle outside slightly purple and the margin

HIA:

Dorstenia elata is variable, such as in the length of the petiole and in the indumentum. The collections Brade et al. 18278 and Sucre 10969 represent a form with a

short(1-1.5 cm long) peduncle, a lingulatereceptacle 0.5-1 cm broad,relativelynarrowstipules,in whichthe venationis inconspicuous,andanindumenplurinervate

TAXONOMIC TREATMENT

183

Fig. 49. Dorstenia elata (from Martius, Fl. Bras. 4(1): t. 55. 1853). Leafy part of stem with inflorescence, staminate flowers (1,2) stamens (7), stigmas (13), exocarp body and embryo (21, 24), fruits (17), empty exocarp (23).

tumof predominantlypatent(ratherthanretrorse)hairs. It was describedas D. sucrei andmightrepresenta distinct infraspecifictaxon.

19. Dorstenia urceolata Schott,Flora4(1): 197. 1821; Miquel, in Martius,Fl. Bras. 4(1): 162. 1853; Bureau,in DeCandolle, Prodr.17:266. 1873;Carauta, Rodriguesia29(44):71, t. 4. 1978. Type.Brazil.Rio de Janeiro:Seffada Estrela,Schotts.n. (W,destroyed, no isotype traced;neotype, Carauta,Rodrigu6sia 29(44): 3. 1978. Brazil. Rio de Janeiro: Mun. Petr6polis,Serrada Estrela,Meio da Serra,10 Oct 1971, Carauta 1409, RB; isoneotypes, CTES, F, GUA, LIL, MBM, MICH, U). Fig. 50 DorstenianervosaDesvaux,MWm.Soc. Linn. Paris 4: 217. 1826;Miquel,in Martius,Fl. Bras.4(1): 161, t. 56. 1853; Bureau,in De Candolle,Prodr. 17: 266. 1873.Type.Brazil.Rio de Janeiro:Mun. Maje, Serrada Estrela,Mandioca,1821, Langsdorif s.n. (holotype, P). Dorstenia nervosa Desvaux var. latifolia Desvaux, MWm.Soc. Linn. Paris4: 217. 1826. Dorstenia nervosa Desvaux forma latifolia (Desvaux) Miquel, in Martius,Fl. Bras. 4(1): 162. 1853. Dorstenia urceolata Schott var. latifolia (Desvaux)

Bureau,in De Candolle, Prodr.17: 266. 1873. Type.As the species.

Dorstenia nervosa Desvaux var. angustifolia Desvaux, M6m. Soc. Linn. Paris 4: 217. 1826. Dorstenia nervosa Desvaux forma angustifolia (Desvaux) Miquel, in Martius, Fl. Bras. 4(1): 162. 1853. Dorstenia urceolata Schott var. angustifolia (Desvaux) Bureau, in De Candolle, Prodr. 17: 266. 1873. Type. Brazil. Without locality, Gaudichaud s.n. (holotype, P). Dorstenia caulescens Vellozo, Fl. Flumin. 55. 1829 ("1825"), Ic. 1: t. 141. 1831 ("1827"), non Linnaeus, 1753; Fielding & Gardner, Sert. P1. 1: t. 13. 1843; Miquel, in Martius, Fl. Bras 4(1): 162. 1853; Bureau, in De Candolle, Prodr. 17: 266. 1873. Dorstenia langsdorffiana Fischer & Meyer, Index Sem. Hort. Petrop. 11: 64. 1846, as a synonym of D. caulescens Vellozo. Type. Fl. Flumin. Ic. 1: t. 141. 1831 ("1827"). Dorstenia caulescens Vellozo var. angustifolia Fielding & Gardner, Sert. P1. 1: t. 13. 1843. Type. Brazil. Rio de Janeiro: nr. Rio de Janeiro, Luschnath s.n. (not traced). Dorstenia caulescens Vellozo var. Iatifolia Fielding & Gardner, Sert. P1. 1: t. 13. 1843. Type. Brazil. Without locality, Riedel s.n. (not traced). Dorstenia urceolata Schott var. variegata Bureau, in De Candolle, Prodr. 17: 267. 1873. Type. Described from material cultivated in the greenhouse of the Mus6um National d'Histoire Naturelle in Paris, Anonymous s.n. (P).

184

FLORA NEOTROPICA

Fig. 50. Dorstenia urceolata (from Martius, Fl. Bras. 4(1): t. 56. 1853). Habit, receptacle (31), staminate flower and tepals (4), stamens (7), pistil (10), stigmas (13), empty exocarp (18).

TAXONOMICTREATMENT Herb, to 25 cm tall; stem 3-5 mm thick, minutely puberulous;intemodes to 2.5 cm long. Leaves in spirals;laminaelliptic to lanceolate,often tendingto subovate, 5-17 x 2-6.5 cm, chartaceous,sometimes variegated;apexacuteto subobtuse;baseobtuseto rounded or to subacute;marginsubentireto repandorto coarsely crenate-dentate;uppersurfaceglabrous;lower surface minutelypuberulousto glabrous,sometimesscabridulous;venationprominentbeneath;lateralveins6-10 pairs, loop-connected;tertiaryvenation(sub)scalariform(to reticulate);petiole 3-9 cm long, minutelypuberulous to glabrous;stipulessubovateto triangular,0.5-1.2 cm long, plurinervate,usually only the midribconspicuous, subacuminate,minutely puberulousto glabrous. Inflorescences with the whole receptaclered-brown; peduncle 6-15 cm long, minutely puberulous;receptacle centrallyto eccentricallyattached,cup-shaped(to urceolate),orbicularin outline, 1.5-2.5 cm diam., the outsideminutelypuberulous,usuallyratherdensely so on prominentribs leading to the bracts,or sometimes glabrous, the margin subcrenate,the fringe to 2 mm broad,minutelypuberulous;bractsin a singlemarginal row, appressed,lanceolateto oblong, to 1.5 mm long, smallerand largerones ? altemating,minutelypuberulous;staminateflowersamongthepistillateones;tepals 2; stamens2; filaments longer thanthe perianth;stigmas equal in length, ca. 0.8 mm long, red-brown.Endocarp body ca. 2.5-5 mm, tuberculate. Distribution (see Fig. 79.16). In East Brasil (Rio de Janeiroand Parana);in moist and shadedplaces; at low elevations.

185 Herb,to 30 (?) cm tall;stemwith a repent,rhizoma-

withretrorse straight touspart,2-4 mmthick,puberulous internodesto 4 cm to uncinatehairs,or subglabrous; to oblong(or long.Leavesin spirals;laminalanceolate ovate to subovate),(3.5-)12-18 x (1.5-)3.5-5 cm, apexacuteto subacumchartaceous to subcoriaceous; (todecurrent); margin inate;baseacuteto subattenuate uppersurfaceglasubentire to faintlycrenate-dentate; brous;lowersurfacesparselyandminutelypuberulous on themainveinsto subglabrous; lateralveins 10-13 tertiary venationreticulate; petipairs,loop-connected; ole3.5-8(-10)cmlong,puberulous withretrorse, straight stipulestri(tocurved),uncinatehairs,orsubglabrous; sparselypuberuangular,0.5-1 cm long,uninervate, Inflorescences: hairsorsubglabrous. louswithretrorse withretrorse 4-10 cmlong,puberulous straight peduncle receptacle slightlyectouncinate hairs,orsubglabrous; inoutlineand discoid,suborbicular centrically attached, ca. 1cmdiam.orellipticto obovateinoutlineand0.8withuncinate 1.2x ca.0.8 cm,theoutsidepuberulous thefringenarandstraight hairs,themarginsubcrenate, rows,ovate,to 1mm row;bractsinca.2 (sub)marginal staminateflowersamong long,minutelypuberulous; aslong thepistillateones;tepals2;stamens2;filaments stigmasnotseen. asorslightlylongerthantheperianth; Endocarpbodyca.2 x 2 mm,tuberculate. Distribution(see Fig. 79.16). In easternBrazil (EspiritoSanto);in moistandshadedplaces;at low elevations.

Specimensexamined. BRAZIL. ESPiRITO SANTO: Pontode Balan9o,10 Oct 1986,Bittancourts.n. (GUA); Representative specimens examined. BRAZIL. Mun.DomingosMartins,Morrodo Chapeu,6 Feb 1981, PARANA: Serrado Mar,VoltaGrande,SerraMarumbi, Kautsky676 (GUA, mixed with anotherspecies, prob13 Feb 1904, Dusen 3548 (P). RiO DE JANEIRO: Mun. ably Dorsteniaturnerifolia). Nova Igua9u,Serrada Bandeira,17 Oct 1977, Araujo The speciesshowsstrongaffinitiesto Dorstenia et al. 1889 (GUA); Mun. Rio de Janeiro,Corcovado, Lagoinha,8 Sep 1865, Glaziou1010 (P); Mun. Rio de urceolata in its vegetative parts. Janeiro,Tingua,nr.Represado Rio Piaba,28 Sep 1960, Pabst5395 (HB,M, MICH);Mun.Maje,Serrada Estrela, DORSTENIA SECTION Mandioca,1 Mar 1823, Riedel s.n. (NY, US). 9B. Dorstenia Linnaeus sect. Dorstenia. Dorsteniaurceolatais a ratheruniformspecies,disSychiniumDesvaux,MWm.Soc. Linn. Paris4: 216. tinctivedueto thered-browncup-shapedreceptacle.The 1826. Dorstenia Linnaeus sect. Sychinium species occasionallyhas variegatedleaves;plantswith Endlicher,Gen. P1. 279. 1837. Dor(Desvaux) such leaves are still in cultivationin severalEuropean stenia Linnaeus sect. Sychinium (Desvaux) botanicalgardens.The species appearsto be closely reFischer& Meyer,IndexSem.Hort.Petrop.11:57. latedto D. hildegardisandmore distantlyto D. elata. 1846.DorsteniaLinnaeussect. SychiniaCarauta, Bradea 2(21): 151. 1976. Dorstenia Linnaeus series Furcatae Lemaire,Ill. Hort. 10: t. 363. 20. Dorstenia hildegardis Carauta,Valente& Barth, 1863, to replacesect. SychiniumDesvaux.Type. Rev. Brazil. Biol. 38(3): 613, t. 2, 3. 1978. Type. Sychinium ramosum Desvaux (= Dorstenia Brazil. Espirito Santo: Mun. Domingos Martins, ramosa(Desvaux)Carauta,Valente& Sucre). Campinho,rd. to Chapeu, 15 Sep 1975, Carauta DorsteniaLinnaeusseriesScapigeraeSprengel,Syst. 1811 (holotype, RB, isotypes F, GUA, HB, K, NY, Veg.3: 777. 1826.Lectotype,Berg& Hijman,Ilicifolia 2: 67. 1999.DorsteniacontrajervaLinnaeus. R, SP, U, UB). Fig. 51

186

FLORA NEOTROPICA

''4''.'

4. ................. . ';8t\r. 2A;.

~~~~~~v; J.'~~~~~~~~~' ?*V..

*

;..

.

.

....

.

.

N

9W.

-..

*

..


..... ................. '*-t.:4 5 ,/:

!:

:

R

-

sR

;

/

!

~~~~~~~~~~~~~~ ffi~~~~~~~~~~~ ||;t :v

Fig. 51. Dorsteniahildegardis.Leafypartof stem withinflorescence(Carauta1811). [By H. Rypkema,Utrecht.]

Dorstenia Linnaeus sect. Eudorstenia series Acaules Endlicher, Gen. P1. Suppl. 4(2): 35. 1847. Lectotype, Berg & Hijman, Ilicifolia 2: 67. 1999.

Dorstenia contrajervaLinnaeus. Dorstenia Linnaeus series Quadratae Lemaire, Ill. Hort. 10: t. 362. 1863. Lectotype, Berg & Hijman, Ilicifolia 2: 67. 1999. Dorstenia contrajerva Linnaeus.

Plants herbaceous, mostly(sub)acaulescent; stem unbranched, oftenpartlysubterranean; internodesoften shortandthenthe leavesrosulate.Leaves in spirals;laminamostlyovateto (sub)cordate, oftenpinnately(tosubpalmately orsubpedately) lobedtoparted; venation(basically)brochidodromous to reticulate; orplurinervate. petiolelong;stipulesbroad,uninervate

TAXONOMICTREATMENT

(ellipticto oboInflorescences mostly? zygomorphic to moreor less irvatein outline),(sub)quadrangular regularlylobedor (in D. ramosa)furcate,bracteate, orappendiculate (in (bractssometimes onappendages) pistillateflowDorstenia ramosa), thefringenarrow; and ersnumerous.Endocarp body mostlytetrahedral tuberculate, sometimes smooth;seedsmall,endosperm present;embryowithflatandequalcotyledons.

187 DorsteniaanthuriifoliaBlake,Contr.U.S. Natl.Herb. 20: 237. 1919; Carauta, Rodriguesia 29(44): 101. 1978. Type. Brazil. Bahia:Rio Gongoji, 1 Oct-30 Nov 1915, Curran130 (holotype, US).

Herb, to 50 cm tall; leafy stem for the greater part supraterranean,to 25 cm long, erect or ascending, 6-12 mm thick, glabrous; internodes to 1 cm long; latexyellow. Leaves in spirals, initially subrosulate; lamina elliptic to lanceolate, 9-27 x 3-12 cm, chartaceous; apex acuminate andcom- to acute; base cordate to rounded to truncate or to subThissectionis confinedtotheNeotropics priseseightspecies:six of themin eastemBraziland acute; margin finely and often ? irregularly crenate-dentwo of themrangingfromMexicoto northemCosta tate to repand; upper surface glabrous; lower surface glaRicaorfromMexicoto Peru,Venezuela,andtheCar- brous; lateral veins 9-12 pairs, loop-connected; tertiary ibbean,respectively. venation often partly scalariform;petiole 10-21 cm long, Mostof the speciesareconfinedto therainforest glabrous; stipules subovate to triangular, 0.5-1 cm long, habitat.Dorstenia contrajervaextendsto driertypes faintly plurinervate, with only the midrib prominent and of forest(semideciduous forest)anddoesoccuralsoin conspicuous, subacuminate, glabrous. Inflorescences with secondarygrowth.Dorstenia drakenaoftenoccursin the receptacle outside green to slightly purplish(-brown) dryforestor scrub,in whichthestemcanbe ? tuberand with the margin and the flowering face (partly) dark ous,butit is notclearwhetherthisspeciesis geophytic purple to purplish-brown, the flowering face initially facundersuch(seasonally)dryconditions. ing downwards, later upwards; peduncle 5-15 cm long, cordate, commonly Therelativelylarge,? distinctly glabrous; receptacle centrally attached, discoid to broadly ofthis incisedleaveswithlongpetiolesarecharacteristic turbinate,the flowering face ? concave, orbicularto ? prosection.Thistypeof leafis associatedwitha reduced nouncedly (quadr)angular in outline, 1-3(-4) cm diam., However,in thissectionthe lengthof theinternodes. the outside minutely puberulous, mainly with retrorse gradually passesinto growthformwithlongintemodes hairs, the margin lobed to crenate, the fringe narrow to the growthform with shortinternodes.Dorstenia almost lacking;bractsin 2(-3) rows on the margin,broadly hasstemswithmostinternodes contrajervaoccasionally ovate to semiorbicular, appressed, to 2 mm long, glabrous long.Dorstenia bahiensis is distinctlysubacaulescent or minutely puberulous; staminate flowers among the piswhenyoung,lessdistinctlyso later.Dorsteniaramosa tillate ones; tepals 2; stamens 2; filaments longerthan the is subdivided intotwosubspecies, onewithlonginterperianth; stigmas equal in length, ca. 0.3-0.4 mm long, nodesandtheotherwithonlyshortinternodes. pale green;minuteandthin to conical hairsamong the flowDorsteniaramosais so similarinitsvegetative parts ers and on the perianths present. Endocarp body ca. 2 x toD. arifolia(cf.Miquel,1853)andtoD.grazielaethat 1.5 mm, tuberculate.

ofitsinflorescence (ebracteate, appenthespecialfeatures of a diculate,andfurcate)do notjustifymaintenance Sect.Dorstenia section(cf.Carauta, 1976,1978a). separate can be regardedas derivedfromsect. Lecanium, in fromthegroupwithbroadstipules. particular

Distribution (see Fig. 79.16). In eastem Brazil (Pemambuco and Bahia); in moist and shaded places: at low elevations.

Representativespecimensexamined.BRAZIL.BAP6rtoSeguro,BR.5, km 18, 14 Sep 1961, Duarte 5660 (HB,NY, RB, U); Mun.Itabuna,CEPLAC,10 Jan 21.DorsteniabahiensisKlotzschex Fischer&Meyer, 1974,Harley15009 (K); Sao Bento,Luetzelburg247 (F, IndexSem.Hort.Petrop.11:64. 1846;Klotzschin M, NY);Mun.Ilheus,CEPEC,19 Jan 1971,Pineiro1006 Schlechtendal,Linnaea14: 288. 1840, nomen; (CEPEC,RB, U); Mun. Ilheus, Castelo Novo, Lagoa Miquel,in Martius,Fl. Bras.4(1): 164.1853;Bu- Encantada,FazendaPontaGrossa,15 Nov 1968, Vinha reau,inDe Candolle,Prodr.17:262. 1873;Valente 90 (CEPEC,K, RB, U). PERNAMBUCO:Maraial,Fazenda & Carauta,Cienc. & Cult. 27(6): 645, t. 1. 1975; Ceu Azul, 13 Oct 1957, Andrade-Lima2712 (IPA); 19 Mar Carauta,Rodriguesia29(44): 101, t. 28. 1978.Type. Quipapa,Vila de Igarapeba,EngenhoBrejinho, 11 (IPA);Rio Formosa,HortoFlorestal Barros-Lima 1966, Brazil. Bahia: Ilheus, 25 Jun 1837, Luschnaths.n. de Saltinho,9 Sep 1954,Falcdoet al. 993 (RB, U); Bo(holotype, LE; isotypes, B, BR). nito,FazendaCucau,15 Dec 1965,Marizet al. s.n. (IPA). DorsteniamartianaMiquel,in Martius,Fl. Bras.4(1): Dorstenia bahiensis is a distinct species, rather 165. 1853; Carauta,Rodriguesia29(44): 101. 1978. Type. Brazil. Bahia:Almanda,nr. Ilheus, variable in the shape of the lamina. It is one of the few Martius953 (lectotype,designatedhere, M; iso- species with yellow latex (also found inD. appendiculata). lectotypes,BM, BR, G, GH, K, L, LE, P, U). The plants are initially acaulescent, but older specimens HIA:

188

FLORANEOTROPICA

Distribution (see Fig. 79.1). In eastern Brazil aredistinctlycaulescent,withthestemto 25 cmlong. ThespeciesshowsaffinitiestoD. elata andD.grazielae (Minas,Gerais,Rio de JaneiroandSao Paulo);in moist inthecharacters of thestemandstipules.Theinitially and shadedplaces; to 1000 m. acaulescent habitandtheshapeof thereceptacle, tendRepresentative specimens examined. BRAZIL. withD. MINAs GERAIS: AntonioDias, UHE.Guilamn-Amorim, ingto be quadrangular, suggestrelationships contrajerva andwithD. ramosa. Nov 1998,L. V CostaBHCB44722(BG).Rio DE JANEIRO:

Valente&Sucre,Atas 22.DorsteniagrazielaeCarauta, Soc.Biol. Riode Janeiro16(2/3):59, t. 1-2. 1973; Valente& Sucre,Rodriguesia Carauta, 27(39):235, t. 2-3, 1974;Carauta, Rodriguesia 29(44):70, t. 3. 1978.Type.Brazil.Riode Janeiro: Parati,Meiode Serrade Parati,31 Oct 1971,Carauta 1411 (holotype,RB; isotypes,F, GUA,K, SP,U).

Mun. Parati,Parati-Mirim,8 Dec 1976, Carauta2222 (GUA,U); Mun.SilvaJardim,ReservaBiol6gicade Pogo dasAntas,Carautaet al. 2956 (RB);Mun.Cachoeirasde Macacu,Serrade Pati, 16 Feb 1977,Carautaet al. 2300 (GUA,RB, U); Mun.Rio Bonito, Bracana,Fazendadas Cachoeiras,22 Jan 1977, Lachette286 (R); Mun. Rio Claro,rd. RJ.16,nr.firsttunnel,16 Mar 1978,Lima262 (RB); Mun. Maje, Paraiso,Centrode Primatologia,19 Oct 1984, Limaet al. 2272 (GUA); Mun. SantaMaria Magdalena,FazendaMaterBoni, ca. 1000 m, 18 Feb 1981,Martinelliet al. 76087(GUA,RB);Mun.Campos, nr. borderwith Mun. SantaMariaMagdalena,Matado ImbE,21 Jan 1976,Peixotoet al. 838 (RB). SAOPAULO: Sao Sebastiao,28 Mar 1892,Edwalls.n. (SP); Ubatuba, 5 Nov 1961, Fontellaet al. 84 (SP).

Herb, to 1.5m tall;stempartlyrhizomatous, supraterranean partto 1.3m long,ascending, 4-10 mmthick, to hirtellous,partlywithunci(retrorsely) puberulous to 1.5(-2.5)cm long.Leaves in natehairs;internodes tosubpanspirals,spaced;laminaovatetosubcordiform In Dorstenia grazielae the leaves arenormallyenin thelowerpartoftenconstricted,10-40 x duriform, to acute;base tire, only occasionally lobed, whereas in D. arifolia 3.5-20 cm,chartaceous; apexacuminate cordatewitha wideto verynarrowsinus,thelobesof- specimenswith entireandincised leaves can be found tenoverlapping; crenate-dentate to side by side and occur in almost equal proportions. margin? irregularly ? deeplylobed;uppersurface repand, sometimes sparsely strigoseto almostglabrous;lowersurface(mainlyon with 23. Dorstenia albertii Carauta,Valente& Barth,Atas theveins)minutelypuberulous orpartlyhirtellous Soc. Biol. Rio de Janeiro 17(2): 63, t. 1-2. 1974, ortohirsuteon uncinate hairsand/orpartlyhispidulous, D. albertorum; Carauta,Rodriguesia 29(44): as themidrib; lateral veins8-10pairs,faintlyorhardly loop75, t. 7. 1978. Type. Brazil. Espirito Santo: venation reticulate tosubscalariform; connected; tertiary Mimoso do Sul, 24 Apr 1972, Sucre et al. 8920 withretpetiole10-35 cm long,minutelypuberulous (holotype, RB). Fig. 52 rorsehairsor partlyhirtellousor sometimeshirsute; 0.7-1.3 cm long, stipulesovateto broadlytriangular, Herb, to 20 (?) cm tall; stem for the greaterpart withthevenationfaintlyfla- supraterranean,erect or ascending, 2-3.5 mm thick, coriaceous,plurinervate, or puberulouswith uncinatehairs;internodesto 0.8 cm bellate,basesubcordate, apexacuminate, puberulous face long. Leaves in spirals, spaced; lamina oblong to partlyhispidulous. Inflorescenceswiththeflowering tored-brown; 5-12 cmlong,minutely subovate,7-18 x 2-7.5 cm, chartaceous;apex acumipurplish peduncle with? retrorse hairsorpartlyhirtellousor nateto subacute;base cordate;margincrenate-dentate; puberulous sometimes hirsute; receptacle centrally attached, discoid, uppersurfacesparselyhispidulous,scabridulous;lower (sub)orbicular (to elliptic)in outline,1-1.8 cmdiam., surfacehispidulousto puberulouswith uncinatehairs sometimes slightlylobed,theoutside minutely puberulous, on the main veins; lateralveins 7-10 pairs, loop-con? distinctly ribbedbyveinsleadingtothemarginal bracts, nected;tertiaryvenation reticulateto subscalariform; themargindentate toappendiculate, thefiinge1-1.5mm petiole 2.5-9.5 cm long, (minutely) puberulous to bractsin onerowon the (subhirtellous)with retrorsestraightand curvedhairs; broad,minutelypuberulous; marginoroftenontheapicesof shortoroccasionally to stipules ovate to subcordiform, 0.3-0.6 cm long, 2.5 cm long filiformappendages, departingfromthe plurinervate,subacuminate,sparsely hispidulous.Inmarginor occasionallybelowthemargin,appressed, florescences green with a red margin;peduncle ca. 3 ovateto triangular, to 2.5 mmlong,distinctlydifferent cm long, minutely puberulousto hispidulous; recepin length,shorterandlongerones? clearlyalternating, taclecentrallyattached,discoid,orbicularin outline,ca. staminate minutelypuberulous; flowersamongthepis- 0.8 cm diam., the outside minutely puberulous to tillateones;tepals2; stamens2; filamentslongerthan hispidulous,? distinctlyribbedwhendry,marginformtheperianth; stigmas(almost)equalin length,0.3-0.5 ing a subcrenaterim, the fringe narrow;bractsca. 10mmlong.Endocarpbodyca. 1.5 x 1.5mm,tuberculate. 12 (almost) in a single submarginalrow, radiating,

TAXONOMIC TREATMENT

189

Fig. 52. Dorstenia albartii. Leafy part of stem with inflorescences (Carauta 1669). [By E. M. Hupkens van der Elst, Utrecht.]

subspathulate,(0.5-)2-3 mm long, hispidulous;staminateflowersamongthepistillateones;tepals2; stamens 2; filamentsalmostas long as the perianth;stigmasequal in length, ca. 0.3 mm long. Endocarp body not seen.

CTES,GUA, HB, HBR, MG, NIT, RBR, RFA, Fig. 53 SP,UNB, USP,US, VIC).

orsupraterranean partof thestem Herb,acaulescent shortorto 7 cmlong,erectto ascending,1-5 mmthick, Distribution (see Fig. 79.16). In eastern Brazil puberulouswith retrorseuncinatehairs;intemodes (EspiritoSanto), only known from the type locality, a short,someof themto 0.8 cm long.Leaves in spirals, (sub)rosulate; laminain outlineovateto subcordiform, moist and shadedplace. (= pseudoalmostentireto 3-lobateormostlytripartite Specimensexamined. Materialfromthe type localx 6-14(-21) cm, membranatrifoliolate), 5-11(-17) ity, cultivatedin the BotanicalGardenof Rio de Janeiro, thelateralonesasymsymmetric, ceous,themidsegment Carauta 1669 (GUA). metric,entireor sometimes2-fid;apicesacuminate; Dorstenia albertii shows strong similarities to D. bases(sub)acute, thepetiolulesto 3 mmlong;margins grazielae. It is mainly distinct in the smallerinflores- subentireto crenate-dentate; uppersurface(sub)cences, shorterstipules, and shorterstamens.It might glabrous;lowersurfaceon theveinspuberulous with meritrecognitiononly at the subspecific level. hairs;lateralveinsof themidsegment (retrorse) uncinate venationreticulate 6-10 pairs,loop-connected; tertiary to subscalariform;petiole 4.5-17(-30) cm long, 24. Dorstenia bonijesu Carauta& Valente,Atas Soc. puberulous to curvedto uncitohirtellouswithstraight Bot. Rio de Janeiro 1(20): 112, t. 1-3. 1983. Type. natehairs;stipulesnarrow-ovate to 0.6 to triangular, Brazil. Rio de Janeiro:Mun. Bom Jesus de Itaba- cmlong,acuminate, withretrorseuncinate puberulous poana,Dtto. Carabu9u,FazendaSao Jorge,nr.Rio hairs.Inflorescencesgreen(?);peduncle6-12 cmlong, Itabapoana,16 Oct 1982, Carauta et al. 4400 (ho- graduallybroadenedinto the receptacle,puberulous centrally lotype, RB; isotypes, BC, CEPEC, COIM, COL, withmostlyretrorse straighthairs;receptacle

FLORA NEOTROPICA

190

Fi,g. 53.

Dosei

bnjs.

Hai

1-;aut

et

a.

attached, turbinate to shallowlycup-shaped, orbicular inoutline,2.5-3 cmdiam.,theoutsidepuberulous with retrorse uncinatehairs,with+ conspicuous, darker coloredvenationwhendry,themargin(sub)crenate, the fringeca. 3-4 mmbroad,minutelypuberulous; bracts in a single (sub)marginalrow, ovate, 0.5-1.2 mm long, shorterandlongeronesalternating, hispidulous; staminateflowersamongthepistillateones;tepals2; stamens2; filamentslongerthantheperianth; stigmas

440.

[B

,E;.

M.

Hukn,a

de

Elt

Utreht.

not seen. Endocarp body ca. 1.5 x 1.5 mm, smooth or tuberculate. Distribution (see Fig. 79.3). In eastern Brazil (EspiritoSanto, Minas Gerais, and Rio de Janeiro);in moist and shadedplaces; at low elevations. Specimensexamined. BRAZIL. ESPtRITOSANTO: Mun. Alfonso Cliudio, Empossado, 17 May 1991, Hatschbachet al. 55586 (SP);Cachoeirado Itapemirim,

TAXONOMIC TREATMENT

MorroGrande,13 May 1993,Mello-Silvaet al. 843 (NY, SPF). MINAs GERAIS: Mun. Governador Valladares, Campode Sementes, 25 Nov 1941, Magalhaes 841 (RB).

25. Dorstenia arifolia Lamarck, Encycl. 2: 3 17. 1786, Illustr. t. 83, fig. 2. 1791; Hooker, Exot. Bot. 1: t. 6. 1822; Bot. Mag. t. 2476. 1824; Loddiges, Bot. Cab. 10: t. 999. 1825; Vellozo, Fl. Flumin. 53. 1829 ("1825"), sub D. drakena; Fischer & Meyer, Index Sem. Hort. Petrop. 11: 60. 1846; Miquel, in Martius, Fl. Bras. 4(1): 166, t. 60. 1853; Carauta,Rodriguesia

29(44): 98, t. 27. 1978. Dorstenia multiformis Miquel var. arifolia (Lamarck) Bureau, in De Candolle, Prodr. 17: 261. 1873. Type. Brazil. Rio de Janeiro: probably nr. Rio de Janeiro, Dombey s.n. Figs. 54, 55 (holotype, P). DorsteniafaniaPaiva,Farmacop.Lisboa20. 1785;Mem. Hist.Nat. Acad. Real. Ci. Lisboa.64. 1790; Sprengel, J. Bot. (Schrader) 2: 238. 1801; Sprengel, Syst. Veg. 3: 777. 1826; Carauta, Rodrigu6sia 29(44): 98. 1978. Type. Not traced, probably destroyed. Dorstenia quadrata Desvaux, Mem. Soc. Linn. Paris 4: 218. 1826; Bureau, in De Candolle, Prodr. 17: 261. 1873. Type. Brazil. Without locality, Anonymous s.n. (holotype, P). Dorstenia cyperus Vellozo, Fl. Flumin. 53. 1829 ("1825"), Ic. 1: t. 140. 1831 ("1827"); Miquel, in Martius, Fl. Bras 4(1): 165. 1853: Wawra, Bot. Ergeb. 128. 1866; Bureau, in De Candolle, Prodr. 17: 261. 1873. Dorstenia riedeliana Fischer & Meyer, Index Sem. Hort. Petrop. 11: 58. 1846, as a synonym of D. cyperus Vellozo; Miquel, in Martius, Fl. Bras. 4(1): 165. 1853; Bureau, in De Candolle, Prodr. 17: 261. 1873. Type. Fl. Flumin. Ic. 1: t. 140. 1831. Dorsteniaficifolia Fischer & Meyer, Index Sem. Hort. Petrop. 11: 58. 1846; Miquel, in Martius, Fl. Bras. 4(1): 167. 1853. Dorstenia multiformis Miquel var. ficifolia (Fischer & Meyer) Bureau, in De Candolle, Prodr. 17: 261. 1873. Dorstenia arifolia Lamarck var. pinnatifida Miquel, in Martius, Fl. Bras. 4(1): 167. 1853, to substitute D. ficifolia Fischer & Meyer; Wawra, Bot. Ergeb. 128. 1866. Dorstenia multiformis Miquel var. pinnatifida (Miquel) Peckolt & Peckolt, Hist. P1. Med. 893. 1890. Type. Brazil. Rio de Janeiro: Corcovado, Riedel et al. 71 (lectotype, designated here, LE). Dorstenia mandiocana Fischer & Meyer, Index Sem. Hort. Petrop. 11: 59. 1846; Miquel, in Martius, Fl. Bras. 4(1): 165. 1853; Bureau, in De Candolle, Prodr. 17: 261. 1873. Type. Brazil. Rio de Janeiro: Mandioca, Riedel s.n. (holotype, LE). Dorstenia tentaculata Fischer & Meyer, Index Sem. Hort. Petrop. 11: 59. 1846; Bureau,in De Candolle, Prodr. 17: 261. 1873; Carauta, Valente & Sucre, Rodrigu6sia 27(39): 256. 1974; Carauta, Rodriguesia 29(44): 96, t. 25. 1978. Brazil. Rio

191

de Janeiro:nr.Rio de Janeiro,Riedel& Luschnath (holotype, LE). Dorsteniapeltata Fischer& Meyer,IndexSem.Hort. Petrop.11: 66. 1846, non Sprengel,1822. Dorstenia fischeri Bureau, in De Candolle, Prodr. 17:277. 1873,as a synonymof D. peltata Fischer & Meyer; Carauta,Rodriguesia29(44): 97, t. 26. 1978. Type. Brazil. Rio de Janeiro:Macae, Riedel s.n. (holotype, LE). DorsteniamultiformisMiquel, in Martius,Fl. Bras. 4(1): 165, t. 59, 60. 1853, nom. illeg., pro parte (forma B). Dorstenia strangii Carauta, Bradea 1(42): 433. 1974;CarautaRodrigu6sia29(44):73, t. 5. 1978. Type. Brazil. Minas Gerais:Mun. Bom Jardim de Minas,Serrada Mira,12 km S of Bom Jardim de Minas,3 Mar1964,Salehs.n. (holotype,HB). Dorstenia maris Valente & Carauta,Trab. XXVI Congr.Nac. Bot. Rio de Janeiro613, t. 82. 1975; Carauta,Rodriguesia29(44): 95, t. 24. 1978. Type. Brazil. Rio de Janeiro:Mun. Angra dos Reis, FazendaJapuiba,19 Mar 1951, Kuhlmann 2610 (holotype, SP). Herb, subacaulescent,with at least the lower part of the stem subterranean; stem 5-15 mm thick, puberulousto hirtellouswith straightanduncinatehairs; internodesshort,occasionally to 1.2 cm long. Leaves in spirals,rosulateto ? spaced;laminanarrowlycordiformto ovateor to subsagittiform,occasionallypeltate, 10-35 x 8-18 cm, chartaceous,entire to pinnatelyor subpalmatelylobedto parted;apex(sub)acuteto acuminate;base cordatewith a wide to narrowsinus, sometimes with overlappinglobes;margin? irregularlyand finely to coarsely crenate-dentate;upper surface glabrous;lower surface (on the main veins densely) minutely puberulouswith straighthairs,intermixedwith sparse to ratherdense longer uncinate hairs; lateral veins 3-9 pairs,in entireleaves usually (faintly) loopconnected;tertiaryvenation subscalariformto reticulate;petiole 8-42 cm long, abaxially,especially in the upper part,minutely puberulous;stipules triangular, 0.3-0.8 cm long, faintlyplurinervate,often ? distinctly carinateandthemarginsinvoluteorrevolute,puberulous with minute straighthairs and slightly longer, often ? retrorseuncinatehairs.Inflorescenceswith the receptacle greenishoutside, flowering face purplishor redbrown;peduncle 9-20 cm long, minutely puberulous with straightto curved, often slightly retrorsehairs; receptacle eccentrically,sometimes almost centrally, attached,discoid,often ? convex, ellipticto suborbicularin outline, 1-2.5 cm long or 0.8-1.5 cm diam., the outside minutely puberulous,the margin entire, ? irregularlylobed, shortly appendiculateor with 10-20, to 3.5 cm long, stipiformto filiforn appendages,the fringe very narrow;bractsin 1-3 rows on the margin, appressedor some of themradiatingand/oron the ap-

192

FLORA NEOTROPICA

Fig. 54. Dorstenia arifolia (from Martius, Fl. Bras. 4(1): t. 59. 1853). Habit (I, II), receptacles (1.31, II.31), margin of receptacle (11.31), staminate flowers (I.1, 1.2, 11.1, II.2), stamens (1.7, 11.7), pistillate flower (I.1. ), pistils (I. 10, 11.10).

TAXONOMIC TREATMENT

193

Fig. 55. Dorsteniaarifolia (fromMartius,Fl. Bras.4(1): t. 60. 1853). Habits,partof receptacle(31), staminate flowers (1), stamens (7), fruit (17), pistils (10).

194 pendages,ovate to oblong, 0.5-1.5 mm long, glabrous; staminateflowers among the pistillate ones; tepals 2; stamens2; filamentslongerthanthe perianth;stigmas equal in length, 0.3-0.5 mm long. Endocarp body ca. 1.5 x 1 mm, distinctlytuberculate. Distribution (see Fig. 79.2). In easternBrazil(from Bahiato Sao Paulo);in moist and shadedplaces, often on sandy soil; to 1800 m. Representative specimens examined. BRAZIL. BAHIA: Without locality, Gillot s.n. (P). ESPiRITO SANTO:Mun. Itaguasu, Jatibocas, 27 May 1972, Brade et al. 18429 (RB, U); Mun. Domingos Martins, Biriracas, 12 Nov 1969, Kautsky 199 (HB); Mun. Domingos Martins, Rio de Jucu Bra9o Norte, Pedra Branca, 27 Jan 1982, Kautsky 724 (GUA); between Arituba and Sao Jose do Calcado, 21 Aug 1982, Rizzini 152 (GUA). MINASGERAIS: Mun. Marlieria, Parque Florestal do Rio D6ce, 18 May 1980, Almeida 187 (RB); Mun. Dionisio, ParqueEstadual Rio D6ce, 29 Sep 1976, Heringer 16615 (K); Juiz de Fora, Krieger 7807 (RB); Praes. Sao Joao Batista, Martius s.n. (M); Tombos, Fazenda Cachoeira, 7 Jul 1955, Mello Barreto 1447 (BHMH, F). RIO DE JANEIRO: Mun. Mangarariba, rd. to Muriqui, 22 Sep 1975, Araujo 800 = Peixoto 606 (RB, U); Mun. Rio de Janeiro, Agulhinha de Copacabana, Morro de Sao Joao, 5 Jan 1963, Carauta 173 (GUA, NY, RB, U); Mun. Itaborai,Serrado Barbosao, 10 Nov 1966, Carauta 346 (RB); Mun. Parati, ParatiMirim, Rio Parati-Mirim, 21 Dec 1976, Carauta et al. 2260 (GUA, U); Mun. Nova Friburgo, Sao Pedro, 18 Oct 1977, Carauta et al. 2707 (GUA, RB); Mun. Barra do Pirai, rd. to Valen9a, km 53, 9 Apr 1980, Carauta et al. 3466 (GUA, RB, U); Mun. Bom Jesus de Itabapoana, Fazenda Sao Jorge, 1 km from Rio Itabapoana, 16 Oct 1982, Carauta et al. 4390 (GUA); Mun. Campos, Morro do Coco, base of Morro do Bau, 13 Dec 1982, Carauta et al. 4420 (GUA); Mun. Maje, Dtto. Guapirim,nr. Centro de Primatologia, 21 Oct 1983, Casari et al. 1103 (GUA); Mun. Itaperuna, Fazenda Sao Jose, 11 Jan 1967, Castellanos 26206 = Strang 917 (F, GUA, HB, RB); Mun. Rio de Janeiro, Rio Comprido, Aug-Sep 1826, Gardner 115 (BM, FI, G, GH, K); Mun. Rio de Janeiro, Floresta de Tijuca, Glaziou 779 (P); Mun. Teres6polis, Cascada do Imbui, Oct 1952, Markgraf 10060 (RB); Mun. Angra dos Reis, rd. to Praia Grande, 29 Sep 1973, Martinelli et al. 67 (NY, R, RB, US); Mun. Petr6polis, Morro do Cuca, 1300-1500 m, 17 Dec 1973, Martinelli et al. 148 (GUA); Mun. Petr6polis, Vale das Videiras, ca. 1800 m, 2 Jul 1975, Martinelli 607 (RB); Mun. Petr6polis, Morro do Cuca, 1500-1700, 10 Oct 1979, Martinelli et al. 6242 (GUA); Mun. Casimiro de Abreu, Corregio da Luz, Pogo Pai Joao, 11 Oct 1982, Pedrosa et al. 611 (RB); Mun. Marica, Itaipu-acu, Morro do Moirao, 18 Feb 1983, Plowman et al. 12856 (F, U), Mun. Niteroi, Morro Alto Moirao, 12 Feb 1985, Plowman et al. 13931 (RB). SAo PAULO: Caraguatatuba, 26 Oct 1974, Carauta 1744 (GUA, K, MICH, SI, SP); Ilha de Sao Sebastiao, 31 Mar 1892, Edwall 1745 (SP); Mun. Ubatuba, Praia do Lam-

FLORANEOTROPICA berto,21 Oct 1981,Peixotoet al. 13062(MBM);between Caraguatatuba and Ubatuba,7 Feb 1968, L. B. Smithet al. 15405 (HB, MICH,US). Dorsteniaarifoliais in its vegetativepartsvery similar to D. ramosa, as D. drakena is to D. contrajerva. Dorstenia arifolia is a variablespecies. The bractscan be appressed,or especiallythe largersubmarginalones can be radiating;they can be sessile or borne on short and i radiatingappendages. Specimens with sessile bractsand those with bractson appendageshave been placedin differentspecies(Carautaetal.,1974a;Carauta, 1978a). Recognitionof distincttaxa appearsnot to be justifiedbecauseof thepresenceof manyintennediates. The distributionof these formssuggests the possibility of geographicalseparationandecologicaldifferentiation, parallelledby the two subspeciesof D. ramosa,but far less clear.Specimens with a shortly appendiculatereceptaclearemorecommonfartherinland(Serrado Mar) and found at higher elevations (400-1800 m) thanthe form with sessile and usually appressedbracts. The peltatelaminaof the type collection of D.fischeri may be regardedas an aberration.The same applies for the lingulatereceptaclesof the type collections of D. maris andD.fi cifolia.

26. Dorstenia ramosa (Desvaux) Carauta,Valente& Sucre,An. XXIII Congr.Nac. Bot. Garanthus158 (1973);Carauta,Rodriguesia29(44): 89, t. 20. 1978; Berg,Proc.Kon.Ned.Akad.Wetensch.,Ser.C, Biol. Med. Sci. 89: 142. 1986. Sychinium ramosum Desvaux,Mem. Soc. Linn.Paris4: 217, t. 12. 1826. DorsteniasychiniumSteudel,Nomencl.ed. 2. 1:526. 1840, as a synonym of Sychinium ramosum. DorsteniamultifornisMiquelvar.ramosa(Desvaux) Bureau,in De Candolle,Prodr.17:261. 1873. Type. Brazil. Without locality, Anonymouss.n. in herb. Desvauxs.n. (holotype,P). Figs. 56, 57 Herb, to ca. 1 m tall, rhizomatous;supraterranean partof the stem to 60 cm long, ascending to procumbent, 4-15 mm thick, puberulouswith straighthairs; intemodesshortor to 5 cm long. Leavesin spirals,rosulate or spaced;lamina(suborbicularto) elliptic to oblong to lanceolate,or ovateto subovate,entireandthen often constrictedin the lowerpartor pinnatelylobedto partedwith 5-3(-1) lobes at each side, 16-35 x 6-30 cm, chartaceous; apex acuminate to subacute; base deeply cordate;margin ? irregularlyand sometimes faintly,finely to coarselycrenate-dentate; uppersurface glabrousor initially sparsely hispidulous; lower surfaceon the(main)veinspuberulouswithminutestraight hairs andusually also with sparseto ratherdense, Tetrorse,uncinateones; lateralveins 8-14 pairs,loop-con-

TAXONOMIC TREATMENT

195

Fig. 56. Dorstenia ramosa (from Martius, Fl. Bras. 4(1): t. 57. 1853). Habit, staminate flowers (11.1, 11.2, II.28), stamens (I.7, II.7), pistillate flowers (II. 2, II.10), stigmas (II.13), fruit and empty excarp (1.18).

196

FLORA NEOTROPICA

Fig. 57. Dorsteniaramosa(fromMartius,Fl. Bras.4(1): t. 58. 1853). Habit,partof receptacle(31), staminate flowers (1,2), stamens(7).

TAXONOMICTREATMENT

197

nected;tertiaryvenationpartlyscalariform in entirelaminas; petiole 17-60 cm long, in the upperpartabaxially puberulous;stipules ovate, 0.5-1.5 cm long, faintly plurinervate, ? conspicuously carinate, the apex subacuminate(in dry material? pungent),the outside ratherdensely,minutelypuberulous,the marginsometimes dentate.Inflorescencespurplish;peduncle9-18 cm long, minutelypuberulous;receptaclefurcate,with the branches3.5-17 cm long and to ca. 0.6 cm broad, often unequalin length, the outside minutelypuberulous with straightand often also with uncinatehairs, the margin with ca. 2 rows of to 2 cm long filiform appendagesof unequallengthandminutelypuberulous, the fringe to ca. 0.5 mm broad;bractslacking,or occasionally presenton the apices of the marginalappendages;staminateflowersamongthepistillateones;tepals 2; stamens2; filaments longer thanthe perianth;stigmas ? unequalin length, 0.1-0.5 mm long. Endocarp body ca. 1.5 x 1 mm, distinctly tuberculate. Dorstenia ramosa can be easily recognized by its peculiarinflorescences,butwithoutthem,it can hardly be told apartfrom D. arifolia. The leaves and the variationof the leaves arestrikingly similarto those of Dorstenia arifolia, which apparentlyled to Miquel's decision (1853) to unite the two species underD. multiformis,a decision accepted by Bureau(1873). Two subspecies can be recognized on the basis of quantitativemorphologicalcharactersanddistribution. The materialdescribedas Dorstenia capricornianais more or less intermediatebut matchessubsp.dolichocaula ratherthan subsp. ramosa. Bracts are normally lacking, but are occasionally presenton the shorterappendages(e.g., in Sucre9991).

Gartenflora5: 87. 1856. Type. Describedfrom materialcultivatedin Hort.Petropolis,collected in Brazilby Riedel (holotype,LE). Dorstenia cerathosanthesLoddiges var. riedeliana Fischerex Regel,IndexSem.Hort.Petrop.(1855): 18. 1855; Regel, Gartenflora5: 87. 1856. Type. Describedfrommaterialcultivatedin Hort.Petropolis,collectedin Brazilby Riedel(holotype,LE). DorsteniaceratosanthesLoddigesformaintegrifolia Wawra,Bot. Ergeb.128. 1866. Type.Brazil.Rio de Janeiro:Corcovado,Wawra& Maly501b (isotype?, LE). Dorstenia multiformisMiquel var. cerathosanthes (Loddiges)Bureau,in De Candolle, Prodr.17: 261. 1873. DorsteniamultiformisMiquel, in Martius,Fl. Bras. 4(1): 165, t. 57, 58. 1853, nom. illeg., pro parte (formaA). DorsteniaficusVellozo,Fl. Flumin.53. 1829("1825"), Ic. 1: t. 138. 1831 ("1827");Carauta,Rodrigu6sia 29(44): 91. t. 21. 1978. Dorstenia vellozoana Lemaire,Ill. Hort. 10: t. 362. 1863, as a synonymof D. ficus. Type.Fl. Flumin.Ic. 1: t. 138. 1831 ("1827"). Supraterranean part of thestemto 30 cm long, (5-) 10-15 mm thick; intemodes short. Lamina to 35 cm long, if incised, then with 5-3(-1) lobes at each side; stipules often to 1.5 cm long. Distribution (see Fig. 79.3). In eastem Brazil(Rio de Janeiro);in humid and shaded places; to 800 m. Representativespecimensexamined.BRAZIL.Rio DE JANEIRO: Mun.Rio de Janeiro,SerraCarioca,Gruta

Geonoma,9 Nov 1967,Carauta484 (GUA,MICH,RB); Mun. Mendes,FazendaSao Jos6 das Paineiras,6 Mar 1980, Carautaet al. 3456 (GUA);Mun.Rio de Janeiro, Corcovado, Gardner 197 (BM, BR, GH, K); Mun. Petr6polis, valley of Rio Bonsucesso, 25 Mar 1971, Key to the subspecies of Dorstenia ramosa Kennedyet al. 897 (RB); Mun. Rio Bonito, Bracana, 1. Stem to 30 cm long; internodesshort....... Fazendadas Cachoeiras,3 Oct 1976, Lachette247 (R); ................................................... 26a. subsp. ramosa Mun. Pirai,Riberaodas Lajes,5 Jul 1983,Peixotoet al. 1. Stem to 60 cm long; internodesto 5 cm 1872 (GUA);Mun. Maj6, Serrada Estrela,Mandioca, subsp. dolichocaula long .26b. Nov 1822, Riedel s.n. (LE, NY, US); Mun. Campo Grande,Macico de Gericino, Serrade Mendanha,27 Nov 1969, Sucre et al. 6380 (= Braga et al. 1770, F, 26a. Dorstenia ramosa (Desvaux) Carauta,Valente GUA, RB); Mun.Nova Iguacu,Tingua,21 May 1980, Vilaca90 (GUA). & Sucre subsp. ramosa. Figs. 56, 57 DorsteniaceratosanthesLoddiges,Bot. Cab. 13: t. 1216. 1827; Bot. Mag. t. 2760. 1827; Schott in 26b. Dorstenia ramosa (Desvaux) Carauta,Valente Endlicher, Cat. Horti Vindob. 1: 241. 1842; & Sucre subsp. dolichocaula (Pilger) C. C. Berg, Fischer& Meyer, Index Sem. Hort.Petrop.11: Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol. 57. 1846. Type. Bot. Cab. 13: t. 1216. 1827. Med. Sci. 89: 148. 1986. Dorstenia Nova Acta fluminensis Walpers,

Phys.-

Med. Caes. Leop.-Carol.Nat. Cur. 19(Suppl.1): 421. 1843.Type.Brazil.Rio de Janeiro:Nr.Rio de s.n. (holotype,B?, not traced). Janeiro,Anonymous Dorstenia ceratosanthes Loddiges var. triloba Regel, Index Sem. Hort. Petrop. (1855): 18. 1855; Regel,

DorsteniadolichocaulaPilger,FeddesRepert.Spec. Nev. RegniVeg. 41: 222. 1937;Carauta,Valente & Sucre, Rodriguesia27(39): 248, t. 9. 1974; Carauta, Rodrigu6sia 29(44):93, t. 23. 1978.Type. Brazil.Rio de Janeiro:Mun. Resende,valley of

198 Rio Preto, Nucleo Colonia Maua, Feb 1931, Kaempfe 366 (holotype, B). Dorstenia ceratosanthes Loddiges var. dissidens Pilger, Feddes Repert. Spec. Nov. Regni Veg. 41: 222. 1937. Type. Brazil. Rio de Janeiro: Mun. Resende, Itatiaia, Maromba, Taqueral, 25 Feb 1936, Brade 15065 (holotype, B, not found; isotypes, ITA, RB). Dorstenia capricorniana Carauta, Valente & Sucre, Ciencia e Cultura28(3): 357. 1976; Carauta,Rodriguesia 29(44): 92, t. 22. 1978. Type. Brazil. Rio de Janeiro: Mun. Rio das Flores, Fazenda Santa Genoveva, 7 Oct 1974, Sucre 7763 (holotype, RB). Supraterranean part of the stem to 60 cm long, 4-

8 mm thick;intemodes to 5 cm long. Laminato 30 cm long, on an average smallerthan in subsp. ramosa, if incised,thenwith (3-)2(-1) lobes at each side;number of lateral veins on an average less than in subsp. ramosa; stipules usually to 1 cm long. Distribution (see Fig. 79.4). In eastem Brazil(Rio de Janeiro, Minas Gerais, and Sao Paulo); in humid and shadedplaces; at 700-1500 m. Representative specimens examined. BRAZIL. MINAsGERAIS:Coronel Pacheco, Fazenda da Liberdade, 12 Feb 1942, Herringer 934 (GUA, SP); Rio Paraibuna, 5 Jul 1970, Heringer s.n. (SP); without locality, SaintHilaire 74 (P). RiO DEJANEIRO: Mun. Resende, Visconde de Maui, Rio Preto, 25 Jul 1971, Carauta 1387 (F, GUA, K, R, RB); Mun. Mendes, Floresta dos IrmaosMaristas,4 Sep 1979, Carauta 3203 (GUA); Mun. Barra do Pirai, Ipiabas, 13 Oct 1986, Carauta 5380 (GUA); Mun. Rio Claro, nr. Lidice, 10 Dec 1974, Gurken 45 (GUA, RB). SAO PAULO:Ubatuba, between Rio Comprido and Rio Cemiterio, 5 Nov 1962, Fontella et al. 84 (US); Barreireos, Fazenda Sao Miguel, 25 Apr 1984, Loefgren et al. s.n. (SP); Serra da Roxaina, 29 Dec 1931, Lutz 736 (US); Barreiro da Bocaina, Mata dos Viveiros, 29 Apr 1926, Hoehne et al. s.n. (GUA, SP).

The collections Gurken45 and Sucre 7763 have features? intermediatebetween the two subspecies.

FLORANEOTROPICA P1.Amer.29, t. 8. 1703 (completedin Plumier,P1. Amer.t. 119.1757). Fig. 58 Dorstenia contrajerva Linnaeus var. houstonii Linnaeus, Sp. P1. 121. 1753; Linnaeus, Hort. Cliff. 32. 1737; Houstoun,Philos. Transact.37: 196, t. 2. 1731; Bureauin De Candolle,Prodr. 17: 259. 1873. Dorstenia houstonii (Linnaeus) Linnaeus, Sp. P1. ed. 2. 176. 1762; Lamarck, Encycl. 2: 317. 1786; Willdenow,Sp. P1. 1(2): 682. 1798; Bot. Mag. t. 2017. 1818; Loddiges, Bot. Cab. 11: t. 1005. 1825. Dorsteniaquadrangularis Stokesvar.integrifoliaStokes,Bot. Mat. Med. 4: 339. 1812, basedon D. houstonii.Type. Mexico.Campeche, Houstouns.n. (holotype,BM). DorsteniaalexiteriaLinnaeus,Syst. Nat. Veg.ed. 10. 889. 1759.Type.Basedon the same specimenas D. houstonii. Dorsteniaquadrangularis Stokesvar.sinuataStokes, Bot. Mat. Med. 4: 339. 1812. Based on Jacquin, Plant.Rar.t. 614. 1790. Dorstenia quadrangularisStokes var.pinnatifida Stokes, Bot. Mat. Med. 4: 341. 1822. Syntypes. St. VincentAndersons.n. (BM)and"WestIndies," Smeathmans.n. (BM). DorsteniapalmataWilldenowex Schultes,Mant.3: 317. 1827, non Engler, 1894; Bureau, in De Candolle, Prodr.17: 259. 1873. Type. Venezuela. "Orinoco,"without locality, Humboldt& Bonplands.n. (holotype, B). Dorstenia maculata Lemaire, Ill. Hort. 10: t. 362. 1863. Dorstenia contrajerva Linnaeus var. macu-

lata (Lemaire)Bureau, in De Candolle. Prodr. 17: 260. 1873. Type.Ill. Hort. 10: t. 362, drawn and describedfrom living materialcollected by Ghiesbreghtin Mexico. Dorstenia contrajerva Linnaeus subsp. tenuiloba Blake, Contr.U.S. Natl. Herb.24: 2, t. 1. 1922. Dorstenia contrajervaLinnaeusvar. tenuiloba (Blake)Standley& Steyermark, Publ.Field Mus. Nat. Hist., Bot. Ser. 23: 44. 1944. Type. Guatemala.Izabal:Sierrade Las Minas,betweenIzabal and Los Amates,31 May 1914, Blake 7803 (holotype, US).

Herb, (sub)acaulescent;stem at least partlysubterranean,3-14 mm thick,puberulousto subglabrous;in27. Dorstenia contrajervaLinnaeus,Sp. P1.121. 1753; temodes usually short, sometimes to 2.5 cm long, in Houstoun,Philos.Trans.37: 196,t. 2. 1731;Linnaeus, slenderrhizomatousstems with all or some of the inHort. Cliff. 32. 1737; Plumier,P1.Amer. 109, 119. temodes long, the partswith shortinternodes? tuber1757; Linnaeus,Syst. Nat. Veg. ed. 10. 899. 1759; ous. Leaves in spirals, rosulate or spaced; lamina Jacquin,P1.Rar. t. 614. 1790; Willdenow,Sp. P1. broadlyovate to cordiformto subhastate,pinnatelyto 1(2):683. 1798:Grisebach,Fl. Brit.W. I. 154. 1864; subpalmatelyor subpedately,variouslylobed to parted Bureau,in De Candolle,Prodr.17: 259. 1873; Ur- with 3-8 lobes at each side or subentire,4-25 x 3.5ban, Symb.Antill. 4: 196. 1905; Burger,Fl. Costa 30 cm, chartaceous,sometimes variegated;apex (or Rica, FieldianaBot. 40: 138, t. 17. 1977; Carauta, apices)acuteto subacuminate;base cordateto acuteor Rodrigu6sia29(44):103,t. 39.1978. Dorsteniaquad- often the laminadecurrent;margincrenate-dentateto rangularisStokes, Bot. Mat. Med. 4: 338. 1812, as subentire; upper surface hirtellous to subhirsute or a synonym of D. contrajerva.Lectotype,Berg, Fl. strig(ill)ose,usually scabrous;lower surfacemainlyon Guianas,Ser.A, Fasc. 11: 34. 1992. Plumier,Nov. the veins puberulouswith patent to appresssed,unci-

TAXONOMICTREATMENT

199

-4.

Fig. 58 Dorstena contrajerva 1 Habit (Contreras 1299). 2. Leaf (Lundel 885) 3. nflorescence (Tuerckheim 310). (By E. M. Hupkcns van der Elst, Utrccht.]

200

FLORA NEOTROPICA

nate,orstraight hairsoralsohispidulous, oftenscabridu- Pochutla,11 Apr 1917, Conzattiet al. 3024 (US); Chilous to scabrous;lateralveins 3-7 pairs,furcately nantla1000m, 1840,Galeotti292 (BR, P);nr.Chiltepec, in themargin;tertiaryvenation 6 km frombridge over Rio Papaloapan,12 Oct 1979, branched, terminating subscalariform to reticulate;petiole 3-20 cm long, Graham1177 and 1178 (MICH); 8 km N of Matias puberulous with? retrorse uncinateandstraighthairs, Romero,nr. La Princesa, 17 Oct 1970, Graham1197 oralsohispidulous; to ovate,to 0.6 (MICH);35 km N of Pochutla,rd. to PuertoAngel, 21 stipulestriangular Oct 1970, Graham1209 and 1210 (MICH);5 km N of cm long, puberulousto hispidulous.Inflorescences MatiasRomero,27 Jul 1958, King 815 (MICH).QUINgreenorwiththemarginpurplish(-red); peduncle5- TANARoo: 8 km N of Uni6n, 110 km SW of Chetumal, 30cmlong,denselyto sparsely puberulous, mainlywith 7 May 1982, Davidse et al. 20154 (U); 40 km SW of ? retrorse uncinatehairs,oralsohispidulous, theapex Chetumal,rd. to Union, 7 May 1982, Davidse et al. broadened intothereceptacle; receptacle eccentrically 20200 (U); 9 km S of Hwy. 186, on Hwy. to ThomAs attached,quadrangular to (orand)? irregularly lobed Garrodo,San Jose de la Montania,9 May 1982,Davidse in outline,(0.5-)1.5-3 cmdiam.,theoutsidepuberu- et al. 20286 (U); 26 km W of Tulum, 19 May 1982, lous,mainlywithretrorse uncinate hairs,oralsohispidu- Davidse et al. 20627 (U); Lake Chichankanab, Gaumer lous,thefiinge0.5-2 mmbroad,minutelypuberulous; 1923(B, BR,F,GH,MO,NY,P,US). SONORA: RioMayo, bractsin ca. 3 rows,on the margin,on the fringe,or CanyonSapopa,14 Oct 1934,H. S. Gentry1026 (F, GH, MO).TABASCO: Mun.Tacotalpa,nr. Tapijulapa,30 Sep alsoa fewbelowthemargin,appressed, semiorbicular 1986, Hammelet al. 15536 (BG); Balancan,La Palma, to ovate,to 1(-1.5) mmlong,puberulousto hispidu- 1 Jun1939,Matuda3276 (F,MICH,NA,NY).VERACRUZ: flowersamongthepistillateones;tepals Nr. Catemaco,21 May 1956, Calzada56 (GH),22 Mar lous;staminate 2; withglobose,conicaland/orminuteuncinatehairs, 1973, Cedilloet al. 138 (BR);Cant6nCordoba,Colonia stamens2;filamentsaslongasor(slightly)longerthan MelchorOcampo,21 Jun 1891, Conzatti175 and 176 theperianth; stigmas(almost)equalin length,0.3-0.5 (GH);Mun.Teocolo,TexoloRavine,1000m, 1 Apr 1974, Dorantes2665 (MICH);nr. San AndresTuxtla,7 Aug 1953,Dressleret al. 12 (BM, MICH,MO, NY, US); 10 km S of Catemaco,Hwy. 180, 14 Oct 1970, Graham Distribution(see Fig. 81.8).FromMexicoto the 1189 (MICH);Zacuapan,Sep 1905,Purpus2806 (F, GH, WestIndiesandthroughCentral Americato Colombia MO,NY, US); Misantla,Jul 1912,Purpus5951 (F, GH, and northernVenezuela (possibly extending to MO, NY, US); Bailos de Carrizal,Aug 1912, Purpus

mm long, white.Endocarpbodyca. 1.8-2 x 1.8-2 mm,

tuberculate.

Chichen Itza, 27Guyana),also in southwestemEcuadorandin Peru 6110 (F, GH, MO, NY, US). YUCATAN: in shade,usuallyin ? wet,sometimesin 28 Feb 1899,Armour1631 (F);Izamal,Jan1917,Gaumer (SanMartin); et al. 23805 (F, GH, NY, US); Chichen Itza, Steere 1273 drytypesof forestor scrub,ofteninrockyplaces,of(BM, F, MICH). tenon limestone,alongwatercourses,alsoin secondGUATEMALA. ALTAVERAPAZ: Chanabal, Apr1889, arygrowth;to ca. 1500m. DonnellSmith1619 (GH,NY, US); Sierrade las Minas,

Representative specimens examined: MEXICO. Jalapa, 1839, Linden37 (G, GH, MICH); 30 Oct 1931,Lundell884 (F, GH,MICH,MO, Tuxpenia, US); Champoton,7-15 Jul 1932, Steere 1882 (MICH). CHIAPAS: 13 kmN of Berriozabal, 9 Oct 1971,Breedlove 20288 (MICH); Laguna Ocotal Grande, 45 km E of Ocosingo, 2 Aug 1954, Dressler 1569 (GH, US); Esquintla,Mt. Ovando,16 Jul 1940,Matuda4216 (GH, MICH,NY); Acacoyagua,4 May 1948, Matuda17778 (F, NY); 1 km S of SanMiguel,13 Jul 1973,McPherson 741 (MICH).CHIHUAHUA: Nr. Batopilas,9 Oct 1898, Goldman238 (GH,NY). HIDALGO: S of Tumazunchale, 17 Jun 1935, Clark6883 (MO,NY). JALISCO: N of La Cuesta, rd. to Talpa de Allende, 15-16 Oct 1960, McVaugh20261 (MICH),20-21 Nov 1960, McVaugh 21205 (MICH).PUEBLA: Mun. Jalpan,El Jingibre, 1 Nov 1966, CruzCisneros1481 (MICH);Las Cruzes,27 Aug 1970, Hernindezet al. 709 (MICH);Ayotoxco,29 Apr 1970, Ventura995 (MICH,MO, US). SAN Luis POTOSI: Hwy. 85, km 15 fromborderwith Hidalgo,20 Nov 1971,Dieterle4234 (MICH);nr.Tamazunchale, 28 Nov 1937, Kenoyers.n. (MO); Mun. Tamanzunchale, Taman,28 Jun 1959, Rzedowski 10997 (MICH);nr. Xilitla, 23 Sep 1954, Sohns 1444 (US). OAXACA: CAMPECHE:

Teleman,Finca Mercedes, 14 Jul 1968, MartinezS. et al. 22754 (BG);TreceAguas,31 Mar 1906,Lewton312 (US); nr. Pancajehe,5 Apr 1939, Standley 70798 (F); Coban, Mar 1908, Tuerckheim 11.2013 (BR, F, GH,

MICH,MO, NY, US, Z). CHIMALTENANGO: Sibaya, 1050 m, 6 Jan 1939, Standley 62280 (F); VolcAnde Fuego, 1200-1600 m, 20 Sep 1942, Standley 52118 (F, US). CHIQUIMULA:VolcanIpala,nr.Amatillo,25 Oct 1939,Steyermark30514 (F). ESQUINTLA: Nr. Esquintla, Mar 1890, DonnellSmith2027 (GH, US); Los Diamantes, Rio Cucunya,8 May 1896, Seler 2512 (GH, NY, US); Rio Guacalate,28 Nov 1938, Standley58244 (F); Volcande Fuego,FincaMonterrey,5 Feb 1939,Standley 64564 (F). GUATEMALA: Finca La Aurora, 1500 m, 1938-1939, Aguilar 337 (F); 12 mi NE of Guatemala City, 1100 m, 23 Jul 1977, Croat41902 (BG); 10 km S of SanRaimundo,1800m, 18 Jan 1938,Standley62861 (F). HUEHUETENANGO:Sierrade los Cuchumatanes, Rio Amelco, Finca San Rafael, 27 Jul 1942, Steyermark 49681 (F, NY); betweenDemocraciaand CanyonChamushu,1000-1300 m, 24 Aug 1942, Steyermark51075 (F);Rio Trapichillo,Pasodel Boquer6n,below La Libertad, 21 Aug 1942, Steyermark51135 (F); Sierrade los between SantaAna Huistaand Rancho Cuchumatanes,

TAXONOMIC TREATMENT

51354 (F,MO).IzABAL: Lucas,26 Aug 1942,Steyermark Cadenas(PuertoMendez),RioGraciosa Dios,16 Sep1969, Contreras9191 (U); 4 mi SW of PuertoBarrios,22 Jul 1977,Croat41833 (BG);nr.Quirigua,15-31 May 1922, Standley24552 (F); nr. Escobas, 2 Jan 1922, Standley 24846 (F); Rio Juyama,SE of Cheyenne,8 Apr 1940, Nr. Jalapa,7-18 Nov Steyermark39125 (F). JALAPA: 1940, Standley77524 (F). PETtN:Nr. La Libertad,11 Jan 1934,Aguilar250 (F, MICH,MO,NY); Vaxactum, 19-22 Mar 1931, Bartlett 12199 (MICH,NY, S, US); Lake Peten, Tayasal, Jun 1933, Lundell 4256 (F, S); Ceibal, 18 Nov 1965, MolinaR. 15837 (F, GH);Cerro Ceibal,betweenmouthof Rio SantaMonicaandmouth of Rio San Martin,30 Apr 1942, Steyermark46120 (F, San Francisco Miramar, Apr US). QUEZALTENANGO: 1905, Pittier 87 (US), 15 Mar 1921, Rojas et al. 151 (MO,US); Colombia,ca. 1000 m, 31 Dec 1934, Skutch 2050 (F, GH,NY, US); VolcanSantaMaria,1300-1500 San m, 8 Jan 1940 Steyermark33768 (F). RETALHULEU:

Martin,Zopolitlan,14 May 1921, Rojas 571 (US); Rio Coyote, 4 km W of Retalhuleu,17 Feb 1941, Standley 87412 (F); Rio Ocosito,W of Retalhuleu,24 Feb 1941, Standley88275 (F); PuebloNeuvo, Mar 1923, Stricker Rd. Antigua-Esquintla, 2.3 343 (US). SACATEP#QUEZ:

mi SW of Alotenango,1300, 26 Jul 1977, Croat42031 (BG);Las Lajas,1200 m, 28 Nov 1938,Standley58131 (F); BarrancaHondo, 1200 m, 11 Nov 1941, Standley 88491 (F). SAN MARCOS:Rio Mopa, below Rodeo, 14

RoSA:Nr. Santa Mar 1949, Standley68781 (F). SANTA Rosa, ca. 1000 m, Jun 1892, E. J. Heyde et al. 2962 (GH,MICH,MO, NY, US); Azagualpa,Oct 1937, E. J. Heyde et al. 6236 (G, GH, MICH,NY, US), 6237 (F, GH,MICH,MO,NY, US); nr. Cuilapilla,23 Nov 1940, Standley78059 (F); S of Guazacapan,Avellenard., 29 Nov-6 Dec 1940, Standley 78606 (F). SOLALA:Santa Barbara,Aug 1891, Shannon 97 (US). SUCHITEPEQUEZ:

Mun. Santa Barbara,FazendaSantaAdelaida, 26 Jul 1972, Lind 10 (WIS);Patutul,Rio MadreVieja, 6 Jan 1939, Standley 62208 (F). BELIZE.Cayo,15 Feb 1931,Bartlett11500(MICH); CorozalDistrict, 1931-1932, Gentle220 (MICH,NY, rd., 10 S, US); ToledoDistrict,ColumbiaR., Chavarrias Oct 1947,Gentle6275 (F,MICH,NY,S, US);CayoDistr., Fairchild,8 Jul 1970, Gillis 9651 (S); Xunantunich,17 Jul 1960, Hedger 20 (F); Honey Camp, 18 Sep 1929, Lundell499 (MO);Cayo, Mar 1933, Lundell4423 (F, MO);Augustine,26 Apr 1969, Proctor30205 (IJ);Rio Grande,Perrett'slanding,7 Jul 1933, Schipp1188 (F, GH, MICH,MO,NY, S, Z); Rio Temash,13 Jan 1935, Schipp 1332 (F, GH, MICH,MO, NY, S, Z). EL SALVADOR.Colina de SantaTecla, Jul 1923, Calder6n1721 (GH, US); Dtto. Sonsonate,Balneario de Atecozol, 20 May 1963, MolinaR. et al. 12469 (F, nr.Ahuachapan, 9-27 Jul 1922, NY); Dtto.Ahuachapan, Standley19784 (GH,S, US); Dtto.SanVincente,nr.San Vincente,2-11 Mar 1922, Standley21393 (US); Dtto. Sonsonate, nr. Santa Emilia, 22 Mar 1922, Standley 22252 (GH, MO, S, US); Dtto. Mirazan, Sierra de Hacaguatiga,FincaGeneralJ. T. Calder6n,2 Jan 1942, Tucker705 (F, MICH,NY, US).

201

HONDURAS. ATLANTIDA: Puente Alto, E of La Ceiba, 19 Jul 1938, Yunckeret al. 8550 (GH, MO, NY, US). COMAYAGUA: CerroAzul de Meambar,12 Aug 1974,Hazlett1968 (CR);6 kmW of Sequatepeque,1200 m, 8 Aug 1936, Yuncker et al. 6382 (F, GH, MO, U, US). COPAN:La Florida, Hda. Espirito Santo, 14 May 1919, Blake 7428A (US); Copan (ruins), 22 Aug 1971, Molina R. 26246 (BM), 9 Jan 1907, Pittier 1837 (US); El Guabo, between Espirito Santo and Las Playitas, 1018 May 1919, Pittier 8531 (US). CORTES:Sierra de Omoa, La Cumbre, 30 Nov 1950, Molina R. 3453 (F, GH); Rio Ulua, nr. Pimienta, 27 Aug 1955, Molina R. 5625 (F); Ocote Arrancado, 50 km N of Lago de Yajoa, 1-30 Nov 1980, Nelson et al. 6019 (BG). GRACIASA Dios: Without locality, Hjalmarson s.n. (S). MORAZAN: Aldea de Suyupa, NW of Tegucigalpa, 1100, 10 Aug 1949, Molina R. 2578 (F, GH). LEMPIRA:Banios de Lepaera, 1000 m, 26 Sep 1963, Molina R. 12977 (F, NY). OLANCHO:Culmi, 17-22 Jul 1978, Nelson et al. 4632 (U). SANTABARBARA:Los Dragos, Rio Chamelec6n, SW of Quimistar, 16-17 Aug 1947, Standley et al. 7331 (F); San Pedro Sula, Mar 1889, Thieme 5492 (GH, US); El Mochito, 23-26 Jul 1976, E. Vargas 737 (MO). YORO:Subirana, ca. 1000 m, Sep 1937, Hagen et al. 1056 (NY). NICARAGUA. ESTELf:La Guayabita, 1360-1400 m, 17 Sep 1982, Moreno 17530 (MO); El Chayote, 11001200 m, 31 Jul 1983, Moreno 21743 (MO). JINOTEGA: 3.8 mi SSE of Yali, between Condega and Jinotega, 6 Aug 1977, Croat 42842 (MO); rd. from Hwy. 3 through La Fundadora,between las Camelias and La Salvadora, 1100-1 150 m, 31 Oct 1979, Stevens et al. 15286 (BG); Rio Bocuy, Salto Kayaska, Stevens et al. 16597 (MO, U); nr. Jinotega, 1030-1300 m, 19 Jun-9 Jul 1947, Standley 9697 (F); E of Jinotega, 1300-1500 m, 5 Jul 1947, Standley 10865 (F). MADRIZ:Ca. 5 km SW of San Juan de Rio Coco, rd. to Telpaneca, 1080 m, 29 Jun 1980, Stevens et al. 17668 (MO). MANAGUA:Sierra de Managua, 20 Aug 1977, Stevens 3453 (MO); Hwy. 8, ca. 2.4 km SW of intersection with Hwy. 2, km 28, 18 Sep 1977, Stevens 3992 (BG). MASAYA:Sierra de Managua, Las Nubes, 24 May 1947, Standley 8736 (F). 0.7 km W of Hwy. 3 on rd. to San Sim6n MATAGALPA: de Palcila, 1200 m, 30 Nov 1980, Stevens et al. 18566 (MO). ZELAYA:Mun. Siuna, Uli Arriba, 9 Aug 1983, E Ortiz 1407 (MO). COSTA RICA. ALAJUELA:Cerro de San Isidro, nr. San Ramon, 6 Jan 1901, Brenes 14391 (GH); Naranjo de Alajuela, 4 Sep 1932, Brenes s.n. (NY);Rio Barranca, 19 Aug 1973, Lent 3603 (CR). CARTAGO:Rio Agua Caliente del Llano, 11 May 1906, Brenes s.n. (NY); nr. Turrialba, 23 Jun 1949, Holm et al. 103 (GH), 1 Aug 1965, Lent 692 (MICH, NY); Las Concavas, SW of Dulce Nombre, ca. 1500 m, 13 Jun 1928, Stork et al. 8 km N of Bagaces, 2531 (MICH, NY). GUANACASTE: 14 Jul 1964, Croat 493 (MO); Filadelfia, 12-14 Jun 1984, Gomez et al. 23050 (BG); between Cafias and Liberia, 14 Jul 1905, Walker148 (GH). HEREDIA:Puerto Viejo, Rio Sarapiqui,Aug 1891, Pittier et al. 6908 (BR, CR). LIM6N: Reserva Biol6gica Hitoy Cerere, 29 Aug

FLORA NEOTROPICA

202

Monte1985, Gomezet al. 23661 (BG). PUNTARENAS: verde, 1500-1550 m, 10 Nov 1984, Haber 1054 (BG); San Luis valley, 1100-1200 m, 29 Sep 1985, Haber et al. 2948 (BG); Cascajal,3 Jul 1949, Holm et al. 229 (GH, NY, P, U); 21 km SE of Puntarenas,Rio Barranca, 20 Aug 1938, Stork et al. 8913 (NA). SAN Jost: Nr. Alajuelita,1000 m, Aug 1894, Pittier et al. 8855 (BR, CR, MICH);El General,1050 m, Jul 1936,Skutch2750 (GH,MICH,NY,S); betweenSanPedroandSanRam6n, 1300 m, 24 May 1913, Tonduz17880 (CR). PANAMA.BOCASDEL TORO:ChiriquiLagoon, 5 Nov 1940, Wedel1530 (GH). CHIRIQUI:CerroColorado,5-8 km fromEscopeta,17 Aug 1977,Folsom4921 (U). COCLi: Vallede Chiquito,25 Jul 1935,Seibert502 (GH, MO, NY). COL6N: Rio Boquer6n, 5 Sep 1977, Berg et al. 414 (BG, U); 18 km S of Colon, Rio Providencia,25 May 1966,Blumet al. 2333 (MO);nr.Achiote, 31 Oct 1967, Correaet al. 424 (MO); 1.5-2 km N of Gamboa,19 Jan 1974,Nee 9508 (IJ, LD);Las Cruces,1 Aug 1935, Seibert 574 (GH, MO, NY). DARItN: Rio Tuqueza,between QuebradaVenadoand Peje swamp, 28 Jan 1967, Bristan 1002 (MO);2-6 km N of Pucro, 22 Jan 1967,Duke13045 (MO);PijiVasal,24 Jul 1977, Folsom4577 (U); Rio Paya,betweenPayaand Boca del Paya, 12 Jun 1959,Sternet al. 272 (GH,LE, MO,US); Rio Chucunaque,above confluencewith Rio Tuqueza, 4 Jul 1959, Stern et al. 914 (GH, LE, MO, US). Los aboveEl Cortezo,25 Oct 1978, SANTOS:Rio Guanquito, Hammel5316 (MO). PANAMA: Between El Llano and Rio Mamoni,15 Sep 1962, Duke 5633 (GH, MO);jct. of Rio-Corsoand Rio Pacora,9 Jun 1967, Duke 11913 (MO) = Oliver 46 (K); nr. TortiArriba,31 Aug 1977, Folsom 5130 (MO, U); Juan Diaz, nr. Rio Tapia, 1-2 Jan 1923, Maxon et al. 6716 (GH, NY, US); Ancon Hospital,14 Feb 1911,Pittier 2748 (GH,NY, US); Isla

COLOMBIA. ANTIOQUIA: Mun. Turbo, Cgto. Curruloa,km 19, 2 Aug 1987, Callejaset al. 4959 (BG, MO); between Turboand Chigorod,Rio Vijagual,25 Mar 1962, Feddema2014 (MICH,NY); nr. Turbo,7 Apr 1945, Haught4548 (US); nr. Nicoli, 23 Jun 1946, Haught4901 (F, MO,US); Dabeiba,Apr 1889,Lehmann 4738 (F, K, US). BOLIVAR: Cartagena, 1919, Bro. Heriberto 7 (US); Turbaco, 6-22 Nov 1926, Killip et al.

14297 (LE, NY, US) and 14355 (F, GH, NY, S, US); Quimari,22 Mar 1922, Sneidern5712 (GH, MO, NY, US). CALDAS: Mun.La Dorada,Cgto. Norcasia,Rio La Miel, 7 Nov 1989, Estradaet al. 582 (COL).CHOc6: Mun.Acandi,Cgto. San Francisco,VeredaCoquital,22 May 1989, Betancur et al. 1192 (MO); Serrania del Darien, W of Unguia, 17 Jul 1975, Gentry et al. 15229

(U), Mun.Acandi,Vrda.Paramo,22 May 1989,Roldan et al. 1189 (COL,MO). MADGALENA: Nr. La Jagua,40 km NE of Chiriguana, 21 Aug 1938,Haught2274 (US), 30 Aug 1943, Haught 3644 (US); nr. Santa Marta, Bonda, 14 Nov, H. H. Smith 1444 (BM, F, GH, LE, MICH,MO, NY, P, S, U, US, WIS); nr. Santa Marta, Calavasa,2 Jan, H. H. Smith 1928 (F, K, MICH,MO, NY,P, S, U, US);Rio Frio,22-30 Jun 1906,Pittier1591 (NY, US); nr. Santa Marta,Minca, ca. 1000 m, May 1852, Schlim 916 (= Linden 916) (BM, BR, G, K, P).

CordilleraMacarena,Guapayita,between Rio Guejarand CainoGuapayita,20-28 Dec 1959, Idrobo et al. 912 (A, US); nr. Villavicencio, Llanos de San Martin,1846, Karstens.n. (BM, G, JE, LE);Cordillera Macarena,Plaza Bonita, 14 Nov 1949, Philipsonet al. 1416 (BM); CordillleraMacarena,Rio Guejar,10 km below confluencewith Rio Zanza,21 Aug 1950, S. G. Smithet al. 1531 (GH, MO, NY, US), nr.Villavicencio, Llanode SanMartin,Triana850 (K, P), Jan1856,Triana 1860 (BM). SANTANDER: Rio Surata,betweenBucaraTaboga, 23-24 Jul 1938, Woodson et al. 1505 (GH, manga and El Jaboncillo, 800-1500 m, 2 Jan 1927, MICH,MO,NA, NY). SAN BLAS:Ailigandiarea,7 Oct Killip et al. 16359 (F, GH, NY, US). TOLIMA: Between 1978, Hammelet al. 4993 (MO, U); Cangandi,27 Mar Hondaand Guaduas,1843, Hartwegs.n. (K, LD). 1986, Nevers et al. 7621 (BG). VENEZUELA. ARAGUA: ColoniaTovar,May 1854, PUERTORICO. Anasco, 12 Oct 1913, Hess et al. Fendler1286 (BR, GH, K, MO, P, US); La Victoria,9 3573 (NY); Quebradillas, 19 Jun 1937, Sargent A-77 Jun 1929, Holt 390 (VEN); La Trinidadde Maracay, (US);Cacey,MorillosR., 5 Oct 1885,Sintenis2294 (BM, Jan-Feb 1913, Pittier 5817 (US), Vogl 785 and 786 (M); G, GOET,K, LE, NY, US); Cacey, "PedroAvila,"22 Mun.Aut6nomoMora, E of Mor6n,,20-21 Jul 1991, Sep 1885,Sintenis2448 (B, BM, GH,JE, LD, MO,NY, W Diaz et al. 509 (BG). DISTRITO FEDERAL: Las QueP, S, US); Bayamon,1887, Stahl 702 (S). seras, nr. Caracas,Nov 1924, Allart 122 (G, NY, US, ST.MAlRTIN/MAARTEN. Withoutlocality,Bernhardi VEN); Rio La Guayara,Birschel s.n. (K); Puertode la 339 (JE);Mt. Paradis,12 Sep 1906, Boldingh2362 (U), Cruz, 1928, Holt 121 and 122 (VEN); Caracas,Jardin 12 Sep 1906, Boldingh 3344 (NY) and 3344B (NY, U). Botanico,24 Jun 1978,Liesner5317 (MO);La Guayara, GUADELOUPE. Mt. St. Louis, Schwartz s.n. (S). Sep, Moritz559 (BM, K, LE). FALC6N: Sierrade San MARTINIQUE. Without locality, 1857, Belanger Luis, Pedrade Agua, 20 Jun 1979, Liesneret al. 7638 261 (P); Marin, Morne Gommier, Duss 1403 (F, MO, (MO);ParqueNacionalQuebradade la CuevaEl Toro, NY, US), Duss 4130 (NY); St. Pierre, 1869, Hahn 856 (BM, G, K, P, S); without locality, 1833, Martin 339 (BM, M), 1841, Plie 782 (P), 1822, Sieber s.n. (GOET). GRENADA. St. Georges, 27 Jun 1904, Broadway s.n. (BM); Snug Corner, 24 Mar 1905, Broadway s.n. (F, NY); without locality, May 1908, Buysman 2540 (U).

META:

21-22 Jun 1979, Liesner et al. 7695, 7844, and 7924

(MO);Sierrade San Luis, betweenCurimaguaand San Luis, Puentede Jobo, 20 Jul 1967, Steyermark99270 (VEN);Dtto. Silva, CerroChiriviche,betweenLizardo and Mallorquines, 4 Sep 1974, Steyermark et al. 110732

(VEN).LARA: Dtto.Moran,4.5 km SWof Guarico,1350 ST. VINCENT.Kings Hill, 31 Jul 1937, Sandwith m, 9 Jun 1980, Davidse et al. 18145 (U); Dtto. Moran, 981 (K); nr. Owia, Sep 1889, H. H. Smith et al. 476 Rio Tocuyo,S of MumocaroAlto, 1300 m, 13 Oct 1974, (BM, GH, K, NY). Steyermarket al. 111061 (VEN). MeRIDA: BetweenSan

TAXONOMICTREATMENT IsidroAlto and SantaCruz de Mora,760-1800 m, 15 May 1944, Steyermark56568 (VEN). MIRANDA: Los Mostazas,on railroadbeyond Los Teques,Nov 1924, Allart 194 (G, NY, US) and 321 (VEN); Guatue,Hda. El Bautismo,1200-1500 m, 29 May 1944,Lasser 1057 (VEN); Petare-Guarenas,20 Aug 1939, LI. Williams 11181 (F). PORTUGUESA: 28-32 km NNE of Guanare, Cerro et al. 127056(U). SUCRE: 2 Nov 1982,Steyermark 6 May 1945,Steyermark Rio Manzanares, Turumuquire, 62684A (F, VEN); Cerro Turumuquire,between Rio Manzanaresand La Trinidad, 800-1300 m, 12 May 1945, Steyermark62768 (F, VEN). TRUJILLO: San Jacinto,27 Aug 1941, Tamayo1701 (VEN).YARACAY: San Felipe, 9 Apr 1946, Burkart16928 (VEN); Sierre de Aroa, 1000 m, Caminoal Chimbaroso,15 Jan 1953, Vareschiet al. 2727 and 2728 (M). ZULIA: rd. Represa Guasare,2 km beforePuerto ManuelOte-Destacamento Delicias,9 May 1984,Berget al. 1498 (U);Dtto.Bolivar, rd. Lara-Zulia, km 70, SE of Puente, 29 Apr 1979, 3.5 km Buntinget al. 7667 (U); rd. Machiques-Sirapta, fromSirapta,23 Nov 1977,Jeffreyet al. 2273 (K);Sierra de Perija,Rio Macoita,Jan 1948, Lasser2570 (VEN). TRINIDAD.Dibe valley, 15 Apr 1920,Brittonet al. 5 Apr 1921, Britton 1759 (GH,NY, US); Chacachacare, et al. 2762 (NY, US); St. Anns, cascade, 24 Oct 1924, Broadway5452 (MO,S, US);DiegoMartin,14 Jan1932, Broadway7882 (MO,S); 2-2.5 km N of St. Josephand Tunapuna, 12 Jun 1975, Raynal 15669 (K, TRIN); Blanchisseuse,3 Oct 1967, Wong87a (TRIN). TOBAGO.Withoutlocality, Anonymous(herb.Ph. Miller)s.n. (BM). ECUADOR.GUAYAS:Nr. Guayaquil,26 Jan 1955,

203 Loddiges, Bot. Cab. 7: t. 677. 1822; Bureau,in De Candolle, Prodr.17: 260. 1873; Burger,Fl. Costa Rica, FieldianaBot.40: 139, t. 17. 1977. Dorstenia ovalis Stokes, Bot. Mat. Med. 4: 338. 1812, as a synonym of D. drakena.Type. Mexico. Veracruz: Veracruz, Houstoun s.n. (holotype, BM). Figs. 59, 60 Dorstenia mexicana Bentham, P1. Hartweg. 51. 1840; Bureau,in De Candolle. Prodr.17: 260. 1873. Type.Mexico. Michoacan:Morelia,1838, Hartweg386 (holotype,K; isotypes, LD, LE). DorsteniacrispataS. Watson,Proc.Amer.Acad.22: 452. 1887. Type. Mexico. Jalisco: Rio Blanco, Jun-Oct 1886, Palmer 39 (holotype, GH; isotypes,G, K, NY, P, US).

Herb, (sub)acaulescent;stem at least partlysubterranean,minutelypuberulousto subglabrous;internodes short.Leaves in spirals,rosulate;laminaovate, sometimes to suborbicular,reniform or subhastate-subsagittate, (2.5-)8-22(-30) x (2.5-)4-22(-30) cm, (2 x 2-4) pinnatilobedto -parted,with2-4 lobes ateachside, sometimes the incisions down to the midrib,the midsegmentoftenlobed,sometimesthelaminaonly 3-lobed or almostentire,chartaceous;apices of the laminaand its segments acuminateto rounded or to acute; base cordate to subcordate,or sometimes to truncate,the laminadecurrentalongtheupperpartof thepetiolewith narrowor lobate wings; margin? irregularlycrenatedentate(to subentire);uppersurfacescabrousto scabriAsplund 15239 (B, G, LD, NY, S), Ruiz & Pav6n s.n. dulous, sparselypuberulousto hirtellousor to strigil(BM, G, MA, P); CerroAzul, N of Chongon, 18 Mar lose; lower surface scabridulous (to smooth), with 1980, Dodson et al. 9668 (SELBY,U). MANABI: With? retrorse,uncinatehairsandlongerstraightor minute, out locality, 1897, Eggerss.n. (P); Guale, 6 Feb 1940, often confined to the veins; lateralveins hairs, curved Haught3064 (F, GH, US). PERU. SAN MARTIN: Rio Huallaga, Juanjui, 9 Jun 4-8 pairs, furcately branchedand terminatingin the 1936, KIug 4390 (F, GH, MO, NY, S, U, US); rd. margin;tertiaryvenation subscalariformto reticulate; km 25, 6 Oct 1984, Maaset al. 5964 petiole (1.5-) 4-16(-24) cm long, 1-2.5 mm thick, Tarapoto-Juanjui, Jun1855,Spruce3921 (BR, retrorselypuberulousto hirtellous;stipulesovate, 0.2(BG,MOL,U); nr.Tarapoto, G, GH,GOET,K, LD, LE,MPU,NY,P, S); nr.Tarapoto, 0.8 cm long, minutelypuberulous.Inflorescenceswith Ule 6509 (K), 10 Dec 1929, LI. Williams5905 (F); the flowering face entirely green, green with a red or Dec 1929,Ll. Williams6832 (F). Rumizapa,nr.Tarapoto, purplefringe,or entirelyred or (dark)purple;peduncle The species is very variable in the dimensions of (2-)4-24 cm long, puberulouswith retrorseuncinate the wholeplant,leaves,andinflorescences,in the length hairsto hirtellouswithstraightto curved,patentto someof the internodes,in the shapeof the receptacle,andin whatretrorsehairs;receptacleeccentricallyattached, the shape of the leaves, these varying from entire to discoid, flat, obovate to elliptic or to suborbicularin variouslyincised. It was not possible to relatesome of outline, (0.5-)1-3.5(-5.5) cm long, 0.5-3(-3.5) cm thevariationclearlyto certainhabitatsor regions.Some broad,the outsideminutelypuberulousto sparselyhircollections placed in D. drakenashow featurestransi- tellous, the marginentireor sometimes slightly lobed tional to those of D. contrajerva. or crenate,the fringe 0.5-1.5(-2) mm broad;bractsin 2-3 rows, I row on the margin,1-2 rows on the fringe, appressed,ovate to elliptic, 0.1-1.2 mm long, puberu28. Dorstenia drakena Linnaeus,Syst. Nat. Veg. ed. lous; staminate flowers numerous, peripheral and 10. 899. 1759, non Vellozo, Fl. Flumin. 53. 1829 amongthe pistillateflowers;tepals2, with minutesub("1825"), Ic. 1: t. 139. 1831 ("1827");Houstoun, globose hairs; stamens 2; filaments longer than the Philos. Trans.37: 196, t. 1. 1731;Lamarck,Encycl. perianth;stigmas? unequalin length,0.2-0.8 mm long. 2: 317. 1786; Willdenow, Sp. P1. 1(2): 683. 1798; Endocarp body 2-2.2 x 1.5-1.8 mm, ? tuberculate.

204

FLORA NEOTROPICA

Fig. 59. Dorstenia drakena.1. Habit(McVaugh20377). 2. Leaf (McVaugh20370). [By E. M. Hupkensvan der Elst, Utrecht.]

TAXONOMICTREATMENT

205

3m7~~~~~~~~c

Fig. 60. Dorsteniadrakena.1. Habit(Rzedowski22773) (formC). 2. Habit.3. Inflorescence(McVaugh19214). 4. Leaf (McVaugh19149). [By E. M. Hupkensvan der Elst, Utrecht.]

206

FLORANEOTROPICA

tela, rd. to Mazatan,ca. 1000 m, 12 Jul 1951, H. S. Gentryet al. 10850 (MICH,NA); rd. Tepic-Jalcocotan, 1300 m, 25 Dec 1970, Grahamet al. 1281 (MICH);15 km SE of Ahuacatlan,rd. to Barrancadel Oro, 11001300 m, 7 Jul 1957, McVaugh15181 (MICH);18 km W of Tepic,9 km E of Jalcocotan,7 Sep 1960,McVaugh Representative specimens examined. MEXICO. 18818(MICH).OAXACA:2 kmS of Niltepec,17Jul 1959, 2 AGUASCALIENTES: Sierradel Laurel, 35 kn SWof Calvi- King 1744 (MICH,NY, US); Isthmusof Tehuantepec, Nr. Tuxtla km E of Zanatepec,21 Jul 1959,King 1904 (MICH,NY, 1lO,25 Jul 1976,Correa163 (MICH).CHIAPAS: Guti6rrez,5 Jul 1990, Reyes Garcia et al. 1770 (F); 8 km US); Rio San Geronimo,Jul 1914, Purpus7137 (F, GH, MO,NY,US);Mun.Pochutla,3 km SW of PuertoAngel, W of Tuxtla Gutierrez, 17 Jul 1990, Reyes Garcia 1951 (F). CHIHUAHUA: Rio Mayo, Guasaremos, 2 Aug 1935, 24 Sep 1965,Rzedowski21160 (MICH);Dtto.TeotitlAn, H. S. Gentry 1551 (F, GH, MO, NY, US); Batopilas, 9 Oct Teotitlande Camino,7 Jul 1938,Schulteset al. 101 (NA). 1898, Goldman 238 (GH, NY, US). COLIMA:Nr. Playa MexIco: Ca. 1 kmW of Cerrodel Campo,rd.to El Salitre de Santiago, rd. to Manzanillo airport,21 Jul 1957, (San Miquel),30 Oct 1970, Graham1259 (MICH);ca. McVaugh15596 (MICH);21 km WNWof Santiago,rd. 5 km SW of Nuevo San Tomas,rd. to Tingambati,1250 to Cihuatlan (in Jalisco),25 Jul 1957, McVaugh15746 m, 2 Nov 1970, Graham1273 (MICH);20 km N of (MICH);SW of Nevado de Colima,2-3 km S of Hda. Amatepec,rd. to Tejupilco,30 Oct 1970, Graham1254 San Antonio, 1200-1250 m, 11 Aug 1957, McVaugh (MICH);Temascaltepec,1750 m, 1 Aug 1933, Hinton 16098 (MICH);Hwy. 110, 3 km W of bridgeover Rio 5078 (GH,MO,NY). PUEBLA: Hwy. 190, 1-2 km E of Tuxpan,27 Aug 1973,Stevenset al. 1805 (MICH).GuA- Tepexco, 28 Oct 1970, Graham et al. 1246 and 1248 NAJUATO: Tepic, 1904,Dugues5 (GH).GUERRERO: Nr. (MICH);rd. Maramoros-Acatlan,km 78, Las Cruces, et al. 709 (MICH);CerroChanIguala, 7 Aug 1945, Alexander et al. 2031 (MICH, NY, 27 Aug 1970,Herndndez US);NW of Chilpancingo,8 km fromjct. with Hwy.95, tepeque,nr.Pilcaya,26 Jul 1943,Whiteet al. 5077(MICH). 1700m, 24 Nov 1971,Dieterle4206 (MICH);Dtto.Mina, SINALOA:Nr.Culiacan,19Aug 1904,Brandegees.n. (US). Patambo,28 Jul 1936, Hinton9172 (F, NY, US); Dtto. SONORA:SanBernardo, Rio Mayo,2 Aug 1959,Arguelles 26 Jun 1939,Hinton14354 107 (B, P, US); Rio Mayo,CanyonSapopa,14 Oct 1934, Galeana,Moreno-Camalote, (NY,US); Rio Balsas,26 Aug 1910,Orcutt4189 (F, GH, H. S. Gentry1026 (F, GH, MO); 18 km E of Alamos, MO, US); 8 km NW of Chilpancingo,rd. to Chichi- Rio Cuchujaqui,12 Sep 1966,Henrickson2432 (MICH); hualco, 1350 m, 7 Jul 1966,Rzedowski22773 (MICH); rd. Alamos-Milpillas,6 km E of ArroyoCuchujaqui,9 Iguala,Canyonde la ManoNegro, 11 Aug 1905, Rose Sep 1971, Keil et al. 8101 (MICH);ArroyoGochico,E et al. 9370 (GH, NY, US). JALISCO:12-15 km SW of of San Bernardo, 5-9 Aug 1935, Pennell 19519 (US). Autlan,23 Jul 1951, H. S. Gentry10946 (MICH,NA); VERACRUZ: Rio Cataxtla,PiedraMovite, 17 km from 18 km S of Talpade Allende, 1200-1700 m, 18-19 Oct jct. of Hwys. 150 & 145, 11 Oct 1970, Graham1172 1960, McVaugh20377 (MICH);Puente Pedro, 8 km (MICH);Mun.PuenteNacional,Matade Caina,25 Sep SW of Tecalitlan, 19 Nov 1970, McVaugh24497 and 1971, Ventura 4329 (MICH). 22 May 1889, Pringle 24498 (MICH);nr. Guadalajara, GUATEMALA. ALTA VERAPAZ: BetweenSanPedro 2498 (BR, F, G, GH, GOET,K, LE, MO, NA, NY, P, CauchAand Senshui,Apr-Sep 1885, Tuerckheim310 US), 11 Jul 1902,Pringle11172 (F, GH,K, MICH,MO, (GH, MICH,NY, P, US); Sasis, 1100 m, May 1908, NY, US); Hwy. 54, Guadalajara-Zacatos,rd. to Las Tuerckheim II.2207 (F, 0, P, US); Rio Cheacte, Finca Camochas,ca. 1700 m, 13 Jul 1971, Oliver et al. 677 VolcAn,4 Mar 1939, Wilson 277 (F). CHIQUIMULA: (U); 15 km S of Autlan,ca. 1900m, 19Aug 1949, Wilbur QuebradaShusho, above Chiquimula, 14 Oct 1940, et al. 2420 (MICH). MICHOACAN:11-13 km SW of Standley 74248, 74349 (F). IZABAL: Gualan,26 May Azalzingan,rd.to DosAguasandAguilla,5-9 Sep 1972, 1919,Blake7698 (US). JUTIAPA: Atescatempa,ca. 1000 Dieterle 4292 (GH, MICH); Dtto. Coalcoman, Villa m, Jul 1894, E. J. Heyde et al. 6382 (F, GH, MICH,NY, Victoria, 18 Aug 1939, Hinton 15119 (GH, NY, US); US). ZACAPA: Rio Hondo,base of Sierrade las Minas, Mun.Tancitaro,rd. Tancitaro-Apatzingan, 30 Jul 1940, 11 Oct 1940, Standley74033 (F); nr. Zacapa,7-16 Oct Leavenworth388 (F, GH, MO, NY); Mun.Apatzingan, 1940, Standley 74248 (F); between Agua Blanca and La Majada, 5 Aug 1941, Leavenworth et al. 1316 (GH, Cumbrede Chiquimula,15 Oct 1940, Standley 74414 MO, P); Cerro de Carboneras,22 km S of Uruapan, (F);Rio Hondo,baseof Sierrade las Minas,10 Oct 1939, Distribution (see Fig. 79.15). From Mexico to southwesternCostaRica (GuanacasteandPuntarenas); in shade, in various types of ? dry forest or scrub, in Mexico often in oak forest, often in rocky places, also in secondarygrowth;to ca. 2000 m.

1100-1200 m, 6-22 Oct 1961, King et al. 4820 (MICH, NY, US). MORELOS:Sierra Grande, nr. Jojutla, 1300

Steyermark 29462 (F). EL SALVADOR. Nr. Salvador, May 1922, Calder6n

m, 13 Jul 1947,Barkelyet al. 2302 (MICH);Amacuzac, 704 (NY, US); Izalco, Jul 1923, Calder6n 1703 (NY, rd to Taxco, nr. border Morelos-Guerrero, 2 Aug 1961, US); Cant6nSanAntonioChavez, 16 Sep 1978, Seiler Feddema 1635 (MICH); 10 km ESE of Cuernavaca, rd. 546 (F); nr. Sonsonate, 18-27 May 1922, Standley to Casantla, Canyon de Lobos, 28 Oct 1970, Graham et 22335 (GH,NY, US); nr. La Uni6n, 16 Aug 1938,Stork al. 1245 (MICH);Huajintlan, Jul 1929,Lyonnet257 (GH, et al. 8752 (GH,MICH,MO, NA, US); Dtto. MirazAn, MO, NY, US); Xochiltepec, Jul 1938, Lyonnet 2163 Montecristo,15 km NE of San Miquel, 10 Dec 1941, (US). NAYARIT: NE of SantaMariadel Oro, 18-20 Aug Tucker508 (F, MICH,NY, US); Dtto. La Libertad,La1959, Feddema731 (MICH);10-12 km W of Compos- guna Calderas, 11 Jun 1992, Villacorta et al. 1107 (MO).

TAXONOMIC TREATMENT

HONDURAS.CHOLUTECA: Llano de Choluteca,2 km fromCholuteca,5 Aug 1955,MolinaR. 5473 (F);nr. Pespire, 18-25 Oct 1950, Standley27183 (F). COMAYAGUA:Rio Humuya,W of Comayagua,27 Jun 1964, Standley 14300 (F). EL PARAiSO: Drainage of Rio Yeguare,betweenMataIndioand Lipaza,25 Jul 1951, Molina R. 4064 (US); valley of Rio Guayambre,Valle de Jamastran,16 Jul 1956, MolinaR. 7506 (F, US); rd. Jagua,3 Jul 1962, Websteret al. Zamorano-Yuscaran, 11932 (U). MORAZAN:Drainageof Rio Yeguare,Santa ClaraCr., 14 Jul 1948, Glassman 1888 (F, NY); Las Mesas, 14 Jul 1947,MolinaR. 309 (F);Rio de la Orilla, SE of Zamorano,25 Aug 1953,MolinaR. 5317 (F, US); Santa Inez, Aug 1943, Rodriguez429 (F); Zamorano, Rio Caparossa,22 Jul 1949,Standley21568 (F). VALLE: Ampala, 12 Sep 1945, Rodriguez3411 (F). NICARAGUA. BOACA:4 km S of Boaquito,21 Oct

207

919 (MO);23 km SE of Liberia,23 Jul 1964, Lent 86 (CR, F, NY); 1 km E of Playasdel Coco, 14 Sep 1973, Solomon630 (CR);nr. La Cruz,23 Aug 1979,Stevens 13606 (BG,F); Catalina,28 Jun 1928, Stork2780 (F). PUNTARENAS: E of Monteverde,29 Oct-2 Nov 1975, Burgeret al. 9892 (F).

Dorsteniadrakenashowsstrikingsimilaritiesto D. contrajerva in thevariationof thevegetativeparts, butthevariation patterns ofthetwospeciesarenotquite similar.Dorsteniadrakenais a variablespecies,even veryvariable,if materialof whichthe identityis not quitecertainis included,as in thepresenttreatment. Somecollections fromGuatemala (Verapaz: Tuerckheim 310 and II.2207 and Wilson277) and two collections from El Salvador(La Libertad:J4llacortaetal. 1107;

andSonsonate: Standley22335)havedistinctlylobed fromtheshapeof thereceptacle, receptacles. Judging Moreno21509 (MO);2 km N of Boaquito,4 Jul 1983, the broad fringe and thepresenceof a broadzoneof Moreno 21548 (MO). CARAZO:Rio Escalante, Estero thesecollections probably peripheral staminate flowers, 22 Jun Aranda Mun. Chacocente, 1984, et al. 87 (MO); 1982, Moreno 18067 (MO); La Cruz, 14 Jun 1983,

Diriamba,13 km SW of Diriamba,10 Jul 1984,Moreno belong to D. drakena.Although in D. contrajervathe 24315 (MO). CHINANDEGA: Amaya, 19-21 Jun 1923,

fringeof thereceptacleis usuallyverynarrowandthe

Maxonet al. 7125 (US); rd. Cinco Pinos-SanFrancisco pistillateflowersoccurnearthemarginof therecepdel Norte,km 5, 28 Sep 191, Moreno11753 (MO);NW tacle,severalcollectionsof thisspecies--collectedin of VolcanCosigiuina,18 Jun 1983, Sandinoet al. 4394 Guatemala (especiallyin Pet6n),Mexico(especiallyin (MO);nr.Chichigalpa,12-18 Jun 1947,Standley11182 Veracruz andYucatan), aswellassomefromBelizeand 8 km S of Cuapa,21 Sep 1983, Nee ElSalvador-havearelatively (F). CHONTALES: (to 1.5mm)broadfringe 28247 (MO);Comarca,nr. QuebradaManiqua,13 Jul of thereceptacleanda ? distinctzone of peripheral 1976, Neill 7438 (MO);Hda.Corpus,W of Juigalpa,5 thecollections staminate flowers.Therefore, mentioned Sep 1982, Stevens 21795 (MO). ESTELI:Chayote, 31 an extreme of D. contrajerva. above might represent Jul 1983, Moreno 21772 (MO); Pan-AmericanHwy., km 157, 1 Aug 1983,Moreno21808 (MO);Kukamonga, Anotherpossibilityto be consideredis hybridization 1 Aug 1983,Moreno21827 (MO).GRANADA:IslaZapa- between D. contrajervaandD. drakena. Anotherpeculiarform(see Fig. 60.2-4) is repretera, Aug 1982, Grijalva864 (MO); nr. Granada,Jun 1869, Levy 45 (K, P), 2 Jul 1923, Maxonet al. 7594 sented by the collections Dieterle 4060 andMcVaugh km 19148, 19149, 19214, 20261, 20262 and21205, made (GH, NY, US); Casa Tejas,rd. Nandaime-Granada, 21, Moreno 17095 (MO). LE6N: 6 km NW of Puerto in Mexicoin the Statesof JaliscoandNayarit.These Momotombo,14 Jul 1984, Grijalvaet al. 3882 (MO); collections arecharacterized tobroadly by suborbicular Rio Sinecapa, 15 Sep 1977, Stevens 3852 (BG), Que- ovatereceptacles, beingoften? lobateandhavinga narbradaLas Ruedas, NW of El Transito, 16 Oct 1937, rowfringeandradiating(marginalandsubmarginal) Rd.Managua-Masachapa, Stevens4717 (BG).MANAGUA: withtransitional featuresis notpresent bracts.Material 14.8 mi S of jct. with Rt. 12, 18 Aug 1977, Croat43728 ofthese the collections studied. The distinctness among (MO);El Portillo,rd. to Masachapa,nr. San Rafaeldel makesthe positionof the collections Sur,9 Jun1983,Moreno21494 (MO);Hwy.10, nr.bridge inflorescences matchthenormalD. drakena) of Rio LaAduana,18Aug 1977,Stevens3374 (BG,MO). (whichin otherfeatures MASAYA: VolcanSantiago,Oct 1975, G6mezet al. 6185 dubious.Itsuggeststhepossibilityof theirrecognition andplasticity variation (CR), rd. Managua-Masaya,km 17.5, 24 Sep 1982, atthespecificlevel.Theextreme mefromcreatin Dorsteniaspeciesrestrain Grijalva1219 (MO);VolcanSantiago,27 Oct 1977,Neill occurring PresaSantaBarbara, inganewtaxon.Morematerial 2794 (BG, MO, U). MATAGALPA: andfieldwork areneeded. San Juande Dios, 22 Jun 1983, Moreno21582 (MO); A thirdform(see Fig. 59), however,certainlybebetween PuertasViejas and Esquipulas,27 Jul 1983, longingtoD. drakena, bythecollections is represented Stevens22321(MO).RivAs:Islade Ometepe,14 Sep 1983, Hinton 5078, McVaugh20376, 20377, 20378, 24518, Moreno22036 and22066 (MO);convergenceof Rio La and Wilburet al. 2420, fromMexico,the Statesof PitaandRio Escalante,3 Aug 1978,Stevens9722 (MO); havea crenate(and Rio Amayo, 20 Aug 1981, Stevenset al. 20582 (MO). JaliscoandMexico.Thereceptacles In other featuresthey partly slightly lobate) margin. Rio Potreroarea,ca. COSTA RICA. GUANACASTE: 18 km SE of Liberia,4 Jul 1968, Burger6123 (CR, F); matchnormalD. drakena.Thecollectionshavebeen 5 km N of Bagaces, 1 Aug 1965, Croat521A (MO);3 madeat 1200-1800m, in oakforest,andmightrepresucha taxonat km N of Canias,FincaLa Pacifica,14 Jun 1971, Gentry senta separate Establishing subspecies.

208

thismomentappearsto be futileconsideringthe extremevariationin thespeciesandtheproblemsstillto be solved. ThematerialdescribedasDorsteniacrispata(see Fig.60.1),butgraduallypassingintonormalD. drakby severalcollections,mainlyfrom ena,is represented Jalisco,andMexico. Mexico,theStatesof Guerrero, Theplantsareoftenrelativelysmall,havethickrhi(oracute) zomes,andthelobesofthelaminaarerounded in attheapex.Thefloweringfaceof theinflorescence thisformis oftenpurpleorreddish. EMYGDIOA SECTION

9C. Dorstenia Linnaeussect. Emygdioa Carauta, Bradea2(21): 151. 1976.Type.Dorsteniabrasiliensis Lamarck.

FLORANEOTROPICA

lamina intemodesshort.Leaves in spirals,subrosulate; ellipticto oblong(to subovate),9-13 x 4.5-6.5 cm, subcoriaceous;apex obtuse;base cordate;margin lowersurto repand; uppersurfaceglabrous; subentire to hirtellous,on thelowerpartof the facepuberulous lateralveins6-7 pairs, venationpinnate; midribhirsute; subscalarivenation ? distinctly loop-connected; tertiary petiole3-18 cm long,hirsutewith formto reticulate; hairs;stipulesovate,0.5-1 cmlong,uninervate, straight hairs. (tohirsute),mainlywithstraight acute,hirtellous Inflorescenceswithatleastthemarginof thereceptacle peduncle2.5-3 cm long,hirsute;receptacle purplish; centrallyattached,discoid,orbicularin outline,ca. 2 cmdiam.,theoutsidehirtellous,themarginentire,the fringeto 3 mmbroad;bractsin 1-2 rowson andbelow ones orsomeof thesubmarginal themargin,appressed ? radiating, triangular to ovate,to 2 mmlong,puberuflowersamongthepistillousto hirtellous;staminate lateones;tepals2;stamens2; filamentslongerthanthe stigmasnotseen.Endocarp body notseen. perianth;

stem small,oftensubacaulescent; Plantsherbaceous, intemodes unbranched; mostly often(partly) rhizomatous, Leavesinspirals, mostly short,if longthestemscreeping. laminaovateto elliptictooblongtosuborbicurosulate; Distribution (see Fig. 79.4). In eastem Brazil sometimes peltate,entire toreniform, larortocordiform (EspiritoSanto),knownonly fromthe type, froma incised;venaorsometimespinnatelyto subpalmately moistandshadedplace. venationusually thetertiary tionpinnateto subradiate, species,notshowbroad, Dorsteniaconceptionisis a distinct long;stipules(rather) reticulate; petiolerelatively ac- ingclearrelationswithotherspeciesof thesection. ovatetotriangular, mostlyuninervate. Inflorescences sometimes? irregularly (andorbicular), tinomorphic onshortappendages), sometimes (bracts lobed,bracteate thefringemostlynarrow;pistillateflowersnumerous. 30. DorsteniacayapiaVellozo,Fl. Flumin.52. 1829 someandtuberculate, bodymostlytetrahedral Endocarp ("1825"),Ic. 1:t. 137. 1831("1827");Miquel,in timessubgloboseandsmooth;seedsmall;endosperm Martius,Fl. Bras.4(1): 167. 1853;Bureau,in De embryowithflatandequalcotyledons. present; Candolle,Prodr.17:262. 1873;Hassler,Annuaire Cons.Jard.Bot. Geneve21: 115. 1919;Carauta, Thissectioncomprises18 species,all in theNeo29(44): 109,t. 32. 1978.Berg,Proc. Rodriguesia and tropics.IthasitsmaincenterintheGreaterAntilles Kon.Ned. Akad.Wetensch.,Ser.C, Biol. Med. adjacentpartsof CentralAmericaandMexico.Inthis Sci. 89: 143,t. 2. 1986.Type.Fl. Flumin.Ic. 1:t. area14speciesoccur;therestof thespeciesarefoundin Fig. 61 137. 1831. SouthAmerica,? centeredin eastemBrazil.Thissection comprisesa few geophyticspecies(suchas D. Herb, withthestem(atleastpartly)subterranean, brasiliensisandD. cayapiap.p.),occurringin relatively (1-)5-2(-20)mmthick,puberulous; internodesshort is not orpartof themlong,stempartswithshortintemodes innon-foresthabitats. Geophytism dryconditions, of the(rhizoma- sometimes succulence bypronounced accompanied lamina tuberous. Leavesin spirals,rosulate; mayberegarded ovateto cordiform tous)stemsasinAfrica.Sect.Emygdioa entireor3-7-lobedto toreniform, as derivedfromsect.Dorstenia.Thegeneralhabitof parted,3.5-13(-17) x 2.5-10(-16) cm, chartaceous, the speciesof sect.Emygdioamatchesthatof themono- oftenvariegated; basecordate; apexobtusetorounded; typic Africansect. Bazzemia(Berg & Hijman, 1999). tosinuateto lobate;upmargincoarselycrenate-dentate per surfaceglabrousor with sparseto ratherdense, minute,conicalhairs,smoothto scabrous;lowersur29. Dorstenia conceptionis Carauta,Bot. Mus. Bot. onthe(main) facesparselytorather denselypuberulous Curitiba17: 1, t. 1. 1974; Carauta,Rodriguesia29 lamina often to hirtellous; the base of towards veins, (44): 104, t. 30. 1978. Type. Bazil. EspiritoSanto: faintly veins 3-7(-8) pairs, venation lateral pinnate; Itarana,Jatibocas,28 May 1946,Bradeet al. 18461 atthe branching) (after terminating or loop-connected (holotype,RB;isotypes,F, GUA,R, SP,U). venationin largeleavespartlyscalarimargin;tertiary theup3-5 mm form;petiole3-15(-25) cmlong,puberulous, Herb,with the stem(mainly)subterranean, thick,hirsuteto hirtellous,mainlywithstraighthairs; perpartusuallyhirtellous;stipulesbroadlyovateto

TAXONOMICTREATMENT

209

Fig. 61. Dorstenia cayapia subspecies. 1. Subsp.paraguariensis.Habit(Hassler 7598a). 2. Subsp. cayapya. Habit(T S. Santos 3008). 3. Subsp.vitifolia. Habit(Irwinet al 15589). 4. Subsp.asaroides.Habit(Harleyet al 10949). [By E. M. Hupkensvan der Elst, Utrecht.]

210

0.2-0.5cmlong,coriaceous ellipticorto semiorbicular, plurinervate, midinconspicuously to subcoriaceous, ribslightlyprominentto plane,usuallyinconspicupartly ous, withobtuseto roundedapex,puberulous, usuallyunisexual, withuncinatehairs.Inflorescences sometimesbisexual,withthe innersurfaceof recepto (blackish-) tacleoralsothepedunclereddish-brown purple;peduncle2-10(-18) cm long, (sparsely)mireceptaclecentrallyto slightlyecnutelypuberulous; centricallyattached,discoid,oftenconcave,orbicular inoutline,(0.5-)1-2 cmdiam.,theoutsidetowardsthe themarginentireorslightlylobed, marginhispidulous, thefringealmostlacking;bractsin ca. 3 rowson the to ovate,to 1 margin,? appressed,broadlytriangular mm long, minutelypuberulous;staminateflowers amongthepistillateones;tepals2; stamens2;filaments longerthanthe perianth;stigmas(almost)equalin length,ca. 0.1-0.5 mmlong.Endocarpbodyca. 3 x 2.5 mm,smooth.

FLORANEOTROPICA 3. Lamina not variegated; lateral veins terminating at the margin; Paraguay, Bolivia and W Brazil ........................ 30b. subsp. paraguariensis

30a. Dorstenia cayapia Vellozo subsp. cayapia. Fig. 61 Dorstenia pachecoleoneana Machado, Brasil-Medico 49/50: 457. 1944; Carauta, Rodriguesia 29(44): 109. 1978. Type. Brazil. Rio de Janeiro:rd. Rio de Janeiro-Sao Paulo Machado 10 (holotype, RB). Dorstenia pseudo-opifera Hassler, Annuaire Cons. Jard. Bot. Geneve 21: 117. 1919. Type. Brazil. Rio de Janeiro: Nr. Rio de Janeiro, Allemdo s.n. (holotype, G).

Stemonly with shortintemodes. Laminanormally ovate to cordiform,ca. 2-1.5 times longer thanbroad, entire(or occasionally3-lobed),to 15(-17) cm long, to 7(-13) cm broad,usually variegated;marginusually Dorsteniacayapiais theonlyspeciesin thegenus coarselycrenate-dentateto sinuateor sublobate;upper thestaminateonesap- surfacesmooth, glabrousor with sparseminute coniwithunisexualinflorescences, pearingbeforethe pistillateones.Bisexualinflores- cal hairs;lateralveins 4-7(-8), often faintly loop-concencesoccur,butshowdistinctdifferencesinthepro- nected;petiole to 15(-25) cm long; stipules to 0.5 cm long. Receptacle (0.5-)1-2 cm diam. andpistillateflowers. portionsof staminate

Thepresentconceptof the specieswas proposed Distribution (see Fig. 80.2). In eastern Brazil by Bureau(1873). Gardner(1843) alreadyindicated (from Bahia to Rio de Janeiro);in places with shade thepossibilitythatDorsteniavitifoliaandD.asaroides or half-shade, associated with the rain forest habitat; differ- at low elevations. areconspecific.Onthebasisof morphological ences,mainlyin theleaves,foursubspeciescanberecRepresentative specimens examined. BRAZIL. BAto beecologically cayapiaappears ognized.Subspecies HIA: Itaimbe, nr. Jequitinhonha, 25 Nov 1970, Emygdio distinctas well.Themajorityof thespecimenscanbe et al. 2919 (RB); Santa Cruz de Cabralia, 3 Nov 1978, placedin one of the subspecies.However,thereare Euponino 363 (NY, RB); 17 km W of P6rto Seguro, features,especiallybe- Reserva Biol6gica Pau-Brasil, 12 Feb 1974, Harley 16140 specimenswith? intermediate tweensubspp..vitifoliaandasaroides,andbetween (K); Mun. Alcoba9a, rd. Teixeira de Freitas-Alcobaca, km 32, 21 May 1980, Mattos Silva et al. 818 (GUA, K); subspp.vitifoliaandparaguariensis. Key to the subspecies of Dorstenia cayapia 1. Laminanot lobed, mostly reniformand somewhatbroaderthan long; stipulesto 0.2(-0.3) cm long; receptacleusuallyca. 1 cm diam.;Centraland AmazonianBrazil 30d. subsp. asaroides and Bolivia............... 1. Laminanot lobed,ovateto cordifonn,mostly longerthanbroad,or palmatelylobedto parted,stipulesto 0.5 cm long; receptacle mostly ca. 1-2 cm diam. 2. Laminapalmatelylobed to parted, mostly almost as long as broad;central Brazil, and Bolivia........... 30c. subsp. vitifolia 2. Laminanot lobed,mostly ca. 1.5-2 times longer than broad. 3. Laminavariegated;lateralveins often faintly loop-connected; E Brazil......... 30a. subsp. cayapia

Mun. Itape, BR.145, km 19, W of Itabuna, 2 Mar 1978, Mori et al. 9348 (RB, U); Itamaraju,Campo Alegre, rd. to Piraja, 14 May 1971, Santos 1633 (CEPEC, U); P6rto Seguro, Parque Nacional Monte Pascoal, 14 Nov 1996,

Thomaset al. 11326 (NY). ESPIRITO

SANTO:

Serrado

Cristalino, 18 Oct 1953, Duarte3725 = Gomes476 (RB); Colatina, Corrego Sao Gon9alo, affluent of Corrego Jose, 5 May 1934, Kuhlmann 300 (RB); Colatina, Estrada Velha, Rio Paucas, 1 Dec 1943, Kuhlmann 6589 (G,

MBM,NY, RB, U, US). MINAs

GERAIS:

BetweenSitio

and Barbacena, 10 Nov 1982, Glaziou 13211 (G, K, NY, P), Nov-Dec 1887, Glaziou 16348 (BR, F, G, IAN, K, P, US). RiO DE JANEIRO: Mun. Petropolis,Serrada Estrela,Meio da Serra, 10 Oct 1971, Carauta 1408 (GUA, RB); Mun. Rio Bonito, Bracana, Fazenda das Cachoeiras, Lachette 170 (R); Mun. Silva Jardim, Pogo d'Antes, nr. Sao Joao, 24 Nov 1976, Oliveira 264 (U); Mun. Maje, Serra da Estrela, nr. Mandioca, 2 Apr 1832, Riedel s.n. (LE); Ilha Furtada,25 Dec 1967, Sucre 2026 (GUA, RB). SAO PAULO: Aparecida, 15 Jun 1917, Porto 247 (RB).

TAXONOMICTREATMENT The distributionof subsp.cayapia largelymatches thatof eastem BrazilianDorstenia species associated with rain forest, whereas the other subspecies are associated with open vegetation, such as cerradoor savanna, or also secondary growth. This subspecies is, moreover,distinctin the variegatedlamina. 30b. Dorstenia cayapiaVellozosubsp.paraguariensis (Hassler) C. C. Berg, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med. Sci. 89: 143. 1986. DorsteniacayapiaVellozo(var.typica)f. paraguariensis Hassler,AnnuaireCons. Jard.Bot. Geneve 21: 116. 1919. Dorsteniaparaguariensis (Hassler) Carauta,Bradea2(37): 256, cum. tab. 1978. Type. Paraguay.Between Rio Apa and Rio Aquidaban, San Luis, 1908, Fiebrig 4296 (lectotype, Carauta, Bradea 2(37): 256. 1978, G; isolectotypes, BM, GH, GOET,K, M, NY, P, US, Z). Fig. 61 DorsteniacayapiaVelloza(var.typica)f. paraguariensis Hasslersubformalobata Hassler,Annuaire Cons. Jard.Bot. Geneve 21: 116, 1919. Type. Paraguay.Depto. Concepcion,nr. Concepci6n, Oct, Hassler 7598 (holotype, G; isotypes, BM, G, GH, K, MPU,NY, P)*. Stem only with short intemodes. Lamina ovate to cordiform,ca. 1.5timeslongerthanbroad,to 12cm long, to 8 cm broad;marginlobate to repand;uppersurface almost glabrous;lateralveins 4-7, terminatingat the margin;petiole to 17 cm long; stipulesto 0.5 cm long. Receptacle0.8-1.8 cm diam. Distribution (see Fig. 80.3). In Bolivia, Paraguay, and the adjacentpart of Brazil; in gallery forest and cerradovegetation;at low elevations. Representative specimens examined. BRAZIL. MATOGROSSODO SUL: Mun. Bela Vista, Rio Guaviral,

23 Oct 1987, Hatschbachet al. 51594 (BG, MBM). CRUZ: BOLIVIA.SANTA Nuflode Chavez,3 km S of Ascenci6nde Guarayos,Arroyode San Joaquin,17 Apr et al. 31879 (CTES);Prov.Chiquitos, 1977,Krapovickas 1 kmNEof Robore,RioRobore,22 Apr1980,Krapovickas et al. 36421 (GUA);Prov.Chiquitos,28 km NE of San Jose, PuertoSan Marcos,27 Apr 1980, Krapovickaset al. 36610 (GUA);Prov.Sara,BuenaVista,29 Dec 1926, Steinbach7665 (BM, U); Prov.Sara,BuenaVista, Rio Surutu,7 May 1981, VdzquezAvila 372 (SI, U)%. PARAGUAY.Depto. Caragautay,Colonia Fortuna, 7 May 1974,Arenas688 (CTES); 7 km fromCaraguatay, betweenRioApaandRioAquidaban,1908,Fiebrig4156, BM,G, GH,K,NY,P,Z, mixedwithmaterialof D. brasiliensis); Depto.Amambay,nr. Bella Vista, 19 Oct 1981, Schinini 21337 (GUA)%;Depto. San Pedro, Colonia Primavera,7 Jun 1956, Woolston684 (K, NY, P). The collectionsmarkedwith % havefeatures?transitional to subsp. asaroides, and those markedwith *

211

havefeatures? transitional to subsp.vitifolia.Subspeciesparaguariensis approachessubsp. cayapia in the

shapeof thelamina,butit appearsto be moreclosely relatedto subsp.vitifolia,andit mayevenproveto be only a varietyof subsp. vitifolia.

30c. Dorstenia cayapia Vellozo subsp. vitifolia (Gardner)C. C. Berg, Proc. Kon. Ned. Akad. Wetensch.,Ser.C, Biol. Med.Sci. 89: 143. 1986. DorsteniavitifoliaGardner,in Fielding& Gardner,

Sert.P1.1:t. 14. 1843;Miquel,inMartius, Fl.Bras. 4(1):170.1853;Bureau,inDe Candolle,Prodr.17: 262. 1873;Carauta, 107.1978. Rodriguesia29(44): Type.Brazil.Goias:betweenAlmas andNatividade, Nov 1839, Gardner3427 (holotype, K; isotypes,

B, BM,BR, Fl, G, P, US).

Fig. 61

Dorstenia morifolia Fischer & Meyer, Index Sem. Hort. Petrop. 11: 60. 1846; Miquel, in Martius, Fl. Bras. 4(1): 168. 1853; Bureau, in De Candolle, Prodr. 17: 262. 1873; Carauta, Bradea 2(37): 218. 1977. Type. Brazil. Minas Gerais: Mun. JeguitibA,nr. On9a, Rianchao da Onca, Langsdorff s.n. (holotype, LE). Dorstenia bryoniifolia Martius ex Miquel, in Martius, Fl. Bras. 4(1): 167. 1853; Martius, Flora (or Bot. Zeit.) 20(Beibl. 2): 128. 1837, nomen; Carauta, Rodriguesia 29(44): 107, t. 31. 1978. Dorstenia cayapia Vellozo var. bryoniifolia (Miquel) Bureau, in De Candolle, Prodr. 17: 262. 1873; Hassler, Annuaire Cons. Jard. Bot. Geneve 21: 116. 1919. Type. Brazil. Mato Grosso: Cuiaba, Martius 212 (holotype, M; isotypes, BR, G, F, L, LE, P). Dorstenia bryoniifolia Martius ex Miquel f. minor Hoehne, Exp. Scient. Rosevelt-Rondon, Annexo 2: 39. 1914. Syntypes. Brazil. Mato Grosso do Sul: Corumba, Fazenda Urucum, Hoehne 5823 and 5824 (SP?-n.v.).

Stemwithshortinternodes, occasionallyalsowith Laminacordiform somelongintemodes. (tosuborbicularorto subreniform), mostlyaboutas longas broad, atmostabout1.5timeslongerthanbroad,usually(3-) 5(-7)-lobedto -parted,to 17cmlong,to 16cmbroad, thelobessometimesalmostrectangular; uppersurface withrathersparse,minute,conicalhairs,sometimes atthe lateralveins4-5(-7), terminating scabridulous; margin;petioleto 15(-25)cmlong;stipulesto 0.5 cm long. Receptacle(0.5-)1-2 cm diam. Distribution (see Fig. 80.4). In centralBrazil and Bolivia;inplaceswithshadeorhalf-shade,oftenon sandy soil; a componentof cerradovegetation;it also occurs as a weed in com andbean fields; at low elevations. Representative specimens examined. BRAZIL. DISTRITO FEDERAL: Brasilia, 20 Sep 1984, Heringer

18650 (GUA). GoiAs: Vila Boa de Goias, Cachoeira

212 Grande,22 Jan 1969, Carautaet al. 721 (GUA, RB); Anapolis, 14 Mar 1979, Hatschbach 42164 (GUA, MBM);Formosa,Rio Tiquira,80 km fromBrasilia,18 Apr 1967, Heringer11408 (IAN, NY, U, UB)%; Serra de Caiapo,37 km S of Caiap6nia,22 Oct 1964,Irwinet al. 7176 (NY, RB); Serrade Rio Preto,ca. 15 km E of Cabeceiras,17 Nov 1965, Irwin et al. 10408 (M, NY, UB)%; Serra dos Pirineus, 50 km S of Corumbade Goias,rd.to NiquelAndia, 25 Jan 1968,Irwinet al. 19155 (NY,UB); MATO GROSSO: Barrado Gar9as,69 km N of Xavantina,FazendaSantaMaria,22 Nov 1969,Eiten et al. 9496 (K, UB);Valede Sonhas,93 kmN of Xavantina, rd. to Aragar9as,9 Nov 1968, Harley et al. 10990 (K, RB, U); Mun.AltoAraguaia,RioAraguaia,16Nov 1973, Hatschbach33281 (MBM);Mun.Baraode Malgago,4 Jan 1979, Maciel et al. 175 (INPA);betweenBuritiand Chapadadas Guimaraes,19 Oct 1973, Prance et al. 19219 (INPA,NY, U). MATO GROSSO DO SUL: Mun. Rio Brilhante,BR.267, Entroncamanta,22 Oct 1970, Hatschbach25087 (M, MBM);Mun.Aquidauana,Piraputanga,6 Oct 1962,Hatschbach30500 (MBM).MINAS GERAIS:Mun. Jequitiba,On9a, nr. RiachAoda On9a, 22 Jan 1977, Carauta2279 (GUA,RB, U); Mun.Unai, Patosde Minas,Meotada Corda,30 Aug 1950, Duarte 3019 (RB)*; Mun. Unai, Patos de Minas, Fazendado Chapador,28 Oct 1961, Duarte5714 (HB, RB, U); Rio Paraopeba,LagoaPreto, 31 Mar 1958, Heringer6343 (SP); 10 km SE of Paracatu,7 Feb 1970, Irwin et al. 26265 (NY, UB); Mun. Ituiutaba,SantaTeresinha,20 Dec 1949, Macedo 2041 (MO); Lagoa Santa, 2 Dec 1863, Warming1938 (C, P). PARANA: Mun. Guaira, ParqueNacional 7 Quedas,4 Nov 1981, Buttura 721 (RB);Mun.Roncador,Cancan,19 Oct 1973,Hatschbach 32898 (GUA, RB). SAo PAULO: Barretos,Rio Pardo, Nov 1917, Frazdo RB 15509 (U); Ibitiunga,Roseira, 14 Jan 1941, Grotta4 (GUA);Piracununga, Bagua9u,8 Jan 1955, Kuhlmann3537 (SP); Moji-Mirim,Vila de Emas,Rio Moji-Mirim,FazendaSantaAnado Baguaqu, 21 Jan 1971, Joly s.n. (GUA); Sao Jose do Rio Preto, Alamedadas Cristalidas,Jardimdos Seixas,23 Jan1974, Menezes22 (RB). BOLIVIA. SANTA CRUZ: Prov. Santiesteban,Gral. Saavedra, 19 Apr 1977, Krapovickas et al. 31524 (CTES, RB); Prov. Sara, Buena Vista, 12 May 1921, Steinbach5654 (G), 29 Dec 1926,Steinbach7665 (BM, F, G, K, MO,NY, S, U)%. The collections markedwith * have featurestransitional to subsp. asaroides, and those markedwith % have features transitionalto subsp. paraguariensis. Subspecies vitifolia is characterizedby the palmately lobed to partedlamina,but it is not always clearly distinguishablefrom subsp. asaroides.

30d. Dorstenia caya pia Vellozo subsp. asaroides (Hooker) C. C. Berg, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med. Sci. 89: 143. 1986.

FLORANEOTROPICA Dorstenia asaroides Gardnerex Hooker, Ic. P1. Ser. 1. 4: t. 399. 1841; Gardner in Fielding & Gardner,Sert.P1. 1: t. 14. 1843; Fischer & Meyer, Index Sem. Hort.Petrop. 11: 62. 1846; Miquel, in Martius,Fl. Bras. 4(1): 168, t. 61. 1853; Carauta, Rodriguesia 29(44): 1 11, t. 33. 1978. Dorstenia cayapiaVellozovar.asaroides (Hooker)Bureau,in De Candolle, Prodr. 17: 263. 1873. Type. Brazil, Cear.: Crato,Gardner2001 (holotype,K; isotypes, BM, G). Fig. 61 Dorstenia asaroides Hookervar. celiae Carauta& Valente,AtasSoc. Bot.Brasil1(20):117,t. 5. 1983.Type. Brazil. Minas Gerais: Mun. Montes Claros, Fazenda Buritis, 29 Dec 1981, Carauta et al. 4006 (holotype, RB; isotypes,BC, GUA,HB, R, SP,U, UNB, US, VIC). Stem often with some of the intemodes or sometimesalmostall ofthem long.Laminarenifonn(to suborbicularor to cordiform),usually somewhatbroader than long, sometimes variegated, margin ? coarsely crenate-dentateto sinuate, sometimes sublobateor to 3-5-fid, to 8.5 cm long, to 10 cm broad;uppersurface with ratherdense, minute, conical hairs, usually scabrousto scabridulous;lateralveins 3-4(-5), terminating at the margin;petiole mostly to 10 cm long, sometimes to 20 cm long; stipules to 0.2(-0.3) cm long. Receptaclemostly ca. 1 cm diam. Distribution(see Fig. 80.5). InnortheimcentralBrazil and Amazonian Brazil; in shade or half-shade, or sometimesin openplaces,oftenon sandysoil;associated with cerradoandsavannavegetation;at low elevations. Representative specimens examined. BRAZIL. AMAPA:PBrto Santana,26 Jan 1961, Emmerich670 =

Andrade707 (HB, R);Macapa,5 km E of P6rtoSantana, 10 Feb 1962, Mattoset al. 9898 (SP); Macapa,rd. to Matapi,km 38, 2 Feb 1955, Pires et al. 4796 (IAN); Igarape Ariramba, 4 Aug 1962, Pires et al. 52339 (IAN, MG, K, NY, SP). AMAZONAS?:Rio Madeira,Ferreira s.n. (P). BAHIA: Maracas, rd. Jequie-Contendas de Sincora, Fazenda Santa Elena, 26 Jan 1980, Vianna 1422= Martins488 (GUA).DISTIuTo FEDERAL: Brasilia, Planaltina, 26 Nov 1975, Heringer 14878 (RB, UB); Brasilia, 6 Oct 1978, Heringeret al. 15598 (UB). GoiAs: Gurupi, 31 Jan 1976, Carauta1875 (RB);betweenAlmas andNatividade,Oct-Nov 1839, Gardner3428 (B, BM,

BR, Fl, G, K, P); Mun.Bom Jesusde Goias,BR.462, 18 Apr 1975, Hatschbach33358 (MBM); nr. Tocantin6polis, FazendaMucambo,28-29 Nov 1981, Jangouxet

al. 1716 (BG); Serrado Rio Preto, 10 km E of Cabeceiras, 19 Nov 1965, Irwin et al. 10501 (MICH, NY); Mun. Presidente Kennedy, 12 km W of Presidente Kennedy, 2 Feb 1980, Plowmanet al. 8279 (GUA, NY, RB). MARANHAO: Ilha de Sao Luis, Jan 1976, Barroso s.n.(GUA);Alcantara,nr.Fontedo Maritiau,25 Jan1976, Carauta 1860 (RB); Timon, 7 km W of Teresina (in Ceara), 28 Jan 1981, Krapovickaset al. 31786 (GUA).

TAXONOMIC TREATMENT

MATOGRosso: Aripuana,nr. HumboldtCenter,below Salto dos Dardanelos, 15-16 Oct 1973, Berg et al. P18539 (INPA, K, U), P18573 (INPA, MG, RB, U); R.12, ca. 5 km S of base camp,ca. 12?49'S,51?46'W,5 Nov 1968,Harleyet al. 10949 (K, P, RB, U); ca. 50 km N of Barra do Gar9as, rd. to Xavantina, 15 Oct 1964, Irwin et al. 6881 (RB); sources of Rio Jatuarana, Dec 1931, Krukoff 1657 (BM, G, U); Cuiaba, 16 Dec 1893, Malme 1212 (GH, US); nr. Olaria, 21 Nov 1967, Philcox et al. 3202 (K). MATO GROSSO DO SUL: Between Anastacio and Palmeiras, 17 Feb 1970, Hatschbach 23731 (MBM); Mun. Aquidauana, Piraputanga, 16 Oct 1972, Hatschbach 30498 (M, MBM, MICH).MINAS GERAIS:Mun. Paraopeba,Fazenda Sao Sebastiao, 30 km from Paraopeba,26 Mar 1956, Heringer s.n. (SP); Corinto, 4 Mar 1970, Irwin et al. 26892 (NY, RB, UB); Engenheiro Dollabella, 8 Nov 1938, Markgraf et al. 3168 (RB); base of Serra Ricardo Franco, 7 Feb 1978, Windisch 1684 (GUA). PARA:Belem, S forest of IAN, 5 Dec 1942, Archer 7935 (IAM, NY, US); Rio Paru do Oeste, Missao Tiryo, Igarape Muneni, 9 Feb 1970, Cavalcante 2349 (MG); Alemquer, Colonia Lauro Sodre, rd. to Igarape do Cipoal, 21 Feb 1953, Fr6es et al. 29385 (IAN, UB); Tapirinha, nr. Santarem, 3 Sep 1927, Ginzberger et al. 640 (F); Rio Tapaj6s, Parana do Tanari, 12 Apr 1924, Kuhlmann 1985 (RB, U); Mun. Conceigao do Araguia, ca. 20 km W of Redengao, 9 Feb 1980, Plowman et al. 8542 (IAN). PERNAMBUCO: Arco Verde, 24 Jan 1972, Andrade-Limas.n. (IPA); Tapera,Aug 1927, Pickel 1413 (IPA); Russinha, 26 Mar 1934, Pickel 3687 (IPA, GH, NY). ROND6NIA:Serra dos Tres Irmaos, sources of Rio Madeira, Mutumparana,29 Dec 1968, Prance et al. 8984 Boa Vista, Kuhlmann (K, NY, RB, U). RORAIMA: s.n.(RB). BOLIVIA. SANTA CRUZ: Prov. Velasco, Parque Nacional Noel Kempff M., Serrania de Huanchaca, rd. to "Los Fierros,"12 Dec 1991, Foster et al. 13940 (USZ); Prov. N4uflode Chavez, 2 May 1977, Krapovickas et al. 32170 (CTES); Prov. Andres Ibainez, 3 km SW of La Belgica, 21 Dec 1992, Nee 43207 (NY, USZ); Portachuelo and Buena Vista, 28 Oct 1980, Zuloaga et al. 1521 (SI, U). PANDO:Nr. Cobija, Jan 1912, Ule 9319 (K, MG).

213

late;laminabroadlyto narrowlycordiform, (1-)3-9 x (1-)2-7 cm,chartaceous; apexobtuse;basecordate with a broadsinus;marginsubentireor faintlycrenate(to subdentate); uppersurfacestrigilloseto substrigose, scabridulous; lowersurfacepuberulous tohirtellous (on themainveins),withuncinateandstraighthairs,scabrousto scabridulous; lateralveins(2-)4-6 pairs,after branching terminating atthemargin;petiole4-14 cm long,puberulous to (intheupperpart)hirtellouswith uncinateandstraight hairs;stipulestriangular, 0.2-0.4 cmlong,acuteto subacuminate, puberulous. Infloreswithuncicences:peduncle2.5-6 cmlong,puberulous nateandstraighthairs,to hispidulous; receptaclecentrallyattached, discoid,angular to substellate inoutline, 0.5-1.6cmdiam.,theoutsidepuberulous withretrorse uncinate hairstohispidulous, thefringeto0.5mmbroad; bractsin ca.3 rowson themarginandonfringe,ovate to semiorbicular, to ca. 0.5 mmlong,puberulous, the largeroftensetulose;staminate flowersamongthepistillateones;tepals2; stamens2; filamentsaslongasor slightlylongerthantheperianth; stigmasalmostequal in length,0.2-0.3 mmlong.Endocarpbodyca. 1.5x 1 mm,tuberculate, palebrown. northern Distribution(seeFig.80.1).InUruguay, Argentina, southern Brazil,andsouthern in Paraguay; shadedandhumidplaces;atlow elevations. specimens examined. BRAZIL.

Representative

PARANA:ParqueNacional do Iguagu,Rio Tamandua,7 May 1949, Duarte et al. 1811 (G, MBM, RB, U); Mun. Cerro

Azul, Rio Bom Sucesso, 22 Nov 1979, Hatschbach 42576 (MBM); Mun. Guaira,7 Quedas, 7 Sep 1981, Hatschbach 43989 (GUA, MBM).

Rio

GRANDE

et al. 50641

bach

Pereci Novo,

Rambo

Pereci

Montenegro,

18 Dec

Branco,

43791

Depto.

1 Jan 1978,

Schinini

Ybytyme,

20 Apr

1946,

ARGENTINA.

Tome,

23

Chimiray, Jan

GUA,

1908,

Candelaria, Depto. 1460

km

NE

et al. 1930

Cristobal (CTES,

20

Sep RB).

Ekman Loreto

Candelaria,

of

1974,

et

12 Sep

1944, Ana,

NY, Montes

26 Aug

Caniguas,

1945, Montes 4024 (K, LP, US).

Puerto

S).

11 Apr Depto.

Mar

San Tome,

Posadas,

(F, MICH,

Santa

(F, K, LIL); Depto.

20

Krapovickas

2017

(LIL,

CORRIENTES:

Tome,

Rio

(GUA);

Benitez,

Depto.

MISIONES:

(WIS). Cora,

13331

Rojas

San

(WIS);

Wachter 699

Cerro

Colonia

do LIL,

Barra Macaco

et al. 20160

Schulz 17477 (GUA, RB).

1970, San

CHACO:

Joao

HBR,

et al. 14111

Amambay,

1980,

15 Aug

(BR,

Itapiranga,

L. B. Smith

1964,

PARAGUAY.

Montes,

Herb,withthe stemsubterranean, (1-)3-7 mmthick, puberulousto hirtellous,mainlywith retrorseuncinate hairs;intemodeslong andshort.Leavesin spirals,rosu-

Uru-

Leopoldo,

Mun.

(LIL);

Rambo

Velho,

(LIL); Sao

42177

SANTA CATARINA: Mun.

(ICN).

Aquidaban,

31. Dorstenia tenuis Bonpland ex Bureau, in De Candolle,Prodr.17: 264. 1873;Valenteet al., Rev. Brasil. Biol. 37(1): 170, t. 2. 1977; Carauta,Rodriguesia 29(44): 123, t. 41. 1978. Type.Argentina. Corrientes:Withoutlocality,Bonpland 973 (holotype, P).

Rio

SUL:

do Montenegro,

Joao

Henz 27552

1944,

23 Jun 1949,

Ivoti,

Mun.

(MBM);

Aug

P); Santa Cruz do Sul, Trombudo,

Subspeciesasaroidesis distinctin itsrenifonnlamina andrelatively small plantparts.Specimenswith large leaves canbe regardedas transitionalto subsp.vitifolia.

DO

guay, 23 Feb 1916, Dusen17759 (S); Mun. Foz do Igua9u, ParqueNacional do Igua9u,Macucos, 13 Oct 1986, Hatsch-

al.

1983, Arroyo

26224

Bonpland, P, S); 216

23

Depto. (LIL);

1946,

Montes

Rico,

27 May

214

FLORANEOTROPICA

Dorsteniatenuisis clearlydistinctfromthetwo other speciesin the southempartof SouthAmerica,D. brasiliensis andD. cayapia. It shows strikingresemblances to D. fawcettii from the GreaterAntilles. D. tenuis is possibly the same as D. cordifolia, described by Lamarck(1786). This namehas been appliedtoD.fawcettii for a long time, as D. cordifolia Swartz.

32. Dorstenia brasiliensis Lamarck,Encycl. 2: 317. 1786; Stokes, Bot. Nat. Med. 4: 338. 1812; Nees von Esenbeck, P1.Medicin. t. 99. 1828; Fischer & Meyer, Index Sem. Hort. Petrop. 11: 62. 1846; Miquel, in Martius,Fl. Bras. 4(1): 168. 1853; Bureau,in De Candolle,Prodr.17: 263. 1873;Peckolt & Peckolt, Hist. P1. Med. 895. 1890; Carauta, Rodrigu6sia29(44): 113, t. 34. 1978. Type. Uruguay. Montevideo, Commersons.n. (holotype, P; isotypes, G, MPU). Fig. 62 Dorstenia tubicina Ruiz & Pav6n, Fl. Peruv. 1: 65, t.

102b.1798;Loddiges,Bot.Cab.18:t. 1784. 1832; Miquel, in MartiusFl. Bras. 4(1): 168. 1853; Grisebach.Fl. Brit. W. I. 154. 1864; Bureau,in De Candolle, Prodr. 17: 264. 1873; Hassler, AnnuaireCons.Jard.Bot. Geneve21: 119. 1919; Carauta,Rodriguesia29(44): 116, t. 36. 1978. Dorstenia brasiliensis Lamarck var. tubicina (Ruiz

& Pavon) Chodat & Vischer, Bull. Soc. Bot. Geneve, Ser. 2, 11: 258. 1920. Type. Peru. Huanuco:Nr. Chinchao,Ruiz & Pav6ns.n. (holotype, MA; isotypes,B, BM, F, G). Dorstenia

pernambucana

Arruda da Camara,

Discursosobrea utilidadeda instituicaodejardins nas principais provincias do Brasil 45. 1810; Schultes,Mant.3: 318. 1827;Carauta, Rodrigu6sia 29(44): 119. 1978. Type. Brazil. Pernambuco: Tapera,Pickel 1336 (neotype, designated by Carauta,Rodriguesia29(44): 119. 1978, IPA). Dorstenia infundibuliformis Loddiges, Bot. Cab. 20:

general index. 1833; Bureau, in De Candolle, Prodr.17:264. 1873.Type.Bot. Cab. 18:t. 1784. Dorstenia montevidensis Fielding & Gardner, Sert.

P1. 1: t. 13. 1843;Bureau,in De Candolle,Prodr. 17: 263. 1873. Type. Uruguay. Montevideo, Tweedie s.n. (holotype, K). Dorstenia tomentosa Fischer & Meyer, Index Sem.

Bot. Geneve21: 120. 1919. Type.Paraguay.Rio Apa, Nov, Hassler 7847 (holotype,G; isotypes, BM, G, K, MICH,MO,NY, P). Dorstenia montanaHerzog, Meded. Herb. Leiden 27: 73. 1915. Type.Bolivia. SantaCruz:Samaipata, Mar 1911, Herzog 1729 (holotype, B; isotypes,G, S). DorsteniabrasiliensisLamarckvar.palustrisHassler, AnnuaireCons.Jard.Bot. Geneve21: 118. 1919. Type.Paraguay.Depto.Central,nr.Lagode Ypacarai,Jun,Hassler3023 (holotype,G; isotype,G). Dorstenia tubicina Ruiz & Pavon var. opifera (Martius)Hasslerf. subexcentricaHassler,AnnuaireCons.Jard.Bot. Geneve21: 119. 1919.Type. Paraguay.Depto. Central,nr. Lago de Ypacarai, Sep, Hassler927 (holotype,G; isotypes,BM, G, K, NY, P). Dorstenia brasiliensis Lamarckvar. guaranitica Chodat& Vischer,Bull. Soc. Bot. Geneve, S6r. 2, 11: 257. 1920. Type. Paraguay.Depto. Cordillera, San Bernardino,Chodat& Vischer247 (lectotype,designatedhere, G). Dorstenia brasiliensis Lamarckforma balansae Chodat,Bull. Soc. Bot. Geneve,Ser.2, 11: 257. 1920.Type.Paraguay. Depto.Guaira,DonaJuana, nr. Villarica,17 Sep 1874, Balansa 1983 (holotype, G; isotype, P). DorsteniasabanensisCuatrecasas,Bol. Soc. Venez. Cienc. Nat. 17: 93. 1956. Carauta,Rodriguesia 29(44): 118. 1978. Type. Venezuela. Guarico: nr. Calabozo,Jul 1954, Aristeguieta& Herndndez 2316 (holotype,F; isotype, VEN). DorsteniaheringeriCarauta& Valente,Bradea2(5): 17, cum tab. 1975;Carauta,Rodrigu6sia29(44): 120, t. 39. 1978. Type. Brazil. DistritoFederal: Brasilia, 15 Nov 1975, Heringer 14350 (holotype, UB; isotypes,GUA, RB, US). Dorsteniaschulzii Carauta,Valente& Araujo,Bull. TorreyBot. Club 103: 173, t. 3. 1976; Carauta, Rodriguesia29(44): 121, t. 40. 1978. Type.Argentina.Formosa:Formosa,7 May 1964, A. G. Schulz 18318 (holotype, in herb.A. G. Schulz; isotypes,F, IPA,LIL, RB). DorsteniaamazonicaCarauta,Valente& Barth,Arq. Jard.Bot. Rio de Janeiro23: 105, t. 1 and t. 2. 1979.Type.Brasil.Goias:Araguaia,Rio BoaVista, FazendaBoa Vista, 28 Jan 1976, Carauta1863 (holotype,RB;isotypes,GUA,HB,MICH,R, US).

Hort. Petrop. 11: 61. 1846; Miquel, in Martius, Herb, with the stem subterranean,(I-)4-15 mm Fl. Bras.4(1): 168. 1853;Bureau,in De Candolle, thick, often (at least partly) d tuberous;mostly all inProdr.17: 263. 1873. Dorstenia brasiliensis Lamarck var. tomentosa ternodes short.Leaves in spirals, rosulate; laminael(Fischer& Meyer)Hassler,AnnuaireCons. Jard. liptic to oblong to suborbicular to (sub)obovate to Bot. Geneve 21: 118. 1919. Type. Brazil. Mato (sub)ovate or to lanceolate, 1.5-18 x 1-7 cm, coriaGrosso:Rio Pardo,Riedel 527 (holotype,LE). ceous to membranaceous;apex roundedto emarginate Dorstenia brasiliensis Lamarck var. major Chodat, to obtuseto acuteor to acuminate;base cordateto trunin Chodat& Hassler,Bull. Herb.Boissier,Ser.2, cate to obtuse;margin? irregularlycrenate(to faintly 3: 350. 1903. Dorstenia tubicina Lamarck (var. genuina) forma lobate);uppersurfacescabrousto smooth,with (somemajor (Chodat) Hassler, AnnuaireCons. Jard. times sparse),minute, conical hairs, also hirtellousor

TAXONOMICTREATMENT

215

Fig. 62. Dorstenia brasiliensis. 1. Habit (Liesner et al 5733). 2. Habit (Steyermark 89331A). 3. Habit (Pereira

245). 4. Habit(Troncosoet al. 2255). [By E. M. Hupkensvan der Elst, Utrecht.]

216 strigilloseto strigose,only strigoseto strigillose,or subglabrous;lower surface at least on the veins puberulous to hirtellous or to subtomentellous,mainly with straighthairs;venationpinnate;lateralveins 6-14 pairs, usually running after branching (1-2 times) almost straightinto the margin,terminatingin ? conspicuous hydathodes,sometimesfaintlyloop-connected;petiole (0.5-)1-9(-17) cm long, puberulousto hirtellouswith straightor often also with uncinatehairs; stipules triangular,0.2-0.7 cm long, coriaceous,uninervate,? distinctly carinate,with sparsehairs mainly on the margin. Inflorescencesgreenor the floweringfacepurplish; peduncle(0.5-)1-12 cm long, puberulousto hirtellous; receptaclecentrallyto slightly eccentricallyattached, discoid to broadlyturbinateto ? cup-shaped,orbicular in outline, 0. 5-3 cm diam., the outside puberulousto hirtellous, the margin usually entire, sometimes + crenate,occasionallysublobate,the fringe(almost)lacking; bracts in 1-2 rows on the margin or sometimes with severalrows, partlybelow the margin,appressed, broadlyovate, 0.2-1 mm long, minutelypuberulousto glabrous;staminateflowers among the pistillateones; tepals2; stamens2; filamentslongerthanthe perianth; stigmas unequalor equal in length, 0.2-0.5 mm long. Endocarp body ca. 1.5 x 2 mm, tuberculate. Distribution (see Fig. 80.6). FromnorthemrArgentinaandUruguayto Bolivia,Peru(HuAnuco, SanMartin), Colombia (Meta), Venezuela, Trinidad,Guyana,and FrenchGuiana;in cerradoand savannavegetation, in relativelymoist places, often in rockyplaces andalong streamsor streamlets;to 1500 m, in Bolivia to 2800 m. Representativespecimensexamined.COLOMBIA. La Serrania,between Rio Meta and Rio Ariari, 26 Nov 1939, Cuatrecasas 7889 (US); Llanos de San Juan, 1844, Goudots.n. (or A.]) (P). VENEZUELA.AMAZONAS: Nr.PuertoAyacucho,rd. to Samariapo,14Apr 1977,Huber604 (U). ApuRE:Dtto. et al. 5658 Munioz,HatoEl Polvero,28 Apr1987,Aymard META:

(BG). BOLivAR: Dtto. Piar,Calceta Larga,NW of El Man-

teco, Jul 1978, Delascio et al. 7227 (U); Dtto. Roscio, NW of Tumeremo,Rio Botanamo, Reserva Forestal Imataca,Feb 1980,Delascio et al. 8803 (VEN);1 km N of Paragua, 28 Jul1978,Liesneret al. 5753(U);RioCaroni, SantaRosa, 1864, Grosourdys.n. (or cat. no. 19) (P); nr. Tumeremo, 15 Jan 1840, Otto 1109 (B, P); Las 89299 (NY,VEN); Chichirras,11 Feb 1961, Steyermark Altiplaniciede Nuria, 47 km N of Tumeremo,12 Feb 1961, Steyermark89331A (GH, K, NY, VEN); Dtto. Cedefio,22.5 km SWof Calcaradel Orinoco,2 Sep 1985, et al. 131195(BG);nr.Miamo,13 Mar1973, Steyermark Unsheims.n. (VEN); El Tigre, Rio Cuchivera,18 Jun 1940, Li. Williams13377 (F, S, US, VEN). CARABOBO: SE of Valencia,El Paih, Trujillo4415 (MY). GUARICO: Calabozo,Est. Biol. Los Llanos,Aug 1965,Aristeguieta

FLORANEOTROPICA 5637 (VEN), 4 Sep 1965, Castillanos 38 (VEN). MONAGAS:Guanajuana, 1843, Funck 107 (LE, P). TRINIDAD. St. Augustin, St. Joseph Savanna, 8 Aug 1973, Adams et al. 13482 (TRIN); Arima, Jun 1878, Purdie 93 (K). GUYANA. Kanuku Mtns., Moco Moco R., 13 Jul 1995, Jansen-Jacobs et al. 4436 (BG, U). FRENCH GUIANA. Nr. Kourou, 4 Apr 1962, Hoock s.n. (U); Piste de St. Elie, km 3.7, 13 Mar 1986, Prevost 2130 (BG). PERU. SAN MARTIN: Tarapoto, Spruces.n. (K). BRAZIL. AMAPA:Macapa,BR.156, km 18, Rabelo et al. 2558 (BG). AMAZONAS: Manaus, Igarape de Santa Maria, 17 Nov 1954, Chagas INPA 391 (INPA, U); nr. Manaus, 20 Jan 1956, Chagas INPA 3345 (INPA); nr. Manaus, Paranary, 17 Mar 1924, Kuhlmann 1649 (F, G, K, L, RB, U); Rio Negro, nr. Sao Gabriel da Cachoeira, Jan-Aug 1852, Spruce 2196 (BM, BR, C, G, GOET, LD, LE, MG, MPU, NY, P). BAHIA: Barreiras, 24 Nov 1954, Black 54-17755 (IAN); Riachao das Neves, 12 Mar 1979, Hatschbach 42121 (MBM); Salvador, 8 Nov 1977, Valeria s.n. (RB). CEARA: Without locality, Allemao 1441 (P, R); Taboleiro, 10 Jan 1834, Luetzelburg 25951 and 26009 (M). DISTRITO FEDERAL: Ca. 12 km W of Taguatinga, rd. to Braslandia, ca. 1250 m, 26 Nov 1964, Irwin et al. 10675 (NY, UB)*; Brasilia, Horto Guara, 5 May 1962, Heringer 8937/1131 (F, HB, LP, PEL, UB)*; Reserva Zool6gica, 30 Mar 1963, Pereira 7457 (HB, MICH). GoIAs: Valley of Rio Parana, Rio dos Macacos, 5 Feb 1967, Duarte 10339 (RB); Serra Caiapo, ca, 20 km S of Caiap6nia, rd. to Jatai, 31 Oct 1964, Irwin et al. 7630 (NY, RB); BR.153, 20 km N of Itumbiana,20 Jan 1978, Krapovickaset al. 33092 (CTES); Goiatuba, 15 Feb 1964, Labouriau et al. 1168 (SP); Mun. Presidente Kennedy, 12 km W of Presidente Kennedy, 4 Feb 1980, Plowman et al. 8389 (MG). MARANHAO: Loreto,Ilhade Balsas,between Rio Balsas and Rio Paruaiba, 3 Apr 1962, Eiten et al. 3972 (K, NY, US); Timon, 7 km W of Terezina (in Ceara), 28 Jan 1981, Krapovickas et al. 37187 (GUA); Ilha de Sao Luis, Jan 1976, Ramos s.n. (GUA). MATOGROSSO:Aripuana, nr. Humboldt Center, below Saltos dos Dardanelos, Rio Aripuana, 16 Oct 1973, Berg et al. P18574 (K, MG, RB, U). MATO GROSSO DO SUL: Mun. Campo Grande, Capao Bonito, 5 Sep 1936, Archeret al. 83 (NA, US); Mun. Rio Brilhante, Entrocamento, 16 Feb 1970, Hatschbach 23664 (MBM); Mun. Sidrolandia, BR.163, 27 Oct 1970, Hatschbach 25262 (MBM); Mun. Ponta Pora, 25 May 1976, Hatschbach 38671 (MBM); Mun. Bela Vista, Rio Guaviral, 23 Oct 1987, Hatschbach et al. 51596 (BG); Corumba, Jan 1892, Moore 963 (BM); Mun. Corumba, Urucum, 22 Oct 1953, Pereiraet al. 440 (U). MINAS GERAIS: Carandai, 18 Nov 1914, Duarte 523 (RB); Serra do Cipo, ca. 1300 m, 9 Dec 1949, Duarte 2274 (RB, U); Paraopeba, Melo, 3 km N of Herto, 26 Nov 1965, Goodland 218 (U); Serra do Cip6, 3 Mar 1958, Heringer et al. s.n. (SP)*; Campo Florido, 6 km from Rio Sao Francisco, 12 Feb 1964, Labouriau 1137 (SP). PARA: Marajo, Mun. Salvaterra, Joanes, 15-16 Mar 1978, Bastos et al. 24 (MG, NY, U)

217

TAXONOMICTREATMENT and175 (MG);Vigia,Campinade Caembe,22 Jan 1950, Black50-8572(IAN);MonteAlegre,1 May 1916,Ducke MG 16113 = RB 1053 (BM, MG, RB, US); Belem,Nov 1897, Huber629 (MG);Mun. Concei9aode Araguaia, 20 km fromRedencdo,9 Feb 1980,Plowmanet al. 8481 (GUA, MG). PARAtBA: Without locality, Coelho de Moraes s.n. 2066 (US). PARANA:Rio CapaoGrande,19 Mar1903,Dusen 8012 (GH,P, US); Mun.PontaGrossa, VilaVelha,21 Oct 1914,Dusen 15695(GH,K, L, MICH, MO,NY, P); Mun.CampoMourao,CampoMourao,11 Dec 1960, Hatschbach7632 (MBM,SP); Mun.Guarapuava, Lagoa Seca, 14 Feb 1986, Hatschbach et al. 50348 (GUA, MBM);Mun. Ponta Grossa,Furnas,13 Dec 1965,Reitzet al. 17518 (GH,NY, P, US). PERNAMBUCO: 60 kmS of Recife,Saltinho,23 Jul 1983,Morawetz et al. 14-23783(BG);Recife,DoisIrm6es,Parquedo Institutode PesquisasAgron6micas,8 Apr 1936, Vasconcelos Sobrinhos.n. (IAN, IPA,SP). PIAut:Oeiras,May 1839, Gardner2310 (BM, F, G, GH, K, LE, P). RORAIMA: S of Cachoeirade Fuma9a,24 Jul 1987, Millikenet al. 513 (BG); Mun. Alto Sumure,Serrade Memoria,nr. FazendaTriunfo,29 Apr 1980,I. A. Rodriguezet al. 675 (IAN).RIOGRANDE DO SUL:Nr.Santanado Livramento, 10 Feb 1972,Cruzs.n. (HB,M, MBM);Guaiba,Fazenda 131 (RB); Sao Maximiano,22 Aug 1974,Maltzenbacher Santiago,EstanciaMundoNovo, 29 Jul 1974, Quintas 1021 (RB); P6rtoAlegre, 1 Dec 1948, Rambo24053 (LIL);Mun. P6rtoAlegre, Vila Maresa,17 Nov 1948, Rambo 38432 (P). SAO PAULO:Moji-Guaciu,Reserva Florestal,FazendaCampininha,28 Nov 1961, Fontella s.n. (GUA,SP);Itaim,20 Oct 1936,Hoehneet al. 36556 Cesar,24 Feb 1921,Kuhlmann (NA,NY);VillaCerqueira RB 14979 (RB); Corumbatai,Reservade Cerrado,29 Dec 1976, Mariniset al. 16 (HB, RB). RioYacuma, Espiritu, BOLIVIA. BENI: Prov.Ballivian, 6 Oct 1921, 4 Mar1987,Beck15014(BG);Rurrenabaque, Rusby817 (K, NY). LA PAZ: Prov.Iturralde,Luisita, 27 Feb 1984,Becket al. 10087 (BG);Prov.SudYungas, SantaAna de Mosetenes,15 Mar 1990, Hinojosaet al. 1063 (BG); Ixiamas,24 Dec 1901, R. S. Williams261 (BM, K, NY, US). COCHABAMBA: Prov. Mizque, nr. Santiago, 2800 m, 14 Jan 1995, Saravia et al. 1200 (BOLV).SANTACRUZ:Nr.SantaCruz,Dec 1910,Herzog delCarmen, 1299(G,S);Prov.Nuflode Chaves,Embocada 40 km SW of Concepci6n,2 May 1977,Krapovickaset al. 32155 (CTES,RB); Prov.Velazco, 12 km E of San Ignacio,6 May 1977, Krapovickaset al. 32272 (CTES, GUA, RB); Prov.Sandoval,San Mattias,18 Apr 1980, Krapovickaset al. 36200 (GUA);Prov. Chiquitos,28 km NE of San Jose, Puesto San Marcos,27 Apr 1980, et al. 36608 (GUA)%;Prov.AndresIbaiiez, Krapovickas rd. SantaCruz-Cotoca,km 12, 27 Jan 1988,Nee 36063 (BG);SantaCruzde la Sierra,27 Oct 1980, Zuloagaet al. 1433 (U). TARIJA: Prov.GranChaco,Cototo,7 km E of Villa Montes, 11-12 Apr 1977, Krapovickaset al. 31181 (CTES,G, GUA, RB). PARAGUAY.Cordillera de Altos, 1 Nov 1902, Fiebrig353 (BM,F,G, GH,GOET,K, M);Depto.Guaira, Villarica,May 1930,J0rgensen4286 (F, MO,NY, US),

Depto. San Pedro, Rt. 3, 10 km SE of San Estanislao, 14 May 1974, Schinini 8895 (GUA, RB); Depto. Chaco, Major Pedro Lagerenza, Rio Timane, I1 Apr 1978, Schinini et al. 15202 (CTES, RB); Depto. Amambay, Cerro Cora, 7 Aug 1980, Schinini et al. 20270 (GUA). ARGENTINA. CHACO:Colonia Benitez, 12 Mar 1972, A. G. Schulz 874 (GUA), 20 Oct 1906, Stuckert 16020 (G). CORRIENTES:Depto. Ituzaingo, Salto del Apipe, 21 Nov 1973, Lourteig et al. 2927 (CTES, G, P); Depto. Empedrado, Estancia Las Tres Marias, 22 Nov 1962, Pedersen 6646 (BR, K, LP, P, S); Depto. San Miguel, 12 km NE of San Miguel, Estancia Curuzu Laurel, 14 Jun 1974, Schinini et al. 9355 (CTES, GUA, RB); Depto. Monte Caseros, 13 km W of Monte Caceros, 10 Sep 1979, Schininiet al. 18293 (GUA).ENTRERios: Concepcion del Uruguay, May 1877, Lorentz 954 (BM, F, G, GH, M, P); Concordia, 19 Sep 1978, Troncoso et al. 2255 (SI, U). FORMOSA:Pirane, 6 Nov 1945, Morel 193 (F)%, 5 Dec 1945, Morel 524 (K)%; Potrero Norty, 28 Oct 1972, A. G. Schulz 18390 (herb. A. G. Schulz, RB)%. MISIONES:Depto. San Ignacio, San Ignacio, Cabrera et al. 29354 (K); Posadas, Bonpland, Jan 1908, Ekman 2026 (K, MICH, NY, S, U); Depto. Candelaria, Loreto, 11 Aug 1944, Montes 190 (U); Iguazu, Puerto Esperanza, 26 Jun 1950, Montes 1026 (M); Apost6les, San Jose, Renvoize et al. 3077 (K); Santa Ana, Jan 1907,

Spegazini 15461 (U). URUGUAY. Depto. Maldonado, Balneario Solis, Oct 1841, Burkharts.n. (SI); Depto. Florida, Timote, Estancia

Santa Clara, 12 Oct 1943, Gallinal et al. 5301 (US); Depto. Montevideo, Atahualpa, 25 Apr 1925, Herter 361 (F, GH, LE, M, MO, S, Z); Depto. Montevideo, Col6n,

La Paz, May 1925, Herter et al. 361a (NY, U, WIS); Depto. Maldonado, Cerra Pan de Azucar, 13 Oct 1970,

Krapovickaset al. 16329 (CTES). Collections markedwith * belong to the form describedas Dorstenia heringeri,andthose markedwith % belong to the form described as D. schulzii (see below). In the presentconceptD. brasiliensisis a polymorphicspeciesandalso the most widespreadin South America. In the southernpart of its area (Argentina, Uruguay,andsouthemBrazil)the speciesis represented by specimenswiththick(coriaceousto subcoriaceous), broadlyelliptic to ovate leaves with on the uppersurface mostly only dense, minute, conical hairs. In the northernpartof the area(Venezuela,Trinidad,French Guiana,the Amazon Basin, and northeasternBrazil) the speciesis representedby specimenswiththin(thinly chartaceous),mostly oblong to subobovateleaveswith mostly ratherlong appressedhairs on the upper surface. In thecentralpartof the area(centralBrazil,Peru, Bolivia andParaguay)most of the specimensaremore or less intermediatein leaf characters.In this central partof the areamany specimens have leaves with an obtuse to subacuteapex, a featureoccasionally found in specimensof the northernand southernpartsof the

218

area.Thevariationin leaf characters is moreor less distinctlyclinal. Altoughthenorthern andsouthernformsarequite distinct,thelackof discontinuities in thevariation rendersrecognition of intraspecific taxaalmostimpossible. However, within Dorstenia brasiliensis two rather

FLORANEOTROPICA

ginpartlyhispidulous, themargin? crenatedueto the basesof the(sub)marginal bracts,thefringe0.25-0.5 mmbroad,glabrousor with sparse,minute,conical hairs;bractsin ca.2-3 rows,on andbelowthemargin oralsoa few on thefringe,appressedorsometimes? radiating,ovateto elliptic,to 0.7 mmlong,minutely puberulous; staminate flowersintermixed withthepistillateones;tepals2; stamens2, filamentslongerthan theperianth;stigmasalmostequalin length,0.1-0.3 mmlong,oftenpurple.Endocarpbodyca. 1.5 x 1mm, ? tuberculate.

distinctformsoccur.Oneof themis foundin central Brazilandhasbeendescribedas D. heringeri.It has manyrowsof bractsbelowthe marginof the receptacle,whilethereceptacle normallybearsca.tworows of (sub)marginal to fiverows.Anbracts,occasionally otherformis foundin northern Argentina, Paraguay, Distribution(seeFig.81.5).InMexico;in forest, MatoGrosso.Ithasbeendescribed Bolivia,andsouthern oftenin rockyplaces,oras a weed;to 700 m. asD. schulziiandis distinctintherather thin,variegated Specimens examined. MEXICO.Withoutlocality, leaves.Considering theconsiderable variation withinthe speciesit seemshardlyjustifiedto giveformalrecogni- Galeotii s.n. (BR), 1841-1842, Karwinski 701 (LE). tionto thesetwoformsbytreatingthemasvarieties. GUERRERO: Achotla, Jul 1926, Reko 4907 (US). HIDALGO:between Huejutla and Macuxtepetla, 21 May Localnames.Venezuela: Bolivar:tusilla.Bolivia: 1942, Moore2912 (GH). SAN Luis POTOSI:Mun. San LaPaz:ere(Moseten); Cochabamba: forother, Antonia,Tanjasnec,11 Nov 1978,Alcorn2157A (TEX); sapirana; morecommonvernacular namesseethegenerictreat- Mun.El Pujal,Rio Tampanon,17 Jul 1939, Chase 7474 ment of Dorstenia, above.

33. DorsteniaexcentricaMoricand, P1.Nouv.Amer. 92, t. 59. 1840;Bureau,in De Candolle,Prodr.17: 263. 1873.Type.Mexico.Tamaulipas: Tampico,7 Feb 1827,Berlandier30 (holotype,G;isotypes, B, F, G, GH,LD, LE, MICH,MO,NY, P, S, U, US). Fig. 63 Herb,withthe stemsubterranean, partly? tuberous,(1-)4 8 mmthick,puberulous withuncinate hairs; internodesshortor a few long. Leaves in spirals, laminaoblongtoellipticorsometimes rosulate; toovate orto suborbicular, 3.9-5 x 1.5-4cm,chartaceous; apex basecordate; rounded; tocoarselyand marginsubentire ratherfaintlycrenate,oftenalsodenticulate; uppersurfacescabrouswithdense,minute,conicalhairs,sometimesintermixed withlongerstiffhairs;lowersurface scabridulousto smooth,puberulouswith curved, straight,and/oruncinatehairs,sometimesalmostconfinedto themainveins;venationpinnate;lateralveins ormostlyafterbranch(4-)6-9, faintlyloop-connected, atthemargin; ingterminating petiole1.5-6(-8)cmlong, withcurvedandoften? retrorse puberulous orstraight hairs;stipulestriangularto ovate,to 0.35 cm long, puberulous. greenorthereceptacle with Inflorescences themarginandthefloweringface(partly)purple;peduncle2.5-12(-18) cmlong,puberulous withshortto minute,straight,curved,and/oruncinate,often? retrorsehairs;receptacleeccentrically attached, discoid, in outline,ca.0.5-1.5 cmdiam.,some(sub)orbicular timesellipticand1.5-2 x 1-1.2cm,theoutsidepuberulouswith? retrorseuncinatehairs,towardsthemar-

(F, MICH,MO,NA, NY); Mun.Cd.Valles,RanchoPago Pago,W of Chantal,nr.jct. with Rio Mesillas and Rio Mico, 15 May 1981, Fryxell et al. 3445 (F, TEX); 18 km S of Valles,2 Jul 1940, Hitchcocket al. 6906 (NA, US); 23 mi S of Valles, 28 Nov 1946, Johnston s.n. (TEX);nr. Nuevo Morelos, 8 Jan 1948, Kenoyeret al. 3731(GH,MICH);Rascon,Aug 1911,Purpus5311 (BM, F, GH,NY, US), 5779 (MO);San Isidro,40 km NW of Valles, 2 May 1959, Rzedowski10398, 12 km SE of Tamasopo, 11 Sep 1967, Rzedowski24428 (MICH). TAMAULIPAS:Sierrade San Carlos,La Sardina,14 Aug 1930,Bartlett10955 (F, LL, MICH,U, US); LasYucas, 40 km NNW of Aldama, 19 Jul 1957, Dressler 1889 (MICH, MO). VERACRUZ: Nr. Tantoyuca, 1858, Ervendberg123 (G, GH, GOET,P); Mun. Chalma,3 km NE of Huejutla,22 Jun 1980, Nee 18431 (F). Dorstenia excentrica is a ratheruniform species, showingsimilaritiesto D. brasiliensis.CollectionReko 4907 (fromGuerrero)is distinctin the presenceof rela-

tivelylonghairsontheuppersurfaceof thelaminaand therelativelylargeandellipticreceptacle. 34. Dorstenia fawcettii Urban,Ark. Bot. 22A(10): 2. 1929;Grisebach, Fl. Brit.W.I. 154. 1864,subD. cordifolia; Fawcett& Rendle,Fl. Jamaica3: 44. 1914, sub D. cordifolia; Rossberg,Notizbl.Bot. Gart.Berlin-Dahlem12: 172. 1934;Adams,Flowering Plants of Jamaica220. 1972. Type. Jamaica. Trelawny,nr. Troy,28 Jun 1904, Harris 8690 (holotype, B; isotypes, NY, P). Fig. 64 Dorstenia marginata Urban & Ekman,Ark. Bot. 22A(10): 5. 1929; Rossberg,Notizbl. Bot. Gart. Berlin-Dahlem12: 173. 1934. Type. Haiti. Mas-

TAXONOMIC TREATMENT

219

Fig. 63. 1. Dorsteniaexcentrica.Habit(Chase7474).2. D. belizensis.Habit(Proctor35982). [By E. M. Hupkens van der Elst, Utrecht.]

220 sif de la Hotte, Morne Delcour, nr. Pestel, 27 Aug 1927, Ekman 8990 (holotype, S; isotypes, B, IJ). Dorstenia haitensis Rossberg, Notizbl. Bot. Gart. Berlin-Dahlem 12: 173. 1934. Type. Haiti. Massif de la Hotte, Tiburon, Tiburon R., 24 Aug 1928, Ekman 10566 (holotype, B; isotype, S). Herb, with the stem rhizomatous, 1-3.5 mm thick, puberulous to hirtellous with ? retrorse, straight and/ or uncinate hairs; intemodes long and short, the stem parts with short intemodes sometimes tuberous. Leaves in spirals, ? rosulate; lamina basally attached, broadly ovate to (sub)cordiform 1-7 x 1-1.5 cm, (thinly) chartaceous; apex subacuminate to obtuse, sometimes rounded; base cordate, sometimes with overlapping lobes; margin subentire to crenate or dentate to sublobate; upper surface with sparse long hairs, often also with minute or short hairs, smooth or ? scabrous, lower surface minutely puberulous with uncinate hairs and hirtellous to subhirsute with straight and uncinate hairs; venation pinnate; lateral veins 3-7 pairs; petiole 1-14.5 cm long, puberulous to hirtellous to subhirsute with straight and uncinate, whitish to brownish hairs; stipules triangular, 0.15-0.5 cm long, uninervate, puberulous to hirtellous. Inflorescences pale green or with at least the margin ofthe receptacle purplish; peduncle 1.28 cm long, puberulous to hirtellous with straight and uncinate hairs; receptacle centrally to slightly excentrically attached, discoid, suborbicular in outline, 0.3-1.8 cm diam., the outside puberulous to hirtellous, the fringe to 3 mm broad, with sparse, minute, swollen hairs; bracts in 1(-2) rows (almost) confined to the margin, all or most of them radiating and on to 2 mm long triangular appendages of different length, ovate, to 1 mm long, shorter and longer ones ? clearly alternating, puberulous to hirtellous, sometimes also with 1-2 rows of minute broadly ovate to semiorbicular bracts on the fringe; staminate flowers mainly peripheral, a few to several among the pistillate ones; tepals 2; stamens 2; filaments longer than the perianth; stigmas mostly equal in length, 0.3-0.5 mm long, usually dark purple to black. Endocarp body ca. 1 mm diam., tuberculate. Distribution (see Fig. 81.1). In Jamaica and Haiti; mostly in crevices of limestone rocks, in shaded and moist places; to 800 m. Representative specimens examined. JAMAICA. St. Andrews, Soyo Falls, Driver's R., 31 Jan 1980, Berg 990 (U); Trelawny, Cockpit Country, Oxford, 13-18 Sep 1906, Britton 427 (F, NY); Trelawny, Trou, 15 Jul 1963, Crosby et al. 770 (F, GH, MICH, NY), 9 Dec 1904, Harris 8808 (BM, F, NY); Hanover, nr. Dolphin Head, 24 May 1906, Harris 9266 (BM F, K, NY, US); St. Elizabeth,Jones R., 25 Apr 1961, Kramer 1740 (U); Portland, nr. Hart-

FLORANEOTROPICA ford,nr. Priestman'sR., 9 Jun 1904, Maxon2549 (GH, NY,US);St. Andrews,nr.Kingston,19Aug 1928,Orcutt 2415 (BM); Trelawny,Windsor(Caves),23 Aug 1955, Powell136 (GH,IJ,MICH);Westmoreland, nr.RatTrap, 9 Apr 1961, Proctor22161 (GH, IJ); St. Andrews,SW of Ecclesdown,1 Sep 1962, Proctor22700 (IJ, MICH, NY, U); St. Catherine,nr.Croft'sMountain,6 Nov 1962, Proctor22779 (IJ);St. Ann, DouglasCastleDistrict,23 May 1965,Proctor26411 (GH,IJ,LL,MICH),Trelawny, NNWofAccompong, 27Apr1956,Stearn951(A,BM,IJ). HAITI. Withoutlocality, Swartzs.n. (BM, M, S). Massifde la Hotte,PetitTroudes Nippes, 12 Jan 1926, Ekman5408 (IJ, S, US); Massifde la Hotte,Jeremie,3 Jul 1928, Ekman10210 (S). Dorsteniafawcettii,D. petraea, andD.jamaicensis constitute a group of very closely related species. The material from eastem Jamaica is slightly different in the smaller receptacles with a narrower fringe. It matches the material from Haiti. Among the collections of D. fawcettii there is one (in P) mentioning on the label Purdie as the collector and Santa Marta (Colombia) as the provenance; a mistake has probably been made. Dorstenia cordifolia Lamarck (1786), based on material from Brazil, has been incorrectly applied to this species by Swartz (1788) and this name has generally been used with Swartz incorrectly as the author.

35. Dorstenia petraea Grisebach, Cat. P1. Cub. 58. 1866; Bureau, in De Candolle, Prodr. 17: 265. 1873; Rossberg, Notizbl. Bot. Gart. Berlin-Dahlem 12: 174. 1934; Sauget&Liogier, Fl. Cuba 2: 55. 1951. Type. Cuba. Oriente: San Juan de Buenavista, 24 Nov 1860, Wright2277 (holotype, GOET; isotypes, B, GH, K, P). Fig. 64 Herb, with the stem rhizomatous, 1.5-3 mm thick, puberulous to hirtellous with straight and uncinate hairs; intemodes short and long. Leaves in spirals, ? rosulate; lamina (sub)peltate, (sub)ovate to elliptic or oblong, sometimes to suborbicular, 1-6.5 x 0.5-3.5 cm, thinly chartaceous; apex obtuse; base (sub)cordate to rounded; margin subentire to ? irregularly and faintly crenate to dentate; upper surface puberulous to hirtellous to subhirsute, smooth to scabrous; lower surface hirtellous with straight hairs; venation pinnate; lateral veins 3-6 pairs; petiole 0.5-9 cm long, puberulous to hirtellous to subhirsute with straight white to brownish hairs; stipules triangular, 0.05-0.15 cm long, uninervate, puberulous. Inflorescences greenish or with at least the margin ofthe receptacle purplish; peduncle 1-9 cm long, puberulous to hirtellous to subhirsute with straight hairs; receptacle centrally attached, discoid, (sub)orbicular in outline, 0.4-1.2 cm diam., the outside puberulous to

221

TAXONOMICTREATMENT

Fig. 64. 1. Dorsteniafawcettii.Habit(Crosbyet al 1182). 2. D. petraea.Habit(Ekman7285).3. D. jamaicensis. Habit(Ekman15001). 4. D. flagellifera. Habit(Ekman8635). [By E. M. Hupkensvan der Elst, Utrecht.]

thefringeto 1mmbroad,with hirtellousto subhirsute, sparse,minute,(sub)globosehairs;bractsin 1 (or 2) all or mostof themradiating, row(s),(sub)marginal, ovate,to 0.6 mmlong,longerandshorterones? distinctlyalternating,puberulousto hirtellous(to subandamongthe hirsute);staminateflowersperipheral longerthan pistillate ones;tepals2;stamens2;filaments stigmasequalin length,ca.0.2 mmlong, theperianth;

whitish (?). Endocarp body subglobose, ca. 1 mm

to smooth. diam.,slightlytuberculate Distribution(see Fig. 81.2). Confinedto eastern Cuba;on steeprocks,alongstreams,in shadedplaces; atlow elevations. S of Bayamo, CUBA.ORmENTe: examined. Specimens beMay1951,Bro.Clemente7594 (GH);SierraMaestra,

FLORANEOTROPICA

222 tween Rio Oro and Rio Bayamo,6 May 1916, Ekman 7285 (F, K, S, US); Nagua,at Rio Yara,5 Jul 1922,Ekman14172(NY,US);SanFelipe,nr.wellsof RioManaca, 1 Aug 1922, Ekman14702 (NY, US); Pico Turquino Loma de Quintin,Nagua, Jul 1922, Bro. Leon 10778 (NY);SierraMaestra,Rio Buey,29 Oct 1941,Mortonet al. 3824 (US); Rio Narou, S of Pico Turquino,31 Jul 1935, Roig et al. 6580 (NY); SierraMaestra,N of La Bayamesa,S of Pablo de Yao, 16 Aug 1951, Webster 4073 (GH, MICH). Dorstenia petraea is a ratheruniform species, apparentlyvery close to D.fawcetti. Itmainlydiffersfrom the latterin a usuallypeltateandthinnerlamina,which may show transitionsto a non-peltatelamina.

36. DorsteniajamaicensisBritton,Bull.TorreyBot. Club35: 567. 1908;Rossberg,Notizbl.Bot.Gart. 12:175.1934;Fawcett&Rendle,Fl. Berlin-Dahlem P1.Jamaica 3:44. 1914;Adams,Flowering Jamaica nr.Mandeville, Manchester: 220.1972.Type.Jamaica 22 Sep 1908, Harris & Britton 10607 (holotype,

NY;isotypes,B, BM, F, K, P, US).

Fig. 64

Specimensexamined.JAMAICA.Withoutlocality, Wolles.n. (GH).Manchester,nr.Mandeville,15-24 Feb 1910, Brown 236 (NY, US), 21-23 Sep 1908, Britton 3737

(GH, NY), 25 Feb 1956, Proctor 11592 (BM, IJ, US). Dorsteniajamaicensisis reminiscentof D. roigii. It differs from the latter in the radiate bracts, arranged almost in a single row. Although the inflorescences are very similar to those of D. fawcettii, the species seems to be less closely related to D.fawcettii than to D. petraea.

37. Dorstenia flagellifera Urban & Ekman, Ark. Bot. 22A(10): 8. 1929; Rossberg, Notizbl. Bot. Gart. Berlin-Dahlem 12: 176. 1934. Type. Haiti. Massif de la Hotte, Morne Rochelois, nr. Miragoana, 18 Jul 1927, Eknan 8635 (holotype, S; isotypes, B, GH, P). Fig. 64

DorsteniahotteanaUrban& Ekman,Ark. Bot. 22A (10): 6. 1929;Rossberg,Notizbl. Bot. Gart.Berlin-Dahlem12: 175. 1934.Type.Haiti.Massifde la Hotte,nr.Baguette,30 Sep 1926,Elrnan7084 (holotype,S; isotypes,A, C, G, GH, IJ, U, US).

0.5-5 mmthick, Herb,withthestemrhizomatous, mostinternodes short,thestem retrorsely puberulous; Leavesin partswithshortintemodesoftent tuberous.

Herb, with the stem rhizomatous, creeping, 1-3.5 mm thick, hirtellous; intemodes predominantly long, the stem parts with short internodes ? tuberous. Leaves spirals, rosulate;lamina peltate, narrowlyto broadly in spirals, spaced to rosulate; lamina peltate, (broadly) ovate,1.5-5.5 x 1.3-4 cm,ratherfirmlychartaceous; ovate, 1.5-5 x 1-3 cm, chartaceous; apex acute to baseusuallyrounded, to(sub)acute; someapexrounded rounded; base cordate to rounded; margin subentire to timessubcordate;margin(coarsely)crenateor subentire torepand; uppersurfacewithminuterigidhairs,t sca- (finely or coarsely) crenate or crenate-dentate; upper surface sparsely, but near and on the margin more brous,on the main veins minutelypuberulousandhirdensely, puberulous to hirtellous or subhirsute; lower venation tellous;lowersurfaceminutelypuberulous; pinnate;lateralveins 4-6 pairs;petiole1.3-13.5 cm surface hirtellous with rather soft straight and uncinate andminutelypuberulous tohirtellous hairs; venation pinnate; lateral veins 4-6(-7) pairs; petilong,? retrorsely withstraightanduncinatehairs;stipulestriangular, to ole 1.5-17 cm long, puberulous to hirtellous, mainly 0.2 cm long, uninervate,puberulousto hirtellouswith with straight hairs; stipules triangular, to 0.15 cm long, straight and uncinate hairs. Inflorescences reddish- uninervate, puberulous to hirtellous. Inflorescences purple;peduncle1-7.5 cm long,i retrorselyand mi- greenish or at least the margin of the receptacle pursometimes cen- plish; peduncle 1.5-9 cm long, puberulous to hirtelhirtellous; receptacle nutelypuberulous, inoutline,0.4-0.8 lous; receptacle centrally attached, discoid, (sub)orbicdiscoid,(sub)orbicular trallyattached, cmdiam.,theoutsidesparsely andminutely puberulous, ular in outline, 0.4-1.1 cm diam., the outside puberulous thefringeto 0.5 mmbroad,minutelypuberulous with to hirtellous, the fringe to 0.5 mm broad, with minute club-shaped hairs;bractsin 1(or2) row(s),marginal, club-shaped hairs; bracts in (2-)3-4 rows on the marmostof themradiatingoron shortmarginalappendages, gin and the fringe or also below the margin, appressed ovateto lanceolate,to 1mm long,shorterandlongerones or the marginal ones i radiating, lanceolate to ovate, to ? distinctlyaltemating, flowers 1 mm long, puberulous; staminate flowers intermixed puberulous; staminate mainlyperipheral,a few amongthepistillateones;tepals with the pistillate ones; tepals 2; stamens 2; filaments theperianth; stigrnas longer than the perianth; stigmas ? unequal in length, longerthan 2;stamens 2;filaments equalin length,0.2-0.3 mmlong,whitishorpurplish 0.2-0.3 mm long, whitish (?). Endocarp body 0.6-0.7 (?). Endocarp body ca. 1mmdiam.,tuberculate. mm diam., distinctly tuberculate.

Distribution (see Fig. 81.7). In Jamaica,only knownfromthesurroundings of Mandeville; in crevicesof limestonecliffs;atlow elevations.

Distribution (see Fig. 81.6). In Haiti, known only from the Massif de la Hotte; on steep rocks, in shaded places; to 400 m.

TAXONOMIC TREATMENT

Specimensexamined.HAITI.Massifde la Hotte,nr. Miragoane,MorneRochelois,24 Jul 1926,Ekman6488 (C, IJ, K, S, U, US), Jul 1927, Eyerdam196 (F, GH, MO, US).

223

Dahlem12:175. 1934;Sauget&Liogier,Fl.Cuba 2:55.t.20. 1951.Type.Cuba.PinardelRio:Mogote de la Jagua,17 Jun1929,Roig&Ascuy2547(holotype,NY). Fig. 65

The species is ratheruniform. It is reminiscentof Herb,withthe stemrhizomatous, 4-8 mmthick, D. nummulariain the inflorescences(shapeandbracts) puberulous to hirtellouswithstraighthairs;intemodes andflowers (unequalstigmasandshortstyle). It is dis- short.Leavesin spirals,rosulate; laminapeltate,ovate, tinctin the predominantlylong intemodes. In the veg- sometimes tosuborbicular subcordiform orsubreniform, etativepartsD.flagellifera largelymatchesD.fawcettii. (3-)5-10 x (2-)3-7.5cm,chartaceous tosubcoriaceous; 38. Dorstenia belizensis C. C. Berg, Proc. Kon. Ned. Akad.Wetensch.,Ser.C, Biol. Med. Sci. 85: 38, t. 4. 1982.Type.Belize.ToledoDistrict,nr.SapoteCamp, ca. 1Okm W of Medina Bank, 23-27 Apr 1976, Proctor35982 (holotype,IJ;isotype,U). Fig. 63 Herb, ? procumbent,with the stem ascending, ca. 2 mm thick,puberulousto hirtellouswith straighthairs; internodes 0.5-1 cm long; uncinate hairs lacking. Leaves in spirals,spaced;laminapeltate,ellipticto oblong to (sub)obovateor sometimesto suborbicular,1.56 x 0.8-3.3 cm, firmly chartaceous,often variegated; apexroundedor sometimesmucronulate;baserounded (to emarginate);marginsubentireto irregularlycrenate or denticulate;uppersurfacepuberulous,scabridulous; lower surfacehirtellousto subtomentose;veins plane above, (rather)prominentbeneath;venation pinnate; lateral veins 4-6 pairs, ? distinctly loop-connected; petiole 1.5-7 cm long, hirtellousto puberulous;stipules narrowlytriangularto almostsubulate,0.1-0.2 cm long, uninervate,puberulous.Inflorescencesgreenish,with the flowering face facing downwards;peduncle 1-2.5 cm long, puberulous;receptacle almost centrally attached,broadlyturbinateto almostdiscoid,(sub)orbicularin outline, 0.5-0.8 cm diam.,the outside (sparsely) puberulous,the fringe to ca. 0.3 mm broad;bractsin ca. 3 rows on the marginandon the fringe, appressed, ovate to triangular,to 0.5 mm long, puberulous;staminateflowersamongthe pistillateones;tepals2; stamens 2; filamentsshorterthanthe perianth;stigmas(almost) equal in length, 0.1-0.2 mm long. Endocarp body ca. 1.5 mm diam.,distinctlytuberculate.

oralmostrounded; apexacutetoveryshortlyacuminate baseroundedto cordate;margin? irregularly dentate to crenateorsubentire; uppersurfacepuberulous with rigidhairs,scabrousto scabridulous; lower surface ontheveinstohirtellous withstraight puberulous, hairs, alsowithnumerous, minute,whiteto brownish,(sub) lateral globose,pluricellularhairs; venationpinnate; veins 3-6 pairs;petiole(1.5-)6-27 cm long,puberulous to hirtellouswith straightanduncinatehairs;stipules broadlyovateto semiorbicular, 0.1-0.25 cm long, withatleastthe uninervate, puberulous. Inflorescences orthereceptacle purplish; peduncle 4-7 cmlong, margin to hirtellouswithstraighthairs;receptacle puberulous inoutline,0.6centrally attached, discoid,(sub)orbicular 1.2cmdiam.,oftenslightlylobed,theoutsidesparsely andminutelypuberulous to subglabrous, thefringeto 1 mmbroad;bractsin ca. 3 rows,on andbelowthe marginandon the fringe,appressed,ovate,to 1 mm staminate flowlong,sparsely andminutely puberulous; ersamongthepistillateones;tepals2, almostfree,with denseminutegloboseto club-shaped hairs;stamens2; filamentslongerthantheperianth; stigmasunequalin length,0.1-0.2 mmlong,whitish(?).Endocarpbody ca.2 mmdiam.,tuberculate. Distribution(see Fig. 81.7). In Cuba(Pinardel Rio);onrocksin shadedplaces;atlow elevations. Specimensexamined. CUBA. PINAR DEL Rio: Sierra Guacamayas,Mogote de la Baliza, 7 Nov 1923, Ekman17973 (F, K, NY, S, U, US). Dorstenia roigii is very close to D. nummularia,

anddiffersfromthelattermainlyin beingmorerobust andhavingdenselypuberulous perianths.

Distribution (see Fig. 81.7). In Belize, only known fromthe type locality;in crevicesof limestonecliffs; at low elevation.

40. DorstenianummulariaUrban& Ekman,Ark. Bot.22A(l0):7. 1929;Rossberg, Notizbl.Bot.Gart. Dorstenia belizensis is a distinct species, probably Berlin-Dahlem 175. 12: &Liogier,Fl. 1934; Sauget relatedto the group of taxa with peltate leaves, comLas Lomas Cuba 2: 55. 1951. Cuba. Villas: Type. prisingD. roigii, D. nummularia,D.flagelligera, and 4 de nr. Rio Jul Ekman Siguanea, Navarro, 1922, D. peltata. 13888(holotype,S; isotypes,B, NY). Fig. 65 39. Dorstenia roigii Britton, Bull. TorreyBot. Club 51: 9. 1924; Rossberg, Notizbl. Bot. Gart.Berlin-

DorsteniaekmaniiUrban,Ark.Bot. 22A(10):7. 1929; Rossberg,Notizbl. Bot. Gart.Berlin-Dahlem12: 175. 1934. Type. Cuba. Santa Clara:Trinidad

224

FLORA NEOTROPICA

Fig. 65. Habitsof Dorstenia species. 1. D. roigil (Ekmnan17973). 2. D. nummularia (L. B. Smithet aL 3376). 3. D. peltata (Ekman1S472). 4. D. erythrandra(Ekmnan 603S). [By E. M. Hupkensvan der Elst, Utrecht.]

Mtns., Rio Hanabanilla,14 Feb 1924, Ekman often ? revolute; upper surface with ratherlong stiff 18463 (holotype,B; isotypes, B, G, NY, S). hairs, scabrous to smooth; lower surface puberulous Herb, with the stem rhizomatous, 3-5 mm thick, to hirtellouswith straightanduncinatehairs;venation minutelypuberulouswith straightand uncinatehairs; pinnatewith 2-5 pairs of lateralveins, or almostradimost internodesshort,the stem partswith shortinter- ate; petiole 0.6-14 cm long, retrorsely and minutely nodes often ? tuberous. Leaves in spirals, rosulate; puberulousto hirtellouswith straightanduncinatehairs; laminapeltate,orbicularor ovate, 0.5-5 x 0.5-3.5 cm, stipulestriangular,to 0.15 cm long, uninervate,sparsely chartaceous;apex roundedto obtuse;base roundedto and minutely puberulous.Inflorescences greenish or truncate;margin subentireor rathercoarsely crenate, with at least the marginof the receptaclepurplish;pe-

TAXONOMICTREATMENT duncle 1.5-5.5 cm long, retrorselyandminutelypuberulous; receptacle(almost) centrallyattached,discoid, (sub)orbicularin outline,0.2-1.1 cmdiam.,oftenslightly lobed,the outsidesubglabrousor sparselyandminutely puberulous, the fringe to 1 mm broad, with sparse minute club-shapedhairs;bractsin ca. 3 rows on and below the marginand on the fringe, appressed,ovate, to 1 mm long, sparselyandminutelypuberulous;staminate flowers mainly peripheraland a few to several among the pistillate ones; tepals 2, subglabrous;stamens 2; filamentslongerthanthe perianth;perianthof thepistillateflowers(almost)glabrous;stigmasunequal in length,0.1-0.2 mm long, whitish(?). Endocarpbody ovoid to subglobose, ca. 1-1.5 mm diam., smooth to slightly tuberculate. Distribution (see Fig. 81.1). In Cuba(SantaClara andLasVillas);in creviceson (wet) limestonecliffs, in deep shade;at low elevations.

225

rals,rosulate; laminapeltate,orbicular to broadlyovate, 1-5.8 x 1-4.7 cm,chartaceous; apexroundedto subacute;baserounded, sometimes truncate orsubcordate; marginsubentireorrepandor (coarsely)crenate;uppersurface minutely puberulous tohirtellous withrather soft, straightto curvedoruncinatehairs,densernear themargin,smooth;lowersurfacepuberulous to hirtellous;venationpinnatewith2-4 pairsof lateralveins, sometimesalmostradiate; petiole1-16.5cmlong,minutelypuberulous ortohirtellous, sometimes subhirsute, with? retrorsestraightanduncinatehairs;stipulestriangular, to 0.2 cmlong,uninervate, puberulous. Inflorescencesgreenish(orpartlypurplish?); peduncle112.5 cm long, minutelypuberulousto hirtellous, sometimeswithretrorsehairs;receptaclecentrallyattached,cup-shaped to turbinate, in out(sub)orbicular line,0.3-1.1 cm diam.,theoutsideminutelypuberulous,thefringewithminuteclub-shaped hairs;bracts in 3-4 rows,on (orjustbelow)themarginandon the to 0.7 mm fringe,appressed,ovateto semiorbicular, toglabrous; long,minutelypuberulous staminate flowersintermixedwiththepistillateones;tepals2, with minutethinto club-shaped hairs;stamens2; filaments of thepistilusuallylongerthantheperianth; perianth lateflowerswitha tubular apexandminutethintoclubshapedhairs;stigmasunequalin length,0.1-0.2 mm long, whitish(?). Endocarpbodyca. 1.5 mm diam., tuberculate.

Specimensexamined.CUBA. LAS VILLAS: Nr. Las Vegasde Matagua,7 Dec 1929, Banghams.n. (IJ);Sierrade San Juan,Mina Carlotta,SW of Cumanayagua, 8 Jul 1941, Howard 5739 (BM, BR, F, GH, K, LE, MICH,MO,NY, 0, TEX,U, US), 11 Jul 1947, Woodet al. 7465 (IJ), 14 Jul 1953, Websteret al. 196 (GH,NY). SANTA CLARA: TrinidadMts., BuenosAires, Gavinas, Aug 1941, Howard 6452 (F, GH, MO, NY, U, US); TrinidadMts.,BuenosAires,Naranjo,10 Apr 1930,Jack 7884 (A, F, NY, S), 24 Jul 1930, Jack 8108 (A, F, NY, S); Lagunade Matagua,LasVegas,E of Cienfuegos,Jul Distribution(see Fig. 81.1). In the Dominican 1929,Bro.Le6net al. 13983(NY),7 Dec 1929,Bangham RepublicandeastemCuba;on steeprocks,in shadeor s.n. (IJ);TrinidadMtns.,BuenosAires,La Sabana,5 Aug in litteron forestfloor;atlow elevations. 1936, L. B. Smithet al. 3376 (S, US). The species is ratheruniforn andclosely relatedto Dorstenia roigii.

Specimensexamined.CUBA.ORIENTE:Baracoa, 22 Nov 1914,Ekman3580(G, K, NY,S), 3 Jun1860, 2225(B, BM,G, GH,LE,MO,P,S). Wright

DOMINICAN REPUBLIC. Puerto Plata, nr. Cabarete,21 Mar 1930,Ekman14561 (S), 27 Oct 1969, 41. Dorstenia peltata Sprengel,Neue Entdeck.3: 22. Liogier16511 (NY);Peninsulade Samana,Boca de Rio 1822; Schultes, Mant.3: 317. 1827; Bureau,in De San Juan,17 May 1930,Ekman15001 (C,GH, K, US); Candolle,Prodr.17:265. 1873;Urban,Symb.Antill. Peninsulade Samana,Loma Zaramagua,9 Jun 1930, 8: 166. 1921;Rossberg,Notizbl. Bot. Gart.Berlin- Ekman15248 (S, U); SamanaBay, Los Haitises,28 Jun Dahlem 12: 174. 1934. Type.DominicanRepublic. 1930, Ekman15472 (C, GH, K, LL, U, US), 14 Apr 1965,Jimenez5050 (NY), 19 Mar 1969, Liogier 14490 Withoutlocality,Berteros.n. (holotype,B; isotypes, (NY);SamanaBay,Bocadel Inferno,2 Feb 1928,Miller G, K, MO, P). Fig. 65 1034 (IJ, US); SamanaBay, San Lorenzo,30 Oct 1909, DorsteniacrassipesGrisebach, Cat.P1.Cub.58. 1866; Taylor95 (NY); Los Haitises, 25 Apr 1985, Zanoniet Bureau,in De Candolle, Prodr.17: 266. 1873; al. 34310 (BG, U). Rossberg,Notizbl. Bot. Gart.Berlin-Dahlem12: to berelatedtoD. roigii. Dorsteniapeltata appears 174. 1934. Type.Cuba.Oriente:nr. La Catalina, Itis distinct inthecup-shaped andthemarked receptacle 1860, Wright2224 (holotype, GOET;isotypes, towards orbicular leaves.Thecollection tendency Miller B, G, GH, LE, MO, NY, P, S). Herb,withthe stemrhizomatous,0.5-7.5 mm thick, minutelypuberulous;intemodesmostly short,the stem partswith shortintemodes ? tuberous.Leaves in spi-

1034differsfromtheothercollectionsinthereceptacle withits shortmarginalappendages.The collections madein Baracoa(Cuba)aredistinctby therelatively smallandslightlyangulatelamina.

226

FLORANEOTROPICA

42. Dorstenia erythrandra Grisebach,Cat. P1.Cub.

hairs;receptacle (almost) centrally attached,discoid, (sub)orbicularin outline, 0.4-1 cm diam., the outside puberulousto hirtellous, the fringe to 2.5 mm broad, Gart. Berlin-Dahlem 12: 175. 1934; Sauget & with sparse,minute, club-shapedhairs;bractsin (2-) Liogier,Fl. Cuba2: 56. 1951. Type. Cuba.Oriente: 3-4 rows on the marginand on the fringe, appressed, MonteVerde,3 Jun 1860, Wright2226 (lectotype, ovate,to 0.5 mm long, sparselypuberulousto glabrous, designated here, GOET;isolectotypes, G, GH, K, butthe largermarginalbractssetulose;staminateflowMO, P). Fig. 65 ers intermixedwith the pistillateones; tepals 2, almost Dorstenia confusa Britton,Bull. TorreyBot. Club free;stamens2; filamentsshorterthantheperianth;stig51: 9. 1924; Rossberg,Notizbl. Bot. Gart.Ber- mas equal in length, ca. 0.4-0.5 mm long, whitish (?). lin-Dahlem12: 176. 1934. Type. Cuba.Oriente: Endocarpbody0.8-1 mm diam.,distinctlytuberculate. Monte Verde, 3 Jun 1860, Wright2226 (holoDistribution(see Fig. 81.7). InCubaandHispaniola; type, GH; isotypes,G, GOET,K, MO, P). in crevices of steep limestone rocks in shaded usually Dorstenia domingensisUrban& Ekman,Ark. Bot. in litteron forest floor; to 1300 m. places, sometimes 3. Notizbl. Bot. 22A(10): 1929; Rossberg, Gart. Berlin-Dahlem 12: 171. 1934.Type.Haiti.Massif Speciemensexamined.CUBA. ORIENTE: mouthof des Matheux,Morne-a-Cabrits, Thomazeau,3 Jul RioYurmi,E of Baracoa,13 Jan 1956,Bro.AlainLiogier 1927,Ekman8547 (holotype,S; isotypes,GH, P). & Morton4079 = Morton& Bro. Alain Liogier 9029 DorsteniamultisquamaeUrban,Ark. Bot. 22A(10): (NY, US). 3. 1929; Rossberg,Notizbl. Bot. Gart. BerlinHAITI. Massif du Nord, Morne Bonpere, 30 Sep Dahlem 12: 174. 1934. Type. Haiti. Trouin,rd. 1925, Ekman 4954 (IJ, S); Massif du Nord, Morne to CarrefourFauche,10 Nov 1924,Ekman2438 Pedregal,6 May 1926Ekman6035 (C, G, K, IJ, LL, S, US); Massifdes Cahors,St. Michelde I'Atalaye,Gorge (holotype,B; isotype, IJ). DorstenianipensisUrban& Ekman,Ark.Bot. 22A- LaBoc, 8 Jun1927,Ekman8380 (S);Massifdes Matheux, 3 Jul 1927, Eyerdam19 (10): 5. 1929;Rossberg,Notizbl.Bot. Gart.Ber- Thomazeau,Morne-a-Cabrits, lin-Dahlem12: 172. 1934. Type.Cuba.Oriente: (F, GH, US); Ile La Gonave,10 Aug 1927,Ekman8882 SierraNipe, MacheteRivulet, 16 Sep 1922, Ek- (A, S), Aug 1927,Eyerdam298 (F,GH,NY,US);nr.Jean Rabal,27 Jan-9 Feb 1929, Leonardet al. 12772 (US). man 15123 (holotype,S; isotypes, B, NY, P). DOMINICAN REPUBLIC. Baoruca Mtns., nr. Dorstenia howardii Bro. Le6n, Revista Soc. Cub. Bot. 5: 77, cum. tab. 1949. Type.Cuba.Oriente: Aceitillar,11 Feb 1970,Bro. AlainLiogier13812 (NY), Sierrade Moa, 15 km SW of Companiade Moa 6 Nov 1969,Bro. AlainLiogier 16713 (NY); Barahona, mill, Jul 1941, Bro. Le6n & Howard 20155 = between Pedernales and Aceital, 8-12 Aug 1946, Howardet al. 8113 (BM, IJ, US). Howard4881 (holotype,GH; isotype, NY). Thespecieshasbasallyattachedleavesin Hispaniola. Herb, with the stem rhizomatous,0.3-4 mm thick, Leonard et al. 12722, a sterile specimen which Only minutely puberulous, predominantly with uncinate almost certainlybelongsto thisspecies,haspeltateleaves. hairs, sometimes glabrescent;most intemodes short, Most collections from Cuba have peltate leaves. The the stempartswith shortintemodes? tuberous.Leaves collection of type Dorstenia multisquamaeis slightly in spirals,rosulate;laminabasally attached,(broadly) ovate to cordiform or sometimes to subreniform,or differentin lackingdistinctlysetulose marginalbracts.

58. 1866;Bureau,in De Candolle,Prodr.17:265. 1873,as D. erythranta);Rossberg,Notizbl.Bot.

peltateand(angulate-)ovateto almostrhombiform,0.74.5 x 0.8-4 cm, chartaceous;apex roundedor to subacute;base (deeply) cordate,or if peltate,thenrounded to obtuse;marginsubentireto finely or coarselycrenate or repand;uppersurfacepuberulouswith minuterigid hairs, sometimes to hirtellous, usually scabridulous; lower surfacepuberulousto hirtellouswith straightand uncinatehairs on the main veins, more sparsely elsewhere;venationpinnatewith 2-6 pairsof lateralveins, to almostradiate;petiole0.5-9(-13) cm long,retrorsely puberulousto hirtellouswithstraightanduncinatehairs; stipulestriangular,to 0.3 cm long, uninervate,puberulous, predominantlywith uncinatehairs.Inflorescences with (always?) at least the margin of the receptacle purplish;peduncle 0.7-8(-16.5) cm long, retrorsely puberulous and hirtellous with straightand uncinate

43. Dorstenia caimitensis Urban,Ark, Bot. 22A(l 0): 4. 1929;Rossberg,Notizbl.Bot. Gart.Berlin-Dahlem 12: 172. 1934. Type. Haiti. Ile GrandeCaimite,24 Aug 1927, Ekman8955 (holotype, B; isotypes, C, G, GH, K, S, U). Fig. 66 DorsteniaasperifoliaRossberg,Notizbl. Bot. Gart. Berlin-Dahlem 12: 272. 1934.Type.Haiti.Massif de la Hotte,Dame-Marie,betweenTroud'Enfer and Faux Cap, 9 Aug 1928, Ekman10497 (holotype, B; isotypes,G, GH, S). Herb, with the stem rhizomatous, 2-7 mm thick, minutelypuberulous;most intemodes short,the stem partswith shortinternodes? tuberous.Leaves in spirals, rosulate;laminabasally attached,ovate to cordi-

227

TAXONOMIC TREATMENT

Fig. 66. 1. Dorstenia caimitensis. Habit (Ekman 10497). 2. D. crenulata. Habit (Wright 1447). 3. D. tuberosa.

Habit(Wright1446). 4. D. rocana. Habit(Ekman19110). [By E. M. Hupkensvan der Elst, Utrecht.]

form, 1-1.5 x 1-5.5 cm, chartaceous;apex roundedto basecordate,sometimesthelobes vety shortlyacuminate; marginfinelyorcoarselycrenateto (crenate-) overlapping; dentate or suberose; upper surface with minute rigid lowersurfaceminutelypuberhairs,usuallyscabridulous; ulouswith straightanduncinatehairs;venationpinnate; lateralveins 4-6; petiole 1.5-11 cm long, +retrorsely to 0.2 cm long,uninervate, stipulestriangular, puberulous;

puberulous. Inflorescences withthemarginof the receptacleandthefloweringface(partly)purplish;perecepduncle0.7-5 cm long, : retrorselypuberulous; discoid,(sub)orbicularinoutline, attached, taclecentrally 0.2-1.2 cm diam.,the outsidesparselyandminutely puberulous,the fringeto 1 mm broadwith minute swollenhairs;bractsin 2-3 rows,on themarginand on thefringe,oftena few belowthemargin,themar-

FLORA NEOTROPICA

228

Thespeciesis distinguished bytheobscuremarginal theothersappressed, ovate ginalonesoften? radiating, to 0.7 mmlong,sparselypuberulous bracts.Itis possiblyrelatedto Dorsteniaerythrandra. to semiorbicular, anda to glabrous;staminateflowersmainlyperipheral fewamongthepistillateones;tepals2; stamens2; filaP1.Wright.1:173. thantheperianth; mentslongerorshorter stigmas equalin 45. DorsteniatuberosaGriesbach, = n. s. 8: 173. 1861; 1860 Amer. Acad. Arts Mem. ca.0.2mmlong,oftenpurplish length, (?).Endocarpbody PI. Cat. Cub. 58. Grisebach, 1866; Bureau,in De ca. 1.5mmdiam.,tuberculate. Candolle,Prodr.17:264. 1873;Rossberg,Notizbl. Distribution(see Fig. 81.3). In Haiti;in crevices 12:172.1934;Sauget& Bot.Gart.Berlin-Dahlem of limestonecliffs,in shadedplaces;atlowelevations. Liogier,Fl. Cuba2: 54. 1951. Type. Cuba.Oriente: nr. MonteVerde,1859, Wright1446 (holotype, Specimens examined. HAITI. Jeremie,W of Trou GOET). Fig. 66 Cochon,9 May 1941, Bartlett17313 (GH,MICH,US); Ile GrandeCaimite,23 Aug 1927, Eyerdam314 F, GH; MO, NY, U, US).

Thisspeciesis ratheruniformandis distinguished on allparts. by theshortindumentum 44. Dorstenia crenulata Grisebach,P1.Wright.1: 173.

1860= Mem.Amer.Acad.Artsn. s. 8: 173. 1861; Grisebach,Cat.P1.Cub.58. 1866;Bureau,in De Candolle,Prodr.17:264. 1873;Sauget&Liogier, Fl. Cuba2: 54. 1951.Type.Cuba.Oriente:Monte 1447(holotype,GOET; Verde,1 Sep1859,Wright isotypes,B, G, GH,MO,K, P). Fig. 66

1-2.5 mmthick, Herb,withthestemrhizomatous, withstraightanduncinatehairs; minutelypuberulous mostintemodesshort,thestempartswithshortintemlamina odesoften+ tuberous. Leavesinspirals,rosulate; to subreniform, 0.7basallyattached, (sub)cordiform 2.2 x 0.9-2.8 cm, chartaceous; apexrounded,sometimestosubacute; sometimes thelobes basedeeplycordate, overlapping; marginsubentireto (coarsely)crenate; to hirtellous;lowersurface uppersurfacepuberulous withstraight anduncinate puberulous tohirtellous hairs; venationpinnate;lateralveins3-4 pairs;petiole0.57.5 cm long, puberulousto hirtellouswith retrorse to 0.15 straightanduncinatehairs;stipulestriangular, cmlong,uninervate, puberulous. Inflorescences greenish (?);peduncle0.6-2.3 cm long,retrorselypuberuloustohirtellous; receptacle centrally attached, discoid, in outline,0.3-0.7 cm diam.,the out(sub)orbicular sidepuberulous withrigidhairs,thefringe to hirtellous to 1mmbroad,minutelypuberulous; bractsin 1 (or2) row(s),almostconfinedto themargin,radiate,ovate, to 0.5 mmlong,longerandshorterones? clearlyalternating,puberulous to hirtellous; staminate flowersperipheral; tepals2; stamens2; filamentsshorterthanthe perianth; perianthof thepistillateflowerspuberulous withclub-shaped anduncinatehairs;stigmasequalin

0.5-2 mmthick, Herb, withthestemrhizomatous, withsparse,minute,rigidhairs;mostinternodes short, thestempartswithshortintemodesoften? tuberous. cordiLeavesinspirals, rosulate; laminabasally attached, form,0.8-2.2 x 1-2 cm, chartaceous; apexrounded; basecordate,sometimeswithoverlapping lobes;margin (coarsely)crenate;both surfaceshirtellousand venationpinnate;lateralveins2-4 pairs; scabridulous; hirtellous withstraight petiole0.6-5.5cmlong,retrorsely anduncinatehairs;stipulestriangular, ca.0.1 cmlong, withrigidhairs.Inflorescence: uninervate, puberulous to hirtelpeduncle2-5 cmlong,retrorsely puberulous length, 0.3-0.5 mm long, whitish (?). Endocarpbody anduncinate louswithstraight hairs;receptacle centrally ovoid,ca. 1-1.2 mmdiam.,(almost)smooth. in outattached, shallowlycup-shaped, (sub)orbicular line,0.5-0.7 cmdiam.,theoutsidepuberulous to hirDistribution (see Fig. 81.3). In Cuba (Oriente); tellouswithrigidhairs,themargincrenulate, thefringe found along stream(let)s;at low elevations. to ca. 0.5 mmbroad,puberulous; bractsin ca. 3 rows onthefringeandonthemargin, appressed, semiorbicular Specimens examined. CUBA. ORIENTE:Monte Verde,PinalCr.,9 Jul 1859, Wright1446 (GH);nr.Monte to ovate,to 0.6 mmlong,puberulous, marginalbracts Verde,24 May, Wright1446 (K), 10 Aug, Wright1446 obscure;stigmasalmostequal,ca. 0.3-0.4 mmlong. (BM);Yunquede Baracoa, 12 Jun 1860, Wright1446 Thematerial Endocarpbodytuberculate. didnotallow (GH);nr.Palenque,8 May, Wright1446 (= 739?);withof floralpartsandfruits. closerexamination out certainprovenance,Wright1446 (B, G, K, LE, MO, Distribution(see Fig. 81.2). In Cuba(Oriente); in forest;atlow elevations. alongstreamlets Specimens examined. CUBA.ORIENTE:PeniaPrieta Joa, 30 Dec 1953, Bro. Alain Liogier 3590 (GH).

NY, P, S, US).

46. Dorstenia rocana Britton,Bull. TorreyBot. Club 51: 8. 1924;Sauget& Liogier,Fl. Cuba2: 54. 1951.

TAXONOMIC TREATMENT

229

Type. Cuba.Las Villas: Rio Cupeyales,Banao Dorstenia cordifolia Lamarck,Encycl. 2: 317. 1786; Mtns., 5 Aug 1918, Bro. Leon & Roca 8092 (holoSwartz,Prodr. Veg.Ind.Occ.34. 1788;Willdenow, type,NY). Fig. 66 Sp. P1. 1(2):682. 1798; Sprengel,Syst.Veg. 3: 777.1826;Schultes,Mant.3:316. 1827;Grisebach, Dorstenia tricolor Urban& Ekman,Ark. Bot. 22A(10): Fl. Brit.W.I. 154. 1862;Bureau,in De Candolle, 8. 1929; Rossberg,Notizbl. Bot. Gart. BerlinProdr. 17: 265. 1873. Type. Anonymouss.n. Dahlem12: 174. 1934. Type. Cuba.Oriente:SierraNipe, nr. Pedra,30 Jun 1924, Ekman19110 in herbde Jussieu(Not "Amerique meridionale" (holotype,S; isotypes,B, G, GH, NY, P, US). traced).Sincethe publicationof Swartz(1788), materialfromtheGreaterAntilles,heretreatedas Herb,withthestemrhizomatous, 0.3-2 mmthick, D. to asD. corfawcettii,hasusuallybeenreferred minutelypuberulous withstraightanduncinate,some-

difolia Swartz.D. cordifolia has been associated timesretrosehairs;mostinternodes short,thestemparts withD. brasiliensisby Sprengel(1826). withshortintemodesoften? tuberous.Leavesin spirals,rosulate; laminapeltateorbasallyattached, broadly ovateto suborbicular, 1-2 x 1-2 cm,chartaceous; apex Dorstenialanei Howard& Briggs,J. ArnoldArbor. 34: 182. 1953. Type.Howard,Briggs, Kamb,Lane rounded,sometimesto obtuse;basein peltateleaves &Ritland69, Cuba.LasVillas:10 km S of Santa rounded,truncateto subcordate, in non-peltate leaves Clara (holotype,A). Identityuncertainbecauseof cordate;marginsubentire to (coarsely)crenate,sometheabsenceof inflorescences inthetypespecimen. timesslightlyrevolute; uppersurfaceminutelypuberulouswithrigidhairs,densernearthemargin,orhirtellous to subhirsutewith straighthairs;lowersurface Dorstenia opifera Martius,in Martius& Spix, Reise Bras.2: 787. 1828;Miquel,in Martius,Fl. Bras. withstraightanduncinatehairsor sparselypuberulous 169. 1853 =D. cayapia Vellozo var. opifera 4(1): hirtellous to subhirsute withstraight anduncinate hairs; Prodr.17:.1873; (Martius) Bureau,inDe Candolle, venationpinnatewith 1-3 pairsof lateralveins,to alChodat& Hassler,Bull.Herb.Boissier,Ser.2, 3: mostradiate;petiole1-4.5 cmlong,minutelypuberu350. 1903 = D. tubicinaRuiz & Pavonvar.opifera lousand/orhirtellouswithretrorsestraightandunci(Martius)Hassler,AnnuaireCons. Jard.Bot. natehairs;stipules(narrowly)triangular, to 0.15 cm Geneve21: 199. 1919.Type.Nottraced. long, uninervate,acuminate,puberulous.Inflorescences withthemarginof thereceptacleandfloweringface Lisbonense20. (partly)purpleorpurplish; peduncle0.7-4.5cmlong,? DorsteniavilellaPaiva,Farmacopea 1785, nomen,Mem. Hist. Nat. Acad. Real. Ci. andminutelypuberulousand/orhirtellous retrorsely Lisboa 1: 64. 1790, descr.; Sprengel, J. Bot. withstraightanduncinatehairs;receptacle slightlyeccentrically attached, deeplycup-shaped, (sub)orbicular (Schrader)2: 239. 1800;Sprengel,Syst.Veg.3: 77. 1826, as D. vitella; Schultes,Mant.3: 317. inoutline,0.2-0.4 cmdiam.,theoutsidepuberulous to 1827,as D. vitella;Miquel,in Martius,Fl. Bras. hirtellous,the fringeto 1.5 mmbroad,withminute 169.1853,subD. brasiliensis;Bureau,inDe 4(1): puberulous club-shaped hairs;bractsin ca.3 rowsbeCandolle,Prodr.17:263. 1873.Type.Nottraced. lowandonthemargin, appressed, ovate,to0.5mmlong, staminate flowersmostlyperipheral; puberulous; tepals 2; stamens2; filaments2 timesaslongastheperianth; Nomina nuda stigmasunequalin length,0.1-0.2 mmlong,whitish The not validly publishedDorstenia following (?).Endocarp body 1-1.2 mm diam., tuberculate. namesweremetwithin literature (andarenotmeantto Distribution(seeFig. 81.6).InCuba;on rocksor bepublishedhere): coralreefs,in shade;atlow elevations. D. aculeatasee Penna,Dic.Bras.Plant.Med.104. Dorsteniarocanais distinguishedby its deeplycup- 1941;ed.3: 151. 1946. D. bifida see Bot. Mag. t. 2760. 1827. inthefeaturesof the Theuniformity shapedreceptacle. seeMiquel,inMartius, D. cordifolia, nonLamarck, thepresenceof is thereasonto disregard inflorescence bothpeltateandbasallyattached leavesasanimportant Fl. Bras.4(1): 165. 1853;Bureau,in De Candolle, Prodr.17:262. 1873. in thegenus. character differentiating D. exaltatasee Miquel,in Martius,Fl. Bras.4(1): 161. 1853. D.fruticulosasee Endlicher, Cat.HortiVindob.1: DoubfIdandExcluded NamesandNominaNuda 241. 1842. Names of UncertainIdentity D. glabrasee Miquel,in Martius,Fl. Bras.4(1): Due to Missing or Incomplete Type Specimens 162. 1853.

230

D. glauca see Miquel, in Martius,Fl. Bras. 4(1): 162. 1853. D. hirta, non Desvaux, see Miquel, in Martius,Fl. Bras. 4(1): 163. 1853. D. hydrocotylesee Miquel,in Martius,Fl. Bras.4(1): 168. 1853. D. longipes see Miquel, in Martius,Fl. Bras. 4(1): 164. 1853. D. macahensis see Glaziou, Bull. Soc. Bot. France 59(Mem. 3): 640. 1913. D. ophidianasee Penna,Dic. Bras.Plant.Med. 105. 1941;ed.3: 152. 1946. D. palmata, non Schultes, nec Engler,see Miquel, in Martius,Fl. Bras. 4(1): 167. 1853. D. paradoxa see Bureau,in De Candolle,Prodr.17: 261. 1873.

FLORA NEOTROPICA

D. placentoidessee Lamarck,Encycl. 2: 317. 1786. D. reniformissee Miquel,in Martius,Fl. Bras.4(1): 168. 1853. D. rotundifolia see Arrudada CGnara,Discusso sobrea utilidadede intituicaodejavelins nas principais provinciasdo Brasil 45. 1810. D. volubilisseeWalpers, Ann.Bot. Syst. 1:736. 1849. Excluded name Dorstenia caulescens Linnaeus,Sp. P1. 121. 1753, cf. Poiret, Encycl. Suppl. 4: 554. 1816 = Procris quinquenerviaSprengel, Syst. Veg. 3: 846. 1826, cf. Weddell,Arch. Mus. Hist. Nat. 9: 339. 1856 = Pilea caulescens(Linnaeus)Urban,FeddesRepert. Spec. Nov. RegniVeg. 15: 163. 1918.

231

TAXONOMICTREATMENT

DORSTENIEAEP.P. ("BROSIMEAE")AND CASTILLEAE("OLMEDIEAE") Additions and Correctionsto Flora Neotropica Monograph 7 INTRODUCTION Botanicalexplorationin thelast25 yearshasinand creasedthenumberof collectionsconsiderably; severalcountries,suchas BoliviaandEcuador,and regionswhichwerefloristicallyverypoorlyknown at thetimeMonograph7 (Berg, 1972)wasprepared amore havebecomemuchbetterknown.Thisprovided accurateknowledgeof rangesof distributionof the taxatreated,butalsoaffectedtheconceptsof species delimitations.Moreover,manynew taxahavebeen

of newdatajustifiesthe discovered. Theaccumulation presentationof additionsandcorrectionsto Monograph7, as supplementto the newly revisedtaxa treatedabove.Thecitationof specimensin thepresent treatmentis moreor less completefor new taxa;for the othersthey largelyfill the gapsin therangesof distributionas knownwhenMonograph7 was written, andin some cases confirmthe presenceof the taxain certainregions.

DORSTENIEAE (continuedfromp. 229) thegeneratreatedasmembersof the"Brosimeae." Comprising Brosimeae (see Berg, Fl. Neotrop. Monogr. 7: 157228. 1972.) The tribeBrosimeaehas been includedin the tribe Dorstenieae (Berg, 1973, 1988, 1989b); see p. 000. The factthatsome woody speciesof Dorsteniafrom Africa have been described as species of Trymatococcus is anindicationof theclose relationshipbetween thegenerapreviouslyplacedin theBrosimeaeandthose in the Dortstenieae.

Brosimum 10. Brosimum Swartz(see Berg, Fl. Neotrop.Monogr. 7:161-208. 1972.) Morethan25 yearsof furtherbotanicalexploration has moderately affected the taxonomy of this genus. Only one new species, Brosimummultinervium,has been discovered,andB. utile subsp. longifoliumhas to be reinstatedas a species, raisingthe numberof Brosimumspecies to 15.

Forseveraltaxa,newrecordsindicaterangeextenpatterns sions.Theserecordsconfirmthedistribution in the withinthegenus:subg.Feroliais concentrated continent distinctly northern partoftheSouthAmerican withtheGuianaShield,andsubg.Brosimum associated hasa morediffuseandwiderdistribution. Somesynonymsareaddedto thoseof thetwosubalicastrum. Govaerts (1996)listed speciesof Brosimum B. refractum (Miquel,in Martius,Fl. Bras.4(1): 108. 1853)withoutindicationthatit is a nomennudum. material, in particular of Theincreaseof herbarium sterilespecimens,indicatesmoreclearlythatseveral thoseof subg.Ferolia,showmore species,inparticular beor less pronounced differencesin leaf characters tweenthejuvenileandadultstateof thetrees.In the juvenilestatetheleavesmaybe larger,theindumentum maybe sparserandlonger,andthenumberof lateral veinslarger.Juvenilefeaturesmayberetainedintothe reproductive state.

Revised key to the species of Brosimum 1. Stipules fully amplexicaul,connate. 13. B. multinervium 2. Lateralveins (30-)40-50 pairs................................................................... 2. Lateralveins to 32 pairs. 3. Laminadensely puberulousaroundthe areolesbeneath. around 4. Laminawith slightly prominentveinlets beneath,white to yellowish-puberulous the areoles .......................................................

14. B. potabile

around 4. Laminawith very prominentveinlets beneath,yellow to brownish-puberulous the areoles .............................................................

15. B. parinarioides

3. Laminasparselyhairy or glabrousaroundthe areolesbeneath. 5. Stipules0.3-0.7 cm long; laminawith a coarsereticulum,the veins prominentbeneath, more or less impressedabove,and the lateralveins loop-connectedfar fromthe margin ............................................................................................................................ 1 l. B. melanopotamicum

232

FLORANEOTROPICA

5. Stipules0.5-4.5 cm long; laminawith a fine reticulum,the veins (at least the smaller ones) slightly prominentto plane beneath,mostly plane above, and the lateralveins loop-connectedclose to the margin. 6. Margin of laminarepand.......................................................... 12. B. longifolium 6. Marginof laminaentire. 11. B. utile 7. Laminalongerthan 15 cm.......................................................... 7. Laminashorterthan 15 cm. 8. Laminausuallygradingto obovate,the base acute to obtuse;lateraland smallerveins slightlyprominentto plane beneath;indumentum mostly 9. B. rubescens ............................................ yellowish;epidermisof the petiolepersistent 8. Laminausuallygradingto ovate, the base truncateto obtuse;lateralveins or also smallerveins mostly prominent;indumentumwhite; epidermisof the petiole often flakingoff...................................................................... 11. B. utile 1. Stipulesfree, not fully amplexicaul. 9. Laminawith the midribimpressedabove; venationof the stipulesprominent,subflabellatefurcate;perianthof the staminateflower well-developed;trees dioecious...........................3. B. lactescens 9. Laminawith the midribprominentto plane or occasionallyslightly impressedabove; venation of the stipulesinconspicuous;perianthof the staminateflowerlackingor vestigial (except in B. costaricanum);trees monoeciousor dioecious. 10. Perianthof the staminateflower well-developed;trees dioecious;Costa Rica and Panama .4. B. costaricanum ........................................................................................................................................... 10. Perianthof the staminateflower vestigial or lacking;trees monoeciousor dioecious. 11. Lamina scabridulousbeneath. 12. Petiole 0.6-1.5 cm long; lateralveins 11-22 pairs;inflorescencesunisexual(or bisexual?);E Brazil......................................................... 7. B. glaziovii 12. Petiole 0.2-0.6 cm long; lateralveins 6-14 pairs;inflorescencesbisexual. 13. Perianthof the staminateflower lacking,substitutedby a perianth-likebract; E Brazil............................................................ 8. B. glaucum 13. Perianthof the staminateflowerpresent,? vestigial;widespread ...............5. B. guianense 11. Laminasmooth beneath. 14. Laminaand leafy twigs glabrousor nearlyso; laminagreenishto grayishwhen dry .1.B. alicastrum .................................................................................................................................... 14. Laminaand leafy twigs mostly distinctlyhairy;laminamostly brownishto grayish when dry. 15. Lamina? scabrousand/or? denselypuberulousabove, at least on the midrib; trees dioecious...................................... 2. B. acutifolium 15. Laminasmoothand glabrescentor sparselypuberulouson the midribabove; trees monoecious. 16. Laminawith the indumentumappressedbeneath..................................5. B. guianense 16. Laminawith the indumentumpartly? patent. 17. Laminausuallywith dentatemargin,mostly grayishwhen dry; in 6. B. gaudichaudii campo cerrado................................ 17. Laminausually with entiremargin,mostly brownishwhen dry; in rain forest................................ 5. B. guianense 1. Brosimum alicastrum Swartz (see Berg, Fl. Neotrop. Monogr.7: 169-173. 1972).

All collections from the Lesser Antilles (Carricou and St. Vincent) and from Trinidad are made from la. Brosimum alicastrum Swartzsubsp.alicastrum. materialin cultivation. Cloudsof pollen arereleasedfrom staminateinfloAdditionalsynonyms: rescences by touching them or by wind (as personally Ficus faginea Kunth& Bouche, Index Sem. Hort. observedin Florida).The peculiarshape of the anther Berol. (1846). 20. 1847. Urostigma fagineum of this subspecies-the thecae fused, forming a circu(Kunth& Bouche)Miquel,LondonJ. Bot. 6: 537. 1847.Type.Hort.Berol.(holotype,B; isotype,U). lar structureperpendicularto the filament anddehiscing laterally-can be regardedas an adaptationto wind Distribution. In CentralAmericafrom Mexico to pollination. The subspecies is (at least regionally) CostaRica andsome of the GreaterAntilles,Cuba,and gynodioecious ratherthandioecious as revealed in an Jamaica;in primaryor secondaryevergreen,semi-ev- autecological study in Veracruz(Mexico) by Peters ergreenor deciduous forest; to 1000 m; regionally or (1983, 1991).Treesproduceeitherpistillateinflorescences locally (as at Maya ruin sites) very abundant. or morphologicallybisexualones of which a relatively

TAXONOMIC TREATMENT

smallnumbersetfruit;withtheincreaseof thesizeof treesthe numberof inflorescencessettingfruitdecreases,andinlargertreesalmostnoneof themsetfruit. morphologically andinitiallyfunctionTreesproducing tendtobecomelargerthan allybisexualinflorescences thoseproducingpistillateinflorescences,a situation of pollenby wind. promotingtransport

233

Venthroughnorthem Distribution.FromPanama ezuelato GuyanaandRoraima(Brazil)andlargely alongtheAndesto BoliviaandAcre(Brazil),withi discontinuous distribution andregionallycommonor

VargasTorres,2 Feb 1991 (st), Gentryet al. 73017(MO, CantonGuayaquil,rd. to Salinas,km QCNE).GUAYAS: 15, 17Aug 1991(q fl-fr),Rubioet al. 1926(AAU,QCNE). LOJA:Rd. Alamor-Puyongo,km 20, 2 Apr 1980 (R fl), Harlinget al. 17767(BG,GB).Los Rifos:CantonVinces, Jaunecheforest, rd. Quevedo-Palenquevia Mocachi, km70,4 Oct1979(st),Dodson et al. 8825 (MO). MORONASANTIAGO:Nr. Taisha, 17 Aug 1985, Warush Juwa Pozo PetroleroMoreteRBAE.4(BG, QCNE).PASTAZA: cocha,70 km E of Puyo,4-21 Oct 1990 (d, Gudifioet al. 982 (BG, MO, QCNE); Curaray,Pozo Petrolero Villano2, 4-19 Aug 1993 (9 fl), Yiradoet al. 172 (BG, Los Encuentros,20 Sep QCNE). ZAMORA-CHINCHIPE: 1978 ($?), Montedesdeoca18-3 (MO,QAME,QCNE). Prov.Pachitea,Dtto. Honoria,rd. PERU.HUANUCO: to Tournavista, 28 Oct 1965(d'), Jenssen49 (MOL,NY). Hda.La Genoa, 1210 m, 4 JUNIN:Prov.Chanchamayo, Mar 1991 (st), Gentry et al. 73377 (MO). LORETO: Prov.Maynas,Rio Amazonas,2 km W of Indiana,15 Feb 1987(? fl + fr), Gentryet al. 55951 (BG, MO,NY). MADRE DEDios: Prov.Tambopata,HermosaChica, 15 Jun 1989 (st), Alexiadeset al. 842 (BG); Prov.Manui, ParqueNacional del Manui,Rio Manu, Cocha Cashu Station, 26 Aug 1984 (st), Foster 9915 (BG). SAN MARTiN:Prov.MariscalCareres,Dtto. TocacheNuevo, Quebradade Tanata,(?), SchunkeV 4414 (MO,MOL, Prov. Zarumilla,Bosque Nacional de US). TUMBES: Tumbes, between P.C. "El Coucho" and P.C. 22 Jul 1992(st), C. Diaz et al. 5061 (BG). "Campverde," 86 of rd. Prov.CoronelPortillo,22 km S of kmn UCAYALI: Pucallpa-TingoMaria,11 Feb 1981 (? fl + fr), Gentry et al. 31225 (MO).

abundant;in evergreen,semi-evergreenanddeciduous

BRAZIL. ACRE: Fazenda Bom Sossego, between

Use. Thissubspecieshasrecentlyreceivedample inconnection asa importance attention with(potential) seeds,as a forage food sourcedueto theprotein-rich asanelementof tree,andasa timbertree.Itis regarded (e.g.,G6mezancient,extant,andfutureagroforestry & SanchezMunioz, Pompaet al., 1987;Pardo-Tejeda 1982). 1977;Peters,1983,1991;Peters&Pardo-Tejeda, lb. Brosimum alicastrum Swartzsubsp.bolivarense (Pittier)C. C. Berg. Additionalsynonym: Mutisia33: Romero, Brosimumbolivarense(Pittier) 2. 1970,basedon HelicostylisbolivarensisPittier, Contr.U.S. Natl. Herb.20: 96. 1918,beingbasionymof B. alicastrumsubsp.bolivarense(Pittier) C. C. Berg.

forest;to 1000m.

Igarapedo CujubimandIgarapeJacamin,27 Sep-7 Oct 1985 (st), Campbellet al. 8606 (BG); Rio Tarauaca, Representative additional specimens examined. 20 Sep Praiado Carapana, COLOMBIA.AMAZONAS: Ca. 2.5 km NE of the mouth SeringalUniverso,Coloca9ao et RORAIMA: 1994 M. al. 860 NY). Silveira (BG, (v), of QuebradaEl Tigre, 22 Sep 1986 (st), M. SdnchezS. et al. 1440 (BG). BOLtVAR: Morales,Cgto. Arenal,rd. SEMAEcologicalReserve,Ilhade Maraca,1 May 1987 BuenaVista-Arenal,20-22 Apr 1966 (a), Foreroet al. (2 fl), Millikenet al. 161 (BG). BOLIVIA. BENI: Prov. Ballivian, rd. Yucumo443 (NY, U); Mun. Cartagena,Hda. Canalete, 12 Aug km 34, 7-14 Jul 1990 (? fl), D. N. Smith 1985 (? fl), Zarucchiet al. 4083 (MO).CESAR: 5 km W Rurrenabaque, of Manaure,13 Jan 1988(st), Gentryet al. 60708 (MO). et al. 14131 (BG, MO). LA PAZ:Prov.Abel. Iturralde, GUAJIRA: 15 km S of Mingueo,21 Aug 1986 (st), Gen- AltoMadidi,28 May 1990(st), Gentryet al. 70721(MO). try et al. 55422 (MO). MAGDALENA: Tucurinca, Dec PANDO:Prov.Madrede Dios, rd. Sena-Rio Beni, km 650 (US). RISARALDA: Rd. 10, 12 Oct 1991 (v), Becket al. 1991 (MO);Cobija,25 1947(V), Romero-Castaneda km 18, Hda.Corsega,12 Feb 1990 Oct 1977 (v), Meneceset al. 2076 (NY). La Virginia-Cerritos, (st), Silverstone-Sopkin et al. 5938 (MO). SUCRE: Estacionde Primatesde Coloso, 17 Nov 1981 (st), Gentry et al. 34813A(MO).VALLE:Nr.Bugua,5 May 1982 2. Brosimum acutifolium Huber (see Berg, Fl. (i fl-fr), Murphy351 (MO). Neotrop. Monogr.7: 173-176. 1972.) ARAGUA: Rd. Maracay-ElLim6nVENEZUELA. Ocumare,24 Sep 1984(?), Berget al. 1525 (NY). LARA: The subspecies that have been recognized can be Dtto. Jimenez,Paso de Angostura,28-31 Jul 1973 (st), maintained,althoughsome collections show intermeSteyermark et at. 107626 (MO, NY, U). SUCRE: diatefeatures.Onecollectioncouldnot be placedin one QuebradaEl Tigre, S of Fila La Baquete,21 Nov 1981 of the subspecies: BRAZIL. AMAZONAS: Mun. Sao (2 fl), Davidseet al. 19225 (MO).ZULIA: Dtto. Mara, 17 Nov Rio Gabriel da Ipanure, Uaupes, Cachoeira, 1 km W of CorpozuliaCampamentoCurichuano,2 Jun et 3235 al. Lima (NY). fl-fr), 1987 (? 1980 (? fl), Steyermark123176 (MO). Thiscollectionshows similaritiesto subsp.acutifoliW of San Mateo, Reserva ECUADOR. ESMERALDAS: Forestalde JardinBotanico, UniversidadTecnicaLuis urm,as in the dimensions of the lamina and the some-

FLORA NEOTROPICA

234

what revolute laminamargins.However, it is distinct in the coarsehairs,makingthe laminascabrousat both sides. The featuresof the indumentum,andthe factthat the collection has been made far outside the known range of subsp. acutifolium,could be indicationsthat this collection representsanothersubspecies.

BOLIVIA.BENI: Prov.VacaDiez, nr.Alto Ivon,21 Feb 1984 (st), Boom 4514 (MO, NY). PANDO: Prov. km 35, 25 Jun Nicolas Suarez,rd. Candelaria-Extrema, 1978 (v), Meneces 736 (NY); 54 km SW of Cobija, Triunfo,30 Jul 1988(v), R. T Penningtonet al. 62 (BG, F, NY). SANTA CRUZ: Prov. Guarayos, 8 km NW of Perseverancia, Rio Negro, 15 Sep 1990 (v), Nee 38798 (BG, MO, NY).

2a. Brosimum acutifolium Hubersubsp.acutifolium. No changes to be reported.

3. Brosimum lactescens (S. Moore) C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 177-179. 1972).

Distribution. In CentralAmerica from southern Mexico to CostaRica, andin SouthAmericaeastof the Andes in theAmazonBasin, the Guianaregion,northDistribution. In the lowerAmazon Basin (eastem westernVenezuela, and easternBrazil from Bahia to Amazonas, Pari, and Maranhao);in non-inundated Santa Catarina;in non-inundatedforest; to 1200 m; regionallyfrequent. forest;infrequent. 2b. Brosimum acutifolium Hubersubsp.interjectum C. C. Berg.

Representative additional specimens examined. BRAZIL. MARANHAO: Mun. Carutapera, 7 Nov 1986 (st), Balee et al. 3006 (BG, NY); Maraca9ume, 13 Jul 1958 (st), Froes 34484 (IAN). PARA:SerraBuritirama, Maraba, 10 Jul 1970 (v), Pires 12427 (IAN); Rio Itacaines,Aug 1970 (st), Pires 12849 (IAN);rd. BelemBrasilia, km 161-250, between Paragominas and Gurupi,19 Aug 1964 (v), Prance et al. 58940 (NY).

2c. Brosimum acutifolium subsp.obovatum (Ducke) C. C. Berg. Distribution. In the upperAmazonBasin, extending to Guyana;in non-inundatedforest; at low elevations; infrequent.

Representative additional specimens examined. COLOMBIA.AMAZONAS: Rio Cahuinari,12 km from the mouth, 10 Sep 1986 (st), M. Sanchez S. et al. 1115 (BG). CAQUETA:Araracuara,27 Nov 1991 (d), Restrepo

et al. 647 (MO);Quebradadel Quinche,Nov 1989 (2), Urrego G. et al. 1243 (NY). META: ParqueNacional NaturalTinigua, SerraniaChamusa,Apr 1991 (? fr), Stevenson301 (MO). VICHADA: ParqueNacionalNatural El Tuparro,ca. 9 km NE of El Tap6n,18 Mar 1985 (?), Zarucchiet al. 3742 (BG, MO, NY). VENEZUELA.AMAZONAS: Depto.Atures,45 km SE of PuertoAyacucho,11 May 1980 (?), Steyermark et al. 122242(NY,U). APuRE:Dtto.SanFernando, RioArauca, 16-17 May 1977 (st), Davidse et al. 13264 (MO, VEN).

Dtto. Paez, Cerro Riberon, between Rio Guapoand Rio Chiquito,8-10 Jun 1977 (c?), Davidse et al. 13735 (NY,U). ZULIA: Rio Catatumba,15 km NE Representative additional specimens examined. of rd., 29 Aug 1972 (?), R. F Smith V7139 (VEN). FRENCHGUIANA.Saul, 21 Jun 1988 (st), Gentry ECUADOR.NAPO:Canton Orellana, Parque Nacional Yasuni,Maxusrd., km 6-9, 12 Dec 1993 (a?), Grijalva et al. 63044 (MO, NY); OyapockR., Trois Sauts, 13 700 (BG, QCNE).PASTAZA:Rd. Coca-Auca,115 km S Aug 1989 (st), Prevost et al. 2784 (BG, NY). ECUADOR. EL ORO: Rd. Piedras-Marcabeli, of Coca,6 km S of Rio Tigiuino,1-9 Apr 1989 (st), Zak Arevalo et al. 10 (QCA).MORONA-SANTIAGO:CentroShuar et al. 4253 (BG, NY, QCNE). Prov. Pachitea, Dtto. Honoria, Rio Yukutais,12 Mar 1990 (? fr), Bennettet al. 4039 (BG, PERU.HUANUCO: Pachitea,nr. Miel de Abeja,26 Sep 1967 (?), Schunke QCNE);Estaci6nExp.El Padmi,12 Sep 1975(st),Little V 2178 (MO, MOL, NY). LORETO: Prov. Maynas, et al. 322 (LOJA,QCNE,US). NAPO:ReservaBiologica Manacamiri,10 Aug 1992 (st), Ayala6439 (MO);Dtto. JatunSacha, 8 km E of Misahualli,30 Dec 1987 (st), LasAmazonas,Rio Napo,nr. Sucusari,5 Mar 1991 (st), Gentry et al. 60191 (BG); Parque Nacional Yasuni, Pipoly 14609 (BG). MADRE DE DIos: Jct. of Rio Ainangu,ParqueNacionalYasuni, 1-15 Feb 1986 (st), andRio La Torre,22 Jul 1985(st), Gentryet Korninget al. 47511 (AAU,QCA,QCNE).PASTAZA:Rd. Tambopata al. 51106 (MO). UCAYALI:Nr. Pucallpa, 13 Sep 1980 Coca-Auca,110km fromCoca,10 km fromRio Tigiiino, 7 Jan 1989 (?), Palacios et al. 3367 (AAU, BG, MO, (2), Sousa 71 (MO, U). BRAZIL. ACRE: BR.317, km 68, 10 Jul 1991 (a'), NY, QAME); Pozo Petrolero Ramirez, 20 km S of Waltieret al. 42 (NY). AMAZONAS:Rio Uaupes, nr. Curaray, 21-28 Feb 1990 (?), Zaket al. 4982 (BG,MO, Campo de Aviagao, 20 May 1975 (st), L. Coelho 15 = NY, QCNE). SucuMBfos: Dureno, 27 Dec 1988 (st), INPA 49896 (BG); rd. Manaus-Porto Velho, BR.319, Gentryet al. 64248 (BG);ReservaFaunisticaCuyabeno, km 265, between Rio Igap6-aquand Rio Jutai, 12 Oct LagunaGrande,11 Mar1990(R fr),Balstev et al. 97121 1974 ($), Prance et al. 22829 (INPA,NY, U). MATO (AAU,BG, QCA,QCNE).ZAMORA-CHINCHIPE: Parque Rio Juruena, 2 Jul 1977 (a), M. G. Silva et al. Nacional Podocarpus, Jamboe Bajo, 1100 m, 3 Nov GROSSO: 3258 (NY). 1996 (st), J. L. Clark et al. 3277 (BG). MIRANDA:

235

TAXONOMICTREATMENT PERU. HuANuco: Prov. PuertoInca, Dtto. Yuyapichis, Reserva Forestal DANTAS, 16-31 Jan 1991 (? fr), Tello 1174 (MO). JUNIN: Prov. Tarma, Dtto. Pichinaki, Alto Cuyani, 12 May 1982 (st), ReynelR. 423 (MOL). LORETO: Prov. Maynas, Rio Nanay, Mishana, 23 Feb 1981 (v), Gentry et al. 31663 (AAU, NY); Prov. Loreto, Rio Marafion,Nauta, 9 Nov 1982 (st), R. Vasquezet al. 3456 (NY). MADREDE DIos: Prov. Manu, Parque Nacional del Manu, Cocha Cashu Station, 24 Jun 1984 (2 fl-fr + e'), Foster 9673 (MIO,NY). PASCo:Prov.Oxapampa,Iscozacin, 28 Aug 1986 (a), Parionaet al. 869 (MO,MOL);Prov. Pozuzo, San Cristobal, 23 Oct 1975 (? fr), Rios T et al. I (MOL). PUNO:Rio Candamo,mouth of Rio Guacamayo, 28 May 1992 (st), Gentry et al. 77280 (MO) BRAZIL. ACRE:Rio Branco, rd. AC.40, km 54, 22 Aug 1989 (?), Souza 144 (BG, NY). ESPIRITOSANTO: Linhares, Reserva da CRDV, 31 Aug 1983 (e), Folli 459 (BG), (v), Folli 671 (BG). GoiAs: Rio Javaes, 27 Jul 1978 (a'), Pires et al. 16239 (NY). MARANHAO: Mun. Mon9io, Rio Turia9u, 6 Feb 1985 (st), Balee et al. 488 (NY); Mun. Santa Luzia, 83 km W of Santa Inez, 29 Mar 1983 (9 fl-fr), E. L. Taylor et al. 1117 (BG). MATO GROSSO: Rio Aripuana, Salto dos Dardanelos, nr. Humboldt Center, 13 Oct 1973 (v), Berg et al. P18473 (MO, NY). ROND6NIA:Mineragao Campo Novo, 14 Oct 1979 (?), Zarucchiet al. 2701 (NY). RORAIMA:Mun. Alto Alegre, Ilha de Maraca, SEMA Station, 14 Jun 1986 (v), Hopkins et al. 752 (MO, NY); Rio Branco, Parana do Marara, 4 Mar 1977 (st), M. R. Santos 152 (NY). BOLIVIA. BENI: Prov. Vaca Diez, 20.8 Km E of Riberalta, 25 Sep 1981 (a), Solomon 6445 (MO); Cachuela Esperanza, 1923 (v), Meyer 326 (NY, U). Prov. Carrasco, Estacion Valle del Sajta COCHABAMBA: UMSS, 1989 (st), J. Le6n s.n. (BG, LPB). LA PAZ: Prov. Abel Iturralde, Upper Madidi, 24 May 1990 (st), Gentry et al. 70506 (MO); Prov. Sud Yungas, nr. Sapecho, 16 Nov 1991 (? fr), Seidel et al. 5602 (BG, LPB). PANDO:Puerto Oro, 74 km SW of Cobija, 19 Aug 1988

PANAMA. CHIRIQUi:Burica Peninsula, nr. border of Costa Rica, 24 Feb 1973 (st), Croat 22153 (AAU, MO, NY). DARIkN: Upper Rio Chucunaque, Tiotuma, 25 Jun 1868 (fl-fr), Duke 15814 (BG). COL6N: Pipeline rd., Rio Medonsa, 5 Sep 1977 (st), Berg 411 (U).

5. Brosimum guianense (Aublet) Huber (see Berg, Fl. Neotrop. Monogr. 7: 181-184. 1972.) Distribution. ThroughoutnorthernSouthAmerica, extendingto PanamaandCostaRica, anddisjunctlyin an areacomprising southernMexico, Belize and part of Guatemala,also in eastern Brazil, in Pemambuco and in Rio de Janeiroand the adjacentpartof Minas Gerais;in primaryand secondarynon-inundatedforest; to 1000m, sometimesin ratherdryor wet habitats; regionallyor locally frequent. Representative additional specimens examined. MEXICO. CHIAPAS:Mun. Ocosingo, Estaci6n Chajul, 30 Oct 1992(fl), MartinezS. et al. 25575 (MO). OAXACA: Mun. Santa Maria Chamalapa, 8-8 km N of Santa Maria, 11 Jun 1985 (fr), Hernandez G. 1217 (MO); Mun. Matias Romero, 2.3 km SSW of Aserradero La Floresta, 15 km S of Esmeralda,28 May 1981 (fl), Wendtet al. 3341 (MO). GUATEMALA. IZABAL:Cadenas (Puerto Mendez),

19 Aug 1969 (fl), Contreras8987 (MO, NY).

NICARAGUA. Rio SANJUAN:2 km NW of Rio Santa Cruz, 24 Feb 1984 (fl), Moreno 23343 (MO). COSTA RICA. ALAJUELA:5 km S of Brasilia, Rio Pizote, 30 Oct 1987 (fl-fr), Herrera 1029 (BG).CARTAGO: 24 km NE of Turrialba, 10 May 1983 (fl), Liesner et al. 15397 (BG).HEREDIA:Rio PuertoViejo, Finca La Selva, 1 Jul 1982 (fl), Hammel et al. 13071 (MICH, MO, NY). LIM6N: 29 km W of Tortuguero, 16 Aug 1979 (fl), Foothills of Davidson et al. 8378 (NY). PUNTARENAS: Cord. de Talamanca,just below Helechales, 1000-1100 (? fl), R. T Penningtonet al. 144 (BG);Prov.Manuripi, m, 30 Mar1984(fl), Davidse et al. 26332 (BG,MO, NY). PANAMA. COL6N: Santa Rita Ridge rd., between between Luz de America and Siriguero, 3 Sep 1991 (? fl-fr), Hinojosaet al. 1423(BG,LPB, MO). SANTACRUZ: Transisthmian Hwy. and Agua Clara, 11 Dec 1973 (flfr), Berg et al. 308 (BG, MO, NY). DARIEN:23 km SE Prov. Ichilo, Rio Ibabo, Reserva Forestal Chore, 5-10 of Jaqu6, 28 Apr 1980 (fl), Garwood 955 (MO). SAN Aug 1990 (st), Neill et al. 9198 (MO). BLAS: El Llano-Carti rd., Sendero Nusagandi, 16 Jun In some Ecuadoriancollections,includingPalacios 1987 (fl), McPherson 11017 (BG, MO, NY). Rio Caqueta,nr.Araracuara, COLOMBIA. AMAZONAS: et al. 3367, the peltate bractsof the pistillateinflores30 Sep 1990 (fl), Alvarez et al. 109 (NY). BOLiVAR: cence are ? impressedand surroundedby a low rim. Mun. San Juan Nepomuceno, Loma de los Colorados, 2 Sep 1986 (st), Cuadros V 3101 (BG). CAQUETA:Nr. 4. Brosimum costaricanum Liebmann(see Berg, Fl. Araracuara, 11 Aug 1986 (fl), Berg et al. 1552 (BG). CHOC6:Mun. Riosucio, Cerrosdel Cuchillo, 21 May 1988 Neotrop.Monogr.7: 180-181. 1972.) (fl), Cdrdenas 2025 (BG); Mun. Quibd6, rd. Quibd6Distribution.In CostaRicaandPanama;in low- Guayabal, 12 Sep 1976 (fl), Forero et al. 2771 (MO). NARINO:Tumaco, Curasanga, 13 Jul 1961 (fl), Acevedo land forest;rare. 12 mi SE of Barranca et al. C.100 (AAU). SANTANDER: Representative additional specimens examined. Bermeja, 1 Oct 1954 (st), Romero-Castanieda4967 (NY). COSTA RICA. PUNTARENAS: Parque Nacional Corco- VALLE:Bajo Calima, ca. 15 km N of Buenaventura, 10 vado, Ollas Trail, 3 Jan 1989 (9 fl), Kernanet al. 864 Apr 1987 (st), Gentry et al. 56751 (BG). VENEZUELA. AMAZONAS:Depto. Rio Negro, (F, MO); Rinc6n de Osa, 6-7 Feb 1974 (v), Liesner1840 middle part of Rio Baria, 21 Jul 1984(fl), Davidse 27585 (AAU, MO, NY).

236 Rio Leon, NE of Bergantin, (BG, NY). ANZOATEGUI: 10 Mar 1945 (st), Steyermark61467 (VEN). MIRANDA: Dtto. Paez, Rio Guapo,CerroRiberon,44.5 km SE of Caucagua,8-10 Jun 1977 (fl), Davidseet al. 13712 (U). SUCRE:Montainade Mochino, 20 km SE of Cumana, et al. 108426(NY).ZULIA: 16 Sep 1973(fl), Steyermark Sierrade Perija,SW of Machiques,31 Aug 1967 (st), Steyermarket al. 99942 (NY). ECUADOR.AzuAY:Nr. Molletura,29 Jan 1990 (flRio Cayapas, fr),Ortiz215 (QCA,QCNE). ESMERALDAS: ZapalloGrande,1-2 Aug 1982 (fl), Kvist et al. 40811 (AAU, NY, QCA, QCNE);W of San Mateo, Reserva Forestalde JardinBotanico, UniversidadT6cnicaLuis VargasTorres,2 Feb 1991(st), Gentryet a. 73032(MO, Pozo PetroleroGarza,ca. QCNE). MORONA-SANTIAGO: 35 km NE of Montalvo,2-12 Jul 1989 (fl-fr),Zaket al. ReservaFloristicaEl Chuncho, 4613 (BG,QCNE).NAPO: 5 km N of Coca, 10-15 Sep 1986 (fl), Neill et al. 7307 (AAU,BG, NY, QCA,QCNE);ReservaBiol6gicaJatun Sacha,8 km E of Misahualli,2 Oct 1986(fl-fr),Palacios Rd. Coca1309 (AAU, BG, NY, QCA, QCNE). PASTAZA:

Auca, km 115 S of Coca, 5 km fromRio Tigiiino, 1-6 Mar 1989 (fl), Zak 4067 (BG, NY, QCNE); Pozo

FLORANEOTROPICA BOLIVIA. BENI: Prov.VacaDiez, nr.Alto Iv6n, 8 Feb 1984(st),Boom4482 (NY);Prov.Ballivian,Estancia El Porvenir,50 km E of Rio Maniqui,18 Nov 1985 (fl), Solomon14785 (BG, NY). LA PAZ: Prov.SudYungas, nr.Sapecho,2 Mar1994 (st), Berg 1706A(BG).PANDO: Prov.Madrede Dios, rd. Sena-RioBeni, ca. km 10, Beck et al. 20426 (BG);Cobija,25 Oct 1977 (fl-fr),Meneces et al. 2078 (NY). SANTA CRUZ: Prov. Ichilo, Parque NacionalAmboro,6 Dec 1991(fr),Nee 41983 (BG,NY).

6. Brosimum gaudichaudii Trecul (see Berg, Fl. Neotrop. Monogr.7: 184-186. 1972). Distribution. Moreor less continuouslyin Central andeasternBrazilfromParato ParandandeasternParaguay and Bolivia and in isolated small areas in the AmazonBasin,inAmazonas(nearHumaita)andin Para (nearAlmeirim);in cerrado(scrub)andsavanna(woodland) and in adjacentdeforested(and depleted) areas; at low elevations;regionally frequent.

Representative additional specimens examined. PetroleroMasaramu,40 km NNE of Montalvo, 1-16 BRAZIL. AMAZONAS: Mun. Humaita,nr. BR.230, km 384 (BG,NY,QCNE).SUCUMBIOS: 2, 2 Jan 1980 (fl), Janssen 129 (NY). BAHIA: Mun. May 1990(fl), Gudinio ReservaFaunisticaCuyabeno,nr. LagunaGrande,1 Apr Barreiras, rd.to airport,11 Jun 1992 (fl), Carvalhoet al. 1988-1 Apr 1990 (st), Valencia67375 (AAU, QCA). 4000 (BG, NY). RONDONIA: 5 km fromVilhena,1 Nov Prov. Pachitea, Dtto. Honoria, 1979 (fr), Vieiraet al. 840 (MO). PERU. HUANUCO: Quebradade Shahuinto,20 Sep 1967 (fl), SchunkeV BOLIVIA.BENI: Prov.VacaDiez, 35 km E of Ribe2162 (NY). LORETO:Prov. Requena, Cafio Supai, nr. ralta, 12 Sep 1981 (fl), Solomon6234 (NY). LA PAZ: JeneroHerrera,23 Nov 1982,Encarnacion25049 (NY). Ixiamas,17 Dec 1921 (fr), Cardenas1938 (NY). SANTA MADRE DE Dios: 39 km SW of Puerto Maldonado, CRuz:Prov.Andr6sIbianez,6 km NW of Terevinto,29 Laguna,Cocacocha,17 Oct 1987 (fl), S. F Smithet al. Aug 1987 (fl), Nee et al. 35850 (MO,NY); Prov.Gua410 (NY). PASCO:Prov.Oxapampa,PuertoBermu'dez, rayos, 5 km SE of Ascencionde Guarayos,7 Sep 1990 16 Jan 1983 (st), Gentryet al. 42052 (BG). (fl), Nee 38634 (BG, NY). Rio Jurua'-mirim,17 BRAZIL. ACRE:Nr.Potangaba, PARAGUAY. Amambay: Parque Nacional Cerro Mun. Cora, 1 Nov 1983 (fl-fr), Hahn 1715 (MO, NY). May 1971 (fl), Maas et al. R13082 (NY). BAHIA: Una,Maruim,30 Apr 1981 (st), Moriet al. 13866 (NY); Mun. Uru9uca,7.3 km N of Serra Grande,on rd. to Itacare,1-12 Jul 1991 (st), Thomaset al. 8269 (NY). 7. Brosimum glaziovii Taubert(see Berg, Fl. Neotrop. ESPiRITOSANTO:Mun. Linhares,ReservaFlorestalda Monogr.7:186-188. 1972). CRVD, 6 Nov 1979 (fl), Folli 159 (BG). MARANHAO: Rio TuriaQu, Ka'aporIndianReserve,31 Jan 1985 (st), No changes to be reported. Balee et al. 135 (NY);Alzilandia,Rio Pindare,12 Dec 1978(fl), Jangouxet al. 443 (NY). MATOGROSSO:Mun. Sao Felix do Araguia,FazendaSanta Cecilia, 17 Oct 1985(fl-fr),Ferreiraet al. 6506 (BG,NY);RioAripuana, 8. Brosimum glaucum Taubert(see Berg, Fl. Neotrop. Monogr. 7: 188. 1972). nr. HumboldtCenter, 10 Oct 1973 (fl), Prance et al. 18375 (NY). PERNAMBUCO:Nr. border of Engo. Distribution. In eastem Brazil (southern Minas SambaquimandJundia,12 Feb 1965 (fl), Teixeira2594 Gerais and Bahia); in forest; apparentlyvery rare. (US). RORAIMA:Ilha de Maraca,Mun. Alto Alegre, SEMAStation,8 Jun 1986 (fl), Hopkinset al. 616 (BG, Additionalspecimen examined. BRAZIL. BAHIA: NY). RONDONIA:Mun.Boa Vista,Rio Uraricoeira,Ilha Mun. Una, rd. Oliven9a-Buerarema, 33 km SW of de Maraca,10 May 1987 (fl), Limaet al. 753 (BG,NY); Olivenqa, 1 May 1981 (st), Mori et al. 13882 (NY). Madeira-Mamorerailway, 13 Sep 1963 (fl), Maguire MINAs GERAMS: CarlosChagas,UHE Mucuri,Feb 1999 rd. BR.101, (fl), L. V Costa BHCB 45813 (BG). et al. 56628 (NY). SERGIPE:Mun.EstAncia, ca, km 19,4, 28 Nov 1993(fl), Amorimet al. 1519 (BG). The second (or possibly third) collection of this SAOPAULO:Portal de Igarata,5 Jan 1985(fl), Gentryet al. 49194 (BG). species differs fromthe materialon which the descrip-

TAXONOMIC TREATMENT

tion in the monograph was based in having a dense indunentumon thelaminabeneath(being,therefore,not scabridulous)anda denserindumentumof the stipules. The materialwas collected from a tree 25 m tall.

9. Brosimum rubescens Taubert (see Berg, Fl. Neotrop. Monogr.7: 189-193. 1972).

237 BRAZIL. ACRE: BR.364, km 40, ReservaINCRA SantaLuzia, Campbellet al. 7097 (BG). BAHIA: Una, FazendaSao Rafael, 5 Nov 1969 (fl), Santos 467 (U). GoiAs: Ilha do Bananal,ParqueNacionaldo Araguaia, 2 km fromMacauiba,23 Sep 1980 (fl-fr), Ratteret al. 4501 (K). ROND6NIA: Mun. P6rtoVelho,Usina Hidroelectrica Samuel, 12-24 Sep 1988 (st), Thomaset al. 6355 (BG). RORAIMA: Ilhade Maraca,SEMAEcological Reserve,27 Sep 1987 (st), Millikenet al. 637 (BG). BOLIVIA. PANDO: Prov.Abuna,Rio Negro,nr.jct. withRioAbunk,7 Jul 1992(st),Gentryet al. 77938(MO).

Distribution. In (largely the northernpartof) the Amazon Basin and the Guianaregion, and disjunctly in the Pacific Coastalregion of ColombiaandPanama, and in eastem Brazil (Bahia to Rio de Janeiro and 10. Brosimum melanopotamicum C. C. Berg (see Goias);in non-inundatedforest;to 1200 m; regionally Berg, Fl. Neotrop. Monogr. 7: 193. 1972). frequent. Representative additional specimens examined. PANAMA.BOCASDELToRo: Nr. Rio, San Pedro, 18 Jan 1975 (st), Gordon 17 (MO). COLON: Santa Rita lumber rd., E of Colon, 22 Dec 1978 (st), Berg 333 (MO). PANAMA:El Llano-Cartird., km 8-12, 12 Dec 1973(st), Nee et al. 8792 (U, NY). SAN BLAS: Cangandi, 27 Mar 1986 (fl), Nevers et al. 7533 (BG, MO). COLOMBIA. AMAZONAS: Rio Caqueta, nr. Araracuara,10-22 Nov 1982 (st), Idroboet al. 11226 (NY). C6RDOBA:Jct. of Rio Tigre and Rio Manso,25 Jul 1988 (st), Gentryet al. 63750 (MO). VALLE:Rio Naya, PuertoMarizalde,20-23 Feb 1943 (st), Cuatrecasas 14030 (F); ca. 20 km NW of Buenaventura,18 Aug 1988 (st), Faber-Langendoenet al. 2054 (BG, MO). VAUPtS:Estaci6nBiol6gica Caparu,3 km N of Lago Taraira,1986-1990 (fl), Defler 324 (MO). VENEZUELA. AMAZONAS:Santa Barbara del Orinoco, 1-2 km S of Trapichote,26 Feb-2 Mar 1976 (st), Berryet al. 2078 (U, VEN);Rio Manapiare,4 km W of Cailo Seje, 18-19 Jun 1876 (st), Berry2219 (U, VEN).BOLiVAR:Mun.Cedeno,55 km S of Jabillal,Apr 1989 (fl), Fernandez5258 (BG, NY); Rio Karuai,be1200 m, tween base of Ptari-tepuiand Sororopan-tepui, 28 Nov 1944 (st), Steyermark60721 (F). ECUADOR. MORONA-SANTIAGO: 20 km N of Taisha,20-23 Aug 1976 (st), OrtegaU. 94 (US). NAPO: ReservaFloristicaEl Chunco,5 km N of Coca, 10-11 Sep 1986 (fl), Palacios et al. 1244 (BG, MO, QCA, QCNE,US); ReservaBiol6gicaJatunSacha,8 km E of Misahualli,6-14 Oct 1988 (fl-fr),Palacios 3065 (AAU, BG, QCNE).PASTAZA:Pozo PetroleroNamoyacu,30 km S of Curaray,13-30 Nov 1990 (fl), Espinozaet al. San 571 (BG, QCNE). SUCUMBiOS: Cant6nShushufindi, Roque, 14 Dec 1997 (st), E. Freireet al. 2849 (QCNE). PERU. AMAZONAS:Prov. Bagua, Dtto. Imaza, Feb 1995 (fl), R. VYasquez et al. 19878 (BG). Yamayakat, andRioLaTorre, MADREDEDIos: Jct.of RioTambopata 23 Jul 1985(st), Gentryet al. 51190 (BG).PASCO:Prov. Oxapampa,Rio Iscozacin, Cabeza de Mono, 10 Jun 1983 (st), Gentryet al. 41789 (BG, MO).SAN MARTIN: Prov. Marsical Careres, Dtto. Tocache Nuevo, Rio Tocache,FundoMelidia, 16 Jul 1974 (fr), SchunkeV 7454 (MO, U).

Distribution. In the Amazon Basin, along the Rio Negro, and in Guyana; in non-inundated forest; uncommon.

Representative additional specimens examined. nr.Cerro VENEZUELA.AMAZONAS: Rio Mawarinuma, de La Neblina Base Camp, 27 Nov 1984 (st), Croat 59369 (MO).

GUYANA.UpperTakutu-UpperEssequiboregion, KamoaMts. 2 km S of KamoaR., 12 Nov 1996 (d), Clarke3125 (BG, US). BRAZIL. AMAZONAS: Nr. Manaus, Estrada do Aleixo, 30 Aug 1973 (st), Berg 247 (AAU);Rio Negro, Jupati,17 Apr 1947 (o), Pires 353 (NY).

11. Brosimum utile (Kunth) Pittier (see Berg, Fl. Neotrop. Monogr. 7: 194-204. 1972). An increase in theAmazonian collections made it clear that Brosimum longifolium has to be reinstated as a species. Several characters, such as the repand margin of the lamina, the persistent epidermis on the petiole and the initially recurved inflorescences, usually occurring in pairs, justify the reinstatement. The other subspecies which have been recognized can be maintained. A number of sterile collections from the upper Amazon Basin (in particular from Peru) could not be included in one of the two Amazonian subspecies, and in some cases it is even uncertain whether they belong to the species.

Ila. Brosimum utile (Kunth) Pittier subsp. utile. Distribution. Northem Venezuela to northem Colombia, extending into Panama (San Blas, Panaml, Chiriqui, Cocle, and Colon); in forest; to ca. 1500 m.

Representative additional specimens examined. PANAMA.CHIRIQUi: E of AudubonCabin,S of Cerro Punta, 12 Jul 1983, 1400-1800 m, (fl-fr), Hamiltonet 7 Feb al. 3832 (BG).COCLP: Rd. La Pintada-Coclesito, 1983 (fl-fr), Hamiltonet al. 2832 (BG, MO). COLON:

2 -38-

TresBrazosSawmill,betweenSaludandRio Indio,15 Jul 1969 (st), Howell 100 (MO). PANAMA:Rd. El LlanoCarti,km 16, 23 Mar 1973 (fl-fr),Kennedyet al. 2922 (MO). SAN BLAS: Nr. El Llano-Carti rd., NW of Nusagandi,18 Jun 1987 (fl), McPherson11060 (MO).

FLORA NEOTROPICA

Distribution. In Panama (Darien, San Blas, and Panama);in forest; at low elevations.

Representative additional specimens examined. PANAMA.PANAMA: Rd. El Llano-Carti,12.7 km, 15 Feb 1975(fl-fr),Moriet al. 4702 (BG, MO).SANBLAS: COLOMBIA. CHOC6: Mun. Turbo, rd. Tapon del Darien,betweenRio LeonandLomasAisladas,kan37, 27 Rd. El Llano-Carti,lkm19.1, 28 Sep 1984 (fl-fr),Nevers Dec 1983(fl-fr),Brandet al. 772 (MO);Mun.Riosucio, et al. 3948 (BG, MO). Urabaregion, Cerrodel Cuchillo, 9 Nov 1987 (fl-fr), The leaf base may varyto subcordate.This subspe30 km E of Carare,29 Cdrdenas691 (MO). SANTANDER: cies co-occurswiththeapparentlymorecommonsubsp. Sep 1977 (st), Gentryet al. 20008 (BG, MO); Mun. PuertoWilches,betweenLa Gomezandkm 80 of Ferro- utile, fromwhich it can be readilydistinguishedby the carrildel Atlantico,22 Apr1960(fl-fr),Romero-Castaniedarounded to subcordate leaf base and the prominent 8395 (NY). VALLE:Bajo Calima,ConcesionPulpapel/ smallerveins on the lower surfaceof the lamina. The 19 Feb 1985(fl-fr),MonsalveB. 668 (MO). latter feature can also be used to distinguish subsp. Buenaventura, darienense from subsp. magdalenense. llb. Brosimum utile (Kunth) Pittier subsp. occidentale C. C. Berg.

Distribution. In the CoastalPacificregionof Ecuador and Colombia,northwardsto Antioquia;in forest; at low elevations.

1 f. Brosimum utile (Kunth) Pittier subsp. ovatifolium (Ducke) C. C. Berg.

Distribution. In the Amazon Basin, largely in its northempart,extendingto FrenchGuiana,Guyana,and Representative additional specimens examined. Venezuela(Bolivar), and with some recordsfrom the Mun.Taraza,Cgto. El 12, 9 COLOMBIA.ANTIOQUIA: southem partof the Amazon Basin; in non-inundated Nov 1987 (fl), Callejaset al. 5491 (BG, NY). CHOCO: to ca. 1100 m. forest; Rio Baudo,QuebradaPaulita,11 Feb-29 Mar 1967 (st), Fuchs et al. 22038 (NY); behindBahia Solano, Puerto Representative additional specimens examined. Mutis, 5 Jan 1973 (fl-fr), (entry et al. 7237 (U). COLOMBIA. CAQUETA:Nr. Araracuara, 23 Jan 1989 NARIF10:Mun. Iscuande,Rio, Sequi6n, 23 Nov 1955 (st), Gentryet al. 64967 (MO). VAuPts:Rio Apaporis, (st), Romero-Castafneda 5490 (AAU, NY). Jinogoje,nr.mouthof Rio Piraparana,15 Jun 1952 (fl), Schulteset al. 16735 (MO) and 16825 (GH). The relativelylong stipules,morenumerouslateral VENEZUELA. AMAZONAS: Rio Yutua,Caiio Grande, veins, and the tendency to have a roundedbase of the SSE of CumbreCamp,1100-1150 m, 26 Dec 1957 (fl), laminaratherthanan acuteto obtuse one, appearto be Maguireet al. 42531 (MO,NY). BOLivAR:Mun. Raul consistent enough to distinguishthis subspecies from Leoni, Rio Kurutu,ca. 11 km above mouth,Apr 1988 (fl), Fernandez4530 (BG, MO, NY); Rio Baria (Rio subsp. utile. SW side of Cerrode La Neblina,16 Feb Mawarinuma), 1985 (fl), Nee 30878 (BG, MO, NY); Rio Casiquiare, llc. Brosimum utile (Kunth) Pittier subsp. allenii betweenmouthof Rio Siapaand mouthof Rio Paciba, 26 Jul 1959 (fl), Wurdacket al. 43633 (NY). (Woodson) C. C. Berg. GUYANA.Potaro-Siparuniregion, IwokramaRain No changes to be reported. ForestReserve,Karupukari, 26 May 1997 (fr), Moriet al. 24620 (BG, US). lld. Brosimum utile (Kunth) Pittier subsp. magdalenense C. C. Berg.

FRENCH GUIANA. SinnamaryR., above Petit Saut,

betweenCr.PlombandCr.Tigre,31 Aug 1993(st),Mori et al. 23490 (BG,NY); Sauilregion,LayonRochaBateau, Distribution. In Colombia, in the Rio Magdalena 15 May 1986 (fl), Sabatier1245 (BG, NY); GrandInini valley, and probably also in Choco; in forest; at low R., 16 Jul 1990 (st), Sabatieret al. 3315 (BG, MO). ECUADOR. MORONA-SANTIAGO: 11.7 km SW of elevations. Taisha,11-15 Sep 1976(fl), OrtegaU 131 (US). NAPO: Additional specimen examined. COLOMBIA. ReservaBiol6gicaJatunSacha,8 km E of Misahualli,8 CHOC6:Rd. Quibd-Tutunendo,14 km NE of Quibdo, Aug 1987 (st), Palacios 1746 (BG, QAME);Afnangu, ParqueNacionalYasuni,2 May 1983 (fl), Lawessonet 17 Jan 1979 (fl), Gentryet al. 24314 (MO). al. 39562 (BG,MO,NY, QCA,QCNE),Jan 1986 (fl-fr), The identityof the additionalcollection consisting SEF 9295 (AAU, NY, QCA, S). PASTAZA: Rd. Cocaof fallen leaves andinflorescencesis not quitecertain. Auca, 110km S of Coca, 10 km fromRio Tigiiino,7 Jan 1989 (st), Palacios et al. 3398 (BG, MO, QAME). SUCUMBiOS: Dureno,28 Dec 1988 (st), Gentryet al. Ile. Brosimum utile (Kunth) Pittier subsp. darien- 64324 (BG); ReservaFaunisticaCuyabeno,Zancudo, ense C. C. Berg. 27 Sep 1991 (st), Palacios et al. 7920 (QCNE).

239

TAXONOMICTREATMENT PERU. AMAZONAS:Rio Santiago,ca. 65 km N of Pinglo, Quebrada Caterpiza, 19 Sep 1979 (fl-fr), Huashikat696 (MO).HUANUCO: Prov.PuertoInca,Dtto. Yuyapichis,ReservaForestalDANTAS,14 Oct 1989 (flfr), KroellS. 619 (BG).JUNIN:Prov.Satipo,SanMartin de Pangoa,Mazoranquiri, 2 Oct 1993 (fl), Rios T et al. 1910 (QCNE). LORETO: Prov. Maynas, Rio Nanay, PuertoAlmendrez,22 Jul 1984 (fl-fr),R. Vdsquezet al. 5321 (NY); Prov.Requena,JeneroHerrera,18 Jul 1986 (st), Spichigeret al. 3064 (NY). MADRE DE DIos: Prov. Manu, Cerro de Pantiacolla, 10-15 km NNW of Shintuya,13 Dec 1985 (st), Fosteret al. 10798 (BG). UCAYALI:Prov. Coronel Portillo, Dtto. Calleria, San Alejandro,6 Sep 1973 (st), CeijasB. s.n. (NY, US). BRAZIL. MATOGRosso: RioAripuanA, nr.Humboldt Center,11 Oct 1973 (fl), Berg et al. P 18429 (NY); Rio Juruena,Cachoeirade Sao Joao da Barra,4 Apr 1977 (fl), Rosa et al. 2055 (NY). BOLIVIA. BENI:Prov. Vaca Diez, nr. Alto Iv6n, 30 Jan 1984 (st), Boom4334 (NY). PANDO:Rio Negro, nr.jct. with Rio Abuna,7 Jul 1992 (st), Gentryet al. 77947 (MO).

BRAZIL. AMAZONAS: Manaus, Estrada do Aleixo, Compania de Plantag6es, 3 Sep 1973 (fl), Berg et al. P 18804 (NY); Mun. Santa Isabel do Rio Negro, nr. Santa Isabel, 10 Aug 1987 (fl), Ferreira 9337 (BG, NY).

The collection Nee 30878 (from Bolivar, Venezuela) has turbinateinflorescences with the staminate flowers confined to its upper part. In young fruiting state the upper part is separated from the lower fruit-containing part by a constriction, due to which the (developing) infructescence resembles that of Trymatococcus

13. BrosimummultinerviumC. C. Berg,Novon6: 230,t. 1. 1996.Type.Ecuador. Pastaza:Rd.CocaAuca,ca. 115 km S of Coca, 4-6 km S of Rio Tigiiino,22-28 Feb1989(d), Zak3955(holotype, QCNE;isotypes,BG, MO). Fig. 67

glabrousorsparselypuberulous; receptaclegloboseto turbinate orto hemispherical. 0.5-0.8 cmdiam.;perianthlacking;stamen1;filamnent ca. 0.5-0.8 mmlong; anther ca.0.2mmbroad;stigmas0.2-2 mmlong;bracts ca.0.5-2mmdiam.,minutely pubermlous. Infiuctescences globose,ca. 3 cmdiam.,brownishatmaturity. Distribution.Inthenorthwestem partof theAmazon Basin, from Colombiato the surroundingsof Manaus;in non-inundated forest;apparently rareand distributed. discontinuously Representative additional specimens examined. COLOMBIA. AMAZONAS:RioCaquetA,Araracuara,Villa Azul, 11 Mar 1989 (st), Londonio et al. 1088 (NY). AMAZONAS/VAUPES:

Rio Apaporis, Jinogoje, 13 Jun

1952 (st), Schultes et al. 16481 (MO, NY), 20 Jun 1952

(fl), Schulteset al. 16825 (GH, MO, NY).

amazonicus.

Tree,to40 mtall.Leafjy twigs3-5 mmthick,sparsely torather denselybrown-puberulous towhitish-pubescent. Laminaellipticto oblongto (sub)ovate,12-30 x 6-19 cm,coriaceous; apexshortly andabruptly acuminate; base cordateto truncate(toobtuse);marginentire,often? 12. Brosimum longifolium Ducke,Arq. Serv. Florest. lower revolute; uppersurfacepuberulous onthemidrib; 1: 3. 1939. Brosimum utile (Kunth) Pittier subsp. to subvelutinous on surfacedenselybrown-puberulous longifolium C. C. Berg, Acta Bot. Neerl. 19: 328. theveins;lateralveins(30-)4-50 pairs,oftensomeof 1970; Berg, Fl. Neotrop. Monogr. 7: 201. 1972. themfurcate; venation tertiary forthegreater partscalariType. Brazil. Amazonas: nr. Sao Paulo de Olivenca, form;petiole0.5-1.5 cm long,denselybrown-puberu17 Feb 1932, Ducke RB 23621 (holotype, RB; lous,theepidermis flakingoff;stipules4.5-9 cmlong, isotypes, B, G, K, P, U, US). to brownto whitish-pubesdenselybrown-puberulous Brosimumfoetidum Ducke, Bol. Tecn. Inst. Agron. cent.Inflorescencessolitaryin the leaf axils,initially 2-6.5cmlong,thelowerpart N. 4: 1. 1945. Type.Brazil.Amazonas:Manaus, deflexed; peduncle sparsely nr. Estradado Aleixo, Ducke 1457 (holotype, puberulous, theupperpartdenselyandminutelypuberRB; isotypes, F, GH, K, NY, US). ulousandwitha fewbracts,theuppermost partbroadof staminate inflorescence discoidto ened; receptacle Tree,to 30 m tall. Leafy twigs 3-5 mm thick, glabrous 1cmdiam.;perianth ca. stamen subturbinate, lacking (?); or with sparse hairs mainly near the scars of the stipules. ca. 0.5 mm ca. mm filament anther 0.2 1; long; long; or elliptic Lamina ovate to subovate to oblong, 8-25 of receptacle pistillate inflorescence subglobose to base (-33) x 3-10(-15) cm, coriaceous; apex acuminate; witha singlepistillate 0.8-1.2 cm subturbinate, diam., to truncate; margin repand; both surfaces glasubcordate flowerin thecenter;stigmas1-2 mmlong;bractsca. 1 brous; lateral veins 8-14(-18)pairs; tertiaryvenationpartly mmdiam.,minutely puberulous. the Use. The fibers of inner bark are used to make bark cloth (Bolivia. Beni; Colombia. Vaupes).

scalariform;

epidermis with

sparse

petiole

long

cm long,

0.8-2.5(-3.5)

persistent; hairs.

stipules

1-4

Inflorescences

the leaf axils, initiallyrecurved;

glabrous,

cm long, usually

peduncle2.5-3.5

glabrous

or

in pairs in cm long,

Distribution(seeFig.82.4).IntheupperAmazon Basin,fromColombia(Vaupes)to Peru(Cuzco);in non-inundated forest.

FLORA NEOTROPICA

240

F ig67Brsmmmlievu

efytimahifoeec

H

oeal130.BHedikBegBegn

Flg. 67. Brosimum multinervium.Leafytwig with inflorescence(Hurtado et aL 1300). [By HendriekeBerg,Bergen.]

Specimens examined. COLOMBIA. AMAZONAS/ QCNE) and 4349 (BG, MO, NY, QCNE). SUCUMBIOS: VAUPtS: Rio Apaporis, Soratama, nr. mouth of Rio Dureno, 30 Apr 1986 (st), Ceron 286 (BG, QAME). PERU. AMAZONAS: Prov. Bagua, Yamayakat, 21 Feb Cananari,14 Dec 1952 (fl), Schulteset al. 14908 (US). ECUADOR. PASTAZA: Rd. Coca-Auca, ca. 120 km 1996 (fl-fr), N. Jaramillo et al. 1223 (MO); Quebrada S of Coca, nr. Rio Tigiuino, 17 Jul 1989 (fl), Gudifio 62 Sasa, 3 Jul 1979 (juv), Kayap 996 (AAU, MO, NY). (AAU, BG, MO, QCNE); rd. Coca-Auca, ca. 115 km S Cuzco: Prov. Quispicanchis, rd. Cuzco-Maldonado, nr. of Coca, 4-6 km S of Rio Tiguiino, 7-9 Jan 1989 (i), Quince Mil, 18 Feb 1991 (st), Nziriez13048 (MO); Prov. Hurtado 1300 (BG, MO, QCNE), 26-31 Jan 1989 (st), Quispicanchis, between Pande Azucar and Quince Mil Neill et al. 8789 (BG, QCNE), 24 May 1989 (v), Rubio Airport, 10 Aug 1991 (st), Ngifez et aL.13941 (BG); Prov. 131 (AAU, BG, MO, QCNE), Zak 3976 (QCNE) and 4005 Paucartambo,Cosniipata,May 1914 (st), Weberbauers.n. (QCNE), 1-6 Mar 1989 (st), Zak 4116 (MO) and 4178 (MOL). HukNuco: Prov. Puerto Inca, Pachitea, 15 Sep (MO), 1-9 Apr 1989 (v), Zak et al. 4244 (BG, MO, NY, 1982 (st), Foster 8851 (MO); Prov. Puerto Inca, Dtto.

241

TAXONOMIC TREATMENT Yuyapichis, Reserva ForestalDANTAS, 1-15 Oct 1990 (flfr),Tello267 (MO).UCAYALI: Prov.CoronelPortillo,Dtto. Calleria, San Alejandro, 2 Sep 1993 (st), Ceijas B. s.n. (MOL), 11 Sep 1973 (st), Ceijas B. s.n. (MOL), 6 Oct 1973 (st), Ceijas B. la (BG); Prov. Coronel Portillo, Carretera Marginal, 22 km S of km 86, 11 Feb 1981 (?), Gentry et al. 31198 (MO, U); Est. Exp. Alexander von Humboldt,23 Feb 1981 (st), Haccha 11 (INPA).

This species is relatedto Brosimumparinarioides and B. utile. It can be easily recognized by the great numberof lateralveins. In the two relatedspecies the numberof lateralveins varies between 12 and 32. Juvenile material(Ceijas s.n. andKayap996) has leaves to 50 x 16 cm, the indumentumon the venation of the lamina beneath is rathersparse, and the number of lateralveins tends to be ca. 30.

14. Brosimum potabile Ducke (see Berg,Fl. Neotrop. Monogr. 7: 204. 1972). Distribution. In the northem part of the upper Amazon Basin; in non-inundatedforest;at low elevations; infrequent. Representative additional specimens examined. COLOMBIA. META: Parque Nacional Natural Tinigua, Serrania Chamusa, Centro de Investigaciones Primatologicas La Macarena,Dec 1990 (fl-fr), P. Stevenson 216 (COL). VAuP*s: Estacion Biologica Caparui,3 km N of

LagoTaraira,1986-1990(fl), Defler316 (MO). VENEZUELA. AMAZONAS:Depto. Atabapo, "pica al norte del campamento,"04?00'N, 67?45'W, Oct 1989 (fl),

Velazco741 (BG). ECUADOR.

NAPO:

Parque Nacional Yasuni, Pozo

PetroleroAmo2, 9-19 Jan1988(st),Neillet al. 8359 (BG,

15b. Brosimum parinarioides Ducke subsp. amplicoma (Ducke) C. C. Berg. Distribution. In the upper Amazon Basin, from Colombiato Bolivia; in non-inundatedforest; apparently ratherrare,but regionally or locally common. Representative additional specimens examined. COLOMBIA.AMAZONAS: Rio CaquetA, nr.Isla Cristina Maria,8 Jun1988(st),Sainchez S. et al. 665 (BG). PERU. LORETO:Jenero Herrera,3 Apr 1986 (st), Spichigeret al. 3066 (NY); Prov. Maynas,Rio Napo, Llachapa,21 Jan 1983(ol), R. Vdsquez et al. 3817 (AAU, BG,NY). BOLIVIA.PANDO:Prov.Abuna,Rio Negro,nr.jct. withRioAbuna,7 Jul 1992(st),Gentryet al. 77962(MO); Prov.FedericoRoman,15 kmSWof mouthof RioNegro, 16 Jul 1992(st),L. Vargaset al. 1018 (BG,LPB). The identity of the sterile andjuvenile specimens from Bolivia is not quite certain.

UnnamedCollections Some sterilecollections could not be placed in any of the recognized species. They were from trees to 25 m tall, with lanceolate to oblong laminas, 8-18 x 1.5-5 cm, with subacute base, the lower surface sparselypuberulouson the veins, with 15-20 pairsof lateralveins and the midrib slightly impressed, with the epidermis of the petiole flaking off, and the stipules ca. 1 cm long. ECUADOR.NAPo:ParqueNacionalYasuni, Maxusrd., km 40, 10 Sep 1994 (st), Aulestia2736 (QCNE);Canton Tena,EstacionBiol6gicaJatunSacha,8 kmE of Mishualli, 21 Oct 1989(st),Palacioset al. 4691 (QCNE).

MO, QAME); Aniangu,ParqueNacional Yasuni,Apr 1985 Prov. Pachitea, (st), SEF8704 (AAU).PERU. HUANUCO: Trymatococcus ca. 20-24 km SE of Puerto Inca, 1 Oct 1988 (fl-fr), Wallnoeffer18-11088 (BG). LORETO:Prov. Requena, 11. Trymatococcus Poeppig & Endlicher (see Berg, Jenaro Herrera, 17 Aug 1982 (fl), Spichiger et al. 3011 Fl. Neotrop. Monogr. 7: 208-216. 1972). (MO);Prov.Maynas,Allpahuayo,Nov 1990 (st), R. Vdsquez By placing Trymatococcus paraensis under T et al. 14741 (BG, MO). BRAZIL. ACRE:ReservaINCRA SantaLuzia, BR.364, amazonicus, the number of species of this genus is

km 40, 5-19 Oct 1984 (st), Campbellet al. 6899 (BG).

reduced to two.

ROND6NIA:P6rtoVelho, Usina HidroelectricaSamuel, 12-

24 Sep 1988(st), Thomaset al. 6502 (BG).

15. Brosimum parinarioides Ducke (see Berg, Fl. Neotrop. Monogr. 7: 205-208. 1972.)

15a. Brosimum parinarioides Ducke subsp. parinarioides. No changes to be reported.

1. Trymatococcus amazonicus Poeppig & Endlicher (see Berg, Fl. Neotrop.Monogr.7: 212-213. 1972, incl. T paraensis). Additional synonym: Trymatococcus paraensisDucke,Arch.Jard.Bot. Rio de Janeiro3: 22. 1922. Revised description: Tree,to 15 m tall, or shrub. Leafy twigs 1-5 mm thick, yellowish to darkbrown,appressed-puberulous

242

FLORANEOTROPICA

tohirtellous to(sub)velutinous, usuallysomeof thehairs SUCUMBIOS: ReservaFaunisticaCuyabeno,nr. Laguna uncinate.Lamina (ellipticto) oblongto subobovate, Grande,11 Mar 1990 (fr), Balslev et al. 97327 (AAU, (2-)7-22(-36) x (1.5-)3-8(-1 1) cm, not or slightly QCA). PERU.LORETO:Prov.Requena,nr.JenaroHerrera, tocoriaceous; inequilateral, chartaceous apex(acuteto) 15 Feb 1988 (fr),Daly et al. 5653 (BG, MO,NY); Prov. acuminateto caudate;baseacuteto rounded(to subAlto Amazonas,Dtto. Lagunas,between Lagunasand cordate);marginentire,planeto revolute;uppersur- Argentina,6 Jul 1972 (fl), McDanielet al. 16447 (NY). ? bullate; SAN MARTIN: Prov.Lamas,rd.Tarapoto-Yurimaguas, faceglabrous sometimes exceptforthemidrib, nr. lowersurfacesparselytodenselyappressed-puberulous Yumbatos,8 Oct 1984(fl), Maaset al. 5999 (AAU,BG). to hirtellousontheveins;lateralveins5-13 pairs;terBRAZIL. ACRE:Mun. Cruzeirodo Sul, Rio Jurua, tiaryvenation partlyscalariform, thesmallerveinsplane IgarapeViseu,5 Nov 1991 (fl-fr),Ferreiraet al. 10595 to prominentbeneath;petiole(0.3-)0.6-2(-2.5) cm (BG). AMAPA: Camaipi, 8 Sep 1983 (st), Mori et al. long,usuallywithuncinatehairs;stipules0.3-0.8 cm 15956 (BG, MO). AMAZONAS: Mun. Itapiranga,Rio long,appressed-pubescent, caducous.Inflorescences: Uatuma,nr. mouth of Rio Pitinga, 27 Aug 1979 (fl), to Ferreiraet al. 868 (NY); Rio Cauaburi,13 Nov 1965 peduncle0.3-0.7(-1.5)cmlong,brown-puberulous oftenwithuncinatehairs;receptacle (fl-fr),Maguireet al. 60178 (MO,NY); Rio Purius,Lago (sub)velutinous, Preto,2 km N of Labrea,26 Jun 1971 (fl), Pranceet al. turbinateto almostfungiform,and0.4-1 cm diam., 13725 (NY); rd. Manaus-PortoVelho, Igap6 A9u, 15 withorwithoutuncinate brownshort-velutinous, hairs; Mar 1974 (fl-fr), Prance et al. 20585 (MO, NY). staminate flowersseveralto many;perianth1-1.5mm ROND6NIA: Mun. P6rto Velho, Usina Hidroel6ctrica long,3-lobedto -parted,puberulous, sometimeswith Samuel,12-24 Sep 1988 (fl), Thomaset al. 6251 (BG). uncinatehairs;stamens3(-2); filaments0.5-1.5 mm RORAIMA: Rio Uraricoera,CachoeiraTocuxema,9 Mar long;anthers 0.2-0.6x 0.2-0.8mm;style3-5 mmlong; 1979 (fl), Pires et al. 16896 (NY); Serrados Surucucus, stigmas3-7 mmlong;bractsovateto suborbiculate, nr.Posto SurucucuMission, 19 Feb 1969 (fl-fr),Prance attached orsubpeltate, basally puberulous. Infiuctescences et al. 10085 (NY). PARA: Rio Cachorro, Serra do 1.5-3.5cmdiam.,crowned withthe(rem- Cachorro,20 Nov 1985 (fl), L. Coelhoet al. 219 (NY); (sub)globose, of staminate nants the) flowersandusuallycovered Rio Jarn,Monte Dourado, 13 Jun 1968 (fr), Oliveira withsparseor shorthairsandsparseto denselonger 4506 (IAN,NY); Rio Jan, rd. to Munguba,18 Apr 1969 (fr), N. T Silva 1871 (BG, NY). uncinatehairs,greenishorbrownish. Trymatococcus

amazonicusis a very variablespeDistribution(seeFig.82.3).IntheAmazonBasin, cies with regardto the dimensions, shape, textureand to Ecuador, mainlyitsnorthern extendpart,westwards venation of the lamina,in the denseness and the color ing to the Guayanapartof Venezuela(Bolivar)and of the indumentum,varying from pale yellow to dark northern innon-inundated through Parato theGuianas; rusty-brown,andin the shapeof the inflorescencesand inundated orinperiodically (varzea,tahuampa) forest; in the size of the fruitingreceptacle.Most of the mateatlow elevations.

rialwest of theRio Negro (Brazil-Venezuela)has lamiRepresentative additional specimens examined. nas with pale yellow, rathersparse and ? appressed COLOMBIA. AMAZONAS: Rio Caqueta,Isla Mariiiame, hairs and plane smallerveins beneath.East of the Rio 23 Sep 1986 (st), Brand et al. 1489 (BG). CAQUETA: Negro material usually has laminas with prominent Nr.Araracuara, 13 Jul 1986 )(fr),Berg et al. 1557 (BG). smallerveins anddenserand patenthairson the lamina VENEZUELA. AMAZONAS:SanCarlosde RioNegro, beneath.The indumentumis usuallydarker,varyingto 4.3 km NNE of Solanord., 10 Jul 1980 (fr),H. L. Clark et al. 7640 (NY);Depto.Atabapo, Rio Puruname, 29 May- dark rusty-brown, and the lamina is usually thicker 5 Jun 1982 (fr), Huber et al. 6391 (NY); SierraParima, (to coriaceous and then often more or less bullate nr. Simarawochi,Rio Matacuni,18 Apr-23 May 1973 above). In this eastern part of the range of distribu(fl-fr), Steyermark 107061 (NY). BOLIVAR: Mun. Sucre, tion-Venezuela (Amazonas and Bolivar) and Brazil Wanaecudunia,5 km W of Santa Maria de Erebato, Feb (northwestemAmazonasandRoraima)-the leavesare 1989 (fr), Fernandez et al. 4932 (BG, MO, NY); Rio relatively large (longer than 10 cm) and the fruiting Canaracuni, 13-26 Apr 1988 (fl), Stergios 11828 (NY). receptaclesrelatively large (to 3.5 cm diam.). More to GUYANA. Upper Demerara-Berbice region, the east, in the Guianas, Brazil (Amapa and northern Berbice R., N of Kwakwani, 17 Apr 1993 (st), Hoffman Para),the leaves tendto be smaller(to 10 cm long) and et al. 3969 (BG). the fruitingreceptacleto 1.5 cm diam.;the indumentum FRENCH GUIANA. Saul, La Fumee Ouest, 10 Jun is mostly i darkbrown and the laminais mostly thick 1988 (fr), Mori et al. 18850 (BG, NY). ECUADOR. NAPO:Cant6nAguarico, Reserva Etnica and ? bullateabove andwith a ? revolute margin.The fact that in the monographjuvenile specimens of the Huaorani, Maxus rd., km 61, 26-30 Oct 1993 (fl), entity recognized as T paraensis (with largerleaves Aulestia et al. 1173 (BG, QCNE); Parque Nacional Yasuni,Tiputini, 31 May 1997 (st), Macia et al. 668 (BG). than adult ones) have been referredto T amazonicus

TAXONOMICTREATMENT

243

is anindication of theweaknessof differentiating characters.Thethreemorphological entitiescircumscribedthe"typical" westernform,thefonnof thecentralpart of therange,includingthetypeof T turbinatus,and theeasternform,includingthetypeof T paraensisareconnectedwithintermediates; it seemsunjustified to recognizethemformally,notevenatthesubspecies level.Theinfructescences, whichoftenremaingreenish,canadhereto mostsurfacesby theuncinatehairs (Dalyet al. 5653),andmaycontribute to exozoochorousdispersal.

(NY); RupununiDistrict, between KuyuwiniLanding and KassikaityuR., 22 Oct 1992 (fl), Jansen-Jacobset al. 3055 (MO, NY). BRAZIL. AMAPA: Munguba,PerimetralNorte, 28 km E of Agua Fria,20 km W of CampoVerde,24 Oct 1979 (fr),Austinet al. 7205 (NY,U); Rio Oiapoque,nr. CachoeiraTresSaltos,2 Sep 1960(fl), Irwinet al. 47968 (BG, MO, NY); Mun. Cal9oene, BR.156, 53-72 km WNWof Cal9oene,11 Dec 1984(fl), Rabeloet al. 2971 (BG,MO);Mun.Macapi,nr.Serrado Navio,3 kmNNW of Serrado Navio (village), 4 Jan 1985 (fr), Rabelo et al. 3182 (NY).

2. Trymatococcus oligandrus(Benoist)Lanjouw (see Berg,Fl. Neotrop.Monogr.7: 214-216. 1972).

12. HelianthostylisBaillon(see Berg,Fl. Neotrop. Monogr.7: 216-218. 1972).

Helianthostylis

Distribution. In the threeGuianasandAmapa Beforethepublicationof themonograph a second (Brazil);innon-inundated lowlandforest. species had been discovered and mentioned in an adRepresentativeadditionalspecimensexamined. dendum.Moreover,Helianthostylissprucei appearsto GUYANA.E of Marayoku,5 Nov 1952 (fl-fr),FD 7550

be common in some partsof Colombia.

Revisedkey to the speciesof Helianthostylis 1. Basal lateralveins departingat (almost)rightangles (80?-90?);pistillodeslongerthan the perianth; 1. H. sprucei stigmasirregularlypapillatein the lower half...................................... 1. Basal lateralveins departingfromthe midribat acute angles (mostly40o60); pistillodesaboutas 2. H. steyermarkii long as the perianth;stigmasnot irregularlypapillate......................................

1. Helianthostylis sprucei Baillon (see Berg, Fl. Neotrop. Monogr.7: 216-218. 1972). AdditionalisotypeofHelianthostylissprucei:(BR). Additionalsynonyms:

Colombia and in western Ecuador;in non-inundated forest;at low elevations.

Representative additional collections examined. COLOMBIA. ANTIOQUIA: Mun. Zaragoza, Cgto. Helianthostylissalzedoi Romero,Mutisia 33: 5, t. Providencia,12 Feb 1971(fl), Soejartoet al. 2797 (COL, 2-6. 1970; Romero,Mutisia 38: 1, t. 1. 1972. GH);Anori,Providencia,29 Apr 1973 (fr), Soejartoet Mun.PuertoWilches, al. 3950 (F, MO).ANTIOQUIA/BOLIVAR: Border,38 km Type.Colombia.Santander: 26 Feb 1967 (a), Bruijn 1515 del W of Barrancabermeja, betweenLa Gomezand km 80 of Ferrocarril Mun.Morales,Cgto.Norosi, BOLIVAR: WAG). (COL, U, 8389 Atlantico,22 Apr 1960,Romero-Castanleda rd. to Tiquisionuevo,9-14 Apr 1985 (e), CuadrosV (holotype, COL). HelianthostylisschultesiiRomero,Mutisia33: 11, t. 2111 (BG, MO). CALDAS: La Dorada, (e), Duque7-8. 1970. Type.Colombia.Choc6:betweenRio Jaramillo 4003 (COL, F) CHOcO:2-4 km NW of CuricheandAlto Curiche,31 Jan 1967(e), Duke Teresita,18 May 1967 (fr), Duke 11026 (MO,NY, US); nr. Quibd6,Apr 1975 (a'), E. Foreroet al. 1135 (COL, 9578 (holotype,COL;isotype, US). Jct. of Rio Tigre and Rio Manso, HelianthostylisguggenheimiiRomero,Mutisia33: MO, U). C6RDOBA: Gentry et al. 63825 (MO);Mun.Ayapel, 27 Jul 1988 (st), 12, t. 9. 1970. Type. Colombia. Caqueta:Rio Caguan,S of Cartagena,27 Apr 1953, Romero- Palotal, 15 May 1949 (fr), Romero-Castanleda et al. Guadas,May 1972 (fr), Castanieda4117 (holotype,COL;isotype, MO). 1664 (COL).CUNDINAMARCA: P La nr.Caseriode Murca, Palma, Diaz s.n. Mun. (COL); 238: Helianthostylissuerpo Romero,Mutisia 3, t. 3. 1972; Jaramillo& Uribe-Uribe,Mutisia 39: 1300 m, 18 Jan 1972 (e'), Romero-Castaniedaet al. Mun. 11359 (COL,F, MO). GUAJIRA:15 km S of Mingueo, 12. 1976. Type. Colombia.Cundinamarca: La Palma, between Caserio de Murcaand La 21 Aug 1986(st), Gentryet al. 55421 (BG).SANTANDER: andCimitarra,3 Aug 1954 (e), Lamb Palma,1300 m, 18 Jan 1972,Romero-CastaniedabetweenErmitafno 156 (COL,MICH).VALLE:Rio Calima,La Trojita,19 et al. 11356 (holotype,COL;isotype,AAU). Feb-10 Mar 1944 (e'), Cuatrecasas16303 (F). Distribution (see Fig. 82.2). In the AmazonBasin, ECUADOR.Los Rios: Rio Palenque Biological largely in its middle partand in northernand western Station, 12 Feb 1974 (st), Gentry9862 (U).

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FLORA NEOTROPICA

PERU. AMAZONAS: Prov. Bagua, Yamayakat, 5 Feb 1996 (a), N. Jaramillo et al. 1106 (MO). PASCO:Prov. Oxapampa, Palcazu, 22 Jan 1984 (? + a"fl), Foster 7955 (BG, F, MO) BRAZIL. ACRE:Upper Rio Moa, nr. FazendaArizona, 24-30 Sep 1984 (st), Campbell et al. 8396, 8401 (BG). Mun. Mara Rio Japura,Canto Galo, 10 Jan AMAZONAS: 1991 (a), Ferreira et al. 10031 (NY); rd. Manaus-P6rto Velho, BR.3 19, km 380, 2 km S of Rio Jutai, 13 Oct 1974, Prance et al. 22863 (U). PA": Rd. Itaituba-Jacareacanga, km 85, 14 Nov 1978 (9), M. G. Silva et al. 3724 (NY); rd. Cuiaba-Santarem, BR.163, km 1135, nr. Igarape Natal, 16 Nov 1977 (2 fl-fr), Prance et al. 25509 (U). RONDONIA:Mun. Porto Velho, Usina Hidroelectrica

the receptacle basally attached, (sub)ovate, passing gradually into numerous subulate to (sub)peltate ones.Infiuctescencesglobose,2-2.5 cm diam., interfloral coveredwith ratherdense uncinatehairsand scattered, but towardsthe base with ? crowded,persistentstaminate flowers.

Tree,to 18 m tall. Leafytwigs 1-2.5 mm thick, minutely puberulous and with sparse to rather dense, longer, uncinate hairs. Lamina elliptic to lanceolate, 5.5-17.5 x 3.5-6 cm, broadestin or above the middle, subcoriaceousto coriaceous;apexacute (sub)equilateral, to acuminate or to obtuse; base (sub)acute; margin (sub)entire;both surfaces glabrous or a few uncinate hairson the main veins and margin;venation slightly prominentabove,prominentbeneath;lateralveins 6-10 pairs,the lower ones departingfromthe midribat acute angles,40'-60'(-80o); tertiaryvenationpartlyscalariform;petiole0.3-0.7 cm long, minutelypuberulousand with longer uncinate hairs; stipules 0.3-0.6 cm long, appressed-puberulousand with or without uncinate hairs.Inflorescences bisexual, ovoid, ca. 1 cm diam.; peduncle 0.5-0.7 cm long, puberulous; receptacle densely andminutelypuberulousandwith longeruncinatehairs;staminateflowersca. 50, mostlyon the lower partof the receptacle;pedicel ca. 1 mm long; perianth ca. 2 mm long, 3-fid, minutelypuberulous;stamens3; filaments 2-2.5 mm long, the upper part deflexed at and afteranthesis;anthers0.3-0.4 x 0.2-0.3 mm, the connectiveratherbroad;pistillode ca. 1.5-2 mm long, branchedor unbranched;pistillate flower one, almost completelyimmersedin thecenterofthe receptacle;stigmas ca. 2 mm long, sparselyhairy;bractsat the base of

modelof Treesor shrubs, with the architectural "Cook" (Halle&Oldeman,1970),dioeciousormono-

Distribution (see Fig. 82.4). In the northernpart of the upperAmazon Basin, Venezuela (Amazonas), andBrazil (NW Amazonas);in non-inundatedforest, locally common.

Specimensexamined. VENEZUELA. AMAZONAS: Depto.Rio Negro,rd. San Carlos-Solano,km 2, 29 Jan Samuel, 13 Aug 1987 (fl), Dionizio et al. 141 (NY). ca. 1992(st),Aymardet al. 9837 (BG);rd.Yavita-Maroa, 2.5-10.5 km W of Yavita,21-28 May 1996 (st), Aymard The featuresof the additionalcollections from Co- et al. 10939, 11006, 11049, 11061, 11076, 11078, and lombia are largely covered by the species description 11127 (MO);Depto.Rio Negro,Rio Mawarinuma, Cerro in the monograph.The identityof the sterilecollection de La NeblinaBase Camp,7 Feb 1985 (st), Boomet al. from Ecuadoris not quite certain. 5679 (BG);betweenYavitaand Pimichin,21 Nov 1953 (fl), Maguireet al. 36315 (NY, U). cimarr6n; Local names.Colombia:Caldas:guAimaro Rio Tea, 12 Jun 1976 (fr),L. BRAZIL. AMAZONAS: Cordoba:castaiio;Cundinamarca:castaiio,suerpo. Coelho455 (INPA);Sao Paulode Olivenga,3 Apr 1944 (fr), Ducke 1537 (F, IAN, NY); Mun. Sao Paulo de Oliven9a,S of the city, 24 Nov 1986 (fl), Ferreiraet al. 2. Helianthostylis steyermarkii C. C. Berg,Acta Bot. 8506 (BG, INPA,US); IgarapeJandiatuba,12 Jan 1949 Neerl.21: 99, t. 1. 1972.Type.Venezuela.Amazonas: (fr), Fr6es 23892 (IAN, NY) and 23898 (IAN). betweenYavitaandPimichin, 1 km fromYavita,21 Oct 1970 (fr), Steyermarketal. 102860 (holotype, U; isotypes, MO, NY, VEN). IV. CASTILLEAE

ecious, withoutuncinatehairs.Leaves alternate,in spirals or (on flowering branches)distichous;laminaentire,venationpinnate,marginentireor dentate;stipules fully amplexicaulto semiamplexicaul.Inflorescences capitate, discoid to cup-shaped with an involucre of basallyattachedbracts,unisexual,multi-to uniflorous; interfloralbractslacking(interfloralbract-likestructures apparentlyderived from tepals). Staminateflowers complete and4-merous or with reductionsof the perianth(to absent)and/orstamens,sometimes with more

than4 tepals;pistillodeusuallyabsent.Pistillateflowers freeor connate, tepals 4, mostlyconnate;ovary usually? adnateto the perianth,stigmas2 andequalin wholewiththepelength.Fruitforminga drupaceous

rianth,withotherflowers,oralsowiththereceptacle; seed large,withoutendosperm;embryothick,with equalcotyledons. Thetribecompriseseightgeneraand55 species.In additionto thegeneraandspecieslisted,discussedor OldWorld genera: treated below,therearetwomonotypic Antiaris (fromAfricato Oceania) and Mesogyne (in butin WestGondwanan, Africa).Thetribeis essentially contrast totheDorstenieae, inAfrica. poorlyrepresented

TAXONOMIC TREATMENT

245

aresylleptically principle, (plagiotropic) lateral branches formedin eachof theleafaxils.Onthesebranches the leavesaredistichously arranged. Thesebranches bearthe evenwhenstill flowers(axillary) andareshed(sometimes bearingleaves)byabscission. Thisleavescharacteristic concavescarsonthemainaxes,andtheshedbranches haveaphallus-shaped base.All axillarybudsareused toformlateralbranches, butlateralaccessory budsmay produce additional (orthotropic) branches. Thephenomenonof self-pruning is mostapparent inthelarge-leaved suchasin specieswiththickandlonglateralbranches, a PerebeaguianensisorP mollis.Mostspeciesproduce ineachleafaxil,butinseveralNaucleopsis branch occupied in Trophis,Olmediaaspera was reducedto a lateral synonymof Trophiscaucana,treatedin the present species(e.g.,N. ulei) theplagiotropic branches areprorevisionof Trophis. ducedin onlysomeof theleafaxils.Lateralbranches Since the publication of the monographon arenotformedinthemonocaultreeletsof Naucleopsis have themonotypicgenusOlmediahasbeen stipularis.Theneotropical speciesof theCastilleae "Olmedieae," buds.TheAfricangenusMesogynehas excluded,andeightnewspeciesandtwonewsubspe- lateraldormant budsinthesameposition,whereastheyare cies havebeendescribed:Perebeamennegae(Berg, thedormant 1978b), Naucleopsis capirensis,N. herrerensis,N. hu- axillaryinthepaleotropical genusAntiaris. As theabsenceof interfloral bractsis characteristic milis, N. straminea, N. velutina, N. naga subsp. mersuchasinterstaminal idionalis, N. ulei subsp. puberula (Berg & Franco, forthetribe,theuseof indications 1996),Pseudolmediagentryi,andP.manabiensis(Berg, bractsforCastillaandPseudolmediaandpseudobracts 1998a).Maquiracostaricana hadtobereducedto sub- forNaucleopsis hasbeenincorrectand/orconfusing. species(Bergin Berg& Franco,1996).Twonames, Bract-likestructures occuringamong"free"stamens Ficusfaginea and Trophisglabrata, were transferred (in CastillaandPseudolmedia)oramong"free"styles fromothertribes,thelattertransferleadingto a name (in Naucleopsis) have to be regardedas "detached" scalesor asinterstaminal change:Pseudolmediaglabrata.Duringthepreparation tepals.Theycanbe indicated of thesecorrections andadditions someadjustments had freepartsof tepalsrespectively. tobemade:inPseudolmediacombining ofP guaranitica of Castilla Thefactthatthestaminate inflorescences withP. laevigata,andof P mururewithP macrophylla, andPseudolmedia arecloseduntilanthesiscanbe reandestablishment of anothernameforoneof thesec- latedto thedisintegration of thebasicfloralconfigurations;in PerebeaglabrifoliaandP mollis,thereinstate- tion,andthereduction of thenumberof tepals,andthe oftwonewcom- lossof theprotectivefunctionof theperianth. mentof P rubraandtheestablishment Thefact binations,P. mollis subsp. lecithogalacta andP rubra that staminateinflorescences are also closed in subsp.glabrifolia; andin Naucleopsis,the description Naucleopsis canbe relatedto somedegreeof disinteof a newsubspecies,N. uleisubsp.subandina,andthe grationof thebasicarrangement of thefloralparts.The reductionof Naucleopsisamarato a subspecies. is in amoreadvancedstatein thepistildisintegration inrecentcol- late inflorescencesin Naucleopsis. In Perebea sect. Newtaxaarestillbeingencountered lections.Thelist of unnamedcollectionsunderNau- Noyera (P. mollis andP rubra) the innerinvolucral thatcurrent andfuturecollect- bractscovertheflowersbeforeanthesisin bothstamicleopsisis anindication newtaxa, nateandpistillateinflorescences. todescribe ingactivities mayprovidematerial Theflowersarequite atleastin thisgenus.Dueto furtherbotanicalexplora- normalin thissection.Theinnerinvolucralbractsare of manyspeciesprovedto elongateandnarrowed tion,rangesof distribution in contrast tothoseof thestamibelargerthanindicated andmappedinthemonograph. nateinflorescences of Naucleopsis andPseudolmedia. Althoughrangeextensionswill be indicatedbelow, Inthesegeneratheyarebroadened. detailscanbe foundin theFloraof theGuianas(Berg An accountof thewoodanatomyof theCastilleae in G6rts-vanRijn,1992),theFloraof Ecuador(Berg ("Olmedieae") has beenpresentedby Mennegaand in Harling& Andersson,1998a),andFloraMeso- Lanzing-Vinkenborg (1977).Thispublication supports americana (Berg,submitted). theexclusionof Olmedia. indicated asthe Thecharacteristic treearchitecture, modelof "Cook"(Halle& Oldeman,1970),hasbeen indetailbyBerg(1977a).Onthe(orthotropic) "Olmedieae"(see Berg,Fl. Neotrop.Monogr.7: 1described In inspirals. 156. 1972.) mainaxesofthetree,theleavesarearranged Fieldwork carried outafterpublication of themonographon "Olmedieae" (Berg,1972)revealedthatall generaof thisgrouphavethearchitectural treemodel of "Cook"(Oldeman &Halle,1970;Halleetal., 1978) exceptforthemonotypicgenusOlmedia.Moreover, the stamensin this genusareof the Urticaceaetype, characteristic forthegenera(presently) arranged inthe tribeMoreae. Thisledtotheexclusionof Olmedia from thetribeandconsequently a changeof thenameforthe tribeto Castilleae(Berg, 1977a).Subsequently, the genusOlmediawasreducedto a sectionof thegenus Trophis (Berg,1988).Astheepithetasperawasalready

FLORANEOTROPICA

246

Olmedia Ruiz& Pav6n(see Berg,Fl. Neotrop.Monogr. P spuria, anothername has to be established for the 7:13-17. 1972.) othersection of the genus. The monotypicgenus has been reducedto a section of the genus Trophisand is treatedagain in the present monograph.

Pseudolmedia

Pseudolmedia Treculsect. Olmediopsis (Karsten)C. C. Berg, comb. et stat.nov., based on Olmediopsis Karsten,Fl. Columb.2:17. 1862.Type.Olmediopsis obliquaKarsten=P laevis(Ruiz&Pav6n)Macbride.

SectionPseudolmedia,characterizedby the absence of hairson the anthersandby oblongoid-capitatepluricellularhairson the lower surfaceof the lamina,comSeveral changes had to be made: the inclusion of prisesP gentryi,P glabrata, P manabiensis,P rigida, inP laevigata;theadoption and P. spuria. Section Olmediopsis, distinguishedby Pseudolmediaguaranitica of anothernameforthe speciesrecognizedasP oxyphyl- the commonpresenceof hairsat the apex of the anthers laria, as Trophisglabrataprovedto be conspecific;the and by globose-capitatehairs on the lower surfaceof reductionof E murureto a synonymof P macrophylla; the lamina,comprisesP hirtula,P laevigata,R laevis, andthe recognitionof two new species, P gentryiand andP macrophylla. P manabiensis (Berg, 1998a). The pluricellularhairs are sometimes brownishto In the monograph,Pseudolmediawas lectotypified reddish, in particularin Pseudolmedia gentryi and P byPfernuginea(Poeppig& Endlicher)Trecul=P laevis rigida. These hairsaremostly whitish andmore easily (Ruiz & Pav6n) Macbride.However,the genus hadal- detectablein driedmaterial.Theinflorescencesaresomereadybeen lectotypifiedwith P havanensisTrecul=R timessubsessile(in P. rigida subsp.rigida andP manaspuria(Swartz)Grisebachby BrittonandWilson(1924). biensis) and the pistillate ones occasionally contain As Pseudolmedia sect. Acomandra Engler includes more thanone flower (P laevigata). 13. Pseudolmedia Trecul (see Berg, Fl. Neotrop. Monogr. 7: 17-38. 1972).

Revisedkey to the speciesof Pseudolmedia 1. Pluricellularhairs (minute!)on the lower surfaceof the laminaoblongoid-,ellipsoid-,or ovoidcapitate;anthersalways glabrousand apiculate. 2. Stipulesto 1.2 cm long. 3. Hairson leafy twigs and laminapartlycrinkled. 4. Tertiaryvenationreticulateto subscalariform; laminaoblongto subobovate;interstaminal W Ecuador scales filiformto subspathulate; 6. P. manabiensis .......................................................... 4. Tertiaryvenationpartlyscalariform(or in small leaves to subscalariform); laminaobscales spathulate;Mexico and CentralAmerica....... 5. P. glabrata long to subovate;interstaminal 3. Hairson leafy twigs and laminastraightor absent. 5. Tertiaryvenationreticulate(or in large leaves to subscalariform). 6. Hairson the leafy twigs aboutequal in lengthor absent;Mexico to Panamaand to GreaterAntilles............................................................. 8. P. spuria 6. Hairson the leafy twigs differentin length;SouthAmericato Costa Rica............. 7. P. rigida 5. Tertiaryvenationpartlyscalariform(or in small leaves to reticulate). 7. Involucralbractsi densely (sub)sericeous;fruitingperianthusually densely hairy (yellowishsericeousor subvelutinous);Mexico to Costa Rica .............. ............... 5. P. glabrata 7. Involucralbractsrathersparselyappressed-puberulous; fruitingperianthsparselyhairy South Americato Costa Rica.................................7. P. rigida (whitishappressed-puberulous); 2. Stipules 1.2-2.5 cm long. 8. Involucralbractsi densely subsericeous;hairson leafy twigs and laminausually partly 5. P. glabrata crinkled;Mexico to CostaRica................................................................ 8. Involucralbractsrathersparselyhairy;hairs on leafy twigs (and lamina)straight. 9. Tertiaryvenationdistinctlyscalariforn;uppersurfaceof the laminahairyon the midrib (or also the lateralveins); laminamostly ca. 10-25 x 5-10 cm; (sub)montane ..........9. P. gentryi 9. Tertiaryvenationreticulate(to subscalariform); uppersurfaceof the lamina (sub)glab7. P. rigida rous; laminamostly 5-15 x 2-6 cm; lowlandsand (sub)montane ................ ................. 1. Pluricellularhairs (minute!)on the lower surfaceof the laminaglobose-capitate;anthersusually hairyat the apex, not apiculate. 10. Leafy twigs with very shortpatenthairsintermixedwith distinctlylongerpatentto appressed ones; tertiaryvenationlargely scalariform.

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11. Lamina? stronglyinequilateral. 12. Longhairson theleafytwigsstraight,yellowto brownish; northern SouthAmerica.......... .......1.P. laevis 12. Longhairson the leafytwigscrinkled,paleyellow;E Brazil.............................................. 2. P. hirtula 11. Laminaslightlyinequilateral to equilateral. 13. Laminausually15-30 cm long;terminalbudnot swollen;stipulesmostly1-1.5 cm long;involucralbractsdenselyyellow sericeous;fruitingperianthbrownishvelutin3. P. macrophylla ous;AmazonBasin........................................................ 13. Laminausually5-15 cm long;terminalbud? swollen;stipulesto 1(-1.5) cm long; involucralbractsnotdenselyhairy;fruitingperianthhirsute,sometimesonly nearthe apex;northernSouthAmerica........................................................ 1. P. laevis 10. Leafytwigswithhairsof aboutequallength;tertiaryvenationreticulateor largelyscalariform 14. Tertiaryvenationreticulate,to subscalariform in relativelylargeleaves;laminamostly 5-15 cm long;indumentum mostlywhitish;Panamato E BrazilandParaguay........... ......... 4. P. laevigata 14. Tertiaryvenation(largely)scalariform; laminamostly15-30 cm long;indumentum yellow; AmazonBasinandE Brazil............................................................ 3. P. macrophylla 1. Pseudolmedia laevis (Ruiz & Pavon) Macbride (see Berg, Fl. Neotrop. Monogr. 7: 21-23. 1972).

Clark et al. 3281 (QCNE). Prov.Mariscal Caceres, Dtto. TocaPERU. HUANUCO: che Nuevo, Fundo Melodia, 16 Jun 1974 (2 fl), Schunke V Additional type of Olmediopsisobliqua: Karsten 7450 (NY). PASCO:PalcazuValley,Iscozacin, 27 Mar 1986 Prov. Coronel Portillo, (st), Salick 7281 (NY). UCAYALI: s.n., d'(isotype, O). Bosque Nacional Alexander von Humboldt, rd. PucallpaDistribution. In northernSouth America, east of Tingo Maria, km 86, 14 Aug 1980 (V?), R. Oliveira 17 the Andes, throughoutthe Amazon Basin, the Guiana (MO). region, andnorthwesternVenezuela;in non-inundated BRAZIL. ACRE:Mun. Brasileia, Seringal Porongaba, 30 May 1991 (? fr), Daly et al. 6805 (MO, NY). AMAforest; to 1100 m. Representative additional specimens examined. ZONAS:Between Maloca and Rio Cauaburi,31 Dec 1965 (d'),N. T Silvaet al. 60784 (NY,U). MATOGROSSO:Rio FRENCHGUIANA.Sail, 22 Jun1988(st), Gentryet al. Aripuana,nr. Humboldt Center, 12 Oct 1973 (V?), Berg et 63056 (NY). al. P18437 (MO, NY). PARA:Rio Jari, Monte Dourado, 3 Depto.Rio Negro, 1 km VENEZUELA.AMAZONAS: Dec 1967 (a), Oliveira 3811 (NY); Rio Xingu, Glaba E of Cerrode La NeblinaBase Camp,Rio Mawarinuma, Bacaja, just below mouth of Rio Bacaja, 25 Nov 1980 (? 25 Feb 1984(st),Liesner16248(MO,NY);SierraParima,

Rio Matacuni,18 Apr-23May 1973(? nr.Simarawochi, fl-fr),Steyermark 107192(NY,U).APURE: ReservaForestal 6 Apr1968(? SanCamilo,7 kmE of LaCeibadel Jordacn, et al. 101931(U). BARINAS: BetweenLa fl), Steyermark andEl Curito,25-26 Aug 1966(st),Steyermark Esmeralda et al. 96550 (NY). PuertoMorona,E of ECUADOR.MORONA-SANTIAGO: Rio Morona,1 Oct 1975(st),Littleet al. 529 (LOJA,MO, NY,QAME,QCNE);Bomboiza,nr.Misi6nSalesiana,10 May1986(2 fl-fr),Baker6984(BG,QAME,QCA,QCNE). 1-15 Feb1986(9 NAPo:Afnangu,ParqueNacionalYasuni, fl-fr),Korninget al. 47566(AAU,MO,NY,QCA,QCNE); ReservaBiol6gicaJatunSacha,8 kmE of Misahualli,17-

fl), Pranceet al. 26438 (MO,NY).

BOLIVIA. BENI: Prov. Ballivian, rd. Yucumo-Rurrenabaque, km 35, Mar-Jun 1990 (st), D. N. Smith et al. 14386 (MO, NY); Prov.Vaca Diez, 18.4 km E of Riberalta, 21 Sep 1981 (a'), Solomon6367(MO, NY). COCHABAMBA: Prov.Carrasco,EstacionValle del SajtaUMSS, 6 Apr 1988 (st),Fedlmeier4 (MO,NY). PANDO:Rio Madeira, 12 km above Abuna, 20 Jul 1968 (a'), Prance et al. 6219 (NY). SANTACRUZ:Prov. Guarayos, 4 km N of Perseverancia,9 Sep 1990 (? fl-fr), Nee 38686 (MO, NY); Prov. Ichilo, Reserva Forestal Chore, Rio Ibabo, 16-18 Aug 199 (e),

Neill er al. 9382 (MO,NY).

The longer hairs are usually more or less patent, in particularon the stipules. However, in several colQCNE),4490 (BG, QAME);rd. Hollin-Loreto,km 40, lections from the southernpartof the range of distriHuamani,1200m, 19Oct1989(? fl), Palacios4735(AAU, bution,in particularin Bolivia andMato Grosso (Bra19- zil), the longer hairs are often ? appressed, even on Moretecocha, BG,MO,NY,QAME,QCNE).PASTAZA: 21 Mar1991(? fl-fr),Gudiuo1386(NY,QCNE);rd.Coca- the stipules. Such specimens can be confused with border),1-9 Apr Auca,km 115 (10 kmS of Napo/Pastaza those of Pseudolmedia macrophylla,in particularits 1989 (v?), Zak et al. 4236 (BG, MO, NY, QCNE). Canton Aguarico, Reserva Faunistica small-leaved form. However, in that form of P. SUCUMBlOS: nr.Zdbalo,21 Nov 1991(9 fl-fr),Palacioset al. macrophyllathe indumentum(of the leafy twigs) does Cuyabeno, 9427 (BG); ReservaFaunisticaCuyabeno,nr. Laguna not consist of very short patent hairs and distinctly longerones. The formof P laevis with appressedhairs Grande, 11 Mar 1990 (st), Balslev et al. 97213 (AAU, often has relatively narrowlaminas;it may co-occur Jamboe Bajo, E border of QCA).ZAMORA-CHINCHIPE: NationalPark,1100m, 3 Nov 1996(st),J. L. with the typical form. Podocarpus 28 May 1989 (d), Palacios 4322 (BG, MO, QAME,

248

FLORANEOTROPICA

InSierraLaMacarena (Meta,Colombia)thefruitingperianth is ellipsoidoroblongoid.Theellipsoidperianthsturnorange-red andtheoblongoidonesyellowgreen.Differentfruitingperiodshavebeenobserved forthetwotypesof fruitingperianth (PabloStevenson, Bogota,pers.comm.).A similardifferencein colorof thefruitingperianthis foundin someotherMoraceae

ca. 0.8-1.5 cm diam., concave; involucralbractsin 68 series,reniformto suborbiculateto ovateto lanceolate, obtuse (to mucronate), yellow-sericeous; filaments (0.05-)0.1-0.4 mm long; anthersca. 1.5-2 mm x 0.40.6 mm,with apicalhairs;interstaminalscales (= tepals) 3-8 mm long, lanceolateto linear,hairy.Pistillateinflorescences 0.2-0.3 cm diam.; involucral bractsin 3-6 species. series, reniformto ovate, obtuse to acute, yellow-sericeous or yellowish to whitish appressed-puberulous; perianth ca. 2-3 mm long, brownish-velutinous to 2. Pseudolmediahirtula Kuhlmann(see Berg, Fl. subsericeous;stigmas7-9 mm long. Fruitingperianth Neotrop.Monogr.7: 23-26. 1972). subglobose to ellipsoid, 1-2.2 cm long, brownvelutinous to sparsely appressed-puberulous,somehasbeenuncertain. Thedateof publication Accordtimes only at the apex.

ingto Carauta (1994),it is 1939insteadof 1940. No otherchangesto be reported.

Distribution (see Fig. 82.6). In the AmazonBasin, mainly the southempartandwestem part,also in eastem Brazil(Bahia);in non-inundatedforest;to 1600 m.

3. PseudolmediamacrophyllaTrecul(see Berg,Fl. Representative additional specimens examined. Neotrop.Monogr.7:26-28. 1972,incl.P murure). COLOMBIA.VAUPAS:Rio Papuri,betweenPapuriand CafioYapu', Apr 1977 (? fr), Patmoreet al. 62 (GH);Rio Additionalsynonym: Vaupes,Pozo Verde,Nov 1975 (st), Roa T 200 (INPA). PseudolmediamurureStandley,Publ.FieldMus. ECUADOR.NAPO: ParqueNacionalYasuni,Pozo Nat.Hist.,Bot.Ser.17: 186. 1937. Newcollectionshavemadeit moreandmoredifficultto separateP macrophylla, typicallywithrobust leafytwigs,relativelylargeleavesanddenselyhairy asfoundintheupperAmazon fruitingperianths, Basin fromPmurure, (PeruandEcuador), typicallywithmore slenderandlesshairyleafytwigs,relatively smallleaves andsparselyhairyfruitingperianths,as foundin the lowerAmazonBasin(ParaandMaranhao). As faras canbe concludedfrommaterialathand,in particular collectionsfromnorthernBoliviaandfromAcreand Rondonia(Brazil),the two entitiesappearto be extremesof a moreorlessdistinctlyclinalvariation from thelowerAmazonBasin(Para)throughthe southern partof themiddleAmazonBasinto Boliviaandfrom therenorthwards to Ecuador.

PetroleroDaimi2, 16-19 Jan 1988(?), Cer6n3459 (BG, QAME,QCNE). PASTAZA:Rio Curaray,Tonliampari, El Guiquita et al. 110 (QCA). MORONA-SANTIAGO: Pincho,61 km NE of Zamora,14 Sep 1975 (?), Littleet al. 337 (LOJA,QAME,US); 16.7 km NE of Taisha,1419 Aug 1976 (st), Ortega U. 43 (US). ZAMORACHINCHIPE: Nr. Sabanilla,19 km E of Loja, 1600 m, 4 Sep 1975,Littleet al. 226 (LOJA,NY, QAME,QCNE, US);betweenZumbiandPaquisha,1000m, 28 Oct 1991 (? fl-fr), Palacios et al. 8803 (BG, COL, QCNE). PERU.AMAZONAS: Rio Cenepa, Huampami, 3 Nov

1972(v),Kayap40 (AAU,NY);Prov.Bagua,Dtto.Imaza, Yamayakat, Feb 1995 (?), R. Vdsquez et al. 19342 (BG). Cuzco: Prov.La Convenci6n,nr. Rio Klause and Rio Maputunuari,18 Jun 1968 (9 fl), Dudley 10290B (F). JUNIN:Rio Negro, N of Satipo, 20 Aug 1960 (Q), Woytkowski 5862 (MO, NY). LORETO:Prov.Requena, JenaroHerrera,22 Aug 1970 (st), Lao M. 14 (U); Prov. Ucayali, Dtto. VargasGuerra,Orellana,Dec 1982 (st), Reviseddescription: ReynelR. 1013 (MOL,U). MADREDEDIos: Prov.Manui, Tree,to 35 m tall.Leaffytwigs(1-)2-9 mmthick, ManuNationalPark,CochaCashuStation, 18 Aug-i9 withyellowto brownishappressed topatentfirmhairs, Sep 1989 (??), Nuniezet al. 11252 (MO,NY); nr. conwithmuchshorterpatenthairs;scars fluenceof Rio Tambopataand Rio La Torre,39 km SW oftenintermixed of the stipulesmostlyprominent.Lamina oblongto of PuertoMaldonado,14 Jul 1987 (?), S. F Smithet al. subobovateor to subovate,(8-)10-33 x 3.5-13 cm, 1018 (MO, US). PASCO:PalcazuValley,Iscozacin,26 slightlyinequilateral, (sub)coriaceous; apexacuminate; Mar1986(st), Salick7297 (BG,NY). PUNO:Rio Tavara, baseacuteto obtuse(to subcordate); uppersurfaceal- 20 May 1992 (st), Gentryet al. 76876 (MO). BRAZIL.AMAZONAS: Rd. Manaus-P6rto Velho,km mostglabrousto puberulous orhirtellousontheveins; 515, 24 Apr 1976, Monteiro et al. 941 (INPA). BAHIA:

lowersurfaceappressed-pubescent to -puberulous to Ilheus, CEPEC, 8 Jan 1970 (a), Santos 542 (CEPEC, sometimesslightlyrugose;lateralveins14- U). MATOGROSSO:Mun. Sinop, BR.080, km 142, 4 hirtellous, 21(-25) pairs;tertiaryvenationlargelyscalariform; Aug 1985 (? fr), Ferreiraet al. 6334 (BG, MICH,NY); petiole0.3-2 cm long;stipules(0.4-)1-1.5 cm long, Rio Arupuana, HumboldtCenter,24 Oct 1973, Berg et yellow-sericeous, caducous.Staminateinflorescences

al. P.19864 (AAU, MO, NY); Mun. Cuiaba,Chapada

TAXONOMICTREATMENT

249

dos Guimaraes,Veude Novia, 21 Oct 1985 (2 fr), Tho- Grande,10 rd. mi fromcontinentaldivide, 6 Dec 1987 mas et al. 4489 (BG, MO,NY). MARANHAO: Ca.45 km (? fr), McPherson11824 (BG, MO). DARIEN: Cerro S of Itauna,18 Oct 1980 (a), Daly 677 (INPA,MICH, Campamiento, trailCoasi-Cana,E of TresBocas,29 Apr MO, NY). PARA:Rio Jani,Monte Dourado,Pilao, 22 1968 (2 fr), Kirkbride1227 (MO, NY). PANAMA: El Nov 1968 (c), N. T Silva 1431 (NY, U). ROND6NIA: Llano-Cartird., km 8.7, 3 Sep 1977 (st), Berg et al. Mun. Costa Marques,9.5 km E of Forte Principeda 404 (BG). Beira,28 Mar 1987 (9 fl), Nee 34519 (BG, MO, NY); COLOMBIA.AMAZONAS: Rio Caqueta, nr.AraraVilhena,25 May 1979(a'),M. G. Silvaet al. 4660 (NY). cuara,11 Sep 1989 (?), Londonoet al. 1326 (NY); nr. BOLIVIA.BENI:Prov.VacaDiez, Alto Iv6n, 1 Feb Leticia,19 Jun 1991 (i?), Rudaset al. 2146 (BG, MO). 1984 (st), Boom 4364 (NY); Prov. Yacuma,rd. Rio AMAZONAS/VAUP]S: Rio Apaporis,betweenRio CanaCuriraba-La Pascana,14 Sep 1987 (st), Sobrevilla1700 nan andRio Oacoa,1-15 Dec 1951(v?), Garcia-Barriga (F). LA PAZ.Prov. Sud Yungas,Chulumani,Rio San 13830 (NY). ANTIOQUIA: Mun.San Luis,Vrda.Arauca, Jos6, 7 Aug 1983 (c), Beck 8557 (BG, MO, NY). 1 Mar 1990 (2 fr), Cardenaset al. 2442 (MO); Mun. PANDO: Prov.Manuripi,rd. Porvenir-Puerto Heath,km Anori,Providencia,20-25 Feb 1976 (? fl-fr), Soejarto 26, 16 Aug 1979 (9), Meneces902 (NY); Prov.Madre et al. 4524 (F, HUA,MO, NY). BOLIVAR:Quebradala de Dios, SantaFe, Rio Manupare,1 Jun 1987,Solomon Culebra,CainoCaribona,betweenjct. of Rio Cancaand 16832 (MO,NY). SANTA CRUZ:Prov.Guarayos,ca. 6 Rio Magdalena,4 May 1987 (st), Gentryet al. 57296 km W of Perseverancia,10 Sep 1990 (V?), Nee 38711 (BG). CHOC6:UpperRio Truand6,La Teresita,19 Jan 1974 (v), Gentry9379 (MO,NY, U). C6RDOBA: Jct. of (BG, MO, NY). Rio TigreandRio Manso,26 Jul 1988(st), Gentryet al. The easternformof Pseudolmediamacrophyllaand 63887 (BG, MO). META: Serraniade Chamusa,Rio P. laevis can have leaves of about the same size. The Duda,Feb 1988(? fr),Stevenson31 (MO). SANTANDER: two taxa can usuallybe distinguishedby the indumen- Mun. PuertoWilches, between La G6ez and km 80 of tum: appressed with hairs not distinctly different in Ferrocarrildel Atlantico, 23 Apr 1960 (??), Romerolengthon the stipulesandleafy twigs in ? macrophylla, Castanieda8402 (MO, NY). VALLE: Bajo Calima,rd. anddistinctlydifferentin lengthandusuallyi patentat to JuanchacoPalmeras,10 Jul 1984 (??), Gentryet al. EstacionBiol6gicaCapuru, least on the stipules in P laevis. The lower surfaceof 47822 (BG, MO).VAUP-ES: 24 Aug 1989 (? fl), Defler 326 (MO);Rio Vaupes,bethe laminais oftendistinctlyrugosein P laevisandonly tween Miraflores and Balso, 15 Feb 1944 (? fl-fr), sometimes slightly so in P macrophylla. GutierrezV et al. 803 (GH) VENEZUELA.AMAZONAS: Depto.Atures,8 kmNW of Yutaje,1000-1050 m, 3 Mar 1987 (d), Liesneret al. 4. Pseudolmedia laevigata Trecul (see Berg, Fl. 21542 (BG, MO). ZULIA:Dtto. Perija, nr. Estaci6n Neotrop. Monogr. 7: 28-31. 1972, incl. P. HidrologicaAricuaisa-Pie, 25 Feb-3 Mar 1982 (Q), Buntinget a!. 10837 (NY). guaranitica). GUYANA.UpperMazaruniR. region, Kamarang, Additionalsynonym: 30 Jun 1987 (st), Boom 8369 (BG). PseudolmediaguaraniticaHasslerhas to be put into FRENCHGUIANA.Saul,nr.EauxClaires,24 May the synonymyof P. laevigata. 1992 (2 fr), Mori et al. 22279 (BG, NY). ECUADOR.LOJA: CantonMacara,jct.of rd.to PapaIncludingPseudolmediaguaraniticahardlyaffects yal and QuebradaPapayal,2 Jan 1989 (??), Palacios the circumscriptionof P laevigata, as the differences 3355 (BG). MORONA-SANTIAGO: CentroShuarYukubetweenthe two taxaas indicatedin the monographare tais, 12Mar1990(? fl), Bennettet a. 4050 (BG,QCNE); very small. Recognition of the two taxa was based on PuertoMorona,Rio Morona,30 Sep-2 Oct 1975 (v?), phytogeographicalconsiderationsratherthanon mor- Little et al. 560 (COL, LOJA, QAME, US). NAPO: phological ones. Morerecentcollecting bridgedmore ReservaFloristicaEl Chuncho,5 km N of Coca, 12-14 or less the gap between the areasof distribution,pre- Oct 1987 (st), Neill et al. 7890 (BG, QAME);Parque NacionalYasuni,Pozo PetroleroAmo 2, 9-13 Jan 1988 sumed to be disjunct. (v), Palacios2406 (BG,NY,QAME);ReservaBiol6gica Distribution (see Fig. 83.3). FromPanamato the JatunSacha, 8 km E of Misahualli, 8 Sep 1992 (st), PacificCoastalregion of ColombiaandEcuador,to the Palacioset al. 10427(BG,QCNE).PASTAZA: Rd.CocaMagadalenaValleyandnorthwestern Venezuela(Zulia), Auca, 110 km S of Coca, 10 km from Rio Tigiuino,7 in the Amazon Basin, in the Guiana region (eastern Jan 1989 (v), Palacios et al. 3355 (MO, NY, QAME, Venezuela,Guyana,andFrenchGuiana),centralBra- QCNE);Pozo PetroleroRamirez,20 km S of Curaray, zil (Minas Gerais), easternBrazil (Bahia), and Para- 21-28 Feb 1990 (? fr), Zak et al. 5071 (BG, QCNE). SUCUMBIOS:ReservaFaunistica Cuyabeno,10 kmdownguay; in non-inundatedforest;to 1500 m. streamfromoutlet of LagunaGrande,4 Apr 1989 (e?), Balslevet al. 84813 (AAU,BG, QCA);Dureno,28 Dec Representative additional specimens examined. PANAMA. BOCAS DEL TORO: Nr. hwy. to Chiriqui 1988 (v), Gentryet al. 64343 (AAU, NY). ZAMORA-

250 CantonNangaritza,Miazi, 21 Oct 1991 (v?), Palacios et al. 8570 (QCNE). PERU.LORETO: Prov.Requena,JenaroHerrera,24 Sep 1982(?), Encarnacion 26133(NY,US);Prov.Maynas, Puerto Almendras, Rio Nanay, 19 Feb 1985 (v), R. Vasquezet al. 6251 (NY). HUANUCO:Prov. Pachitea, Sungaro,W of PuertoInca, 14 Sep 1982 (? fl), Foster 8784 (MO,NY). MADRE DE Dios: Prov.Manui,Parque Nacionaldel Manu',CochaCashuStation,Aug-Sep 1989 (? fl), Nuiiezet al. 11438 (MO).PASCO:PalcazuValley, Iscozacin,27 Mar1986(st),Satick7366(MO,NY).PuNo: Rio Tavara,17 May 1992(st), Gentryet al. 76731(MO). SAN MARTIN: Prov. MariscalCaceres, Dtto. Tocache Nuevo, FundoMelodia, 16 Jun 1974 (v), SchunkeV 7461 (MO, NY). BRAZIL. BAHIA: Rio Gongoji,1 Oct-30 Nov 1915, FEDERAL: 20 km S of Brasilia, Curran20 (US).DISTRITO on rd.to Belo Horizonte,23 Sep 1965 (? fl-fr),Irwinet al. 8606 (MO, NY, U). MATO GROSSO:Rio Aripuana, rd. HumboldtCenter-RioJuruena,km 4, 10 Oct 1973 (st), Berg et al. P18417 (MO,NY); Serrado Roncador, 84 kmN of Xavantina,4 Jun1966(d'), Irwinet al. 16593 (NY,U); Sarare,4 Aug 1978 (c?), Pires et al. 15335 (F); 10 km S of Xavantina,4 Sep 1967 (? fl), Richardset al. R.598(NY).MINAsGERAIS:LagoaSanta,21 Oct 186310 Oct 1864 (? fl-fr), Warming1941 (C). ROND6NIA: Rio Guapore,SantaF6, 6 May 1982 (e), M. G. Silva et al. 5913 (NY). RORAIMA: Rio Mucajai,Posto Mucajai, 19 Mar1972 (2 fl), Pranceet al. 11122 (NY,U). BOLIVIA. BENI: Prov. Vaca Diez, rd. RiberaltaCachuelaEsperanza,24 Sep 1993(st),Michelet al. 2264 (BG). LA PAZ: Prov.Iturralde,Luisita,5 Mar 1984 (st), Becket al. 10210(NY);Prov.AbelIturralde, AltoMadidi, 30 May 1990 (??), Gentryet al. 70780A(MO). PANDO: Prov.Manupiri,rd. Porvenir-PuertoHeath,km 26, 16 Aug 1979 (d'?), Meneces 900 (NY); Prov.Abuna,Rio Negro,nr.jct. with Rio Abung,4 Jul 1992(st), Gentryet al. 77794 (MO). AMAmBAY: PARAGUAY. CerroCora,8 Dec 1978(? fl), Bernardi18970 (AAU, NY), 1 Nov 1983 (? fl), Hahn 1734 (BG, MO). CHINCHIPE:

The collection McPherson11824 (Panama)is very unusual, deviating from the characteristicuniflorous pistillateinflorescenceby having 2-3 basally connate flowers, one or all of them producingfruit.

5. Pseudolmedia glabrata (Liebmann) C. C. Berg, Novon 6: 241. 1996 (see Berg, Fl. Neotrop. Monogr.7: 31-33. 1972., subP oxyphyllaria). Synonyms: TrophisglabrataLiebmann,Kongel.DanskeVidensk. Selsk. Skr.Naturvidensk.Math.Afb., Ser. 5, 2: 314. 1851.Type.Mexico.Palenque,nr. Misantla, Mar 1841 (st), Liebmann14274 (lectotype,Berg, Novon 6: 241. 1996, C). Also: BrosimumramonenseStandley,Pseudolmedia malacocarpaStandley& L. 0. Williams,P mollis

FLORANEOTROPICA Standley,P. simiarumStandley & Steyermark, and P oxyphyllariaDonnell Smith. Distribution. FromsouthernMexico (Chiapasand Veracruz) to Costa Rica; in humid forest; to 1500 (-1 800) m. Representative additional specimens examined. MEXICO. CHIAPAS:Mun. Ocosinga, LagunaOcotal Grande,ca. 1100 m, 14 Apr 1967 (? fl-fr), Breedlove 15707 (MICH);Mun. Berriozabal,13 km N of BerriozAbal,15 May 1973 (? fr), Breedlove35299 (MO,NY). OAXACA:Tuxtepec,CerroSebastopol,31 Aug 1967 (e), Penningtonet al. 9080 (NY); RumboNorte,4 km from CerroMann, 25 Feb 1976 (6'), Shapiro395 (MICH). GUATEMALA. ALTA VERAPAZ: Sebol, 14 Apr 1964 (? fr), Contreras4281 (NY). BAJA VERAPAZ: Ninlo Perdido,Rio San Jose, 30 May 1977 (? fl-f), Lundellet al. 20966 (BG, NY). HONDURAS.ATLANTIDA: Above Mataras,8 Apr 1980 (? fr), Hazlett3393 (MO). GRACIOSA Dios: Ca. 3 kmNWof Bulebar,23 Mar1981(R fl-fr),Saunders1100 (NY). COMAYAGUA: MontaniaLa Choca, nr. Quebrada El Zope, 1500m, 14 Apr 1957 (2 fl-fr),MolinaR. 8151 (NY). OLANCHO:Rd. San Esteban-BonitoOriental,7 Feb 1987 (a'), Croat et al. 64399 (BG, MO). NICARAGUA. BOACO:CerroMombachito,4 km NE of Comoapa, 18 Jan 1981 (? ?), Hahn et al. 454 Macizos de PefiasBlancas,20-21 (MO). MATAGALPA: Jan 1982 (6), Stevenset al. 21399 (MO)ZELAYA:Cerro Baka,ca. 6.5 km E of Rio Coperna,19 Mar 1979(Y fl), Pipoly 4934 (MO). COSTA RICA. ALAJUELA: Reserva Forestal San Ram6n,ca. 40 kmfromSanRam6n,21 Mar1986(q fl-fr), Almedaet al. 5684 (NY).GUANACASTE:CerroNubes,ca. 2 km E of Silenciode Talaran,26 Jan 1985 (e'), Grayum et al. 4990 (BG, MO, NY). In Costa Rica the indumentumof the leafy twigs and the laminais more often patentand the leaf venationbeneathis oftenmoreprominentthanin otherparts of the range.

6. Pseudolmedia manabiensis C. C. Berg, Fl. Ecuador 60: 98. 1998. Type. Ecuador. Manabi: Cerro Montecristi, 15 Oct 1995 (o), Cornejo et al. 4660 (holotype, GUAY;isotype, BG). Tree,10 m tall. Leafy twigs 1-3 mm thick, yellowish-tomentoseto hirtellous,the hairsdistinctlydifferent in lengthandpartlycrinkled.Laminacoriaceous,oblong to subobovate,2.5-1 1 x 1-4.5 cm, slightlyinequilateral; apexacuminate;base subacuteto obtuse;marginentire, ? revolute; upper surface puberulous to tomentose, mainlyon the veins; lower surfacetomentoseto hirtellous; lateralveins 12-18 pairs;tertiaryvenationreticulateto subscalariform;petiole 0.2-0.4 cm long, tomentose to hirtellous; stipules 0.8-1.2 cm long,

25 1

TAXONOMICTREATMENT

caducous.Staminateinfloyellowish-(sub)sericeous, rescencessolitaryorinpairsintheleafaxils,subsessile, (immature) 0.5-0.7cmdiam.;involucre withca.7 rows of semiorbicular to ovate,yellowish-subsericeous to appressed-puberulous bracts;stamens(immature) with filamentsca.0.2-0.3mmlong,anthers ca.0.7-0.9mm long,apiculate, glabrous; interstaminal scales(=tepals) 1-1.3mmlong,filiformtonarrowly subspathulate, the upperpartwithdenseandrelativelylonghairs.

18Mar1985(ei), Galeanoet al. 667 (COL).MAGDALENA: Cgto. Manaure,Finca Los Venados, 1400-1700 m, 9 Mar 1959 (c), Romero-Castanieda 7575 (US), (? fl-fr), 7532 (MO,US); Mun. SantaMarta, Romero-Castanleda betweenCerroQuemadoand Cincinatti,1400-2600 m, 19 Apr 1959 (9?), Romero-Castaileda 7801 (MO,NY). VALLE:SantaHelena,above Topacio,1930 m, 10 Dec 1985 (st), Gentryet al. 53023 (BG, MO); rd. CaliBuenaventura,Cgto. Elvira, 1900 m, 23 Jun 1996 (c), Giraldo-Gensini et al. 723 (MO), 25 Aug 1996 (2 fl),

Distribution(see Fig. 82.6). In westernEcuador Giraldo-Gensiniet al. 768 (MO).

VENEZUELA. DISTRITOFEDERAL:Cerro El Avila, 1700m, 24 Jul 1991 (e), Meieret al. 199 (BG). LARA: Becauseof thesimilarities in theindumentum this BetweenCubiroand Escalera,10-15 km SE of Cubiro, speciescanbe regarded to berelatedto Pseudolmedia 1600-2000m, 7 Jul 1974 (c), Steyermarket al. 110192 glabrata,fromwhichitdiffersintheshapeof thelamina (NY, U, VEN). MONAGAS:NW of Caripe,aboveLa Sa(oblongto subobovate), thelargelyreticulatetertiary banade las Piedras,1500m, 5 Apr 1945 (c), Steyermark 61846 (NY). PORTUGUESA:15 km E of Chabasquen, venation,andthe narrow(filiformto subspathulate) 67 kmNNWof Guanare,1450-1520m, 29-31 Oct 1982 interstaminalscales. Pseudolmediamanabiensismay (st), Steyermark et al. 126845 (MO, NY). ZULIA:Sierra proveto be endemicto thecloud(gariua) forestof the de Perija,CaiioHelena,Jul 1975 (? fr), Delascio C. et

(Manabi);in cloud(mist)forest;atca. 700m.

lowmountainsin westernEcuador.

al. 348 (U). ECUADOR. CARCHI: Rio Branco, above Chical

1300-1500 m, 25 Sep 1979 (v?), Gentryet al. 26557

7. Pseudolmedia rigida (Klotzsch & Karsten) (MO).GUAYAS: ReservaEcologicaManglaresChurute, Cuatrecacas (see Berg, Fl. Neotrop.Monogr.7: CerroPanchoDiablo, 5 Jan 1997 (ei), Cornejoet al. 5544 (BG). 33-36. 1972). Theinflorescences tendtobe subsessileratherthan entirelysessileastypicalforthegenus.A short,to ca. 1.5 mm long, bracteatepedunclecan sometimesbe detected.Subsessileinflorescences appearto bemost commonin thenorthernpartof therangeof distribution.Themontaneandsubmontane subspeciesandthe lowlandsubspeciescan stillbe distinguished as distinctentities,althoughthemorphological differences arebecominglessclear. 7a. Pseudolmedia rigida (Klotzsch & Karsten)

Cuatrecacas subsp.rigida. Distribution.IntheAndes,fromBoliviato Venezuela,extendingto theCoastalMountain Range,also in CostaRicaandPanama; in (sub)montane forest;at (700-)1400-2200m.

PERU.HUANUCO: Prov.Pachitea,ca. 20-24 km SE of PuertoInca, 1050 m, 3 Jul 1988 (oi?), Wallnoeffer 11-3788 (BG). PASCO: Prov. Oxapampa, Palmazui,

2150 m, 3 Oct 1984(9 fl), D. N. Smithet al. 8638 (AAU, BG, F, MO, NY). BOLIVIA. COCHABAMBA: Prov.Carrasco, Kharawasi,

2100 m, 2 Sep 1994 (st), AsbunC. s.n. (BG).

7b. Pseudolmedia rigida (Klotzsch & Karsten) Cuatrecacassubsp. araguensis C. C. Berg. Distribution. In northwestern Venezuela; in submontaneforest; at 700-1600 m. Representative additional specimens examined. VENEZUELA. ARAGUA:RanchoGrande,ca. 1000 m, 19 May 1983 (st), Berg s.n. (NY). BARINAS: Mun.

Pedreza,betweenAlto del AguaandLos Piedrones,1250 m, 21 Feb 1955(9 fl), Bernardi2010 (NY).CARABOBO: Rio San Gian, E of Los Tanques,S of Borburata,750Representative additional specimens examined. 1100 m, 31 Mar 1966 (c?), Steyermarket al. 95395 15-18 km NNW of Ospino,700COSTARICA.SANJost: CerrosEscazui-La Carpintera, (NY). PORTUGUESA: et al. 127009(MO). Rio Suarez,betweenCedralandB. Lourdes,1700-1900 800 m, 1 Nov 1982(st), Steyermark m, 4 Mar1994(o), Lepizet al. 192 (NY);Cant6nAserri, ZULIA:Dtto. Mara,nr. Puesto El Bosque, 10-15 Nov Cerrode Escazui,Tarbaca,Rio Su'arez,1800-2000 m, 1982 (? fl), Buntinget al. 12326 (MO). 30 Dec 1993 (ae), Moraleset al. 2174 (MO). PANAMA. CHIRIQUl: 13 km from Rio Sereno, Finca Hartmann,1400-1500 m, 14 May 1991 (? fr), 7c. Pseudolmedia rigida (Klotzsch & Karsten) McPherson15335 (F, MO). Cuatrecacassubsp.eggersii (Standley)C. C. Berg. CUNDINAMARCA:Pacho,Vrda.La EsmeCOLOMBIA. Correctionof the description: ralda,1980 m, 10 May 1964 (e), Garcia-Barriga18049 (NY); rd. Bogota-La Lagunade Pedro Palo, 2100 m, The fruitingperianthcan be shorterthan 1.3 cm.

252

Distribution. In the Pacific Coastal region from Panamato Ecuadorandin theupperAmazonBasinfrom Ecuadorto Bolivia; in non-indudatedforest;to 1000m.

FLORA NEOTROPICA BOLIVIA. SANTACRUZ: Prov.Velasco, 10 km SE of buildingsof EstanciaFlorde Oro,25 Jun 1991 (V?), Nee 41382 (BG, MO, NY).

Specimens? transitionalto subsp.rigida aremarked Representative additional specimens examined. with *. PANAMA. DARIEN: La Laguna ridge, between Rio Tacarcunaand Rio Tapalisa,7 Feb 1975 ($ fr), Gentry et al. 14185 (MO);ca. 17 km E of Pucuro,17 Oct 1987 (q?), Hammelet al. 16277 (MO);Rio El Cupe, ca. 20 8. Pseudolmedia spuria (Swartz)Grisebach(see Berg, km SSW of Boca de Cupe, 29 Apr 1990 ($ fl-fr), Fl. Neotrop.Monogr.7: 36-38. 1972.) McPherson14989 (EAP,MO). Distribution. In Central America, from central COLOMBIA.CHoc6: Mun.Turbo,rd. Rio LeonLomas Aisladas, km 47, 29 Feb 1984 (e'), Brand et al.

Antilles;in everMexicoto Panamaandin theGreater

1005 (MO);trail Unguia-CerroMali, lowest slopes of Serraniade Darien,20 Jan 1975(q fl + fr), Gentryet al. 13731 (MO).

green moist forest; to ca. 900 m.

ECUADOR. EL ORO:Machala, Balneario Los Cocos,

1968 (? fl), Vivareet al. 663 (LOJA).ESMERALDAS: Anchayacu,Eloy Alfaro,Mayronga,10 May 1993 (e'), Between Pennington et al. 13694 (NY,QCNE). GUAYAS: Las Americanasand Daule, 30 Sep 1952, Fagerlindet al. 318 (S). IMBABURA: Nr. Lita, 28 Apr 1949 (? fr), Acosta Solis 12313 (F). Los Rios: Cant6n Vinces, Jaunecheforest, between Mocachi and Palenque, on EsteroPefiafiel,24 Jan 1981 (q fr), Gentryet al. 30739 (AAU, MO, S); Pichilingue,2 Aug 1962 (st), Jdtivaet al. 297 (MO). MANABI:Rio Jama,San Isidro, 11 Aug 1982 (st), Pearsall 1071 (AAU). MORONA-SANTIAGO: 52 km NE of Zamora,Estaci6nExp. El Padmi,11 Sep 1975 (? fl), Littleet al. 297* (COL,LOJA,NY,QAME, US); Pozo PetroleroGarza,ca. 35 km NE of Montalvo, 2-12 Jul 1989 (e), Zaket al. 4425 (BG). NAPO:Ca. 10 km NW of Coca,Feb 1980 (st), Berg et al. 1084 (AAU, HuatiBG, GB, QCA,TUR,U); rd.Coca-Loreto-Hollin, cocha, 11 Jan 1989 (q fl), Palacios3568 (BG, MO,NY, QAME,QCNE);ReservaBiol6gicaJatunSacha,8 km E of Misahualli, 23-31 Jan 1989 (d?), Cer6n 5977 (BG,

MO, QCNE);ParqueNacionalYasuni,Pozo Petrolero Amo 2, 9-13 Jan 1988 (a'), Cer6n et al. 3294 (BG, MO,

QAME).PASTAZA: Pozo PetroleroGarza,ca. 35 km NE of Montalvo, 2-12 Jul 1989 (a?), Zak et al. 4425 (BG,

MO,NY, QCNE).PICHINCHA: Rd. Toachi-LasPalmas, km 5, 1000 m, 21 May 1983 (?), Dodson et al. 13739 Tarapoa, Pozo Petrolero (F, MO, QCNE). SUCUMBIOS:

Representative additional collections examined. MEXICO.CAMPECHE:CampoExperimentalForestal 19 Jan 1966 (st), El Tormento,rd. Escarcega-Cadelaria, Cha/velaP ES.1615 (WIS). CHIAPAS:Chapingo, Campo

ExperimentalForestalEl Tormento,15 Mar 1968 (a'), Pennington et al. 9564 (NY). PETEN:Tikal,rd. to Remate,25 Feb GUATEMALA. 1961 (d), Contreras 1999 (NY). CerroMusun, above NICARAGUA. MATAGALPA:

Salto Grandeof Quebradanegra,20-21 Apr 1977 (c), Neill 1724 (MO). ZELAYA:El Zapote, 40 km NE of Nueva Guinea, 25-31 Mar 1984 (e'), Sandino 4925

(MO);ca. 14.3 km N of El Empalme,rd. to Rosita,27 Apr 1978 (ae), Stevens8380 (BG, MO). COSTA RICA. ALAJUELA:Upala,Dos Rios, 8 Apr 1744 ($ fr), Herrera 1744 (BG, MO). PUNTARENAS: ParqueNacionalCorcovado,18 Aug 1988 (? fl), Kernan 834 (BG, MO, NY). PANAMA.COLON: SantaRita, 18 Mar 1969, Sucre et al. 4832 (U). PANAMA: BarroColoradoIsland,6 Jun 1981 (? fl), Foster 3915 (NY).

JAMAICA.St. Catherine,ca. 2.5 km SE of Sligoville, 16 Apr 1971 (ae), Proctor 31729 (NY). DOMINICAN REPUBLIC. BARAHONA: 3-4 kn N of

Paraiso,11Apr 1985(a), Gentryet al. 50719 (BG,NY). Los Haitises,ca. 8 kmW of Bocadel Infierno, ELSEIBO: 24 Apr 1985 (? fl-fr), Zanoniet al. 34288 (BG, NY). It is noteworthy that the species has not yet been collected in Honduras.

Mariann3, 23 Apr 1990 (st), Ceronet at. 9443 (BG); Dureno,27 Dec 1988(st), Gentryet al. 64235(BG,MO). ZAMORA-CHINCHIPE: Cant6nNangaritza,nr. Shaime, 9. Pseudolmedia gentryi C. C. Berg, Fl. Ecuador60: 31 Jul 1993 (st), Gentry80898 (MO); 27 km NE of 93. 1998. Type.Peru.SanMartin:Rd. Rioja-PomaZamora,rd. to Guaizimi, 17 Aug 1975 (q fl), Little et cocha,km 291, Venceremos,13 Feb 1984 (d), Genal. 426* (LOJA,QAME,QCNE,US). tryet al. 45529 (holotype, MO; isotype, BG). PERU. LORETO:Rio Tascha Curaray, 18 Sep 1972 (? fl-fr), Croat20368 (NY); Rio Mazan,just above La Libertad, 10 Jul 1976 (a'), Gentry et al. 16630 (NY);

Prov. Maynas,Rio Momon, nr. Iquitos, 10 Mar 1973 McDaniel16949 (NY); Prov.Maynas,Indiana,14 Feb 1990 (a), R. Vizquezet al. 13480 (BG). BRAZIL. AMAZONAS:Rio Puru's,between Aiapua

Tree, to 25 m tall. Leafy twigs 3-6 mm thick, drying

purplishbrown to blackish, yellowish to whitish ap-

(a'),

to strigillose,withhairsof i difpressed-puberulous ferentto almostequallength.Laminacoriaceous,ob-

and Moibanda, 18 Jun 1971, Prance et al. 13359

long to elliptic, (7-)10-24(-38) x 4.5-11(-17) cm, (atthebase);apexacumislightyto distinctlyinequilateral nate;base subacuteto roundedto subtruncate;margin

(MICH,NY).

TAXONOMIC TREATMENT

253

uppersurentire(orwhenjuveniledentateto crenate); A sterileBoliviancollectionwithsmallleaves(6onthemidriboralso 11 x 2-4 cm) might also representthe species: facesparsely appressed-puberulous thelateralveins,andinitiallywithdensebrownglobose- COCHABAMBA: Prov.Carrasco,Kharawasi, 2400 m, capitatepluricellular hairs;lowersurfaceyellowishto 4 Sep 1994(st),AsbunC. 35 (BG) to strigillose ontheveins, whitishappressed-puberulous oftenalsowithsomelongerhairs,(initially) withdense, Unnamed Collections pluricellular hairs; brownish, ovoid-toellipsoid-capitate, Twocollectionsof Pseudolmedia fromthePacific venationscalarilateralveins(13-)18-24pairs;tertiary form;petiole0.7-1.2(-2)cmlong,yellowishtowhitish Coastalregionof Colombiacannotbe placedin one to strigillose; stipules1.2-2.5cm of the speciesof sect.Pseudolmedia.TheymatchP appressed-puberulous long,rathersparsely,yellowishto whitishappressed- rigidain the relativelylong stipules,butareclearly tertiaryvenationandthe to strigillosemainlyon themidribandto- differentin the scalariform puberulous dentate margin of the lamina. wardsthe baseoutside,puberulous inside,caducous. Shrub or tree, to 15 m tall. Leafjytwigs 2-3 mm Staminate inflorescences 2-4 in the leaf axils, strigilloseto strigoseto subhirtellous ca.0.4cmdiam.;involucre with thick,? angulate, (sub)sessile, (immature) (broadly) ovate,sparsely, (the longerhairsoften intermixedwith sparseand ca.6-7 rowsof semiorbicularto bracts; (im- minutepatenthairs),witha few conspicuouslenticels yellowishtowhitish,appressed-puberulous mature)stamenswithfilamentsca. 0.2-0.3 mmlong, just below the nodes. Laminasubcoriaceous,lananthersca. 6-7 mm long, apiculate,glabrous;inter- ceolateto oblong,8-24 x 3.5-7.5 cm,broadestabove base(sub)acute;margin tonarrowly staminal scales(= tepals)oblanceolate sub- themiddle;apexacuminate; towards the dent(icul)ate apex; upper surface on theupperpartwithrelatively shorthairs. spathulate, to strigilloseon themidrib;lowersurface puberulous Distribution(see Fig. 82.6). In Ecuador(Napo) strigilloseto strigoseon themainveins,to appressedandPeru(SanMartinandCuzco),possiblyalso Bo- puberulous on the smallerveins;lateralveins 17-22 forest;at(250-) pairs;tertiary livia;in lowermontaneorpremontane venationscalariform; petiole0.4-0.8 cm 1150-1850(-2400)m. long,strigoseto strigillose;stipules0.8-1.5 cmlong, Specimens examined. ECUADOR NAPO:31 kanE of sparselystrigilloseon andalongthe midrib.Stamito 4 togetherin the leafaxils,ca. rd. Tena-Baeza, on rd. to Coca, 1150 m, 23 Dec 1988 (st), nateinflorescences Gentryet al. 64070 (BG, MO);Cant6nEl Chaco,Rio Quijos, 0.5 cm diam.,subsessile;involucrewith6-7 rowsof ca. 0km S ofReventador,lS00 m, 3-5 Oct 1990 (st),Palacios (broadly)ovate, yellowish appressed-puberulous 6441 (BG, QCNE);rd. to Loreto,4 km W of Rio Payamino, bracts;interstaminal scales(= tepals)yellowishapMisionJaponesa,250 m, 3 Aug 1986 (st),Palacios etal. 1131 pressed-puberulous; anthersapiculate.

(BG);Cant6nEl Chaco,El Pacto,9 kmfromrd.Hollin-Loreto, at GuaguaSumaco, 1500 m, 7 Sep 1988 (e), Zaket al. 3787 (BG, MO, NY). PICHINCHA:Maquipucuna, 5 km E of Nanegal, 1550 m, 11 Feb 1991 (st), Gentryetal. 73232 (MO). PERU. Cuzco: Prov. Paucartambo, San Pedro, KosniipataValley, 1100 m, 9 Sep 1991 (st), Nuinezet al. 14227 (BG). SANMARTIN: Rd. Rioja-Pomacocha,km 291, Venceremos, 1850 m, 9 Feb 1984 (st), Gentryet al. 45235 (BG, MO), 10 Feb 1984 (st), Gentryet al. 45331 (BG, MO), 11 Feb 1984 (st), Gentry et al. 45454 (BG), Gentry et al. 45475 (MO).

COLOMBIA.CHOC6:Mun. Mutata,rd. Mutataca. 3 kmbeforeRio Sucio, 100-150m, Pavarandogrande, 8 Dec 1982,Bernalet al. 437 (COL);Mun.SanJose del RioTorito,FincaLosGuaduales, 630-830m, 5 Mar Palmar, 1980(d'), Foreroet al. 6704 (COL,MO).

Perebea 14. Perebea Aublet (see Berg, Fl. Neotrop. Monogr. 7: 38-61. 1972).

ThisspeciesresemblesPseudolmedia macrophylla Additionalcollectionsmadesincethepublication fromthe in its vegetative parts.It canbe distinguished 7 affectedthe taxonomyof the genus, latterby theellipsoidto ovoidheadsof thepluricellar of monograph trichomeson the laminabeneath,usuallyabundantly in particularsect. Noyera. A new species has been present.The stipulesof P gentryiarelongerthanthose

added,PerebeamennegaeC. C. Berg (1978b). Descrip-

arelessdenselyhairyoutsideandare tionsof somespeciescouldbemademorecompleteor of P macrophylla, material the precise (such as for P. longepedunculata and P. inside.In(sub)juvenile sparsely puberulous In manycasestherangesof distribution marginof the laminais ? pronouncedlydentateto tessmannii). relatedto P becamebetterknown.The unnamedcollections listed crenate.Pseudolmedia gentryiis probably rigida,fromwhichit differs,e.g.,in thepredominantlyin Monograph7 are now accommodated:Woytkowski venationandthelargerleaves. scalariform 5534 in P tessmanniiand the othersin P mennegae. tertiary

FLORANEOTROPICA

254

Revisedkey to the speciesof Perebea 1. Pluricellularhairs (minute!)of the lower surfaceof the laminaglobose-capitate;inner involucral bractsdistinctlylonger than the outerones and incurvedbefore anthesis;stigmasfiliform. 2. Leafy twigs and lower surfaceof the laminadensely hairywith the areolesdistinctlyconcave; 8. P. mollis ....................................................... AmazonBasin and Suriname 2. Leafy twigs and lower surfaceof the laminasparselyhairywith the areolesplane or slightly 9. P. rubra concave;AmazonBasin and Guianas....................................................... inner involucral 1. Pluricellularhairs (minute!)of the lower surfaceof the laminaoblongoid-capitate; bractsnot distinctlylongerthan the outerones and not incurvedbefore anthesis;stigmastongueshaped, band-shapedor semicircular. 3. Stipules 0.5-1.5 cm long. 4. Lateralveins of lamina6-14 pairs,the tertiaryvenationreticulateor in leaves longer than 15 cm to scalariform. 5. Laminausually ca. 15-30 cm long, the tertiaryvenationscalariform;upperAmazon 6. P. mennegae Basin ................................................. 5. Laminausually ca. 5-15 cm long, the tertiaryvenationreticulateto (sub)scalariform. 6. Inflorescenceswith 1 flower;perianthof pistillateflower narrowlytubular;inflorescence-bearingshort-shootsto 1 cm long; shrubsto 5 m tall; AmazonianPeruand Ecuador ..............................................

7. P. humilis

6. Inflorescences(usually)with 2 or more (to 10 or 20) flowers;perianthof pistillate short-shoots1-2 mm flower broadlytubularto urceolate;inflorescence-bearing 4. P. angustifolia long; shrubsor trees;CostaRica to Bolivia....................................................... 4. Lateralveins of lamina 13-26 pairs,the tertiaryvenationscalariform. 7. Peduncleof staminateand pistillateinflorescenceat least 1 cm long; marginof lamina dentate; Amazonian Peru.................................................

5. P. longepedunculata

7. Peduncleof staminateand pistillate inflorescenceat most 0.5 cm long, or that of staminateinflorescencesto 1.5 cm long, then marginof lamina(sub)entire;Costa Rica to N Venezuelaand to Acre (Brazil).................................. ........................ 3. P. xanthochyma 3. Stipules 1.5-6 cm long. 8. Involucralbractsextendingto the upperpartof the peduncle;upperAmazonBasin.... 2. P. tessmannii 8. Involucralbractsnot extendingto the upperpart of the peduncle. 9. Stipules2-6 cm long; lenticelsconcentratedin the upperpartof the internodes;north1. P. guianensis ern South Americaand Panama ................................................. 9. Stipulesto 2.5 cm long; lenticelsi evenly scatteredon the internodes;CostaRica to 3. P. xanthochyma N Venezuelaand to Acre (Brazil).............. ...................................

1. PerebeaguianensisAublet(seeBerg,Fl.Neotrop. Monogr.7:42-48. 1972.)

VENEZUELA. AMAZONAS:Depto. Rio Negro, middle part of Rio Bar6a, 29 Jun 1984 (c), Davidse et al. 26860 (MO, NY). Ingeneralthesubspeciesrecognizedin themonoECUADOR. PASTAZA: Cant6n Pastaza, Pozo graphcanstillcanbetoldapart, eveninthenorthwesternPetrolero Villano 2, 23 Jul 1992 (d), Palacios 10261 Cant6n Nangaritza, partof theAmazonBasinwhereallsubspecies,except (BG, QCNE). ZAMORA-CHINCHIPE: for subsp.castilloides,co-occur(sometimeseven in Miazi, 29 Jul 1993 (st), Gentry 80709 (QCNE); Rio Nanthesamelocality).Ina smallnumberof collectionsfea- garitza, Shaime, 1000 m, 26 Oct 1991 ($ fl-fr), Jaramillo turesaremixed,possiblyas a resultof hybridization. 14431 (COL, NY). GUYANA. Upper Takutu-Upper Essequibo region, Aishalton, 14 Jul 1996 (d), Clarke 2192 (BG, US); Upper Takutu-UpperEssequibo region, Rewa R., 17 Feb 1997 la. Perebea guianensis Aublet subsp. guianensis. (? fl-fr), Clarke 3626 (BG, US). BRAZIL. AMAZONAS:Rd. Manaus-Caracarai, km Distribution. Throughoutthe Amazon Basin, in 148, 27 Sep 1973 (?), Berg et al. P.18131 (NY). FrenchGuiana,andwesternSuriname;usually in nonRORAIMA:Rd. Manaus-Caracarai, km 346, Igarape inundatedforest;to 1800 m. Jundia, 26 Nov 1976 (de), Prance et al. 24319 (NY). Representative additional specimens examined. PERU. AMAZONAS: Rio Santiago, Caterpiza, 7 Nov COLOMBIA.CAQUETA: TrailAraracuara-LasPeiias, 1979 (d), Huashikat 1188 (MO). Cuzco: Prov. Cuzco, 11 Aug 1986 (d), Berg et al. 1556 (BG).PUTUMAYO: 4 Dtto. Camisea, Campamento San Martin-C., 11 Jan 1997 km NE of PuertoLeguizamo,Nov 1948 (c), Jaramillo ($ fl),AcevedoR. et al. 8583(BG, US); Prov.Quispicanchi, M. 614 (BG, COL). Camanti, Miniri, 17 Oct 1990 (a^), limand 1042 (MO).

TAXONOMIC TREATMENT LORETO: Prov. Maynas, Dtto. Las Amazonas, nr. Sucusari,

22 Feb 1991 (2 fr), Pipolyet al. 13337(MO). MADREDE DIos: Prov. Manu, 2-3 km W of Atalaya, 12 Dec 1983 (o'), Fosteret al. 7438 (MO).PASCO:Prov.Oxapampa, Palcazu valley, Rio San Jos6, 12 May 1983 (d), D. N. Smith 3976 (BG, MO). PUNO:Rio Tavara, 17 May 1992 (st), Gentryet al. 76738 (MO); Prov. Carabaya,nr. Sangaban, 18 Oct 1984 (e), Maas et al. 6067 (MO). BOLIVIA. COCHABAMBA: Prov. Carrasco, Estaci6n Valle del Sajta UMSS, 29 Oct 1991 (c), Garciaet al. CA.368 (MO, USZ). LA PAZ: Prov. Nor Yungas, 4 km NE of Incahaura, 1500-1530 m, 24 Jan 1984 (st), Gentry et al. 44579 (MO); Prov. Abel Iturralde,Alto Madidi, 22 May 1990 (st), Gentry 70372 (MO).

255

et al. 936 75 km E of Puyo, 4-21 Oct 1990 (d), Gudinio (BG, MO, NY, QCNE). SUCUMBfos:17 km W of Lumbaque,4 Nov 1974 (st), Gentry12423 (MO); ReservaFaunistica Cuyabeno, Laguna Zancudo Cocha, 28 Sep 1991 (c'), Palacios et al. 7783 (QCNE). BRAZIL. AMAZONAS: Mun. Limoeiro, Reserva Ecol. Juami-Japura, 24 Apr 1986 (9 fr), Ferreiraet al. 7190 (NY). PERU. AMAZONAS: Rio Cenepa, QuebradaChichijamenti, 24 May 1973 (2 fr), Ancuash 464 (AAU, MO, NY); Quebrada Saa, 16 Feb 1973 (? fl-fr), Kayap 383 (MO, NY, U). Cuzco: Prov. Quispicanchis,ca. 22 km from Tanquimayo, 6 Mar 1965 (ei), Vargas C. 16208 (US). LORETO:Prov. Requena, nr. Jenaro Herrera,15 Feb 1988 (? fr), Daly et al. 5649 (MO, NY); Prov. Maynas, Indiana, 13 Jul 1990 (9 fr), R. Vasquez et al. 14137 (MO).

The collection Berg et al. P.18131 is hirsuteon the The collections Dik 256, Gentry 12423, and R. mainveins ofthe laminabeneathandon the leafytwigs, moreor less similarto the BoliviancollectionsBuchtien VYsquezet al. 14137 have leaves much narrowerthan 1567 (see Fl. Neotrop.Monogr.7: 46) andGentry70372, normal,the stipulesareoften shorterthan4 cm, andthe and to the Peruviancollection D. N. Smith 3976; the staminateinflorescences have short peduncles. They indumentumrecalls thatof subsp. hirsuta.In northern could representa hybridbetween subsp. acanthogyne AmazonianPeru (Amazonas and Loreto) a form with and subsp. guianensis. smallleaves(mostlyca. 10-25 long) andrelativelyshort stipules(often2-2.5 cm long),representedbythecollections Huashikat1188 andPipoly et al. 1333 7, is com- Ic. Perebea guianensis Aublet subsp. hirsuta C. C. Berg. mon.In the adjacentpartof Ecuador(Morona-Santiago andZamora-Chinchipe)some collections with unclear Distribution. In the northwestempartof the Amaidentityhavebeenmade.Theymatchsubsp.guianensis zon Basin,fromEcuadorto Loreto(Peru)andthe adjain the size and shape of the lamina and length of the centpartof Brazil(Amazonas);in non-inundatedforest; stipules,but in the indumentumthey show similarities at low elevations. to subsp.hirsuta,althoughthe yellow hairsareshorter Representative additional specimens examined. thannornal for the lattersubspecies.These collections Puerto Morona, Rio ECUADOR. MORONA-SANTIAGO: are as follows: ECUADOR. MORONA-SANTIAGO:Morona, 30 Sep 1975 (e), Little et al. 507 (LOJA, NY, Cant6nGualaquiza,Cuangos,20 km E of Gualaquiza, QAME, QCNE, US). NAPO:Ca. 10 km NW of Coca, 29 19 Jul 1993 (? fl-fr),Gentry80210 (QCNE).ZAMORA- Jul 1981 (c), Berget al. 1076 (AAU, BG, GB, MO, NY, CantonNangaritza,Pachicutza, 18 Oct TUR, U); Canton Archidona, rd. Hollin-Loreto, between CHINCHIPE: 1991 (o), Palacios et al. 8303 (MO, QCNE). Avila and Loreto, Huiruno, 24 Nov 1989 (ei), Hurtado 2672 (BG, MO); Reserva Biol6gica Jatun Sacha, 8 km E of Misahualli, 22 Sep 1989 (st), Palacios 4522 (BG, lb. Perebea guianensis Aublet subsp. acanthogyne QAME, QCNE). PASTAZA:Rd. Coca-Auca, nr. Rio Tigiuino, 9 Jan 1989 (?), Palacios 3484 (BG, MO, NY, (Ducke) C. C. Berg. QAME); 20 km S of Curaray, 21-28 Feb 1990 (Y fl), Distribution. In the northwesternpartof the Ama- Zak et al. 5067 (BG, QCNE). SUCUMBIOS:Lumbaqui, zon Basin,fromColombiato southemAmazonianPeru; 1000 m, 13 Aug 1975 (? fr), Little et al. 164 (MO, QAME, QCNE). in non-inundatedforest; at low elevations. PERU. AMAZONAS: Rio Cenepa, 25 Jan 1973 (R flRepresentative additional specimens examined. fr), Kayap 234 (MO, NY, U). LORETO:Prov. Maynas, Rio Caqueta, nr. Araracuara, Rio Ampiyacu, nr. Pebas, 5 Apr 1978 (? fl-fr), PlowCOLOMBIA. AMAZONAS: 10-22 Nov 1982 (V), Idrobo et al. 11325 (NY). man 6666 (AAU, MO, U, US), 6 Apr 1977 (Q fl + oi), ECUADOR. NAPO: Rd. Coca-Auca, ca. 76?54'W, Plowman 6669 (U, US), (e), Plowman 6670 (U, US), 0?44'S, 26 Feb 1980 (ei), Brandbyge et al. 30052 (AAU, (? fl-fr), 6671 (MO, U). SAN MARTfN: Prov. Mariscal BG, MO); ParqueNacional Yasuni, Pozo Petrolero Daimi Caceres, Dtto. Tocache Nuevo, mouth of Rio Mishollo, 2, 26 May-8 Jun 1988 (ei), Cer6net al. 3851 (BG, MO, 25 Jul 1973 (e), SchunkeV 6403 (AAU, MO, NY); Prov. QAME, QCNE); Canton Orellana, Parque Nacional Mariscal Caceres, Cerro de Palo Blanco, 8 Jul 1974 (e), Yasuni, Maxus rd., km 52.5-52.7, 5 Sep 1993 (ei), Dik Schunke V 7231 (MO, NY, U). 256 (MO); Parque Nacional Yasuni, Pozo Petrolero Amo Plowmannoted for his collections 6669-6671, that 2, 9-19 Jan 1988 (ei), Neill et al. 8360 (BG, MO, NY, QAME, QCNE). PASTAZA:Pozo Petrolero Moretecocha, the staminate inflorescences are borne on the upper

256

FLORANEOTROPICA

andthepistillateonesonthelowerbranches. two cited fromLoreto,andthe Ecuadoriancollections branches In Espinoza et al. 615 andPalacios et al. 8303, show feaInmostcollectionstheleavesarerelativelynarrow. a fewcases(suchas inKayap 234, Palacios 3484, and turestransitionalto subsp. acanthogyne. SchunkeV 6403)the leavesresemblethoseof subsp. pseudopeltata. le. Perebea guianensis Aublet subsp. castilloides (Pittier)C. C. Berg. Id. Perebea guianensisAublet subsp.pseudopeltata (Mildbraed)C. C. Berg. Completionof the description: Pistillate inflorescences2.5-3 cm diam.;peduncle

bractsin 8-10 series,broadly 1.3-2cmlong;involucral acuminate to acute;flowtonarrowly ovateto subulate, ca. 10mmlong,4-fid,freeparts ersca.40-80;perianth 3-5 mm long,narrow; style4-5 mmlong,stigmas2.74 x 1.2-1.5 mm.Infructescences4-8(-1 1) cmdiam.; fruitingperianth1.5-2 cmlong,yellow-velutinous.

Distribution. From northwestern Colombia to Panama;in forest;mostly at low latitudes,butrecorded from elevations to ca. 1850 m in Antioquia (Colombia).

Representative additional specimens examined. PANAMA.PANAMA: El Llano-Cartird., km 16, 3 Sep 1977(de),Berget al. 406 (NY,U). SANBLAS: El LlanoCartird.,rd. to Rio Pingandi,31 Jan 1985 (de),Neverset al. 4782 (BG, NY). COLOMBIA. ANTIOQUIA: Mun. Guatape,Vrda. Santa Rita, ca. 1850 m, 6 Dec 1984 (q fl), Albert de Distribution. In the northwesternpartof theAma- Escobaret al. 4950 (HUA);Mun.Amalfi, 18 km E of zonBasin,fromEcuadorto Loreto(Peru);in non-in- Amalfi,rd. to Monosand Portachuelo,1500 m, 23 May undatedforest;to 1250m. 1989 (o), Daly et al. 6060 (BG, MO, NY). CHOC6: Between Quibdo and Tutunendo, rd. to Carmen de Representative additional collections examined. 8 1982 (a'), Albertde Escobaret al. 2194 Atrato, Aug. ECUADOR.MORONA-SANTIAGO: Cord.de Cutucui,rd. La (NY); Mojarra, just upriverfromIstmina,5 Nov 1983 Mendez-Morona,2 Feb 1989 (? fl), Werffet al. 10296 (ei), Juncosa 1245 (BG, MO, NY). (BG, MO, QCNE),4 Feb 1989 (e), Werifet al. 10378 (BG, MO, QCNE).NAPO: Rd. Coca-Auca,km 50-60, 29 Jul 1981 (? fr), Berg et al. 1088 (AAU,NY, S); Rio Payamino,2-10 km S of rd. Loreto, 14-15 Dec 1987 2. Perebea tessmannii Mildbraed (see Berg, Fl. Neotrop. Monogr. 7: 48-50. 1972). (a'), Cer6net al. 3035 (AAU,BG, MO,QAME,QCNE); Huashito, 20 km N of Coca, 3-21 Nov 1989 (? fl), Revised description: Espinoza100 (BG,MO,NY,QCNE);CantonArchidona, Tree,to 20 m tall, dioecious. Leafy twigs 2-7 mm rd. Hollin-Loreto,km 25, ChalluaYacu, 1230 m, 1019 Nov 1988 (a'), Hurtadoet al. 947 (BG, MO,QCNE); thick,yellow-subsericeousto substrigoseto hirtellous Aniangu,ParqueNacionalYasuni,30 May-21 Jun 1982 to hirsute or to subvelutinous. Lamina elliptic to ob(a'), SEF 8716 (QCA,NY). PASTAZA: Cant6nPastaza, long to (sub)ovateor sometimes to subobovate,(10-) 20 km S of Curaray,13-30 Nov 1990 (a), Espinozaet 15-40 x (6-)8-18 cm, slightly inequilateral,subcoriaal. 615 (MO); 2 km NE of Mera, 1100 m, 27 Feb-19 ceous to chartaceous;apex acuminate;base subcordate Mar 1985 (? fl-fr), Neill et al. 6073 (AAU, BG, MO, to deeply cordatewith overlapping lobes; marginenNY, QCA, QCNE);rd. Coca-Auca, 110 km S of Coca, tire (to repand);uppersurfaceglabrescent;lower sur10 km fromRio Tigiiino,7 Jan 1989 (a'), Palacioset al. face sericeousor hirtellousto hirsuteon theveins, other3375 (AAU, BG, NY); confluenceof Rio Quiwadoand wise puberulous beneath, the pluricellular hairs 21 Rio Tiwaeno, Ap 1981 (? fl-fr), Davis et al. 1009 lateralveins 14-22(-24) pairs;teroblongoid-capitate; (BG, NY, QCA). SUCUMBlOS: Reserva Faunistica venation tiary largely scalariform;petiole 0.4-1.8 cm Cuyabeno,nr. LagunaGrande,1 Apr 1988-1 Apr 1990 long; stipules 1.5-2.5 cm long, yellow-sericeous, ca(st), Valenciaet al. 68569 (AAU, QCA). PERU.AMAZONAS: Rio Cenepa,mouthof Kayamas ducous.Staminateinflorescencesup to 7 together,0.8Cr., 4 Dec 1972 (a'), Berlin 448 (NY, U); Quebrada 1.5 cm diam.; peduncle 1.5-3 cm long, bracteateto4 Jul 1976(? fl-fr),Kayap1030 (AAU,MO, wardsthe apex;involucralbractsin 4-5 series,broadly Huampami, NY). LORETO: Rio Napo, 16 Sep 1972 (? fl), Croat ovate to subovate, acuminateto acute, subsericeous; 20256 (NY, MO); Altura Tuta Pishco, Prov. Maynas, perianth1.2-2 mm long, 4-fid to 4-parted,membranaRio Gueppi, 15 May 1978 (? fl), Gentryet al. 21906 ceous, hirtellousto puberulous;stamens(3 or) 4; fila(NY, U). ments 1.7-2.2 mm long; anthersca. 0.4 x 0.2-0.4 mm. In partof the materialplaced underthis subspecies Pistillateinflorescencessolitary,1-2 cm diam.;peduncle the reticulumis plane or almost plane beneath,in an- ca. 1.5-2.5(-3.5) cm long, bracteatetowardsthe apex; otherpartthe reticulumis more or less prominent,of- involucralbractsin 4-5 series,broadlyto narrowlyovate, ten with the areolesi bullate.Some collections, as the appressed-puberulous to subsericeous; flowers ca.

TAXONOMICTREATMENT 20-50; perianth3-5 mm long, lobed, densely yellowpuberulousto subvelutinous;style ca. 2 mm long; stigmastongue-shaped, ca 1.5mm long,acute.Infiuctescences 3-5.5 cm diam.; fruiting perianth ellipsoid to obovoid, ca. 1-1.6 cm long, yellow-hirtellous to subhirsute,orangeto red at maturity. Distribution. FromAmazonianEcuadorto Amazonian Peru (Loreto to Madrede Dios) and to Brazil (Acre); in non-inundatedforest;to 1000 m. Representative additional specimens examined. ECUADOR.MORONA-SANTIAGO:70 km NE of Zamora, Pachicutza,1000 m, 14 Sep 1975 (d), Little et al. 340 (COL,LOJA,NY,QAME,QCNE,US);6 kmSEof Gualaquiza,16 Sep 1975(? fl-fr),Littleet al. 381 (COL,QAME, QCNE,US). NAPO: ReservaBiol6gica JatunSacha, 8 km E of Misahualli,17-24 Feb 1988 (? fr), Cer6n3699 (BG, MO, NY, QAME);ParqueNacionalYasuni,Pozo PetroleroDaimi2, 26 May-8 Jun 1988 (a'), Cer6net al. 4133 (AAU, BG, MO, NY, QAME,QCNE);Huashito, 20 kmN of Coca,3-21 Nov 1989 (a6),Gudifio209 (BG, MO, QCNE).PASTAZA: Rd. Coca-Auca, 115 km S of Coca, 10 km S of borderwith Napo, 26-31 Jan 1989 (e), Neill et al. 8796 (BG, MO, NY, QCA); Canton Pastaza,nr.Villano,3 Dec 1991(a), 7ipazet al. 417 (MO). SUCUMBIOS: Cant6nLago Agrio, Dureno,29-31 Dec 1987 (9 fl), Cer6net al. 3055 (MO,NY, QAME,QCA, QCNE).ZAMORA-CHINCHIPE: Cant6nNangaritza,nr. Shaime,31 Jul 1993 (st), Gentry80937 (QCNE). PERU. LORETO: Prov.Maynas,Rio Amazonas,nr. Indiana, 13 Feb 1989 (d), Gentryet al. 65708 (NY). MADRE DE DIOS:ParqueNacionaldel Manui, Rio Manu, CochaCashuStation,23 Nov 1980 (a), Foster5826 (F, U); Prov.Tambopata,rd. Iberia-Ifiapan,km 17, 1 Mar 1982 (? fl-fr), Ruiz C. 23 (MOL).SAN MARTfN: Prov. ca. km 55, 6 Dec 1997 SanMartin,rd.Tarapoto-Chazuta, (a'), Berg et al. 1785 (BG, COL, MOL);Saposoa, 20 Prov. 5534 (MO,US).UCAYALI: Oct 1959(st), Woytkowski CoronelPortillo,Arboretumvon Humboldt,2 Dec 1978 (9 fr), Diaz et al. 678 (BG). BRAZIL.ACRE:Cruzeirodo Sul, Rio Moa, 21 Aug 1983 (a'), Croatet al. 62463 (INPA,MO). This species appearsto be closely relatedto Perebea longepedunculata.

3. Perebea xanthochyma Karsten (see Berg, Fl. Neotrop. Monogr. 7: 50-52. 1972). Additionaltype of R xanthochyma:Karstens.n., ? (isotype, 0). Distribution. From Costa Rica to western Ecuador,to southernAmazonianPeru,andto Brazil(Acre), andin northernVenezuela(Carabobo);in non-inundated forest;to 1100 m, occasionallyto ca. 1500 (Antioquia, Colombia) or to ca. 1800 m (Carchi,Ecuador).

257 Representative additional specimens examined. COLOMBIA. ANTIOQUIA: Mun. San Luis, Piedra del CastrillOn, 1300-1700 m, 17 Sep 1988 (a), Betancur et al. 699 (BG, MO), BOLIVAR: La Raya, Quebrada La Culebra, 4 May 1987 (st), Gentry et al. 57331 (BG, MO). CAQUETA: Mun. San Vincentedel Caugan,Vrda.El Recreo, Finca Albania, Rio Tubia, 31 Jan 1990 (e), Betancur et al. 1698 (AAU, MO, NY); Araracuara,11 Nov 1988 (a'), P. Palacios et al. 2172 (BG). CAUCA:Rio Micay, Caliche, 26 Feb 1943 (a'), Cuatrecasas 14205 (F). CHOC6: Rd. Quibd6-Guayabal, Rio Duata, 25 Apr 1975 (e), Forero et al. 1202 (COL, MO, NY, U). NARINO: Mun. Tumaco, rd. Tumaco-Pasto, km 63, 19 Feb 1984 (e'), Benavides4265 (MO).VALLE:Bajo Calima, ca. 15 km N of Buenaventura, 14 Feb 1983 ($ fl-fr), Gentry et al. 40251 (BG, MO). VAUPIIS: Estacion Biol6gica Caparui, 3 km N of Lago Taraira, 26 Jul 1988 (2 fl-fr), Defler 318 (MO). VENEZUELA. CARABOBO: Mun. Aut6nomo Mora, Rio Mor6n, 3-5 May 1991 (2 fl), W Diaz et al. 215 (BG). ECUADOR. CARCHI:Reserva Indigena Awa, Chical, 1700-1900 m, 20-29 Jul 1991 (? fr), Quelal et al. 201 Nr. Taisha, 14-15 Oct (QCNE). MORONA-SANTIAGO: 1975 (a), Little et al. 734 (LOJA, US); nr. Bomboiza, 29 Aug 1985 (st) ShakaimRBAE.16a(NY). NAPO:Parque Nacional Yasuni, Pozo Petrolero Amo 2, 9-19 Jan 1988 (a'), Neill et al. 8357 (BG, MO, NY, QAME, QCNE); nr. Reserva Floristica El Chuncho, 6 km from Rio Payamino, 15 Dec 1987 (? fr), Palacios 2285 (BG, MO, NY, QAME, QCNE, US). PASTAZA:Rio Curaray, nr. mouth of Rio Namoyacu, 14-18 Aug 1985 (v?), Palacios 607 (BG, NY, QAME, QCA, QCNE); Pozo Petrolero Villano 2, 2 km from Villano, 3 Dec 1991 (Q fl-fr), lipaz et al. 451 (BG). PICHINCHA: 2 km S of La Independencia, rd. La Concordia-Puerto Quito, km 12, 29 Jul 1984 (a'), Dodson et al. 14612 (AAU, BG, MO, NY, QAME, QCNE). SUCUMBIOS: Lumbaqui, 8 May 1987 (st), Pennington et al. 12224 (BG, QAME, QCA); Reserva Faunistica Cuyabeno, nr. Laguna Grande, 1 Apr 1988-1 Apr 1990 (st), Valencia al. 67341 (AAU, QCA). ZAMORACant6n Nagaritza, Pachicutza, 17 Oct 1991 CHINCHIPE: (a), Jaramillo13939 (GB, QCA, NY). PERU. AMAZONAS: Rio Cenepa, nr. Huampami, 1978 (a'), Ancuash 1289 (BG); Quebrada Wampushik-enta, 13 Jun 1973 (a'), Kayap 926 (NY). Cuzco: La Convencion, Quillabamba, Salespampa, rd. to Kiteni, Rio Urubamba, 1100 m, 28 Oct 1986 (st), Nuniezet al. 6279 (MO). LORETO: Quebrada Tahuayo, above Tamishiyaco, 30 Sep 1972 (a), Croat 19732 (GH, MO, NY); Prov. Loreto: Airico, 11 Dec 1984 (a'), R. Vasquez 6093 (BG, NY). MADREDE DIOS: Tambopata Nature Reserve, jct. of Rio Tambopata and Rio La Torre, 3 Jun 1987 (st), Gentry et al. 58134 (BG); Tambopata,Hermosa Chica, 9 Feb 1989 (st), Alexiades 250 (BG). BRAZIL. ACRE: Reserva INCRA Santa Luzia, rd. BR.364, km 40, 5-19 Oct 1984 (st), Campbell et al. 7203 (NY) and 7554 (BG); Cruzeiro do Sul, Rio Jurua, 27 Oct 1966 (? fl), Prance et al. 2901 (F, INPA, NY, U). AMAZONAS:Mun. Limoeiro, Reserva Ecol. Juami-Japura,

258 24 Apr 1986 (d), Ferreiraet al. 7184 (BG, MG, MO, NY); Rio Solim6es, Sao Paulo de Oliven9a,Barriode Odorio,17Aug 1973(a), Lleraset al. P1 7371(MO,NY). BOLIVIA. LA PAZ: Prov. Abel Iturralde,Alto Madidi,24 May 1990 (st), Gentryet al. 70519 (MO). The peduncle, in the monograph described as to 0.3(-0.6) cm long, is sometimes to ca. 1.5 cm long in staminateinflorescences, as in westernColombiaand Ecuador,in AmazonianPeru(Loreto,Madrede Dios) and in Brazil (Acre), and for pistillate inflorescences even to 2.5 cm long in materialcollectedin Amazonian Peru (Loreto) and Brazil (Acre and Amazonas). The axillaryspursusuallyremainshortbut may sometimes become to 1.5 cm long, as in some large-leavedcollections fromColombia(Valle),e.g., Cuatrecasas16629.

4. Perebea angustifolia (Poeppig & Endlicher)C. C. Berg(seeBerg,Fl.Neotrop.Monogr.7: 52-54. 1972). Distribution (see Fig. 82.5). From Costa Rica to westernEcuador,to northwestern Venezuela(Zulia)and to Amazonian Brazil (Acre) and Bolivia (Pando, La Paz); in non-inundatedforest;at low elevations. Representative additional specimens examined. COSTARICA.HEREDIA: Rio Sarapiqui,FincaLa Selva, 27 Jul 1980 (a), Hammel9338 (NY). LIM6N:Refugio Gandoca-Manzanillo, Gandoca, 27 Jan 1987 (e), Corcovado Grayumet al. 8015 (BG,NY). PUNTARENAS: NationalPark,trailLlorana-RioCorcovado,21 Jul 1977 (e'), Hartshorn1884 (BG). PANAMA.COL6N:SantaRitaRidge,ca. 10kmfrom TransisthmianHwy., Porter et al. 4790 (AAU, NY). DARIEN:CerroPirre,800-1500 m, 9-10 Aug 1967 (? fr), Duke 13766 (AAU, NY). PANAMA:Ca. 20 km NE of Chepo, 12 Dec 1973 (R fl-fr), Berg et al. 334 (BG); El Llano-Cartird., km 9, 22 Mar 1975 (a'), Moriet al. 5164 (AAU, NY). SAN BLAS: El Llano-Carti rd., km

FLORANEOTROPICA de Cutucu,trailLogrofio-Yuapi, Nov 1976 (? fr), Madison et al. 3334 (US). NAPO: ParqueNacional Yasuni, Pozo PetroleroDaimi2, 26 May-8 Jun 1988 (e), Cer6n et al. 3863 (MO);Afiangu,ParqueNacionalYasuni,115 Feb 1986(e), Korninget al. 47752 (AAU,BG, QCA, QCNE);ReservaBiologicaJatunSacha,8 kmfromPuerto Mishualli,1-15 Sep 1987 (a), Palacios 2004 (BG, MO, NY, QCA,QCNE).PASTAZA: Pozo PetroleroMasaramu, ca. 40 km NNE of Montalvo,1-16 May 1990 (? fl-fr), Gudifio318 (BG, MO,NY, QCNE).PICHINCHA: 12 km E of PatriciaPilar,31 Jul 1984(st), Dodsonet al. 14759 (BG). SUCUMBIOS: Cant6nGonzaloPizarro,nr. Campo Bermejo6 Norte, 30 km NW of Lago Agrio, Cer6net al. 9399 (QCNE);Rio Aguarico,5-7 km upstreamfrom SantaCecilia, 2 Feb 1969 (a), Duke 16074 (BG, MO). PERU. HUANUCO:Prov. Pachitea, Dtto. Honoria, BosqueNacionalde Iparia,Rio Pachitea,20 km above confluence with Rio Ucayali, 21 Feb 1967 (? fr), SchunkeV 1653 (F), 24 Apr 1967 (? fr), SchunkeV 1879 (F). LORETO: Rd. Iquitos-Nauta,km 44, 11 Dec 1988 (a), R. Vdsquezet al. 11468 (BG, MO);CaballoCocha,Aug 1929 (? fl), LI. Williams2048 (F). MADRE DE DIos: 39 km SW of PuertoMaldonado, nr.confluence of Rio La Torreand Rio Tambopata,25 May 1987 (? fl), Gentryet al. 57651 (AAU,BG,MO,NY); Rio Mana, CochaCashuStation,22 Oct 1980 (? fl-fr),Foster5626 (F). PASCO:Prov.Oxapampa,Dtto. Huancabamba, Siete de Junio,1 Aug 1982($ fl-fr),ReynelR. 589 (BG).PUNO: Rio Tavara,16 May 1992 (? fl), Gentryet al. 76663 (MO).SAN MARTIN: Rio Huallaga,Chazuta,Apr 1935 (a'), Klug 4088 (F, K, MO, NY, US). UCAYALI: Prov. Coronel Portillo, Bosque Nacional Alexander von Humboldt,rd. Pucallpa-TingoMaria, km 86, 22 Feb 1980 (a'), Begazo 95 (BG, MO). BRAZIL.ACRE: UpperRio Moa, FazendaArizona, 10-16 Oct 1985 (? fl-fr), Campbellet al. 6481 (BG). BOLIVIA. PANDO: 74 km SW of Cobija, 12 Aug 1984(? fr),R. T Penningtonet al. 109 (BG,F). LA PAZ: Prov.AbelIturralde, Alto Madidi,29 May 1990(a), Beck et al. 18237 (BG).

26.5, 9 Apr 1985 (? fl-fr), Neverset al. 5265 (BG).

Individualsof this species startto flower as shrubs of 1-3 m height, but may become trees to 20 m tall NacionalNaturalAmacayacu,22 Feb 1991(? fr),Rudas throughoutthe rangeof distribution,or occasionallyto et al. 1378(BG,MO).CHOC6: CaboCorrientes, Quebrada 35-40 m tall.Thetertiaryvenationvariesfrom? clearly Blanca,28 May 1974 (a'), Warner305 (F,MO,NY,US). in VALLE:Rio Calima,Quebradade la Brea, 18-22 May scalariform relativelylargeandbroadlaminasto (almost)reticulatein relativelysmallandnarrowlaminas. 1946 (a), Cuatrecasas21188 (F). VENEZUELA. ZULIA:Rio Guasare,Serraniade The inflorescence-bearingshort-shootsremainshort, 1-2 mm long. Twoformscanbe distinguished:themost Perija,9 May 1983 (st), Gentry41159 (BG, MO). ECUADOR. ESMERALDAS:Cant6n San Lorenzo, widespreadand common form with rathersparse apMataje,21 Sep 1992 (e?), Aulestia et al. 518 (AAU, pressedindumentumon the leafy twigs, lower lamina QCNE);Fila de Bilsa, 7 km E of SanJose de Bilsa, 28 surface,andfruitingperianth,anda less commonform Jan 1991 (e), Gentryet al. 72814 (F, MO,QCNE).Los (represented Cer6n by et al. 3863, Gudino 318, and Rios: Cant6nQuevedo,CerroCentinela,Montainasde Palacios 2004) which is densely (sub)tomentose to Ila, 19 Jan 1991(a?), Palacioset al. 7433 (MO,QCNE). on the mentioned. pubescent parts Los RIoS/PICHINCHA:El Centinela, crest of Montafias de Ila, rd. PatriciaPilar-24de Mayo,km 12, 6 Feb 1979 COLOMBIA. AMAZONAS:Mun. Leticia, Parque

(a'), Dodson 7335 (MO, QCNE, US). MORONASANTIAGO: PuertoMorona,E of Rio Morona,1 Oct 1975

(st), Littleet al. 521 (LOJA,QAME,QCNE,US); Cord.

5. Perebea longepedunculata C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 54. 1972).

TAXONOMICTREATMENT Completionof the description: Pistillate inflorescences solitary,0.1-1 cm diam.; peduncle 1.2-2.2 cm long, with 1 or 2 bractsin the upperpart;involucralbractsin 4-5 series,broadlyto narrowly ovate, appressed-puberulous;flowers ca. 7-15; perianth ca. 3 mm long, dentate, densely yellowpuberulous;styleca. 1mm long;stigmastongue-shaped, 1-1.5 mm long,acute.Infructescences1.5-2.5 cm diam.; fruitingperianthellipsoid to subglobose,ca. 1-1.2 cm long, rathersparselyyellowish-puberulous,orangeto red at maturity;fruitellipsoid to subglobose, ca. 0.60.8 x 0.4-0.7 cm.

259

stipules0.5-1.5 cmlong,yellowish-subsericeous, caducous. Staminateinflorescences numerous, succes-

sivelyformedon unbranched orbranched,to 0.8 cm longshort-shoots, withstrigillosescales,thereceptacle 0.1-0.25 cm diam.;peduncle0.2-0.4 cm long, appressed-puberulous; involucrewith 8-15 ovate,appressed-puberulous to subsericeous bractsin 4-5 rows; flowers2-6; perianth ca. 1mmlong,3-4-fidto-parted, thetepals? cucullate,glabrousorpubenulous; stamens 3-4, if 4, thenoftenoneof them? reduced,free,about aslongastheperianth; filamentsbroadorratherslender.Pistillate inflorescences solitary in the leaf axils;

inpeduncleca. 0.3 cm long,appressed-puberulous; volucrewithca. 10ovate,appressed-puberulous bracts in3 rows;flowers2-6; perianth tubular, ca.4 mmlong, 4-lobed,puberulous; stigmaslingulate,ca.0.1cmlong. Representative additional specimens examined. Fruiting perianth ca. 1 cm long, red, densely PERU. LORETO:Prov. Requena, nr. JenaroHerrera, puberulous; seedca. 0.7 x 0.5 cm. Distribution. In Amazonian Peru (Loreto, San Martin,and Ucayali); mostly in floodplain forest or restingas,but also on terrafirme;at low elevations.

SupayCocha,25 Nov 1988(? fr),Daly et al. 5769 (BG, MO,NY); Prov.Maynas,betweenIquitosandRio Itaya, MoenaCanlo,7 Jan 1976 (e fl-fr), Gentryet al. 15676 (AAU, BG, F, MO, NY); Prov. Requena, nr. Jenaro Herrera,Rio Iricahua,12 Dec 1985 (a'),Peters141 (BG, MO,NY); Prov.Maynas,Rio Itaya,nr. SanAntonio,18 Feb 1977 (? fr), Revilla 2371 (BG, F, MO, NY). SAN MARTiN: Prov.MariscalCareres,Dtto. Uchiza, 24 Jul 1974 (a'), SchunkeV 7734 (BG, MO); Prov.Mariscal Caceres,Dtto. TocacheNuevo, between PuertoPizana andQuebradade Salas,ca. 50 km N of TocacheNuevo, 23 Dec 1981 (o?), Plowmanet al. 11674 (BG, F, GB, GH, MO). UCAYALI: Canchahuayo,29 Nov 1985 (e), R. Vasquezet al. 7023 (BG, MO).

Distribution(see Fig. 82.1). In easternLoreto (Peru), the adjacentpart of Brazil (Amazonas), and Colombia (Caquetd);in non-inundcated forest; at low elevations. Representativespecimensexamined.COLOMBIA. AMAZONAS: Rio Caqueta, nr. Araracuara, 23 Aug 1990 (e),

Vester17 (BG). CAQUETA: Araracuara, 21 Jan 1984

(2 fr), Gentryet al. 64902 (F, MO). BRAZIL. AMAZONAS:Sao Antoniode l9a, 27 Sep 1906 (c), Ducke MG 7698 (BM); nr. Sao Paulo de

Oliven9a,IgarapeCamatia,(st), Fr6es23999 (IAN);Sao Paulode Olivenca,(v), Fr6es 24074 (IAN); Mun. Sao Paulo de Olivenca, 26 Oct-l Dec (o), Krukoff 8768 (BM, The laminais sometimes subentire,such as in colF, MICH,U, US); Rio Cunhua,Deni Indian village, lectionPlowmanetal. 11674.A sterilecollection,Nuiiez 6?43S, 66?47'W, 28 Nov 1971 (c), Pranceet al. 16458 et al. 12165 (BG) from Madre de Dios (Prov. (INPA,NY);Rio Javari,Estiraodo Ecuador,22 Oct 1976 Tambopata),might belong to this species. (st), Prance et al. 23993 (NY, U, US). PERU. LORETO:Prov.Requena,JenaroHerrera, AugSep 1976(e), Bernardi16240(U);nr.mouthof RioNapo, 6. PerebeamennegaeC. C. Berg,ActaBot. Neerl. 14 Sep 1972 (oi), Croat20169 (AAU, F, MO, NY, U); 27:12,t.2. 1978.Type.Brazil.Amazonas: RioJavari, Maynas,SWof Iquitos,rd.to Quistococha,18 Nov 1975 72049'W,5008'S,1 Aug 1973 (ei), Lleras et al. (v), Davidson3282 (F,LAM);Prov.Maynas,Rio Nanay, P.16988 (holotype,INPA;isotypes,F, K, NY,U, Mishana,30 May 1978 (? fr), Gentryet al. 22346 (F), 20 Mar1982(o), Gentryet al. 36510 (F,MO,U), 29 Jul US). 1982(st), Gentryet al. 38020 (BG, MO);Prov.Maynas, Shrub,to 5 m tall,probablydioecious.Leafytwigs QuebradaSucursari,Rio Napo,below Mazan,7 Jul 1983 1.5-5 mmthick,withminutepatenthairs,intermixed (o'), Gentryet al. 42716(BG, MO); Prov. Requena,Jenaro distinctlylonger,mostly appressed,yellowish to gray- Herrera,25 Feb 1987 (st), Gentryet al. 56647 (MO); Prov. Maynas, Rio Yahuasyacu, 15 Jul 1990 (ei), ish, stiff hairs.Laminaoblong,(8-)15-30 x (4-)7-14.5 subcoriaceous tocoriaceous; Grandezet al. 1718 (BG, MO); Prov.Maynas,Puerto cm,slightlyinequilateral, baseacuteto obtuse; Almendras, 11 Nov 1984 (oi), Maas et al. 6260 (MO, apexacuminateto subcaudate; U); Prov. Maynas, Rio Nanay, Mishana, 18 Aug 1978 (e),

marginsubentireto ? distinctlydentate; uppersurface McDanielet al. 22088 (F); Prov.Maynas,Rio Nanay, to strigilloseon themid- Mishana, Santa Maria de Nanay, 30 Sep 1990 (c), Pipoly sparselyappressed-puberulous

to et al. 12607 (BG, F, MO); Prov. Maynas, Dtto. Las rib;lowersurfacesparselyappressed-puberulous strigilloseon the midribandlateralveins; lateralveins 6-15 pairs;tertairyvenationlargely scalariform;peti-

tostrigillose; ole0.4-1.2cmlong,appressed-puberulous

Amazonas, nr. Sucusari, 5 Mar 1991 (c), Pipoly et al.

14520 (BG, MO);Prov.Maynas,SantaMariade Nanay, 14 Mar 1991 (e), Pipolyet al. 14947 (BG); Prov. Maynas,

260

FLORANEOTROPICA

Pebas,Rio Ampiyacu,19 Jul 1976 (a), Revilla939 (BG, MO); Prov.Maynas,Rio Nanay, Mishana,5 Jun 1979 (c?), Rimachi Y 4436 (NY); Prov. Maynas, Puerto Almendras,9 Nov 1983(d'), R. Vdsquez et al. 4588 (BG); Prov.Maynas,rd. Iquitos-Nauta,km 5, 28 May 1987 (a'), R. Vcsquez et al. 9003 (BG); Pebas, 13 Oct 9976 (a'), R. Vasquezet al. 9776 (BG);Prov.Maynas,Puerto Almendras, 16 May 1990 (? fl-fr), R. Vdsquezet al. 10537 and 13731 (BG);Prov.Maynas,Rio Nanay,Santa Mariade Nanay,8 Aug 1990(a), R. Vasquez et al. 14197 (BG).SANMARTIN: Dtto.TocacheNuevo,Rio Huallaga, PuertoPizana, 20 Dec 1970 (a), SchunkeV 4593 (F, NY, U). UCAYALI:Prov.CoronelPortillo,Rio Abujao, 10 Dec 1978 (? fl-fr), C. Diaz et al. 751 (MO).

Prov.Abuna,Nuevo Mundo, 2 Jul 1992 (st), Gentryet al. 77607 (MO).

PANDO:

Specimensof Perebeahumilisarenotalwayssubshrubs,butcanbetreeletsto 4 m tall,andspecimensof P angustifoliacanstartfloweringas shrubsortreelets of 3 m tall.Thetwospeciescannotbe toldapartbythe habit.Besidessome(oftennoteasilydetectable) features of flowersandinflorescences, thetaxacanbe mostly disinguishedby thecommonpresenceof patenthairs onthelaminabeneathandtheleafytwigsinP humilis, whereasthehairsof theseplantpartsaremostlyappressedinP angustifolia. Inthesparselyhairyformof P angustifoliathehairson thestipulesareusuallynot Perebea mennegae,togetherwith R humilisandR (almost) confinedtothemidribof thestipuleastheyare longepedunculata, belongs to a group of species of inP humilis. Theinflorescence-bearing short-shoots can shrubs,subshrubsor treeletsto 6 m tall, occurringin a becometo 1cm longandareslenderinP humilis,but small area in northernAmazonian Peru and adjacent areusuallynotmorethan1 mmlonginP angustifolia.

parts of Brazil (Amazonas) and Ecuador (Napo and Pastaza)and in Colombia(CaquetA).

7. Perebea humilis C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 54-56. 1972). Correctionof the description: Subshrub,less than0.5 m tall to treelet,to 4 m tall.

The collections Ceron et al. 9399 and Schunke V

7208matchP agustifolia intheirindumentum, butthey havethecharacteristic slenderelongateshort-shoots and the mostlyuniflorousstaminateinflorescencesof P. humilis.Thisis anotherindicationhowcloselyrelated thetwotaxaareandthatadditional collections mayprove thatonlyrecognition atthesubspecific levelisjustified.

Distribution. In the upperAmazon Basin, in Peru 14B.PerebeaAublet sect.Noyera(Trecul)Engler,in (Loretoto Madrede Dios) andEcuador;in undergrowth Engler&Prantl,Nat.Pflanzenfam. 3(1):84. 1889. of non-inundatedforest; to 1050 m.

Increased collectionshavecreatedproblemsin dis-

Representative additional specimens examined. tinguishing thethreemorphological entitiesrecognized ECUADOR.NAPO:Nr.Nuevo Rocafuerte,26 Feb 1981 in the monograph: Perebea mollis subsp. mollis, P (? fl), Jaramilloet al. 4255 (MO,QCA).PASTAZA:Rio Curaray,mouthof Rio Namoyacu,14-18 Aug 1985 (fl), mollis subsp.rubra,andP glabrifolia. Sterilematerial Neill et al. 6613 (BG, MO, QAME, QCA, QCNE). of P glabrifoliacan only be told apartfromsterile of P mollissubsp.rubraby thelowersurface Cant6nGonzaloPizarro,CampoBermejo, material SUCUMBiOS: 30 km NW of Lago Agrio, 1050 m, 30 Mar 1990 (a'), of thelamina,whichis planeinP glabrifolia,whereas Cer6net al. 9399 (BG, QCNE);SantaCecilia, 30 Mar theareolesare? concavein thematerialreferred to P 1972 (9 fl), Dwyeret al. 9757 (BG, QCA,MO). mollis subsp.rubra,morepronouncedlyso in P mollis PERU.LORETO:Prov.Maynas,betweenIndianaand subsp. mollis. mouthof Rio Napo, 18 Feb 1981(fl), Gentryet al. 31377 InPerebeaglabrifoliatheperianths of thepistillate (BG, MO, NY, U); Rio Itaya,Soledad,20-22 Sep 1929 (fl), Killipet al. 29574 (F), (fl), Killipet al. 29576 (US). flowersarefree andhirsute,as they arein P mollis material referred toP mollis MADREDE DIOs:Rio Manui,Cocha CashuStation, 14 subsp.mollis.Thepistillate periOct 1980 (a'), Foster 5539 (MO, F); nr. confluenceof subsp.rubrahas connateminutelypuberulous Rio Tambopataand Rio la Torre,39 km SW of Puerto anths.However,materialwiththis featurecanbe diMaldonado,24 Jan 1989 (fl), S. F Smithet al 1556 (F, videdintotwogroups:onewithsparseindumentum, US). PASCO:Prov. Oxapampa, 9 km SW of Puerto largelysimilarto thatin P glabrifolia,andthe other Bermudez,Rio Nochos, 7 Sep 1982 (a'), Foster 8600 withdenseindumentum. Theformergroupconsistsof (BG,F,MO,NY).SANMARTIN:Rd.Tarapoto-Yurimaguas, materialfromthe Guianas,northernPara,and the km 66, 27 Apr 1986 (st), Knappet al. 7241 (MO);Prov. Manausregion;the latterof materialfromtheupper MariscalCaceres,Dtto. TocacheNuevo, Cerrode Palo AmazonBasin,Colombia, Ecuador, andPeru.ThemaBlanco, 8 Jul 1974 (a), Schunke V 7208 (BG, MO). UCAYALI: Prov.CoronelPortillo,Dtto. Iparia,nr.Iparia, terialfromtheupperAmazonBasindiffersfromthe materialreferredto P mollissubsp.mollisin theab26 Aug 1968 (fl), Schunke V 2692 (NY, U, US). BOLIVIA.

LA PAZ: Prov. Abel Iturralde, Alto

Madidi, 29 may 1990 (a), Beck et al. 18237 (MO).

senceof a clearlybullateuppersurfaceof thelamina,a tendencyof havingtheindumentum onthemidribbe-

TAXONOMIC TREATMENT

261

hirtellousto hirsuteor neath,ontheleafytwigs,andonthestipules? appressed, 2.5 cm diam.;fruitingperianth fruitalmostentirelyadnateto theperianth. anda tendencytohaverelativelylongstipules,to 3 cm puberulous; it is notclear material Twosubspeciescanbe recognized. long.Dueto thelackof adequate whetherfeaturesof the staminateflowerscouldconentities.The themorphological tribute to distinguishing material 8a. Perebea mollis (Poeppig & Endlicher) Huber tepalsarefreeoralmostfreein thestaminate subsp.mollis. of the(rather)glabrousentitiesanddistinctlyconnate in thehairymaterial;thisdifferencemayproveto be Reviseddescription: solutioncouldbeto consistent.Themostfar-reaching Longerhairs on thestipules,ontheleafytwigs,and treatfourentitiesas subspeciesof a single species. on the midribof the laminabeneathpatent.Lamina solutionseemsto beto However,themostsatisfactory usuallyto 20 cmlong;uppersurfaceof thelaminadisunitethemtwobytwounderdifferentspecies,distinct tinctlybullate,hirsuteto hirtellousto strigose,usually in the characters by someratherweakdifferentiating ? scabrous. Perianthofthepistillateflower free,hirtelandinthedenseparts,suchasinthevenation vegetative lousto hirsuteon thelowerpart. nessof indumentum. theAmaDistribution(seeFig.83.2).Throughout zonBasin,butmainlyin thesouthempart,extending Huber(see tothesouthernmost 8. Perebeamollis(Poeppig& Endlicher) partof Surname;innon-inundated Berg,Fl. Neotrop.Monogr.7: 558-60. 1972). forest;atlow elevations. Representative additional specimens examined. Reviseddescription: Tree,to 30mtall,mostlydioecious. Leafytwigs 1.5- SURINAME. Sipaliwini savanne, 23 Jan 1970 (st), 4.5(-7) mmthick,withveryshortpatenthairsinter- Oldenburgeret al. 1137 (U). PERU. MADREDE DIOS: Prov. Tahuamanu, Dtto. topatent, longer,appressed mixedwithdense,distinctly brownishto yellowto whitishhairs.Laminaellipticto Iberia,rd. Iberia-SanLorenzo,km. 5.5, 10 Dec 1981 (MO), 18-25 Dec 1985 (? fr), oblong to (sub)ovate,6-36 x 2.5-14 cm, slightly (e), C. Diaz 1765-109 ReynelR. 1101 (MOL). apexacuteto to coriaceous; chartaceous inequilateral, Rio Negro, Barcelos, 28 Apr BRAZIL.AMAZONAS: baseobtuseto subcordate; 1952 (e), Fr6es 28340 (U). MARANHAO: acuminateto mucronate; Rd. Tocantinsuppersurface Bel6m, km 345, 13 Nov 1963 (o), N. T Silva 57793 marginentireto repandto dent(icul)ate; strig(ill)oseto hirsute,exceptforthemidribglabres- (NY,U). MATOGROSSO: Rio Aripuana,below Saltodos cent, ? bullateand ? scabrousto almostplaneand Dardanelos,nr.HumboldtCenter,14 Oct 1973 (6), Berg smoothabove;lowersurfacedenselyyellowish-hirsute et al. P.18525 (F, MO, NY, U). PARA: Rio Xingu, area withpatentorpartly? ap- of AssuriniIndians,10 Jun 1986 (st), Balee 2312 (NY); to hirtellousto puberulous pressedhairs;lateralveins(12-)15-22pairs,loop-con- Rio Parudo Oeste,regiaode Tiri6s,4 Apr 1962, (? fr), nected1-1.5 mmfromthe margin;tertiaryvenation Fittkauet al. INPA12912 (U). BOLIVIA.BENI:Prov.VacaDiez, nr.Alto Iv6n,Aug (0%S) petiole0.1-1cmlong;stipules largelyscalariform; 1992 (q fl),Bergeron513 (BG), 20 Feb 1984 (st), Boom 1-2.5(-3) cm long, yellow-hirsuteto subsericeous, 4505 (NY). PANDO:Rio Madeira,oppositeAbuna, 19 sometimessubpersistent.Staminateinflorescencesup Nov 1968 (st), Prance et al. 8676 (F, NY, U). SANTA to 8 togetherin the leaf axils,0.8-1.6 cm diam.;peCRUZ:Prov.Velasco,ParqueNacionalNoel KempffM., in6-9rows,broadly bracts duncle1-3cmlong;involucral Lago Caiman,14 Jan 1997 (? fl-fr), Garvizuet al. 288 to narrowlyovateto lanceolateto linear,acuteto cau- (BG), 21 Nov 1996 (d'), Wallace 82 (BG). to hirtellous,theinnerones date,appressed-pubescent to 0.8 cm long andincurvedbeforeanthesis;flowers numerous; perianth1.7-2.5 mmlong,2-3-lobedto - 8b. Perebea mollis (Poeppig & Endlicher) Huber attheupperend;stamens2-3; filaments fid,puberulous subsp. lecithogalacta (R. E. Schultes) C. C. Berg, 2-3 mmlong;anthers0.8-1.3 x 0.2-0.3mm.Pistillate Fl. Ecuador60: 86. 1998. Castilla ulei Warburgf. by solitaryorsometimesaccompanied inflorescences lecithogalactaR. E. Schultes, Bot. Mus. Leafl. 12: 1-2 staminateones, 0.8-1.2 cm diam.;subsessileor 128. 1946. Perebealecithogalacta (R. E. Schultes) bractsin 4-7 rows, peduncleto 0.5 cmlong;involucral R. E. Schultes, Bot. Mus. Leafl. 13: 298. 1949. broadlyto narrowlyovateto lanceolate,acuteto cauLonger hairs on the stipules, the leafy twigs andat to hirtellous,theinnerones date,appressed-pubescent incurvedbeforeanthesis;flowers6-30, freeorbasally leastonthemidribof thelaminabeneath? appressed, hirtel- sometimesmost(longer)hairspatent.Laminausually 2-3 mmlong,entireto4-dentate, perianth connate; lous,setosenearthebaseornot;styleca. 1.5mmlong, longerthan20 cm;uppersurfaceof thelaminaslightly smooth.Perianthofthepis2-3(-5) mm long.Infiuctescences1.5- bullatetoplane,strigillose, stigmas glabrous;

262 tillateflowerin the lowerpartconnate,puberulous(also in fruit). Distribution (see Fig. 83.2). In the upperAmazon Basin, from Colombia to CentralPeru; in non-inundatedforest;at low elevations. Specimens examined. COLOMBIA. AMAZONAS: Mun.Leticia,ParquenacionalNaturalAmacayacu,Agua Pudre, 11 Nov 1991 (a'), Pipoly et al. 15858 (U). Rio Caqueta, Isla Morrocoy,20 Jun 1988 (st), M. Sanchez S. et al. 903 (BG). VAUPtS:Rio Vaupes, PuertoNare, 18 Feb 1944 (? fl-fr), Schultes 5798 (ECON,K, US, type collection). CAQUETA:Rio Caguan,La Argentina, 3 Mar 1977 (st), Roa 635 (UDBC). META:Mun.Vista Hermosa,caserio Piiialito, Nov 1973 (st), Mosquera UDBC 10848 (UDBC). ECUADOR. MORONA-SANTIAGO: 16.7 km NE of Taisha,14-19 Aug 1976 (st), OrtegaU. 50 (US). NAPO: Yucard., 3 km fromAucaoil fieldrd., 22 Mar1980 (a), Brandbyge et al. 30302 (AAU, BG, MO, NY, QCA);

ParqueNacional Yasuni,Helipuertode Amo Sur, 1619 Jan 1988 (? fl + d), Ceron3456 (BG, GB, MO, NY, QAME,QCNE,US); ParqueNacionalYasuni,Lagunas de Garzacocha,22 Sep 1988 (? fl-fr),Ceronet al. 5099 (AAU, MO, QCNE);Aniangu,ParqueNacionalYasuni, 1-15 Feb 1986 (st), Korninget al. 47609 (AAU,QCA), 30 May-21 Jun 1982 (a'), SEF 8656 (AAU, MO, NY,

QCA); Reserva Biol6gica Jatun Sacha, 8 km E of Misahualli,8 Aug 1987(st),Palacios1762 (BG,QAME); ReservaFloristicaEl Chuncho,5 km N of Coca, 9 Oct 1987(st),Palacios2102 (BG,QAME,QCNE).PASTAZA: Rd. Coca-Auca,ca. 115 km S of Coca,nr.Rio Tigiiino, 29 Apr 1989(st),Rubio57 (BG,QCNE);Cant6nPastaza, Pozo PetroleroDanta2, 50 km SSE of Curaray,1-20 Oct 1990 (a'), Espinoza et al. 447 (BG,MO, NY, QCNE); Pozo PetroleroMoretecocha,75 km E of Puyo, 4-21 Oct 1990 (a), Gudino et al. 1068 (MO, QCNE). SUCUMBiOS:Reserva Faunistica Cuyabeno, nr. Laguna

Grande, 1 Apr 1988-1 Apr 1990 (st), Valenciaet al. 68505 (AAU, QCA). PERU. HUANUCO:Prov. Pachitea, Dtto. Honoria, BosqueNacionalde Iparia,Quebradade Macuya,W of Rio Pachitea,6 May 1968 ($ fr), SchunkeV 2573 (F, MO,MOL,NY, US). LORETO: Prov.Maynas,Quebrada Yanomono,Rio Amazonas, above Rio Napo, 15 Nov 1979 (a), Gentryet al. 28074 (MO, U); Prov.Maynas, between Indianaand mouthof Rio Napo, 10 Jul 1983 (st), Gentryet al. 42848 (BG, F, MO); Prov.Maynas, Dtto.LasAmazonas,nr.Sucusari,1 Mar1991(st),Pipoly et al. 13979 (MO);Rio Mazan,Gamitacocha, 25 Feb 1935

FLORANEOTROPICA

collectionSchunkeV2573theindumentum is moreor less transitional to thatof subsp.mollis,possiblydue to retentionofjuvenilefeatures. 9. Perebearubra(Trecul)C.C.Berg,Fl. Ecuador 60: 87. 1998.NoyerarubraTrecul,Ann.Sci.Nat.Bot. Ser.3, 8: 136. 1847.Perebeamollis(Poeppig& Endlicher) Hubersubsp.rubra(Trecul)C.C.Berg, ActaBot. Neerl. 18: 463. 1969;C. C. Berg,Fl. Neotrop.Monogr.7: 60. 1972. Tree,to 20 m tall,mostlydioecious.Leafjytwigs34 mmthick,with(sparse)veryshortpatenthairsintermixedwith(rather) sparsedistinctlyandlongermostly ? appressed yellowto brownishhairs;peridermflaking off or persistent.Laminaelliptic to oblong to (sub)ovate,8-20(-30) x 4-9(-14.5) cm, slightly inequilateral, chartaceous to coriaceous;apexacuminate;baseobtuseto subcordate; margin(sub)entire; uppersurfacesparselystrig(ill)oseto hirtellousto subglabrous, sparselyhirtellouson themainveinsor onlythemidrib,glabrescent, smoothorfaintlybullate; lowersurface(rather) puberulous andwithlongeryellowtobrownish, mostly? appressed hairs;lateralveins 12-16pairs,loop-connected 2-4 mmfromthemargin; tertiary venationlargelyscalariform; petiole0.3-0.6cm long;stipules0.5-1(-1.5) cmlong,yellow-subhirsute to subsericeous.Staminate inflorescences up to 7 to-

getheron minutespursin the leaf axilsorjustbelow theleaves;peduncle0.7-1 cmlong,slender,minutely puberulous; receptacle0.4-0.7 cmdiam.,discoid;involucrewithbractsin ca. 5 rows,thebractsminutely puberulous,the outerones ovate andca. 1.5-3 mm long,theinneroneslanceolateand3-8 mmlong,with theupperpartincurved; tepals3-4, (almost)free,1-1.5 mmlong,minutelypuberulous; stamens2-4; filaments ca. 1.5mmlong;anthersca.0.8-1 x 0.4-0.5 mm.Pistillate inflorescences solitaryor sometimes accompa-

niedbyasmanyas3 staminate ones,ca. 1-1.5cmdiam.; subsessileorpeduncleto 1.5cmlong;involucral bracts in 6-9 rows,broadlyto narrowlyovate,acuteto caudate,appressed-puberulous to subsericeous; flowers ca.5-10,freeorconnate; perianth 2.5-3 mmlong,entire to 4-dentate,puberulous to hirtellousto hirsute,if the perianth free,thensetosenearthebase;style1-1.5mm long,glabrous, stigmas2-3.5 mmlong.Infructescences brownto yellowish (a'), J. M. Schunke 319 (F, US); Prov. Maynas, Indiana, 2.5-4 cmdiam.;fruitingperianth 1 Dec 1988 (a), R. V6squezet al. 11275 (MO). PASCO: hirtellous to hirsuteorpuberulous. Prov.Oxapampa,Rio Iscozacin, Palcazu, 10 Jan 1984 Twosubspeciescanbe recognized.

(? fr), Foster et al. 7851 (BG, F, MO). BRAZIL. AMAZONAS:Rio Solim6es,Amatura, 13Oct 1945 (9 fl), Ducke 1789 (K, NY, RB, US).

9a. Perebearubra(Trecul)C. C.Bergsubsp.rubra.

Injuvenile materialthe hairs on the midribof the lamina beneath are patent as in subsp. mollis. In the

of laminaoftenslightlybullate,areolesof lowersur-

Peridermof leafy twigs persistent. Uppersurface

TAXONOMICTREATMENT

263

face concave. Pistillate inflorescences(sub)sessile;pistillate flowers connate;perianthpuberulous. Distribution (see Fig. 83.1). In Suriname,French Guiana, Brazil (northern Para and Amazonas, near Manaus);in non-inundatedforest;at low elevations.

Distribution (see Fig. 83.1). In the upperAmazon Basin (Brazil, Peru,and Ecuador);in non-inundatedforest;at low elevations. Additional collections examined. ECUADOR. NAPO:Canton Orellana, Reserva Floristica El Chuncho, 5 km N of Coca, 23 May 1993 (st), Palacios 10821 (MO). PERU. LoRETo:Prov. Maynas, Dtto. Las Amazonas,

Representative additional specimens examined. FRENCHGUIANA.AratayeR., SautPapara,29 Sep 1983 Rio Napo, nr. Sucusari, 22 Feb 1991 (d), Pipoly et al. (e), Granville5653 (U); Saul, La Fumee Oeste, 6 Apr 13372 (MO), Nov 1990 (st), Pipoly et al. 14329 (MO); 1983 (9 fl-fr), Mori et al. 15546 (MO, NY); Sinnamary Prov. Requena, Jenero Herrera, Mar 1984 (st), Spichiger R., abovePetitSaut,betweenCr.PlombandCr.Tigre,27 et al. 1750 (BG); Prov. Maynas, Allpahuayo, Nov 1990 (st), R. Vdsquez et al. 14791 and 15170 (MO). Aug 1993 (o), Mori et al. 23401 (BG, NY). BRAZIL. ACRE: Upper Rio Moa, Fazenda Arizona, BRAZIL.AMAZONAS: Mun.Manaus,ca. 90 km NNE of Manaus,DistritoAgropecuaria da SUFRAMA,Fazenda 24-30 Sep 1984 (st), Campbell et al. 6401 and 8451 P6rto Alegre, 9 Dec 1989 (od),Camposet al. 49 (BG, GH, (BG); Serra da Moa, 1 May 1971 (? fl-fr), Maas et al. R NY, US). PARk: MonteDourado,GlebaMonteDourado, 12699 (F, NY, U). AMAZONAs: Mun. Jutai, Rio Copatana, 15 Feb 1988 (? fl + e), Pires et al. 1997 (NY); Rio Jan 26 Oct 1986 (? fl-fr), Daly et al. 4146 (BG, MO); Rio Estirio

region, rd. Planalto A-Tinguelim, km 6, 7 Mar 1970 (e),

Javari,

N. T Silva 2935 (F, NY, U).

et al. P17226 30 Sep 1976 Rd. P6rto

9b. Perebea rubra (Trecul)C. C. Bergsubsp.glabrifolia (Ducke)C. C. Berg,Fl. Ecuador60: 87. 1998. Noyera glabrifolia Ducke, Bull. Mus. Hist. Nat. (Paris),Ser 2, 4: 722. 1932. Perebea glabrifolia (Ducke) C. C. Berg, Acta Bot. Neerl. 18: 463. 1969; C. C. Berg, Fl. Neotrop. Monogr.7: 60. 1972. Peridermof leafy twigs often flaking off. Uppersurface of lamina often plane, the areoles of lower surface plane. Pistillate inflorescences with ca. 1-1.5 cm long peduncle; pistillate flowers free; perianthhirtellous to hirsute,towardsthe base setose.

de Ecuador,

8 Aug

1973

(q fl-fr),

Lleras

(F, MO, NY, US). MATO GROSSO: Aripuank, Gomes et al. 266 (INPA). RONDONIA: (e),

Velho-Cuiaba,

1 km NE of Ariquemes,

13

Aug 1968 (V), Forero et al. 7038 (MO, NY, U).

Maquira 15. Maquira Aublet(see Berg, Fl. Neotrop.Monogr. 7: 62-75. 1972). Additionalcollections, especially those made in western Ecuadorand various localities in the upper Amazon Basin, have led to combining Maquira guianensis andM. costaricana at the specific level. Furtherbotanical exploration revealed for all taxa widerranges of distributionthanknown before.

Revisedkey to the speciesof Maquira 1. Tertiaryvenationof laminalargely scalariform 2. M. sclerophylla ........... ................................. 1. Tertiaryvenationof laminareticulate. 2. Peduncleof staminateinflorescencesto 0.5 cm long, usuallybracteate;pistillateinflorescenceswith 4. M. coriacea one flower;stigmasfiliform....................................... 2. Peduncleof staminateinflorescencesoften longerthan0.5 cm, withoutbracts;pistillateinflorescences with severalto many free flowersor with few connateflowers(rarelyone flower);stigmastongueshaped. 3. Pistillateinflorescenceswith severalto many free flowers;stamensfree; lower surfaceof lamina 1. M. guianensis smooth,the marginof laminaweakly or not callose..................................... 3. Pistillateinflorescenceswith a few connateflowers(rarelyone flower);stamensconnateat the base; the marginof laminadistinctlycallose.... 3. M. calophylla lower surfaceof lamina(usually)scabridulous,

1. MaquiraguianensisAublet(see Berg,Fl.Neotrop. no longerpossible to recognize these two taxaas disMonogr.7: 64-68. 1972,incl.M. costaricana). tinct at the species level, but only as subspecies. Maquiraguianensis andM. costaricana couldiniandgeo- la. M. guianensis Aublet subsp. guianensis. bothmorphologically tiallybereadilytoldapart, graphically.But featuresof collectionsmadesincein Tree,to 25 m tall.InvolucralbractsdenselybrownwesternEcuadorandintheupperAmazonBasinmakeit ish-puberulous. Peduncleofthe staminateinflorescence

264 0.5-2 cm long. Peduncleofthepistillate inflorescence

FLORANEOTROPICA al. 1898 (BG, MO,NY); Rio Le6n, 15 km W of Chigo-

0.5-1.6cmlong;pistillateflowersusuallymorethan20 rod6, 12 Mar 1962 (9 fr), Feddema1842 (MICH,NY). Rio Caqueta,CainoPaujil,28 Nov 1988 (st), (to50).Fruitingperianth subobovoid, often? distinctly CAQUETA: M. Sanchez S. et al. 1711 (BG). CAUCA:Rio Micay, ribbed,brownvelutinous,witha ? truncateapex. Guayabal, 25 Feb 1943 (a), Cuatrecasas 14135 (F); Rio

Distribution(seeFig.84.3).IntheGuianas,east- Micay,El Chacjajo,27 Feb 1943 (9 fl-fr), Cuatrecasas andAmazonian Brazil(Amapa, 14264 (F). CHoc6: Teresita,18 May 1967 (? fr), Duke emVenezuela (Bolivar), Para,MatoGrosso,andRoraima);in non-inundated 11040 (MO, NY, US); rd. San Jose de Palmar-N6vita, 20 Feb 1977 (d), Forero et al. 3041 (MO,U). NARINO: forest;atlowelevations.

Tumaco,Rio Rosario,5 km above SantaMaria,20 Jun Representative additional specimens examined. 1955 (e), Romero-Castafieda 5179 (MO, NY). VENEZUELA. BoLivAR: RioCaura,RaudalGuanagujafia, VENEZUELA. BOLIVAR:Dtto. Roscio, "El Abismo," 9-26 May 1988 (od), Stergios et al. 12610 (BG, NY). 23 Oct 1985 (? fl-fr),Holst et al. 2446 (BG, NY, US); GUYANA.UpperDemerara-Berbice region,Berbice Dtto. Piar,Rio Acapan,Guarumo,5 km W of AmaruayR., AroiamaMiningCo., 17 Apr 1993(st),Hoffmanet al. tepui, 16 Apr 1986 (a), Liesner et al. 20037 (MO). 3968 (BG). ECUADOR. ESMERALDAS: Mataje,RioMataje,9 Sep BRAZIL.MARANHAO: Mun.Mongao, Rio Tutia9u, 1991 (), Jaramillo 13876 (NY, QCA); Rio Cayapas, 4 kmNW of Urutawy,11 Feb 1985(st), Balee et al. 608 ZapalloGrande,19-25 Oct 1983 (? fl), Barfod et al. (NY); Mun. Carutapera,GurupiunaKa'apor Indian 48442 (AAU, BG, QCA). GUAYAS:Cord. Chong6nReserve,5 Nov 1986 (st), Balee et al. 2861 (NY). MATO Colonche, Jul 1996 (a), Bonifaz et al. 3410 (BG). Los GROSSO:Rio Aripuana, nr. Humboldt Center, 17 Oct Rios: RioPalenque BiologicalStation,rd.Quevedo-Santo 1973 (e), Berg et al. P.18608 (MO, NY). ROND6NIA: Domingo de los Colorados,km 56, 16 Sep 1977 (a), Mun. P6rtoVelho,Usina HidroelectricaSamuel,12-24 Berg et al. 437 (AAU, NY), 24 Oct 1974 (a'), Gentry Sep 1988 (9 fl), Thomaset al. 6202 (BG). RORAIMA: 12011 (MO,QCA, S, U). MANABI:Cant6nPedernales, Trail Surucucu-Uaica,between Maita and Paramiteri, 10 km E of Pedernales, 19-21 1996 (a'), J. L. Clark et 17 Feb 1971 (? fr),Pranceet al. 10578 (MICH,NY,U). al. 2718 (QCNE). NAPO:Reserva Etnica Huaorani, The clearly disjunct occurrencein northernMato Grosso, as in Bagassa guianensis, is noteworthy.

lb. Maquira guianensis subsp. costaricana (Standley) C. C. Berg, Novon 6: 235. 1996. Tree,to 15(-30) m tall. Involucral bracts (rather) Peduncleof thestaminate sparselywhitish-puberulous. inflorescence 0.2-0.8 cm long. Peduncle of thepistillate inflorescenceto 0.8 cm long; pistillateflowers ca. 10-35. Fruitingperianth ellipsoid to obovoid, not or hardly ribbed, (sub)glabrous or sparsely to densely yellowish-puberulous,with an obtuseto roundedapex. Distribution (see Fig. 84.3). From Nicaragua throughwestem Colombiato westernEcuador,and in the upperAmazonBasin, extendingto easternVenezuela (Bolivar); in non-inundatedforest; mostly at low elevations, in Roraima(Brazil) to 1800 m. Representative additional specimens examined. PANAMA.BoCAS DEL TORO:Rd. to Chiriqui Grande, CerroPilade Arroz,10 Mar1986(? fl), McPherson8764 Rd.La Pintada-Coclesito, 7 Feb 1983 (MO,NY). COCLA: (d'), Hamiltonet al. 2828 (MO,NY). DARI*N: Rio Ucurganti, 7 Jul 1967 (e), Bristan 1188 (MO, NY). PANAMA: Ca. 20 km NE of Chepo,12 Dec 1973(? fl), Berget al. 317 (BG);BarroColoradoIsland,18 Apr 1970(at),Croat 9786 (MICH,MO,NY, U); El Llano-Cartird.,km 10.8, 27 Dec 1974 (ci), Mori et al. 4137 (AAU, MO, NY, WIS). COLOMBIA.ANTIOQUIA: Mun.Mutata,rd.Mutata-

Pavarand6,km 1, 27 Apr 1987 (9 fl-fr), Fonnegraet

Maxus rd., km 111-112, 1-4 Sep 1994 (a'), Dik et al.

1581 (BG);ParqueNacionalYasuni,Pozo PetroleroAmo 2, 9-19 Jan 1988 (a'), Neill et al. 8349 (BG, MO, NY, Rd. Coca-Auca,110 km S of QAME,QCNE).PASTAZA:

Coca,7 Jan 1989(? fl), Palacioset al. 3383 (AAU,BG, MO, NY, QAME). SUCUMBfOS:Reserva Faunistica Cuyabeno,nr. LagunaGrande,1 Apr 1988-1 Apr 1990 (st), Valencia et al. 68019 (QCA) and 68242 (AAU).

PERU.Cuzco: Prov.Paucartambo, Cosfiipata,May 1914 (? fl), Weberbauer6957 (F, GH, MOL).LORETO: Prov.Maynas,Dtto. Las Amazonas,"RocaEterna,"17 Aug 1991 (a'), Grandez et al. 2858 (BG); Prov. Maynas,

Rio Nanay,SantaClara,12 May 1977 (st), Revillaet al. 2544 (MO);Prov.Requena,JenaroHerrera,25 Jan 1985 (st), Spichiger et al. 3057 (NY). MADRE DE Dios:

NatureReserve,confluenceof Rio La Torre Tambopata andRioTambopata, 27 May 1987(st), Gentryet al. 57727 (AAU,BG, MO,NY); Prov.Manu,Rio Manu,Puestode Pakitsa, 6 Oct 1987 (st), Sobrevilla et al. 1889 (F). BRAZIL.ACRE: PortoAcre, 2 Nov 1993 (? fl-fr), Daly et al. 8010 (NY); Rio Branco, 30 Oct 1981 (a'), R. F Silva et al. 101 (INPA). AMAZONAS: Mun.Sao Paulo de Oliven9a,BelemCr.,26 Oct-l Dec 1936(a), Krukoff

8642 (BM, K, LE, MICH, MO, NY, P, U, US); Rio Maturaca,between PalmitoCampand TatuCamp,21 Dec 1965 (? fr), N. T. Silva et al. 60702 (NY, U). RORAIMA:Serrados Surucucus,1800 m, 15 Feb 1969 (9 fl), Prance et al. 9954 (NY). BOLIVIA.PANDO:Prov.Nicolas Suarez,San Jose, 14 Jan 1983 (a'), Fernandez Casas et al. 8249 (NY).

In western Ecuadorthe fruiting perianthis often sparsely to densely yellow-puberulous; some of the specimensfromthis area(Rio PalenqueBiological Sta-

TAXONOMIC TREATMENT

265

tion)havebeendistributed undertheunpublished name Maquiragrandis Gentry.

InCentralAmericathepistillateinflorescences oftencontainmorethan20 flowers,whereasin western Ecuadorandeastof theAndestheyusuallyhaveless than20 flowers,oftenca. 10 or evenless. Moreover, eastof theAndestheleavestendto berelativelysmall, oftenabout10cmlong,with7-10 pairsof lateralveins. InCentral AmericaandwesternColombia, thetrees appearto startfloweringwhensmall,a featureshared with some other species of the same tribe, e.g., Helicostylis tomentosaandPerebeaangustifolia.

2. Maquirasclerophylla(Ducke)C. C. Berg (see Berg,Fl. Neotrop.Monogr.7: 69. 1972).

VENEZUELA. AMAZONAS:Depto. Atabapo, Rio Putaco,RaudalXikita-pora,Feb 1990 (c'), Ferndndez 7035 (BG,NY); Dtto. Rio Negro, 1-4 km E of Cerrode La NeblinaBase Camp,Rio Mawarinuma,10 Feb 1984 (2 fl-fr), Liesneret al. 15824 (BG, MO). FRENCHGUIANA. Stationdes Nouragues,basin of AratayeR., 7 Mar1987(st), Sabatier1540 (BG,MO); Piste de St. Elie, 18 Aug 1987 (st), Sabatieret al. 1747 (BG, MO). ECUADORNAPo:CantonOrellana,Huashito,20 km N of Coca, 3-21 Nov 1989 (a), Rubio 298 (BG, NY, QCNE);ReservaBiol6gica JatunSacha,3-4 Jun 1988 (a'), Neill et al. 8478 (BG, MO, NY, QCNE);Aiiangu, ParqueNacionalYasuni,1-15 Feb 1986 (q fr), Korning et al. 47481 (AAU,QCA,QCNE).PASTAZA: 10 km from Napo border,Estaci6nPetroleroTigiuino2, 9 Jan 1989 (? fl), Palacios 3453 (BG, MO, NY, QAME, QCNE). SUCUMBtOS: CantonGonzaloPizarro,Rio Tigre, 900-

Distribution.InthethreeGuianasandthenorthern 1100 m, 18-21 Feb 1987 (? fr), Palacios et al. 1598 (BG, MO, QAME, QCNE); Reserva FaunisticaCuyabeno, toAmazonas nr.LagunaGrande,1 Apr 1988-1 Apr 1990 (st), Valencia partoftheAmazon Basin,fromParA (Brazil) (Venezuela), andapparently disjunctlyin thesouthern 67840 (AAU, QCA). partof the AmazonBasin (Rond6nia,with a single PERU. AMAZONAS: Rio Santiago, nr. Caterpiza, 21 record);in non-inundated forest;atlowelevations. Nov 1979 (? fr), Huashikat1359 (MO, U). HUiNUCO: Representative additional specimens examined. VENEZUELA.AMAZONAS:Karohi, 3 Nov 1975 (6'), Lissots.n. (VEN, U). GUYANA.RupununiDistrict, between Kuyuwini Landingand KassikaityuR., 23 Oct 1992 (e), JansenJacobs et al. 3072 (MO, NY). FRENCHGUIANA.Sauil, 8 Oct 1991 (e), Moriet al. 22002 (BG, NY). BRAZIL. AMAZONAS: Rd. Manaus-Caracarai, BR.174, km 159, 20 Sep 1974 (at), Prance et

al. 22716

(MO, NY, U); Rio Maturaca,betweenMalocaand Rio Cauaburi, 31 Dec 1965 (2 fl-fr), N. T Silva et al. 60779

13 Nov 1978 (NY,U). PARA:Rd. Itaituba-Jacareacanga, (9 fl-fr), M. G. Silva et al. 3707 (MG,NY);Rio Jari, Monte Dourado,between Pilao and Repartimento,Nov 1968 (? fl-fr), N. T?Silva 1354 (NY,U). ROND6NIA:Mun. P6rto Velho,UsinaHidroelectrica Samuel,12-24 Sep 1988(st), Thomas et al. 6570 (BG).

3. Maquira calophylla (Poeppig & Endlicher)C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 70-72. 1972). Distribution. In the upper Amazon Basin, from Venezuelato Bolivia, and in French Guiana;in noninundatedandperiodicallyinundatedforest;at low elevations.

Prov. Puerto Inca, Dtto. Yuyapichis, Reserva Forestal DANTAS,16-31 Jul 1989 (a'), KroellS. 514 (NY), 1630 Jun 1990 (st), Tello 839 (MO). LORETO:Prov. Maynas, Dtto. Iquitos, Rio Nanay, trail Shihua-Momoncillo, 14 Mar 1977(a), RimachiY 2879 (MO,NY). MADRE DE DIos: TambopataNature Reserve, nr. jct. of Rio La Torre and Rio Tambopata, 24 May 1987 (st), Gentry et al. 57533 (BG, MO). PASCO:Palcazu valley, 27 Nov 1984 (a), Hartshornet al. 2665 (BG, MO). PUNO:Rio Candamo, nr. mouth of Rio Guacamayo, 25 May 1992 (st), Gentry et al. 77192 (MO). BRAZIL. ACRE: Upper Rio Moa, Fazenda Arizona, 24-30 Sep 1984 (st), Campbell et al. 8429 (BG). ROND6NIA: Minera9ao Campo Novo, ca. 100 km SW of Ariquemes, 14 Oct 1979 (? fl), Zarucchi et al. 2700 (GH, MICH, MO, NY, U). BOLIVIA. BENI: Cachuela Esperanza, 1924 (? fl-fr), Meyer339 (U). PANDO: 74 km SW of Cobija, PuertoOro, 13 Aug 1988 (q fr), R. T. Pennington et al. 117 (BG).

The collectionKrukoff8642,in the previousmonographreferredto Maquira calophylla, belongs to M. guianensis subsp. costaricana.

4. Maquira coriacea (Karsten)C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 73-75. 1972.)

Distribution. In the Amazon Basin, the Orinoco Basin, the northernRio ParaguayBasin, in northwestRepresentative additional specimens examined. em andeasternVenezuela,andapparentlydisjunctlyin COLOMBIA. AMAZONAS:ParqueNacional Natural Goias (Brazil), where known from a single record;in Amacayacu,nr.Mocagua,20 Jan 1988 (9 fr), Gentryet inundatedforest. al. 60810 (BG, MO); Rio Caqueta, nr. Araracuara,12 periodically Dec 1990 (st), Vester182 (BG); Leticia,Sep 1963 (e), Representative additional specimens examined. Schultes s.n. (ECON). COLOMBIA.AMAZONAS: Rio Caqueta,QuebradaEl

266 Engefio,9 May 1988(st), M. SdnchezS. et al. 297 (BG). VAUPtS:Nr. Mitu, 24 Mar 1970 (q fr), Soejartoet al. 2379 (HUA). VICHADA: ParqueNacional Natural El Tuparro,ca. 9 km NE of El Tap6n,(? fl), Zarucchiet al. 3746 (BG, GH, MO, NY). VENEZUELA. AMAZONAS: Depto.Atures,Serrania Batata,24 kmNEof SaltoColorado,Oct 1989(st),SanojaAngelet al. 3306 (BG). APuRE:Dtto. R6mulaGallegos, Lagunadel Termino,6 Mar 1979 (a'), Davidse et al. 16064 (MO, U). BOLIVAR: Between mouth of Rio 3-10 Aug 1985(9 fr),Horner NichareandCailoSarrapio, et al. 163 (MO). ECUADOR. SUCUMBIOS: Reserva Faunistica Cuyabeno,nr. LagunaGrande,1 Apr 1988-1 Apr 1990 (st), Valenciaet al. 68725 (AAU, QCA). PERU. LORETO: Prov. Requena, Rio Ucayali, 3212 (BG, NY). Yucuruchi,21 Aug 1982(a), R. Vdsquez and MADRE DE Dios: Nr. confluenceof Rio Tambopata Rio La Torre,39 km SW of PuertoMaldonado,Laguna Cocacocha,20 Jan 1989 (? fr), S. F Smithet al. 1445 (F, NY, US). BRAZIL.AMAPA: Estradado IgarapePacoval,NE of Macapa,13 Nov 1979 (9 fl-fr),Austinet al. 7384 (MO, NY);Rio Cupixi,2 Nov 1976(st),N. A. Rosa 1024 (NY). GoIAs:Rio Araguaia,Caseara,17 Aug 1978 (e'), N. T Silva4841 (MO,NY,U). MARANHAO: Mun.SantaLuzia, ca. 83 km W of SantaInez, 13 km E of Entrocamento, 26 Mar1983(a), Tayloret al. 1079(NY).MATOGRosso: Rio Aripuana,aboveAndurinafalls, nr. HumboldtCenter, 19 Oct 1973 (? fl), Berg et al. P 18663 (MO,NY); Mun. Pocone, 1 km NW of PortoJoffre, 26 Oct 1985 (a'), Thomaset al. 4626 (BG, MO,NY); Mun.Juara,45 km SW of Juara,FazendaMaringa,27 Oct 1987 (? fl), Tsugaruet al. B.1174 (NY). RORAIMA: Withoutlocality, "areapantanosa,"16 Feb 1977 (9 fl), N. A. Rosa et al. 1514 (NY). BOLIVIA. BENI: Prov. Vaca Diez, 20.8 km E of 25 Sep 1981(e), Solomon rd.to Guayaramerin, Riberalta, 6450 (BG, MO, NY). LA PAZ: Prov. Abel Iturralde, Luisita, 25 Feb 1984 (? fr), Beck et al. 10015 (BG). PANDO:Prov.Manuripi,Rio Manuripi,5 km fromSan Silvestre,20 Oct 1989 (a), Beck et al. 19564 (BG).

FLORANEOTROPICA Additionaltype: TrymatococcusguanabarinusDuarte, Rodriguesia 23/24: 55. 1964),as Trimatococcus guanabarinus. Type.Brazil.Rio de Janeiro,Estradado Sumare, Lagoinha,Duarte 5658 (holotype, RB). Distribution. In northernSouth America, eastern Brazil (northof Rio de Janeiro),and Panama;in noninundatedforest;to 1600 m. Representative additional specimens examined. PANAMA. BOCASDELToRo: Laguna de Chiriqui, Nuri,

19 Mar 1993 (9 fl-fr),Fosteret al. 14605 (F, SCZ),(a), Foster et al. 14609 (F, SCZ); rd. to ChiriquiGrande, CerroPila de Arroz,10 Mar 1986 (a), McPherson8751 (BG, MO);Chiriquicito,PuntaPefna,7 Jun 1967 (? flfr), Lewis et al. 2188 (MO, U). COLOMBIA. AMAZONAS:Rio Popeyaca, 25 Feb

1952 (? fl-fr),Schulteset al. 15639 (US). ANTIOQUIA: Mun.Cocorna,Vrda.La Pifiuela,rd. to San Francisco, 1 Sep 1991(? fr),G. Canas499 (AAU,MO);Mun.Anon', above Providencia,24-31 May 1973 (d"), Soejarto et al. 4104 (GH,MO, NY). BOLtVAR: La Raya, Quebrada La Culebra, between jct. of Rio Cauca and Rio Magdalena,5 May 1987 (st), Gentryet al. 57391 (BG). CHOC6:Mun. Riosucio, Uruba region, Cerro del Cuchillo, 13 Apr 1988 (e"), Cdrdenas1610 (JUAM). C6RDOBA: Jct. of Rio Tigre and Rio Manso,Paramillo National Park, 28 Jul 1988 (st), Gentryet al. 63856 (MO).META: Macarena,Serraniade Chamusa,RioDuda, Feb 1988 (? fl-fr), Stevenson25 (MO);Macarena,Rio Juejar,24 Dec 1950 (? fl), Schulteset al. 11159 (GH). NARI1SO: Mun. Iscuande, Rio Sequi6n, 22 Nov 1955 (a"), Romero-Castanieda5483 (COL). SANTANDER:

CampoCapote,30 km E of Carare,30 Sep 1977 (? fl), Gentryet al. 20099 (MO, NY); Rio Carare,between Ermitanloand Cimitarra,31 Jul 1954 (a), Lamb 142 6 Dec (MICH).VALLE:Ca. 10 km N of Buenaventura, 1981 (st), Gentry35387 (MO). VENEZUELA. TACHIRA: Rd. El Vigia-La Fria, Coloncito,30 Oct 1978 (st), Marcano-Bertiet al. 2-1078 (MER). ECUADOR. ESMERALDAS: San Miguel, Rio Grande,

22 Nov 1992 ("), Tipazet al. 2319 (BG). MORONASANTIAGO: Cordillerade Cutucui,rd. Mendez-Morona, Helicostylis 2 Feb 1989 (a), Werff 10290 (BG, MO, QCNE); 16. HelicostylisTr6cul(see Berg,Fl. Neotrop.Monogr. Bomboiza, Mision Salesiana-Shuar,6 Nov 1986 (st), Neill et al. 7455 (QCNE).NAPO:CantonFranciscode 7: 75-92. 1972). Orellana,rd. to Los Zorros,Pozo PetroleroJaguarI, 23 Furtherbotanical explorationhas not changed the Oct 1988 (? fl + a), Palacios 3201 (BG, MO, NY, taxonomy of the genus. For all species known, the QAME,QCNE);San Jose de Payamino,1-7 Dec 1983 rangesof distributionhavebecomemoreextensive.The (e), Batslevet al. 4573 (AAU,BG, NY, QCA);Reserva Biol6gica JatunSacha, 8 km E of Misahualli,22 Sep most prominent range extensions are found in 1989 (a"), Palacios 4503 (BG, MO, NY, QAME). Helicostylis tomentosa (to the Pacific Coastal region PASTAZA:Rio Curaray,Lorocachi,20 May 1980 ($ fland Panama)andH. pedunculata (to Bahia). fr), Brandbygeet al. 31044 (AAU, BG, MO, QCA); Cant6nPastaza,Pozo PetroleroMasaramu,40 km NE of Montalvo, 1-17 May 1990 (2 fl-fr), Espinoza 235 1. Helicostylistomentosa(Poeppig& Endlicher)Rusby (NY,QCNE).SucuMBios:ReservaFaunisticaCuyabeno, (see Berg, Fl. Neotrop. Monogr. 7: 79-81. 1972). nr.LagunaGrande,11 Mar1990(a), Balslevet al. 97064

TAXONOMIC TREATMENT

(AAU,BG, QCA,QCNE).ZAMORA-CHINCHIPE: Canton Nangaritza,Rio Nangaritza,Miazi, 1000-1100 m, 11 Dec 1990 (o), Palacios et al. 6754 (BG, MO, QCNE). PERU. AMAZONAS: N of Rio Cenepa,nr. mouthof QuebradaCikan-inci,30 Dec 1972 (? fl-fr),Berlin 757 (NY). HUANUCO: Prov.LeoncioPrado,Dtto.JoseCrespo y Castilla,Quebradade Sangapilla,nr.Aucayacu,4 Mar 1978 (%), Schunke V 9987 (BG,NY, U). LORETO:Prov. Requena,3 km E of JenaroHerrera,28 Nov 1988 (a),

267

fl-fr), Garcia-Barriga 13945 (NY). CAQUETA:Nr. 22 Nov 1988 (e), M. SanchezS. et al. 1657 Araracuara, (BG). VAUPtS:Rio Apaporis,nr. RaudalYayacopi(La Playa), 18 Feb 1952 (? fr), Schulteset al. 15466 (U). San Carlos de Rio NeVENEZUELA. AMAZONAS:

gro, 3 May 1979 (? fl-fr),H. L. Clark7176 (MO,NY); Rio Cunucunuma,3 Nov 1950 (e), Maguire et al. Cerrode La Neblina 29368 (NY,U); Rio Mawarinuma, Base Campon Rio Baria,26 Jan 1985 (? fl), Nee 30549 Daly et al. 5779 (NY). (MO, NY). BRAZIL.BAHIA:Rd. PortoSeguro-Eunapolis,km CantonNangaritza, ECUADOR ZAMORA-CHINCHIPE: 5, 18 Feb 1972 (de), Eupunino224 (U); Mun.Urusuca, Miazi, 1000m, 9 Dec 1990 (e fl), Neill et al. 9614 (BG, 7.3 km N of SerraGrande,on rd. to Itacare,7 May 1992 MO, QCA, QCNE). (d' fl-fr), Thomaset al. 9187 (NY). ESPIRITOSANTO: PERU. AMAZONAS:Prov. Bagua, Dtto. Imaza, Linhares,ReservaFlorestal,da CVRD,2 Feb 1985(? fl), Kampaenza,25 Sep 1994 (o), N. Jaramilloet al. 505 Peixotoet al. 3210 (BG, MO).MARANHAO: Turia9u,rd. (BG). LORETO:Prov. Requena, Jenaro Herrera, 21 Nov BR.106,km 6, 30 Nov 1978(? fl), N. A. Rosaet al. 2751 1988 ($ fl-fr), Daly et al. 5802 (BG, MO, NY); Prov. (MO). MATOGROSSO:Mun. Vila Bela da Santissima AltoAmazonas:nr.Pongode Manseriche,Rio Maranion, 4 km S of Rond6niaState 19-21 Oct 1962 (ae), Wurdack2325 (NY). Trinidade,rd.Vilhena-Cacera, line, 3 Nov 1985(q fl), Thomaset al. 4796 (BG).MINAS BRAZIL. AMAZONAS: Mun. Manaus, Distrito Reserva1501,26 Nov 1988(9 fl-fr),Boom GERMS:CoronelPacheco,EstaqaoExperimental de Cafe, Agropecuaria, Rio Uatuma,13 Aug 29 Nov 1942 (e), Heringer1137 (US). RORAIMA:Be- et al. 8655 (NY);Mun.Itapiranga, tweenMaitaandParamiteri, 18 Feb 1971(? fl-fr),Prance 1979 (2 fl), Ferreira et al. 354 (MICH, MO, NY). et al. 10602(MICH,NY,U); Rio Mucajai,PostoMucajai, ROND6NIA:38 km from Vilhena, 29 Oct 1979 (? fl), Vieiraet al. 762 (NY). 26 Feb 1971 (2 fr), Prance et al. 11240 (NY,U). BOLIVIA.BENI: Prov.VacaDiez, nr.Alto Iv6n, 29 Jan 1984 (? fr), Boom4313 (NY);CachuelaEsperanza, Nov 1923, Meyer 336 (MO, U). COCHABAMBA: Prov. 3. Helicostylis elegans (Macbride) C. C. Berg (see Carrasco,Estaci6nValle del SajtaUMSS, 31 Oct 1991 Berg, Fl. Neotrop. Monogr. 7: 83-85. 1972). (st), Villegas203 (MO).LAPAZ:Prov.Larecaja, Consata, Distribution. In the upper Amazon Basin, from 7 km fromMapiri,1400 m, 14 Dec 1981 (2 fl-fr),Beck 4949 (MO);Prov.NorYungas,10 km N of Caranavi,1 Colombia to northernMato Grosso (Brazil); in nonNov 1984 (a'), Nee et al. 30299 (AAU, MO, NY). PANDO: inundatedforest;at low elevations. Prov.Abuna,Nuevo Mundo,2 Jul 1992 (st), Gentryet Representative additional specimens examined. al. 77734 (MO). COLOMBIA. VAUPtS: Falls of Rio Yurupari,24-26 Several collections from northern Colombia Oct 1952 (e), Garcia-Barriga14978 (MO, NY). VENEZUELA. AMAZONAS:Ca. 10.5 km W ofYavita, (Antioquia,Bolivar,andCordoba)show featuresremi28 May 1996(st),Aymardet al. 11086 niscentof Helicostyliselegans. Morefertilematerialis rd.Yavita-Maroa, Mun.ForaneoAripao,upperRio Caura, needed to establish the identity of these collections, (MO).BOLIVAR: 1 km below Araguafia,1 May 1988 (a'), Aymardet al. representingeithera short-hairyform of H. tomentosa 6735 (BG). or a somewhatdifferentform of H. elegans, occurring ECUADOR.NAPO: Estacion Exp. INIAP,Reserva disjunctly. FloristicaEl Chuncho,5 km NW of Coca, 2 Oct 1987 Flowering trees of this species are normally me- (st), Palacios 2035 (BG, MO, QCNE). SucuMBios: Reserva dium-sized,but some collections from Panama,Lewis FaunisticaCuyabeno,nr. LagunaGrande,11 Mar 1990 et al. 2188 andMcPherson8751, aremadefromshrubs (st), Balslev et al. 97056 (AAU, BG, QCA,QCNE). PERU. AMAZONAS: Rio Santiago, Quebrada or treelets (to 5 m tall). Caterpiza,2 Jan 1980 (a), Tunqui531 (BG, MO),4 Jan 1980 (? fl-fr), Tunqui 538 (MO). LORETO: Prov. 2. Helicostylis scabra (Macbride) C. C. Berg (see Maynas,Dtto. Iquitos,Rio Nanay,rd. to Picuruyacu,18 Oct 1978 (e), Rimachi Y 4029 (MO, NY); Prov. Berg, Fl. Neotrop. Monogr. 7: 82. 1972). Requena,JenaroHerrera,30 Jan 1986 (! fl-fr),Spichiger andRio BetweenShiringamazu Distribution. In the upper Amazon Basin from et al. 3042 (NY).PASCO: Carachama,13 Jun 1986 (a'), Hartshornet al. 2990 Colombia to Rond8nia (Brazil); in non-inundated for(AAU, BG, MO). est; to 1000 m. BRAZIL. ACRE: 1-6 km NNW of Boca de AcreairRepresentative additional specimens examined. strip, 22 Sep 1966 (a), Prance et al. 2500 (INPA,NY, Rio Apaporis, beCOLOMBIA. AMAZONAS/VAUP*S: Rd. Manaus-P6rtoVelho, BR.319,km U). AMAZONAS: tween Rio Kananariand Rio Pacoa, 1-15 Dec 1951 (e 265, betweenRio Igapo-a9uand Rio Jutai,12 Oct 1974

268

FLORANEOTROPICA

(? fl), Prance et al. 22824 (NY, U); rd. ManausItacoatiara, km 74, 3 Sep 1965 ('), Rodrigues 7088

Kukle 11 (BG); rd. Manaus-Itacoatiara,Reserva Florestal Ducke, 5 Nov 1996 (d), Ribeiro et al. 1849 (BG, INPA).

(NY); N of Rio Negro, 2 km above Tapuruquara,18 This species is closely relatedto Helicostylis Oct 1971 (c), Prance et al. 15391 (MO, NY, U). MATO GRosso: Rd. fromHumboldtCenter,Rio Aripuana,rd. pedunculata. Morematerialis neededto confirmthat to Rio Juruena, 17 Oct 1973 (oi), Berg et al. P.18613

(MO, NY, U). PARA:Mocambo, EMBRAPAForest Reserva, 16 Nov 1984 (st), Gentry49122 (MO).

4. Helicostylis pedunculata Benoist (see Berg, Fl. Neotrop. Monogr. 7: 85-87. 1972).

thetwotaxacanbe treatedas speciesor shouldbe regardedas subspecies. 6. HelicostylisheterotrichaDucke (see Berg, Fl. Neotrop.Monogr.7: 87-90. 1972.)

Distribution.Inthenorthwestem partof theupper

Distribution. Fromthe Guianas(easternSuriname AmazonBasin;in non-inundated forest;atlowelevaand French Guiana) through Amapi to eastern Part tions. (Brazil),anddisjunctlyin Bahia(in theAtlanticCoastal Additional specimen examined. COLOMBIA. forest); in non-inundatedlowland forest. CAQUETA: Araracuara,Aug 1988 (R fr), M. Sanchez S. Representative additional collections examined. 909 (BG); Rio Caqueta, nr. Cafio Prujil, 20 Nov 1988 (st), M. SanchezS. et al. 1792 (BG). BRAZIL. AMAPA:ReservaINCRARio Falsino, 15-20 Aug 1983 (st), Campbell et al. 10159 (BG); Mun. Maza-

Thesmallnumberof additionalcollectionsmade gao, FelipeIX area,3 Feb 1987 (? fl), Pires et al. 1574 (NY).BAHIA: Mun.Una,Maruim,1 May 1981(st),Mori since 1972 suggests that this species is rare. et al. 13885(NY);P6rtoSeguro,ReservaFlorestalCVRD, 14 Jul-1988 (9 fr), Folli 772 (BG); rd. to Canaviviras, 7. Helicostylistovarensis(Klotzsch&Karsten)C.C. Berg(see Berg,Fl. Neotrop.Monogr.7: 90-92. Uru9uca,7.3 km N of SerraGrande,on rd. to Iracare, 1-12 Jul 1991 (st), Thomaset al. 8540 (NY). PARA: 1972).

Camasa, 22 Oct 1971 ('),

T S. Santos 1403 (U); Mun.

MonteDourado,3 Mar1988(9 fl), Pireset al. 2026(NY); rd. Capanema-Maranhao, BR.22, km 64, nr. Piritoro,4 Nov 1965 ('), Prance et al. 1950 (MO, NY); rd. Cuiabi-

Santarem,BR.163, km 1350, 11 km N of Rio Ariri,24 Nov 1977 (? fl + o), Prance et al. 25723 (AAU, MO,

NY); Rio Jari,Estradado Munguba,km 2, 7 Mar 1970 (9 fl + e), N. T Silva 2941 (NY).

5. Helicostylis turbinata C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 87. 1972). Distribution (see Fig. 84.1). In the upperAmazon Basin; in non-inundatedforest;at low elevations.

Distribution. FromCostaRicato theCoastalRange of Venezuelaand throughthe Andesto Bolivia;in montaneor submontaneforest;at ca. (400-)6002000(-2500)m. Representative additional specimens examined. PANAMA. COCLII:Cerro Pilon, 5 km NE of El Valle, 800-1050 m, 14 Jun 1975 (d), Mori 6630 (U). DARII?N: Cerro Mali, nr. Colombian border, 1400-1500 m, 2122 Jan 1975 (? fl), Gentryet al. 13781 (MO, U), (d'), Gentryet al. 13782 (MO, U). PANAMA:CerroJefe, ca. 900 m, 22 Dec 1978 (e fl), Berg 390 (F, MO). VERAGUAS: NW of Santa Fe, 18 Jun 1975 (? fl), Mori 6709 (AAU, MO, U). COLOMBIA. ANTIOQUIA: Mun. Campamento,Vrda. El Alto, 1810 m, 9 Sep 1989 (S fl), Callejas 8397 (BG, NY); Roca del Peniol,Aug 1947 (? fl), Daniel 4006 (F, US). HUILA:Mazizo Colombiana, San Augustin, 2420 m, 27 Aug 1958 (d'), Idroboet al. 2896 (P).NORTEDE SANTANDER: Sarare region, Quebrada del Sorrarito, ca.

Representative additional specimens examined. ECUADOR. NAPO: Parque Nacional Yasuni, Pozo PetroleroDaimi2, 26 May-8 Jun 1988 (9 fl-fr), Cer6n et al. 3837 (BG, QCNE);Pozo PetroleroAmo 2, 9-13 Jan 1988 (9 fl), Palacios 2405*(BG, NY, QAME, QCNE).SUCUMBiOS:Cant6nShushufindi,San Roque, 14 Dec 1997 (st), E. Freireet al. 2835 (QCNE);Reserva 1950 m, 22 Oct 1941 (? fl), Cuatrecasaset al. 12579 FaunisticaCuyabeno,nr. LagunaGrande,1 Apr 1988- (F). VALLE: Rio Anchicaya, between Pavas and 1 Apr 1990 (st), Valencia et al. 68597 (QCA).

Miramar, 350-450

m, 15-16 Apr 1943 (9 fl-fr),

PERU.LORETO: Prov.Maynas,Dtto.LasAmazonas, Cuatrecasas 14424 (F); Rio Sanquinini, La Laguna, 1400 nr. Sucusari,21 Feb 1991 (st), Pipolyet al. 13271 (BG); m, 10-20 Dec 1943 (9 fl), Cuatrecacas15545 (F). Prov.Maynas,Miraflores,Quebrada Tamshiyuca, 22 Feb VENEZUELA. SUCRE:Peninsula de Paria, Cerro 1986 (9 fl + c), R. Vdsquezet al. 7207 (AAU, BG, NY). BRAZIL. ACRE: FazendaBom Sossego, between

Igarap6do Cujubimand IgarapeJacamin,27 Sep-7 Oct 1985 (st), Campbell et al. 9312 (BG). AMAZONAS: Mun. Manaus,ca. 90 km NNE of Manaus,9 Nov 1989 (ei),

Humo, Rio Santa Isabel, between "Los Positos" and La Roma, 700-800 m, 11 Aug 1966 (9 fl), Steyermarket al. 96344 (F). TRUJILLO:Between Escuque and La Mesa de San Pedro, 1300-1650 m, 20-23 Feb 197 (? fl), Steyermark 104709 (F).

TAXONOMIC TREATMENT ECUADOR. ESMERALDAS:Canton San Lorenzo, Alto Tambo, 650 m, 23 Sep 1965 (d), Little et al. 21126 (F, QAME, US). CARCHI: Rio Blanco, above Chical, 13001500 m, 25 Sep 1979 (9 fl-fr), Gentry et al. 26555 (AAU, QCA, U); Gualpi'Alto 900-1000 m, 18-25 May 1985 (a), Thomsen 58822 (NY, QCA, QCNE). NAPO: Canton El Chaco, Rio Quijos, Finca La Ave Brava, 1800-1900 m, 7-10 Sep 1990 (9 fl), Palacios 5344 (BG, F, NY, QCNE); Canton Archidona, Cord. de Guacamayos, nr. Sarayacu, 1500 m, 12 May 1990 (2 fl-fr), Palacios et al. 5092 (BG, F, NY, QCNE). SUCUMBtOS: Canton Gonzalo Pizarro, Reventador, 1800 m, 23 May 1990 (2 fl-fr), Cer6n et al. 9868 (QCNE). ZAMORA-CHINCHIPE: Podocarpus National Park, nr. El Tambo, 2210 m, 13 Jul 1993 (d'), Gentry 79922 (QCNE); trail from Loja-Zamora rd. to Rio Santiago, 4 May 1995 (? fl), Pedersen et al. 104206 (LOJA). PERU. AMAZONAS: Prov. Luya, Cerro Huiscocunga, between CamporredondoandTullanya,2350 m, 3 Sep 1989 (9 fl), C. Diaz et al. 3717 (BG, MO); Serraniade Bagua, ca. 12-18 km E of La Peca, 1800-1950 m, 14 Jun 1978 (? fl), Gentry et al. 22909 (AAU, U). HUANUCO: Prov. Pachitea, ca. 26-28 km ESE of PuertoInca, ca. 1750 m, 13 Sep 1988 (a), Wallnoefer113-13988 (BG); Cerrosdel Sira, RioYuyapichis, ca. 1290 m, 13 Jul 1969 (? fr), Wolfe12307 (F, NA). BOLIVIA. LA PAZ: Prov. Franz Tamayo, 44 km SW of Apolo, rd. Calabatea-Correo, 1650 m, 9 Jun 1990 (e), Gentryet al. 71033 (BG);Prov.NorYungas,SerraniadeBella Vista, 16 km N of Carrasco,37 km N of Caranavi,1500 m, (2 fr), Solomon et al. 12694 (BG, NY).

The absence of collections from the middle part of Ecuadorsuggests the possibility of a gap in the distribution,similar to that in Sorocea trophoides. A recent collection, made from a shrubof 1-2 m tall, shows general similarities to Helicostylis tovarensis, but is clearly distinct in the small laminas(4-7 x 0.7-1.2 cm) and in the very shortly pedunculate staminateinflorescences.It almostcertainlyrepresents a distinct taxon, probably at the subspecific level: PERU. SANMARTIN: Prov.Rioja,Dtto. PardoMiguel, between Aguas Verdesand rd. to San Pablo, 1220 m, 23 Jun 1998 (a), I. Sanchez V et al. 9456 (BG).

269 la. Castilla elastica Sess6 subsp. elastica. No changes to be reported.

lb. Castilla elastica Sesse subsp. costaricana (Liebmann)C. C. Berg. No changes to be reported.

lc. Castilla elastica Sesse subsp. gummifera (Miquel) C. C. Berg. No changes to be reported.

2. Castilla ulei Warburg (see Berg, Fl. Neotrop. Monogr. 7: 100-102. 1972). Distribution. In the AmazonBasin, most common in its upperpart,presentlyalso known from Ecuador; in non-inundatedforest; at low elevations. Representativespecimens examined. ECUADOR. NAPO: Reserva Biologica Jatun Sacha, 8 km E of Misahualli, 21 Oct 1989 (st), Palacios et al. 4679 (BG,

QAME,QCNE);ParqueNacionalYasuni,Pozo Petrolero Amo2, 9-13 Jan1988(st),Ceronet al. 3228 (BG,QAME, SanRoque,14 CantonShushufindi, QCNE).SUCUMBIOS: Dec 1997 (st), E. Freire et al. 3005 (QCNE);Reserva nr.LagunaGrande,15Mar1990(st), Faunistica Cuyabeno, Valenciaet al. 355 (QCA). BOLIVIA.PANDO:54 km SW of Cobija,Triunfo,4 Aug 1988 (a'), R. T Pennington et al. 82 (BG); Prov.

Manuripi,rd.Cobija-Chive,km 15, 29 Sep 1991(? fl-fr), Perry et al. 289 (BG, MO).

3. Castilla tunu Hemsley (see Berg, Fl. Neotrop. Monogr. 7: 102-104. 1972). No changes to be reported.

Naucleopsis Castilla 17. Castilla Sesse (see Berg, Fl. Neotrop. Monogr. 7: 92-104. 1972.) Recent botanical exploration has hardly affected the genus. Most additionalcollections have been made within the known ranges of distribution.Only the extension of the rangeof Castilla ulei to AmazonianEcuador is noteworthy. 1. Castilla elastica Sess6 (see Berg, Fl. Neotrop. Monogr. 7: 94-100. 1972.)

18. Naucleopsis Miquel (see Berg, Fl. Neotrop. Monogr. 7: 104-144. 1972). Additional synonym: UleodendronRauschert,Taxon31: 560. 1982, to replace AcanthosphaeraWarburg(1907), nom. illegit.,

alreadyput into the synonymyof Naucleopsisby Ducke(1922). Furtherbotanicalexplorationhas affectedthis genus more than the other revised genera, in particular by new recordsfrom CentralAmerica(CostaRica and Panama),fromthe Pacific Coastalregion of Colombia andEcuador,andtheNaporegion.Fivenew speciesand

FLORANEOTROPICA

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flowers twonewsubspecieshavebeendescribed. Naucleopsis lost.Thismayalsoexplainwhythestaminate amarahadto bereducedto a subspeciesofN. ulei,and areenclosedby the relativelylargeinnerinvolucral thenameof oneof thespecieshadto be changedfrom bracts.Inothercases,asin CastillaandPseudolmedia, closeduntilanthesisis correlated N. mello-barretoi to N. oblongifolia due to rules of havinginflorescences of theperianth. priority.Naucleopsisulei subsp.subandinais described withreduction in thepresentmonograph. However,betweentheperipheral scale-likestrucInmorerecentdescriptionsI haveabandoned inflorescence, whichcanberegarded the turesofthepistillate term"pseudobracts" forall,mostlyaculeate, whichcan processes astrueinvolucralbracts,andthestructures andused beregardedas freepartsof thetepals,oneoftenfinds ontheupperpartof thepistillateinflorescence instead"freepartsoftheperianth." Insomeof thespecies aculeateprocesseswhichdo notclearlybelongto the withthestyle involucreorto theflowers. allthesefreepartsaredistinctly associated Withinthegenusa numberof speciesarecharacterandthuscanbe easilyrecognizedaspartsof theperithatpeelsoff in smallflakes.The of tepalsis oftenmorethan4, thebasic izedbytheperiderm anth;thenumber numberof thefamilyandtribe,andvariesfrom4 to 8 specieswiththisfeatureappearto represent a groupof distribution or 10.Evenwiththisincrease,a distinctarrangement relatedspecieswitha largelynorthwestern of freepartsof theperiantharoundstylescanstillbe in SouthAmerica(linkedwiththe northernAndean present,butsometimesoneora fewfreeperianth parts region):theupperAmazonBasin,thePacificCoastal occurdispersed amongtheflowers.Inmanyothercases regionof SouthAmerica,andCentralAmerica.This the freeperianthpartsareloosely arranged,andthe groupcomprisesthespeciesnumbered15-22. numberof thesepartsamongthestylesis oftenmore Latexof severalspeciesof Naucleopsisis usedto than10perflower(orstyle).Asinterfloral dartpoison.Theuseof latexof N. oblongifolia bractsseemto prepare belackingintheCastilleae,it is notlikelythatthepro- (= N. mello-barretoi)was reportedby Prance(1972a, cessesrepresent bracts,butratherthatfloralconfigura- 1972b)andBissetandHylands(1977) fortheMaku' tionsbecameobsolete,ashappened lesspronouncedly IndiansinAmazonas (Brazil)anddiscussedbyShrestha in thestaminate inflorescences of thegenus. et al. (1992)forthePacificCoastalregionof ColomThestaminateflowersof Naucleopsisoftenhave bia. The local namespalo venono, veneno, venenillo, morethanthebasicnumberof 4 tepalsandoftenless venenu teiug, andvenueo, used for N. chiguila and/or thanthebasicnumberof 4 stamens,whichareconse- N. naga subspp.naga andmeridionalisin the northern quentlyoftennotinthenormalantitepalous positionof partof thePacificCoastalregionof Ecuador, alsosugthe stamens,anda consistentfloralconfigurations is gestthatthesespeciesyieldpoison.

Revisedkey to the speciesof Naucleopsis 1. Laminadistinctlyhairy beneath;upperAmazonBasin. 2. Base of the laminaacuteto subacute......................................................................... 1. N. macrophylla 2. Base of laminaobtuseto cordate. 3. Stipulesca. 1.5 cm long; free partsof the perianthof the pistillateflower scale-like.......2. N. riparia 3. Stipules0.5-1 cm long; free partsof the perianthof the pistillatefloweraculeate.........3. N. velutina 1. Laminaglabrousbeneath,or if hairy,then only on the midrib,or the whole surfacescabridulous. 4. Leafy twigs with partswith distinctlyshorterinternodes,in these partsthe nodes bearingonly (subpersistent)stipules. 5. Treeletsto 3 m tall; stipules0.5-1.5 cm long; staminateinflorescences0.2-0.3 cm diam.; free partsof tepalsof pistillateflowersto 0.7 cm long in fruit;upperAmazonBasin........20. N. humilis 5. Trees to 25(-40) m tall; stipules0.8-2.5(-3.5) cm long; staminateinflorescences(0.2-)0.51.5 cm diam.;freepartsof tepalslongerthan 1 cm (to 5 cm long) in fruit;AmazonBasinand W Ecuadorto CostaRica................................................................ 19. N. ulei 4. Leafy twigs without parts with distinctly shorterinternodes,the nodes (normally)bearing normal leaves. 6. Peridermoften ? wrinkled,peeling off in small flakes. 7. Marginof both laminaand stipules(at least partly)revolute,but the base of the lamina 15. N. herrerensis involute;upperAmazonBasin............................................................. 7. Marginof laminaplane or revolute,the stipules not revolutenor the base of of the lamina involute. 8. Stipulesto ca. 1 cm long; tertiaryvenationreticulate. 9. Lateralveins 15-23 pairs;involucralbracts? denselyhairy;AmazonBasin... 21. N. krukovii 9. Lateralveins 8-17 pairs;involucralbractssparselyhairyto (sub)glabrous; Pacific Coastalregion and CentralAmerica.

TAXONOMICTREATMENT

271

10. Laminacoriaceous,the base roundedto obtuse,the apex acuminateto sub......... 17. N. capirensis caudate,the marginusuallyrevolute;Costa Rica to W Ecuador 10. Laminasubcoriaceous,the base (sub)acute,the apex (sub)caudate,the marginplane; W Colombiato Panama................................................... 22. N. straminea 8. Stipules(usually)longer than 1 cm; tertiaryvenation(usually)partlyscalariform. 11. Stipules 1.5-2.5 cm long; petiole 0.5-1 cm long; W Ecuador ...........................16. Nl. chiguila 11. Stipules0.8-1.5 cm long; petiole 0.8-2.2 cm long; Hondurasto W Ecuador ....... 18. N. naga 6. Peridermmostly smooth,or if ? wrinkled,then persistent. 12. Laminaca. 50-75 cm long; stipules ca. 1.5-3.5 cm long, subpersistent;middle Amazon Basin............................................................... 14. N. stipularis 12. Laminashorterthan 50 cm, or if longer(to ca. 55 cm long), then the stipulescaducous. 13. Leafy twigs + densely hairy,tomentoseto velutinous. 14. Laminadistinctly inequilateral. 15. Stipules (usually)subpersistent;peduncleof staminateinflorescence0.2-0.6 cm long; only stigmasvisible at anthesis;free partsof tepals aculeatein fruit;upper AmazonBasin ................................................. 12. N. inaequalis 15. Stipulescaducous;peduncleof staminateinflorescence0.5-1.6 cm long; both stigmasandtepalsvisible at anthesis;free partsof tepalscushion-shaped to pyramidate(to aculeate)in fruit;Guianasand Brazil (Amapa)....................... 11. N. guianensis 14. Laminaequilateralor slighty inequilateral. 16. Petiole 1-1.5(-2) mm thick. 17. Pistillateinflorescencewith 1-4 flowers;peduncleof staminateinflorescence 0.1-0.2 cm long; AmazonBasin. 18. Involucralbractsof staminateand pistillateinflorescencesand the free parts of the perianth of the pistillate flower yellowish-velutinous .... 13. N. concinna

18.Involucralbractsof staminateand pistillateinflorescencesand the free partsof the perianthof the pistillateflower sparselywhitish-puberulous ..................................................................................................... 9. N.

ternstroemiiflora

17. Pistillateinflorescenceswith 5-20 flowers;peduncleof staminateinflorescence 0.5-1.2 cm long; Guianasand Brazil (Amapa) 11. N. guianensis ........... 16. Petiole 2-4 mm thick. 19. Laminawith sparseyellowishhairson the midrib(and lateralveins) beneath; southwesternAmazon Basin.............................................. 5. N. jamariensis 19. Laminaglabrous,subglabrousor minutelyhispidulousbeneath. 20. Lamina(usually)scabridulousbeneath;free partsof the perianthsof pistillateflowers aculeate;involucralbractsof staminate'inflorescences sparselyhairy;lower AmazonBasin........................................... 4. N. caloneura 20. Laminasmoothbeneath;free partsof the perianthsof pistillateflowers cushion-shaped;involucralbractsdensely hairy;upper AmazonBasin ................................................................................................................ 13 .N. concinna

13. Leafy twigs glabrousor sparselyhairy. 21. Laminausually shorterthan 20 cm; leafy twigs mostly thinnerthan 4 mm. 22. Lateralveins in the middleof the laminastraightor nearlyso and arched2-4 mm from the margin;Guianasand Brazil (Amapa)................................. 12. N. guianensis 22. Lateralveins in the middleof the laminacurved,or if (almost)straight,then arched 1-2 mm from the margin. 23. Petiolesand leafy twigs dryingblackish;pistillateinflorescenceswith 1-4 (-5) flowerswith cushion-shapedfree partsof the tepals; staminateinflorescences 0.4-0.8 cm diam., the peduncle0.1-0.4 cm long; upperAmazon Basin and E Brazil.............................. 10. N. oblongifolia 23. Petioles and leafy twigs dryingbrownishor yellowish;pistillateinflorescences with at least 5 flowersand free partsof the perianthaculeate;staminate inflorescencesusually more than 0.8 cm diam. and/orthe peduncle longer than 0.4 cm. 24. Stipulesusuallylongerthan 1 cm, stiff and pointedwhen dry; northem 6. N. glabra South America........................... 24. Stipulesshorterthan 1 cm, not stiff and pointedwhen dry;upperAmazon 8. N. pseudonaga Basin........................... 21. Laminausuallylongerthan20 cm long; leafy twigs mostly at least 4 mm thick. 25. Pistillateinflorescenceswith 1-4 flowers;peduncleof staminateinflorescence 9. N. ternstroemiiflora 0.1-0.2 cm long; upperAmazonBasin................................. 25. Pistillateinflorescenceswith more than 4 flowers;peduncleof staminateinflorescence longer than 0.2 cm.

272

FLORANEOTROPICA 26. Stipulesto ca. 1 cm long; lateralveins often not distinctlyloop-connected; 8. N. pseudonaga upperAmazonBasin................................................ 26. Stipulesusually 1-3 cm long; lateralveins distinctlyloop-connected. 27. Stipules 1.5-3 cm long; leafy twigs 4-12 mm thick;pistillateinflorescences 1.5-3.5 cm diam. (at anthesis);upperAmazonBasin .............7. N. imitans 27. Stipules0.8-2 cm long; leafy twigs 2-4.5 mm thick;pistillateinflorescences 0.7-1.8 cm diam.(at anthesis);mainlylower and middleAmazon basin or widespread. 28. Leafy twigs denselybrownish-puberulous and the stipulesbrownish to yellowish-subsericeous; involucralbracts? densely hairy;margin of the laminaoften revoluteand lateralveins often straight ....... 4. N. caloneura 28. Leafy twigs and stipules(sub)glabrous,or if hairy,then whitishor pale yellow; involucralbractssparselyhairy;marginof the lamina usuallyplane and the lateralveins usuallycurved..........................6. N. glabra

1. Naucleopsis macrophylla Miquel (see Berg, Fl. Neotrop. Monogr. 7: 113-114. 1972).

Tree,to 25 m tall.Leafytwigs 3-4 mm thick,brown-

(sub)velutinous. Laminaoblong,22-40 x 8-14 cm, coriaceous;apexacuminate; baseobtuseto rounded; Distribution. In the central part of the Amazon uppersurfacesparsely onthemidrib;lower puberulous Basin, in Brazil and Bolivia; in non-inundatedforest. surfacebrownish-(sub)velutinous on theveins;lateral venationlargelyscalariform; Representative additional specimens examined. veins18-26pairs;tertiary BRAZIL. ACRE:Mun. Cruzeiro do Sul, Rio Jurua, petiole0.5-1 cmlong,brown-velutinous; stipules0.5IgarapeViseu, 5 Nov 1991 (? fl), Ferreiraet al. 10610 1 cm long,denselybrownishto whitish-hirtellous to (BG). MATOGROSSO: Rio Aripuana,below Sato dos Dardanelos,nr.HumboldtCenter,5 Oct 1973 (d), Berg Mun.P6rtoVelho, et al. P18534 (MO,NY). ROND6NIA: Vila Caneco-MineraqioJacunda,26 Oct 1979 (? fl-fr), Vieiraet al. 289 (MICH,NY). BOLIVIA.BENI:Rio Beni, CachuelaEsperanza,Jan 1924 (d), Meyer255 (MO, NY, U). PANDO:Rio Madeira, opposite Penha Colorada,20 Nov 1968 (9 fr), Prance et al. 8704 (NY).

2. Naucleopsis riparia C. C. Berg (see Berg, Fl. Neotrop. Monogr.7: 114-116. 1972). Additionto the description: Pistillate inflorescencessometimeson shortleafless branchlets. Distribution. In the upperAmazon Basin, in Brazil (Amazonas andAcre) and Peru (Loreto); in riverside andterrafirme forest. Additionalspecimensexamined.Brazil.ACRE: Santa Lucia,40 km E of Cruzeirodo Sul, 14 Oct 1987 (9 flfr), Pruskiet al. 3476 (BG, MO, NY). AMAZONAS: Rio Jutai,8 Nov 1975 (9 fl-fr),L. Coelho et al. 401 (U). PERU. LORETO: Varaderode Mazan from Rio Amazonasto Rio Napo,21 Aug 1972(? fr), Croat19381 (NY); Prov.Maynas,Rio Napo, QuebradaSucusari,30 May 1991 (Q fr), Gentryet al. 74328 (MO).

3. Naucleopsis velutina C. C. Berg, Novon 6: 241. 1996. Type. Peru. Loreto: Prov. Maynas, Iquitos, Allpahuayo,EstacionExperimentalde IIAP,23 Aug 1990 (d), R. Vasquez etal. 14263 (holotype, MO).

subvelutinous.Staminateinflorescences 2-5 together

intheleafaxilsandbelowtheleaves,0.8-1.3cmdiam.; peduncle0.3-0.5 cmlong,bracteate; involucral bracts in ca. 7 series,broadlyovateto semicircular, densely to subvelutinous; yellowishto brownish-puberulous perianthca. 2.5-3 mmlong,tepals4-6, basallyconnate(orfree),? denselyyellow-hairy. Infructescences 6-8 cm diam.,(sub)sessile;involucralbractsin 6-8 series, ovate to lanceolate, yellow-strigose to theinneronesto2 cmlong;flowers/fruits subsericeous, numerous; freepartsof thetepalsdispersed,aculeate, yellow-strigoseto subsericeous; styleca.4 mmlong; stigmas2-3 mmlong,tongue-shaped. Distribution (see Fig. 83.5). Only known from AmazonianPeru(Loreto);in non-inundatedforest.

Specimens examined. PERU.LORETO: Prov.Maynas, Quebrada Yanomono,Rio Amazonas,abovemouthof Rio Napo,5 Nov 1979(? fr), Gentryet al. 27466 (BG,MO).

Thisnewspeciesis relatedto NaucleopsismacrophyllaandN. riparia.It differsfrombothin theshort stipules.Thepistillateinflorescences areclearlydifferentin theabsenceof distinctperianths, theslenderaculeate,yellow-subsericeous, looselyarranged freeparts of the tepals, and the short stigmas.

4. Naucleopsis caloneura (Huber)Ducke (see Berg, Fl. Neotrop. Monogr. 7: 116-119. 1972). Distribution. In the lower Amazon Basin, in Brazil, fromMaranhaoandParatothe Manausregion,also in Acre and MatoGrosso, and in Bolivia (SantaCruz);

TAXONOMICTREATMENT in non-inundatedforest;regionally (as in the Manaus region) very common.

273 Representative additional specimens examined. COLOMBIA. AMAZONAS:Rio Caqueta,nr. mouthof

Quinche,5 May 1988(st), M. SdnchezS. et al. Quebrada 198 (BG). ANTIOQUIA: Mun.Guatape,nr. borderMun. SanRafael,1890m, 1 Oct 1982(? fr),Albertde Escobar Acre, km 33, 12 Oct 1982, Ferreira et al. 2872 (MO, et al. 2388 (BG, HUA); Segovia, 21 Jul 1979 (? fr), NY). MARANHAO: Rd. Belem-Brasilia, kmn179, Oliveira Renteriaet al. 1717 (COL,MO);Mun.Anori,Rio Anori 753 (IAN). MATOGRosso: Rio Aripuana, below Salto valley, nr. Providencia, 24-31 May 1973 (2 fl-fr), dos Dardanelos,nr. HumboldtCenter,14 Oct 1973 (a), Soejartoet al. 4070 (A, HUA,MO,NY). BOLtVAR: Mun. Berg et al. P18518 (MO, NY). PAR,k: Mun. Oriximina, Achi, QuebradaLa Culebra, 21 May 1983 (? fr), SerraSacazinho,10Nov 1987(d' fl), Ferreira 9506 (BG, Cuadros V 1559 (COL); Guimari, 5 May 1949 (v), BR.163, km 1004, 14 MO, NY); rd. Cuiaba-Santarem, Sneidern5807 (S), (e), Sneidern5808 (COL,S). CHOC6: Nov 1977 (? fr), Prance et al. 25386 (NY). Mun.Riosucio,Cerrosdel Cuchillo,rd.Cidon-LaCumbre BOLIVIA. SANTACRUZ:Prov.Velasco, Lago Caiman, Sureste,23 Jun 1988 (2 fr), Cdrdenas2108 (MO);2-4 22 Oct 1996 (9 fl-fr), Painter 62 (BG). km NW of Teresita,18 May 1967 (a' fl-fr),Duke 11043 A scabridulouslower surfaceof the laminais com- (MO,NY, US). META: Rd. PuertoLosada-LasMarimmon in materialcollected nearManaus,less common bas, Vrda.PuertoLosada,25 Feb 1988(a'),Callejaset al. 4 kmW of LaMacain materialfromPara.ThematerialfromPari also tends 5918 (BG,MO);Mun.LaMacarena, rena, 10 Aug 1988 (? fl-fr), Callejas 7032 (BG, MO). to have smaller leaves than that from the ManausreSANTANDER:CampoCapote(CarareOpon),2 May 1977 gion. The small-leavedform,in which the pistillatein(2 fl-fr),Renteria460 (COL,MO); 15 km W of Puerto florescencesoften have few flowers, can be easily con- Berrio,13 May 1949(? fl-fr),Scolniket al. 19005 (US). fused with Naucleopsis ternstroemiiflora,but can be ECUADOR.NAPO: San Pablo de los Secoyas, 29 distinguishedfromthe latterby the dense indumentum Aug 1981 (d), Brandbygeet al. 36237 (AAU,MO, NY, of the involucralbractsandthe petiole, mostly longer QCA, U); Estaci6n Exp. INIAP-Payamino, 5 km N of than 1 cm. Coca, 18-26 Feb 1986 (d), Zaruma 442 (BG, MO, QAME);ParqueNacionalYasuni,Pozo PetroleroAmo 2, 9-13 Jan 1988(e), Cer6net al. 3223 (AAU,BG, MO, 5. Naucleopsis jamariensis C. C. Berg (see Berg, Fl. NY, QAME,QCNE).PASTAZA: Rio Curaray,nr.mouth of Rio Querano,3 Sep 1985(? fr),Neill et al. 6829 (BG, Neotrop. Monogr. 7: 119. 1972). MO, NY, QAME,QCA, QCNE).SUCUMBtOS: Reserva Completionof the description: FaunisticaCuyabeno,nr. LagunaGrande,11 Mar 1990 Staminateinflorescencesup to 5 togetherin the leaf (2 fl-fr),Balslevet al. 97441 (AAU,BG, QCA,QCNE). PERU. AMAZONAS: Rio Santiago,ca. 65 km N of axils; peduncle 0.5-0.8 cm long; involucralbractsin 4-5 series, ovate, obtuse, the outer ones pale yellow Pinglo, 24 Mar 1980 (? fl-fr), Huashikat2350 (MO, the innerones only ciliolate;pe- U). HUANUCO:Prov. Pachitea, Dtto. Honoria, rd. to appressed-puberulous, rianthca. 1 mm long; tepals4-5, cucullate,thickenedat Ayamiria,3 km fromMiel de Abeja,20 Jan 1967 (9 fr), SchunkeV 1548 (MOL,U). LORETO: Prov. Requena, the apex, ciliolate; stamens3-5; filamentsca. 0.5 mm Dtto. Puinahua,CochaYarina, 11 Jul 1985 (2 fl-fr), long; antherselliptic, 0.8-1 x 0.7-0.8 mm. Grandezet al. 498 (MO);Prov.Maynas,Dtto. Iquitos, Distribution. In the southwesternpartof the Ama- Rio Mom6n,San Andres,24 Mar 1977 (a), RimachiY Isla Bafnos,29 zon Basin, in Rondonia and Acre; in non-inundated 2921 (NY); Prov.Ucayali,Canchahuayo, MADRE DE et 6999 R. al. Nov 1985 Vasquez (a'), (NY). forest;apparentlya rarespecies. Dios: 15 km ENE of PuertoMaldonado,18 Dec 1989 Additional specimen examined. BRAZIL. ACRE: (? fl-fr), Gentryet al. 68952 (BG); Prov.Tambopata, F.E.A.,nr. Coloca9aoApui, 29 Nov 1991 (e), Sothers Dtto. Las Piedras,AlbergueCuzco Amazonico,17 Jan et al. 66 (NY). 1991 (? fr), Timanaet al. 1245 (BG, MO). UCAYALI: SEof Pucallpa,nr.jct. of RioPachiteaandRioYuyapichis, 7 Oct 1985 (? fl), Morawetzet al. 116-71085(BG);nr. 6. Naucleopsis glabra Spruce ex Pittier,Contr.U.S. Pucallpa, 19 Nov 1965 (a'), SchunkeV 950 (U). BRAZIL. PARA: Rd. Altamira-Itataituba,29 Oct Natl. Herb. 13: 440. 1912. (see Berg, Fl. Neotrop. 1977 (d), Berg et al. BG.754 (NY). Monogr.7: 119-123. 1972). BOLIVIA.BENI: Rio Beni, CachuelaEsperanza,Oct of the Coderequiredthatthe au- 1924 (i), Meyer267 (NY,U); Prov.VacaDiez, 20.8 km Strictinterpretation thorshiphas to be assignedto Pittierandnot to Baillon. E of Riberalta,25 Sep 1981 (a'), Solomon 6444 MO, (NY). PANDO: Prov.Abuna,Rio Negro,nr.jct. with Rio Distribution. In theupperAmazonBasin,fromCo- Abuna,4 Jul 1992 (st), Gentryet at. 77803 (MO);Prov. lombiato Bolivia,disjunctlyalsoin Surinameandnorth- Nicolas Suarez, SW of Cobija, Rio Naraueda,2 Aug forest;atlow elevations. 1982 (e), Sperlinget al. 6471 (BG, NY). ernColombia;in non-inundated Representative additional specimens examined. BRAZIL. ACRE: Mun.RioBranco,rd.Rio Branco-Prto

274 Just as in Naucleopsis imitans, in particularif the petiole is relativelylong (to 3 cm), theupperandlower partof thepetioleare? different:thelowerpartis thicker andin drymaterialdarkbrownandthe surfaceslightly fissured;and the upperpartis thinnerand pale brown and smooth when dry. Materialof Naucleopsis caloneura with a smooth lower surface of the lamina can be confused with N. glabra specimens, which have hairy leafy twigs and stipules. The two taxa can be usually distinguishedby the lateralveins, (almost) straightin the formerand ? curvedin the latter,andby the color of the indumentum of the twigs and stipules,brown(ish)in the formerand pale yellow to whitish in the latter.

7. Naucleopsis imitans (Ducke) C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 123-125. 1972). Distribution. In the upperAmazonBasin (Colombia, Peru,Brazil,Ecuador);in non-inundatedforest;at low elevations. Representative additional specimens examined. COLOMBIA. AMAZONAS: Rio Caqueta, nr. Villa Azul, 2 Nov 1989 (2 fl), Andel et al. 384 (BG); Rio Cahuinari, 13 km W of its mouth, 15 Sep 1988 (st), M. Sdnchez S. et al. 1252 (BG). VAUP9S:Estaci6n Biol6gica Caparu, 3 km N of Lago Taraira, 25 Mar 1988 (st), Defler 325 (MO), (6'), Defler 328 (MO). ECUADOR. NAPO:Huashito, 20 km N of Coca, 321 Nov 1989 (%), Rubio 377 (BG, QCNE); Aiiangu, Parque Nacional Yasuni, 1-30 Apr 1985 (9 fl), Korning et al. 58645 (AAU, BG, NY, QCA); Reserva Biologica Jatun Sacha, 8 km E of Misahualli, 1-15 Sep 1987 (? fl-fr), Palacios 1995 (BG, NY, QCA, QCNE). SUCUMBiOS: Sacha Lodge, 3 km NW of Ainangu, 5-12 Jun 1995 (V), J. L. Clark et al. 1162 (QCNE); Dureno, 27 Dec 1988 (st), Gentry et al. 64303 (BG). PERU. LORETO:Prov. Maynas, Yanamono, between Indiana and mouth of Rio Napo, 27 Jul 1980 (9 fr), Gentry et al. 29132 (NY); Mishayacu, nr. Iquitos, OctNov 1929 (2 fl), Klug 257 (F, NY, US, in Monograph 7: 144, under unnamed collections). MADREDE DIos: Nr. confluence of Rio Tambopata and Rio La Torre, 39 km SW of Puerto Maldonado, 16 Jan 1989 (? fr), S. F

FLORANEOTROPICA

Itis notquitecertainwhetherNaucleopsisimitans canbemaintained asa distinctspecies;thedifferences betweenit andN. glabraarelargelyquantitative, N. imitansbeingin all partsmorerobustthanN. glabra. 8. Naucleopsispseudonaga(Mildbraed) C. C. Berg (see Berg,Fl. Neotrop.Monogr.7: 125. 1972). Completion of thedescription: Infructescences subglobose,ca. 14cmdiam.when dry,to 20 cmdiam.whenfresh;involucralbractsto 2 cm long; free partsof the tepalsspine-liketo subpyramidate,5-8 mm long, densely puberulousto subhispidulous; fruitsca. 3.5 x 2.5 cm. IntheupperAmazon Distribution. Basin,fiomLoreto (Peru)to Pando(Bolivia)andAcre(Brazil),andapparently inVaup6s(Colombia), disjunctly knownfrom asinglerecord; innon-inundated forest;atlowelevations. Additional specimens examined. COLOMBIA. VAUPtS:Estaci6nBiologica Caparu, 3 km N of Lago Tataira,22 Feb 1990 (? fl), Defler 320 (MO).

PERU.Cuzco:Prov.Cuzco,Dtto.Camisea, CampamentoArmihuari,28 Jan 1997 (? fr), AcevedoR. et al. 9248 (BG, US). HUANUCO: Prov. Puerto Inca, Dtto. Yuyapichis,ReservaForestalDANTAS,1-14 Sep 1990 (a'), Tello 201 (MO, NY) and 353 (MO).LORETO:Prov. Maynas,trailbetweenIndianaand Mazan,4 Mar 1979 (? fl + fr), Gentry et al. 25388 (MO, NY, U); Prov.

Maynas,nr. Indiana,14 Feb 1989 (? fr), Gentryet al. 65771 (BG, MO). MADREDE DIOS:Parque Nacional del Manu,Rio Manfi,CochaCashuStation,20 Oct 1979 (? fl-fr),Gentryet al. 27045 (F, NY, U), 29 Jan 1977 (? fr), Fosteret a. 6033 (F), 11 Oct 1980 (a'), Foster5521 (F), 23 Aug 1984 (ao), Foster 9886 (BG, MO). BRAZIL. ACRE:Mun. Sena Madureira,Rio Macaua, Seringal Joaci, 22 Sep 1994 (a'), Daly et al. 8329 (NY);

Mun. Cruzeirodo Sul, Rio Tejo, SeringalIracema,12 Mar 1992 (? fr), Ferreiraet al. 10799 (BG, NY); Rio Muru,6 km aboveconfluencewith Rio Tarauaca,16 Sep 1968 (e), Prance et al. 7285 (NY, U). BOLIVIA. PANDO:Prov. Nicolas Suarez, Cobija, 20

Jun 1978 (? fl), Hartshorn713 (NY).

Thelateralveinsareloop-connected ratherfar(2-4 mm)fromthemargin,andtendto be faintlyorhardly Smithet al. 1302 (NY). loop-connected evenin theupperpartof thelamina. BRAZIL. ACRE: Rio Acre, ca. 30 km below Rio Thisfeaturein combination withtheshort(usuallyto Branco, 19 Oct 1980 (? fl), Nelson 744 (BG). AMAZONAS: ca. 1cmlong)stipules helpstoseparate thisspeciesfrom Mun. Fonte Boa, Parana do Mamupina, 3 Nov 1986 (d? otherspecieswithsmallto medium-sized leaves.The fr), Ferreira et al. 8391 (BG); Rio Pur'us, nr. Boca do collectionfromColombiadifferssomewhat intherelaAcre, 20 Sep 1966 (6'), Prance et al. 2426 (INPA, MICH, tively largeleaves andthe stipules,beingdistinctly NY); Rio Javari, 7 hrs. above Paumari, 16 Oct 1976 (6'), longerthan1 cm(toca. 1.5cm long)andstiff,resemPrance et al. 23836 (MICH, NY, U). blingthoseof Naucleopsisglabra.

The petiole is often differentiatedinto a lowerpart, which is thickerandin drymaterialdark-brownwith a ? crackedsurface,and an upperpart,which is thinner and in dry materialpale brown and smooth.

9.Naucleopsis ternstroemiiflora C.C.Berg (Mildbraed) (seeBerg,Fl.Neotrop.Monogr.7: 125-128.1972).

TAXONOMIC TREATMENT

275

VENEZUELA.AMAZONAS: Nr. San Carlosde Rio Distribution. In the upper and middle Amazon Basin, fromEcuadorto Para(Brazil);in non-inundated Negro, H. L. Clarket al. 7361 (MO). ECUADOR. SUCUMBIOS: Rd. Lago Agrio-Tarapoa, forest;at low elevations. km 27, 2 Aug 1975(6), Littleet al. 29 (QAME,QCNE); Representative additional specimens examined. ReservaFaunisticaCuyabeno,nr. LagunaGrande,1 Apr BRAZIL. AMAZONAS:Mun. Manaus,90 km NNE of 1988-1Apr1990(o), Valenciaet al. 67331(AAU,QCA). Manaus, 13 Nov 1989 (9 fr),Camposet al. 21 (BG,NY); NAPO:Rd. Hollin-Loreto,Rio Huataraco,800-1000 m, rd. Manaus-P6rtoVelho,BR.319,km 510, 17 Oct 1974 Ceronet al. 7485(QAME);ParqueNacionalYasuni,Pozo (? fr), Prance et al. 23002 (NY, U). MATOGROSSO: PetroleroDaimi2, Cer6net al. 4135 (QAME).PASTAZA: Rio Aripuana, nr. Humboldt Center, 25 Oct 1973 (d'), Rd. Coca-Auca,km 115, nr. Rio Tigiiino,4 May 1989 Berg et al. P.19881 (MICH,MO); Mun.Alta Floresta, ($?), Neill et al. 8994 (BG, MO, QAME). 17.5 km N of Alta Paraiso,23 Sep 1985 (a), Thomaset PERU. LORETO: Prov. Maynas, Dtto. Iquitos, al. 4119 (BG,NY). PARA:Rd.Santarmr-Cuiabi, BR.163, Mishana,3 Dec 1976 (? fl-fr), Davidson5218 (MICH, km 113, 15 Nov 1977 (e), Prance et al. 25430 (BG, NY); NY, U); Prov.Maynas,PuertoAlmendra,10 Feb 1977 Rio Xingu,GlabaBacaja,belowmouthof Rio Bacaja,22 (e), Revilla2304 (MO,NY, U); Prov.Requena,Jenaro Nov 1980 (9 fl-fr), Prance et al. 26398 (NY). Herrera,10 Dec 1980 (q fr),R. Vdsquez1041 (MO,NY, ROND6NIA:Mun. P6rtoVelho, BR.164, km 164, 7 Oct U). UCAYALI:Prov. Coronel Portillo,Yarinacocha,31 1979(? fr), Vieiraet al. 311 (MICH,MO,NY).RORAIMA: Mar 1988 (q fl-fr), R. Vasquezet al. 10446 (BG, MO). BRAZIL. ACRE:FazendaBom Sossego, between Mun. Novo Airao, Rio Camanau,nr. Aldeia Mare, 31 Igarapedo Cujubimand IgarapeJacamin,27 Sep-7 Oct Aug 1989 (st), Milliken577 (MO). 1985(q fl-fr),Campbellet al. 9328 (BG);Mun.Cruzeiro Fertilematerialof this species can be easily distin- do Sul, Igarap6Humaiti, 22 Mar 1992 ($ fr), Daly et guished from Naucleopsis concinna by the sparse al. 7572 (BG, NY). AMAZONAS: Mun.Novo Airao,Rio indumentumof the involucralbractsandthe aculeate Camanaii,nr.AldeiaMare,2 Sep 1989 (a'), R. Milleret free partsof the perianthof the pistillateflower.Sterile al. 626 (NY); Rio Negro Basin, Rio Uneiuxi,MakuiInmaterialof both species can hardlyor not be separated. dianvillage, 23 Oct 1971 (6), Pranceet al. 15563 (NY, alsoresemblesthe small- U). BAHIA:Itaibo,rd. to Aquarema,29 Oct 1970 (ae), T Naucleopsisternstroemiiflora S. Santos1226 (U). ESPIRITOSANTO:Linhares, Reserva leaved formof N. caloneura (in Para),which often has Florestalde CVRD, 18 Sep 1979 (a"), Folli 118 (BG). few (down to 4) pistillateflowers in the pistillateinflo- MINAsGERAIS:Mun.Caratinga, FazendaMontesClaros, rescence. Nauacleopsis ternstroemiifloracan be dis- 8 Feb 1980 (R fr), Nishimura50 (GUA, U). tinguishedfromthelatterby theverysparseindunentum Sterilematerialof this species can be distinguished on the involucralbractsandthe relativelyshortpetiole, to 1 cm long. The stipules of N. ternstroemifloratend fromNaucleopsis ternstroemiifloraand small-leaved to be subpersistentandtheepidermisof thepetioletends specimens of N. concinna by the glabrousor sparsely puberulousleafy twigs, stipules,andpetioles, the latter to flake off. dryingblackish.Some specimensof N. oblongifolia(in Loreto,Peru)have dense indumentumon the perianth 10. Naucleopsis oblongifolia (Kuhlmann)Carauta, of the pistillate flowers, rathersimilar to that in N. Albertoa3(24): 262, t. 1-2. 1994. (see Fl. Neotrop. concinna. The numberof flowers of the pistillate inMonogr.7: 131-133. 1972, subN. mello-barretoi). florescence is usually up to three in the form with (sub)glabrousperianths,but often threeto four,rarely As the correct year of publication of Ogcodeia to five in theformwithdenselyhairyperianths;thenumoblongifolia by Kuhlmannwas established, namely ber of flowers is normallyat least five in N. concinna. 1939 insteadof 1940, the nameadoptedfor the species Use. The latex is used to prepare arrow poison in the monograph,Naucleopsis mello-barretoi,hadto (Prance, 1972a, 1972b); see also the generictreatment be replaced. of Naucleopsis, above. Distribution. In the northwesternpartof the Amazon Basin from Colombia(Caqueta)to Peru(Ucayali) to Acre andAmazonas(Brazil), also in easternBrazil; 11. Naucleopsis guianensis (Mildbraed)C. C. Berg in non-inundatedforest;at low elevations. (see Berg,Fl. Neotrop.Monogr.7: 130-131. 1972). Representative additional specimens examined. Distribution. In the three Guianas and Brazil AMAZONAS: Mun. Leticia, Parque (Amapi); in non-inundatedforest;at low elevations. Nacional NaturalAmacayacu, 12 Apr 1991 (9 fl-fr), Representative additional specimens examined. Pipoly et al. 15165 (BG); Rio Caqueta,nr.Araracuara, 26 Feb 1991 (? fl-fr), Vester269 (BG). CAQUETA: GUYANA.Maburaregion, Kurupukarimain, km 27, Araracuara,23 Jan 1984 (st), Gentry et al. 64964 (MO). 29 Nov 1993 (q fr), Ek et al. 1027 (NY). COLOMBIA.

276 FRENCHGUIANA.Saul, La FumeeWest,31 Aug 1989 (d), Mori et al. 20757 (BG, NY); OyapockR., Armantabo Cr., 3 Jul 1969 (d), Oldeman B.2427 (CAY);PassouraCr.,30 Sep 1992 (2 fl-fr),Sabatieret al. 4074 (NY).

12. Naucleopsis inaequalis (Ducke) C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 128. 1972). Extension of the description: Infiuctescences 2-2.5 cm diam.; free parts of the tepals aculeate,to 0.3 cm long, minutelypuberulous.

FLORANEOTROPICA La Torreand Rio Tambopata, 3 Jun 1987 (d'), Gentryet al. 58140 (BG, MO);Prov.Tambopata,TambopataReserve,26 IknS of PuertoMaldonado,5 Nov 1979 (? fl), Hartshorn2408 (BG, MO, NY). PASCO:Palcazuivalley, 9?50'-10?45'S,68?00'-68?30'W,7 May 1985 (st), Hartshornet al. 3723 (MO).UCAYALI: SE of Pucallpa, nr.jct. of Rio Pachiteaand Rio Yuyapichis,7 Oct 1985 (e), Morawetzet al. 18-71085 (BG). BRAZIL. ROND6NIA:Mun. Santa Barbara, BR.364,

km 120, 25 May 1982 (a'), Teixeiraet al. 739 (BG, F, MO, NY). BOLIVIA. BENI:Rio Beni, Cachuela Esperanza, Sep

1923 (6'), Meyer178 (BG, U), Oct 1923 (?), Meyer347 (U). PANDO: Prov.Abuna,Rio Negro, nr.jct. with Rio Distribution. In the middle part of the Amazon Abuna,7 Jul 1992 (st), Gentryet al. 77908 and 77949 Basin, from the Manaus region to Pando (Bolivia); (MO);Rio Abufia,14 Nov 1968 (? fl-fr), Prance et al. mostly in inundatedforest(vdrzea),sometimeson terra 8421 (MO, NY). firme. The indumentumof the leafy twigs and stipules is Additional specimensexamined. BRAZIL.ACRE: almost similarto thatof Naucleopsis terstroemiiflora, Mun. Bujari,Rio Branco,Riozinhodo Andira,25 Mar butthe involucralbractsof both staminateandpistillate 1995 (? fl-fr), Pardoet al. 78 (NY); Mun. PortoAcre, inflorescences are densely hairy, whereas they are Rio Acre, ReservaFlorestalHumaita,2 Nov 1993 (a), sparselyhairy in N. ternstroemiiflora. M. Silveira et al. 700 (NY). RONDONIA:Nr. Alvarada do Oeste, 30 Apr 1987 (? fl-fr), Ferreira8955 (NY). BOLIVIA. PANDO:Rio Abuna, 14 Nov 1968 (o), 14. Naucleopsis stipularis Ducke (see Berg, Fl. Prance et al. 8411 (BG, NY). Neotrop. Monogr.7: 133-136. 1972). 13. Naucleopsis concinna (Standley) C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 133. 1972). Correctionof the description: Pistillate inflorescences with (4-)5-40 flowers; perianthyellow-velutinous. Distribution. In the upper Amazon Basin, from Ecuadorto Bolivia andto MatoGrosso(Brazil);in noninundatedforest;at low elevations. Representative additional specimens examined. COLOMBIA. AMAZONAS: Rio Caqueta,S of mouthof QuebradaBocaduche, 17 Aug 1944 (a), Dulmen256

(BG); Rio Caqueta, nr. Araracuara,5 Sep 1990 (st), Vesteret al. 53 (BG). VAUPES:Estaci6n Biol6gica Caparu,3 km N of Lago Taraira,1986-1990 (? fl-fr), Defler 319 (MO).

Additionto the description: Tree,to 12 m tall. Distribution. In the middle part of the Amazon Basin, from the Manaus region to eastern Para, and southwardsto Mato Grosso and Rondonia;in non-inundatedforest. Representative additional specimens examined. BRAZIL. AMAZONAS: Rd. Manaus-Itacoatiara, 9 Nov 1973 (st), Stewardet al. P.17649 (NY); rd. Manauskm 26, ReservaFlorestalDucke, 1 Feb 1996 Itacoatiara, (? fl-fr), Ribeiro et al. 1800 (INPA).MATOGROSSO: Rio Aripuana,nr. HumboldtCenter,20 Oct 1973 (st), Berg et al. P19807 (MO,NY). ROND6NIA: Mun.P6rto Velho,RepresaSamuel,7 Jun 1986 (st), Thomaset al. 4996 (NY); Mun. Ariquemes,Minera9aoMibrasa,20 May 1982 (? fr), Teixeiraet al. 622 (INPA,NY).

Puerto Morona, ECUADOR. MORONA-SANTIAGO:

E of Rio Morona,1 Oct 1975(ad),Littleet al. 522 (COL, 15. Naucleopsis herrerensis C. C. Berg, Novon 6: LOJA, MO, NY, QAME, QCNE, US). SUCUMBiOS: 237, t. 6. 1996. Type.Peru.Loreto:Prov.Requena, ReservaFaunisticaCuyabeno,nr. LagunaGrande,21 Jenaro Herrera,arboretum(tree 7/65), Aug-Sep Mar 1991 (st, juv), Berg 1644A (AAU, BG, QCA). 1976 (a), Bernardi 16201 (holotype, G; isotype, PERU. LORETO: Prov. Maynas, Rio Blanco, 3 BG). Fig. 68 (Johnson40 hp) hoursfromTamshiyacu,15 Mar 1978 (q fr), C. Diaz et al. 174 (MO);Boquer6nPadreAbad, Tree,to ca. 25 m. Leafy twigs 1.5-3.5 mm thick, 26 Mar 1962 (2 fr), Schunke V 5830 (F, MO); Prov. peridenn(in Maynas, Rio Nanay, Mishana,20 Jan 1985 (? fr), R. sparselyto densely appressed-puberulous; x 1.3off. Lamina 6.5-24 olderparts) flaking lanceolate, Visquezet al. 6146 (BG, MO). MADREDEDIos: Parque NacionalManu,Rio Sotileja,9 Oct 1986 (? fl), Foster 6 cm,coriaceous;apexacuminateto subacute,theacumen 11732 (F, NY); TambopataNatureReserve,jct. of Rio oftenacute;baseacuteto obtuse;marginpartly? revolute,

TAXONOMICTREATMENT

277

Fig. 68. Naucleopsisherrerensis.1. Leafy twig with staminateinflorescences.2. Tepaland stamen(Bernardi 16201). [By HendriekeBerg, Bergen.]

butalwaysinvoluteatthebase;bothsurfacesglabrous; Distribution (see Fig. 83.6). In the northwestern or AmazonBasin (Naporegion);in non-inundatedforest. venationreticulate lateralveins12-26pairs;tertiary petiole0.5-1.8cmlong,appressedpartlyscalariform; Specimens examined. COLOMBIA. CAQUETA: theepidermis often?flakingoff;stipules1- Morelia,4 Nov 1941 (9 juv), SneidernA.1283 (MICH, puberulous, Stami- NY); nr. Araraquara, (st), Torres D. et al. 1103 (NY). appressed-puberulous. 1.5cm long,(brownish) intheleafaxils, PUTUMAYO:Rio Putumayo, Puerto Asis, 14 Aug 1964 nateflornce esondistinctshort-shoots 1cmdian.;peduncle0.2-0.3 cmlong; (9 fl-fr), Vogelmann et al. 1302 (COL, U, US). upto6together,ca. ECUADOR.NAPO:ParqueNacionalYasuni,Maxus involucral bractsinca.5 series,broadlyovatetosemicir1.5- rd., km 40, 5 Jun 1994 (st), Aulestia2335 (QCNE),10 culartosuborbicular, appresed-puberulous;perianth 2 mmlong;tepals4-5, freeorbasallyconnate;stamens Sep 1994 (9 juv), Aulestia2754 (QCNE),14 Sep 1994 oblongtoellip- (st), Aulestiaet al. 2772 (QCNE),km 108, 11 Jan 1995 0.3-0.4mmlong;anthers 3-4;filaments tic, 0.6-0.8 x 0.3-0.4mm. Pistillate inflorescences(ju-

(9), Aulestia et al. 2991 (QCNE), km 52.5-52.7, 5 Sep 1993 (st), Dik 262 (QCNE),km 53-60, 6 Oct 1993 (st),

(immature!) venile!)belowtheleaves.Infructescences Dik 595 (QCNE);ParqueNacionalYasuni,Pozo Petrolero ca.2-7 cmdiam.,sessile;involucralbractsin ca. 6 se- Daimi 2, 26 May-8 Jun 1988 (d), Ceron et al. 4135 flowers (MO, QAME,QCNE);Cant6nArchidona,rd. Hollinries, broadlyto narrowlyovate,puberulous; to Loreto,Rio Huataraco,800-1000 m, 23-30 Aug 1989 tosubulate, freepartsof thetepalsaculeate numerous; I cm long, minutelypuberulousto subhispidulous; (st), Ceron et al. 7485 (BG, QAME, QCNE); Parque style5-6 mmlong;stigmassubulate,2-3 mmlong. NacionalYasuni,Pozo PetroleroAmo 2, 9-19 Jan 1988

278

FLORANEOTROPICA

(2 fl-fr), Neill et al. 8335 (BG, NY, QAME,QCNE); subsessile; involucral bractsin 7-8 series,broadlyovate ReservaFloristicaEl Chuncho,5 km N of Coca,23 May tonarrowly lanceolate, to 3 cmlong,acute,puberulous; 1993 (C), Palacios 10791 (QCNE). PASTAZA: Mission

Shandia,Rio Jatunyacu,17 Aug 1957 (? fl), Barclay 4982 (COL). SucuMBiOS:ReservaFaunisticaCuyabeno, nr.LagunaGrande,19-21 Mar1991(st), Bergs.n. (BG); Cant6nShushufindi, Pucapeila,4 Dec 1997(st),E. Freire et al. 2567 (QCNE);ReservaFaunisticaCuyabeno,nr. LagunaGrande,1 Apr 1988-1 Apr 1990 (st), Valencia et al. 67804 (AAU). PERU.AMAZONAS: Rio Cenepa,E of Huampami,19 Dec 1972 (v), Berlin 641 (AAU, MO, NY); upperRio Cenepa,21 Feb 1973 (? fr),Kayap394 (NY). LORETO: Prov.Requena,Dtto. Sapuena,JenaroHerrera,1971 (st), Flores la (BG);Prov.Maynas,Rio Nanay,Mishana,28 Feb 1979 (st), Gentryet al. 25244 (MO,U), 6 Jan 1983 (st), Gentryet al. 39201 (BG, MO);Prov.Maynas,Rio Nanay,Mishana,12-13 Jan 1983 (d' fl), Gentryet al. 39472 (BG, MO); Prov. Requena,JenaroHerrera,23 Feb 1987 (st), Gentryet al. 56423 (MO);Prov.Maynas, Rio Yavinete,tributaryof Rio Putumayo,Bellavista,15 Apr 1978 (st), Haxaire3865 (MO);Prov.Maynas,Dtto. Las Amazonas,nr. Sucusari,4 Mar 1991 (st), Pipoly et al. 14359 (MO);Prov.Requena,JenaroHerrera,25 Aug 1976 (6), Revilla 1185 (MO, NY, U); Prov. Maynas, Dtto. Las Amazonas,Rio Sucusari,7 Nov 1989 (?), R. Vasquezet al. 13092 (BG).

flowersnumerous; freepartsof thetepalssubulate, 0.20.5cmlong,puberulous; styleto 0.5 mmlong;stigmas filiform,8-10 mmlong. Distribution (see Fig. 83.5). Confinedto thePacific lowlandsof Ecuador;in wet forest;at low elevations. Additional specimens examined. ECUADOR. N of Lita,N of Rio MiraandE of Rio Baboso, CARcHI: 1 Jul 1994 (st), Boyle et al. 3293 (BG). ESMERALDAS: CantonSan Lorenzo,ReservaIndigenaAwa, Ricaurte, Balsareino, Rio Palabi,15-29 Apr 1991 (st), Rubioet al. 1409 (BG, QCNE);Canton Eloy Alfaro, Luis Vargas Torres,Rio Santiago,Playa de Oro, 1-2 Nov 1993 (2 fl), Tiradoet al. 247 (QCNE).Los Rios: Rio Palenque Biological Station,rd. Quevedo-SantoDomingode los Colorados,km 56, 2 Oct 1976 (st), Dodson et al. 6333 (AAU, MO, QCA, U), 11 Aug 1977 (st), Dodson et al. 6733 (MO),2 Oct 1976 (st), Dodson et al. 6471 (MO, QCA),13 Feb 1974(st), Gentry9886 (MO).PICHINCHA: ReservaFlorestalENDESA,rd. Quito-PuertoQuito,km 113, 10 km N of the rd., 19 May 1987 (a') P. Acevedo et al. 1721 (QCA,NY).

17. Naucleopsiscapirensis C. C. Berg, Novon 6:

This species can be recognized by the relatively 235, t. 5. 1996. Type. Panama.Panama:W of narrowlaminawith the margin(partly)? revolute,but Interamerican Hwy.,nr.Capira,ca. 60 km W of towardsthe base distinctly involute. It is possibly re1Mar1985 Panama City, on rd.to CerroCampana, lated to Naucleopsis ulei, resemblingin its vegetative (d'), McPherson 6913 (holotype, MO; isotypes, parts the small-leaved subsp. amara. However, it is BG, BM, CR,MEXU,PMA). Fig. 69 clearly differentin the shape and size of the staminate inflorescences, the absence of very short intemodes, Tree, to 15 m tall. Leafy twigs 2-3 mm thick, andthe distinctlyloop-connectedlower lateralveins of (sparsely)puberulous; peridermflakingoff. Lamina the lamina. oblong, (5-)8-15(-2 1) x (l.7-)3-5.5(-7.5) cm, coria16. Naucleopsis chiguila Benoist (see Berg, Fl. Neotrop. Monogr. 7: 136. 1972). Revised description,includingthe staminateinflorescences: Tree,to 25 m tall. Leafy twigs 2-4 mm thick, grayish, yellowish or brownishappressed-puberulous; peridermflaking off. Laminaoblong to lanceolate, 11-23

baseobtuseto ceous;apexacuminateto subcaudate; rounded;marginoften? revolute;uppersurfaceglabrous(orsparselypuberulousat thebaseof themidrib);lowersurfaceglabrousorsparselypuberulous on (thebaseof) themidrib;lateralveins 10-16 pairs;tertiaryvenationreticulate; petiole0.2-0.8 cm long,the epidermis flaking off; stipules 0.3-1 cm long, puberulous,caducous.Staminateinflorescencesin the

leaf axils andbelowthe leaves,ca. 1-1.5 cm diam.; x 4.5-8.5 cm,(sub)coriaceous; involucral base peduncle0.2-0.3 mmlong,notbracteate; apexacuminate; to lateralveins16-18 bractsin 4-5 series,broadlyovateto semicircular bothsurfacesglabrous; (sub)acute; theouteronesminutelypuberulous, the suborbicular, pairs;tertiaryvenationpartlyscalariform;petiole 0.51cmlong;stipules1.5-2.5cmlong,puberulous atthe innerones only ciliolate;perianthca. 2 mm long;tepals ontheupper minutely puberulous base, caducous. Staminateinflorescences 0.5-0.8 cm 2-7, often? cucullate, diam.;peduncle0.4-0.6 cm long, bracteatetowardsthe part;stamens3-5; filaments0.6-0.8 mmlong;anthers apex; involucral bracts in 4-5 series, broadly to nar- oblong to elliptic,0.7-1 x 0.3-0.4 mm. Pistillate inflorowly ovateto oblong,ciliolate;centerof thereceptacle rescences 1.5-3 cm diam., subsessile or peduncle to 1.2-1.4mmlong;tepals6-8, free,often 0.6 cm long; involucral bracts in 6-8 series, broadly conical; perianth somewhatcucullate,minutelypuberulous; stamens4- to narrowly ovate, minutely puberulous to submur6;filaments0.4-0.7 mmlong;antherselliptic,0.4-0.6 iculate, the innerones to 1.5 cm long; flowers numerx 0.4-0.5 mm. Pistillate inflorescencesca. 2 cm diam., ous; free parts of the tepals spine-like to pyramid-

TAXONOMICTREATMENT

279

0.5 mm Fig. 69. Naucleopsiscapirensis.1. Leafytwigs with staminateinflorescences(McPherson6931). 2. Leafytwigs with pistillateinflorescence.3. Infructescence(Haberet al. 1960). 4. Staminateflower and stamens(McPherson 6931). [By HendriekeBerg, Bergen.]

Distribution (see Fig. 84.2). From Costa Rica to ate,minutelypuberulous to submuriculate; style2-5 mm long; stigmasfiliform, 4-7 mm long. Infruc- the PacificCoastalregion of ColombiaandEcuador;in tescences(3-)4-8(-l0) cm diam.,free partsof the wet forest; to ca. 1800 m. tepalsto 1 cm long. Specimens examined. COSTA RICA. ALAJUELA:

280

FLORANEOTROPICA

(QCNE);ReservaIndigenaAwa,GualpiChicoarea,1330 m, 20 Jan 1988 (2 fl-fr),Hooveret al. 2827 (BG, MO, QCA);Cant6nTulcan, ReservaIndigenaAwa, Gualpi Medio,900 m, 21 May 1992 (?), Quelalet al. 576 (BG, QCNE);CantonTulcan, Gualpi Alto, 1800 m, 15-28 Jun 1991 (o), Rubioet al. 1723 (BG, QCNE),(? fl-fr), Rubioet al. 1746 (BG, QCNE);Cant6nTulcan,Reserva IndigenaAwa,Chical,23-27 1992 (? fl-fr), 7iipaz1074 (BG, QCNE);Cant6nTulcan,TobarDonoso, 650-1000 m, 19-28 Jun 1992 (o), YTpaz et al. 1309 (QCNE),(? Nr. fr), Jipazet al. 1362 (QCNE).CARCHI/ESMERALDAS: Lita,20 May 1987(? fl-fr),Werff9525 (BG,MO,QCNE) and9527 (AAU,BG, QCNE).ESMERALDAS: Cant6nSan Lorenzo,ReservaIndigenaAwa, Alto Tambo,22 Mar Rio San Lorenzo, 24 Mar 1991 (a), Alvarado 113 (MO); 1993 (? fl-fr),Aulestiaet al. 1499 (BG, QCNE);Canton ParqueNacional Guanacaste,Estaci6n Cacao, 4 Jun San Lorenzo,ReservaIndigenaAwa, Rio Mira, 10 km 1990 (? fl-fr), Barrelier 35 (MO); ParqueNacional W of AltoTambo,16-26 Mar1991(ae),Rubioet al. 1268 Guanacaste,Estaci6nPitila,4 Apr 1991 (oF),P Rios 353 (BG,QCNE);Cant6nSanLorenzo,ReservaIndigenaAwa, (MO). HEREDIA: Parque Nacional Braulio Carillo, Ricaurte,Balsareilo,Rio Palabi, 15-29 Apr 1992 (st), Estaci6nMagsay,16 Jun 1991 (st), Aguilar 167 (MO). Rubio et at. 1409 (BG). PUNTARENAS:1 mi due S of SanVitode Java,ca. 1150m, These specimens differfrom the CentralAmerican 18 Aug 1967 (a'), Raven 21897 (BG, F, MO, PMA). in thefreepartsof thetepalsbeingdistinctlybroadones PANAMA.DARIEN:TrailfromCanato Colombian border,Rio Setiganti, 19 Apr 1980 (st), Gentryet al. enedatthebase,theconicalstigmas,thenarrowerlamina 28590 (MO); Cerro Campamiento,E of Tres Bocas, (lanceolateto oblong), andlongerpetioles. In addition, Cuasi-Canatrail,29 Apr 1968 (9 fl-fr),Kirkbrideet al. the inflorescences and infiuctescences appearto be 1226 (MO, NY). PANAMA: Trail Zamora-Campana smaller,down to 0.5 cm and ca. 2.5 cm diam. respecNationalPark,14 May 1992 (9 fl-fr), Correa8911 (F, tively.Furtherexplorationmay providematerialelimiSCZ);CerroCampana,aboveSu-LinHotel, 16 Sep 1971 nating the morphologicaldiscontinuitiesbetween the (9 fr), Gentry1854 (MO). groups of specimens cited above. Naucleopsis capirensis is relatedto N. naga, from The collections cited from Costa Rica andPanama which it differs, e.g., in the absence of bracts on the are quite uniform. Several specimens from Colombia of the staminateinflorescences, the shorter peduncle and Ecuadorprobablybelong to this species andmay tepalsof the pistillateflowers, andthe shorterstipules. representtwo subspecies:

ReservadeArenal,RioPeinasBlancas,Quebrada Aguagata, FincaVillalobos,1000 m, 20 Apr 1990 (st), Bello et al. 2218 (MO);ReservaForestalSan Ram6n,Campamento Rio Lorencito,900-1000 m, 2 Sep 1989 (st), G6mezLaurito 11828 (F); San Carlos, Peiias Blancas, 10 Jul 1985 (9 fl-fr), Haber et al. 1960 (BG, MO); Reserva Biol6gicaMonteverde,Rio PefiasBlancas,15 May 1987 (? fl), Haberet al. 7033 (BG,MO), 13 June1987(? fr), Haberet al. 7221 (MO);ReservaBiol6gicaMonteverde, FincaNovo, 23 Jul 1987 (9 fr),Haberet al. 7339 (BG, MO);ReservaBiologicaMonteverde,Brillante,1500 m, 8 Sep 1987 (? fr), Haber et al. 7507 (MO); Upala, BijaguaEl Pil6n,CerroLa Carmela,8 Jul 1988(? fl-fr), Herrera 2018 (BG, F, MO,TEX).GUANACASTE:Tileran,

COLOMBIA. ANTIOQUIA: Mun. Frontino,Cgto. Nutibara,Rio Cuevas, 1800-2000 m, 17 Mar 1984 (a'), D. Sanchez et al. 60 (BG, MEDEL), 1700 m, 18 Mar 1985 (? fr), D. Sdnchez et al. 166 (BG, MEDEL); Mun.

Frontino,Vrda.Venados,ParqueNacionalNaturalLas

18. Naucleopsis naga Pittier (see Berg, Fl. Neotrop. Monogr.7: 136-139. 1972).

Revised description: Tree,to 30 mtall. Leafytwigs2-6 mmthick,sparsely (COL). andminutelyappressed-puberulous; peridermflaking These specimensdiffer fromthe CentralAmerican off. Laminaoblongto lanceolateor to subobovate,(8-) ones in the following: the trees becoming to ca. 35 m 15-30(-40) x (2.5-)4-10(-12) cm, coriaceous; apex tall, the fully matureinfructescencesbecoming prob- acuminate(to subcaudate)or to acute;base roundedto ably about 10 cm diam. with the free parts of tepals acute;uppersurfaceglabrous;lower surfaceglabrous on theveins;lateralveins becoming to 2.5 cm long and the fruits ca. 2.5 x 1.5 orpuberlous to subhispidulous cm. The stigmasareconical andsimilarto those found 15-28 pairs;tertiaryvenationscalariformto reticulate, in the collections cited below.The petiole can be to 1.5 the smallerveins plane or ? prominent;petiole 0.7-2.2 cm long. The staminateinflorescences aresubsessile. cm long, the epidermisflakingoff; stipules0.8-2(-3.5) cm long, (sparsely) appressed-puberulous,caducous. COLOMBIA. NARINO: Mun. Barbacoas, Cgto. Staminate inflorescences up to 3 together in the leaf Altaquer,Vrda.El Barro,ReservaNaturalRio Nambi, axils or below the leaves, ca. 0.5-2 cm diam.;peduncle 1325 m, 5 Dec 1993 (st), Betancur et al. 4631 (COL). 0.2-0.7 cm long, bracteatetowardsthe apex(sometimes VALLE: Alto Rio Anchicaya,Yatacue, 17 Jul 1984 (? peduncularbracts almost included in the involucre); fl), Gentry et al. 48260 (BG, COL, JUAM, MO). ECUADOR. CARcHI: ReservaIndigenaAwa, Chical, involucralbractsin ca. 5-7 series,semicircularto subor1150 m, 19-28 Feb 1993 (? fl-fr), Grijalvaet al. 600 bicular,the outerones minutelyappressed-puberulous, Orquideas, 880 m, Jun 1988 (a' juv), H. Triana et al. 56

281

TAXONOMICTREATMENT the inner ones only ciliolate; perianthca. 2 mm long; tepals 3-7, free, ? cucullate, puberulous;stamens 4; filaments 0.6-1 mm long; anthersoblong to obovate, 0.8-1 x 0.3-0.4 mm. Pistillate inflorescences 1.5-3.5 cm diamn., sessile;involucralbractsin ca. 6 series,semicircularto ovate to oblong, (the innerones) to 1.2 cm or to 2.5 cm long, minutelypuberulousto muriculate; flowers numerous;free partsof the tepals subulateto cylindrical,0.3-0.5 cm long, minutelypuberulous;stigmas subulate,3-6 mm long. Infructescences4-12 cm freepartsof the tepalspyramidateto subulate,to diamn.; 0.5 cm or to 2.5(-4.5) cm long, ? curved. Two subspecies can be recognized.

18a. Naucleopsis naga Pittier subsp. naga. Laminaglabrouson the lowersurface;smallerveins plane;stipulesto 3.5 cm long. Pistillate inflorescences with the innerinvolucralbractsto ca. 1.2 cm long. Distribution (see Fig. 84.1). From the Atlantic Coastal region of Honduras (Atlintida) to Pacific Coastal region of Colombia (Antioquia, Choco, and Valle); in wet forest;to ca. 2000 m. Representative additional collections examined. Rio SAN JUAN:20 kanNE of El Castillo, NICARAGUA. 18-21 Apr 1978 (st), Neill et al. 3497 (MO). CaminoRaizde Hule,SE COSTARICA.CARTAGO: of Platanillo, 1200-1400 m, 1 Jul 1976, Croat 36813 (MO).HEREDIA:La Selva,nr.PuertoViejo,29 Jun 1971 (V),Gentry1052 (MO),13 May 1970(d), Hartshorn885 (MO).LIM6N: Hda. Tapezco, 29 km W of Tortuguero, 26 Aug 1979 (q fl-fr),Davidsonet al. 8859 (MO,NY). PUNTARENAS: 1 mi S of San Vito de Java,ca. 1150 m, 18 Aug 1967 (o), Raven21897 (BG, F, MO). PANAMA.DARItN: CerroPirreca. 800-1500 m, 910 Aug 1967 (9 fr), Dukeet al. s.n. (MO);Cana-Altos de Nique,trailbetweenRio SetegantiandRioAltoTuira, 1050 m, 18 Apr 1992 (st), Foster 14115 (F). PANAMA: CerroAzul,nr.FincaVegade Melo, 13 Nov 1975,Ching Ho et al. 45 (MO,NY); CerroCampana,above Su-Lin Hotel, 16 Sep 1971 (q fl-fr), Gentry1845 (MO).

27 Mar 1985(? fr),MonsalveB. 832 (MO),2 Aug 1989 (? fl-fr), Monsalve B. 3178 (BG, MO). ECUADOR. CARCHI: San Marcos valley, 20 Nov 1983 (st), Barfod et al. 48912 (AAU); Canton Tulcan, Reserva Indigena Awa, Tobar Donoso, 650-1000 m, 1928 Jun 1992 (9 fl-fr), 7ipaz et al. 1394 (QCNE), 1800 m, 17-27 Aug 1992 (? fl), 7ipaz et al. 1811 (QCNE), 17-27

Aug1992(? fl),lipazetal. 1865(BG,QCNE).ESMERALDAS: Canton San Lorenzo, Reserva Etnica Awa, Centro Mataje, 21 Sep 1992 (? fr), Autestiaal. 539 (AAU, QCNE);Canton San Lorenzo, Reserva Indigena Awa, Ricaurte, Centro Pambilar, 21 Jan 1993 (? fl), Auslestia et al. 1016 (AAU, BG, MO, QCNE); Reserva Indigena Awa, Rio Mira, 10 km W of Alto Tambo, 16-26 Mar 1991 (? fl-

fr), Rubio et al. 1190 (QCNE). Uses. Latex used as poison for blowgun arrows, also used for big game (Barfod et al. 48912); see also the generictreatmentof Naucleopsis, above.

18b. Naucleopsis naga Pittier subsp. meridionalis C. C. Berg, Novon 6: 238. 1966. Type. Colombia. Nariino:La PlanadaReserve, nr. Ricaurte,21 Dec

1987($ fr), Gentryet al. 59669 (holotype,PSO; isotypes, BG, COL, MO, SI). Lamina puberulous or subhispidulous (and then scabridulous)on the lower surface;the smallerveins ?

prominent; stipulesto 1.5cmlong.Pistillateinflorescences with the innerinvolucralbractsto 2.5 cm long. Distribution (see Fig. 83.6). On the westem slopes of theAndes fromAntioquia(Colombia)to Azuay(Ecuador);in forest; at ca. 600-2 100 m.

Specimens examined. COLOMBIA. ANTIOQUIA: Mun. Frontino, corr. Nutibarra, source of Rio Cuevas, 1700-2000 m, 18 Jul 1987 (? (fl-fr), D. Sanchez et al. 1479 (MEDEL).NARI*o: La PlanadaReserve, nr.Ricaurte, 1800 m, 25 Jul 1986 (st), Gentry et al. 55101 (BG), 15 Nov 1987 (9 fl-fr),Restrepoet al. 409 (MO).RiSARALDA: Pereira, Jan 1963 (st), Goitia UDBC 631 (UDBC). ECUADOR. AZUAY: Between Jesus Maria and Molleturo, ca. 12 km from Guayas border, 1200 m, 16 Jul 1977 (e), Boeke2184 (NY).BOLIVAR: Hda. Changuil, COLOMBIA. ANTIOQUIA:Mun. Amalfi, Vrda. El Sector la 47, Aug 1995 (2 fl-fr), Bonifaz et al. 3240 (BG). Oso, 1590m, 30 Sep 1988(st),Betancuret al. 903 (MO); CARCHI:Canton Mira, El Carmen, rd. to Chical, 1600Mun. Frontino,Cgto. La Blanquita,rd. Nutibara-La 1800 m, 10 Feb 1992 (a), Palacioset al. 9674(BG, QCNE), Blanquita, km 10-18, Callejas et al. 6645 (MO). 2000-2200 m, Palacios et al. 9691 (BG, QCNE), 18BOLivAR: Serraniade San Lucas,Vrda.La Libertad,20 25 Aug 1994 (? fl-fr), Tirado et al. 1306 (QCNE). Jul 1987 (st), Cuadros3670 (MO).CHOC6: Nr. Quibd6, COTOPAXI: Rd. Quevedo-Latacunga, 3 km E of El Pal4 May 1985 (q fr), Cogollo2088 (MO);Alto de Nique, mar, 800 m, 5 Apr 1980 ("), Dodson et al. 10240 (BG, borderColombia/Panama,1350-1520 m, 19 Apr 1980 MO, QCNE). ESMERALDAS: Rio Lita, 4.5 km WNW of (7 fr), Gentryet al. 28636 (F). VALLE: Ca. 16 km NW Lita, 600 m, 8 May 1987 (a), Daly et al. 5150 (BG, of Buenaventura, 21 May 1987 (st), Faber-Langendoen MO, NY, QCA).

et al. 635 (MO);SantaHelena,aboveTopacio,1950 m, The two subspecies occur sympatricallyin Ecua11 Dec 1985 (d?), Gentryet al. 53156 (BG, MO);Bajo Calima,Dindoarea,7 Feb 1989(st), Gentryet al. 65531 dor andColombia,althoughthereseems to be a differ(BG); Bajo Calima,Concesi6nPulpapel/Buenaventura, ence in preferenceto elevation.The differentiationinto

282

leavesandonewithleaveshairy anentitywithglabrous beneathis similarto thesituationfoundinNaucleopsis entiulei.However,inN. nagathetwomorphological thaninN. ulei,andmayproveto tiesaremoredifferent be distinctatthespecieslevel. Ducke(see Berg,Fl. 19.Naucleopsisulei (Warburg) 7:139-142.1972,incl.N.amara). Neotrop. Monogr. Additionalsynonym: Uleodendronulei (Warburg) Rauschert, Taxon 31: 560. 1982, based on Acanthosphaeraulei Warburg.

Therangeof distribution hasexpanded considerably withnewdiscoveriesin CostaRica,Panama,andthe andEcuador. For PacificCoastallowlandsof Colombia thismateriala new subspecieshadto be recognized, subsp.puberula.Newcollectionsmadethedistinction betweenNaucleopsisulei andN. amaramoreandmore

thelatterhadto bereducedto a subspeproblematical; cies of the former.A fourthtaxon,still ratherpoorly known,is includedas subsp.subandina.

FLORANEOTROPICA puberulous;style 2-7 mm long; stigmasband-shaped, (3-)7-lOmm long.Infiuctescences3-15 cm diam.;free partsof the tepalssubulateto narrowlypyramidate,1-5 cm long, ? curved;fruit ca. 2-3 x 2-2.5 cm. Key to the subspecies of Naucleopsis ulei 1. Laminaminutelypuberulousto subhispidulous (and then ? scabridulous)beneath; leafy twigs normallywithoutdistinctly shorterintemodes;CentralAmericaand Pacific Coastalregion........... 19d. subsp. puberula 1. Laminaglabrousor only sparselypuberulous on the veins;leafy twigs with partswith distinctlyshorterintemodes,the nodes of these partsonly bearingstipules. 2. Base of laminausually cordateor subcordate;lower lateralveins usually distinctly loop-connected;stipules to 2.5 (-3.5) cm long, often subpersistent; laminausually20-50 x 7-18 cm; lateral veins (20-)25-35 pairs; upper Amazon 9a. subsp.ulei Basin.1 2. Base of laminaroundedto acute;lower lateralveins not or only faintly loopconnected;stipules mostly caducous. 3. Laminausually 15-30 x 5-12 cm; lateralveins 17-25 pairs;stipulesto 1.5(-2.3) cm long; free partsof the tepals of the pistillateflowersin fruitto 5 cm long;mainlylowerand middleAmazonBasin.... 19c. subsp.amara 3. Laminausually30-47 x 7.5-13 cm; lateralveins 25-33 pairs;stipules mostly 1.5-3 cm long; free parts of the tepalsof the pistillateflowers in fruitto 3 cm long; subandean Peruand Bolivia.... 19b. subsp. subandina

Reviseddescription: Tree,to 25(-40) m tall;intemodesoftendistinctly differentinlength,thepartswithshortintemodesbearingonlystipules.Leafytwigs4-12 mmthick,brownorsubglabrous; periderm ishto yellowish-puberulous irregularly wrinkled,flakingoff in verysmallflakes. tooblanceolate, (1O-)15Laminaoblongtosubobovate apexacuminate; 40(-60) x 3-15(-2 1)cm,coriaceous; basecordateto roundedto subacuteorto acute;upper or misurfaceglabrous;lowersurface(sub)glabrous nutely puberulousto subhispidulous (and then on theveins;lateralveins(13-)17-35 scabridulous) pairs,atleastinthelowerpartof thelaminamostlynot venationpartlyscatertiary distinctlyloop-connected; Duckesubsp.ulei. to reticulate; lariform petiole0.4-1.5cmlong;stipules 19a.Naucleopsisulei (Warburg) to patent0.8-2.5(-3.5)cmlong,yellowishappressed Shortenedinternodes (bearingonly stipules) usupuberulous,mainlyat the base, often curved,often Laminarelatively ally conspicuously present. large,20onpartsof theleafytwigswith subpersistent (especially 50(-60) x 7-1 8(-20) cm;basecordateto rounded(to shortintemodes). Staminateinflorescencesin the leaf lowersurface glabrous (exceptfortheminute, axilsorbelowtheleaves,upto 4 together,(0.2-)0.5- subacute); brown or oronlyhairy whitish, pluricellular trichomes) 1.5cm diam.,subsessileorpeduncleto 1.2cm long; at lateral veins the base of the lower midrib; (20-)25bractsin ca.6receptacleslightlyconcave;involucral ? distinctlyloop-connected; 35 pairs, mostly stipules 7 rows,reniformto suborbiculate, obtuse,sparselyto cm (1-)2.5-3.5 long, often subpersistent. Staminateinflowers densely yellowish appressed-puberulous; ca.2 mmlong,4- florescencessubsessileorpeduncleto 0.2cmlong.Free slightlyconnateatthebase;perianth stamens1-4;an- partsofthe tepalsofpistillateflowersto 4 cm longinfruit. 7-fidto -parted,sparselypuberulous; Distribution.IntheupperAmazonBasin;in nonthers0.5-1 x 0.24.3 mm. Pistillate inflorescencesbeforest;to 1200m. low the leaves, (0.7-)1-3.5 cm diam., sessile or inundated bractsin ca.7 series,reniform to subsessile;involucral Representative additional material examined. deltoidtoovatetolanceolate, obtuse,grayish-puberulous COLOMBIA.AMAZONAS:Mun.Leticia,ParqueNacional sometimesfew(3- NaturalAmacayacu,28 Apr 1991 (2 fl-fr),Pipoly et a). (topubescent); flowersnumerous, 5?);freepartsof the tepalssubulate,0.2-2 cm long, 15590 (MO), 19 Aug 1991 (? fr), Rudas et al. 2943

TAXONOMICTREATMENT

283

Type.Bolivia.La Paz:"NewBrazil,"13 Jun 1902 (st), R. S. Williams1660 (holotype,NY). A subsp. ulei laminae basi acuta vel subacuta;a ECUADOR.NAPO: Rd. Coca-Auca,km 50-60, 27 Jul 1981(st), Berget al. 1094 (AAU,NY,QCA);Reserva subsp.amarafoliis majoribusbinatim25-33 nerviis;a Flon'sticaEl Chuncho,5 kmN of Coca, 10-15 Sep 1986 subsp.puberulalaminasubtusglabralaevi diversa. (d'), Neill et al. 7295 (BG, NY, QAME,QCA, QCNE); Shortened internodes usually present, but not conAniangu,ParqueNacionalYasuni,30 May-21 Jun 1982 spicuously. Lamina 23-47 x 7.5-13 cm; base acute or (e'), SEF 8568 (MO,NY,QCA);ReservaBiol6gicaJatun E of Misahualli,18-30May1985(v), Zaruma subacute (to rounded); lower surface glabrous; lateral Sacha,8 kam et al. 215 (BG,MO,NY,QAME,QCA,QCNE).PASTAZA: veins 25-33 pairs, at least in the lower part of the lamina Pozo PetroleroMoretecocha,75 km E of Puyo, 4 Dec often not distinctly loop-connected; stipules (0.8-)1.51990 (e), Gudinio1157 (BG, MO, NY, QCNE);Pozo 3 cm long, mostly caducous. Free parts of the tepals of PetroleroGarza,35 km NE of Montalvo,2-12 Jul 1989 the pistillateflowers to 3 cm long in fruit. (? fr), Zak et al. 4731 (BG, MO, NY, QCNE). Distribution (see Fig. 83.4). In subandean Peru SucuMBsos:ReservaFaunisticaCuyabeno,1 Apr 19881 Apr 1990 (st), Valenciaet al. 68667 (AAU, QCA); and Bolivia, from San Martin and Ucayali (Peru) to Rio Auguarico,SE of DestacamentoLagartoCocha,30 Cochabamba (Bolivia), also in Acre (Brazil); in Aug 1979 (? fl), Holm-Nielsenet al. 20266 (AAU,BG, submontane and lowland forest; to 1500 m. (BG); TrapecioAmazonico,Rio Loretoyacu,Oct 1945

(e), Schultes 6713 (COL).

Canton Nangaritza, Cord. QCA). ZAMORA-CHINCHIPE:

Specimensexamined. PERU. HUANUCO:Pachitea, Codo de Pozuzo, 21 Oct 1982 (? fl-fr), Foster 9342 (MO). MADRE DE Dios: Ca. 20 km W of Puerto PERU. AMAZONAS:Rio Santiago, ca. 65 km N of Maldonado,rd. to Quince Mil, 23 Aug 1977 (? fr), Pinglo, QuebradaCaterpiza, 15 Mar 1980 (? fl-fr), Gentryet al. 19692 (MO,US). PAscO:Prov.Oxapampa, Huashikat2289 (MO);QuebradaHuampi,14 Apr 1973 PuertoBermiidez,Dec 1980 (? fl-fr),Fosteret al. 8022 (st), Kayap635 (NY). HuANuco:Maquizapa,carretera (BG, F, MO,MOL);Prov.Oxapampa,PuertoEsperanza, Monz6n, 12 Feb 1966 (st), SchunkeV 1067 (ECON, 13 Oct 1980 ($ fl-fr), ReynelR. 6 (MOL);Prov.OxaNY). LORETO: Prov.Alto Amazonas,Andoas,21 Nov pampa,Dtto. PuertoBermuidez, PuertoVictoria,15 Nov 1980(? fl), R. Vasquez et al. 802 (BG,MO);Prov.Loreto, 1980(st),ReynelR. 70 (MOL);Prov.Oxapampa, Iscosazin, et al. 3422 (BG, 4 Jul 1982(st),ReynelR. 648 (BG).PUNO:RioCandamo, Nauta,9 Nov 1982 (? fl-fr),R. Vdsquez F, MO,NY). nr.mouthof Rio Guacamayo,25 May 1992 (st), Gentry BRAZIL. ACRE:Mun. Mancio Lima, Rio Moa, et al. 77188(MO),29 May 1992(st), Gentryet al. 77328 FazendaBoa Vista, 12 Oct 1989 ($ fl-fr),Ferreiraet al. (MO). SAN MARTfN: Nr. Aguaytia, 4 Jul 1949 (st), 10018 (NY). RONDONIA:Mun. P6rto Velho, Usina Mathiaset al. 3571 (F, MO). UCAYALI:Prov.Coronel Hidroelectricade Samuel, 15 Jun 1986, Ferreira7461 Portillo,Dtto.Raimondi,Atalaya,7 Mar1981(st),Reynel (BG, NY, US); Madeira-Mamorerailway,km 133, 15 R. 141 (BG, MOL). Sep 1963 (9 fl-fr), Maguireet al. 56661 (F, NY, U), BRAZIL. ACRE:ReservaINCRASantaLuzia,5-19 (a'), Maguireet al. 56669 (NY, U). Oct 1984 (st), Campbellet al. 7061 and 7601 (BG). Prov.Abuna,Rio Negro,nr.jct. BOLIVIA.PANDO: Prov.Carrasco,Estaci6n BOLIVIA. COCHABAMBA: with Rio AbunA,7 Jun 1992 (st), Gentryet al. 77975 Valledel SajtaUMSS,31 Oct 1990 (q fl), Atahuachiet (MO); Rio Madeira,opposite Abuna, 9 Jul 1968 (o), al. 34 (MO). del C6ndor,above Rio Nangaritza,Pachicutza,4 Dec 1990 (? fl-fr), Neill et al. 9513 (BG, MO, QCNE).

Prance et al. 5720 (MICH, NY).

Thestaminateinflorescencesofthe materialof subsp. ulei collected in Ecuadortend to be largerthan those collected in Brazil, Bolivia, and Peru. The infructescences show considerablevariationin size in relation to the numberof flowers (andfruits)per inflorescence.

19b. Naucleopsis ulei (Warburg)Ducke subsp. subandina C. C. Berg, subsp. nov. Typus.Peru. Madrede Dios: ca. 5 km from Puerto Maldonado,nr. Rio Tambopata,24 Jan 1976 (? fl-fr), Gentty et al. 16275 (holotype, MO; isotypes, F, NY).

Subspecies subandina differs from the typical subspecies in the usually acute to subacute base of the lamina, and (therefore) at least the lower lateral veins not distinctly loop-connected, the usually caducous stipules, and inconspicuous occurrence of shortened intemodes. The leafy twigs are relatively slender. Subspecies subandina differs from subsp. amara in the larger leaves (mostly longer than 30 cm) with more lateral veins (25-33 pairs), and in the shorter free parts of the tepals of the pistillate flower in fruiting state. The infructescences appear to be usually large. Some of the label data suggest that this subspecies is associated with submontane forests.

FicusoblanceolataRusby,Bull. NewYorkBot. Gard. 6: 498. 1910, non F oblanceolataRusby,Bull. New YorkBot. Gard.4: 446. 1907, renamedF 19c. Naucleopsis ulei (Warburg) Ducke subsp. amara (Ducke) C. C. Berg, comb. et stat. nov. Basionym. katherinaeA. D. Hawkes,Phytologia3: 32. 1948.

284 Naucleopsis amara Ducke, Arch. Jard.Bot. Rio de Janeiro 4: 6. 1925. Uleodendron amarum (Ducke) Rauschert,Taxon 31: 560. 1982. Type. Brazil. Para:Rio Tapajos,nr. cataractsof Mangabal, 14 Dec 1919, (? fr), Ducke MG 16757 = RB 13029 (lectotype, Berg, Fl. Neotrop. Monogr. 7: 142. 1972, RB).

FLORANEOTROPICA 19d.Naucleopsisulei (Warburg) Duckesubsp.puberula C.C.Berg,Novon6:240.1996.Type.PanamaPanamA: ca. 20 NE of Chepo, 12 Dec 1973 (? fl), Berg & Nee 313 (holotype, MO; isotypes, AAU, BG).

Shortenedinternodes sometimes present.Lamina (sparsely) puberulous to subhispidulous (and then scabridulous) on the lower surface; base cordate to Shortenedinternodesusually present,butnot con- rounded(to obtuse); lateralveins ca. 25-35 pairs, the spicuously.Laminarelativelysmall, (l0-)15-30(-50) lower lateralveins distinctly loop-connected;stipules x 5-13 cm; base acute to rounded(or to subcordate); 1.5-3.5 cm long, subpersistent.Staminate infloreslower surface (sub)glabrousor minutely puberulous cences with to 1.2 cm long peduncle.Freeparts of the on the veins; lateralveins 17-25(-28) pairs,at least in tepals ofthepistillateflowers to 2.5 cm long in fruit. the lower partof the laminaoften not distinctly loopDistribution (see Fig. 83.4). From Costa Rica to connected;stipules 0.8-1.5(-2.3) cm long, mostly cawesternEcuador;in wet forest; at low elevations. ducous.Freeparts of the tepals of thepistillateflowers to 5 cm long in fruit. Specimensexamined.COSTARICA.LIM6N:Reserva IndigenaTalamanca,rd.Amburi-Cachabri, 27 Jun 1989 Distribution.IntheAmazonBasin,mainlythemiddle (? fl), Chac6n 8 (BG, F, MO). PUNTARENAS: Rancho and lower partof the basin; in non-inundatedforest. nr.Rinc6n,11Jan1993(st),Gentryet al. 78665 Quemado, Representative additional specimens examined. and 78727 (MO); Reserva Forestal Golfo Dulce, Osa COLOMBIA. AMAZONAS:Rio Cahuinari,31.5 km W Peninsula, 15 km W of Rinc6n, 3 Jun 1988 (? fl-fr), of its mouth,10 Sep 1988(st), M. SdnchezS. et al. 1114 Hammelet al. 16990 (MO);ParqueNacionalCorcovado, (BG). CAQUETA:Araracuara,11 Nov 1988 (st), P. Gira de Corcovado,Arco de Piedras-Rio Corcovado, 16 Jun 1989 (? fl-fr), Kernan1150 (BG, F, MO). Palacios et al. 2171 (BG). PANAMA.COCL*:9.4 km above El Cope, 20 Jan BRAZIL. AMAPA:Rio Araguari,5 hrs above Rio 1978 (st), Croat44752 (MO).COL6N:SantaRitaRidge Murur6,24 Aug 1961 (st), Pires et al. 50461 (MG,NY, Hwy.andAguaClara,11 Dec U). AMAZONAS:DistritoAgropecuara,90 km NNE of rd.,betweenTransisthmian Manaus,Reserva1501, 25 Nov 1988 (? fl-fr),Boomet 1973 (d), Berg et al. 301 (AAU, BG), Berg et al. 307 al. 8650 (NY), 11 Jan 1992 (e fl-fr), Oliveira et al. (AAU,BG, F, MO,NY); SantaRita EastRidge,23 Mar 305 (NY);Rio Javari,7 hrsabovePaumari,16 Oct 1976 1968(? fl), Correaet al. 891 (MO);SantaRitaRidgerd., (?), Prance et al. 23841 (NY); Rio Negro, Rio Jaua- betweenTransisthmianHwy. and Agua Clara, 11 Mar pen, EstiraoTacuera,24 Feb 1977(? fl-fr),M. R. Santos 1970 (? fl), Dressler 3858 (MO), 11 Dec 1973 (e), Dressleret al. 8825 (MO);SantaRitaRidge,20.7 kmfrom 98 (NY). PERU.LORETO:RioAmazonas,abovemouthof Rio TransisthmianHwy., 22 Mar 1992 (st), Foster14079 (F, SantaRita Ridge road,4 mi fromTransisthmian Napo, Yanomono,22 Mar 1982 (? fl), Gentryet al. SCZ); 11 Dec 1973 (a), Gentryet al. 8825 (MO);Santa Hwy., 36547 (MO,NY); Prov.Maynas,Rio Nanay,Mishana, RitaRidge, 11 Jan 1987 (R fl), McPherson10262 (BG, 6 Jan 1983 (st), Gentryet al. 39095 (MO). MO).PANAMA:CerroJefe, Altos de Pacora,18.2 km E Subspecies amara differs fromthe typical subspe- of CerroAzul (village), 5 Jan 1975 (? fl), Gentryet al. cies in the relatively small leaves, mostly less than30 13416 (MO, U); 10 km NE of Altos de Pacora,6 Mar cm long, the smallernumberof lateralveins,the smaller 1975 (? fl-fr), Moriet al. 4946 (MO). SANBLAS: stipules(mostlyto 1.5cm long) andthe longerfreeparts Cangandi,16 Dec 1985, Nevers et al. 6516 (MO). COLOMBIA. ANTIOQUIA: Nr. Villa Arteaga, 6 Dec of the tepalsof the pistillateflowers in fruitingstate(to 1948 (? fl-fr), F L6pezet al. 31 (MEDEL,US); Mun. 5 cm long). The base of the laminais often subcordate, Zaragoza,Cgto. Providencia,11 Feb 1971(? fl), Soejarto and,as a consequence,the lower lateralveins are(usu- et al. 2760 (COL,GH); Mun. Riosucio, Urabaregion, ally) not distinctly loop-connected. The presence of Cerros del Cuchillo, 13 Aug 1987 (? fr), Cdrdenas286 partsof leafy twigs with distinctly shorterinternodes (JUAM, MO), 11 Sep 1987 (q fl-fr), Cdrdenas 445 only bearing stipules is less conspicuous in subsp. (JUAM),16 Nov 1987(st), Cdrdenas867 (JUAM,MO), amarathanin subsp. ulei, as suchpartsarenot present 15 Jan 1988 (? fr), Cardenas 1032 (JUAM,MO), 23 on all branches,the numberof shortenedintemodes is Mar 1988 (? fl), Cardenas1459 (JUAM,MO), 30 Jun less, andthe intemodesnot very short.Subspeciesulei 1988 (? fl-fr) Cdrdenas2287 (JUAM). CHOCO:Rd. 3 km W of Tutunendo,8 Jan 1981 is found in the upperAmazon Basin and subsp.amara Quibd6-Tutunendo, (st), Gentryet al. 30334 (BG, JUAM);Rio Mecana,ca. extends from the upper to the lower Amazon Basin. 10 km E of Mecana,7 Mar 1983 (? fl), Gentryet al. These two subspecies are neither geographicallynor 41060 (BG, COL,JUAM,MO). C6RDOBA:Jct. of Rio morphologicallyclearlyseparated. Tigre and Rio Manso, 28 Jul 1988 (st), Gentryet al.

TAXONOMICTREATMENT 63892 (BG, MO). NARIfo: Nr. Tumaco, Rio Rosario, 5

km above Santa Maria, 21 Jun 1955 (9 fl), RomeroCastanieda5194 (COL). ECUADOR. ESMERALDAS:Canton San Lorenzo, ReservaEtnicaAwa,CentroGuadualito,20-29 Jul 1992 (? fl-fr),Autestiaet al. 43 (QCNE);Cant6nSanLorenzo, ReservaEtnicaAwa, Ricaurte,CentroPambilar,21 Jan 1993 (9 fl-fr), Aulestia et al. 1009 (AAU, BG, MO, QCNE);Mataje,6 Sep 1991 (d), Jaramilloet al. 13819 (QCA); Eloy Alfaro, CharcoVicente, Rio Miquel de Cayapas,20-27 Mar 1993 (? fl-fr), Mendezet al. 157 (QCNE);Cant6nEloyAlfaro,SanMiguel,Rio Cayapas, 3-5 Sep 1993 (st), Palacios et al. 11145 (QCNE); CantonSan Lorenzo,ReservaIndigenaAwa, Ricaurte, 19-24 Oct 1992(q fl), Tipazet a. 2071 (QCNE);Canton Eloy Alfaro, Reserva Ecol6gica Cotacachi-Cayapas, CharcoVicente,Rio SanMiguel,20-31 Sep 1993(? fl), lirado et al. 450 (QCNE);CantonEloy Alfaro,Reserva EcologicaCotacachi-Cayapas,Luis VargasTorres,2327 Oct 1993 (9 fl-fr), 7iradoet al. 514 (QCNE);Cant6n Rio Eloy Alfaro,ReservaEcologicaCotacachi-Cyapas, Santiago,Angostura,17-26 Jul 1994 (st), Tiradoet al. 1073 (QCNE).

20. Naucleopsis humilis C. C. Berg, Novon 6: 238, t. 7. 1996. Type. Ecuador.Pastaza:Rio Curaray,nr. LagunaGarzayacu,20-26 Aug 1985 (ci), Neill et al. 6663 (holotype, QCNE; isotypes, BG, MO, Fig. 70 QAME, QCNE). Treelet,to 3 m tall; internodesoften distinctlydifferent in length, the parts with short internodesonly bearingstipules.Leafytwigs 3-5 mm thick,appressedpuberulous;peridermflaking off. Laminasubobovate to oblanceolate,(9.5-)20-45 x (2-)5-10 cm,coriaceous; apex subcaudateto acuminate;base subacuteto obtuse (to roundedto subcordate);margin entire;uppersurface glabrous;lower surfacepuberulouson the base of the midrib;lateralveins ca. (1 5-)25-30 pairs;tertiary venationlargely scalariformto largelyreticulate;petiole 1-2.5 cm long, minutelypuberulous;stipules0.51.5 cm long, striate,yellowish appressed-puberulous, subpersistentor caducous.Staminateinflorescenceson to 0.4 cm long spurs,ca. 0.2-0.3 cm diam.;peduncleca. 0.2 cm long, bracteate;involucralbractsin ca. 5 series, broadlyovate to oblong, the innerones glabrous;flowers 4-6; perianth2-2.5 mm long; tepals 3-5, basally connate,glabrous;stamensI or 2; filaments3-3.5 mm long, thick; anthers elliptic, 0.5-0.6 x 0.3-0.4 mm. Pistillate inflorescencessolitaryin the leaf axils, ca. 1 cm diam., sessile; involucral bracts in ca. 6 series, broadlyovate, minutelypuberulous;flowers few; free partsof thetepalssubulateto aculeate,0.3-0.6 cm long, minutelypuberulous;stigmasband-shaped,ca. 3 mm long. Infructescencessubglobose2-3 cm diam.,yellow

285 at maturity;free parts of the tepals 0.54.7 cm long; fruits 3-6, ca. 0.64.7 cm long. Distribution (see Fig. 83.4). In the northwestern Amazon Basin (Napo region); in riverine, often periodically inundatedforest, or in non-inundatedforest. Nr. Specimens examined. COLOMBIA. CAQUETA:

20 Nov 1991(st), Duivenvoorden et al. 1207 Araracuara, et al. 2319 (BG), (BG), 5 Dec 1991 (st), Duivenvoorden 8 Dec 1991 (st), Duivenvoordenet al. 2449 (BG, MO); Rio Caqueta,1 km above the mouthof Rio Caguan,29 4151 (COL,MO,NY), Apr 1953 (a), Romero-Castanteda 4152 (COL). (? fl-fr), Romero-Castanleda ECUADOR.NAPO:ParqueNacionalde Yasuni,Lagunasde GarzaCocha,22 Sep 1988 (? fr), Cer6net al. 5098 (AAU, MO, QAME,QCNE);Rio Yasuni,80 km upriverfromNuevo Rocafuerte,17 Sep 1977 (? fl-fr), Lorocachi,31 Foster3714 (QAME,QCA,S). PASTAZA: May 1980 (a'), Jaramilloet al. 31580 (AAU,MO,NY). PERU. LORETO:Prov. Maynas, Varaderode Mazan,

fromRioAmazonasto Rio Napo,22 Aug 1972(e'), Croat 19523(MO);Prov.Maynas,CaserioMishana,30 km SW of Iquitos,16Aug 1980(? fl-fr), Foster4341 (BG);Prov. Maynas,Quebrada Yanamono,ExploramaTouristCamp, 5 Nov 1979 (2 fr), Gentryet al. 27483 (BG, MO), 13 Nov 1979 (st), Gentryet al. 27962 (MO), 27 Jul 1980 (a"),Gentryet al. 29149 (MO), 18 Feb 1981(9 fr), Gentryet al. 31402(MO),4 Jul 1983(a'),Gentryet al. 42552 (BG, MO), 8 Jul 1983 (? fl), Gentryet al. 42798 (BG, MO);Prov.Maynas,1 km S of Indiana,17 Jun 1987(a), Gentryet al. 54637 (MO);Prov.Maynas,Rio Amazonas, ca. 2 km W of Indiana, Explorama Inn, 12 Feb 1987 (?

fr), Gentryet al. 55731 (BG, MO), 18 Feb 1988 (R flfr), Gentryet al. 61645 (MO),20 Feb 1988 (a'), Gentry et al. 61767 (BG, MO);Prov.Maynas,Rio Amazonas, nr. Indiana,15 Feb 1989 (a), Gentryet al. 65793 (BG, MO);Prov.Maynas,CaserioGamitana,Reservadel Rio Mazan, 21 Jun 1990(o), Grandezet al. 1608 (BG,MO); Prov.Maynas,Dtto. LasAmazonas,nr. Sucusari,20 Feb 1991 (a), Pipoly et al. 13044 (MO);Prov.Maynas,Rio Amazonas,Yanamono,26 Jun 1984 (? fr),R. Vasquezet al. 5173 (MO);Prov.Maynas,Quistococha,27 Sep 1984 et al. 5601 (BG, MO);Prov.Maynas, (Qfr), R. V&squez Indiana,15 May 1989 (? fl-fr),R. V4squezet al. 12138 (BG, MO); Prov. Maynas, Santa Maria de Nanay, QuebradaYarina,22 May 1989 (? fl-fr), R. Vazquezet al. 12236(BG,MO);Prov.Maynas,Indiana,23 Mar1990 (o), R. Vdsquez et al. 13634 (BG, MO), 11 Jul 1990 (i fl), R. Vasquezet al. 14085 (BG, MO). Naucleopsishumilisis closely relatedto N. ulei,from which it differs, e.g., in the small size of the trees, the smallerstipules,andthe smallerinflorescences.Inmost of the collections the base of the laminais acuteto obtuse.Insomecollections,Pipolyetal. 13044andRomeroCastaneda 4151, the base of the laminais subcordateor narrowlycordate,andthey areregardedas belongingto this specieslargelybecauseof the lengthof the stipules.

286

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Fig. 70. Naucleopsis humilis. 1. Leafy twigs with pistillate inflorescence (Gentry et al 61645). 2. Parts of twigs with only stipules (R. Vdsques et al. 5173). 3. Leafless twigs with staminate inflorescences. 4. Staminate flowers (Neill et al. 6663). [By Hendrieke Berg, Bergen.] 21. Naucleopsis krukovii (Standley) C. C. Berg (see Berg, Fl. Neotrop. Monogr. 7: 142-144. 1972). Distribution. In the upper Amazon Basin, from Ecuadorto Bolivia andto eastem Para(Brazil);mostly in non-inundatedforest, sometimes in inundatedforest (varzea);at low elevations.

Representative additional collections examined. COLOMBIA. AMAZONAS: Mun. Puerto Narihlo, Parque Nacional Natural Amacayacu, 21 Mar 1991 (st), Rudas San Jose de Guaviare,Vrda. et al. 1755 (MO). GUAVIARE: San Francisco, 29 Nov 1977 (9 fl-fr), H. Delgado 28 (UDBC). VAUP]S: Estaci6n Biol6gica Capar'u,3 km N of Laog Taraira, 4 Feb 1989 (d?), Defler 327 (MO).

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ECUADOR. MORONA-SANTIAGO: Rio Morona, Puerto Morona, 30 Sep 1975 (? fl-fr), Little et al. 501 (COL, LOJA,QAME, QCNE, US). NAPO:Ca. 10 km NW of Coca, 27 Jul 1981 (st), Berg et al. 1078 (NY, MO, QCA); Reserva Biologica Jatun Sacha, 8 km E of Misahualli, 14 Aug 1987 (st), Palacios 1851 (BG, QAME); Estacion Exp. INIAP-San Carlos, Reserva Floristica El Ahuano, 8 Sep 1986 (d), Zaruma 601 (AAU, BG, MO, NY, QCA, QAME, QCNE); Aniangu, Parque Nacional Yasuni, 1-15 Feb 1986 (st), Korning et al. 47544 (AAU, QCNE); Parque Nacional Yasuni, Pozo Petrolero Amo 2, 9-19 Jan 1988 (st), Neill et al. 8305 (BG, MO, Rio Cuyabeno,ca. 75?55'W, QAME, QCNE). SUCUMBiOS: 0?10'S, 25 Feb 1980 (st), Berget al. 1069 (AAU, QCA); Dureno, 27 Dec 1988 (st), Gentry et al. 64241 (BG). PERU. MADREDE Dios: Pantiacolla, Serania across, Rio Alto Madre de Dios from Shintuya, 28 Oct 1979 (? fl-fr), Gentry et al 27288 (AAU, NY). PASCO: Prov. Oxapampa, Iscozacin, 8 Aug 1981 (st), Foster 4647 (F). PUNO:Rio Tavara, 19 May 1992 (st), Gentry et al. 76856 (MO). SAN MARTfN: Prov. Mariscal Careres, Dtto. Tocache Nuevo, 10 Nov 1969 (6'), Schunke V 3608 (NY, MO, MOL, U), 20 May 1975 ($ fl-fr), Schunke V 8515 (AAU, MO, NY, U). BRAZIL. ACRE:Rio Branco, P6rto Velho Hwy., km 22, 30 Mar 1979 (9 fl-fr), Albuquerque et al. 1292 (MO, NY, U); rd. Brasileia-Assis Brasil, km 8, 1 Nov 1980 (? Rd. Manausfl-fr),Lowrieet al. 665 (NY). AMAZONAS: Caracarai, BR.174, km 159, 20 Sep 1974 (? fl-fr), Prance et al. 22713 (INPA, NY, U). ROND6NIA:Rio Jamari, Cachoeira de Santa Cruz, 28 Jun 1965 (? fl-fr), Pires et al. 9954 (NY) and 9955 (NY, US). BOLIVIA. LA PAZ:Prov.Abel Iturralde,Alto Madidi,

Tree,to 15 m tall. Leafy twigs 1.5-2.5 mm thick,

to hirtellous; peridennflakingoff.Lamina puberulous ellipticto oblongto subobovate,6-15 x 1.8-5 cm, marbase(sub)acute; apex(sub)caudate; subcoriaceous; ginplane;bothsurfacesglabrous;lateralveins 8-13 pairs;tertiaryvenationreticulate;petiole0.3-0.8 cm theepidermis? flakingoff;stipules long,puberulous, or subpersistent 0.5-1 cm long,sparselypuberulous, Pistillate caducous.Staminateinflorescencesunknown. intheleafaxilsorjustbelowtheleaves, inflorescences ovate withbroadly 1.2-2cmdiam.,subsessile; involucre to oblongbractsin ca. 5 rows,theinneronesi scariwhendry); ous andsubglabrous (andstraw-colored flowersca. 5-10; free partsof the tepalssubulate, ca.0.3-0.6 cmlong sparselyandminutelypuberulous, whendry);stigmasca.2 mmlong. (andstraw-colored 2.5-4 cmdiam.;freeparts subglobose, Infi-uctescences of thetepalsto 1cmlong,subulate(to aculeate). Distribution(see Fig. 83.5). In the northemPapart cificCoastalregionof Colombiaandtheadjacent of Panama,andin CostaRica;in forest;to 1000m.

Specimens examined. COSTA RICA. HEREDIA: ParqueNacionalBraulioCarrillo,transecttrail, 1000m, 25 Nov 1992 (st), Boyle 1248 (MO). PANAMA.DARIEN: Between Cana and Altos de Nique, trail Rio Seteganti-RioAlto Tuira,19 Apr 1992 (st), Foster14254 (F, SCZ). COLOMBIA.ANTIOQUIA: Mun.Mutata,rd.to Pavarandogrande,3 km beyond Rio Sucio, 8 Dec 1982 (? fr), Bernal et al. 436 (COL, JUAM); between Villa 22 May 1990(? fl-fr),Gentryet al. 70400(MO).PANDO: Arteagaand Chirorod6,El Tigre, 1 Oct 1961 (? fl-fr), Prov. Nicolas Suarez, SW of Cobija, Rio Naraueda, 31 Cuatrecasas et al. 26142 (COL,US).

Jul 1982 (? fl-fr), Sperling et al. 6425 (BG, NY).

The freepartsof the perianthof the pistillateflower can be to 1 cm long and may vary from distinctly aculeate to pyramidateto almost cushion-shaped.Their indumentummay varyfromrathersparselypuberulous to subvelutinous.The remarkablecircularholes in the lamina(possibly caused by some fungus) arefound in almost all additionalcollections. Reproductionsometimes startsin treeletsof about3 m tall. The collection D. N. Smith 3900 largely matches Naucleopsis krukovii,butthe pistillateinflorescenceis distinctin thevery short,ca. 1 mmhigh,cushion-shaped free partsof the tepals and peripheral"pseudobracts." PERU. PASCO:Prov.Oxapampa,nr.confluenceof Rio Palcazu and Rio Iscosacin, 23 Apr 1983 (9 fl-fr), D. N. Smith 3900 (MO).

22. Naucleopsis straminea C. C. Berg, Novon 6: 240, t. 8. 1996. Type. Colombia. Choco: 31 km E of Quibdo,ca. 14 km E of Tutunendo,14 Jun 1982 (9 fl-fr), Gentry et al. 36924 (holotype, COL; isotypes, AAU, BG, JUAM, MO).

Fig. 71

NaucleopThisspeciesis relatedto theAmazonian sis krukovii,fromwhichit differsin thesmallernum-

berof lateralveins(8-13 vs. 15-23pairs)andthelonger andmoreslenderfreepartsof thetepalsof thepistillate flower(subulateand1cmlongin fruitvs. conicaland to 0.7 cm in fruit). Unnamed Collections 1. The collections Ferreiraet al. 10104 andHekker etal. 10088showsimilaritiestoNaucleopsispseudonaga

in thefeaturesof thelamina(as lackingclearlylooplateralveins)andthepistillateinflorescence. connected However,theperidermis flakingoff, thestipulesare ca. 1.5-2.5 cm long,andthe stigmasshort,ca. 3 mm long.Thelaminasoftenhavecircularholes(asnormal thatthesecollecinN. kukovii).Theseareindications tionsbelongto anundescribedspecies,whichmight with includethecollectionGentryetal. 36547,referred some doubt to N. ulei subsp. amara. ECUADOR. SUCUMBlOS: Reserva Faunistica Cuyabeno,nr.LagunaGrande,5 Nov 1987(? fl), Hekker et al. 10088 (U).

288

FLORANEOTROPICA

Fig. 71. Naucleopsis straminea.Leafy twig with fruitingpistillate inflorescence(Gentryet al. 36924). [By HendriekeBerg, Bergen.]

BRAZIL. ACRE:Mun. Mancio Lima, Rio Moa, Serra do Divisor, Fazenda Boa Vista, 15 Oct 1989 (? fl-fr), Ferreira et al. 10104 (NY).

2. Some collectionsfromAmazonianEcuadorcould not be placed in any of the recognized species and almost certainly representa new one. Tree.Leafy twigs

1.5-2.5 mm thick, brownish-puberulousto subhispidulous, dryingred-brown, onmanyormostnodes of theolderbranches aborted lateralbranches; periderm persistent.Lamina oblongto lanceolate,6-18 x 1.24.5 cm,subcoriaceous; apexacuminate to subcaudate; baseacuteto subobtuse; uppersurfaceglabrous;lower

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289

5. ThecollectionLittle6639,collectedfroma shrub, surfaceglabrousor sparselypuberulousat the base; juvenilefeatures,couldnot exhibiting lateralveins12-16pairs,? curvedandloop-connected; andpresumedly petiole0.6-1.1cmlong,ca. beassociated tertiary venationreticulate; speciesknown withoneoftherecognized 1 mm thick, sparselypuberulous,dryingpurplish; fromwesternEcuador.Itbelongsto thegroupof taxa of theleafytwigsflakesoff.The stipules0.6-0.8 cmlong,sparselyappressed-puberu-inwhichtheperiderm intheleafaxilsorbelow oblanceolate lous.Pistillateinflorescences lamina(15-25 x 3-5 cm)hasca.20pairs the leaves,subsessile,ca. 1 cm diam.;bractssemi- of lateralveins anda subcaudateapex;it is sparsely flow- appressed-puberulous orbicular to ovate,0.2-0.6 cmlong,puberulous; on themidribbeneathandthe ers7-13; freepartsof theperianthsubulate, ca.0.5 cm marginis ? revolutetowardsthesubacutebase. long, puberulous;stigmas tongue-shaped,thick, ECUADOR. EL ORO: Piedras, 20 Jun 1943 (st), coarselypapillate. Little 6639 (F, US). ECUADOR. NAPO: ReservaHuaorani,Maxus rd., km 111, 30-31 Aug 1994 (? fl), Dik et al. 1535 (BG, 4 QCNE). SUCUMBtOS: CantonShushufindi,Pucapefna,

Itmightbelongto thesametaxonasthecollections listedunder3.

Dec 1997 (st), E. Freire et al. 2638 (QCNE);Cant6n Shushufindi,San Roque,21 Dec 1997 (st), E. Freireet al. 3061 (QCNE).

V. FICEAE:AN INTRODUCTION 3. ThecollectionsFrancoet al. 1267andGentryet al. 63845resemblethecollectionDiketal. 1535,inthe leavesandtheshortstipules, relatively smallandnarrow butthelamina(somewhat) thickeranditsmarginis (or tendsto be) i revolute.Moreover,the scarsof the stipulesareprominent.

Treesor shrubs, oftenwithaerialroots(in hemi-

monoeciousor gynoepiphytesandroot-climbers), dioecious maleorfemale),withwaxyglan(functionally dularspotson the laminabeneathor on the nodesof inspihairsrare.Leavesalternate, leafytwigs,uncinate laamina orsubverticillate; ralsordistichous, subopposite COLOMBIA. CHOC6: Mun. San Jose del Palmar, entireor incised;venationpinnateto subpalmate to Vrda.La Holanda,1100 m, 14 Jan 1983 (st), Francoet subtrinervate; marginentireordentate;stipulesfully al. 1267 (MO). C6RDOBA: Paramillo National Park, at amplexicaul or semiamplexicaul to lateral.Infloresjct. of Rio TigreandRio Manso,28 Jul 1988 (st), Gen- cences bisexual(andpronouncedly protogynous)or try et al. 63845 (BG, MO). These collectionsbelongto with an urceolate receptacle, unisexual, (functionally) the groupof species in which the peridermof the leaf enclosingtheflowersentirelyduringanthesisandthe twigs is flakingoff. orificetightlyclosedwithbracts,multiflorous, bracte4. The collection Merano et al. 4509, froma tree. ate,thebractsneverpeltate.Staminateflowers:tepals perid- 2-6; stamens1-5;pistillodeabsentorpresent.PistilLeafytwigs 3-5 mmthick,sparselypuberulous; ermflakingoff.Laminaoblongto lanceolate,14-25 x lateflowers free;tepals3-5 (ormore),freeorconnate inlength; 5-7 cm,coriaceous, (always)convex;apexacuminate; (ornearlyabsent);ovaryfree;stylesdifferent base subacuteto rounded;margintowardsthe base stigmas1 or2, equalorunequalin length,? cohering flowers.Fruitanacheneora revolute;bothsurfacesglabrous;lateralveins23-27 withthoseof neighboring with seed embryowithflat small, endosperm; drupelet; terpairs,theloweronesoftenfaintlyloop-connected; and equal cotyledons. ? prominent beneath; petiole tiaryvenationreticulate, 0.5-0.8 cmlong,glabrous;stipulesca. 1.2cmlong,? of onlyonegenus,Ficus.ThemonoThetribeconsists curvedto slightytwisted,(very)sparselypuberulous typicNewCaledonian whichhas genusSparattosyce, ca.5 cmdiam.; beenincludedin thetribeby Corner(1962),hasmuch atthebase.Infructescences(sub?)sessile, involucral bractsin ca.7 rows,broadlyovate,sparsely incommon NewGuinean withthemonotypic genusAntito ca. aropsis, andit cannotbereadilyassociatedwithFicus, freepartsof thetepalspyramidate, puberulous; 0.5 cmlong,sparselypuberulous. in spiteof thepresenceof anurceolatereceptacle. rd. ECUADOR. LOJA: ParqueNacionalPodocarpus, to Zamora,San Francisco,2100 m, 4 May 1995 (q fr), Merano et al. 4509 (LOJA).

This collectionapparently representsa montane speciesbelongingto thegroupof speciesinwhichthe holesinthelamina(caused flakesoff.Circular periderm by somefungus),as commonlyfoundin Naucleopsis alsooccurin thematerialof thistaxon. krukovii,

19. Ficus Linnaeus,Sp. P1. 1059. 1753;Miquelin Martius,Fl. Bras.4(1):83-106;Burger,Fieldiana Darwiniana Bot.40: 140-187.1977;VdzquezAvila, 23(3/4):605-636. 1981;Berg,VazquezAvila& Kooy,ActaAmazonica14(1/2,Supl.):159-194. 1-365. 1989;Berg, 1984;Carauta,Albertoa2(1): Fl. Guianas,Ser.A, Fasc.11:35-63. 1992.

290 UrostigmaGasparrini,Nov. Gen 7. 1844. Miquel,LondonJ. Bot. 6: 525. 1847. Pharmacosycea

FLORANEOTROPICA

diversityof the genus Ficus createsa situationof withtheothergenerain Moraceaewith inequivalence compariandphytogeographical regardto systematical sons.Thesituationcanbe easedby recognitionof adequatesubdivisionswithinFicus,whichcanbe then genera.Suchsubdiviwithothermoraceous compared sionsornaturalgroupsof specieshavebeenindicated by Berg(1989a, 1998b). Themostspecioseentitycomprisesca.280monospecies(placedin hemiepiphytic eciousandessentially in Corner's about classification): subgenusUrostigma 100 speciesareneotropical(andareplacedin sect. Americana). Anothergroupof monoeciousspeciesis andcomprisesabout75 species essentiallyterrestrial in Corner's clas(includingsubgenusPharmacosycea (andplaced about20 of theseareneotropical sification);

and or hemiepiphytic Treesor shrubs, terrestrial with aerialroots,monoecious, thenwithanastomosing waxyglandularspots,usuallyoneortwo,at thebase of thelaminabeneath;uncinatehairslacking.Leaves marvenation inspirals; laminaentire; pinnate; alternate, Inflorescences gin entire;stipulesfully amplexicaul. withthereceptacle(syconium)subtendedby a whorl (ostiole)cirof 2 or3 (basal)bracts,theapicalaperture cularwith interlocking(ostiolar)bracts;interfloral to several; bractspresent.Staminateflowersnumerous tepals2-6, freeorconnate;stamens1 or2; pistillode sessile presentorabsent.Pistillateflowers numerous, tepals2-4, freeorconnate;stylesof diforpedicellate; ferent length in the same inflorescence;stigmas in sect. Pharmacosycea). Whether sect. Pharmaco2 or 1.Fruita smallachene. (sub)filiform, sect.Oreosycea(Miquel) sycea andthepaleotropical Someof the OldWorldspecies(of whichseveral canbe keptin a single related and are Corner really intotheNeotropicsandsomeof havebeenintroduced doubtful (cf.Herreet al., 1996). is somewhat subgenus species)deviatefromthe asneotropical themdescribed largely(gyno)the comprises of genus The remainder basedontheneotropical representapresentdescription 15 theSycomorus dioecious only species (of species; tivesof thegenus.IntheOldWorldtheplantscanbe of In the (gyno)dioecigroup monoecious. group) are oftenroot-climbers; spotscan waxyglandular climbers, the occurin variouspositionson thelowersurfaceof the ousspeciesmajorsubdivisionscanbe recognized: lamina,e.g.,inaxilsof lateralveinsotherthanthebasal Ficusgroup(largelysect.Ficus),withabout60 species two of veinsor(also)onthenodes of treesorshrubs;theSynoeciagroup,comprising onesand/orinfurcations sect. as Kalosyce of root-climbers (known groups of leafy twigs;the leaves can be distichous,(sub)with Endlicher), andsect.Rhizocladus thestipulescan (Miquel)Corner oppositeorsometimessubverticillate; a the group (comprisSycidium total 80 species; about at funcbe lateral;theplantsareoftendioecious, least containingeither ing sect. Sinosycidium Corner and sect. Sycidium tionally"male,"withinflorescences staminate flowersand(non-functional) pistillateflow- Miquel),withabout100speciesof shrubs,trees,and erswithshortstylesor (functional)pistillateflowers climbers;andtheSycomorusgroup(includingsubg. withlong styles(andneuterflowers);theostiolecan Sycomorus (Gasparrini)Miquel, sect. Adenosperma bractscanbelacking,theouter Corner,sect.NeomorpheKing,andsect.Sycocarpus interfloral beslit-shaped; bracts,a distinct Miquel),withabout150 speciesof treesandshrubs, canbear(lateral) surfaceof receptacle (geocarpic). orflagelliflorous whorlof basalbractscanbe lacking;theperianthcan manyof themcauliflorous Thetwoneotropicalsubdivisionsof Ficusspecies be almostlacking;thenumberof stamenscanbe more withpaleotropical entitiesconcenthantwo;andthefruitletscanbe distinctlydrupaceous. showrelationships reThe genus comprisesapprox.750 species,with tratedin theeasternpartof theAsian-Australasian flora African Ficus Distinct connections with the gion. 105inAfrica, 500-550 speciesinAsiatoAustralasia, arelacking.Thisfactappearsto bereflectedin thedisandapprox.120in theNeotropics. The tremendousdiversity,not only in vegetative tributionof thegenusin theNeotropics.Incontrastto structures,but also in reproductiveones, inflores- thegroups(tribes)showingdistinctrelationswiththe flora,Ficusis moreclearlyassocirecognition Africanmoraceous cences,flowers,andfruits,couldwarrant Americaandthenorthern Andeanreinthemiddleof atedwithCentral of manygenera,ashasbeenattempted the past century(Gasparrini,1844, 1845; Miquel, gionthanwiththe SouthAmericancontinent,where centerscanberecognized:easternBra1847-1848).However,in 1867Miquelagainmerged twosecondary region. inthesinglegenusFicus,forwhich zil andtheGuiana(orGuayana) allgenerarecognized Thetribe(andgenus)has a distinctset of characthe most recentsubdivisionhas been proposedby Corner(1965).Thissubdivisionneedssomeremodel- ters,settingit clearlyapartfromtherestof theMoraling(Berg,1989a,1998).Inadditionto morphological ceae.Mostof thesetraits,if not all, arerelatedto the of thepollinatingfig uniquepollinationsystem. theclassification considerations, wasps,as proposedby Wiebes(1994), indicatesthe Theinflorescence (syconium,fig)hasanurceolate Thesizeand receptacle needto reconsider Corner's classification. Theflowtheflowersatanthesis. fullyenclosing

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ersareonlyaccessibleto thetinypollinating fig wasps Dueto thisphenologicalfloweringpatternthegenus (andsomeotherinsects)whichmanageto getthrough ensuresa continoussupplyof maturesyconia,foodfor theostiole.Theostioleorentranceto thefig cavityis many(arboreal) animalsin manyhabitats. barred by (interlocking) bracts.Thepollinators usually Manypaleotropical specieshavebeenintroduced lose theirwings(anddamagetheirantennae) on their in theNeotropicsas omamentals.Mostof the introwayintothecavityof thesyconium,andtheretheyre- ducedspeciesbelongto thesubgenusUrostigma. Bemainencaged. causeof theabsenceof pollinators, introduced species Thestylesof thepistillateflowersaredifferentin (e.g.,asornamental trees)donotreproduce. However, length(heterostyly). Inthegroupsof monoeciousspe- inseveralplaces,e.g.,inBermuda Islands,Cuba,Florida cies (to whichallneotropicalspeciesbelong)all figs (Ramirez& Montero,1988;Kaufmannet al., 1991; containstaminate andpistillate flowers,andheterostyly Nadelet al., 1992),andRiode Janeiro(deFigueiredo to berealizedthroughandduringtheontogeny etal., 1992),severalspeciesintroduced fromAsia(e.g., appears of thesyconium.Intermediate stylelengthsoccurand Ficus altissimaBlume andE microcarpaLinnaeusf.) theovariesof thedenselypackedpistillateflowersare havestartedto reproduce,as thepollinatorshavererankedindifferentlayers.Dueto differences in length centlyarrived(byplane?)andbecomeestablishedin of thepedicelor alsoof thelength(andshape)of the placeswithsufficientlylargestandsof thesespecies. Thepresentstateof taxonomicknowledgeof the in onelayeratanthesis ovary,thestigmasarearranged indications of thenumber andcohereintoa synstigmatic layer.Thissynstigma is genusallowsratheraccurate anessentialrequisiteforthepollination apparently sys- of speciesforthetwo neotropicalsectionsof thegeis reasonably well tem(as brieflydescribedin the Introduction). In the nus.Formostspeciesthedistribution by regroupsof dioeciousspecies(all confinedto the Old known.Thisknowledgehasbeenaccumulated World)partof thetrees("male"ones)bearfigs con- gionalstudiesof thegenussuchasfortheFloraMesoarea(Berg,unpubl.)andtheCaribbean retainingstaminateflowersand short-styledflowers, americana inaddition tofloristicaccounts whereasothertrees("female" ones)bearfigswithonly gion(Berg,unpublished) long-styledflowers(andneuterflowerin positions of FicusforCostaRica(Burger,1977),Trinidadand wherestaminate flowerscouldbe expected).Atanthe- Tobago(Berg&vanHeusden,1982),theNetherlands sis theovariesarearranged in a singlelayerto produce Antilles(Berg, 1980),Venezuela(Berg& Simonis, (Berg,1992),andBrazilianAmazonia a coheringlayerof stigmas.However,thedeveloping 2000),theGuianas et (Berg al., 1986), complemented by studieson the fruitsmaybe arranged in morethanonelayer. species ofArgentina (VizquezAvila, 1981)andon speTheanthesisof thestaminate flowersis delayeduntil thefruits(seeds)areripe;thisis characteristic forthe ciesof Brazil(Carauta,1989). Someconsiderable taxonomicproblemsawaitfurgenusandessentialforthepollinationsystem. Smallwaxyglandular spotsareomnipresent in the therstudy.Theyarelargelyrelatedtosomespeciescomgenusandarefoundatvariouspositionsontheleaves plexes,themostprominentandwidespreadonesbeentities complex,comprising andin severalOldWorldgroupsevenontheleafytwigs. ing:theFicusamericana F F provisionally identified as americana Aublet, andiTheyareprobably buttheirfuncbiologically important, F F cola DesStandley, Dugand, guianensis greiffiana tion is unknown.Theymightplaya role in emitting substances forattracting thepollinators atlongdistance. vaux, F mathewsii(Miquel) Miquel, andF subapicuThesmallseedscanremaindormant foralongtime. lata (Miquel)Miquel;andthe F citrifoliacomplex, entitiesprovisionally identifiedasF amaLightis (mostly)neededforgermination. Theproper- comprising F zonica brittonii (Miquel) Miquel, Boldingh,F citritiesof theseedsmakethemsuitableforlong-distance F F eximia folia Miller, dugandii Standley, Schott,and inremoteplaceswill dispersal. However,establishment F subandina Dugand. behampered of treeslarge bytheneedof apopulation enoughto sustainthepollination systembasedona sperelation. cies-specificplant-animal 19A. Ficus Linnaeussubg.Urostigma(Gasparrini) Thegeneralpattemof thephenologyof monoecious Miquelsect.AmericanaMiquel,LondonJ.Bot.6: (thusof allneotropical) speciesis to floweryear-round 525. 1847. toprovidecontinuous breedingsitesforthepollinators. Ficus Linnaeussubg. Urostigma(Gasparrini) Miquel, Thesetinyinsectslive foronlya fewdaysandhaveto Ann. Mus. Bot. Lugduno-Batavum 3: 260. 1867. withpistillateflowersatanthesisto findinflorescences UrostigmaGasparrinisect. AmericanaMiquel,Lonlay theireggs (andcarryoutpollination)withinthat don J. Bot. 6: 525. 1847. shortperiod.Individual treeshavedifferentflowering oftenwith t constantintervalsin flowerprorhythms, Treesor shrubs, (basically) hemiepiphytic; waxy is largelysimultaneous. glandularspot 1, at the base of the midribbeneath.Induction;theflowerproduction

292

FLORANEOTROPICA

onshort- den)kindlycheckedtheLatindiagnosisandMireya florescences usuallyinpairs(ormoretogether with2 basalbracts; stami- Correa(SmithsonianTropicalResearchInstitute, subtended shoots);receptacle aboutlocalitiesin the nateflowerswith 1 stamenandwithouta pistillode; Panama)providedinformation formerCanalZone.Theauthoris muchindebtedtothe pistillateflowerswith 1 stigma. W.C.Burger(FieldMuof thiscontribution, reviewers Thissubdivisioncomprisesabout100species.The Chicago),J.Kallunki(NewYorkBotanicalGarseum, pollinatingfig waspsbelongto thegenusPegocapus. den),andM. Nee (NewYorkBotanicalGarden),for carefulandcriticalreadingof thetextandforvarious toimprovethecontents.For andsuggestions 19B.FicusLinnaeussubg.Pharmacosycea(Miquel) comments material of thefollowinghercontribution the present Miquelsect.Pharmacosycea. bariahas been examined:A, AAU,B, BG, BHMH, Ficus Linnaeus subg. Pharmacosycea (Miquel) Miquel, Ann. Mus. Bot. Lugduno-Batavum3: BOLV,BR,C, CAS,CAY,CEPEC,COL,CR,CTES, CUVC, DUKE, EAP, ECON, F, FI, G, GB, GH, 299. 1867. Ficus Linnaeus subg. Pharmacosycea (Miquel) GOET,GUA,GUAY,H, HB,HBR,HUA,IAN,ICN, Miquelsect. Pharmacosycea;Corner,Gard.Bull. IJ,INPA,IPA,ITA,JAUM,JE,L, LAM,LD,LE,LIL, StraitsSettlem. 17: 406. 1960. LL,LOJA,LP,LPB,M, MA,MBM,MEDEL,MER, MG,MICH,MOL,MPU,MY,NA, NY, 0, OXF,P, withoutaerialroots;waxy Trees,(mostly)terrestrial, PEL,PMA,PSO,Q, QAME,QCA,QCNE,R, RB, spots2, in theaxilsof thebasallateralveins glandular RSA, S, SCZ, SELBY,SI, SP, SPF, TEX, TRIN, (ormoretogether beneath. Inflorescencesusuallysolitary TULV,TUR,U, UB,UDBC,US,USM,USZ,VALLE, on short-shoots); receptaclesubtendedwith 3 basal WIS,Z. Theauthoris indebtedto flowerswith2 stamensandapistillode VEN,VIC,WAG, staminate bracts; their and curatorsfor makingthe material directors oftenpresent;pistillateflowersusuallywith2 stigmas. availableduringvisitsand/orby loans. Thissubdivisioncomprisesabout20 species.The pollinatingfig waspsbelongto thegenusTetrapus.

LITERATURECITED Allemao, F. F. 1857. Artocarpeae.Soaresia nitida - spec.

ACKNOWLEDGMENTS

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Theauthoris indebtedfordirectandindirectfinanFoundation cialsupportprovidedby theNetherlands the forAdvancement of TropicalResearch(WOTRO), novo. Revista Brazil. 1: 368-370. forAdvancement of PureReNetherlands Organization Andrade,J. Cardosode. 1984.Observac6espreliminares search,andthe NorwegianCouncilfor Scienceand sobrea eco-etologiade quatrocoleopteros(Chrysoof Humanities (NAVF),whichallowedaccumulation melidae,Tenebrionidae,Curculionidae)que dependem da Embauba (Cecropia lyratiloba var. nana) andbyfieldworkduringapedatabyvisitsto herbaria na restingado Recreiodos BandeiranCecropiaceae, riodof about20 years.Thefacilitiesprovidedby The tes, Rio de Janeiro.RevistaBrazil.Ent. 28: 99-108. New YorkBotanicalGardenin the periodOctober Andresen, E. 1999. Seed dispersal by monkeys and 1996-March1997finalizedthepresentstudiesandare the fate of dispersedseeds in a Peruvianrainforest. Theauthoris verymuchingratefullyacknowledged. Biotropica31: 145-158. debtedto J.P.P.Carauta (Riode Janeiro),whogener- Aublet,J. B. C. F. 1775. Histoiredes plantesde Guiane Didotjeune, Paris. Francaise.Pierre-Fran9ois ouslygaveaccessto a wealthof materialandinformaH. E. 1860. Note sur une nouvelle espece du tion regardingBrazilianDorstenia speciesandwho Baillon, genre Sorocea. Adansonia1: 212-213, t. 6. shouldhavebeenco-authorfortheDostenia section . 1875a.Ulmacees.Hististoiredes plantes6: 137havebeenmade hadhe wishedto be.Theillustrations 216. L. Hachette,Paris. . 1875b. Stirpes exoticae novae. Adansonia 11: by HendriekeBerg(Bergen),J. Brinkman(Utrecht), 292-312. E. M.HupkensvanderElst(Utrecht),andP.Pardoen H. & S. S. Renner.1989. Diversityof EcuadorTheOlafGrolleOlsensFundof theUniver- Balslev, (Utrecht). ian forests.In L. B. Holm-Nielsen,I. C. Nielsen & of severalof sity of Bergenfinancedthepreparation H. Balslev (eds.), Tropicalforests- biologicaldyThelistof collectionscitedforadditheillustrations. namics,speciationand diversity:287-295. Academic Press, London. hasbeenprepared tionsandcorrections byAxelHansen (Bergen).TheauthorthanksElviraBalslev(Aarhus) Barker,H. D. & W. S. Dardeau.1930. Moraceae.In H. D. Barker,Flored'Haiti.Dept. Agr., Port-au-Prince. andJ.P.P.Carauta forthetranslations Bawa, K. S. & P. A. Opler. 1977. Spatialrelationships (RiodeJaneiro) of the abstractinto SpanishandPortuguese,respecbetweenstaminateand pistillateplantsof dioecious tropicalforest species. Evolution31: 64-68. tively.RupertBameby(TheNewYorkBotanicalGar-

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LITERATURE CITED GattungDorstenia.J. Bot. (Schrader)2: 238-239. . 1821. Neue Entdeckungenim ganzen Umfang der Pflanzenkunde2. FriedrichFleischer,Leipzig. . 1822. Neue Entdeckungenim ganzen Umfang der Pflanzenkunde3. FriedrichFleischer,Leipzig. . 1826.Systemavegetabilium.Ed. 16. 3. Dietrich, Gottingen. phaneroStandley,P.C. 1919.Studiesof tropicalAmerican gams- no. 3. Contr.U.S. Natl. Herb.20: 172-220. . 1929. Studies of Americanplants - II. Publ. Field Mus. Nat. Hist., Bot. Ser.4: 301-345. . 1929. A new species of Sorocea from Colombia. Trop.Woods 19: 39. . 1933. Poulsenia,a genus of trees of the family Moraceae.Trop.Woods 33: 4-5. . 1935.A new SoroceafromBrazil.Trop.Woods 43: 18. . 1940. Studiesof Americanplants- IX. Publ. Field Mus. Nat. Hist., Bot. Ser.22: 3-62. . 1943. Moraceae.In R. F. Woodson& R. W. Schery, Contributionstowarda Flora of Panama. VII. Miscellaneouscollections, chiefly by H. von Wedel,in Bocas del Toro.Ann. MissouriBot. Gard. 30: 85. & J. A. Steyermark.1944. Studies of Central Americanplants- IV. Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 31-109. & L. 0. Williams. 1950. Plantas nuevas hondurenias y nicaraguenses.Ceiba 1: 74-96. . 1952. Ocho generos de arboles y & arbustosnuevosparaCentroAmerica.Ceiba3: 24-35. Steudel,E. G. 1840. Nomenclatorbotnanicus.Ed. 2, 1. J. G. Cotta,Stuttgart. . 1841. Nomenclatorbotanicus.Ed. 2, 2. J. G. Cotta, Stuttgart. Stokes,J. 1812.A botanicalmateriamedica4. J. Johnson, London. Swartz,0. P. 1788. Nova generaet species plantarum prodromus.M. D. Holmiae, Stockholm. Tenore, M. 1845. Catalogo Orto botanico di Napoli. Typografiadell Aquiladi V. Puzziello,Napoli. Trecul,A. 1847. Sur la famille des Artocarpees.Ann. Sci. Nat. Bot., Ser. 3, 8: 114-172. Urban,I. 1905. Floraportoricensis.Pp. 193-352 in I. Urban,Symbolaeantillanae4. Borntraeger,Berlin. . 1918. XXIV. Sertum antillanumz.V. Feddes Repert.Spec. Nov. Regni Veg. 15: 156-171. . 1929. Moraceae.Plantaehaitensieset domingenses novae vel rarioresVI. a cl. E. L. Ekman 1924-1928 lectae. Ark. Bot. 22A(10): 1-9. & E. L. Ekman.1929. In I. Urban,Moraceae. Plantaehaitensieset domingensesnovaevel rariores VI. a cl. E. L. Ekman1924-1928 lectae. Ark. Bot. 22A(10): 1-9. Valdeyron,G. & D. G. Lloyd. 1979. Sex differences and floweringphenologyin the commonfig, Ficus

299 carica L. Evolution 33: 673-685. Valente,M. da C. & J. P. P. Carauta.1974. Dorstenia brasiliensisLamarck(Moraceae).Estudosanatomicos e taxonOmicos. An. XXV Congr. Nac. Bot. Mossoro: 89-94. & . 1975. Dorstenia bahiensis Klotzsch ex Fischeret Meyer(Moraceae),considerac6estaxonomicase anatomicas.Ci. & Cult. 27(6): 645-650. & . 1977. Comparaqio das especies brasileirasde Dorstenia(Moraceae).Trab26 Congr. Nac. Bot., Rio de Janeiro:597-633. & 0. M. Barth.1977.Comentarios sobre 9 algumas especies de Dorstenia L. (Moraceae)da Americado Sul Meridional.Revista Brasil. Biol. 37(1): 167-173. & J. M. A. Braga.1998.Dorsteniaerecta ,Vellozo (Moraceae).Caiapia-erecto.Anatomiafoliar e taxonomia.Albertoa4(25): 329-333. VazquezAvila, M. D. 1981. El genero Ficus (Moraceae) en la RepublicaArgentina.Darwiniana23(3/ 4): 605-636. . 1985. Moraceasargentinas,nativasy naturalizadas. Darwiniana26: 289-330. Vellozo, J. M. da Concei9ao.1829. Flora fluminensis. TypographiaNationali,Rio de Janeiro. . 1831.Florafluminensisicones. Senfelder,Paris. Walpers,W. G. 1843, Moraceae.Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol.Nat. Cur. 19(Suppl. 1): 42 1-422. . 1848-1849. Annalesbotanicessystematicae1. FridericiHofmeister,Leipzig. Warburg, O., 1913.Urticacees.In A. F. M. Glaziou,Liste des plantes du Bresil Centralreceuillies en 18611895. Bull. Soc. Bot. France59(Mem.3): 639-647. Wawra,H. 1866.BotanischeErgebnisseder Reise seiner Majestatdes Kaisersvon MexicoMaximilianI. nach Brasilien.Verlagvon CarlGerold'sSohn, Wien. Weddell,H. A. 1856-1857. Monographiede la famille des Urticacees.Arch.Mus. Hist. Nat. 9: 1-592. . 1869.Urticaceae.In A. P. De Candolle,Prodromus systematis naturalis 16(1): 32-235(1-64). Treuttelet Wiirtz,Paris. Welle, B. J. H. ter, K. Bonsen & P. Detienne. 1992. Woodand timber.Moraceae.In A. R. A. Gorts-van Rijn (ed.), Floraof the Guianas,Ser. A., Fasc. 11: 141-191. Koeltz Scientific Books, Koenigstein. Wheeler,W. M. 1942. Studieson neotropicalant-plants and theirants. Bull. Mus. Comp.Zool. Harvard90: 1-252. Agaoninae(polWiebes,J. T. 1994.The Indo-Australian linatorsof figs). Verh.Kon. Ned. Akad.Wetensch., Afd. Natuurk.,TweedeSect., 92. Willdenow,C. L. 1789. Species plantarum1(2). G. C. Nauk, Berlin. . 1806. Speciesplantarum4. G. C. Nauk,Berlin. Woodson,R. E. & R. W. Schery1960. Floraof Panama, Moraceae.Ann. MissouriBot. Gard.47: 114-178.

300

FLORANEOTROPICA

Fig. 72. Distributionof taxa. 1. Morusceltidifolia.Opencircles,native;solid circles, introduced.2. M. insignis. 3. Trophis mexicana.

DISTRIBUTIONMAPS

301

T involucrata. oftaxa.1.Circles,Ihophis Fig.73. Distribution 3. T caucana.4.Maclura cuspidata; 2. T racemosa. triangles, brasiliensis.

302

FLORANEOTROPICA

of taxa.1. Maclura tinctoria subsp.tinctoria. 2. M. tinctoria subsp.mora. 3. Triangles,Bagassa Fig. 74. Distribution guianensis; circles,Batocarpus costaricensis.

DISTRIBUTIONMAPS

303

Fig. 75. Distribution of taxa. 1. Triangles, Clarisia biflora; circles, C ilicWolia. 2. Batocarpus orinocensis. 3. B.

amazonicus.

FLORA NEOTROPICA

304

le~~~~~~~~~~o

Fig.

76.

Distribution

of

taxa.

1.

4.S.guilleminiana. 3.Sorocea racemosa. 2.Clarisia hilarii. Sorocea bonplandii.

DISTRIBUTIONMAPS

F~~~~~~~~~~~~

305

-

Fig. 77. Distributionof taxa. 1. Triangles,Soroceapubivena subsp.pubivena; circles, S. pubivena subsp. oligotricha. 2. Circles, S. steinbachii; triangles, S. muriculata subsp. uaupensis; crosses, subsp. uaupensis vel aff. 3. S. muriculata subsp.

muriculata.4. S. briquetii.5. Triangles,S. affinis;circles,S. sarcocarpa.6. S. trophoides.7. S.pubivenasubsp.hirtella.

306

FLORANEOTROPICA

I

rm~~~~~~~~~~r

of taxa.1. Circles,Sorocearuminata; triangles,S.jaramilloi.2. Semi-solidcircles,S.spruceisubsp. Fig.78. Distribution triangles,subsp.saxicola;solidhexagon,bothsubsp.saxicolaandsubsp.sprucei; sprucei;crosses,subsp.subumbellata; opencircles,S. duckei.3. Circles,Poulseniaarmata;triangle,Poulseniasp.

307

MAPS DISTRIBUTION

AAX

-

.

N,~~~~~~~~~~~~~ ....

of taxa.1. Dorstenia grazielae. 2. D. arifolia. 3. Circles,D. ramosa subsp.ramosa; triangles,D. Fig. 79. Distribution bonijesu. 4. Circles,D. ramosa subsp.dolichocaula; triangle,D. conceptionis. 5. Circles,D. appendiculata; triangle,D. brevipetiolata. 6. D. turnerifolia. 7. D. hirta. 8. D. bowmaniana. 9. D. setosa. 10. Circles,D. aristeguietae; triangles,D. colombiana; solidhexagon,D. panamensis. 11.D. lindeniana. 12. Circles,D. milaneziana; triangles,D. carautae. 13.Triangle,D. uxpanapana; semi-solidcircles,D. choconiana; opencircles,D. choconiana vel aff. 14. Circles,D. umbricola; triangles,D. peruviana. 15. D. drakena. 16. Semi-solidcircles,D. bahiensis; open circles,D. urceolata; triangle,D. hildegardis; cross,D. albertii.

308

FLORA NEOTROPICA

4. of taxa.1. Dorsteniatenuis.2. D. cayapiasubsp.cayapia.3. D. cayapiasubsp.paraguariensis. Fig. 80. Distribution D. cayapiasubsp.vitifolia.5. D. cayapiasubsp.asaroides.6. D. brasiliensis.

DISTRIBUTION MAPS

----

309

--

-

-

----

-

Fig. 81. Distribution of taxa.1. Semi-solidcircles,Dornteniafawceuifi;triangles,D. nwmmularia;opencircles,D. peltata. 2. Circles,D. petraea; triangles,D. crenulata. 3. Circles,D. caimitensis; triangle,D. tubemosa.4. Circles,D. elata; triangles,D. contensis. 5. D. excentrica. 6. Circles,D.flagellifera; triangles,D. mocana.7. Semi-solidcircles,D. erythrandra; triangle,D. jamaicensis; opencircle,D. roigii; cross,D. belizensis. 8. D. contrajerva.

310

FLORA NEOTROPICA

amazonicus. 4. Circles, sprucei.3. Ikymatococcus Fig.82. Distibution oftaxa.1. Perebeamennegae.2. Helianthostylis Brosimum multinervium; triangles, Helianthosnylis steyermarkii. 5. Perebeaangustifolia.6. Circles,Pseudolmedia macrophylla;triangles, P. gentryi; hexagon, P manabiensis.

DISTRIBUTION MAPS

311

pA~~~~~~~~~~~~

) rubrasubsp.glabrifolia.2. Circles,P of taxa.1. Circles,Perebearubrasubsp.rubra;triangles, Fig.83. Distribution ) mollissubsp.lecithogalacta. mollissubsp.mollis;triangles, 3. Pseudolmedia laevigata.4. Triangles, Naucleopsisuleisubsp. subandina;circles,N. uleisubsp.puberula;hexagons,N. humilis.5. Hexagons,N. velutina;circles,N. chiguila;triangles, N. straminea.6. Circles,N. herrerensis; triangles,N. nagasubsp.meridionalis.

312

FLORANEOTROPICA

of taxa.1. Circles,Naucleopsis naga subsp.naga; triangles,Helicostylis turbinata. 2. Triangles, Fig. 84. Distribution Naucleopsis capirensis; hexagonandcircles,N. capirensis vel aff. 3. Triangles,Maquira guianensis subsp.guianensis; circles,M. guianensis subsp.costaricana.

NUMERICAL LISTOFTAXA

313

NUMERICALLIST OF TAXA 1.Morus Linnaeus 1-1. M.celtidifoliaKunth 1-2. M. insignis Bureau 2. TrophisLinnaeus 2-1. T mexicana (Liebmann) Bureau 2-2. T cuspidata Lundell 2-3. T racemosa (Linnaeus) Urban 2-4. T caucana (Pittier) C. C. Berg 2-5. T involucrataW.Burger

3. Maclura Nuttall 3-1.M. brasiliensis (Martius)Endlicher 3-2.M. tinctoria (Linnaeus) Steudel a. subsp. tinctoria b. subsp. mora (Grisebach)VizquezAvila

4. Bagassa Aublet 4-1. B. guianensis Aublet 5. Batocarpus Karsten 5-1. B. costaricensis Standley & L. 0. Williams 5-2. B. amazonicus (Ducke) Fosberg 5-3. B. orinocensis Karsten 6. Clarisia Ruiz & Pav6n 6-1. C. biflora Ruiz & Pavon 6-2. C. racemosa Ruiz & Pavon 6-3. C. ilicifolia (Sprengel) Lanjouw & Rossberg

7. Sorocea Saint-Hilaire 7-1. S. guilleminiana Gaudichaud 7-2. S. bonplandii(Baillon)W.Burger,Lanjouw&WesselsBoer 7-3. S. hilarii Gaudichaud 74. S. pubivena Hemsley a. subsp.pubivena b. subsp.oligotricha(Akkermans& C. C. Berg)C. C. Berg c. subsp. hirtella (Mildbraed)C. C. Berg 7-5. S. steinbachii C. C. Berg 7-6. S. muriculata Miquel a. subsp. muriculata b. subsp. uaupensis (Baillon) C. C. Berg 7-7. S. briquetii Macbride 7-8. S. trophoidesW.Burger

7-9. S. affinisHemsley 7-10. S. sarcocarpa Lanjouw& Wessels Boer 7-11. S. ruminataC. C. Berg 7-12. S. jaramilloi C. C. Berg 7-13. S. sprucei (Baillon) Macbride a. subsp.sprucei b. subsp. saxicola (Hassler) C. C. Berg c. subsp. subumbellata C. C. Berg 7-14. S. duckei W.Burger

8. Poulsenia Eggers 8-1. P. arnata (Miquel) Standley

8-2.P sp. 9. Dorstenia Linnaeus 9-1.D. appendiculata Miquel 9-2. D. turnerifolia Fischer & Meyer 9-3. D. hirta Desvaux 9-4. D. brevipetiolata C. C. Berg 9-5. D. contensis Carauta& C. C. Berg

9-6. D. bowmaniana J. G. Baker 9-7. D. carautae C. C. Berg 9-8.D. milaneziana Carauta,Valente & Sucre 9-9.D. setosa Moricand 9-10. D. choconiana S. Watson 9-1 1. D. uxpanapana C. C. Berg & T. Wendt 9-12.D. umbricolaA. C. Smith 9-13. D. peruviana C. C. Berg 9-14. D. aristeguietae Cuatrecasas 9-15.D. colombiana Cuatrecasas 9-16. D. panamensis C. C. Berg 9-17. D. lindeniana Bureau 9-18.D. elata Hooker 9-19.D. urceolata Schott 9-20 D. hildegardis Carauta,Valente & Barth 9-21. D. bahiensis Fischer & Meyer 9-22. D. grazielae Carauta,Valente & Sucre 9-23. D. albertii Carauta,Valente & Sucre 9-24. D. bonijesu Carauta& Valente 9-25. D. arifolia Lamarck 9-26. D. ramosa (Desvaux) Carauta,Valente & Sucre a. subsp. ramosa b. subsp. dolichocaula (Pilger) C. C. Berg 9-27. D. contrajerva Linnaeus 9-28. D. drakena Linnaeus 9-29. D. conceptionis Carauta 9-30. D. cayapia Vellozo a. subsp. cayapia b. subsp.paraguariensis (Hassler)C. C. Berg c. subsp. vitifolia (Gardner)C. C. Berg d. subsp. asaroides (Hooker) C. C. Berg 9-31. D. tenuis Bureau 9-32. D. brasiliensis Lamarck 9-33.D. excentrica Moricand 9-34. D. fawcettii Urban 9-35. D. petraea Grisebach 9-36. D. jamaicensis Britton 9-37. D. flagellifera Urban & Ekman 9-38.D. belizensis C. C. Berg 9-39. D. roigii Britton 9-40.D. nummulariaUrban & Ekman 9-41.D. peltata Sprengel 9-42. D. erythrandraGrisebach 9-43.D. caimitensis Urban 9-44.D. crenulata Grisebach 9-45.D. tuberosaGrisebach 946. D. rocana Britton ADDITIONSAND CORRECTIONS

10.Brosimum Swartz 10-1. B. alicastrum Swartz a. subsp. alicastrum b. subsp. bolivarense (Pittier) C. C. Berg 10-2. B. acutifolium Huber a. subsp. acutifolium b. subsp. interjectumC. C. Berg c. subsp. obovatum(Ducke) C. C. Berg 10-3. B. lactescens (S. Moore) C. C. Berg 10-4. B. costaricanum Liebmann 10-5. B. guianense (Aublet) Huber

FLORANEOTROPICA

314 10-6. B. gaudichaudii Trecul 10-7. B. glaziovii Taubert 10-8. B. glaucum Taubert 10-9. B. rubescens Taubert

C. C. Berg 10-10.B. melanopotamicum 10-1 1. B. utile (Kunth) Pittier a. subsp. utile b. subsp. occidentale C. C. Berg c. subsp. allenii (Woodson) C. C. Berg d. subsp. magdalenense C. C. Berg e. subsp. darienense C. C. Berg f. subsp. ovatifolium (Ducke) C. C. Berg

10-12.Brosimum longifoliumDucke C. C. Berg 10-13.B. multinervium 10-14. B. potabile Ducke 10-15. B. parinarioides Ducke

a. subsp.parinarioides b. subsp. amplicoma (Ducke) C. C. Berg

a. subsp.rubra b. subsp.glabrifolia (Ducke)C. C. Berg 15. MaquiraAublet 15-1.M. guianensis Aublet a. subsp.guianensis b. subsp.costaricana (Standley)C. C. Berg 15-2. M. scierophylla (Ducke)C. C. Berg 15-3.M. calophylla (Poeppig& Endlicher)C. C. Berg 15-4. M. coriacea (Karsten) C. C. Berg

16.Helicostylis Trecul 16-1.H. tomentosa (Poeppig& Endlicher)Rusby 16-2.H. scabra (Macbride) C. C. Berg 16-3.H. elegans (Macbride)C. C. Berg 16-4.H. pedunculata Benoist 16-5. H. turbinata C. C. Berg

16-6.H. heterotricha Ducke 16-7.H. tovarensis (Klotzsch& Karsten)C. C. Berg 17. Castilla Sesse

11. TrymatococcusPoeppig & Endlicher 11-1. T amazonicus Poeppig & Endlicher 11-2. T oligandrus (Benoist) Lanjouw

17-1.C. elastica Sesse a. subsp.elastica

12. Helianthostylis Baillon 12-1. H. sprucei Baillon

b. subsp. costaricana (Liebmann) C. C. Berg c. subsp. gummifera (Miquel) C. C. Berg 17-2.C. uleiWarburg

12-2.H. steyermarkii C. C. Berg Trecul 13.Pseudolmedia 13-1. P laevis (Ruiz & Pavon) Macbride

13-2.P hirtulaKuhlmann Trecul 13-3.P macrophylla 13-4.P laevigataTrecul 13-5. P glabrata (Liebmann)C. C. Berg

13-6.P manabiensisC. C. Berg 13-7. P rigida (Klotzsch & Karsten)Cuatrecasas a. subsp. rigida b. subsp. araguensis C. C. Berg c. subsp. eggersii (Standley) C. C. Berg 13-8. P spuria (Swartz) Grisebach

13-9.P gentryiC. C. Berg 14.PerebeaAublet 14-1. P guianensis Aublet a. subsp. guianensis b. subsp. acanthogyne (Ducke) C. C. Berg c. subsp. hirsuta C. C. Berg d. subsp.pseudopeltata (Mildbraed)C. C. Berg e. subsp. castilloides (Pittier ) C. C. Berg 14-2. P tessmannii Mildbraed 14-3. P xanthochymaKarsten 14-4. P angustifolia (Poeppig & Endlicher)C. C. Berg

C. C. Berg 14-5.P longepedunculata 14-6. P mennegae C. C. Berg 14-7. P humilis C. C. Berg 14-8. P mollis (Poeppig & Endlicher)Huber a. subsp. mollis b. subsp. lecithogalacta (R. E. Schultes) C. C. Berg 14-9. P rubra (Trecul) C. C. Berg

17-3.C. tunuHemsley 18.Naucleopsis Miquel 18-1.N. macrophyllaMiquel 18-2.N. riparia C. C. Berg 18-3.N. velutina C. C. Berg 18-4.N. caloneura (Huber)C. C. Berg 18-5. N. jamariensis C. C. Berg 18-6.N. glabra Spruceex Pittier 18-7.N. imitans (Ducke)C. C. Berg 18-8.N. pseudonaga (Mildbraed) C. C. Berg 18-9.N. ternstroemiiflora(Milbraed)C. C. Berg 18-10.N. oblongifolia (Kuhlmann) Carauta 18-11.N. guianensis (Mildbraed) C. C. Berg 18-12.N. inaequalis (Ducke)C. C. Berg 18-13.N. concinna (Standley)C. C. Berg 18-14.N. stipularis Ducke 18-15. N. herrerensis C. C. Berg 18-16.N. chiguila Benoist 18-17.N. capirensis C. C. Berg 18-18.N. naga Pittier a. subsp.naga b. subsp. meridionalis C. C. Berg 18-19.N. ulei (Warburg)Ducke a. subsp. ulei b. subsp.subandina C. C. Berg c. subsp. amara (Ducke) C. C. Berg d. subsp.puberula C. C. Berg 18-20.N. humilis C. C. Berg 18-21N. krukovii(Standley)C. C. Berg 18-22.N. straminea C. C. Berg 19. Ficus Linnaeus

LISTOFEXSICCATAE

315

LIST OF EXSICCATAE Thislist containsthecollectionsexaminedforrevisedgenera(1-9) andthecollectionscitedforthegenera forwhichadditionsandcorrections arepresented (10-18). Abarca,5143 (3-2a). Abbott,W.L., 757 (2-3). Acevedo,C. M. et al., C.100(10-5). Acevedo,P.et al., 1721(18-16). AcevedoR., P.et al., 8583 (14-la); 9248 (18-8). AcostaSolis,M.,6197(1-2);6428(7-10);6453(3-2a);6933 (1-2);6941 (7-8); 7383 (1-2); 11660(1-1); 12292(710); 12313(13-8c); 13601(8-1); 13676(7-10); 13681 (6-2); 13733(6-1);13950(3-2a);13964(8-1). Adams,C. D. [etal.],6767(9-34);7356,8231(3-2a);9269 (9-34);11805(3-2a);12826(2-3); 13482(9-32). Acevedo,P.etal., 1583(2-3);7417 (7-6a). Agostini,G. et al., 1829(3-2a). Agudelo,C. A. et al. 895 (1-2). AguilarH., M., 250 (9-27). Aguilar,R., 649 (9-10). Aguilar,J. I., 316 (1-1);337, 1330(9-27);1645(2-1). Aguilar,167(18-17). Alain,Bro.(=A. Liogier(etal.)),146,467 (2-3);2550,4945 (3-2a);9672, 19024(2-3). Albin C., J., 5372 (2-3). Albertde Escobar, L.et al.,2194(14-le); 2388(18-6);2585 (2-3);2765 (7-8);4950 (14-le). Albuquerque, B. W.de, 1292(18-21). Alcom,J. B., 1846,2016(3-2a);2068(2-3);2157A(9-33); 3294(3-2a). Alcom,P.et al., 17 (2-4). Alencar,L., 554 (7-6b). AlexanderE. J. [etal.],497 (9-27);2031, 2056 (9-28). Alexander,R. C., s.n. (3-2a,2-3). Alexiades,M. [et al.l, 40 (7-7); 109, 114 (5-1); 118 (8-1); 121 (7-7); 150 (5-1); 155(8-1);200 (7-7);250 (14-3); 334(5-1);363(8-1);441,452(5-1);467(6-1);477 (8-1); 541 (5-1); 542 (6-2); 544 (5-1); 719 (8-1); 736 (6-2); 793(5-1);842(10-1b);953(5-1);1211(6-1);1248(7-5). Allart,A., 122, 194,321 (9-27). Allemao,Fr.F.[etal.], 1437(3-2a);1441(9-32);s.n. (3-2a, 9-30a). Allen,C., 168(3-2a);304 (9-27). Allen, P. H. [et al.], 1736 (2-4); 3741 (6-1);4614 (9-27); 5059 (7-4a);5059 (7-9);5207 (6-1);5492 (7-9);5601 (3-2a);5948(5-1);5955(6-1);5971(5-1);5973(9-10); 6895 (3-2a);7299 (2-3). Almeda,F.et al., 5618(7-8);5684(13-5). Almeida,J. [et al.],49 (9-18);152, 187(9-25);210 (9-22); 223 (6-3);290 (7-1); 1842, 1965(9-30a);2349 (7-3). AlstonA. H.G. [etal.], 115(2-3); 186(9-25);6018(3-2a). Alvarado,C., 113(18-17). Alvarez,E. et al., 109(10-5); 120(7-5); 1301(7-6a). Alverson,W.S. [etal.], 1879(7-8);3282(2-5);3277(7-8). Alves,L. C., 12,212 (9-26a);217 (9-25). Alves,L. M. et al., 12 (7-2). Amaral,I. L. [et al.], 357 (7-1);739 (7-6a);IG-2-2-73,IG2-5-164(6-2). Amorin,A. M. A. et al., 438, 1454 (7-3); 1519 (10-5); 1941(9-1). Ananach,L., RBAE.178(5-3). Ancuash,E., 100(5-3);464 (14-lb);377(7-4a);1018(8-1); 1289(14-3). Andeer,M.,s.n. (9-32). Andel,T.van,384 (18-7);500 (7-4b).

Anderson,W.R. et al., 5781,5791, 5795 (2-3). Andrade,A. Gravede [et al.], 9 (9-25);37 (9-26a);515 (926b);707(9-30d);895(9-26b);936(9-25);2283(9-21). Andrade-Lima, D. de, 2712,49-80 (9-21);49-202,49-203, 49-204,69-5534(7-1);s.n. (9-30d). Andre,E., 1090(9-27). Andreasen, M. et al., 639, 650 (9-28). Andreata,R. H. P. [et al.], 183,395 (9-25);438 (9-2);624 (9-19);927 (7-3). Andrews,L. M., 534a,534c (9-27). Angeli,C., s.n. (9-26a). Anisits,J. D. 2316(9-32);2317 (9-30b);2371(9-32);2571 (9-30b);2605 (9-32);2696 (9-30b);2696 (9-32);2859 (9-30b). Anonymous,s.n. (9-2, 9-3, 9-6, 9-20, 9-25, 9-26a,9-27,930a,9-31,9-34). Antonio,T.,622 (9-27); 1240(9-16); 1735(7-9);3206 (61);4036 (7-9). ApesteguiC., G., 108(5-2). Apui,B., 27 (7-8). Aranda,M. et al., 37 (2-3);87 (9-28). Araquistain, M.,976 (2-3);2844 (3-2a);3179 (6-1);3781, 3823 (2-3);3317(7-4a). Araujo,A. P.de, 230 (7-1). Araujo,D. S. Dunnde [et al.], 152 (9-25); 199 (9-3);800 (9-25);1217(9-3);1461(9-25);1852(9-26a);1875(93); 1889(9-19);2250(9-22);4760 (9-18);5782(9-22); 5882 (9-25).

ArbelaezS., G. et al. 2041 (7-13a);2289 (7-8). Arbo,M. M. et al., 737 (9-32);2859 (7-2). Archer,W.A [et al.], 83, 4141, 4804 (9-32);7720 (6-2); 7935(9-30d);8032,8197 (6-3). Arechavaleta, J., 3005(9-32). Arenas,P.,688 (9-30b). Arevaloet al., 10 (10-3). Argent,G. et al., 6515, 6594,6926 (7-1). Arguelles,J., 107, 112(9-28). Argueta,L., 161(2-3). L. [etal.],1980(9-14);2316(9-32);3084,3214 Aristeguieta, (7-13a);5619, 5637 (9-32);5946 (9-14);6035 (3-2a); 6919,7057(7-13a);7361 (7-6b). A. V., 1631,1673(9-27). Armour, Arnason,T. [etal.],17018(3-2a);17068,17756(2-3);17757 (3-2a). Arnoldo,Bro.,6, 1088, 1104, 1632 (7-13a);1649(3-2a); 2106, 2106A (7-13a). Arroyo, L. et al., 137 (5-2). ArrudaJunior,see Ponce.

Arsene,Bro.G., 1142,5834,6585, 10594,s.n. (1-1). Arvigo, R., 65, 264 (2-3).

AsbunC., M. G. [et al.],35 (13-9.note);s.n. (13-8a); Asplund,E., 8911, 9149 (2-4); 9387 (2-3); 12915 (5-3); 12915 (5-2); 12916 (5-3); 12916 (5-2); 12949, 13194

(6-2); 13420(2-5); 14619(2-3); 15330(7-13c); 15239 (9-27);15361(7-13a);15389,15575(7-13c);18085(32a); 19818, 19819 (2-4). Atahuachi,M., 34 (18-19b). Atwood, J. T. [et al.], AN. 184, 7744 (2-1). Aublet, J. B. C. F., s.n. (4-1). Aubreville,A., s.n. (2-3).

Augusto,Bro.B., 298 (3-2a).

316 Aulestia,C. et al., 43 (18-19d);518 (14-4);539 (18-18a); 1009(18-19d);1016(18-18a);1072(7-4b);1174(11-1); 1499(18-17);2754,2772 (18-15). Aulestia,M. [etal.], 1517,1626,1681(7-5);1765,1844(74b); 1893 (7-5); 1985 (7-6a);2492 (7-5); 2702 (5-3); 2736(10-unnamed); 2753,2802(7-5);3376(7-6a);3385 (7-7a);3505,3552A(7-6a);3580 (7-4b). Austin,D. F.et al., 7205 (11-2);7384 (15-4). Avendailo,S. et al. 297, 298 (9-28). Aviles,S., 13,22(8-1);26 (7-9);52, 84 (7-13);98, 98a(2-4). Ayala,F.,535 (2-3);932 (3-2a);1647(7-4c); 1788(7-6a); 3358 (3-2a);5899(7-14);6439 (10-2c). AymardC., G. [et al.], 1147, 2623 (7-13a);5658 (9-33); 5777(7-6b);6735(16-3);6743(7-6a);7492(6-3);7804, 8070,8183(7-6b);9711(7-6);9837(12-2);9841,9995 (7-6);10031(7-6);10939,11006,11049,11061,11076, 11078,11127(12-2). Badillo,V., 1424,1431(7-6b). Badini,J. [etal.], 1824,s.n. (9-25). Bahamon,M., 1339(2-3); 1607(1-2). Bahia,R. P.,36 (6-3). Bailey,L.H.et al.,301(2-4);483 (7-9);491(2-4);519(8-1); 538 (7-9);669 (2-3). Bailey,V.,366 (1-1). Baitello,J. B. et al., 133(7-2). Baker,C. F.,21, 784 (2-3). Baker,M.A. et al., 6319,6774 (5-3);6984 (13-1). Balansa,B., 178 (9-32); 1969(7-13b);1970(3-2a);1971 (7-2); 1983, 1984 (9-32); 3192 (7-13b);3267 (3-2a); 3282 (7-2);4596 (3-2a). Baldwin,J.T.,3560 (7-4b). Balee,W.L. [et al.], 16 (6-2); 135(10-5);488 (10-3);523 (4-1);608 (15-la); 1697(3-2a);1771(4-1); 1826(6-3); 2248,2277(5-2);2298(4-1);2312(14-8a);2322(6-3); 2370 (4-1);2426, 2444 (6-3);2861 (15-la); 3006 (102b);3498 (4-1);3501 (6-3). Balick,M. et al., 2070,2450, 3168 (2-3). Balslev,H.et al., 4573 (16-1);84813(13-4);97004(7-4c); 97056 (16-3);97064 (16-1);97121 (10-3);97213(131);97327(11-1);97441(18-6). Bang,M., 1652,1737(3-2a). Bangham,W.N., 520 (2-4);558, 479 (7-9);s.n. (9-40). Barbosa,C., 6469,6633 (2-3). Barbosa,M. R., 53, 119(9-25). BarbosaRodrigues,A., 47 (4-1). Barcia,J. C. Coutinho,448 (9-25). A. S. [etal.],3313(1-2);3505(3-2a);4982(18-15). Barclay, Barclay,G. W.,47 (7-2); s.n. (9-27). Barfod,A. et al. 48442 (15- lb); 49812 (18-18a).

Barkley,F.A. et al., 2302, 36032,7496 (9-28). Barnes,C. R. et al., 258 (9-28). Barrelier, M., 35 (18-17). Barreto, H. Mello, 1447, 1515, 1802 (9-25); 7270, 7271

(3-2a); 7380 (9-30c); 9130 (3-2a); 10539 (9-30c); 12231 (9-32).

Barrier,S., 4047 (6-3);4830 (5-2). K. et al., 1653(9-27);2529,2605, 3858,4006A Barringer, (9-4a). 11(9-21); Barros-Lima, Barros,W.D. de, 1034(7-2). Barroso,G. M.,s.n. (9-30d). Bartlett,A. W.,8383(7-4b). H.H.[etal.],10775(1-1);10955(9-33);11124(1-1); Bartlett, 11500(9-27);11867(2-3);11984,12199,12273(9-27); 12354, 12375 (2-3); 12575 (3-2a); 16588 (2-3); 16592 (7-9); 17313 (9-43).

FLORANEOTROPICA Bastos, A. de Miranda [et al.], 24, 175 (9-32); s.n. (9-25, 9-30d). Beach, J. H., 1423 (2-5). Beaman, J. H., 5250 (9-27); 5717, 5910 (2-1); 6194 (8-1). Beard, J. S., 526, 1330 (3-2a). Beard, P., 1177 (3-2a). Beck, S. G. [etal.], 1209(1-2); 1991(10-lb); 2199 (1-2);2501, 2501A, 2501B, 2587, 2602 (7-13b); 2731, 4868 (3-2a); 4949 (16-1); 5078 (7-13b); 6903 (2-4); 8557 (13-3); 8558 (2-4); 10015 (15-4); 10087 (9-33); 10210 (13-4); 11643 (3-2b); 12254 (5- 2); 13590 (3-2a); 13684 (6-1); 13719 (7-5); 15014,15351 (9-33); 16686,16702,16702A(8-1); 16910 (3-1); 18237 (14-7); 18325 (9-13); 19220 (7-5); 19564 (15-4); 20059 (7-6a); 20065 (5-2); 20087 (6-1); 20106 (6-3); 20117 (6-1); 20135A (7-1); 20143 (6-1); 20154 (7-6a); 20176 (5-1); 20211 (6-3); 20254A (7-6a); 20375 (6-2);20300 (7-1);20354 (6-3);20356 (7-6a);20375 (6-2); 20426 (10-5); 20428,20439 (6-2); 22288 (3-2a). Begazo, N., 18 (6-2); 87 (6-1); 91 (6-2); 95 (14-4). Belanger, G. P., 261, 583 (9-27). Belem, R. P. et al., 689 (9-2 1); 884 (7-3); 1148 (9-2 1). Bellido, Ceijas L., 65 (6-2); s.n. (8-1). Bello, E. [et al.], 19 (3-2a); 416 (7-8); 438 (3-2a); 731, 758 (2-1); 841 (7-8); 2218 (18-17). Bena, P., 1302 (4-1). Benalcazar,C. et al., 220 (2-4). Benavides, 0. de, 4265 (14-3). Benitez de Rojas, E., 897 (3-2a). Bennett,B. C. et al., 3502 (5-3); 3507 (2-4);4039 (10-3);4050 (13-4); 4092 (7-4b); 4095 (5-3); 4121 (7-4b); 4446 (2-3). Benoist, R., 517, 518 (4-1); 3047 (8-1). Benson, D., 92, 112, s.n. (3-2a). Bequaert,J., 12, 48 (9-27). Berg, C. C. [et al.], 210, 211 (7-3); 213 (9-3); 214 (9-26a); 247 (10-10); 301 (18-19d); 302 (7-11); 307 (18-19d); 308 (10-5); 309 (7-1 1); 313 (18-19d); 317 (15-1b); 333 (109); 334 (14-4); 336 (7-1 1); 356 (8-1); 390 (16-7); 403 (711); 404(134); 405 (7-11); 406 (14-le); 411 (10-4); 413 (2-4); 414 (9-27); 415 (9-16); 435 (6-2); 437 (15-Ib); 442, 443 (1-2); BG.754 (18-6); 990 (9-34); 1031, 1047 (2-4); 1068 (6-2); 1069 (18-21); 1076 (14-ic); 1078 (18-21); 1079, 1080 (5-3); 1084 (13-8c); 1086 (7-5); 1087 (5-3); 1088 (14-id); 1089 (7-5); 1094 (18-19a); 1097 (7-6a); 1112 (2-3); 1144 (9-8); 1150 (7-3); 1306 (3-2a); 1308, 1496 (2-3); 1498 (9-28); 1504 (6-1); 1509 (3-2a); 1525 (10-1b); 1552 (10-5); 1556(14-la); 1557 (11-1); 1591 (7-5); 1616 (24); 1706A(10-5); 1717 (2-4); 1785 (142); P.18131 (14-la); P.18137 (9-32); P.18349 (7-6a); P.18385(4-1); P.18417(13-4); P.18429(10-1 f); P.18437 (13-1); P.18468 (7-6a); P.18473 (10-3); P.18517 (7-6a); P.18518 (18-4); P.18524 (5-2); P.18525 (14-8a); P.18527 (7-1); P.18534 (18-1); P.18539 (9-30d); 18549 (7-6a); P.18573(9-30d);P.18574(9-32);P.18608(15-la); P.18613 (16-3); P.18663 (15-4); P.18804 (10-12); P.19805 (7-1); P.19807(18-14); P.19864(13-3); P.19881(18-9); P.19937 (7-1); s.n. (5-3, 6-1, 6-2, 6-3,7-3, 7-8, 13-8b, 18-15). Berger, C., s.n. (9-27). Bergfeldt, 794a (9-27). Bergeron, S., 513 (14-8a). Berlandier,J. L., 30 (9-33); 172 (2-3); s.n. (9-33). Berlin, B., 448 (14-Id); 641 (18-15); 757 (16-1). Bernal, P., 5 (2-4); 526 (6-1). Bernal, R. et al., 436 (18-2 1); 437 (13-unnamed). Bernardi,A. L., 18 (9-26a); 545 (1-1); 1314 (3-2a); 1886 (61); 1938 (2-3); 2068 (1-2); 2177 (2-3); 2353 (8-1); 3317 (6-1); 6983 (2-3); 7009 (6-2); 16270 (7-4c); 16278 (6-2); 18062 (7-2); 18374 (7-13c); 18419 (3-2a);s.n (2-3,9-25).

LISTOFEXSICCATAE A. L., 18 (7-1). Bernardo, Bernoulli,G. [et al.], 526, 2543, 2544 (9-27). Berry,P.E., 342, 359, 367 (7-13a);752 (7-6b);2078,2219 (10-9);3485,3528(7-13a). Bertero, C. G., s.n. (2-3, 3-2a, 9-4 1). Bertoni,M., 1199,2431 (3-2a);2885,3715,3762 (7-2). Betancur,J. et al., 699 (14-3);903 (18-18a);1192(9-28); 1698(14-3);2207 (7-4b);4631 (18-17). Bettfreund, K.,273 (7-2). Beyrich,H. C., 346 (9-19);s.n. (9-25). Bianco,F.,87 (3-2a). Biolley,P.et al., 6908(9-27). Birschel,M.,s.n. (9-15,9-28). Black,G. A. [et al.], 46-41 (7-1);46-69 (7-6a);46-354(74c); 47-1027 (4-1);47-1077,47-1715,47-1728(6-3); 50-8572(9-32);50-10516(3-2a);54-17755(9-32). Blake,S. F.,7428A(9-27);7698 (9-28);7803(9-27). BlakeWhite,R.,s.n. (9-27). Blanchet,J. S., 258 (3-2a);267 (7-3);280 (3-2a);568(7-3); 772, 919 (9-21); 1034(9-32); 1168(7-3); 1960(9-18); 2390 (9-21);3064, 3301 (3-2a);s.n. (3-2a,6-3, 9-9, 921, 9-32). Blanco,C.A., 101,509 (6-2);854 (7-13a);905 (2-3). Blum,K. E. [et al.], 1216(7-9);2011 (24); 2333(9-27). Boeke,J. D. et al., 1278(7-7);2184 (18-18b). Boldingh,I., 2362, 3344, 3344B (9-27);3932, 4702, s.n. (3-2a). Bondar,G., 2162 (7-1);s.n. (9-3, 6-2, 6-3, 7-3). Bonifaz,C. et al., 3069 (7-12);3240 (18-18b);3352 (8-1); 3407 (7-12);3410(15-lb). A. J.A.,973 (9-31);1512(2-3);3102,3350(1-1); Bonpland, 4485 (2-3);s.n. (2-5, 3-2a,3-2b,7-2). Boog, W., s.n. (7-3). Boom, B. M. [et al.], 4024 (7-1); 4030, 4109 (7-6a); 4313 (16-1); 4316 (7-1); 4334 (10-1f); 4364 (13-3a); 4482 (10-5); 4489 (7-1); 4505 (14-8a); 4514 (10-2c); 4955 (7-1); 5019 (3-2a); 5679 (12-2); 6243, 6696 (7-13a); 8369 (13-4); 8650 (18-19c); 8655 (16-2); 10597 (7-6b). Borbas, E., s.n. (9-30b, 9-32). Borrell, J. R., 194 (3-2a). Boster, J. S., 67 (7-4c).

Botteri,M. [etal.], 852 (1-1); 1014(2-1); 1336,1337,s.n. (1 -1).

BoudetFernandes,H. 0., 34 (9-26a);1379, 1579(9-19); 1730,1863A(9-16).

Bourgeau,E., 189(1-1);2069 (2-3);2520 (1-1);s.n. (2-3). Bowie,J.et al., 65 (9-19);363 (6-3);s.n. (6-3,9-26a,9-25). Box, H. E., 2004 (3-2a).

Boyle,B. et al., 1248(18-22);3293(18-16);3469(7-12). Brade,A. C. [et al.], 4579,7231 (9-3); 7874 (9-6); 9825 (918); 10303 (9-26b); 12700 (7-2); 12705 (7-3); 13030 (9-34); 14152 (9-2); 15017 (9-3); 15065 (9-26b); 15438 (9-30c); 15768 (9-25); 15786 (7-1); 16801 (9-25); 17439 (9-26b); 18242 (9-2); 18242 (9-3); 18278 (9-18); 18278, 18429 (9-25); 18441 (7-3); 18461 (9-29); 18516 (9-18); 18621 (9-19); 18683 (9-26); 19756 (9-3); 19756 (9-6); 19913 (9-1); 20570 (9-25).

Braga,P. 1. Soares[et al.], 6 (9-25);58 (9-22);398, 406, 514, 989 (9-25); 1001 (7-3); 1087 (7-2); 1128, 1139, 1163, 1187 (9-25); 1293 (9-26a); 1448 (9-18); 1452 (98); 1706 (9-26a); 1770 (9-27a); 2206,2207 (9-25); 2428, 2457 (9-26b); 3999 (7-3);s.n. (9-18,9-25). Brand, J. et al., 350 (2-4); 465 (7-9); 483, 685, 686 (2-4);

718, 724 (2-3);772 (10-1la); 1005(13-8c);1186,1282 (7-9); 1442 (6-2); 1463 (7-6); 1489 (11-1).

Brandbyge,J. et al., 30052 (14-1b);30240 (7-4b);30302 (14- 18b); 30398 (22-5); 30650 (7-4c); 30815 (7-6a);

317 30822 (7-4c); 30841 (7-6a); 30870, 30898 (7-4c); 31039 (7-6a); 31044 (16-1); 31072 (7-4c); 31093 (7-6a); 31238 (7-4c); 31416 (7-6a); 31432 (7-4c); 31436 (7-6a); 31499 (7-4b); 31631 (7-5); 31654 (2-3); 32071, 33419 (2-5); 36237 (18-6). Brandegee,T. S., 2316 (2-3); s.n. (9-28). Breedlove,D. E. [et al.], 6591 (2-3); 9393 (1-1); 9760,11952 (2-1); 12884 (2-3); 15707 (13-5); 18374 (1-1); 20288 (9-27); 22238 (2-3); 24074 (2-1); 25244 (3-2a); 25630, 26109,27926,29309 (2-3); 29773 (9-17); 30982,32818, 32900 (2-1); 33290 (2-3); 33660, 34379 (2-2); 34503, 35040, 35182,35247 (2-1); 35299 (13-5); 35310 (2-1); 39787 (1-1); 46740 (2-1); 47409 (9-17); 48708, 49324 (2-1); 49587 (9-17); 49684,49882,49952,50454,50743 (2-1); 50648 (2-3); 50802 (1-1); 52681 (2-1); 55402 (918); 56954, 57128, 57173 (2-1); 57334 (9-17); 57481 (2-1); 58326 (2-2); 66322 (9-17). Brenes,A. M., 18 (9-27); 3992, 3992a (3-2a); 4332, 5488C, 6235, 6364 (7-8); 6504 (9-27); 12207 (7-4a); 12319 (24); 12558, 12558a (3-2a); 12560 (9-27); 12971 (3-2a); 14391 (9-27); 17459 (3-2a); 18920 (9-4a); 18921 (9-27); 18922, 18923, 18924, 18925, 18927 (3-2a); 18949, 18951 (7-8); 18952 (2-5); 19340 (3-2a); 20447, 20450 (6-1); 20826 (9-4a); s.n. (9-27). Breteler, F. J., 3903 (3-2a); 4001 (2-3); 4064 (3-2a); 4333 (7-13a); 4555 (3-2a); 4573 (2-3); 5054 (6-2). Bristan, N., 1002 (2) (9-27); 1115 (7-9); 1119 (2-3); 1188 (15-1b); 1231 (7-11); 1456 (2-4); 1459 (5-1). Brito, J. M. de et al., 13 (7-4c). Brito Junior,J. B. Ferreirade, s.n. (9-26b). Britton, N. L. [et al.], 247 (9-27); 407 (3-2a); 427 (9-34); 898 (3-2a); 1035 (2-4); 1759 (9-27); 1793, 2220 (3-2a); 2238 (9-34); 2553, 2598 (2-3); 2762,2762 (9-27); 2853 (2-3); 3737 (9-36); 7337 (2-3); 7649, 9557, 10020 (32a); 15135 (2-3). Broadway,W. E. [et al.], 677, 1871 (3-2a); 3933 (9-27); 4072 (3-2a); 4371, 4772 (9-27); 5100 (3-2a); 5452 (9-27); 7398 (3-2a); 7882 (9-27); 8046, 8084,8086 (3-2a); 8194 (9-27); s.n. (3-2a, 9-27). Brongniart,A. T., 1867 (9-27). Brooke, W. M. A., 5797 (3-2a). Brown, M., 104 (3-1). Bruijn,J. de, 1056 (6-2); 1206, 1330 (2-3); 1488, 1498 (6-2); 1515 (12-2); 1541 (74b); 1553, 1568, 1658, 1659 (6-2). Buchtien, O., 647, 1357 (3-2a); 1567 (14-la); 1658 (7-5); 1948 (9-13); 3147 (1-2).

Buckley,S. B., s.n. (1-1). Bunting, G. S. [et al.], 713 (2-3); 2095 (8-1); 5032, 5038 (32a); 5180 (2-3); 5792, 6297, 6307, 7372 (3-2a); 7379 (7-13a); 7667 (9-27); 7682, 8402, 10153, 10199 (3-2a); 10201 (7-13a); 10313 (6-1); 10837(134); 11549 (3-2a); 11985 (2-3); 12326 (13-8b); 12854 (2-3); 13233 (7-13a). Burch, D., 4581 (9-10). Burchell, W. J., 882, 883 (9-25); 883 (9-26a); 1071 (7-3); 1124 (9-32); 2870 (7-3); 6360 (3-2a); 6383 (7-1); 7718, 8090 (9-30d); 8283 (9-30c); 8363 (9-32); 9181 (9-30d); 9611 (4-1). Burger,W. C. [et al.], 148, 150 (2-3); 4290 (7-4a); 4429 (21); 4472 (9-10); 4563 (7-8); 4660 (2-4); 5082 (7-4a); 5434 (9-10); 5500 (7-4a); 6123 (9-28); 6618, 6939 (23); 7126,7131,7201 (9-10); 7794 (7-8); 7802 (2-1); 8081 (2-5); 8849 (9-10); 8899 (7-9); 8920 (2-5); 8989 (2-4); 8997 (2-5); 9013 (8-1); 9585 (7-8); 10046,10049 (7-4a); 10483 (9-10); 10488 (7-4a); 10496 (23); 10518, 10614 (2-4); 10886 (2-5); 10948 (2-3); 10984(24); 11280 (910); 11291, 11294 (2-4); 12090 (7-8); 12272 (2-4). Burgos, J. A., 59 (6-2).

318 Burkart, A. E., 14682(7-2);16392(9-27);20575,s.n. (9-32). Busey,P.,318 (2-4);748A(7-14a);748A,775 (2-4). Buysman,M., 2540 (9-27). BW(Boschwezen,Surinam); 4821 (6-2). Cabrera,A. L., 3847, 10815(9-32);21103 (3-2b);29354 (9-32);29906,32219(3-2b). Cabrera,E. [et al.], 3313, 4194 (2-3); 5052, 6383, 6527, 6677,6803 (3-2a); 7721 (9-27); 8590,8988,8994,9013, 9132,9523 (3-2a). Cabrera,I. et al., 875 (3-2a); 2400 (6-3); 2662 (6-2); 5086 (1-2).

Caffrey,G. M., 56 (2-3). Cain,S. A., 63 (5-1). Caldaza,J.I. [etal.],2 (2-1);44 (8-1);56 (9-27);411 (3-2a); 623 (2-3); 763, 1761, 1824 (2-1); 1836 (3-2a);2227 (2-3); 2833 (9-28); 3104 (2-1); 4118 (2-3); 5410,5519 (9-27); 5568 (2-3); 5620 (9-27); 5655, 5657, 5700 (2-3); 5705 (2-4); 6230 (9-27); 7328, 9703 (2-1).

Calder6n,S., 438 (3-2a);845 (2-3);901 (3-2a);1065,1441, 1698 (2-3); 1721 (9-27); 1749 (2-1); 1991 (3-2a); 2178 (2-3).

Callaway,R.,440 (9-28). Callejas,R. [etal.],2267(7-8);2497(1-2);4763(2-4);4854 (7-4b);4880(2-4);4959(9-27);4979(2-4);4982(5-1); 5188 (2-3); 5406 (3-2a);5491 (10-lib); 5657 (2-3); 5705, 5845 (24); 5918 (18-6); 6645 (18-18a); 7032 (186); 7087,7126 (7-5); 8176 (24); 8264 (1-2); 8397 (16-7); 8412 (2-4); 10942 (9-27). Camargo,O., 1758, 62409,62163 (7-2).

Camp,W.H.,E-933,E-951(2-5). Campbell,D. G. et al., 6174, 6202 (5-3);6361 (7-7);6401 (14-9b);6481 (14-4);6899 (10-14);6504, 6552 (7-7); 6526 (5-3);6818 (6-2);6828, 6837, 6902 (5-3);6907, 6909, 6949 (7-4b);7061 (18-19b);7097 (10-9);7203 (14-3);7345, 7542 (7-5);7554 (14-3);7601 (18-19b); 7675 (74b); 7752, 7977 (5-3);7998 (6-2);8054 (5-2); 8068,8081, 8163,8166,8189 (7-4b);8213(7-5);8302 (7-4b); 8396 (12-1); 8400 (7-4b); 8401 (12-1); 8429 (15-3); 8457 (14-9b); 8504, 8555, 8565 (7-4b); 8606

(10-1b);8872(7-5);8923(7-6a);9035,9172(7-5);9312 (16-5); 9322 (6-2); 9328 (18-10); 9325 (6-2); 9607 (7-5);

9861 (6-2); 10159(16-4); 10486(3-2a);10849(7-4b); 10886 (7-5); 11001 (3-2a); 12894 (6-2); 12887 (5-3); 14316(5-2); 14542 (4-1).

Campos,J. F.B. M., 10 (9-32). Campos,M. T.etal., 21 (18-9);49 (14-9a). Canias,D. A. G., 499 (16-1);633, 760, 1713(7-4b). CandidoE., E., 37 (2-3). Capucho,P.,365, 510 (4-1). Carauta,J. P. P. [et al.], 95, 157, 173, 174, 290 (9-25); 292

(9-26a);346 (9-25);347 (9-22);354, 358 (9-26b);369 (7-3);484, 498 (9-26a);511 (9-25);607, 641 (9-26a); 650 (7-1);697 (9-25);721 (9-30c);795 (7-3);944 (925); 1067(9-21); 1067(9-30d);1085(9-3);1099(9-2); 1277 (9-7); 1377 (9-6); 1383 (9-25); 1387 (9-26b); 1400 (9-26a); 1405 (9-25); 1408 (9-30a); 1409 (9-19); 1411

(9-22);1413(9-25);1460(9-33);1494(9-25);1504(73); 1517(9-3); 1583(9-26b);1668(9-18);1669(9-23); 1687(9-2);1689(9-26a);1695(7-3);1744(9-25);1745, 1747(9-3);1771(9-26b);1775(9-21);1776(9-25);1777 (9-19); 1777 (9-2); 1810 (9-3); 1811 (9-20); 1837 (926a); 1849 (9-3); 1860 (9-30d); 1863 (9-32); 1873, 1875, 1877, 1878 (9-30d); 1883, 1888, 1979(9-26a); 2111 (9-

25);2125(9-26a);2129(9-3);2222(9-22);2260(9-25); 2279(9-30c);2292(7-2);2300(9-22);2498(9-3);2707, 2777 (9-25); 2800 (9-18); 2865 (9-25); 2877 (9-32); 2893

FLORANEOTROPICA (9-25);2956 (9-22);2970 (9-25); 3075 (9-30c);3203 (9-26a);3279 (9-18);3290 (9-22);3312, 3423 (9-25); 3456 (9-26a);3466, 3679 (9-25);4006 (9-30d);4292, 4332, 4390 (9-25);4400 (9-24);4419 (9-18);4421 (92); 4464 (9-7);4701, 5399 (9-25). Cardenas, D., 286, 445 (18-19d);691 (10-1la); 867, 1032, 1449 (18-19d); 1461 (6-2); 1572 (5-1); 1610 (16-1); 1727,2008 (7-11);2025 (10-5);2108 (18-6);2121 (711);2287 (18-19d);2442 (13-4). Cardenas, M., 1697,1706(7-13a);1757(7-7);1938(10-6). Cardona,F., 1344(7-6b). Cardoso,T., 195(9-28). Carlson,M. C., 2445,2669 (2-1);3230 (2-3). Carpenter, R., 75 (8-1). Carrasquilla, L., 3019, 3020(3-2a). Carrillo,T.,28 (3-2a). Carter,A. et al., 1210(2-3). Carvalho,A. M. V. de et al., 1159(9-9);2034 (7-3);4000 (10-6). Casaretto, G., 591 (7-1). Casari,M.B. [etal.],238,395,442(9-26a);806, 1103(9-25). L. [etal.], 12,44 (6-1). Castafno-Rosalba, Castellanos,A., 38 (9-32); 15008 (9-32); 24885 (3-2a); 26206(9-25);26250(9-26b);s.n. (9-3, 9-32). Castelnau,F.de,s.n. (6-3). Castillo,A., 146(7-15a);636 (7-13a). CastilloC., G. et al., 126(2-1);564, 644 (2-3);803, 2316 (9-27);2317 (2-3). Castro,D., 2619 (9-28);2628 (3-2a). Castro,M., 67 (2-3). P.[etal.],361(6-1);362,1007,1008,1009,1010 Cavalcante, (6-2); 1018(9-30d);1536 (6-2); 1536 (6-3); 1666 (32a); 1760(6-3);2349 (9-30d);2754 (6-2);2896(3-2a). Cazalet,P.C. D., 5058(8-1);7540(7-6a);7560(2-5);7684 (7-5). CedilloT., R. [et al.], 10 (2-3);98, 117 (2-1); 138 (9-27); 151,180, 199(2-1);1889(2-3);2613(2-1);2710(6-1); 2742,3062(2-1);3793 (2-3). CeijasB., L., la (10-13);s.n. (10-1If, 10-13). Celestino,Bro.,46 (9-27). CerratoB., C.A., 15 (9-10). Cer6n,C. E. [etal.],204 (2-3);262 (2-4);286 (10-13);348 (6-1);452,484(24); 1667(6-1);2161(2-4);2651(5-3); 2697(7-5);3035(14-id);3055(14-2);3077(2-3);3223 (18-6);3228(17-2);3294(13-8c);3347(7-4b);3456(148b);3459(13-3);3608(7-5);3699 (14-2);3837 (16-5); 3849(5-3);3851(14-ib);3863(14-4);4003(7-4b);4133 (14-2);4135(18-15);4523(2-4);5098(18-20);5099(148b);5364 (7-5);5452 (5-3);5621 (2-3);5977 (13-8c); 6005(5-3);6107(7-4b);6279(5-3);6372(6-1);6818(12);7208(7-5);7229(7-4b);7278(7-5);7295(6-1);7302 (7-5); 7482 (7-4b);7485 (18-15); 7563 (7-4b); 8923 (1-2);8926(7-8);9399(144); 9443(13-8c);9868(16-7). Cervi,A. C. et al., 2368 (7-2). Chac6n,A., 124(9-27);243 (2-5);347 (7-8);703 (2-5);984 (6-1); 1103(9-10). Chacon,l. A. [etal.], 8 (18-19a);287 (7-9a);401 (2-5);800 (7-4a);948 (2-3);1228(9-10);1393(7-4a);1820(7-8). Chagas,J.,229 (7-6a);391, 3345(9-32);21496,s.n. (7-6a). ChavelasP.,J.et al.,ES-208,ES-124,ES-69(2-3);ES-1615 (13-8). Chapman, A., 2311 (9-27). Chase,N. H., 7474 (9-33). Chatrou,L. W.et al., 30 (7-7). Chaussy,F., 16(3-2a). ChavezA., R., 273 (3-2a). ChazaroB., M., 376 (2-1).

LIST OF EXSICCATAE

Chiang,F., 12(3-2a);249,406 (2-3). ChingHoet al., 45 (18-18a). Chodat,R. [et al.],247, 247B (9-32);s.n. (7-2). Churchill,H.W.et al., 3992(7-9);4117(7-11);4452(2-5). Cid,C. A., see Ferreira, C. A. Cid Clark,H. L. et al., 7176 (16-2);7361 (18-10);7404 (7-6b); 7640(11-1). Clark,J.L. [etal.],417,524 (7-12);1162(18-7);2718(15lb); 3277 (10-3);3281 (13-1). Clark,0. M., 6883 (9-27). Clarke,D., 2192 (14-la); 3125 (10-10);3626(14-la). Claussen,P., 1 (3-2a);92 (9-2); 156 (9-30d);195(9-26a); 346 (9-32);s.n. (9-30d). Clemente,Bro.,4594 (9-35);5802 (2-3). Coelho,D., 6007 (6-2). Coelho,J. de Moraes,693-A, 1073(7-1);2066 (9-32). Coelho,L. et al., 15(10-2c);67 (7-1);219 (11-1);401 (182); 455 (12-2);1817(6-2);3083(7-6a). Coello,F.,217, 221 (7-5). Cogollo,A. [et al.], 508, 684, 773, 835, 1002, 1002, 1048 (7-8); 1253(7-9b);1471, 1599(7-8); 1663(6-2); 1672 (7-4b);1789(6-2);1917(7-8);2088 (18-18a). Colaris,W., 1050(9-25). Colchester,M. E. M., 2382 (6-3). Collenette,C. L., 131(6-1). Colonello,G., 952 (3-2a);1228(7-13a). Cominote,J.,s.n. (9-26a). Commerson, P.,s.n. (9-32). Conejos,J., 85 (6-2). Constantino, D., s.n. (9-2). Contreras, E.,40 (3-2a);120(2-3);1299,1584,1585(9-27); 1902(2-3); 1999(13-8);2172(2-1);2414 (9-27);2797 (2-3);2842(9-18);3347,3535(2-1);4281(13-5);4282, 4303 (2-1);4338 (9-18);4812 (2-1);5303(9-18);5443 (2-3);6624,6636,6655(2-1);7163(2-3);8987(10-5); 9052,9174, 9186 (2-3);9191 (9-27);9193 (8-1);9483 (2-1);9625(2-3);10476(2-1);10683(9-18);11526(2-1). Conzatti,C. [etal.], 175,176,3024(9-27);3064(2-1);3687 (9-27);4802 (9-10);9308 (9-17). Cook,M. T.,42 (3-2a). Cook,0. F. [et al.], 13 (2-3); 124, 145, 146(9-27);744 (32a); 1719(6-1). Cooper,G. P. [et al.], 4, 4a (2-3);28 (7-4a);88 (8-1); 107 (7-4a);132(8-1);174(74a); 232 (6-1);237(3-2a);326, 349 (2-3);455 (8-1);554 (7-9). Cordeiro,I. [etal.], 185(6-3);613 (9-7). Cordeiro,J., 352 (7-2). Cordeiro,M. R., 1079,1290(6-3). Cordoba,J. J.,265 (7-4a). Cornejo,J. C. et al., 219 (1-2). Cornejo,X. et al., 1335,1806(7-13c);3464(8-1);3479(12); 4218 (7-10);4231 (2-3);4660 (13-6);4926 (7-8); 5021(7-13c);5087(2-3);5544(13-8b). CorreaA., M. D. [etal.],424 (9-27);891 (18-19d);1696(81);1994(2-4);1696(8-1);2019,3541,3966(2-3);4165 (2-4);8911(18-17). Correa,S., 163(9-28). Cosson,E. St.-C.,18 (3-2a). Costa,A., 2988 (3-2a). Costa,L.V.,45813(10-8). Cowan,C. P.,2147, 2306 (3-2a);2907 (2-1);3082(7-8). Cremers,G., 8207 (8-2). Crist6bal,C. L. [et al.], 1378(3-2b);1930,s.n. (9-31). Croat,T.B. [etal.], 493A(9-27);512A(9-29);4106 (7-9); 4276, 4597, 4643 (2-4);5765 (7-9);5792 (2-4);5822, 5897(8-1);5973 (2-4);6115 (7-9);6190 (7-13);6205, 6306 (7-9);6411, 6582, 6994 (2-4);7703, 7914, 8179

319 (8-1); 8457 (7-9); 8533 (8-1); 8878 (3-2a); 8901 (2-4); 8994,9280 (8-1); 9301 (7-13); 9331 (8-1); 9786 (15-lb); 10168 (2-4); 11441 (2-3); 11598 (8-1); 12501 (7-9); 12698 (2-4); 13275 (7-9); 14451 (2-4); 14977, 15247 (2-3); 15384 (2-4); 15576, 15577, 16197 (7-9); 16540 (2-3); 16545, 16625 (7-9); 17191 (7-11); 17366, 17383 (7-9); 17885 (2-3); 19245 (7-5); 19381 (18-2); 19523 (18-20); 19732 (14-3);19532A (74b); 20122 (7-5); 20246 (2-3); 20256 (14-id); 20357 (7-4b); 20368 (138c); 20772 (5-3); 21074 (2-4); 21960 (7-9); 22093A (51); 22139 (7-4a); 22153 (10-4); 22158A (6-2); 22195A (7-4a); 22459B (5-1); 22482A (7-4a); 22490,22499 (24); 22557B (7-4a);27435 (7-9); 34646 (24); 34722 (2-4); 35043 (7-11);35113 (7-4a);36708 (9-10);36813 (18-18a); 38099 (7-9); 39399,39400,40636 (9-27); 41449 (9-10); 41833, 41902, 42031 (9-27); 42607 (9-10); 42842 (927); 43089 (2-1); 43728 (9-28); 43780, 43852, 43984, 44035 (9-27); 51634 (7-5); 51641 (2-4); 55324 (3-2a); 58355 (2-4); 59369 (10-10); 62457 (7-5); 62463 (14-2); 62819 (2-3); 63465 (2-1); 64399 (13-5); 64836 (2-3); 64589 (9-10); 72531 (7-6a). Croizat, L., 762 (6-1). Crosby, M. R. et al., 329 (3-2a); 770, 1182 (9-34). Crosse, R. W., 12578 (6-1). Crow,G. E. et al., 6937 (9-28). Crud-Maciel,6 (9-22). Crueger,H., s.n. (3-2a, 9-27). Cruxent,J. M., 46,47 (7-6b); 299, 300 (6-3). Cruz, J. A. da, 148 (9-27); s.n. (9-32). Cruz, R., 1 (2-3). CuadrosV., H. et al., 597 (2-4); 653 (2-3); 654, 1008 (2-4); 1097 (2-3); 1222 (7-13a); 1229, 1309 (2-4); 1559 (186); 2108 (7-4b); 2111 (12-1); 2120 (7-13a); 2415 (8-1); 3101 (10-5); 3670 (18-18a); 3676 (3-2a); 3846 (2-4); 3881 (2-3); 3893 (9-27). Cuatrecasas,J. [et al.], 3672 (2-3); 7889 (9-32); 9415, 9841 (3-2a); 10160(7-13a); 10489(3-2a); 10618(8-1); 10840 (2-3); 11099 (2-5); 12579 (16-7); 13573 (2-1); 14030 (109); 14117(2-3); 14135(15-1b); 14205 (14-3); 14227(2-3); 14264 (15-1b); 14342 (8-1); 14424 (16-7); 14836 (7-4b); 14857 (2-3); 14891 (7-8); 15148 (6-2); 15204 (7-4b); 15278 (6-1); 15340 (7-13a); 15487, 15493 (6-1); 15508 (9-15); 15545 (16-7); 15572 (7-8); 16303 (12-1); 16348, 16810, 16853, 16858, 17622 (2-3); 17669 (8-1); 21401 (2-3); 18405, 18777 (1-2); 21188 (14-4); 21318 (6-1); 21735 (1-2); 24000 (6-2); 26142 (18-22); 26164 (2-5). Cuello, N., 333, 336, 337 (6-3). Cufodontis, G., 200 (7-4a). Cunningham,A., s.n. (9-25, 9-26a). Curran,H. M. [et al.], 7 (3-2a); 27 (7-13b); 96 (6-2); 102 (2-3); 130 (9-2 1); 145 (3-2a); 186, 205 (2-3); 248 (7-3); 372 (2-3); 601, 683, 875, 1022 (3-2a). Current,D., 31 (3-2a). Daemon, 16 (9-3). Dahlgren, B. E. et al., 43363 (9-27); s.n. (9-28). Daly, D. C. et al., 483 (6-3); 677 (13-3); 732, 1212, 1223 (6-3); 1486 (7-1); 1644 (6-3); 1677 (7-1); 1805, 1932 (6-3); 4146 (14-9b); 4310 (6-3); 5150 (18-18b); 5649 (14-1b); 5657 (7-14); 5653 (11-1); 5664 (3-2a); 5766 (2-3); 5769 (14-5); 5779 (16-1); 5802 (16-2); 6060 (14le); 6243 (7-6a); 6438 (2-4); 6805 (13-1); 7114 (7-5); 7119 (7-6a); 7383 (5-2); 7519 (8-1); 7572 (18-10); 7838 (6-2); 8000 (7-1); 8010 (15-1b); 8015 (7-6a); 8025 (77); 8140 (5-2); 8329 (18-8). Daniel, H., 3390, 3978 (2-5); 4006 (16-7). Daniels, A. G. H. et al., 1079, 1081 (6-3).

320 Danin,A., 76-19-1(3-2a). D'Arcy,W.C., 1945,2682(3-2a);3193(9-27);4041(2-3); 10863(1-2); 15104,15106(7-9). Darwin,S. P.,2313 (9-27);2383, 2472 (2-3). Daubenmire, R., 31 (9-28). Davidse,G. [etal.],2447(7-8);12820,12841(7-13a);13264 (10-3);13712(10-5);13735(10-3);16064(15-4);16972 (7-6a); 17136(7-6b); 18145(9-27); 18398, 18399(32a); 18518, 18553 (8-1); 19225 (10-1b); 19227 (8-1); 20154,20200,20286,20349 (9-27); 20356 (8-1); 20627

(9-27);24237,24263,24471(7-8);26244(9-10);26332 (10-5);26860(14-la); 27585 (10-5);29708,29896(927); 32199, 32292 (2-1); 34481 (9-10);35518 (3-2a); 35875 (9-17).

Davidson,C. [etal.], 3679(2-4);3282(14-6);4875 (3-2a); 5218 (18-10); 6780, 6857 (7-4a); 8378 (10-5); 8859 (18-18a). Davila, P. et al. 1982 (9-28).

Davis,E. W.et al., 1009(14-Id). Davis,D. H., 1696(7-13a). Davis,J.A. W., 1029(6-3). Davis,P.H. et al., 60827 (7-2). Dawe,M. F.,479 (3-2a);882 (6-2). Deam,C. C., 6283,6378 (3-2a). Defler, S., 316 (10-14); 318 (14-3); 319 (18-13); 320 (18-

8); 324 (10-9); 325 (18-7); 326 (13-4); 327 (18-21); 328 (18-7).

Degener,O., 26719(9-27). Del Aguilo, M. et al., 5 (6-2); 81 (6-1).

DelascioC., F.et al., 348 (13-8a);2823 (6-3);7227, 8803 (9-32).

Delforge,H.,s.n. (9-25). Delgado,E., 294,419 (3-2a). Delgado,H.,28 (18-21). Delgado,L., 64, 329 (6-3);375, 964 (7-6b). Denslow,J. S., 79-125(2-5). Desvaux,A. N. (herb.);s.n. (9-3, 9-26a,9-30d). Devia A., W., 451, 611 (2-3); 2296 (2-4). Devia U., J., s.n. (6-2). Diaz, A., s.n. (3-2a).

Diaz,C. [etal.], 174(18-13);678 (14-2);751 (16-6);173095 (7-7); 1765-109(14-8a);1792-4(6-1);2967 (1-2); 3717 (16-7);3148 (9-13);5061 (10-1b). DiazS., C. et al.,43 (7-6a);673 (7-1);1638(6-2);1698(62); 2029 (3-2a);2103, 2158 (7-8); 2507, 2968 (3-2a); 2967 (1-2).

Diaz,S. et al., 3224 (7-13c). Diaz,W.et al., 215 (14-3);509 (9-27). Dickson,J.D.,89 (2-3);1207(3-2a);1327(2-3);1422(2-1). Didrichson,F.,s.n. (7-2). Dieterle,J. V.A., 4000, 4060, 4206 (9-28);4234 (9-27); 4292 (9-28). Dik, A. [et al.], 114 (7-5); 256 (14-1b); 595 (18-15); 617, 1147 (7-5); 1199 (7-4b); 1243 (7-5); 1259 (7-4b); 1535 (18-unnamed); 1581 (15-1b).

Dionizia,F.et al., 34 (6-3);89 (7-6a);107(7-1);109(7-6a); 141 (12-1); 156 (7-5).

Diquet,L.,s.n. (9-28). Dodge,C. K.,44 (1-1). Dodge,C. W.et al., 17365(2-4). Dodson,C. H. [et al.], 1423(2-1);5235 (6-1);5386 (8-1); 5450 (7-10);5931 (2-4);5994 (6-1);6183 (8-1);6333, 6471 (18-16); 6628 (7-10); 6733 (18-16); 7075 (8-1); 7098 (6-1); 7335 (14-4); 7492 (7-10); 8785 (6-2); 8809

(3-2a);8825(10-Ib);8850(6-1);8984(7-10);9601 (713c);9668(9-27);9804 (7-10);9981(6-2);9988(8-1); 10240(18-18b);10278(7-12); 10320(8-1); 10600(2-

FLORANEOTROPICA 4); 11346, 11454, 12346 (7-13c); 12837 (6-1); 13739 (13-8c); 13806 (7-12); 14382 (7-8); 14612 (14-3); 14740 (6-1); 14751, 14972 (8-1); 15849 (6-1). Dombey, J., s.n. (2-5, 9-25). Donselaar,J. van, 1097 (4-1); 1221, 1300,2082,3318 (6-3). Dorantes, B., 2614 (2-1); 2708 (9-27); 2762 (8-1); 2911 (2-1); 3401 (8-1); 4041 (2-1). Dorantes, J. [et al.], 810, 1085, 1274, 1394, 1407 (3-2a); 2665 (9-27); 5254 (2-3). Dorr,L. J. et al., 7523 (1-2). Doyen, C., 12 (2-4). Dressler, R. L. [et al.], 12, 1569, 1783 (9-27); 1889 (9-33); 2372 (2-3); 3136 (7-9); 3461 (7-11); 3858 (18-19c); 4068 (2-4); 4343 (2-3); 4690,4698 (9-17); 5196 (2-4); 8825 (18-19d). Dryer,V. J., 1001 (7-9a); 1538, 1545, 1546 (7-8). Duarte,A. Pereira[et al.], 151 (9-30a); 182 (9-3); 200, 303 (9-25); 384 (9-26a); 410 (9-25); 523 (9-32); 793 (7-3); 1540 (9-18); 1575 (9-3); 1780 (7-2); 1811 (9-31); 2274 (9-32); 3019 (9-30c); 3366 (3-2a); 3623 (9-18); 3725 (9-30a); 3779 (7-1); 3895 (3-2a); 3980 (9-18); 4050 (73); 4164 (9-2); 5299 (7-1); 5438 (7-3); 5466 (9-25); 5658 (16-1); 5660 (9-21); 5709 (9-30d); 5714 (9-30c); 5839 (9-26b); 5969 (6-2); 6369 (3-2a); 6635,6636 (7-1); 6739, 8579 (3-2a); 10239 (9-30d); 10339 (9-32); 10339A (930d); 10756 (9-30c); 19695 (9-3); s.n. (9-3, 9-18). Duchessaing, P. D., s.n. (9-27). Ducke, A., [et al.], 59 (6-2); 114 (7-1); 193 (7-14); 225 (32a); 231 (6-2); 557 (7-6a); 557 (7-14); 810 (7-1); 946 (4-1); 947 (5-2); 948 (6-2); 949 (7-5); 1040 (7-6a); 1236 (4-1); 1457 (10-12); 1478 (3-2a); 1530, 1531, 1532 (714); 1537 (12-2); 1726 (6-3); 1789 (14-8b); 1834 (7-6a); 2058, 2131 (3-2a); 2172 (6-2); 2193 (3-2a); 2251 (6-1); 2625 (3-2a); 2885 (7-1); 2885 (6-3); 3636, 3648 (3-2a); 4899 (6-3); 4938 (7-1); 4957 (3-2a); 6153 (6-3); 6805, 6961 (7-1); 6977 (7-5); 7458 (7-6a); 7655 (7-5); 7670, 7671 (7-6a); 7698 (14-6); 7959 (7-6a); 8296 (4-1); 8845 (7-1); 8957 (7-6a); 9169 (7-1); 9190 (6-3); 11006 (5-2); 12489 (3-2a); 13029 (18-19c); 13042(7-1); 13051,13052, 13054, 13055 (6-2); 13077 (5-2); 14978 (6-3); 15133 (41); 15270 (6-3); 15922 (5-2); 16059 (6-3); 16606 (6-2); 16757 (18-19c); 16924 (5-2); 16993 (6-2); 17044 (7-5); 17204 (6-2); 17527, 17528, 18271 (4-1); 18318 (7-6a); 18323, 18324, 18325, 18326 (6-3); 19453 (4-1); 19473 (6-3); 19474, 19475, 19476 (7-1); 23621 (10-12); 23994, 23995 (7-14); 25255, 25256 (7-6a); 25257 (7-1); 35613 (7-14); 35620 (7-6a); s.n. (6-2,9-3, 9-32). Dudley, T. R., 10290B (13-3); 11494 (2-4). Dugand G., A., 57 (3-2a); 154 (7-13a); 160 (3-2a); 490 (713a);496 (3-2a);590 (7-13a); 1073, 1084 (3-2a); 1213 (713a);3545,3546,5570,5972 (3-2a); 6321,6904 (7-13a). Dugues, A., 5 (9-28). Duivenvoorden,J. F. et al., 1207,2319, 2449 (18-20). Duke, J. A. [et al.], 366, 1403, 1437,2276 (7-9); 3536, 3636 (2-3); 3879 (2-4); 3896,4156,4230 (7-9); 4386,4387 (24); 4736 (2-3); 4744,4778,5159,5198,5230,5424,5546 (7-9); 5630, 5636 (2-4); 5653 (9-27); 5805 (7-9); 5814 (8-1); 5836 (7-9); 8102 (2-5); 8392 (2-3); 8423 (7-9); 8445 (24); 8575 (7-9); 8768, 9257 (2-4); 9578 (12-1); 9875 (7-9); 10249 (2-3); 11026 (12-1); 11040(15-ib); 11043 (18-6); 11301 (7-11); 11409 (2-3); 121912 (2-4); 11913 (9-27); 11928 (2-4); 12214 (7-9); 12449 (3-2a); 13045 (9-27); 13090A, 13250 (2-4); 13278 (3-2a); 13525 (7-9); 13546 (2-4); 13583 (5-1); 13766 (14-4); 13908 (2-3); 14001, 14067, 14077, 14294 (7-9); 14352 (9-27); 14463 (2-2); 14601 (5-1); 14629 (2-3); 14754, 14871 (7-9); 15814 (10-4); 16074 (14-4); s.n. (18-18a).

LIST OF EXSICCATAE

Dulmen,A.van,256 (18-13). Dunlap,V.C., 16,40 (2-3). Duque-Jaramillo, J.M.,2580,3600(3-2a);3895(8-1);4003 (12-1);4115A(6-1). Dusen,P.K.H.,2159(9-26b);3548(9-19);3548(9-7);4196, 8012, 10819(9-32);13002(7-2);13193(9-32);14383, 14617(9-7); 15695(9-32);17034(3-2a);17283(7-2); 17759(9-31);s.n. (9-26b,9-34). Duss,A., 1403(9-27); 1404(3-2a);4130 (9-27). Duss,T., 1208(3-2a). Dutra,J.,202 (3-2a). Dwyer,J. D. [et al.], 12, 146 (2-4);245 (9-27);441 (2-3); 443, 1171 (3-2a); 1353(2-3); 1403, 1437(7-9); 1827 (2-4);2276 (7-9);2518 (3-2a);3371, 4010 (2-4);4230 (7-9);4364A,6259,6633,6927,7169(2-4);8826(7-9); 9757(14-7);10841,10841(2-1);14463(2-2);s.n.(7-9). Ebinger,J. E., 51, 149(2-4); 161(7-11); 169, 1126(2-4). Echeverna,J.A., 1024(9-27); 1356(9-10);4162 (9-27). Edwall,G., 1745(9-25); 1768(9-3);2255(9-32);4496 (93);s.n. (9-22,9-30c). J. B., P-43(2-1);387 (3-2a);415,415,460 (6-1); Edwards, 644 (2-3);664 (3-2a). Edwards,M.T., 664 (3-2a);667 (2-3). Eggers,H. F.A., 651, 1797,5359, 5553 (3-2a);13274(927); 14319(7-10); 14359(2-5); 14410(7-10); 14430 (2-3);14640,15642(7-10);15651(8-1);s.n.(8-1,9-27). Eiten,G. et al., 3972 (9-32);5244(7-6a);9179(7-1);9452 (9-30d);9496 (9-30c);9542(9-30d). Ek,R. C. et al., 1027(18-11). Ekman,E. L., 553 (3-2a);2016 (9-32);2017 (9-31);2166 (3-2a);2438 (9-42);2981 (2-3);3580 (9-41);4954 (942);5408(9-34);6035 (9-42);6407(3-2a);6488,7084 (9-37);7285(9-35);7370(3-2a);8285(2-3);8380,8547 (9-42);8635(9-37);8882(9-42);8955(9-43);8990(934);9005,9387(2-3);10210(9-34);10497(9-43);10566 (9-34);12455(3-2a);12836(3-2a);13888(940); 14172 (9-35);14561(9-41);14702(9-35);15001(9-41);15123 (9-42);15248,15472(9-41);16877(3-2a);17973(9-39); 18463(9-40);19110(9-46);115408(9-34). Elburg,J., 13403(4-1). Elcoro,S., 68 (7-6b);776 (9-32). Elias,Bro.,257, 1246(3-2a). Elias,T. S., 1786(2-4). Ellenberg,H., 2463 (7-7);2511 (6-1). Emmerich,M., 298 (9-28);670 (9-30d);897 (9-25);2390 (9-21);4043 (9-30d). Emrich,K.,1136a-b,33936(3-2a). Emrick,G. M., 133(2-3). Emygdiode Mello Filho, L. et al., see Mello Filho, L. Emygdiode. Encarnacion,F., 1072 (3-2a); 1073, 1277, 1279 (7-14); 25049 (10-5);26007 (2-3);26087(2-4);26133(13-4); 26212(2-3);26494(7-14). Eynden,V.vandenet al., 842 (2-3). Enriquez,0. G., 806 (2-3). Emst,A., 1710(9-14). L.C., 113, 123(9-33);198(9-27);s.n. (9-28). Ervendberg, Espejel,I. et al., 38 (2-1). Espina,[etal.], 1 (3-2a);A.33 (8-1);73 (7-13a). Espina,J.et al., 1002,2551,2761,3002 (2-3);3848(2-4). EspinalT., S., 1205,1211(5-1); 1784(8-1). Espinoza,S. [et al.], 100 (14-id); 148 (6-2); 235 (16-1); 447 (14-8b);571 (10-9);615 (14-Id). Estrada,J. et al., 582 (9-27). Eugenio,J., 518 (7-2). Euponino,A., 224 (16-1);363 (9-30a).

321

Eyerdam, W.J., 19(9-42);196(9-37);298(9-42);314(9-43). Eynden,V.vanderet al., 686 (5-3);842 (2-3). D. et al.,635 (18-18a);751(7-12);2054 Faber-Langendoen, (10-9).

Fabris,H.et al., 44, 7256,7780 (9-32). Faden,R. B. et al., 76-127(9-27). Fagerlind,F. G. et al., 316 (2-4);318 (13-8c);2376, 2403 (2-3);2445,2503 (7-10);2528 (2-4);2665 (7-10). Falcao,J. I. Almeidade et al., 993 (9-21). Fanshawe, D. B., 239 (FD 1975)(4-1);337 (FD3073),338 (FD3074)(7-4b);502 (FD3238) (6-3);641 (4-1);736 (6-3); 1144(FD 3880) (7-4b);1303(6-3); 1647, 1812 (4-1);3484 (FD7080)(3-2a);3556 (4-1). Farenholtz, H.,345 (3-2a). Farias,G. L., 343 (7-1). Farney,C. et al., 274 (7-3); 1866(7-6a). Fay,J. J. et al., 852 (9-27). FD(= ForestDepartment, BritishGuiana); 2975(4-1);3073, 3074(74b); 3377(4-1);3880(7-4b);4383,4548,5827 (4-1);7080 (3-2a);7152 (4-1);7629 (7-6a.note);7891 (7-4b). Feddema,C., 257, 731, 965, 1335, 1353, 1448, 1635 (928); 1842(15-lb); 2014 (9-27);2123,2161 (9-28). Fedlmeier, C., 2 (5-2);4 (13-1). Felber,R., 199(2-1). Fendler,A., 363 (3-2a);710, 1286(9-27). Fernandes, A., 438 (9-25);786 (9-26a);1828,2077 (3-2a). Fema'ndez, A., 310(2-3);2778(2-5);4301(7-6b);4530(101If);4614(6-3);5258(10-9);5281,7001,7002(7-6b); 7035 (15-3);7150(7-6b);7492 (6-3). Fernandez Casas,J. [et al.], 3576,4207, 4252, 7361 (7-2); 8249(15-Ib). FernandezN.,R.,1412,1418(2-1). Y et al., 407 (7-6b) Fernandez, Femandez,16508,16561(3-2a). Fernon,B. E., s.n. (1-1). Ferreira, A. R. [et al.], 295 (9-3);424 (9-22);512 (9-25); 920 (9-19);1729(9-6);s.n. (9-3, 9-30d). Ferreira, C.A. Cid[etal.],354(16-2);868(11-1);2559(7-5); 2569(7-6a);2573(7-4b);2648 (6-2);2679 (7-5);2766 (3-2a);2814(7-5);2822 (3-2a);2865,2875 (6-3);2907 (7-1);2957 (3-2a);3093 (7-7);3103 (7-5);3622 (7-1); 4555 (6-3);4841 (6-1);5088 (7-4c);5313(7-4b);6334 (13-3);6506 (10-5);7184 (14-3);7190 (14-1b);7461 (18-19a);7715 (7-6); 8391 (18-7); 8506 (12-2);9337 (10-12);9506 (18-4); 8276 (7-4c);9667 (7-6a);9808 10031(11-1);10104(18-unnamed); (7-1);10018(18-19a); 10261(5-2);10595(11-1);10601(18-1);10799(18-8). Ferreira, E., 5924 (7-1). Ferreyra, R., 4361 (2-5);4373, 12669,17007(2-4). Ferris,R. S., 8805 (1-1). Feucht,S. P.,637 (2-3). Feuillet,C., 715 (7-6b). Fiebrig,K., 182(7-lSb);353 (9-32);411 (3-2a);951 (7-2); 983 (3-2a);4156 (9-30b,9-32);4156a,4295, 4296 (930b);4394 (3-2a);5386 (7-2);6242(3-2a);6404(7-2). Filho,S. F., 17(7-2). Fischer,F. E. L. von (herb.);312 (9-25);s.n. (2-1, 6-3,7-3, 9-3,9-25). Fittkau,12912(14-8a). Fleischmann, A., 487 (2-5). Flores,A., 4829 (7-13a). Flores,L. et al., 15,41 (6-1). Flores,R. S., 10 (2-3). J. et al., 1633(6-3). Florschiitz, P.et al., 2805 (7-6a). Florschiutz,

322

FLORANEOTROPICA

Galarza,M. I., s.n. (5-2). Galeano,G. et al., 667 (13-8a);709 (5-3). Galeotti,H.G.,290,291 (9-27);293 (9-28);332(1-1);7063 (2-3);s.n. (9-27,9-33). Galleoti,H. G., 291 (9-27);332 (1-1);7063 (2-3). Gallinal,et al., 5301 (9-32). Galue,N., 112(7-13a). Fonnegra,R. et al., 1664 (3-2a); 1898 (15-1b); 1967 (2-4); Galvao,R., 391 (7-3). 2120 (6-1);2166 (8-1);2188 (7-9). Gamero,J. C., 1311(9-32). FonsecaZ.,A., 105(9-28). Gandoger, M., 44, 45 (4-1). Fonseca,A. S., 217 (7-3). Garcia-Barriga, H.,3675 (5-3);7519, 10659(3-2a);10988 (7-8); 11030(2-3); 11030(6-1); 11766(3-2a); 13830 Fonseca, S. G., 1008 (9-30c). (13-4);13945(16-2);18049(13-8a). Fontella, J. Pereira [et al.l, 82 (9-3); 84 (9-22, 9-26b); s.n. (9-32). Garcia,J., 186(2-3). Forero,E. [et al.], 427, 429 (7-9);443 (10-1b);615 (7-9); Garcia,N., 4 (3-2a). 692 (2-4);718(8-1);771 (10-5);1135(12-1);1202(14- Garcia,V.et al., CA.368(14-la). 3); 1863 (2-4); 1898 (9-27); 3041 (15-1b); 3817,4158, Gardner, G., 115(9-25);116(9-3);197(9-26a);621 (9-18); 4619, 4874 (2-3); 6361 (7-6a); 6361 (7-4b); 6704 (132001(9-30d);2002(3-2a);2116(9-3);2310(9-32);2310 unnamed);6922,6940 (2-3); 7038 (14-9b); 7673 (7-3); (9-30d);3427(9-30c);3428(9-30d);4570 (9-25);5629 (7-3);5859(7-2);s.n. (9-3, 9-26a). 8979 (7-9). Forero, L., 913 (3-2a). Garibaldi, C., 246 (7-9). Fosberg, F. R., 19235 (3-2a); 28976 (2-3); 28986 (3-2a); Gamier,Bro.A., 920, 1727(3-2a);3019, 4196 (2-1). 47800 (1-2). Garvizu,M. et al., 288 (14-8a). Foster,R. B. [etal.], 699 (8-1);963, 986, 893 (2-4);988 (2- Garwood,N. et al., 79, 123 (2-4);955 (10-5); 1069(2-3); 1111(3-2a);1124(2-5). 3); 1063(7-9); 1133(8-1); 1207, 1395(7-9);3377 (23); 3714 (18-20);3875 (6-1);3915 (13-8);4573 (5-3); Gaudichaud Beaupre,C., 90 (3-2a);93, 94, 95 (9-25);96 (9-26a);262 (7-1);264 (9-30b);273 (3-2a);277 (9-30c); 4647 (18-2 1); 5060 (7-8); 5187 (7-5); 5210 (7-7); 5220 670(3-2a);977,978(7-3);1086(3-2a);1088(7-3);1089 (7-5); 5225 (7-8); 5256 (7-7); 5521 (18-8); 5539 (14-7); (7-1);1090,1090bis(7-3);1091(9-26a);1093,1094,1095 5616(3-1);5626 (14-4);5691 (5-2);5778(9-13);5815 (7-5); 5826 (14-2); 6003 (7-7); 6033 (18-8); 7438 (14-la); (9-25);s.n. (3-2a,7-1, 7-2, 7-3, 9-3, 9-19,9-25, 9-26a). 7443(9-13);7851(14-8b);7955(12-1);8514(7-7);8600 Gaumer,G. F. [etal.], 698 (9-27);726 (3-2a);1923(9-27); (14-7);8691(7-4c);8716(7-1);8796(8-1);8784(13-4); 23302,23320(3-2a);23805(9-27). 8802(18-19b);8845(24); 8851(10-13);8853(7-5);9172 Gay,C., 2165 (1-1);s.n. (9-32). (1-2);9180(6-1);9342(18-19b);9381(9-13);9673(10- Gehrt,A., 8238 (7-3);28366(9-32). 3); 9711,9716, 9718 (6-1);9886 (18-8);9915(10-1b); Gentle,P.H., 18 (3-2a);30 (9-27); 143(3-2a);220 (9-27); 10798 (10-h lf); 10944,11025 (8-1); 11398 (5-1); 11730 831 (3-2a);1447,2704, 3097 (2-3);3211 (2-1);2530, (7-5); 11732 (18-13); 11939 (7-5); 12059 (7-7); 12176 4785 (9-27);4786,4812,4813, 4981,5161 (2-3);6275 (9-13); 12469 (7-13a); 12621 (9-13); 13168, 13382 (7(9-27);6624,6998 (2-3);7234 (2-1);7447,7781 (2-3). 5); 13719(7-1); 13940 (9-30c); 14079 (18-19d); 14115 Gentry, A. H. [etal.],449 (8-1);805,808 (3-2a);919 (9-28);

Floster,B., 164(3-2a). Foldats,E., 145A(6-3). Folli, D. A., 118 (18-10); 159 (10-5);395 (6-2);459, 671 (10-3);772 (16-4);784, 828 (6-3);2056 (7-1). Folsom,J. P. [et al.], 1713(2-5);4577, 4921 (9-27);4975, 4983 (7-9);5130(9-27);5149,5244(7-9);8042(9-10); 8829,8902,8911,9130 (2-5);9209 (7-4a).

(18-18a); 14254 (18-22); 14538 (8-1); 14605, 14609 (16-1); 14698 (7-4c).

Foumier,L.A., 822, 979 (6-1). FrancoR., P.et al., 1267(18-unnamed); 2859 (1-2). Frankie,G. W., 5la, 51c, 68a, 68c (7-4a); 83a, 107a, 148a

(3-2a);21Oa,210c,432a(2-3);632 (2-4). Frazao,A., 361 (9-25);13048(7-3);s.n. (7-3,9-3, 9-26a,930c, 9-32). Freire,E. et al., 2567 (18-i5); 2635 (16-5);2638 (18-unnamed);2849(10-9);3005(17-2);3061(18-unnamed). Freire-Fierro, A., 2280 (7-8). Freitas,M.A. et al., 115(6-2);454 (7-1). Freyreiss,G. W.,s.n. (9-26a,9-30a). Friedrichs, E., 30699(7-2). E. von,s.n. (7-4a). Friedrichsthal, Froehner, C., 32, 350 (2-4). Fr6es,R. de Lemos[et al.], 2 (7-14); 132,410, 852 (6-2); 1783(4-1);1869(6-3);11804,11849(3-2a);12209/122 (7-14);20969,20970(3-2a);21448(7-6a);22148(7-4b); 22163(7-6a);23120(7-13a);23516(6-3);23822(7-5); 23822 (7-6a); 23892, 23898 (12-2); 23999, 24074 (14-

6);28340(14-8a);29385(9-30d);29629(7-14);29629 (7-6a); 29645 (3-2a); 30297, 30546, 31068, 31101, 31204, 31250 (6-3); 32173 (4-1); 33426 (6-3); 33650, 33710, 33782, 33789 (7-1); 34484 (10-2b); 34905 (6-3). Fryxell, P.A. [et al.], 3520, 3556A (3-2a); 3775 (2-3). Fuchs, H. P. et al., 22038 (10-1 lb). Fuertes, M., 11, 190 (3-2a); 1086 (2-3).

Funcke,N. [et al.], 107(9-32);247, 595 (3-2a).

1052 (18-18a); 1660 (2-4); 1712 (3-2a); 1732, 1756 (79); 1845 (18-18a); 1854 (18-17); 2316,2678,3078,3178, 3300, 3323, 3326 (7-9); 3371 (2-5); 3653 (2-1 or 3-2a);

3808(24); 6276(7-8);6378(8-1);6471(7-9);6554(711);6733 (2-4);7237 (10-1lb); 7543 (2-3); 8825 (1819d);8978(3-2a);9379 (13-4);9687 (6-2);9808(2-4); 9845(8-1);9862(12-1);9886(18-16);9911(7-10);9935 (8-1); 9964 (6-2); 9965 (6-1); 10202 (2-4); 12011 (15lb); 12158 (7-10); 12423 (14-1b); 13409 (2-4); 13416 (18-19d); 13731 (13-8c); 13781, 13782 (16-7); 14185 (13-8c); 14408,14470(9-32); 15153 (3-2a); 15229 (9-27); 15336 (3-2a); 15676 (14-5); 15682 (7-5); 15946 (2-4); 16275 (18-19b); 16630(13-8c); 19692 (18-19b); 16765, 17168, 17489 (2-3); 17765 (8-1); 18956 (7-6a); 19034 (7-4c); 20008 (10-lla); 20099 (16-1); 20288 (2-4); 21068 (7-5); 21285 (3-2a); 21906 (14-Id); 22031 (7-5); 22346 (14-6); 22909 (16-7); 24314 (10-1 Id); 24734 (713a); 24947, 25072, 25074 (7-5); 25239 (7-4c); 25244 (18-15); 25388 (18-8); 26555 (16-7); 26557 (13-8a); 26819 (7-1); 26824 (7-5); 26877 (8-1); 27014 (6-2); 27045 (18-8); 27131 (3-1); 27288 (18-2 1);27466 (18-3); 27476 (6-2); 27483 (18-20); 27521 (3-2a); 27772 (5-1);

27815(7-4c);27962(18-20);27980,28048(7-4b);28074 (14-8b); 28245 (2-4); 28590 (18-17); 28636 (18-18a); 29021 (6-2); 29093 (3-2); 29132 (18-7); 29149 (18-20); 29540 (2-5); 29732 (7-5); 29732 (7-6a); 29733 (5-3); 30334 (18-19d); 30683 (7-10); 30724 (6-2); 30739 (13-

8c);30741(3-2);30804(2-3);31198(10-13);31225(10lb); 31239(8-1);31377 (14-7);31402 (18-20);31470

323

LISTOFEXSICCATAE (7-6a);31663(10-3);32060(3-2a);32262(2-1);32312 (6-1);32361,32484,32519,32546 (2-1);34813A(10-lb); 34825A,34829A(7-13a);35387 (16-1);36006 (5-1); 36510(14-6);36547(18-19c);36924(18-22);38020(146);38153(3-2a);39095(18-19c);39201,39472(18-15); 40115(3-2a);40241(14-3);41060(18-19d);41159(144);41522(2-4);41789(10-9);42052(10-5);42134(5-3); 42552(18-20);42689(7-4b);42716(14-6);42733(5-1); 42798(18-20);42816(5-1);42848(14-8b);42927(5-2); 43036 (7-6a);43047 (7-4c);43081 (5-1);43136 (5-2); 43311 (8-1); 43344 (6-2); 43358 (6-1); 43457 (7-7); 43465, 43483 (5-1);43511 (7-7);43548 (6-1);43679 (6-2); 43770 (5-2); 43926 (6-1); 43957 (6-2); 44024, 44017(2-1);45008(7-13a);44579(14-la);45235(133b);45293(6-1);45349(7-8);45331(13-9);45422(7-8); 45454,45475,45475(13-9),45529(13-3b);45552(76b);45617(5-2);45655(6-1);45692(6-2);45778,45802, 45842(7-7);45864(6-2);45883(7-7);46039,46161(62);46552(7-6b);47224(7-4b);47608(8-1);47678(1-2); 47822(13-4);48260(18-17);48499(8-1);48530(5-1); 48532,48545,48585(7-4a);48709(7-8);49122(16-3); 49194(10-5);49309(9-3);50719(13-8);51106(10-2c); 51170(6-2);51190(10-9);51506(6-2);51851A,51858, 51856(7-2);52209(7-4c);52994(6-1);53023(13-6a); 53038,53078(6-1);53156(18-18a);53759,53808(2-3); 53930 (1-2); 54048 (2-4); 54056 (7-8); 54147 (2-4); 54582(7-4b);54637(18-20);54638(3-2a);55007(6-1); 55101 (18-18b);55104, 55352 (1-2);55384, 55384A, 55390, 55396 (2-3); 55421 (12-1); 55422 (10-1b); 55456A(2-3);55731(18-20);55782(6-2);55786(2-3); 55871 (6-2); 55878 (7-4b);55951 (10-1b);55967 (912);55969(7-4b);56216(3-2a);56217(7-5);56288(74c); 56423(18-15);56501(7-4c);56647(14-6);56751 (10-5);57142(3-2a);57176(1-2);57259,57280,57281 (7-4b);57296(134);57301(74b); 57331(14-3);57332, 57342(7-4b);57391(16-1);57533(15-3);57537(7-7); 57563(5-2);57651(14-4);57653(5-1);57727(15-1b); 58063(6-1);58130(6-2);58134(14-3);58140(18-13); 58292 (3-2a);58459 (8-1); 58501 (5-1); 59298 (7-2); 59669(18-18b);59713(1-2);60163,60174(5-3);60191 (10-3);60207 (5-1); 60567A (7-13a);60708 (10-1b); 60810 (15-3);60814 (7-6a);61057 (3-2a);61645(1820); 61767 (18-20);61921 (74c); 62022 (5-3);63044 (10-3);63056(13-1);63332(7-4c);63750(10-9);63825 (12-1);63845(18-unnamed); 63856(16-1);63887(13-4); 63892(18-19d);64070(13-9);64117(6-1);64184(7-5); 64235(13-8c);64241(18-21);64242(7-5);64248(10-3); 64303(18-7);64324(10-1If);64343(13-4);64902(146); 64964(18-10);64967(10-1If); 65417(1-2);65531 (18-18a);65708 (14-2); 65771 (18-8); 65779 (9-12); 65793(18-20);65957(7-5);68492(7-12);68612(6-2); 68695 (5-1); 69049 (7-1);69249 (6-3);68952 (18-6); 69916 (7-8); 70228 (6-2); 70341 (6-1); 70371 (7-5); 70372(14-la);70399(7-5);70400(18-21);70506(10-3); 70519 (14-3);70555 (7-1); 70559 (7-7);70569 (2-4); 70721(10-1b);70780(13-4);71033(16-7);72439(8-1); 72721(3-2a);72814(14-4);73012(7-10);73017(10-lb); 73032(10-5);73073(7-10);73077(6-1);73083(3-2a); 73232(13-9);73239(1-2);73320(7-1);75311(7-13b); 73377(10-lb);74328(18-2);75568(7-1);76663(144); 76731(13-4);76738(14-la);76763(9-13);76856(1821);76876(13-3);77188(18-19b);77192(15-3);77280 (10-3);77328 (18-19b);77607 (14-7); 77734 (16-1); 77794(134); 77803(18-6);77908(18-13);77938(10-9); 77947(10-1lf); 77962(10-lSb);77965,77975(18-19a); 79922(16-6);80210(14-la.note); 78665,78727(18-19d); 80581(7-8);80709(14-la);80898(13-8c);80937(14-2).

Gentry,H. S., 736M(2-3);770 (9-28); 1026(9-27); 1551 (9-28);3641,3657(1-1);5147,5582(2-3);6130(3-2a); 10800,10906(9-28). Geoy,F.,s.n. (3-2a). Gereau,R. E. et al., 3460 (2-5);3461 (6-1). Gevieski,A., 8 (7-2). A. B., 36 (9-27); 258 (9-28); 260, 646, 676 Ghiesbreght, (9-27). Gilbert,E. J., 112(9-32). Gillis,W.T., 9651 (9-27). Gillot,F.X., s.n. (9-25). Gilly,C. L. et al., 355, 361 (2-3). GimateL.,J., 907 (2-3). Ginzberg,S., 76-225(2-3);s.n. (8-1). A. et al., 640 (9-30d);854 (6-3). Ginzberger, Givens,F.M., 3192(9-27). Glassman,S. F., 1888(9-28);2030, 2650 (3-2a). Glaziou,A. F. M. [et al.], 73 (7-3); 110(6-3); 11OA(7-3); 766, 766A(3-2a);777 (9-2);778, 778, 779 (9-25);859, 879 (6-2);1010(9-19);1014,1015(9-26a); 109,1137 (7-3); 1470(6-3); 1471(7-1); 2705 (9-3); 3112 (7-3); 3114(9-30a);4200(9-18);5989(7-1);7831(9-25);7832 (9-25);7851 (7-2);7851 (7-1);7880 (3-2a);8079 (930a); 8079A (9-32); 8080 (9-19); 8289 (6-2); 8933, 11557(9-18);11565(7-2); 11566, 11567(7-3); 11571 (9-32); 12070 (9-30c); 12172 (6-3); 13211 (9-30a); 13212(9-30c);13212(9-30d);14277(9-30a);14281(32a); 16348(9-30a);16356(7-1); 18473, 18493(7-2); 18494,18595(7-1);20489 (6-2);20495(9-18);22142 (9-30c);22143(9-30d);s.n. (6-3,7-2, 9-26a). Glenboski,L. L., C.242(6-2). Glocker,344, 583 (3-2a). Goeldi,A., 666 (7-2); 1587(4-1). G6es,0. Campos[etal.], 32, 34, 35 (9-25);40, 117(7-2); 132(9-25). Goitia,D., 631 (18-18a). Goldman,E. H.,238 (9-28). Gomes,J. Correa,436 (9-18);476 (9-30a). Gomes,M. et al., 266 (14-9b). G6mez,L.D. [etal.],6185(9-28);9872(9-10);20467,20470 (2-5);20945(9-10);23042(3-2a);23050(9-27);23059 (3-2a);23562,23600,23630(9-10);23661(9-27);23739 (7-4a);23744(9-27);24109(2-4);24139(9-10). J. [et al.], 748 (7-4a);8670 (7-8); 9545, G6mez-Laurito, 9861, 10243, 10256 (9-10); 10538 (7-8); 11480 (2-1); 1 1693 (24); 11828 (18-17); 11888 (2-1). Gomez-Pompa,A., 4552 (8-1). Gonggrijp, J. W., 5201 (9-32).

Gonzales,E., 46 (2-3). GonzalesG, J. 37, 138(3-2a). GonzalezQ., L., 28 (2-3); 1468(9-27). J.G.,5558(3-2a);7100(2-3);7374(9-28); Gonzilez-Ortega, 7493(2-3). Gooding,E. G. B., 287 (3-2a). Goodland,R., 218 (9-32). Gordon,17(10-9). Gottsberger, G.,s.n. (7-14). Goudot,J., I (3-2a);s.n. (1-2, 2-5, 3-2a,7-9b,9-27, 9-32). Goulding,M., 2, 14a(7-14);47a (3-2a);103, 1137(7-14); 1486a (3-2a).

Graham,S. A., 162(9-28). W.L.[etal.], 1162(9-27);1172(9-28);1176,1177, Graham, 1178,1189,1190(9-27);1194(9-28);1197,1209,1210 (9-27);1211,1213,1217,1220,1236,1241,1245,1246, 1248, 1254, 1255, 1260, 1273, 1281, 1289, 1303, 1321, 1366, 1367, 1371 (9-28); 1435 (9-27, 9-28); 1469, 1470

(9-27);1487,1520, 1523, 1526(9-28).

324 Grandez,C. et al., 389,429 (7-5);498 (18-6);1488(7-4c); 1521, 1581(7-5); 1608(18-20); 1718(14-6);2053 (32a);2858 (15-1b). J.-J.de [etal.],T.618,1553,2310,3977,B.4283, Granville, B.4622(6-3);5237(8-2);B.5406,5434(6-3);5653(149a);6960, 10438(6-3). Gray,C. E. et al., 2952 (9-28). M. H.[etal.],2110(2-3);2365,2375(7-4a);2628, Grayum, 2629(2-5);2633 (7-4a);3372 (9-10);3551(2-5);4379 (7-4a);4990(13-5);5397(7-8);5485,5709(7-4a);5720, 5743 (2-4);5818 (9-10);5923 (5-1);6167, 7948 (2-5); 8015(14-4);8348(7-4a);8668(8-1);9140(9-10);9671 (6-2); 10142(9-10). J. M.,449 (9-27). Greenman, Gregg,J., 374,624 (1-1). Gregory,L. E. [et al.], 1856(6-1); 11496(9-32). Grenand,P.,675 (6-3). A. [etal.], 278, 421 (2-1);600 (18-17);605 (2-1); Grijalva, 700 (10-2c);783 (3-2a);864,996, 1219(9-28);1239(32a); 1300,2382, 2431 (2-3);2534 (3-2a);2753 (9-28); 2757(3-2a);3360,3555(2-3);3685(2-1);3882(9-28). R. de, 13 (9-27);s.n. (9-32). Grosourdy, Grubb,P.J. et al., 1539(2-4). Gruener, G., 273 (7-2). et al., 20 (9-26a). Guapyassu, E. [etal.],62 (10-13);103(5-3);209 (14-2);248(7Gudinlo, 5);318(14-4);384(10-5);936(14-1b);982(10-tb);1068 (14-8b);1157(18-19a);1386(13-1);1831,2132 (2-3). Guevara, A., 5 (7-9). Guilding,L., 7 (3-2a). Guillemin,J. B.A., 131(7-1);144(3-2a);195(9-26a);256 (9-25);752 (9-4);847 (3-2a);s.n. (3-2a,7-1, 9-25). Guillen,R., 2229(7-13b). Guiquita,et al., 110(13-3). Gurken,L. C. [et al.], 2 (9-8);24 (9-22);37 (9-2);45 (926b); 119, 120 (9-25); 149(9-18); 150(9-2); 1199(93);s.n. (9-18,9-23). Guttie'rrez,E. et al., 554 (7-1).

G. et al., 749 (7-6a);759 (6-3). Guttierrez, V.G. et al., 803 (13-4). Guttierrez, Guzman,M. et al., 186(3-2a);1343(9-28). Haber,W.A. [etal.],22, 26 (3-2a);364,365 (2-1);666, 668 (9-10);773,806,960(7-8);1054(9-27);1076(9-10);1649 (7-8);1702(9-10);1960(18-17);2581,2935(2-3);2948 (9-27);2951(7-8);2968(9-27);3710,3712(9-10);4498 (2-1);4797(9-10);4980(9-27);5426(9-10);5667(7-8); 6667(2-1);7033,7221,7339(18-17);7429,7481(6-1); 7507(18-17);7756(9-10);7848(2-3);8207(2-1);8443 (2-3);9236(9-10);9907(7-8);9941(9-28);10719(7-8). Haccha,O., 11(10-13). Haenke,T., 1468, 1469(2-3). Hage,L. J., 136(7-3). Hage,J. L. et al., 1595,1671(9-21). Hagen,C. vonet al., 1056(9-27);1130, 1177(2-3). Hahn,L., 104,856, 1128(9-27);s.n. (2-3, 9-27). Hahn,W.et al.,454 (13-5);707 (7-13c);1715(10-6);1734 (13-4);2625,2731, 2752 (7-2). Halle,F., 1092(6-3). Hamilton,C. et al.,943 (7-8);1148(2-4);2753(9-lO.note); 2828 (15-tb);2832,3832 (10-1la); 3916 (7-9). Hammel,B. [et al.], 1121(2-4);2051 (7-8); 3221 (9-16); 4993 (9-27); 5085 (9-tO.note);5316 (9-27);7775 (74a);8080,8129(2-5);9598(6-1);9667(2-3);9338(144); 10021, 10020 (2-3); 12113 (2-1); 13071 (10-5); 13821 (2-1); 14054 (8-1); 14082 (2-5); 15535 (2-3); 15536(9-27);16178,16179(2-4);16277(13-8c);16690

FLORANEOTROPICA (2-3); 16838 (8-1); 16990 (18-19d); 17312(1-2); 17692

(5-1); 17828(2-5); 17859,17947(9-10);18241(5- 1). Hampshire, R. J. et al., 124(2-1). Happ,G. B., 31 (3-2a). Handro,O., 19 (9-32);s.n. (9-25). Hansen,B. et al., 1508,1528(9-28). Hansen,B. F.et al., 7380 (9-27);7483, 7484 (3-2a). Hansen,O.,79, s.n. (3-2a). Happ,G. B., 31 (3-2a). Harley,R.M.[etal.],8345(9-1);10054,10649(7-1);10949 (9-30d);10990(9-30c); 15009(9-21); 16140(9-30a); 17553, 17899(7-3); 18170(7-1); 18170, 18335(7-3); 18355(7-1); 18438(7-3). Harling, G. [etal.],3617(2-4);4499(7-10);4676(8-1);4692 (6-1);5000 (7-10); 11117(7-4b);11685(2-3); 13049, 13878(2-4);17651(7-6a);17767(10-lb);18208(3-2a); 19387(2-4);21069 (7-13c);22313 (7-8);24174(2-3); 24190(7-8);24269(2-4);24741 (7-10);24985(2-4). Harmon,P., 127(2-3); 163(6-1). Harmon, W.E. [etal.],3382(2-3);3542(3-2a);5779(9-27). Harris,E. M. et al, 1133(7-13a). Harris,S. A., EC-4(3-2a). Harris,W.,660,5231 (3-2a);5550,6075(2-3);8690,8808, 9266 (9-34); 10211(2-3); 10607(9-36); 10858(2-3); 12627(9-34);s.n. (3-2a). Harrison,s.n. (9-3).

Hart,J. H., 623 (3-2a);3796 (9-27). C.V.,38 (9-27);51 (9-24);974 (1-1). Hartman, R. L., 12042,12064,12513(2-4). Hartman, Hartshom, G. S., 713 (18-8);885 (18-18a);925 (7-4a);952, 960,961,1072(2-3);1091,1094,1323,1409(2-5);1477 (6-1); 1566(2-5); 1664 (5-2); 1691, 1692(6-1); 1793 (7-8); 1884(14-4);2004 (5-2);2073 (6-2);2106, 2107 (2-3);2401 (7-1);2408 (18-13);2460 (7-2);2665 (153); 2990(16-3);3723 (18-13). Hartweg, Th.,386(9-28);514,519 (1-1);1387(3-2a);1390 (1-2); s.n. (9-27). Hassler,E., 542 (9-32);569, 678 (7-2); 927 (9-32); 1071 (7-13b);1366, 1519, 1519a(3-2a);3023 (9-32);3214 (7-2);3368(3-2a);3438(7-2);6242(3-2a);6404(7-2); 6417 (3-2a); 6455 (7-13b); 6917 (3-2a); 7329, 7329a (3-2b); 7338 (7-13b); 7338a (7-lSb); 7598, 7598a (930b); 7847 (9-32); 9413, 9413a (7-2); 9706 (7-15b);

11883, 11883a,11883b(7-2); 12125, 12125a(3-2a); 12203 (7-2); 12222, 12222a (7-13b). Hatch, W. R. et al., s.n. (9-27).

W.H., 1131,1435, 1436(3-2a);1489(7-8). Hatheway, Hatschbach,G. [et al.], 1147 (9-7); 1521 (7-2); 3194 (932); 5185 (9-7);5474 (9-32);6396 (3-2a);7242, 7243 (7-2); 7632 (9-32); 7887 (9-7); 11234 (7-2); 11356, 11638(9-32);11661(7-2); 12038, 13446(9-7); 14078 (3-2a); 14137(9-7); 14629(7-2); 15254(9-7); 15261 (7-2); 15700 (9-7); 16600, 19318 (7-2); 19910 (9-7);

19999(7-2); 22024 (9-32);22461 (9-7); 22592 (7-2); 23664 (9-32);23741 (9-30d);25079 (3-2a);25087 (930c);25262(9-32);25780(9-3);26006(9-7);29708(72); 30421(9-32);30498(9-30d);30500(9-30c);30893 (9-7);31156(9-32);31827(7-2);32898(9-30a);33018 (9-32);33281 (9-30c);33296 (9-30d or 9-32); 33358 (9-30d);33745(9-7);33750,37042(7-2);37727(9-32); 37772 (9-30c);38550 (9-7); 38671 (9-32);40219 (32a);42121(9-32);42123(9-30d);42164(9-30c);43252 (9-32);45198,45245,45300(7-2);47136(3-2a);48790, 49883 (7-1);49887 (6-3);49956 (9-18);50348(9-32); 50587(7-2);51594(9-30b);51596(9-32);55586(9-24). Haught,O., 1864(2-3);2274,3064(9-27);3487(2-5);3644 (9-27);4224 (2-3);4309 (3-2a);4548 (9-27);4808 (2-

LISTOFEXSICCATAE

325

(9-30c);12911(3-2a);28318(7-2);31010(3-2a)36556 5); 4819 (2-3);4901 (9-27);5555,5564 (2-5). (9-32);s.n. (9-25,9-26a,9-26b,9-30c,9-32). Hauman,L., s.n. (9-31,9-32). Hoffman,B. et al., 1177, 1545,3632,3268(7-4b);3959(6Hawkins,K., 6 (3-2a). 3); 3968(15-la); 3969 (11-1). Haxiare,C., 3865 (18-15). L.R., 1224,1225(3-2a);1310(2-4);2525(2-3); Holdridge, Hayden,S. M.V., 1032(7-9). 6233(2-4). Hayes,S., 8 (7-13);658 (2-3);682, 684 (7-9);952 (3-2a); Holm-Nielsen,L. et al., 272 (1-2); 21090, 21678 (2-4); s.n. (2-4). 22709(7-5). Hazlett,D. L. [et al.], 1968(9-27);2356 (2-1);2413, 2645 Holm,R. W.et al. 94 (2-4); 103, 229 (9-27);346 (9-10); (2-3); 3156 (2-1); 3393 (13-5); 5147 (9-10); 5199 (9389 (7-4a). 27);5243 (7-8). Holmes,W.C. et al., 3485 (3-2a). Heath,M. et al., 79 (2-2); 111(2-1). Holst,B. K. et al., 2446 (15-1b);4555 (9-17). Hedger,J. N., 20 (9-27). Holt,E. G. et al., 121, 122,390 (9-27). Heithaus,R.,305 (9-28). Holton,1.F.,234, 250 (3-2a). Hekker,F.et al., 10088(18-unnamed). Holway,E. W.D., 520, 634 (9-27). Hemmendorff, E., s.n. (9-30c). Hooker,W.(herb.);s.n. (3-2a). Henkel,T.W.et al, 1233(7-4b). Hooper,E. D. M., s.n. (3-2a). J., 2432(9-28). Henrickson, Hoover,W.S., 144(2-1);827 (18-17). Hensold,N., 1101(7-9). Hopkins,M. J. G. et al., 752 (10-3);616 (10-5); Henz,E., 27552(9-31). J.W.,837 (3-2a). Homemann, Bro.,4 (9-27). Heriberto, E. P.[etal.],21 (9-25);398 (7-1);735,785 (9-32); Homer,C. et al., 163(15-4). Heringer, 920(9-25);934(9-26b);1131(9-32);1137(16-1);1447 Houllet, R., s.n. (7-3). (9-30c); 1649(7-1); 2488 (9-25);2986 (9-26a);5349 Houssin,J.,4 (6-2). (7-2);5469,5516(3-2a);5557(7-1);5641(9-32);6343, Houstoun,W.,s.n. (9-28);1730(9-27). 6368 (9-30c);7358 (3-2a);8937 (9-30d);8937(9-32); Howard,R.A. [et al.], 69 (9-nom.dub.);488 (9-10);4881 (9-42); 5739, 6452 (9-40); 8113 (9-42); 8290, 8810, 9234(9-30c);9450,9714(9-32);10732,11408,13091, 14226 (9-30c); 14350 (9-32); 14878 (9-30d);14878, 8812, 1357313691(3-2a);14168(9-34);14557(3-2a). 14880(9-32);15598(9-30d);16615(9-25);16660(9-30c); Howell,J. H., 1 (8-1); 100(10-1la). 17112(9-21);18650(9-30c);s.n. (9-26b,9-30d,9-32). Hoyos,J. et al., 130(7-6b). (I0-l lf); 928(2-3);1188(14-la);1190(2Huashikat,V.,696 H.A. von,5153 (3-2a). Hermann, 3);1346(7-6a);1359(15-3);1703,1835(2-3);1960,1997, Hema'ndez G.,J., 661 (3-2a). 2069 (7-6a);2289 (18-19a);2350 (18-6);2355 (5-3). Hernandez G., H., 241 (6-1);266 (8-1); 1217(10-5). E.A., 5 (9-27). Hubbard, Hemandez,H. M. et al., 415 (9-27). HemrndezM., R [et al.], 405 (9-28);685 (9-27);709 (9- Huber,J., 1404,4300,4339(3-2a);4367 (6-2);6894,6965 (6-3);6977(7-1);6977 (7-5);s.n. (9-32). 28);709 (9-27);1573(2-1);5688 (1-1). Huber,O.,200 (8-1);604 (9-32). Hernandez X., E. H. et al., 295 (2-1). D. LI.,249 (2-3). Herrera Ch.,G. [etal.], 11(2-3);428 (9-10);482 (2-1);561 Hugh-Jones, (7-8);575 (2-1);681 (7-8);841 (7-9);1029(10-5);1043 Humboldt,F. H. A. von (& A. J. A. Bonpland),570, 1484 (9-27); s.n. (3-2a). (7-9); 1087(9-10); 1261(7-8); 1352(2-3); 1448, 1741 (9-10); 1744(13-8); 1789(2-3); 1934(2-5);2018 (18- Hunnewell,F.W.,9874 (9-26a). A. F.,5830 (3-2b). 17);2136 (6-2); 2265 (2-5); 2588 (2-1);3004 (9-27); Hunziker, Hurtado,F. et al., 947 (14-id); 952 (6-1); 1300 (10-13); 4313 (5-1). 1316(6-2);1382(6-1); 1702(7-5); 1870, 1884(7-4b); H. [etal.],202 (7-8);291 (2-3);618 (7-8);709 (2Herrera, 1963(2-4); 1973,2190 (7-5);2234 (74b); 2235 (7-5); 4); 806 (2-3);975 (7-11); 1664(7-4b). 2255(7-4b);2260(7-5);2270(6-1);2287,2307(7-4b); Herter,G. [etal.], 361, 361a,361c(9-32);5028 (3-2a);s.n. 2310, 2486, 2508, 2543 (7-5);2672 (14-ic); 2693 (5(9-32). 3); 2763(2-4). Herzog,T., 393, 668 (3-2a);682 (5-2); 1299(9-32); 1423 (3-2a);1526(3-2b);1630,1630(7-1);1729,s.n. (9-32). IbarraM., G. [et al.], 434, 450 (2-1);529 (6-1);541 (8-1); Hess,W.E. et al., 3573 (9-27). 548 (2-1);646 (8-1);720,1430 (6-1); 1476(2-1);1777 Heyde,E.J. [etal.],89,362, 2962,6236,6237(9-27);6239, (8-1); 1894(6-2);2260, 2348,2436 (2-1);2522 (6-1). 6329 (2-1);6382 (9-28). T. S., 671 (7-13b);672, 921 (3-2b);2583 (9-32). Ibarrola, Heyde,N. M. et al., 128a(4-1). Idrobo,J.M. [etal.],912 (9-27);2896 (16-7);8629(7-4b); Hiegro,G. S., 11 (3-2a). 10022(6-1); 10158(7-13a);11226(10-9); 11325(14G. et al., 479 (3-2b);479 (3-2a). Hieronymus, lb); 16583(8-1). Hijman,M. E. E. et al., 211 (4-1);228 (6-3). E., 89 (3-2a);236 (8-1). Hinojosa,1.et al., 1963(9-32);1096(6-2);1102(8-1);1423 Ijjasz-Madriz, Iltis,H. H. et al., 168(1-1);603 (9-28). (10-3). G.B., 1298,1464(9-28);3165,3181,3186(2-3);3462, Irgand,et al.,s.n. (9-31,9-32). Hinton, 3613,3627(1-1);3643(2-3);3843(1-1);4940,5078(9- Irvine,D., 424,694 (7-4b);818 (7-6a). 28);6750(2-3);9172(9-28);10298(2-1);10590(9-28); Irwin,H. S. [et al.], 2254 (6-3); 5515 (9-30c);6790, 6881 (9-30d);6910 (7-1);7176, 7620 (9-30c);7630 (9-32); 11540(2-3); 14354,15119(9-28);15901,16291(2-3). 8606 (13-4); 10408 (9-30c); 10408, 10501 (9-30d); Hiriart,P.et al., 137(1-1). 10568(9-30c);10675(9-32);10994(9-30c);11020(9Hitchcock,A. S. et al., 6906 (9-33). 32); 11968,15589(9-30c);16593(13-4);19024(9-30c); J.A., s.n. (9-27). Hjalmarson, 19091 (7-1); 19155 (9-30c); 21611 (9-30d); 25413, Hladik,A., 99 (2-4);212 (8-1);406 (2-4);411 (8-1);450, 26265(9-30c);26892(9-30d);47968 (11-2). 483 (2-4);544 (7-13);545 (2-3). Hoehne,F.C. [et al.],4657 (3-2a);5679 (6-2);5823,5824 Izawa,K., 7a (6-1).

326 Jack,J. G., 5002, 5064, 5658, 5842, 5998 (2-3);6062 (32a);6777 (2-3);7884, 8108 (9-40). Jacques,J., 31 (2-3). Jahn,A., 503 (3-2a). Jahn,G., 2 (6-3). M. J. et al., 3055 (11-2);3072 (15-2);3767 Jansen-Jacobs, (4-1);3957,4317 (7-13a);4436 (9-32). Janssen,A.129(10-6). Janzen,D. H., 10111(3-2a);10646(2-3); 11912(2-5). Jangoux,J.etal.,443 (10-5);1644(6-3);1716(9-30d);1744 (9-32). Jaramillo,D., 236 (7-13a). Jaramillo, J. [etal.], 2628 (7-6a);4255 (14-7);6597,6791, 7055 (7-12); 13819(18-19d);13905(6-1); 13939(143); 14431(14-la); 14641(1-2); 14693(7-4a); 14698, 14704(7-6a);14763(7-8); 16040(7-5); 16252,16258 (74b); 16321,16451,16524(7-5); 17182(10-2c);17280 (8-1);30702(7-4a);30774(7-4c);30951,31109,31323 (7-6a);31326,31328 (7-4c);31580(18-20);31743(23); 31748(7-6a). Jaramillo A., J. L., 49, 133(6-1). N. et al., 1106(12-1);1223(10-13). Jaramillo, JaramilloM., R. et al., 614 (14-la); 2080 (2-3). Jardim,A. et al., 359 (3-1);614 (6-2). Jardim,J. G. et al., 1706, 1737(9-2);28517(7-3). Jativa,C. et al., 223, 296 (6-1);297 (13-8c);301 (8-1). Jeffrey,C. et al., 2273 (9-27);2368 (2-3). Jenman,G. S., 2141(3-2a);2480(7-4b);5067,5869(3-2a). Jenssen,E., 49 (10-1b). Jesu's,J. 0. de, 65 (3-2a). Jiggins,C. et al., 175(7-8). Jimenez,J., 5050(9-41);5165 (2-3). JimenezL., O., 32 (9-27);s.n. (9-10). JimenezM.,A.,597(3-2a);1023(9-10);1051(7-14a);1091 (2-5); 2453 (9-10);3237 (1-2); 3279 (7-4a);3280 (910);3624,3629,3631,3678 (2-3). JimenezM., O., 528 (2-1);733 (7-8). Jimenez,Q. [etal.], 783 (7-4c);825 (7-8). JimenezSaa,H., 1839(8-1). J0rgensen,P.,1503,2156,2157 (7-13b);2159(3-2b);3835 (3-2a);3954,3984(7-2);4286(9-32);4502,4503(7-13b). Johanessen,C. L., 325 (3-2a). Johansen,H., 3 (3-2a). Johnson,H., 514 (2-3). Johnston,A., 230 (9-27). Johnston,I. M., 1582(7-13);1584(2-4);1693(7-13);1703, 1733(2-4). Johnston,J. R., 998 (9-27);1358(1-1). Johnston,M.,s.n. (9-33). Johnston,M. C. [etal.],5690B,5733(3-2a);8263, 11929a (1 -1).

Joly,A. B., s.n. (9-3, 9-30c). Jones,G. C. et al., 3130 (9-17). Jones,G. N., 22731,22734(1-1). Jones,J. et al., 9255 (2-4). Jones,M. E., 59 (9-28). Jonsson,G., 969a(7-2). Josse,C., 581, 601 (3-2a). Jouy,P.L.,s.n. (9-28). Judziewicz,E. L. et al., 5138 (2-3). Juncosa,A., 1731(2-3); 1245(14-le); 1851(2-4). Jurgensen, C., 941 (2-1). Juzepczuk,S., 5530,6690 (3-2a). Kalbreyer,W, 851 (9-27).

Kallunki,J. et al., 532 (9-9);562 (6-3);708 (7-3). Karsten,H.,s.n. (2-3, 3-2a,6-1, 9-25, 9-27).

FLORANEOTROPICA Karwinski,L. B., s.n. (9-27, 9-28). Karwinsky,W. von Karwin, 700 (9-27); 701 (9-33); 791, 793 (2-3); 802, 802b, 802c (3-2a); 1430 (1-1); 7938 (23); s.n. (9-28). Kautski,R. A., 199 (9-25); 383, 606 (9-18); 676 (9-20); 711 (9-18); 724, s.n. (9-25). Kawasaki, M. L., 281 (7-4b); 341 (5-2); 387 (7-1). Kayap, R., 40 (13-3); 107 (2-3); 113 (5-3); 125 (2-4); 136 (2-3); 159 (5-3); 184 (7-4c); 234 (14-ic); 362 (7-4c); 383 (14-1b); 394 (18-15); 611 (7-4c); 635 (18-19a); 926 (14-3); 996 (10-13); 1030 (14-Id). Keil, D. et al., 8101 (9-28); 9223 (9-27). Kennedy,H. et al., 366 (24); 897 (9-26a); 1130,1133 (7-9); 1211 (7-9); 1630 (9-10); 1830 (7-9); 1849, 1865 (2-4); 2124 (7-4b); 2778 (2-4); 2922 (10-hla); 3158 (7-11); 4574 (9-10). Kennedy, P. B., 7038 (1-1). Kenoyer,L. A. [et al.], 322 (8-1); 323 (2-5); 324 (7-9); 3515 (3-2a); 3731 (9-33); 3770,4067 (2-3); s.n. (9-27). Kern, P. M., 822 (9-10). Keman,C. [et al.], 15 (2-3); 224 (7-4a); 374 (6-1); 375 (8-1); 406 (7-4a); 536 (2-3); 588, 596, 608 (5-1); 686 (7-4a); 834 (13-8); 864 (10-4); 866, 871 (6-2); 918A (6-1); 945 (8-1); 994 (6-2); 1072 (6-1); 1093, 1098 (7-4a); 1106 (3-2a); 1150 (18-19d); 1207 (6-2). Kesber,E., 395 (2-3). Killeen, T., 2877 (8-1); 2973 (7-1); 2977, 3679 (2-4); 3726 (7-7); 4109 (1-2). Killip, E. P. [et al.], 3000 (3-2a); 11479 (2-3); 12129 (7-8); 14297, 14355 (9-27); 16230 (3-2a); 16359 (9-27); 21013 (3-2a); 23464, 23816, 25057, 25369 (2-4); 26485, 27531 (7-5); 29574, 29576 (14-7); 27685 (2-3); 29639 (9-12); 35392, 35580 (2-3); 38232 (7-13a); 39584 (32b); 45593 (2-3). King, R. M. [et al.], 755 (9-28); 815 (9-27); 1744, 1849, 1904,4820 (9-28). Kings, W., 408 (3-2a). Kirkbride,J. H. [et al.], 493 (7-1 1); 554 (2-4); 569 (7-4a); 1226 (18-17); 1227 (13-4); 1393 (7-1 1); 1397, 1670 (24); 1962 (9-15); 2572 (2-3). Klein, R. M., 21 (7-2); 160 (3-2a); 824, 1553, 1572, 1586, 1602, 1688, 1750, 2259 (7-2); 6985 (9-7); s.n. (7-2). Klitgaard,B. et al., 278 (7-8). Klug, G., 153 (7-6a); 257 (18-7); 739 (7-4c); 2079 (2-3); 2607 (3-2a); 3566 (7-8); 3698 (6-1); 3751 (2-3); 3787 (2-4); 3794 (2-3); 3984 (7-6a); 4088 (14-4); 4161 (7-7); 4390 (9-27); 5907 (2-3). Knapp,S. [etal.], 1039 (9-27); 1496 (7-8); 2044 (2-5); 2400, 2451 (2-4); 2472, 4130, 4539 (7-9); 4561 (6-1); 4589 (2-4); 4990 (7-8); 5142 (7-1 1); 5473 (6-1); 5610 (7-8); 5824 (2-4); 5902 (7-1 1); 7085 (2-3); 7241 (14-7); 7245 (8-1); 7288 (2-3); 8459 (7-4c). Knight, D., 1008 (2-3); 4037 (7-13); s.n. (2-3). Knobloch, I., 96 (1-1). Koch, R. G., 4841 (9-28). Kock, S. D., 7750 (1-1). Korning, J. et al., 47180 (9-12); 47481 (15-3); 47511 (103); 47544 (18-2 1); 47566 (13-1); 47609 (14-8b); 47752 (14-4); 58632 (7-4c); 58645 (18-7). Kral, R., 6143 (3-2a). Kramer,K. U., 1740 (9-34). KrapovickasA.[etal.], 12368, 13428, 13907, 14065, 14218, 15103, 15445, 15972, 16179, 16329 (9-32); 16884 (7-2); 17905 (9-32); 19924 (7-13b); 20400 (9-32); 23237 (926a); 23311,24198,24585 (9-32); 26224 (9-3 1); 26411 (7-2); 31181,31204,31454 (9-32); 31524 (9-30c); 31786 (9-30d); 31879 (9-30b); 32155 (9-32); 32170 (9-30d);

LISTOFEXSICCATAE (9-32);36421 32272,32779,33092,35121,36131,36200 (9-30b);36608(9-32);36610(9-30b);37187(9-32). Kress,W.J.et al.,3809(7-4a);4356,4040,4789,4897(7-8). Krieger,L., 7807 (9-25). KroellS., B., 187,203 (7-7);514 (15-3);619 (10-l lf); 728 (5-3);772 (5-1). Krug,H., 2833 (3-2a). Krukoff,B.A., 1054(6-3);1304(7-1);1309(6-3);1314(76a); 1349(7-1); 1450(6-2); 1657(9-30d);1669(6-3); 4570 (3-2a);4954 (7-5);5008(6-1);5084 (7-4c);5173 (6-2);5250,5290,5352(7-6a);5380(6-2);5513(7-6a); 5785(7-5);5969 (7-1);5977,5982 (7-6a);5991,6033, 6049 (6-3); 6143, 6223 (7-1); 6327 (6-2);6422 (5-2); 6628 (6-2);6849 (6-3);6988 (7-6a);7039 (7-1);8157 (7-4b);8312(6-2);8520 (7-6a);8642(15-lb);8648(74c); 8666 (6-2);8768 (14-6);8849 (7-1); 10109(6-2); 10135, 10277(2-4); 10282(8-1); 10388(2-4); 10410 (6-1); 10533(6-2);10667(8-1);10679(6-2);10728(61);10744(6-2); 10813(6-1); 10819(7-5);10819(6-3); 10932(7-5). J. G., 89,95 (3-2a);127(9-18);178(3-2a);256 Kuhlmann, (7-6a);300 (9-30a);400 (7-6a);673 (6-3);735 (7-1); 1103(7-6a);1399(5-3); 1517(7-4c); 1638(7-1); 1649 (9-32); 1739(9-3); 1906(6-3); 1985(9-30d);2076 (71);2510 (9-25);3537 (9-30c);6458 (6-3);6490 (9-8); 6500 (9-18);6589 (9-30a);13055(6-2); 19292(6-3); 19816(5-3); 19831,19853(6-3);34261(3-2a);s.n. (92,9-3, 9-18, 9-26b,9-30c,9-30d,9-32). Kiihn,E., 21 (3-2a). Kukle,P., 11(16-5);82 (7-6a). Kummerow, R., 1099(9-32). Kuntze,O.,s.n. (3-1). Kvist,L. P.et al., 40255 (8-1);40713 (6-2);40811 (10-5); 48985(8-1);49060(3-2a).

327

W.C. [etal.], 388, 1316(9-28). Leavenworth, Legname,P.R. et al., 7232 (3-2a). Legrand,C. D., 1146,2251,2652, 3438,4013 (9-32). Lehmann,F. C., 2326, 2382 (3-2a);3873 (2-4);4738 (927);5093,7337,9063 (3-2a). Leite,J. E.,413 (9-32);415, 416 (7-2);2330 (3-2a). Lems,K., 104(6-3);5508 (9-27);s.n. (6-3). Lent,R. W.,86 (9-28);691 (9-10);692, 1143(9-27);1419 (7-4a);1791(9-10);2243(24); 3043(9-10);3367(2-1); 3603(9-27);3611(7-8);3632, 3724 (9-10). Leon,Bro. [et al.], 2668, 6610 (3-2a);7078, 7431 (2-3); 8092(946); 10778(9-35); 12362, 12680(2-3); 13983 (9-40);20041,22375(2-3). LeonC., J.,s.n. (8-1). Leon,H.,252 (5-1);383 (7-9);600 (2-4); 1329(7-4b). Leon,J., 848 (7-9a);874 (9-28);877 (7-8); 1691, 1701(74a); 1749(24); 1760(9-27);1885(6-2);26692,26693 (9-27);3221 (2-3);4203 (6-1);s.n. (10-3). Leonard,E. C. [etal.],4163 (3-2a);7603, 7780 (2-3);471, 10135,11330(3-2a);11360(2-3);11630(3-2a);12772 (9-42.note);13123,13129,13130,13368(3-2a). Leoni,L., 3111,3112 (6-3). Lepiz,J. F.et al., 192(13-8a). Leprieur, F.R. M. (herb.);s.n. (9-27). J.,41 (3-2a). Lescarboura, LeSueur,H., 570 (2-3);598 (1-1). Levy,P.,45 (9-28). Lewis,G. P., 1524(6-3). Lewis,M.,40611,40685A(7-1). Lewis,W.H. et al., 2188 (16-1);3271 (7-4a);5568 (2-3); 5568(2-1);9784,9785,9786(3-2a);10514(5-3);11303 (3-2a);11363(5-3); 11507(8-1); 11641(5-3); 11735 (7-5); 13628(3-4); 14039(6-2). Lewton,F.L., 312 (9-27). Lhotsky,J., 11(7-2);s.n. (9-25). F.M., 14212,14213,14216(1-1);14224(3-2a); Liebmann, L. [etal.], 1137,1168(9-32). Labouriau, 14225(1-1); 14259(6-1); 14260(2-1); 14267, 14268, Lachette,P.,170(9-41a);171(9-3);247(9-26a);286(9-22); 14269, 14270, 14271(2-3); 14272(3-2a); 14274(13287,288,289 (9-25). 5); 14276(2-1);14298(3-2a);14300,14302,14318(9Laguna,A., 265, 391 (3-2a);s.n. (2-3). 27); 14349,15110(2-1); 15111(2-3);s.n. (2-1, 2-3, 3Lamb,F.B, 141(16-1);156(12-1). 2a, 6-1, 9-27). Lamotte,S., 450,470 (3-2a). Liesner,R. L. [etal.], 11(2-4);969 (7-9); 1347(24); 1840 L. R., 2071(9-19);2414 (7-2). Landrum, (104); 1984(7-4a);2277,2403(3-2a);2909(5-1);2939 Lang,H. et al., s.n. (7-4b). (74a); 2971,3106,4461,4958(2-3);5317(9-27);5753 J. H., 3111 (3-2a). Langenheim, (9-32);6219, 6275, 7367 (7-6b);7430, 7474, 7579 (7Langlasse,E., 214 (9-28). 6a);7638,7695,7844,7924 (9-27);8300,8430(3-2a); Langman,1.K., 3586 (2-1). 8431(9-27);12438(7-13a);13635(74b); 13934(7-6b); G. H. von [etal.],s.n. (7-3, 9-3, 9-19, 9-21, 9Langsdorff, 14436(9-10);14856(7-8);15286,15332(7-4a);15397 25, 9-26a,9-30c). (10-5);15824(15-3);16248(13-1);17840(7-6b);17846 Lanjouw,J. et al., 2433 (4-1). C. H. [etal.], K-24(9-27);36630(9-10). (6-3); 18285,18372(7-6b);20037(15-lb); 21542(13Lankaster, 4); 26090,26688(9-10). LannaSobrinho, J.de Paula,27 (3-2a);139(9-3);176(7-3); 278 (9-26a);1237(7-2);1746(9-26a);1813,1819(6-3); Lillo,M.,7239,7243(3-2b);7245(3-2a);7277(7-2);10828, 10845(3-2b). 1865(9-26b);s.n. (9-32). Lim,A. P.,31 (7-9). Lanz,C. et al., 644 (8-1). Lao,E.A. [etal.], 2a, 22 (8-1);289 (2-4);312, 315 (3-2a); Lima,A., 73 (9-25);227 (9-30c);262 (9-22);265 (9-25); 1165(9-18);2271 (9-3);2272 (9-22);2357 (9-18);53335 (1-2);441 (2-3);448 (8-1);517 (7-9). 1227(7-1). LaoM., R. et al., 14 (13-3);1664(5-2). Lasser,T., 1057(9-27);1194,1248(3-2a);2054(2-3);2570 Lima,J. [etal.], 202 (7-6a);209 (7-5);750, 787 (7- 13a). Lind,K., 10(9-27). (9-27). J. [etal.],3685,3712(4-1);4778(6-3);5127(9R.M.,881(3-2a);1144(9-27);1328,1678(3-2a). Lindeman, Laughlin, 26b);5178 (6-3);5475 (7-2); 6161, 6458, 6776 (6-3); Lavastre,Bro.B. A., 1928(3-2a). 8891(3-2a);8985,20535(7-2). Lawesson,J. E. et al., 39466(9-12);39562(10-1lf). Lindeman,J. & J. H. de Haas,225 (7-2); 576, 777 (3-2a); A. E., 730 (7-4b). Lawrance, 790,900, 1245(7-2);1671(3-2a);1719(7-2); 1853(3Lawton,R. O., 1292, 1293(2-1). 2a); 1988,2225,2842, 3380 (7-2). Lazor,R. L., 5476, 5483 (2-4). J.& E.A. Mennegaet al., 125(7-6b);134(6-3); Lindeman, LBB(= LandsBosbeheer,Surinam),9466, 13403(4-1). 403 (4-1). Leao,et al., 229 (9-3).

328 Linden,J.J., 34 (2-3);37,63 (9-27);64 (9-17);916 (9-27); 1619(1-2); 1647(2-1);s.n. (3-2a,9-27). Lindley,J. (herb.);s.n. (9-26a). Lino,A. M., 37/72 (6-2). Lindman,C. A. M., A.2411 (9-30c); 2495 (9-30d);3855 (9-32). Lins,27 (9-26a). Liogier,A. M. [et al.], 3590 (9-44); 5079, 13812 (9-42); 14490,16511(9-41);16713(942); 23674(2-3). Lira,C. M. S., [etal.], 278 (9-3);349, 411 (9-26a). Lisboa,P.et al., 4668 (9-25). Lissot,J.,s.n. (15-2). Lister,J. R. A. et al., 2133 (6-3). Little,E. L. [et al.], 164 (14-ic); 226 (13-3);297 (13-8c); 305 (3-2a);314 (6-2);322 (10-3);337 (13-3);340 (142); 341 (6-1);381 (14-2);389 (5-3);413 (2-4);426 (136c);434 (6-2);458 (3-2a);507 (14-ic); 521 (14-4);522 (18-13);529 (13-1);560 (13-4);571 (6-1); 652 (6-2); 660 (5-3);734 (14-3);754 (6-1);6192 (7-10);6204 (61);6219, 6222 (6-2);6235 (3-2a);6240 (6-2);6297 (81); 6375 (7-10);6453, 6462 (3-2a);6477 (6-1); 6478 (7-10);6482,6499(8-1);6510(3-2a);6516(2-4);6531 6735 (8-1);7261, 7311 (3(6-1);6639 (18-unnamed); 2a);8472,8481(2-4);9564, 9569(2-3);9572,9651(75);9670(2-3);9778(3-2a);15088(7-13a);15108,15187 (3-2a);15565(1-2);15882,15883(3-2a);16154(2-3); 16211(8-1); 16407(3-2a);16457(3-2);20010(7-10); 20144,20270(2-3);21017(7-10);21126(16-7);21170, 21183 (6-1); 21188 (6-2); 21203 (8-1); 21232 (6-1); 21233(3-2a);21910,25215,25232(2-3);25287(6-1); 25362(2-3);26025(6-1);40166 (3-2a). Lizot,J.,s.n. (7-6b). Lleras,S. etal., P.16931(7-5);P.16988(14-6);P.17067(76a);P.17226(14-9b);P.17371(14-3);P.19439(7-6a). Lobo,M. G.A. et al., 84 (6-3). Loefgren,A., 502 (9-32);1366(8-2); 1421(7-2);s.n. (9-7, 9-26b,9-30d). Loiselle,B. A., 203 (7-4a).

FLORANEOTROPICA 19647 (2-2); 19647 (2-1); 19722 (2-2); 19814 (2-3); 20023(9-17);20237,20502,20541(2-3);20674,20719 (2-1);20956(2-2);20966(13-5);20983(2-1);s.n. (9-27). Luschnath, B., 50 (9-21);239 (9-25);s.n. (6-3, 7-1, 7-3, 93,9-19, 9-21,9-25, 9-26a). Luteyn,J.L.et al.,540, 1560(9-27);9116(2-4);9218(2-3). Lutz,736 (9-26b);820 (9-25). Lyonnet,S., 257, 2163 (9-28).

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di, s_ L.

n.

(I

- 1.

329 Meier,W. [et al.], 199 (13-8a);2299 (9-14); 2807 (1-2); 3188 (2-3). Meijer,W.et al., 69 (2-3);724 (3-2a). Mejia,M. [etal.], 13262,14762,26060(3-2a). Melherson,G.A., 741 (9-27). M6linon,M., 38, 57, 124,630, s.n. (4-1). Mell,C. D., 685 (2-1);s.n. (2-3). Mello,F., 18,40, 1509(7-14);2961 (6-3). MelloFilho,L.Emygdiode [etal.],357(9-26a);1316(9-25); 1488(7-1);2390(9-21);2673(9-26a);2919,3093(9-30a). Mello,J.C. de, s.n. (9-30c). Mello-Silva,R. et al., 80 (7-2);840 (7-3);843 (9-24). MendezG.,A., 8029 (2-1). Mendez,P.et al., 43 (7-4b). Mendez,R. [etal.], 74 (7-9); 116(7-12). MenendezL., F.et al., 150,381 (2-3). Mendez,157(18-19d). Mendoza,P.et al., 97 (5-3). Mendon9a,R., 673 (3-2a);877, 905, 1087(7-1). Meneces,E. [etal.],2 (8-1);344 (5-2);381 (8-1);424, 643 (6-2);656 (6-1);736 (10-2c);793 (5-2);879 (6-2);900 (13-4);902 (13-3);903 (8-1);2004(5-2);2076(10-1b); 2078(10-5). Menezes,M. de, 22, 644 (9-30c). Merano,B. et al., 459 (18-unnamed). Merida,T., 16(8-1). Mexia,Y, 930 (3-2a);1142, 1701(2-3); 1701(2-1);4475 (3-2a);5158 (6-3); 5189 (3-2a);5303 (7-1); 5329 (32a);5379(7-1);6262,6303(2-3);7220,8133(2-4);8260 (7-7); 8303 (9-13); 8844 (2-3).

MeyerDrees,E., 26 (6-2);78 (3-2a). Meyer,G.,9(6-3);87(7-1);102(7-6a);105,112,112,172(63); 178(18-13);221 (6-3);255 (18-1);267 (18-6);326 (10-3);330 (3-2a);339 (15-3);347 (18-13);s.n. (7-6a). Meyer,T., 162,1060,3185,4959(3-2b);5642(3-2a);6739 (7-2); 10778 (9-32); 11856 (3-2a); 12691, 15021, 15946, 18591 (3-2b). Meyer, W. C., 157 (2-3). Michel, R. de et al., 1040, 1042 (7-13a); 2264 (13-4). Miers, J., 1963,2639 (7-3); 2720 (9-26a); 3130 (9-25); 3246

(9-26a);3387 (7-3);3572 (9-25);3684 (7-3);3854 (32a);3910 (7-1);3966, 3983, 3984 (7-3);4571 (9-25); 8087(7-3);s.n. (3-2a,9-6, 9-18, 9-25, 9-26a). Miller,G. S., 1034(9-41); 1180(3-2a);1426, 1550(9-34); 1742(8-1). Miller,J. S. et al., 967 (7-13);1593(7-6b);3235 (9-28). Miller,P.(herb.);s.n. (9-27). Miller,R. et al., 626 (18-10). Milliken,W. [et al.], 57, 95 (6-3); 103, 146 (7-13a);161 (10-Ib);513 (9-32);526 (6-2); 577 (18-9);580 (4-1); 586 (7-13a);637 (10-9). Miranda, F.,5969(2-3);6238 (2-1);6339(6-1);7028(2-2); 7471,7650 (2-1).

Mirili,R., 4 (9-25). Mocino,J. M. et al., s.n. (9-28). Mohr,C., 465 (2-1);s.n. (1-1). Molfino,J. F.,s.n. (9-32). MolinaR., A. [et al.], 238 (2-1); 266 (3-2a);309 (9-28); 909, 1108a, 1300, 1443 (2-1); 2497 (2-3); 2578, 3453 (9-27); 3810 (2-3); 4064, 5317, 5473 (9-28); 5625 (9-

27); 6379, 6529 (2-1);7461 (2-3); 7506 (9-28);8101 (2-3); 8151 (13-5); 8188 (2-1); 10455 (9-10); 10476, 12185, 12267 (2-1); 12469, 12977 (9-27); 13875 (2-1);

14300(9-28);15413,15429(2-3);15765,15803,15837 (9-27); 20713 (2-3); 20847 (9-10); 21613,21756 (2-1); 24596,24623 (2-3); 25656 (9-10); 26246 (9-27); 27525

(9-28);27970(9-10);30769(2-3).

330 M. E., 162(2-3). Mondrag6n, MonsalveB., M.,668 (10-1la); 832,3178 (18-18a). Montaldo,P.,3366,3420 (3-2a);3504 (2-3). Montes,J. E., 190 (9-32);216 (9-31);900 (9-32);931 (931); 1026(9-32);1326,1460, 1654,2163,2558(9-31); 3890(9-32);4024 (9-31). Montes,M.N. et al., 443, 3074 (3-2a). Monteiro,0. P.et al., 941 (13-3). Montgomery, G., 83 (7-13);90 (2-3); 127(2-4); 196(8-1). Moore,H. E. [etal.], 180(9-29);2919(9-33);2930(3-2a); 5017(9-28). Moore,S., 566 (7-1);963 (9-32). MoraV.,C. E., 8 (1-2). Moraes,J. C. de, 692A (7-1). Moraes,M., 655 (7-13b);2237 (7-13a). Morales,J. F.,35, (7-8); 177(1-2). MoralesR., J., 599 (9-27). Morales,R. et al., s.n. (9-10);2174 (13-8a). Morawetz,W.et al., V.2-101085(8-1);X.3-211085(7-1); V.10-151085(6-2);11-31085,13-21888(7-6a);14-23783 (9-32);14-181085(6-2);14-23783(9-32);14-41085(74c); 14-171085(2-4);15-21088(7-6a);18-26985(7-4c); 18-71085(18-13);116-71085(18-6). Morel,I, 193, 524 (9-32); 1291 (7-13b);3746, 3926 (32a);3954 (7-13b);6518,6668 (3-2b). Moreno,P.P.[etal.],60, 1115(2-1);1185(2-3);1243,1389, 1547,1671,1690(3-2a);1714(2-3);1759(3-2a);2868 (2-1); 3162, 5225 (2-3); 7503, 7584, 8162, 8471 (2-1); 8760, 8847 (9-28); 8903, 8948, 3453 (3-2a); 9097, 9146 (2-1); 9370, 9814, 9934, 10008, 10093, 10126 (9-28);

10495(3-2a);10590(2-3);11087(3-2a);11159,11479, 11753,11895(9-28);12035(2-3); 15501(2-1); 15797 (2-3); 16023(2-1); 16601,16628(9-28);16648(3-2a); 16650, 16665a, 16683 (9-28); 16919 (3-2a); 16942, 17095(9-28);17195(2-3);17530(9-27);18067,18126, 18613(9-28);19259,20074,20268,20328 (2-1);20717, 20803,21136 (2-3); 21186 (2-1); 21494,21509,21541, 21548, 21582 (9-28); 21633 (2-3); 21669 (2-1); 21743 (9-27); 21772,21808,21827 (9-28); 22033 (2-3); 22036,

22066(9-28);22995(2-3);23343(10-5);24227,24204 (9-28);24240(2-3);24315,24335,24809(9-28);24840, 24855, 25473 (2-3); 25505 (9-10).

Mori,S. [etal.],241 (9-28); 1965(2-3); 1979,3084,3872, 3887 (7-9); 4042 (2-3); 4137 (15-1b); 4702 (10-1 le); 4946 (18-19d); 5164 (14-4); 6630, 6709 (16-7); 5182 (6-1); 5190 (2-4); 6247 (2-1); 6391 (2-3); 6468 (2-4); 6984, 7033 (7-1 1); 7542 (2-1); 7693 (7-9); 7960 (2-4); 8789 (6-3); 9309 (7-1); 9348 (9-30a); 9587 (7-1); 9631

(3-2a);10146(7-3); 10659(6-3); 11196(3-2a);11344, 11503, 12015a, 12095 (7-3); 13866 (10-5); 13885 (164); 13882 (10-8); 14117, 15380 (6-3); 15546 (14-9a); 15956, 18850 (11-1); 20757 (18-11); 21593 (6-3); 21750 (7-14); 22002 (15-2); 22279 (13-4); 23401 (14-9a); 23490, 24620 (10-1 If). Morillo, G. et al., 2128, 2131 (9-14); 4269 (3-2a). Moritz, J. W. K., 387 (3-2a); 559 (9-27); 1703 (1-2); s.n. (9-14).

Morton,C.V.[etal.],2393 (9-27);2478(2-1);2539(9-27); 2719(8-1);3402,3824(9-35);9029(9-42);9695(2-3). Morton,E. C., 2054, 2055 (1-1). Morton,J.,s.n. (3-2a). Mosen,H., 31 (9-26a);2943 (9-3);3796 (7-3);4391 (7-2). Moses,T.,65 (6-3). Mosquera,E., 10848(14-8a). Mota, R., 137544, s.n. (7-1).

Mowbray,R. N., 702, 703 (2-3, 2-4). Muehlberg,G., 17 (7-4a).

FLORANEOTROPICA Mueller,F.,92 (9-7);584 (1-1);4169 (7-2);s.n. (3-2a,7-2, 9-7,9-27). Mueller,C. H.,2727 (1-1). Murphy,H. [etal.], 351 (10-ib); 352 (7-13a);627 (2-4). Murphy, N. et al., 1283(9-10). Murray, N. J. et al., 1430(9-28). Mutis,J., 588, 589, 1139,2958 (5-1);2958, 2992 (2-3). Nadeaud,J.,s.n.(3-2a,6-3,7-1, 7-3,9-2,9-25,9-26a,9-32). NarvaezM., M. et al., 443 (3-2a);853 (2-3). Nascimento,J. R.M., 1302.3648(6-2). Nascimento,0. C., 481 (7-1). Nash,G.V.,206 (2-3). Navarro,G. et al., 388 (3-2b). NavarrodeAndrade,6519 (7-2). Nee,M. [etal.], 1983(10-5);3443(3-2a);6792,6888(2-4); 6908 (7-9); 6986 (2-4); 7371 (7-9); 7585, 7716 (2-4); 7778 (7-9); 7951 (8-1); 8666 (7-9); 8694 (2-4); 8792 (10-9);8949(2-4);9087(8-1);9241(2-4);9408(9-27); 9410, 10919 (8-1); 16897 (2-3); 17562 (3-2a); 17785 (8-1); 18421 (9-27); 18431 (9-33); 22377 (2-3); 23260 (1-1); 23608 (8-1); 26130, 26299 (1-1); 26681, 26706 (2-1); 27848 (6-1); 27852, 27961 (2-3); 28247 (9-28); 28716,29224 (2-3); 29753,29810,29872,29945,29955 (2-1); 30299 (16-1); 30549 (16-2); 30878 (10-Ilf);

31391,31397(6-3);32474 (8-1);34242(3-2b);34443 (3-2a);34473(4-1);34519(13-3);35148(8-1);35430 (2-4); 35669 (3-2b); 35729,35813 (7-13b); 35850 (10-6); 36040 (2-4); 36063 (9-32); 36848 (8-1); 37036 (3-2b); 37250 (8-1); 37261 (7-5); 37283 (2-4); 37937 (3-2b);

37940, 37941, 37960, 37988, 38026, 38214 (3-2a); 38634 (10-6); 38686 (13-1); 38711 (13-3); 38798 (102c); 38815 (3-2a); 38900, 39087 (2-4); 39158 (8-1); 39206 (2-4); 39262 (8-1); 39208 (2-4); 39537, 39586 (3-2a); 39827 (2-4); 39985 (3-1); 40044 (3-2b); 40091 (3-2a); 40107 (7-1); 40155 (3-1); 40212 (6-2); 40458 (3-2b); 41382 (13-8c); 41815 (5-2); 41872 (7-1); 41935 (7-7); 42145 (3-1); 42457 (7-6a); 43184 (3-1); 43207

(9-30c);44649,45025,46733 (3-1). Neill,D.A. [etal.],402,603 (3-2a);1208(2-3);1589(2-1); 1724 (13-8); 1804,2298 (2-1); 2391 (3-2a); 2794 (9-28); 2855 (3-2a); 3192 (2-3); 3497 (18-18a); 3813 (2-1); 6040 (7-4c); 6059 (7-6a); 6073 (14-id); 6235 (6-1); 6241, 6391 (7-5); 6456 (2-3); 6494 (7-5); 6497 (5-3); 6503 (6-

1);6517(8-1);6613 (14-7);6663(18-20);6829(18-6); 7046(2-3);7162(6-1);7261(3-2a);7295(18-19a);7307 (10-5); 7342, 7343 (6-1); 7381, 7382,7437 (2-4); 7438

(9-28);7442(5-3);7455(16-1);7471(3-2a);7529(6-1); 7759 (2-1); 7843, 7859 (7-5); 7890 (13-4); 7901 (5-3); 7907 (7-5); 8117 (6-1); 8223 (7-4b); 8305 (18-21); 8307 (6-2); 8335 (18-15); 8349 (15-ib); 8357 (14-3); 8359

(10-14);8360 (14-1b);8478 (15-3); 8482 (5-3); 8789 (10-13); 8796 (14-2); 8859 (6-2); 8994 (18-10); 9032 (6-2); 9155 (2-4); 9198 (10-3); 9368 (8-1); 9382 (13-1); 9384 (6-2); 9447 (6-1); 9513 (18-19a); 9614 (16-2). Nelson, B. W. [et al.], 489 (7-5); 719 (6-2); 730 (7-7); 744

(18-7);794 (7-5);794 (7-4b);799 (7-6a);848 (7-5). Nelson,C. et al.,2520(2-1);4504 (2-3);4632 (9-27);5196, 5374 (2-1); 6019 (9-27); 6665, 6752, 7323, 8769 (2-3); 9742, 9768 (3-2a).

Nelson,E. W.,66, 2350, 2437 (2-3). Netto, L., 209 (7-3); 321 (3-2a).

Neuwied,PrinceMaxim.,s.n. (3-1, 9-21, 9-30c). Nevers,G. de [et al.], 3618 (2-4);3948 (10-1le); 4643 (81);4782 (14-le); 5130 (8-1);5265 (14-4);5748 (8-1); 5772 (7-11);6104 (7-9); 6516 (18-19d);7358 (7-4b); 7455, 7527 (7-1 1); 7533 (10-9); 7621 (9-27).

LISTOFEXSICCATAE Nevling,L. et al., 2563, 2584 (9-27);3893 (2-1). Newman,R., 90 (3-2a). Niederlein,G.,287, 1717,2206,2214,22146(7-2). Nishimura, A., 8 (6-2); 17 (7-1);50 (18-10). Norris,D. H. et al., 13675,13676,1399(9-28). Novaes,C., 23849(3-2a). Novaes,J. da Silva,s.n. (9-1). P. [et al.], 5543 (9-13);6279 (14-3);6349 (3-2a); Nun'iez, 7156 (1-2); 10148 (5-2); 10193 (6-2); 10278 (3-2a); 10305(6-1); 10472(7-7); 10528(8-1); 10630,10714, 10789, 10850, 10889, 10894, 10977, 11040, 11043, 11055, 11136(7-7); 11146(7-5); 11219(7-7); 11252 (13-3);11438(13-4);11806,11860,12164(7-7);12165 (14-5.note);12226(7-7);12279,12677(7-1);12949(913);13048,13941(10-13);14128(2-4); 14227(13-9). OcampoS., R.A., 1647, 1876(7-4a);s.n. (9-10). Occhioni,P.[etal.],242 (9-2);243(9-26a);610(9-25);1016 (9-18);1108(9-19). Oellgaard,B. et al., 34866,35152,35179(2-4). Oersted,A. S., 5855 (9-28); 14293(6-1); 14345(8-1);s.n. (2-1,2-3, 6-1). Oldeman,R. A. A., 342 (6-3);B.628, B.642 (4-1);T.668, 1132,2048,2056(6-3);B.2427(18-11);2676,B.3064, B.3076,B.3173,B.4177,4283(6-3). F.H.F.et al., 127 (7-6a);1137(14-8a);1402 Oldenburger, (7-6a). Oliveira,A. A. et al., 305 (18-19c). Oliveira,J.E. de, 310 (6-3);753 (18-4);949 (6-3);1018(41);1072,1120(6-3);1147(7-1);1179,2422,2681(6-3); 2857, 2919 (7-1);2980 (7-4c);4506 (11-1). Oliveira,P.I., 668 (7-2). Oliveira,R., 17 (13-1);31 (7-6a). Oliveira,R. F.de, 237 (9-3);264 (9-30a);265 (9-22);453, 541 (9-25);3811 (13-1). Oliver,R. L. [et al.], 677, 916 (9-28);2312 (9-27);2362, 2364(2-4). Onishi,E. [et al.], 17, 239 (9-30c);499 (7-1);821 (9-30c); 1278(7-1). Opler,P.A., 283 (9-28);839 (3-2a);925 (7-4a);1657(2-5). J. R., 1033, 1056(2-4). Oppenheimer, Orcutt,C. R., 2415 (9-34);4189 (9-28);4232 (3-2a). O'Reilly,et al., 46 (9-30d). Orjuela,P., 102(2-3). OrozcoC., J. M.,406 (2-3); 1679(6-1). s.n. (9-32). ORSTOM, OrtegaU.,A., 30 (5-3);43 (13-3);50 (14-8b);86 (5-3);228 (2-3); 131(10-1If). Ortega,R. et al., 232 (2-1). OrtegaO., R.V.,823, 1455(9-27). Ortiz,B. et al., 89 (2-1). Ortiz,F., 1407,1969(9-27). Ortiz,L., 167(7-7). Ortiz,215 (10-5). Osario,H.,s.n. (9-32). Osten,C.,2892,3006,3086,3555,5145,5270(9-32);8158, 9146,s.n. (7-2). Otto,E., 1109(9-32). PabonE., M., 1022(7-6a). Pabst,G.,4304, 5394(9-26a);5395,7379 (9-19);8812(930c);55477,55478(9-26a). Painter,L., 62 (18-4). Palacios,M.A. et al., 3409 (9-30c). Palacios,P.A. et al., 1838 (7-4b); 1949 (6-2); 2171 (1819c);2172 (14-3). Palacios,W. [et al.], 261 (7-5);262 (7-6a);514 (5-3);607

331 (14-3);928(5-3);949(2-4);1031(6-1);1055(2-3);1084 (7-5);1131(13-9);1244(10-9);1291(6-2);1301(5-3); 1309(10-5); 1425(6-1); 1441(2-4); 1453(6-2); 1598 (15-3);1613(7-5);1663(7-5);1683(5-3);1685(7-4b); 1746 (10-1If); 1748 (5-1); 1762 (14-8b);1798 (5-1); 1837(5-3);1838(74b); 1851(18-21);1900(7-5);1982 (6-2); 1995(18-7);2004 (14-4);2012 (9-12);2014 (74c);2035(16-3);2037,2051 (7-5);2102 (14-8b);2104 (7-5);2110 (5-3);2117,2139 (7-5);2140 (7-4c);2230, 2260(7-5);2285(14-3);2405(16-5);2406(13-4);2594 (8-1);2793(6-1);3065(10-9);3201(16-1);3227,3265 (5-3);3296(3-2a);3352(7-4b);3355(13-4);3366(6-2); 3367(10-3);3375(14-Id);3383(15-1b);3388(6-2);3398 (10-1if);3453(15-3);3484(14-ic);3568(13-8c);4026 (8-1);4249(7-5);4322(13-1);4467(6-1);4503(16-1); 4505(5-3);4522(14-Ic);4679(17-2);4681(7-4b);4691 (10-unnamed); 4695(5-1);4712(1-2);4735(13-1);4780 (7-5);4782(7-4b);4947(7-5);4951(7-6b);5092(16-7); 5101 (7-5);5116 (7-6a);5322 (1-2);5344 (16-7);6441 (13-9);5983,6126,6228 (1-2);6229 (6-1);6480 (5-3); 6562(6-1);6754 (16-1);7433 (14-4);7566 (5-2);7699 (2-3);7715 (5-2);7720 (2-3);7783 (14-1b);7920 (101lf); 8303(14-la.note);8570(13-4);8705(6-2);8803 (13-3);9427 (13-1);9570 (6-1);9674, 9691 (18-18b); 10040(7-8);10101(7-4b);10261(14-la);10285(9-12); 10427 (13-4); 10791 (18-15); 10821 (14-9b); 11122 (6-2);11145(18-19d);11336(6-2);11356(7-12);12044 (7-6a);12411(6-1). Palmer,E., 29,39 (9-28);89 (3-2a);140(9-28);142(3-2a); 143, 149, 158 (1-1); 337, 454, 543 (2-3); 552 (3-2a); 563 (1-1);1425(3-2a);1443(9-28);s.n. (3-2a). Paredes,R., 718 (6-2). Pardo,M. de et al., 78 (18-12). Pariona,W.et al., 869 (10-3);1021(5-3). Parlatore,s.n. (7-3). Parodi,D., 3267(3-2a). Pastore,U. et al., 157(7-2). 9157(3-2b). Pastrana, V.M., 83 (2-3). Patinlo, Patmore,P.N. et al., 62 (13-3). Pavetti,C.V.,267, 524, 711 (3-2a);789 (7-13b). Pavon,J.A., s.n. (2-5, 9-28). Pearce,R., 2804(9-32). Pearsall,D. M., 1071(13-8c). Peck,M. E., 675 (2-3). Peckolt,G., 17(7-3);s.n. (6-3). Peckolt,T.,s.n. (9-2). Pedersen,T. M., 443 (7-13b);472 (7-2); 847, 856 (3-2b); 1208,1208a(7-13b); 1778,2780,3978,4353(7-2);5132 (9-32);5142(3-2a);5224(9-32);5890(3-2a);6646(932);8420(7-2);9839 (7-13b). Pedrosa,A. et al., 611 (9-25); 612 (9-18); 672 (9-25); 1020 (9-18).

Peixoto,A. Luna[etal.],606 (9-25);838(9-22);3128(6-2); 3200 (6-3);3210 (16-1);3436 (7-3);4012 (7-1);4051 (6-2). Penna,T.M. P.A. et al., s.n. (9-26a). Pennell, F. W., et al., 202 (3-2a); 5778 (7-9b); 19519 (9-28). Pennington, R. T. [et al.], 62 (10-2); 82 (17-2); 109 (104); 117 (15-3); 144 (10-4). Pennington, T. D. [et al.], 158 (6-1); 6350 (8-1); 9080 (135); 9114,9148 (8-1); 9200,9292 (2-3); 9329 (8-1); 9500 (2-3); 9531 (2-1); 9564 (13-8); 9640,9641 (3-2a); 10526 (2-4); 10589, 10643 (6-2); 10679 (7-10); 10689 (3-2a); 10720 (7-13c); 10786 (3-2a); 12224 (14-3); 12258 (62); 13694 (13-8c); 13727 (6-2); 13933, 13939, 13997, 14107, 14147, 14205, 14499 (7-10); 14925 (6-1).

332

FLORANEOTROPICA

(6-1); 12101(2-4); 12411(9-10); 12453(3-2a);13453 Peredo,I., s.n. (3-2b). (7-4a);15407(3-2a);15477(7-4b);15665(2-3);16894 Pereira,B. A. S., 1184(7-1). (7-6b);s.n. (2-3, 7-4a). Pereira,E. [et al.], 245 (9-32); 261 (7-13b);436 (9-30c); 440 (9-30c, 9-32); 573 (9-26b);574 (9-26a);1824(7- Pizzioli,W.,332 (7-1). 1); 2117 (9-2); 2285 (9-18);4176, 6935 (9-25);6995 Pivetta,811 (7-2);1212(9-32); 1213(9-31). (9-26b);7191 (7-1);7192 (7-2);7195 (9-25);7195 (9- Plee,A., 27 (3-2a);112,782 (9-27);825, 848,s.n. (3-2a). Plowman,T. [etal.], 2780 (9-26a);2831 (7-3);4174 (2-5); 26a);7457 (9-32). 5131(7-3);5471(7-13c);5904(9-13);6666,6669,6670, Pereira,N., s.n. (3-2a). 6671 (14-ic); 8279 (9-30d);8389, 8481 (9-32);8542 Perez,L., 433 (7-2). (9-30d);10085(9-1); 11270(5-2); 11430(2-4); 11674 Perez,R., 719 (1-2). (14-5);12613(7-6a);12856(9-25); 13921(7-1). E. et al., 6512 (3-2a);6740 (7-6a). Perez-Arbelaez, Poeppig,E., 1267(2-5);2325(7-6a);2384(3-2a);2563(7PerezJimenez,L.A., 226 (2-3). 5); 2581, 2581B(7-6a);2582 (6-3);2582 (7-6a);2683 Perkins,J. R., 1196(3-2a). (7-1);s.n. (3-2a,6-3, 7-6a). Peron,M., 351 (7-2). Pohl,J. B. E., 740 (3-2a);s.n. (3-2a, 6-3, 9-2, 9-19, 9-20, G. S., s.n. (6-3). Perrottet, 9-3Id). Perry,A. et al., 289 (17-2);737, 760 (7-1). Poiteau,P.A., 6 (6-3);s.n. (2-3, 6-3). PeshaB., V.,71 (5-1). PonceC., F,77, 123,215 (2-1);232 (2-3);237 (8-1). Pessoalde HortoFlorestal,137547(7-3). PoncedeArrudaJunior,G., 2 (9-3). Peters,C., 141(14-5);158(2-3). Poole,J. M., 1939(7-6b). Petrov,I., 78, 89, 100(4-1). Popenoe,W.,845 (8-1). Petterson,P.M. et al., 6666, 6847 (2-3). Porter,D. M. et al., 4790 (14-4). Pfeifer,H. W.et al., s.n. (2-1). Philcox,D. et al., 3059(7-1);3202,3427 (9-30d);3554 (7- Porto,P.de Campos[etal.],247 (9-30a);629 (9-2);631 (926a);791 (7-2); 783 (9-26b); 1836,2100 (7-2); 2204 1);4272 (9-30c). (9-32);13047(7-2);s.n. (9-30c). Philipson,W.R. et al., 2, 1859(9-27);2097 (2-3). Phillips,0. et al., 69 (6-1); 103, 138,351, 416, 420, 450, Potter,D., 5177 (3-2a). Poveda,L.J.,29 (8-1);37 (7-4a);57 (6-2);127(2-4);203(3451, 577 (7-7). 2a);319(2-3);856,987(2-1);1092,1713(5-1);s.n. (7-8). Pickel,B. J., 142(3-2a);510 (9-21); 1336(9-32);1413(930d);3580(7-1);3687(9-30d);4183(9-32);5357,5518 Powell,D., 136(9-34). Powell,H., 87 (9-27). (7-2);5582 (9-32);s.n. (3-2a). R.S., 1006(9-21);1219,1497(7-1);1534(6-3);1803 Pozetta,G. L., s.n. (9-30c). Pinheiro, (3-2a);1812(9-8);1825(9-18);2031(7-3);2071(3-2a). Prance,G. T. [et al.], 1780(6-3); 1950(16-4);2317 (7-5); 2344(7-1);2426(18-7);2500(16-3);2733(7-4b);2733 Pinkley,H.,237 (8-1). (7-1);2769, 2775 (7-5);2884 (6-2);2901 (14-3);5720 Pinto,P.et al., 107(8-1). (18-19a);6219 (13-1);6572 (7-4b);7180 (6-3); 7270 Pipoly,J.etal., 4934 (13-5);5089,5215,6031,6051,6126, (6-1);7285 (18-8);7430 (7-5);7489 (7-1);7531 (7-5); 6181(2-1);6718,6873(6-3);12607(14-6);12464(74b); 7616(7-6a);7619(7-5);7989(7-1);8984(9-30d);8421 13018(7-4c);13271(16-5);13336(6-2);13337(14-la); (18-13);8676(14-8a);8704(18-1);8955(18-12);9954 13372(14-9b);13552(6-1, 7-4c); 13626(5-3); 13738 (15-1b);10085(11-1);10488(6-2);10535(6-3);10578 (7-4c);13979(14-8b);14223(5-3);14305(7-6a);14329 (15-la);10602(16-1);10622,11115,11116(6-3);11122 (14-9b);14359(18-15);14422(5-3);14428(7-5);14520 (134); 11240(16-1);12417(7-5);13359(13-8c);13386 (14-6);14609(10-2c);14614(6-1);14625(6-2);14681 (3-2a); 13725 (11-1); 15563 (18-10); 16458 (14-6); (5-3); 14947(14-5);15165(18-10);15590(18-19a). 18248(7-1); 18349(7-6a);18375(10-5);19011(7-1); Pirajada Silva,35 (9-21). 19219(9-30c);19247(7-1);20585 (11-1);22713 (18Pirani,J.R.[etal.],2434(7-3);2533(6-3);2808,2885,2913 21);22716(15-2);22824(16-3);22829(10-2c);22836 (7-3); 2952 (9-1); 3068 (9-8); 3378 (7-3). (18-7);22863(12-1);22889(6-2);22944(7-6a);23002 Pires,J.Mur9a[etal.],71 (7-6a);118, 170(6-3);224 (7-1); (18-9); 23430 (7-14); 23841 (18-19c);23871 (7-6a); 248 (7-4b);254 (7-1);259, 309 (7-4b);311 (7-1);338, 23993(14-6);24016(74a); 24017(7-6a);24285(7-1); 343 (7-4b);346 (7-6a);353 (10-10);518(7-4b);541(724319(14-1a);25386(18-4);25401(7-6a);25430(186a);555 (7-6b);556 (7-4b);1329,1334(7-6a);1505(39); 25508 (7-6a);25509 (12-1);25723 (16-4);25884, 2a); 1567 (7-1); 1574 (16-4); 1730, 1841 (6-3); 1997 25888,26381(6-3);26398(18-9);26438(13-1);26567 (14-9a); 2026 (16-4); 2062, 3154 (6-3); 3539 (7-6a); (7-1);29467,58678 (6-3);59186, 59293 (7-1);59316 4559 (6-3); 4796 (9-30d); 6150 (7-1); 6873 (4-1); 9954, (3-2a);59361(9-30c);58940(10-2b);59532(7-1). 9955 (18-2 1); 12041, 12101 (2-5); 12427 (10-2b); 12747 (6-3); 12813 (6-2); 12849 (10-2b); 12879 (6-3); 12965 Prevost,M. F., 435 (6-3); 2130 (9-32);2221, 2510 (4-1); 2784(10-3). (6-2); 13049 (6-3); 13111 (7-1); 16239 (10-3); 17254 (7-13a); 17337 (3-2b); 17879 (6-3); 50461 (18-19c); Pride,A., s.n. (9-32). Pringle,C.G., 19,707(1-1);2498(9-28);6791(1-1);7562, 51938(4-1);52339(9-30d). 7566(9-28);7750(1-1);11172(9-28);13197,s.n. (1-1). Pittier,E., 74 (3-2a). Pittier,H. [etal.l, 74 (3-2a);87 (9-27);325, 326(9-10);803 Prior,A., 303,s.n. (2-3). Proctor,G. R., 4140, 4503, 4642, 7167, 7294, 7995, 8590 (24); 1044 (3-2a); 1591, 1837 (9-27); 1918 (3-2a); 2255 (9-34);8678,10216(3-2a);11592(9-36);18177(3-2a); (2-3); 2612,2619 (3-2a); 2748 (9-27); 3045 (7-8); 3627 20588(2-3);22161,22233 (9-34);22304 (2-3);22457 (2-3); 3731 (8-1); 3856, 4034, 4113 (7-9); 4202 (8-1); (9-27); 22700, 22779, 23018 (9-34); 24221 (3-2a); 4707 (2-4);4710 (3-2a);5291,5425(2-4);5695(3-2a); 26422, 27699 (9-34);27982 (3-2a);29409, 30205 (95817(9-27);6359(3-2a);6578(2-4);6636,6637,6652 27); 30269, 31106 (2-3); 31729 (13-8);32502 (3-2a); (2-3);6747(2-4);6908(9-27);7870(7-13a);8091,8531 32665,32752(9-34);35958(8-1);35982(9-38);36165 (9-27);8659(2-3);8855,8856,9074,9388,9389,9524 (2-1);37071(9-34);38353(2-3). (9-27);9898 (9-10);9898 (9-27); 11266(7-4a);12082

LISTOFEXSICCATAE Pruski,J. et al., 3462 (7-2);3476 (18-2). Pujupet,J. O., RBAE.1051(3-2a). Purdie,W.,93 (9-32);s.n. (9-27,9-34, 3-2a). Purpus,C. A., 1919,2316 (2-3);2806 (9-27);4386, 5049 (2-3); 5311 (9-33);5501 (1-1); 5725 (9-27);5779 (933); 5951 (9-27);5955, 5988 (2-3); 5989 (6-1);6110 (9-27);6492(6-1);6900,6960(9-27);7091(2-1);7137 (9-28); 8200 (2-3); 9030 (3-2a); 10837, 11044(2-3); 11070(2-1);12072,14273,16318,s.n. (2-3). Purseglove,J.W.,P.6375(3-2a). Quelal,C. et al., 201 (14-3); 419 (7-5); 521 (7-12); 576 (18- 17). Quevedo,R., 248, 361 (5-2);503 (3-2a). M.et al., 36 (8-1). Quintanilla, Quintas,A. Taveres,1021(9-32). QuinteroD.,A., 185(6-1). QuirosC., M., 795, 23248 (3-2a);23249, 23250 (9-27); 27668(7-4a);s.n. (3-2a). Rabelo,B. V.et al., 2351 (4-1);2971 (11-2);2558 (9-32); 3182(11-2). RaimundoS. P.,Fr.,1093(9-9). Rambo,B., 1308(3-2a);2246(9-31);24053(9-32);27131, 28982(7-2);30084(9-32);32844(3-2a);37737,38280 (7-2);38432(9-32);38586,38604(3-2a);42117,42432, 42980(9-31);43725,43746(7-2);43791(9-31);43820, 44225,48800,48977(7-2);49142,49143(3-2a);57281 (7-2);61222(9-31). Ramia,M. [etal.], 1921,WT.3534(3-2a). RamirezC., W., 1 (2-3). Ramirez,I. et al., 249 (7-6a). RamirezR., M., 27 (2-4). RamirezS., 105(2-1). Ramos,G. H.,s.n. (9-32). Ramos,J. et al., 53 (7-1). Ramos,J. E. et al., 850 (7-13a). Ratter,J.A. et al.,2504(9-30c);2662(9-30d);4501(10-9); 5408,5649,5675(6-2). Rau,W.,s.n. (7-2). Raunkiaer, C., 2525,s.n. (3-2a). Raven,P.H.,21558(7-4a);21699(9-10);21775(7-8);21897 (18-17). Raynal,A., 15669(9-27). Reark,J. B., 248, 276 (1-2);635, 653 (7-8);694 (7-4a). Record,S. J. et al., H.40(2-1). Regnell,A. F.,190,191(9-26a);192,193(9-25);411(9-32); s.n. (9-25).

333 Reveal,J. L. et al., 4176 (9-28);4898 (2-4). Revilla,J.,596 (7-14);685, 790 (3-2a);845 (7-5);939 (146); 1185(18-15);1663(7-4c);2304 (18-10);2371 (145); 2544(15-lb);4270 (7-6a). Reyna,M.L.[etal.], 159, 1420(2-3). ReynelR., C., 6 (18-19b);25 (7-1);70 (19-19b);135(6-2); 141(18-19b);251 (7-7);292 (7-1);318 (5-3);360 (5-1); 423 (10-3);478 (7-1);488 (2-4);589 (14-4);648 (1819b);782 (5-2);932 (7-7);971 (7-5);1013(13-3);1047 (6-2);1048(7-1);1101(14-8a);5216(7-1);5217(7-4c). Ribeiro,B. G. S. et al., 1294,1296(7-1). Ribeiro,J., 21, 26, 44 (6-2); 109(29-25);131(9-30a). Ribeiro,J. E. L. S et al., 1800(18-14);1849(16-5). Riberao,R., 385 (9-18). Richard,A., s.n. (7-3). Richard,L. C. (herb.);s.n. (3-2a,6-3). Richards,R. et al., 598 (13-4). Ricksecker, A. E.,482 (3-2a). Ridley,H.N., s.n. (9-25). Riedel,L. [etal.], 14(9-21);22,71 (9-25);104(9-26a);159 (7-1);303 (7-3);308 (9-18);309 (9-25);359(9-18);506 (9-25);507 (9-19);508 (9-30a);509 (9-3);527 (9-32); 539 (3-2a);597 (7-1);691 (9-30c);759 (9-9); 1011(92); 1010(9-3); 1013(6-3); 1184(7-13b);1191(9-32); 1275(9-19);1862(3-2a);1931(9-32);2062 (3-2a);s.n. (6-3,7-2, 7-3,7-13b,9-3, 9-18,9-19,9-21,9-25,9-26a, 9-30a,9-32). Riise,A. H.,s.n. (3-2a). Rimachi Y, M., 195(3-2a);607(2-3);2027(6-3);2512(7-5); 2879(15-3);2921(18-6);3959(7-5);4229(7-6a);4334 (3-2a);4436(14-6);7095(7-6a);7787(3-2a);8573(7-5). Rimbach,A., 187,505 (2-3). Rios,D. E., 211 (2-3). RiosT.,J., 1 (10-3);2 (7-13a);270 (3-2a);910 (10-1If). Rios,P.,353 (18-17). Rivera,G., 682 (2-3). Rizzini,C. M., 152(9-25);10063,86692(7-2). RoaT.,A, 200 (13-3);635 (14-8a). Robert,A., 784 (3-2a). Robles,R. [et al.], 1239, 1262, 1288, 1640, 2620 (7-4a); 2654(2-3);2693 (2-5);2767 (2-1);2773 (2-5). Robleto,W.,23 (3-2a);206 (2-3);784, 1209(3-2a);1355 (2-3); 1458(9-28);1657(2-3). Rocha,127, 171,298,428 (9-26a). W.[etal.],61,62 (9-25);66, 161(9-26a);225(7Rodrigues, 6a);569 (7-6a);675 (9-32);963 (7-6a);963,987 (7-4b); 2695,2723(7-14);2793(3-2a);3855,3962(7-6a);4234 (6-3);4716 (7-6a);4743,4744 (7-6b);5519(6-2);5594 (7-1);5892(7-14);5893(3-2a);7274, 7348(7-4c). RodriguezC.,A., 149(9-28). RodriguezF.,C., 24 (1-1). Rodriguez,I.A., 14,117 (3-2b);264 (9-32);366 (7-2);460 (3-2a);537 (7-2); 1002, 1031(3-2b). Rodriguez,J.V.,214 (3-2a);632 (9-28);2122,3380(3-2a); 3411(9-28);3478(3-2a). Rohweder, O., 3613 (3-2a). Roig,J. et al., 2547 (9-39);6580 (9-35). Rojas,(=Tonduz& Rojas);151, 152,571 (9-27). Rojas,T. (partlyin Hassler,E.);480 (7-13b);1142(9-32); 5028 (3-2a);7989 (9-31); 10509,10542(9-32);11649

Reed,A., 50 (8-1). Rego,1.F.,et al., 739 (6-2). Rehder,A., 272,496 (1-1). Reiche,K.,294 (9-28);s.n. (1-1). Reitz,R.[etal.],43 (3-2a);139(9-7);524(7-2);1152(9-32); 1720(9-7); 2086 (7-2); 3164 (9-7);3733, 3798 (7-2); 4071 (3-2a);5036(7-2);5759,5922(9-7);6088,6778, 6822, 7012, 7068 (7-2);7327 (9-7);7567 (3-2a);8977 (7-2); 17518(9-32);s.n. (3-2a). Reko,B. P., 3519 (2-1); 3519 (2-3);3590 (8-1);4907 (933);4998 (2-3);6291 (6-2). Renjel,R., 18(3-2a). Renson,C., 112(2-3);203 (9-28). Renteria,E. [et al.], 460, 1717(18-6); 1764(3-2a);1920, (7-13b); 12127, 12283 (3-2b); 12632, 12825, 12904 (92037 (2-3);2686, 2724, 2823 (74b); 2829 (2-3);3633 32); 12945, 12945a (3-2b); 13060 (9-32); 13331 (9-31); (6-1). 13652a(3-2b);13954(3-2a); s.n. (9-32). Renvoize,S. A. [etal.], 3018, 3077 (9-32);3184,3241 (7- RoldAn,F.J. et al., 1189(9-27). 2); 3252(3-2a). Romero-Castanieda, R., 565, 646 (7-13a);650 (10-lb);722 Restrepo,C. et al., 409 (18-18b). (8-1); 1063, 1090 (3-2a); 1107 (7-13a); 1664 (12-1); Restrepo,D. et al., 647 (10-3). 2656 (8-1); 4117 (12-1); 4151,4152 (18-20); 4967 (10-

334 5); 5194 (18-19d); 5483 (16-1); 5490 (10-1 lb); 5586, 6176 (8-1); 6427 (7-9); 7030 (6-1); 7532, 7575, 7801 (13-8a); 8389 (12-1); 8395 (10-1 la); 8402 (13-4); 9037, 9806, 11147 (3-2a); 11165 (2-3); 11356, 11359 (12-1). Rondon, 4481,4504 (7-13b). Rosa M., 570, (9-32). Rosa, N. A. [et al.], 1024, 1514 (15-4); 1684 (7-1); 2055 (10-llf); 2751 (16-1); 3035 (6-3). Rosales, J. et al., 624 (7-13a). Rosas, A. et al., 354 (7-4b). Rosas R., M., 269 (1-1); 1016, 1065, 1213 (2-1). Rose, J. N. [et al.], 3322 (9-28); 3719 (2-3); 4672 (3-2a); 5289 (1-1); 7399, 9370 (9-28); 14428 (2-3). Rose Innes, R., 305 (3-2a). Rosengurtt,B., B-2129 (9-32). Rossbach, G. B., 3720 (7-4a). Rotta, C. La et al., 680 (7-1 1). Rovirosa,J. N., 339 (2-1); 478 (9-27); 569 (3-2a); 881 (9-27). Rowel,'C. M., 2063, 2952 (9-28). Ruano, M., 545 (9-27). Rubio, D. et al., 57 (14-8b); 131 (10-13); 298 (15-3); 787 (6- 1); 1190 (18-18a); 1268 (18-17); 1352 (7-4a); 1399 (8-1); 1409 (18-16); 1723, 1746 (18-17); 1926 (10-Ib). Rudas, A. et al., 1378 (14-4); 1687 (7-6a); 1755 (18-21); 1782 (7-4b); 1785 (5-3); 1801 (74b); 2146 (134); 2943 (18-19a). Rudio, s.n. (9-25). Rueda, R. et al., 416 (7-13a). Rugel, F., 3176 (2-3). Ruiz, H. & J. Pavon, s.n. (1-2, 2-5, 3-2a, 6-1, 6-2, 7-4a, 927, 9-32) Ruiz C., J. [et al.], 1 (7-7); 12 (6-1); 23 (14-2); 1090 (7-6a); 1519 (7-16). L. [et al.], 762,763 (7-3); 763 (2-3); 1242 (6-1); Ruiz-Tera6n, 1247 (2-3); 1247 (7-3). RuizZ., T.et al., 1147(2-3); 1774, 1874, 2705,2968,3089, 3249, 3257, 75-0033, 75-0048, 75-0051 (7-13a). Rusby, H. H., 564 (2-4); 751 (5-2); 817 (9-32); 1011 (2-4); 1284 (3-2a); 2217, 2578 (7-6a). Rutt, et al., RB.67 (7-4b). Ruokulainen, K. et al., 8456 (6-3); 9467 (8-1). Rzedowski, J., 4778, 6894 (2-3); 10398 (9-33); 10997 (927); 12375, 19662 (1-1); 21160 (9-27); 21160 (9-28); 22144 (1-1); 22773 (9-28); 24428 (9-33); 27080,27081 (2-3); 33938 (2-1); 34959 (3-2a). Saa, J., 405 (7-13a). Sabatier,D. [et al.], 28 (5-2); 1245 (10-1 If); 1549 (15-3); 1653 (4-1); 1747 (15-3); 3315 (10-llf); 3900 (5-2); 4074 (18-11). Sacco, J., 2480 (9-25). Sacramento,s.n. (9-3, 9-30a). Saer, J., 142 (3-2a). Sagot, P., 522, 522bis, 922bis (4-1); 530 (6-3); s.n. (4-1). Sagra, R. de la, 287 (2-3); 478 (3-2a); 608 (2-3); s.n. (2-3, 3-2a). Saint-Hilaire,A. F. C. P. de, 54, 55, 56, 57 (3-2a); 58, 59, 60, 61 (9-32); 62 (9-3); 63, 64, 65, 66 (9-25); 67 (9-32); 68 (9-25); 69,70 (9-30c); 70 (9-30d); 71 (9-30a); 72 (932); 72 (9-30c); 73 (9-3); 74 (9-26b); 75, 75bis (9-26a); 76 (9-25); 77, 78bis (9-26a); 79 (9-25); 79 (9-26a); 92 (7-3); 93 (7-2); 94, 95 (7-3); 110 (9-25); 131 (7-1); 278 (7-3); 2150 (9-32); s.n. (9-3, 9-26b). Salas, J. B., s.n. (9-28). Saldanha,J. de (et al.); 1933 (3-2a); 4958 (9-26a). Saldias P., M., 131, 467 (7-5); 1233 (5-2). Saleh, J. E., s.n. (9-25).

FLORANEOTROPICA Salick,J.,7281(13-1);7297(13-3);7331(5-3);7366(134). Salvoza,F.M., 966 (7-9). Salzmann,P., 1838,s.n. (3-2a). A. et al., 39 (3-2a);50 (7-13c). Samaniego, Sampaio,A. J. de, 843 (3-2a). SanchezS., D. [et al.], 60, 166 (18-17);840 (2-4);928 (78); 941 (7-4b);1479(18-18b);1607(7-4b). SanchezV.,I. et al., 8332 (7-5);9456 (16-7.note). SanchezS., M. et al., 198 (18-6);297 (154); 475, 663 (74b);665(10-15b);903(14-8b);1114(18-19c);1115(103); 1252 (18-7); 1440 (10-1b); 1657 (16-2); 1711 (15lb); 1792 (16-6). Sanchez, P. et al., 513 (2-1); 563 (2-5). Sandeman,C., 4279 (1-1); 4962 (2-4).

Sanders,A. C. et al., 9982 (2-3). Sandino,J.C. [etal.],725 (3-2a);1055(2-1);2934A(2-3); 3171 (3-2a);3319 (9-10);3673 (3-2a);4093, 4223 (23); 4394 (9-28);4451 (3-2a);4491 (2-3);4925 (13-8); 4606,4688,4727,4810,4860,5053

(2-3).

Sandoval,E. et al., 956 (2-3). Sandwith,N. Y, 981 (9-27).

E. et al., 3305 (7-6b);3306 (15-4). Sanoja-Angel, Santos,J. P.et al., 48 (6-2). Santos, J. V., 2831, 2916 (2-1).

Santos,M. R., 98 (18-19c);131(7-6a);152(10-3). Santos,T. S. dos [et al.], 223 (6-3);386 (7-3);467 (10-9); 542 (13-3);543 (9-6); 760 (9-5); 829 (9-30a);892 (98);966 (9-18);1086(6-3);1166(7-1);1205(9-3);1226 (18-10);1255(9-8); 1265(9-5); 1308(3-2a);1403(164); 1437 (9-3); 1439 (9-9); 1503 (7-1); 1539 (9-18); 1633

(9-30a);1636(9-18);1638(9-2);1743(9-5);1963,2039 (6-3); 2070 (9-18); 2154 (9-9); 2384 (9-25); 2768 (9-8);

3008(9-30a);3183 (9-1);3326 (7-3);3558 (9-1). Santos,E. B. dos et al., 4119 (6-3). Saravia,C. et al., 867 (3-2a). Saravia,E. et al., 1200(9-32). Sargent,F.H.,A-77 (9-27). Sastre,C. et al., 512 (7-14). Saunders,J.,797 (9-27); 1100(13-5). Scarff,J. G., 96 (2-3). Schaffner, W.,89 (1-1). Schatz,G. E. et al., 230 (2-1); 889a (9-30d); 1036 (2-3); 1055, 1056 (74a); 1100 (2-1); 1119 (9-27). Schenck, H., 1425 (7-2); 1661, 3035 (7-3). Schiede, C. J. W. [et al.], 31 (2-3); 33 (3-2a); 1118 (2-3); s.n. (2-3, 3-2a). Schinini, A. [et al.], 5545, 6688, 7177, 7490, 8330, 8711, 8800, 8848, 8874, 8882, 8895, 9014 (9-32); 9091, 9093 (9-30b); 9173,9355,9420, 11517 (9-32); 12320 (3-2b); 13639 (7-13b); 14584, 14917, 14918, 14946, 15202 (932); 15358(7-13b); 16043 (3-2b); 16204(7-13b); 18030, 18085 (3-2b); 18293 (9-32); 20160 (9-31); 20270,20369 (9-32); 20546,21337 (9-30b); 25609 (9-32).

Schipp,W.A., 1010,1074(2-3);1154,1155(8-1); 1188(927); 1261, 1263 (2-1); 1276 (9-17); 1332 (9-27); 1334 (2-1).

Schlim,L., 916 (9-27). Schmalzel,R. J., 675, 767 (7-9); 1160 (2-4); 1279 (7-9); 1319 (2-4).

Schmidt,E., 2800,4890,4965 (9-31). Schnee,L., 1151(3-2a). Schneider, M.,213 (3-2a). Schott,A., 607, 847 (3-2a). Schott,H.W.,46 (9-27);517 (3-2a);s.n. (9-19). Schreiter, R.,26, 3412,5026, 11351,11443,11444(3-2b). Schubert,B. G. [et al.], 866 (9-27); 891 (7-4A); 1151,1210 (9-10); 1310 (2-5).

LIST OF EXSICCATAE

Schultes,R. E. [et al.], 46 (7-6a);101 (9-28); 139 (7-6a); 3617(8-1);5189(6-1);5417(6-3);5798(14-8b);5858j (7-14);6028 (3-2a);6171 (74b); 6713(18-19a);7169, 7171 (3-2a);8150 (7-1); 8273 (7-14);8325, 8412 (74b);8547, 10128(7-6a);11159(16-1);14908(10-13); 15466(16-2);15639(16-1);16481(10-12);16735(101If); 16825(10-12);18664(9-27);s.n. (15-3). Schulz,A. G.,874 (9-32);1834(3-2b);8861(9-32);16475 (3-2b);17477(9-31);18318,18340,18390,18561(9-32). SchunkeV.,J.,871, 884 (7-7);950 (18-6);953, 1012(2-3); 1067(18-19a);1096(2-3);1548(18-6);1653,1879(144); 2147 (3-2a);2162 (10-5);2178(10-2c);2200(2-3); 2311(3-2a);2573(14-8b);2692(14-7);3330(2-3);3339 (2-4);3365(7-7);3608(18-21);3816,3826(5-2);3895 (7-4c);3999 (2-4);4100, 4425 (3-2a);4430, 4494 (23); 4573 (7-5);4593 (14-6);4632, 4827 (7-4c);5150, 5604 (2-4); 5660 (2-3 or 7-5); 5660 (7-6a);5830 (1813);5907(2-3);6191(5-3);6230(6-1);6232(8-1);6381 (7-6a);6403 (14-ic); 6415 (7-5);6468 (2-3);6500 (24); 6506 (7-4c);6510 (3-2a);6584 (2-4 or 2-3); 6586 (2-3or7-5);6601(7-5);7187(7-4c);7208(14-7);7231 (14-ic); 7250(7-5);7450 (13-1);7453(5-3);7454(109); 7461 (13-4);7608 (2-4); 7734 (14-5);7776 (2-4); 8297(7-5);8359(7-4c);8411(8-1);8515(18-21);8531 (5-3);9585 (2-3);9630 (7-8);9956 (6-2);9987 (16-1); 10623(3-2a);10670(24); 13671(5-3);14122(9-27). Schunke,J. M., 319 (14-8b). Schwabe,W.,s.n. (7-6b). Schwacke,W., 12 (9-25);7153 (9-2); 11390(7-2); 11395 (6-3). Schwartz,G. J., 35, 869 (7-13b);4009, 4513, 4569, 4763, 4909,4934, 6153,7160 (7-2);7351,7360 (3-2a);8650 (3-2b);8681, 10488(7-2). 9080(9-2). Schwerdtfeger, Schwindt,E.,4966 (7-2). Scolnik,R. [et al.], 1209, 1212(3-2a); 19005(18-6);s.n. (9-32). Scottsberg, C., s.n. (3-2b). Secci6nCatastroForestalde Honduras, s.n. (1-1). Seemann,B., s.n. (3-2a,8-1). SEF(= Studieson Ecuadorian Forest);8542(5-3);8565(74b);8568(18-19a);8586(7-4b);8656(14-8b);8684(74b);8704(10-14);8733(7-4b);8776(5-3);8864(5-1); 9008,9039,9043 (7-4b);9107,9133 (5-1);9229(5-3); 9230(7-4b);9295 (10-l lf); 10134(5-1);10151(74b); 10358(5-1); 10404(7-4b). Segadas,F.,s.n. (9-19). Sehnem,A., 3232 (3-2a); 3899, 4070 (7-2); 5076, 5086 (3-2a). Seibert,R. J.,464 (7-9);502 (9-27);566 (7-9);574 (9-27); 576(7-9);1533(8-1);1581,1583(2-3);1671,1672(24). Seidel,R. [etal.],2181,2238(7-5);2247(24); 2868(6-2); 3067 (8-1);4391 (9-13);4636 (7-5);4882 (6-2);5308 (6-1);5433(2-4);5458 (6-2);5602(10-3);5904(6-2). Seiler,R., 546 (9-28). Seixas,E. M.,s.n. (9-32). Seler,C. et al., 2512 (9-27). Sello(w);F., 178(7-2);232 (7-1);756 (7-3);757,787 (7-1); 977,1184(7-3);1185(7-1);1220,1235,2152(7-2);2144, 2156(3-2a);2188,2195(7-2);s.n. (6-3,7-2,7-3, 9-25). Sermeflo,A., s.n. (2-1). Sesse,M.et al., 328 (9-27);330 (9-28);594 (9-27);594bis, 595, 596 (9-28);708 (9-27);4645, 4646 (3-2a);4647 (2-3). Sexton,0. J.,s.n. (7-9). Seymour,F.C., 4339 (2-3). Shakaim,RBAE.16a(14-3).

335

Shank,P.[et al.],24 (8-1);4384 (7-4a);4827 (7-9). Shannon,W.C., 97 (9-27). Shapiro,G., 395 (13-5). Sharp,A. J.,45445,46185 (1-1). Shattuck, O., 14(2-4);184(8-1);507 (2-4);829 (2-3);1021 (2-4); 1104(8-1); 1164(2-3). Shepherd,G. J.,4089 (9-32). Shepherd,J. D., 139, 191(2-3). ShilomTon,A., 2170(1-1);2735,3023(9-27);3680,3784 (2-3);3852(2-1);4346,4388,4683,6493,6514,6557, 6576, 7029, 7216 (2-3);7309 (2-1); 7949 (2-3); 8029 (2-1). Shimek,B. et al., 118(9-28). Shunsuke,T.et al., B.1846(7-6a). Sieber,F.W., 105(3-2a);339, 414, s.n (9-27). Sigueira,R., 3657,4022 (6-3). Silva,A., 124(6-3);365 (9-30d);464 (5-2). Silva,A.S. L.daetal., 54, 123(7-1);921 (6-2);1753(7-1). Silva,I. A., 26 (7-1). Silva,J.A., 312 (7-1). Silva,J. M., 284 (9-7). Silva,M. [et al.], 309, 754, 787, 796 (7-14); 1106 (6-3); 1326(7-1). Silva,M. F.et al., 1275(7-6b). Silva,M. F.F.et al., 1135(9-32). Silva,M. G. [et al.], 1409,2492 (6-3);3258 (10-2c);3425 (6-3);3707(15-2);3724(12-1);3945,4000(7-1);4269 (6-2);4660 (13-3);5913 (13-4). Silva,M. Mattos[et al.], 136 (9-1); 171(9-25);475 (9-1); 818 (9-30a);1428(9-1). Silva,N. T. [et al.], 6, 171(6-3);386 (7-1);549 (6-3);556 (5-2);600 (6-2); 1144, 1171(6-3); 1354(15-2); 1395 (6-3);1431(13-3);1591(7-1);1625(3-2a);1689(6-3); 1871(11-1);2553(4-1);2935(14-9a);2941(16-4);3329 (4-1);57793(14-8a);58429(7-3);60618(7-4b);60702 (15-1b);60779(15-2);60784 (13-1);60808(7-6b). Silva,P.E. Nogueiroet al., 33 (7-1);89 (3-2a). Silveira,F.R. da [etal.],s.n. (9-25) Silveira,M. et al., 688 (7-7);700 (18-12);860 (10-1b). P.A. [etal.l, 1098(7-8);3173(24); 3209, Silverston-Sopkin, 3591, 3659,4122 (7-13a); 5165,5660,5716,5808 (7-8). Simonis, E. et al., 24 (9-25); 5938 (10-1b). Simpson, D. et al., 53 (8-1); 60 (6-2).

SinacaColin,S. et al., 618 (8-1). Sintenis,P.,995,2126(3-2a);2294(9-27);2382(3-2a);2448 (9-27);3760(3-2a);6309 (2-3). Skutch,A. F.,925, 1511(2-1);1986(2-3);1986(2-1);2003, 2004 (6-1);2050 (9-27);2094, 2383, 2386 (2-1);2526 (9-10);2750 (9-27);3006 (9-10);3779 (1-2);4582 (910);4668,4674 (2-4);4960 (5-3);5374(9-10). Smeathman, H.,85, s.n. (9-27). Smith, A., 127, 141, 391 (7-8); 519 (1-2); 802 (7-8); 1004

(9-10);1423(1-2);1558(2-1);1895(9-10);1951,2313, 2720,2857,4227 (7-8).

Smith,A. C., 1917(9-10);3543 (4-1). Smith,C. E. et al., 3298 (8-1);3333 (2-4). Smith,C. L., 133(3-2a). Smith,C.W.,601 (6-2); 12519/25(4-1). Smith,D. [etal.], 574 (7-4a);1213(2-4). Smith,D. N. [et al.], 1213 (2-4); 2492 (6-2); 3125 (24); 3131 (7-4c); 3149 (1-2); 3900 (18-21.note); 3976 (14-

la); 3998 (7-4c);4168, 4382 (1-2);4650, 4651 (6-1); 4749, 4900 (1-2); 5164 (7-4c); 5172 (7-7.note);6173 (3-2a); 6649 (3-4); 8027 (1-2); 8638 (13-8a); 8650 (1-2); 13169 (8-1); 13411 (1-2); 13568, 13727 (6-2); 13810 (2-4); 13831 (8-1); 13866 (6-2); 13879 (1-2);

13981(8-1); 14131(10-1b);14159(6-1); 14175(8-1);

336

FLORANEOTROPICA

P.C.[etal.],7331(9-27);7557(3-2a);7595(9-10); 14205(6-1); 14232,14248(3-1); 14386(13-1);14387 Standley, 8375(2-3);8736(9-27);9436(9-28);9697,9697(9-27); (7-7); 14310,14331(7-13.note);14381(6-1). 9965 (2-1);10053,10865(9-27);11182,11186(9-28); Smith,J. Donnell,1619,2027 (9-27);6378 (3-2a);6777, 11274,11468(3-2a);11590,11736(9-28);13471,14114 6784(9-10). (2-1);15191(9-28);17745(3-2a);17842a,18431(3-1); Smith,F.D., 221 (7-13a). 18683(2-3);19204(9-27);19307,19688(3-2a);19784 Smith,H.H. [etal.],424 (6-1);425 (3-2a);476, 1444(9-27); (9-27);19797(2-3); 19981(9-27);20183 (2-1);20258 1752(2-3);1928,2027(9-27);2411(7-13a);2732(2-3). (3-2a);20680(2-1);20720,20795(3-2a);20824(9-28); Smith,L. B. et al., 3376 (9-40);6152 (9-7); 11823(3-2a); 21030(3-2a);21343(9-27);21404(2-3);21568(9-28); 12355(9-7);12605,12659(3-2a);14111(9-31);15405 21878,22036(9-27);22251(3-2a);22252(9-27);22335 (9-25). (9-28);22340(2-3);22358,22359(9-28);22395(3-2a); Smith,N., 128(7-14). 22458(2-3);22562(3-2a);23381,23459(9-27);23468, Smith,R. V.,V.4171(2-3);K-6260,K-6261(7-13a);7139 24351(3-2a);24552,24846(9-27);24850(9-17);25502, (10-3). 25595,25684(9-27);25735(24); 25968(9-28);26162, Smith,S. F. [et al.], 410 (10-5); 1018(13-3); 1302(18-7); 26833(3-2a);26983(9-27);27179(2-4);27183(9-28); 1445(154); 1556(14-7). 27228 (2-4);27287 (2-3);28200 (3-2a);28410 (7-9); Smith,S. Galen.[et al.], 1189(3-2a);1531(9-27). 28415(8-1);28890(3-2a);29286,29354(9-27);29381 Sneidern,K. von, 1094, 1171(2-3);A.1283(18-15);5169 (2-3);5712(9-27);5807, 5808(18-6). (3-2a);29510(9-27);29659 (2-4);29963(7-9);30235, 30296(2-3);30301(3-2a);30970(2-3);31013,31023(2Snethlage,E. H.,s.n. (7-1). 4);31315(7-9);31357(8-1);31454(2-4);31543(3-2a); Sobral,M. et al., 5169,5173, 4521 (7-2). 31697,31703(2-4);38808(1-2);39932(2-1);40121(2Sobrevilla,C. et al., 1700(13-3);1899(15-1b). 3);40168(3-2a);40797(7-9);40919(8-1);40943(2-4); Soejarto,D. D. [et al.], 794 (7-4c);2379 (15-4);2760 (1841053(7-9);41080(2-3);42176(3-2a);43272,43305(719a);2797 (12-1);3633 (6-1);3931 (7-4b);3950 (121);4070 (18-6);4092 (7-4b);4099 (2-3);4104 (16-1); 8);45271(2-1);46422,46436,47816(3-2a);48591(23);51028,51146(3-2a);51156(7-8);52647(9-10);52812 4178 (7-4b);4524 (13-4);7256 (2-3). (2-3);53114(9-10);53841,54312,54727,54741,54766, Soels,K., KSW-4=y-37(8-1). 54997 (2-3);54997 (3-2a);54677 (6-1);55024 (3-2a); Sohns,E. R., 1444(9-27). 55377(6-1);55601 (9-10);56771 (2-1);58131,58244 Solano,J., 50, 56 (7-4a). (9-27);60197(2-1);62208,62280(9-27);62563(1-1); Solheim,S. L., 1626(3-2a). 62861,63503(9-27);64165(3-2a);64564(9-27);64766 Solis,M., 92 (3-2a). (2-1);64847(9-27);64995(2-1);65029(1-1);65815(2Solomon,J. C., 630 (9-29);3530 (7-5); 6289 (6-3); 6426 1);65384(1-2);66635(3-2a);68333(1-1);68640(1-2); (7-6a);6234(10-6);6367(13-1);6444(18-6);6445(1068663,68756(2-1);68781(9-27);68933,68956,70185, 3);6450 (15-4);6516 (7-1);11201,12076(1-2);12694 70403 (2-1);70489 (9-10);70793 (2-3);70798 (9-27); (16-7); 14155(8-1); 14166, 14175(2-4); 14191(6-1); 14224(2-4); 14226(7-5); 14648(3-1 p.p.); 14721(771411(9-10);71595 (2-1);72483 (2-3);72910 (9-27); 13b);14785(10-5); 16668(1-2); 16832(13-3); 19197 72867(8-1);72959(9-10);72975(9-17);72984(9-27); 73022(9-10);73629,74013(3-2a);74033,74248,74349, (7-4a);19305(9-10). 74414(9-28);75250(3-2a);77524,78059,78192,78205, SoriaS., M.A., 22 (6-2). 78606(9-27);78679(3-2a);79291,79513(2-3);79523 Sosa,D. et al., 101, 168(3-2a). (3-2a);79524(2-3);79528(9-27);79606,79610(3-2a); Sosia,N. et al., 2315 (7-13b). 79986,81255,81359,81968(1-1);83368,84572(2-1); Sothers,C.A. et al., 66 (18-5). 84999(7-8);84999,84999(2-2);85036(1-2);85623(2SotoNuniez.J. C., 2934 (1-1). 2);85623(2-1);85950,86288(1-2);86697,86808,86994, Sousa,J., 89 (6-2);71 (10-2c). 87012(2-1);87412,88275 (9-27);88452 (2-3);88459 Sousa,M. [etal.], 2275, 11973,12381(8-1). (9-27);88549,88729(2-3);88941(9-27);89114(2-1); Souza,J. M.A. de et al., 144(10-3);157(7-5). 89114(2-3);89574,89897(2-1);91635,91677(9-17); Souza,S. A. da M., 323 (7-6a). 91691(2-1);91728(9-17);92292(9-10);s.n. (1-1). Souza,43 (9-26a);202 (3-2a);269,318 (9-26a);339 (9-25). Soza,D. et al., 70 (2-1). Starry,D. E., 84 (7-9). Sparre,B. [et al.], 723, 1637(9-32); 14469,14496,15490 Steam,W.T.,951, 965 (9-34). Steere,W.C., 1273(9-27);1352(3-2a);1640(9-27);1649, (7-10). 1815(3-2a);1882,2485 (9-27). Spegazzini, C., 15454, 15455 (7-2); 15461 (9-32). Spellman,D. et al., 173(9-27). Steggerda,M., 1 (9-27);24c (3-2a). Sperling,C. R.et al.,6400(6-3);6425(18-21);6471(18-6). Stein,B. A. et al., 2575 (7-5);2777 (7-8);3931 (7-9). 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[et al.], 136 (3-2a); 180 (7-9);210 (2-3);233 Stanford,L. R., 6906 (9-33). (7-9);272 (9-27);364 (7-9);390 (9-27);421 (7-9);593

LISTOFEXSICCATAE (9-27); 662 (7-11); 662 (7-9); 728 (24); 754 (3-2a); 770, 771 (7-9); 914 (9-27). Stevens, F. L., 12 (7-9); 1065 (2-5). Stevens, P. L., 5882 (3-2a). Stevens,W. D. [et al.], 1343 (9-28); 1398 (9-27); 1426, 1805, 2644, 3374 (9-28); 3453 (9-27); 3852 (9-28); 3992 (927); 3999 (2-3); 4717 (9-28); 4974 (9-10); 6722, 9171 (2-1); 9251,9665,9716 (3-2a); 9722 (9-28); 9945,10787 (2-3); 11778 (3-2a); 12209 (2-3); 13149, 13226, 13227 (3-2a); 13606 (9-28); 14742 (2-3); 15286, 16597 (9-27); 16843, 16912, 16982 (2-1); 178422 (3-2a); 17668, 18566 (9-27); 18882 (2-1); 20131 (9-28); 20416 (2-1); 20582 (9-28); 20940 (6-1); 21174 (2-1); 21831 (9-27); 22171, 22174 (2-1); 22162,22262 (3-2a); 22321,22363 (9-28); 23412 (9-10); 23444 (7-4a); 25552, 25840 (9-17). Stevenson, J. A., 2407 (2-3). Stevenson, N. S. et al., 140, 153 (2-3). Stevenson,P.,25(16-1); 31 (13-4); 69,71 (6-2); 216 (10-14). Steward,W. et al., P.17649 (18-14). Stewart,R. M., 518 (1-1). Steyermark,J. A. [et al.], 17142 (2-4); 17494 (7-9); 29198 (1-1); 29462 (9-28); 30514 (9-27); 31511 (2-3); 33499, 33636 (2-1); 33768 (9-27); 35461 (2-1); 36300, 37046 (1-2); 37426,37576 (2-1); 38607 (9-17); 39057,39122, 39123 (9-10); 39125 (9-27); 39157 (2-1); 39289 (9-17); 39451, 39932 (2-1); 39972 (9-17); 41663 (9-10); 41801 (7-8); 41995 (9-10); 42003 (9-27); 42744 (2-3); 43588 (2-2); 43588 (2-1); 43644 (2-2); 43664 (2-1); 44542, 44732 (9-17); 45445 (2-1); 45505 (8-1); 45516 (9-17); 45955 (8-1); 46070 (9-17); 46120 (9-27); 46630,46658, 46687,46730,47333,47349,47392 (2-1); 47670,47774, 49549 (2-3); 49681 (9-27); 50515a (1-1); 51030 (2-3); 51075 (9-27); 51185 (2-3); 51135, 51354 (9-27); 51856 (1-1); 52069 (2-1); 52118 (9-27); 53781,53817,54570, 56473 (1-2); 56568 (9-27); 57579 (8-1); 60721 (10-9); 61158 (8-1); 61467 (10-5); 61846 (13-8a); 61949 (3-1); 62352 (3-2a); 62689A, 62768 (9-27); 86310 (7-13a); 86332 (74b); 86720 (3-2a); 89299, 8933 IA (9-32); 91841 (9-14); 92080 (8-1); 95395 (13-8b); 96344 (167); 96550 (13-1); 97521 (9-14); 98263 (8-1); 98674, 98852 (1-2); 99270 (9-27); 99683,99726 (2-3); 99865A (6-1); 99873 (8- 1);99942 (10-5); 100 140 (3-2a); 101253 (1-2); 101931 (13-4); 101957 (7-13a); 102044 (2-3); 102135 (7-13a); 102227 (3-2a); 102249, 102409 (2-3); 102860 (12-2); 104709 (16-7); 107061 (11-1); 107192 (13-1);107063 (7-6a); 107428 (6-3); 107625, 107625 (23); 107626 (10-1b); 107825 (3-2a); 108426 (10-5); 109910 (3-2a); 110192 (13-8a); 110724 (3-2a); 110732 (9-27); 110968 (3-2a); 111061 (9-27); 111807 (7-15a); 113749 (7-13a); 113900 (7-6a); 113942 (6-3); 120264 (3-2a); 122172 (7-6b); 122181 (7-4c); 122242 (10-3); 122348 (7-6b); 122349 (6-3); 122595 (7-13a); 122651 (9-27); 122765 (2-3); 122949 (7-13a); 123044 (9-27); 123135 (3-2a); 123176(10-1b); 123188,123122,123191 (9-27); 123207 (3-2a); 123209, 123301 (2-3); 123304 (3-2a); 123308 (2-3); 123361 (7-13a); 124341 (7-6b); 126845 (13-8a); 127009 (13-8b); 127056 (9-27); 127141 (7-13a); 129387 (7-6b); 131195 (9-32). Stijfhoom, E., 659 (3-2a); 695 (7-1). Stoffers, A. L., 245, 295, 1320 (3-2a); 5112 (7-13a). Stordy, R. J., s.n. (1-1). Stork, H. E. [et al.], 16,46 (2-3); 2531, 2780 (9-27); 3389 (3-2a); 4227 (7-8); 8752 (9-28); 8913 (9-27); 9506 (2-3). Strang, H. E., 408 (7-2); 627 (9-26b); 917 (9-25); 961 (926b); 1025, 26612 (7-2). Stricker,R., 343, 344 (9-27).

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338 (7-1); 10439(6-3);10708(9-8); 10857(7-3);11078(98); 11290(7-1); 11326(9-30a). Thomsen,K.,58822(16-7);58829(7-12). Thompson,C., 40 (9-27);853 (13-8a); Thomson,R., 1384(3-2a). Thorne,R. F. [et al.],40218 (9-27);40536,40540(6-1). Tillett,S. S. et al., 45495 (7-4b). M. [et al.], 1042(14-la); 1243,1245(18-6). Timan'a, Tinney,C., 76-113,76-170(3-2a). Tipaz,G. [et al.], 417 (14-2);451 (14-3); 1041, 1044 (712); 1074 (18-17); 1240 (7-12); 1309, 1362 (18-17); 1394, 1811, 1865 (18-18a);2071 (18-19d);2319 (161); 2556 (6-2); 2581 (7-4b);2595 (7-4b);2599 (6-2); 2603, 2610 (7-4b);2624, 2661,2689 (6-2). Tirado,M. et al., 172(10-1b);247 (18-16);450 (18-19d); 457,493 (6-2);514, 1073(18-19d);1306(18-18b). Toledo,S. C., s.n. (9-32). Tomas,Bro.,1823(2-3). Tonduz,A. [et al.], 151, 152 (9-27); 6683 (2-4); 6751, 6751bis,8124(74a); 8338(6-1);8563(74a); 8855(927);9339, 9389, 9524 (9-10);10122(9-27); 11918(12); 12802, 12844 (7-4a); 12883 (6-1); 14357 (7-4a); 17880(9-27);s.n. (9-10). TorresD., M. et al., 1103(18-15). Torres,J. H. et al., 2848 (3-2a). TorresC., R. et al., 259 (2-3);6886 (3-2a). Traill,J. W.H., 712, 714 (7-6a). Treacy,J.et al., 143(5-3);185(74b); 337 (5-3);534 (7-7). Tressens,S. G. et al., 3071 (3-2a). Triana,H. et al., 56 (18-17). Triana,J. J., 835 (5-3);836 (3-2a);850 (9-27);852 (8-1); 857 (24); 870(1-2);1857(3-2a);1860(9-27);1861(81); 1863(2-4);3605, 3606(2-3);s.n. (2-3, 2-4, 8-1). Trinta,E. Fromm,2276 (9-25). Trinta,Z., 1343(9-25). Troll,C., 277 (9-27);472 (2-3); 1393(9-32). Troncoso,2255 (9-32). Troon,F.vanet al., 16297(6-2); 16297(6-1). Troth,R. O., 1549, 1550, 1551, 1568(7-13a);1579(3-2a). Trujillo,B., 2224, 3786 (3-2a);4415 (9-32);6443, 12214 (3-2a). Tsugaru,S. et al., B.1174(15-4). Tucker,J. M., 508 (9-28);705 (9-27). H. von, II-43(9-17);310 (9-28);751 (9-10); Tuerckheim, 1231(2-1);1709(9-10);II-2013(9-27);II-2013(9-28); 2133(9-10);II-2167,11-2169,II-2170(2-2);11-2207(928);5611 (9-10);7667 (9-17);8581 (2-3);s.n. (9-27). TunOrtiz,R., 11(2-3);234 (9-27);294,929 (2-3);1200(32a);1480(2-3);1630(2-1);2008(2-3);2539(2-1);2559 (3-2a). Tunqui,S.,4 (24); 130,224(2-3);668(5-3);671,813(7-6a). Tumer,B. L., 1948, 1984(9-28). Turner, M., 110(3-2a). Tweedie,F.,85 (3-2a);1299(9-25);s.n. (7-2, 9-32). Tyson,E. L. [et al.], 1453, 1767 (7-9); 2973 (2-4);4686, 4699, 4722 (7-9);5451, 5455, 6209 (2-4);6732 (2-3); 6910 (24); 7152 (1-2).

FLORANEOTROPICA Urrego,G. L. E. et al., 1243(10-3); Urville,J. S. C. Dumontd', s.n. (7-2). Utley,J. et al., 1037(9-10);4826 (2-1);4858 (7-8).

Valencia,R. et al., 311 (7-6a);354 (7-4c);355 (17-2);357 (7-4b);358,360(7-4c);361 (7-1);362(7-5);2809(1-2); 67331(18-10);67341(14-3);67375(10-5);67656(6-2); 67804(18-15);67840(15-3);68019(15-lb);68505(148b);68597(16-5);68569(14-Id);68654(14-8b);68667 (18-19a);68725(15-4). Valente,M. de Conceicao,21 (9-26b). Valerio,M., 88 (7-8); 1306,1307(3-2a). ValerioR., J.,214,2122, 3380,3470 (3-2a). Vareschi,V.et al., 2717, 2718 (9-27). VargasA., J., 34 (3-2a). VargasC., C., 8523,8535 (3-2a);16208(14-1b). Vargas,E. et al., 737 (9-27);739 (3-2a);2083 (2-4). Vargas,I. G.et al. 1119 (6-2);1208(5-2);1279(1-2);2560, 2574, 2601(3-2a);2604 (5-2). Vargas,L. et al., 720, 727 (6-3);786 (7-1); 1018(10-lSb). Vasconcellos,J. de Morales,171(3-2a). VasconcellosSobrinho,(9-32). Vasquez,B., 276, 999 (8-1);6093 (14-3). Vasquez,R. [et al.], 268, 609 (7-6a);742 (3-2a);802 (1819a);881 (2-3);1041(18-10);1529(3-2a);1669(2-4); 2498 (7-4c);2499 (7-6a);2982 (7-5); 3422 (18-19a); 3396 (2-3);3426 (5-3);3456 (10-3);3571 (5-3);3817 (10-lSb); 3984 (7-4c);4144 (2-3); 4340, 4383 (7-5); 4588(14-5);4695(7-4c);4958 (3-2a);4994(7-5);5027 (3-2a);5173 (18-20);5214 (2-3);5321 (10-1If); 5484, 5508(7-5);5601(18-20);5692(7-5);6002(7-4c);6073 (6-1); 6146 (18-13);6223 (7-4c); 6251 (13-4); 6281, 6486, 6507 (7-5);6680 (5-2);6891 (3-2a);6914 (7-7); 6996(2-3);6999(18-6);7023(14-5);7207(16-5);7762, 7777A(7-4b);7905(7-5);8211(7-4c);8302(6-1);9003 (14-5);9374, 9497, 9544, 9560, 9574 (7-4c);9679 (7.. 6a);9918(3-2a);9968,10027(7-4c);10102(2-3);10437 (6-2);10446(18-10);10515(3-2a);10537(14-5);10544 (6-2); 10667(5-2); 11275(14-8b);11309(5-2); 11468 (14-4);11927(5-3); 12004(6-2); 12032(7-14);12172 (6-1);12236(18-20);12520(7-4b);12525(7-5);12539 (7-1); 12541,12566(7-6a);12697(7-5); 12754(7-6a); 12786(7-4c); 12915(3-2a); 13065(7-6a);13092(1815);13118(7-5); 13523,13616(7-6a);13634(18-20); 13480(13-8c);13731(14-5);14085(18-20);14137(14lb); 14197(14-5);14263(18-3);14741(10-14);14791, 15170(14-9b);19342(13-3);19878(10-9). Vaulthier,134, 161, 191(3-2a);s.n. (3-2a,7-3). Vavrek,M. et al., 1 (3-2a). VizquezA., E., 34 (6-2); 179(1-2);213 (7-2). VizquezAvila,M., 304 (3-2a);346 (1-2);378 (3-2a);386, 389 (8-1);456 (5-2). VizquezT.,M., 1732(2-1);2236 (2-3);3883 (2-1). VazquezT.,V., 197,215, 529 (2-3). Vecchi,O., 216 (3-2a). Velazco,J., 203 (7-6b);741 (10-14). Velez,M. C. et al., 908, 914 (8-1);988, 1022(6-1). Veillon,J. P., 17.V(6-1);77 (2-3). Ugent,D., 128, 129(2-3). Velloso,H. P.,917 (7-3). Ule,E.,376(9-30d);757(7-1);1291(7-2);4369(7-1);4378 Ventur,P.83 (2-3); 174(2-1). E. et al., 299 (2-3);2074, 4645 (3-2a). (9-18);4383 (7-3);4943 (9-3);4992 (9-26a);5267 (7- Ventura, 1);5714 (3-2a);5771, 5869, 5870 (7-6a);5871, 5871a Ventura A., F., 732 (2-3); 995 (9-27);999 (2-1); 1325 (3(7-5);6412(2-5);6509(9-27);6838(2-3);8115(7-13a); 2a); 2489 (9-28);3102 (2-1); 3342, 3495 (2-3); 3650 9317(6-2);9319(9-30d). (3-2a);4329(9-28);4559,4745 (2-3);5116(2-1);5273 L. M. et al., 1512(3-2a). Underwood, (9-27); 5735 (3-2a); 8100 (2-1); 9240, 15074 (2-3); 16191(3-2a);19166,20157 (2-3);20964,20974(2-1); Unshelm,s.n. (9-32). 21442(6-1);21239(2-3). Uribe-Uribe, L., 1221(7-9b);1573(2-5).

LISTOFEXSICCATAE Venturi,S., 16(3-2a);5041,5396, 5551 (3-2b). VeraSantos,J., 2831 (2-1). H.[etal.],17(14-5);53(18-13);269(18-10);399(6-2). Vester, Verlet,983 (1-1). Vianna,M. C. [etal.],822 (9-26a);828(9-3);1201(9-26a); 1442(9-30d);1542(9-26a). Vieira,G. et al., 37 (7-6a);289 (18-1);291 (7-6a);311 (189); 322 (7-1);375 (7-6a);512 (6-3);530 (3-2a);557 (71);571 (7-6a);762 (16-2);813 (6-3);840 (10-6). Vieira,J. N., 74 (9-8). Vilaca,A., 90 (9-26a);105(9-25). Villacorta,R. et al., 252 (2-3); 1107(9-28). VillegasH.,A., 71 (2-1). Villegas,M.,203 (16-1). Vincelli,P.C., 384 (2-4); 1139(7-4b);1187(7-6a). Vinha,S. G. da,90 (9-21). VivarC., F.,814 (7-8); 1547,2345 (1-1). Vogelmann, H.W.et al., 1302(18-15). Vogl,C., 785, 786 (9-27). Wachenheim, H.,74 (4-1). J. L., 699 (9-31). Wachter, W.H. [etal.], 35466(3-2a);71063(9-27). Wagner, Walker,F., 1262(9-27). J.W., 141(9-28);148(9-27);149(9-28);334 (2-1). Walker, Wallace,R., 82 (14-8a). Wallnoefer, B., 11-031188(7-7);11-090188(7-1);11-3788 (13-8a);13-181088,13-21888(7-6a); 13-221088(7-5); 15-221088(7-6a); 15-27988(74c); 16-81088(7-6a); 18-11088(10-14);113-13988(16-7). Waltier,F.dasC. S. et al., 42 (10-2c). Warming,E., 373 (9-27);674, 1903 (3-2a); 1937 (9-32); 1938(9-30c);1939(3-2a);1940(9-6);1941(13-4);1949 (7-1); 18811(7-2);s.n. (3-2a,9-30c). WarushJuwa,R., RBAE.4(10-lb). D. C. et al., 693 (2-4). Wasshausen, Watson,S., 296 (9-10). WawravonFernsee,H. [etal.],256 (9-21);429 (9-25);501b (9-26a). Weber,33366(7-2). Weberbauer, A., 1853(7-1); 1912(6-2);2057 (6-1);7040 (1-2); 7057 (6-1); 7059 (2-4); s.n. (6-2, 10-13).

Webster,G. L. [et al.], 196 (9-40);4073 (9-35);5056 (32a);5149(9-34);5642(2-3);8074(3-2a);8324(9-34); 9245 (3-2a); 11932 (9-28); 12522 (3-2a); 12701 (9-10); 16814 (2-4); 16901 (7-8); 23342 (7-4c); s.n. (3-2a).

Weddell,H.A., 122(7-1); 165,291, 742, 790 (9-25);2578 (3-2a);2882 (9-30c);3280 (3-2a);3434 (9-32);s.n. (925, 9-32). Wedel,H. von, 196(2-3);598-A(7-9);896,941 (2-4);961, 962, 1003 (7-9); 1043 (2-4); 1090 (7-4a); 1094 (2-4); 1103, 1390 (7-9); 1530 (9-27); 1582 (2-4); 1599 (2-3); 1715 (7-9); 1719 (2-5); 2075 (2-3); 2736 (2-4). Weir,J.,44 (9-3); s.n. (9-3, 9-19).

339 Whitefoord, C., 1392(9-34). Whiteford,H.N. et al., 14, 59, 70 (3-2a). Widgren,J. F., 499 (9-25);608 (9-30a);729 (9-26a);s.n. (7-2,9-25,9-30a,9-32). Wiggins,I. L., 11017(1-2); 11764(1-1). Wilbur,R. L. [etal.], 1450,1537,1621,2420(9-28);14930 (9-10).

Wiley,J. R., 551A (9-27). Wilkison,E., 2446 (1-1). Williams,LI.[etal.], 115(2-3);201 (7-5);501,672,821 (32a);2086 (6-1);2092 (7-4b);2357 (74a); 2441 (3-2a); 2697,2965(7-4b);3079,3380(6-2);4598(3-2a);4658, 5243(2-3);5334 (5-2);5435 (3-2a);5905 (9-27);6254 (3-2a);6471 (6,1); 6542 (2-3);6544 (2-4);6573 (2-3); 6803 (7-7);6819 (2-4);6832 (9-27);6855 (3-1);7195 (2-4);7743(6-1);8010(7-6a);8040,8050,8217(3-2a); 8426(8-1);9520(2-1);9617,9821,10031(3-2a);10719 (2-3);11181(9-27);12040(7-15a);13191(6-2);13377 (9-32);13488,14532,15079,15184(7-6b). Williams,L. 0. [etal.], 12483(3-2a);13089,14432,14459 (9-10);14815,15601,15799(2-1); 16450(7-8);16764 (3-2a);16850(2-1);18562(3-2a);23350,23839,24941, 24963,27620,27825,29170,29222(2-1). Williams,R. S., 260 (9-32);412 (7-8);938 (9-27). Willis,W.D. [et al.],229 (9-28). Wilson-Browne, G.,431, 432 (4-1). Wilson,C. L., 277 (9-28). Wilson,E.T., 1629(2-3). Wilson,P.,54, 250 (2-3);587, 1062(3-2a);9386 (2-3). Windisch,P.,1684(9-30d). Wingfield,R., 5730,8037, 10071(7-13a). Winzerling, H., II-4(3-2a). Wisniewsky, A., s.n. (4-1). J.T.et al., 8291 (7-9). Witherspoon, Wolfe,F., 12307(16-7). Wolle,J.,s.n. (9-36). Wong,W.,87a(9-27). Wood,C. F. [et al.],7465 (9-40). Woodbury, R., 3426,s.n. (2-3). Woodson,R. E. [et al.], 866 (2-3);899 (3-2a);1383(7-9); 1505(9-27);1924(2-3). R. H. et al., 323 (7-9);372, 604,666 (2-4). Woodworth, Woolston,A. L., 684 (9-30b);705 (7-2);1026(3-2a);1313 (9-32). Woytkowski, F.,1317(3-2a);5114(8-1);5534(14-2);5645 (2-3);5862(13-3);5865(6-2);6295,6317,7055(3-2a); 7088(7-7);7274(2-2);7374 (2-5);7628 (7-4c);35079 (3-2a). Wright,C. [et al.], 589, 592, 594 (2-3); 1439(3-2a);1446 (9-45); 1447 (9-44);2224, 2225 (9-41); 2226 (9-42); 2227 (9-35);s.n. (2-3, 3-2a). Wright,P.C., 83 (7-13b). H. R., 1381a (3-2a). Wullschlaegel, R. et al., 1163(3-2a). Wunderlin, J. J. [et al.], 2117 (2-3);2325 (16-2);43455 (7Wurdack, 6b);43633(10-1If). Wydler,H., 291 (3-2a);375 (2-3).

Weiss,G., 178(8-1). Welch,W.H., 19824(7-9). Wendland, H., 921 (9-10). Wendt,T. [etal.], 2764 (7-8);3341 (10-5);3671 (8-1). Werdermann, E., 2318,2542 (7-13b);2653, 2655(3-2a). Yanez,A. P.et al., 1306(6-1). S., 713 (3-2a). Werff,H. vanderet al., 9525, 9527 (18-17);10290(16-1); Yepes-Agredo, 10296, 10378(14-id); 10846, 12497 (1-2); 13127 (2-4). Young,H. J. et al., 323 (7-1). Wessels Boer, J. G., 1069 (7-6b); 1080, 1879 (6-3). Young,K. et al., 5005 (1-2). Wetmore,R. H. et al., 46 (2-4). Youngken,H., 114(8-1). White,D. A., 167,215 (3-2a). Yuncker,T. G. [et al.], 4558 (9-10);6171 (2-1);6382 (9White,0. E. [et al.], 892 (6-2); 1011(2-5); 1061(6-2). 27);8493(9-10);8550 (9-27);18562(9-34). White,S. S., 2893, 2925a,3749, 4847 (1-1);5077 (9-28); 5432 (3-2a). Zabala,R. et al., s.n. (6-1).

340

FLORANEOTROPICA

Zak,V. [etal.],2705,2880, 2933,3215 (1-2);3787(13-9); 3955 (10-13);3974 (5-3); 3976 (10-13);4067 (10-5); 4116, 4178 (10-13);4236 (13-1);4244 (10-13);4253 (10-2c);4349(10-13);4425(13-8c);4613 (10-5);4731 (18-19a);4949 (7-4b);4982 (10-3);5015 (5-3); 5020 (7-5); 5067 (14-ic); 5071 (13-4); 5099 (7-6a);5101, 5162, 5163(7-4b);5170 (5-3). ZamoraV.,N., 759 (6-2);996 (3-2a);1491(7-4a). Zanoni,T.et al., 149977(3-2a);17686(2-3);21899(3-2a); 25206,26104 (2-3);34288 (13-8);33365(2-3);34310 (9-41);35468,35830(2-3). Zardini,E. [et al.], 2982, 3156, 3172 (7-2);5493 (7-13c); 5940,6915,7085,7291,7926,8487,9750(7-2);33250, 33758,33866(7-13b).

Zarucchi,J. L. et al., 2375 (2-3); 2667 (4-1); 2698 (7-1); 2700(15-3);2701,3742(10-3);3746(15-4);3817,3800 (7-5);4083 (10-1b);4122, 4650,4663 (3-2a). Zaruma,J. [et al.], 50 (6-1); 172 (7-5); 215 (18-19a);223 (7-5);290,361(6-1);379 (5-2);442 (18-6);601(18-21); 684, 685 (2-3);687, 689 (2-4);769 (5-3);803 (2-4). Zehntner, 590,626,4053, 4077 (3-2a). Zent,S., 785-12(6-3). Zetek,J.,3684 (3-2a). ZolaB., M. G., 404 (2-1). Zuloaga,F. 0. [et al.], 648 (9-31); 1433 (9-32); 1421 (930d);1522(9-32). ZuluagaR., S., 757 (5-1); 1412(2-4); 1478(7-9). Zuiniga, R., 74 (2-4);237 (7-4a).

INDEXOF LOCALNAMES Numbersfor eachlocalnamereferto theNumberedList of Taxa.Numbersfollowedby -- indicatevernacularnamesnot linkedto a specificspecieswithinthatgenus. abasbabi(8-1) aji(6-2) almorabranca(6-3) amapa-rana (4-1) amarillo(3-2a) amateblanco(8-1) amorabranca(3-2a,6-3) amoreira(3-2a) apii(9--) ayamoyegui(7-6a) bacasse(4-1) bagasse(4-1) bainhade espada(7-1, 7-2) barbade tamagaz(2-3) barbodillo(9--) barboria(9--) bejucomora(3-1) bibobetubu (7-11) billo (6-2) bobo(6-2) boisd'orange(3-2a) borejore(2-4) caapia'(9--)

cafecillo(7-6b) caiapia(9--)d caimbemi(do (7-14) caimbe(7-14) caimbe'(7-14)

cajiman(6-2) cambapunta(3-2a) cancorosa(7-2) capia(9--) capinuriblanca(7-7) caraipe(7-14) caraipedo igap6(7-14) carapa(9--) carapia(9--) carapicicade fohlamiuda(7-2) carapicica(7-2) caripa(9--) camero(8-1) carpales(9--) castanio (12-1)

castaiio(12-1) catagiva(3-2a) caudillocolorado(6-1) caxapia(9--) caxim-mirim (7-2) cegoolho(7-2) cha-cox(2-3) chagane(8-1) chanaique (2-1) chanchama (8-1) chara(7-13a) charo(7-13a) charoamarillo(2-3) charoblanco(2-3) charomacho(7-4b) charomorado(2-3, 6-1) charonegro(6-1) charo(7-13a) charonegro(7-13a) chatchi(2-3) chatojioji(7-7) cherry(2-3) chicalminia(6-1) chicle(5-1, 6-1) chileamate(8-1) chimicua(5-1, 5-3) chirimoya(8-1) chischillo(2-1) chupa-chupa (9--) cincho(7-2) ciruela(6-2) cocua(8-1) comawantzaui (7-6a) comede pescado(8-1) contra-erva (9--) contrahierba(9--) contrayerba(9--) copey(7-13a) corbon(8-1) corocho(8-1) cousajo(6-1) cow wood(4-1) cucua(8-1)

damagua(8-1) damajaguablanca (8-1) damaque(8-1) dicrinajai(7-4c) dinde(3-2a) encira(3-2a) ere(9-32) etona-tewe(7-7) figueiraterrestre(9--) figueirilha(9--) figueirilho(9--) f6lhade serra(6-3, 7-2) frutode pande monte(5-1) fustete(3-2a) fustic(3-2a) ganchincho-canxim (7-2) gele bagasse(4-1) gigant6n(7-13a) guachilamo(8-1) guacora(7-13c) guaimaro(7-13a) guaimarocimarron(12-1) guaimarolechoso(7-13a) guaimaronegro(2-3, 7-13a) guar(6-2) guarilama (8-1) guaritota(8-1) guariuba(6-2) guariubaamarela(6-2) guariubade caranazal(6-2) guaymaromacho(2-3) guillo(1-2) gusanero(5-1) higueronhabillo(8-1) hihounahi(6-3) hojatinta(2-3) houyoufi(6-2) huaguito(1-2) hualina(6-3) huanchal(2-1) huasca(1-2) huehue(6-2)

cucua (8-1 )

huichilama (8- 1)

INDEXOFLOCALNAMES ikicaspi(7-5) ikor-wala(8-1) incira(3-2a) insira(2-3, 3-2a) insiraamarilla(3-2a) insiracaspi(3-2a) ishangamasha(24) isirillo(6-1) jacabrava(7-7) janete(6-3) janita(6-2, 7-14) jaquitiade leite(7-3) jawahedan(4-1) jopi (7-4c) juandiegoamarillo(6-2) juremade espinho(3-2a) k'arap'acha (8-1) kal-dtsi(8-1) kamush(8-1) katowar(4-1) kauhoedoe(4-1) kaw-oedoe(4-1) kirinir6ki(81-) kusuknash (2-3) kusuknashi kumpari(7-6a) lechecaspi(5-2) lechero(7-13a) lecherocolorado(2-3, 6-1) lechilla(6-1) leilapo(3-2a) letterhout (6-3) liga-liga(9--) liga osso (9--) limao-rana (3-2a) limulona(3-2a) linguade onca(7-1) mahana(8-1) maitin(6-1) maja-nei(8-1) maja-sehi(8-1) maja-seji(8-1) majagua(8-1) mam6nde leche(7-13a) man-redhout (6-3) manchinguilla (7-7) manletterhout (6-3) marfil(2-3) mariamolle(7-2) mascajorojo(6-2) mashonaste(5-1, 5-2, 5-3, 6-1, 6-2) mashonaste amarillo(6-2) mashonaste blanco(6-1) mashonaste negro(5-2, 5-3) mashonastillo (2-4, 5-3, 7-5, 7-6a) masicaran (6-1) masicar6n(2-3) mastate(8-1) matopalo(8-1) mazitoto(8-1) miguelillo(7-12) mora(2-3, 3-1, 3-2a,3-2b) moraamarilla(3-2a) morachica(3-2a,3-2b) moracolorada(3-2a,3-2b)

341 moradel campo(3-2b) morade la pampa(3-2b) moragrande(3-2a) moranegra(1-1) moraccachocolate(6-3) moral(1-1, 3-2a,6-2) moralbasto(6-2) moralbobo(6-2) moralcomidade mono(6-2) moralfino(3-2a) moreira(3-2a) moreiraamarela(3-2a) moro(3-2a) morode la tierra(3-2a) murare(6-1) murure(5-2, 6-2) mururemole(5-2) mururehuasa (5-2) nanduli(9--) nandpi (7-2, 7-13a) naranjillo (6-2) naudipa(7-2) neandipamei(7-13b) niandyra(7-2) nisperoblanco(6-1) fiandypami (7-2) niepo(7-6b) oboye(7-5) odon(4-1) odoun(4-1) oeman-redhout (6-3) oiticica(6-2) ojoche(6-1) ojochemacho(2-3, 5-1) ojochillocolorado(2-3) omeganiko(8-1) orticica(6-2) paloblanco(1-1) palocerano(6-1) palochicle(6-1) palode brujo(7-6a) palode mora(3-2a) palovenono(18--) paloedoesje(3-2a) pandilla(5-3) paude serra(7-1) pelacara(6-2) petaquilla(7-10) pikiu(5-3) pintode Cerro(7-13c) pitiu(2-3, 5-3) pitiuk(5-3) pittiu(5-3) pitu(5-3) pituca(5-3) pituk(2-3) punte(8-1) puruhi(7-4b) quechoamarillo(5-2) rajito(1-2) ram6n(2-3) ram6ncolorado(2-3) ramonde bestia(2-3)

ramoncillo(2-1) ramoon(2-3) rumicaspiyura(2-3) sachachimicua(7-4c) sachaishpingo(5-2) sachapaparagua (5-3) sachavacamicuna(6-1) sande(2-3) sapanamarillo(1-2) sapirana(9-32) sapito(7-13b) savaleta(6-1) serraleiro (7-2) shitapachkusuknash (7-6a) sintaque(7-7) siticica(6-2) soroca(7-2) sorocaba(7-2) soroco(7-2) sota(1-1, 3-2a) suchu(7-2) suerpo(12-1) sunquirimsuwe(6-2) tachiore(8-1) tachori(8-1) taiuva(3-2a) tajiiba(3-2a) tajuva(3-2a) takatakaramio (7-4b) tamamuri (7-7) tapaguemohue (7-6a) tapoguene(7-6a) taraiko'i(4-1) tareka'y(4-1) tarope'(9--)

tata-yba (3-2a) tata-yba-pta (3-2a) tata-ybA-sayu (3-2a) tata-yyva(3-2b) tataguiba(3-2a) tataiba(3-2a) tatajuba(3-2a,4-1) tatajyva(3-2a,3-2b) tatayiba(3-2a) tatayieva(3-2a) tementayi(7-5) tentemoeyi(7-7a) teiui(9--)

teiu-a9u(9--) tetsi(8-1) tillo serrano(6-1) trempa(7-1) tsajanki(3-2a) tsakaya(7-4c) tujiguarce(7-4b) tulpar(6-2) tulpay(6-2) tulpayblanco(6-2) turuuru (7-5) tusilla(9-32) tuwue(4-1) uchipituk(2-3) uriamen(7-3) urupi(6-2)

149

FLORA NEOTROPICA

uvillo(3-2b) uvode monte(7-14) uyapitu(5-3)

venenu teiu (18--) venueo (1 8--) viecrinajae(74b)

(7-4b) vegotahuemo venenillo(18--) veneno(18--)

we-y6 (7-13a) white ramon (2-3) wica (6-1)

xamamore(6-3) xamaroenai(6-3) xanajoxo (7-6a) xoqueshequere (7-1) yanchama (7-1, 7-7, 8-1) yawahedan(4-1) yax-ox (2-1, 2-3)

INDEXOF SCIENTIFICNAMES arenominanudaordoubtfulnames.Pagenumbersin boldface Synonymsarein italics. Namesin [brackets] ormaps. indicateprimarypagereferences.Pagenumberswithanasterisk(*) indicatepageswithillustrations Acanthinophyllum,79 strepitans, 79, 88 ilicifolia, 88 var.cuyunensis, 90 var.guianensis,90 var.micrantha, 90 var.paraensis, 90 spruceana, 90 Acanthosphaera,269 ulei, 282 Aliteria, 79, 88 sagotii, 79, 90 Annonaceae, 10 Anonocarpus, 71 amazonicus, 71, 74 peruvianus, 71, 76 Antiaris, 9, 15, 244 toxicaria, 5 Antiaropsis, 11, 14, 67, 68, 289 Artocarpeae, 4, 9, 14, 15, 17, 20, 67, 6g, 138 Artocarpoideae, 4 Artocarpus,8, 10, 12, 20, 67, 68, 143 altilis, 22, 143 communis, 143 heterophyllus, 13, 22, 143 Bagassa, 5, 6, 7, 12, 67, 68 guianensis, 15, 16, 17, 18, 20, 21, 69, 70*, 302* sagotiana, 68, 69 tiliifolia, 69 Balanostreblus, 92, 98 ilicifolius, 92, 98 Batocarpus,6, 7, 69, 71, 72 amazonicus,14, 71, 72, 74, 75*, 303* costaricensis, 20, 71, 72, 73*, 302* maranhensis, 71, 74 orinocensis, 71, 76, 77*, 303* Bleekrodea, 9, 11, 14, 25 Bosqueiopsis, 10, 144 Brosimeae, 4, 231 Brosimum, 5, 7, 8, 10, 11, 12, 14, 15, 17,21, 144,231 subg. Brosimum, 231 subg. Ferolia, 5, 7, 18, 144, 231 acutifolium, 233 subsp. acutifolium, 234 subsp. interectum, 234 subsp. obovatum, 234 alicastrum,231, 232

subsp.alicastrum,8, 9, 12, 17, 21, 232 subsp.bolivarense, 20, 233 bolivarense, 233 costaricanum, 17, 19,22, 235 foetidum, 239 gaudichaudii, 16, 17, 19,236 glaucum,16, 19,22, 236 glaziovii,16,236 guianense,17,20, 235 lactescens,14, 17,20, 234 longifolium,18,239 237 melanopotamicum, multinervium, 231,239, 240*,310* parinarioides, 17,241 subsp.amplicoma,241 241 subsp.parinarioides, potabile,18,241 ramonense, 250 231 [refractum], rubescens,17,20, 237 utile,20, 237 subsp.allenii,17,238 subsp.darienense,17,238 subsp.longifolium, 231, 239 subsp.magdalenense, 17,238 subsp.occidentale,17,238 subsp.ovatifolium,238 subsp.utile,237 Broussonetia, 5, 12, 15,25, 53 brasiliensis, 53, 54

grevei,25 69 papyrifera, plumeri, 58 tinctoria, 56

58 [xanthoxylon], zanthoxylon,58 Broussonetieae, 4 Bucephalon, 32, 33, 34 mexicanum,32,34,40 racemosum,32 Calpidochlamys,33 drupacea, 33

4 Cannabineae, 4 Cannaboideae, Cardiogyne,53 africana, 53, 54

Castilla,7, 9, 10, 269, 270 elastica,5, 21, 269 subsp.costaricana,17,269

subsp.elastica,17,269 subsp.gummifera,17,269 tunu,8, 17,269 ulei,21, 269 f. lecithogalacta, 261 Castilleae,7, 8, 9, 14, 15, 16, 17, 20, 68,244 Cecropia,4, 6, 10 polyphlebia,38 Cecropiaceae, 4, 8, 10, 15, 16 Celtideae, 4

Celtis,7 Celtoideae,15 Cephalotrophis,32 javanica, 32 Chlorophora,53 brasiliensis, 54 mollis, 59 mora, 66 reticulata, 67 scandens, 54 tinctoria, 53, 56

var.[acuminatissimaj, 59 f. [glabrescens], 59 subsp. mora, 66 var.mora, 66 var.ovata, 59

58 subsp.[xanthoxylon], subsp. zanthoxylon, 58 var.affinis, 59 var.zanthoxylon, 58

var.[xanthoxylon], 59 f. polyneura, 59

f. miqueliana, 59 f. tataiiba, 59 Clarisia,6, 7, 8, 9, 12, 67, 72, 79 88 sect.Acanthinophyllum, sect.Clarisia,80 biflora,12,17,20,71,79,80,81*,303* subsp. mexicana, 80 colombiana, 80

ilicifolia,11, 17,20, 79, 80, 88, 89*, 100, 138,303* var.cuyunensis, 90 var.guianensis, 90 var.micrantha, 90 var.paraensis, 90 mattogrossensis, 80 mexicana, 80 mollis, 92, 105 nitida, 84

343

NAMES INDEXOFSCIENTIFIC panamensis, 80,84

racemosa,8, 14, 15,20, 71, 79, 80, 84, 85*, 138,304* spruceana, 90 strepitans, 88 var.cuyunensis, 90 var.guianensis, 90 var.micrantha, 90 var.paraensis, 90 urophylla,40

4 Conocephaleae, 4 Conocephaloideae, 4 Conocephalus, Coussapoa,4,16 rekoi, 138 Craterogyne,144 Cudrania, 24, 53 javanensis, 53

Dorstenia, 4, 5, 6, 8, 9, 11,12,14,15,16, 17,20,21,22,143,144,145,148* sect. Acauloma, 152 sect. Bazzemia, 152, 208

sect.Dorstenia,152,185 series Acaules, 186 series Caulescentes, 156 sect.Emygdioa,145,151,152,153,208 sect. Eudorstenia, 152

sect.Kosaria,151, 152 sect. Lecanium,15, 144, 151, 152, 156, 187 152 sect.Lomatophora, sect.Nothodorstenia, 144, 152, 157 sect.Sychinium, 144,151, 185

152 sect.Xylodorstenia, series Furcatae, 185 series Quadratae, 186 series Scapigerae, 185

[aculeata],329 albertii,188, 189*,307* 188 [albertorum], alexiteria, 198 amazonica, 214

annua,150 anthuriifolia, 187

appendiculata,146, 147, 152, 157, 158*,187,307* argentata, 149,150, 157 arifolia,146,150,151,152,187, 188, 191,192*, 193*,307* var.pinnatifida, 191 aristeguietae, 19, 175, 177*, 307* asaroides, 210, 212 var.celiae, 212 asperifolia, 226

var.palustris,214 var.tomentosa,214 var.tubicina,214 f. balansae,214 22,151,161,162*,307* brevipetiolata, bryoniifolia, 211 f. minor, 211

caimitensis,226, 227*,309* 198 capricorniana, caulescensLinnaeus,230 caulescensVellozo,183 var.angustifolia,183 var.latifolia,183 carautae,165, 166*,307* cayapia,9, 14, 16, 17, 19, 146, 147, 150, 151, 153,208,210,214 16,149,150,151, subsp.asaroides, 209*,210,211,212,213, 308* subsp.cayapia,16,17, 150,209*, 210,211,308* 209*,210, subsp.paraguariensis, 211, 212,308* subsp.vitifolia,209*, 210, 211, 212,213,308* var.asaroides, 212 211 var.bryoniifolia, var.[opifera],229 f. paraguariensis, 21 1

subformalobata,211 ceratosanthes, 151, 197

224*,226,228, 309* erythrandra, [exaltata],229 151,153,218,219*,309* excentrica, faria, 191

fawcettii,153,214, 218, 221*,222, 309* ficifolia, 191 ficus, 197 fischeri, 191, 194

flagellifera,221*,222, 223, 309* fluminensis, 197

229 [fruticulosa], [glabra],229 [glauca],230 gracilis, 165

146,147,187,188,189,307* grazielae, haitensis, 220 heringeri, 214, 217, 218

hildegardis,152,185, 186*,307* hirta,149,150,159,160*, 161,162, 307* [hirta],230 hispida, 159 f. minor, 159 hotteana, 222 houstonii, 198 howardii, 226

230 [hydrocotyle],

var.trilobata, 197

infundibuliformis,214

f. integrifolia,197 choconiana,157, 168, 170*, 172, 175, 179,307* var.integrifolia,168 colombiana, 17,146,178,179*,307* conceptionis,151, 153,208, 307* confusa, 226

contensis,162, 164*,309* contrajerva, 16, 17, 19,20, 144,146, 147, 149, 150, 152, 153, 155, 185,186,187,194,198,199*, 203, 207,309* subsp. tenuiloba, 198 var.houstonii, 145, 198 var.maculata, 198 var.tenuiloba, 198 cordato-acuminata,168

[cordifolia],145,214, 229 crassipes, 225

crenulata, 227*,228, 309* crispata, 203, 208 cyperus, 191

djettii,147

bahiensis, 146, 149, 150, 187, 307* barnimiana,146 belizensis, 146, 151, 153, 219*, 223, 309*

dolichocaula, 197

[bifida],229

var.varoniifolia, 159 f. minor, 159

var.riedeliana,197

astyanactis,149

boliviana, 175 bonijesu, 5, 189, 190*, 307* bowmaniana,163, 165, 307* brasiliensis, 17, 19, 145, 146, 150, 151, 153, 208, 214,218, 308* var.guaranitica, 214 var.major, 214

var.hispida, 159

var.[maculata],161

var.dissidens,198 226 domingensis, 152, 157 dorstenioides, drakena,145, 149, 150, 151, 152, 153,187, 191,194,203,204*, 205*,207, 307* ekmanii,223

elata,146, 149, 150, 152, 157, 162, 182, 183*, 185,309* embergeri,150 erecta, 159, 161

jamaicensis,220,221*, 222,309* lagoensis, 163, 165

[lanei],229 langsdorffiana, 183

letestui,146 151,153,179,181*, 182, lindeniana, 307* longifolia, 182

[longipes],230 230 [macahensis], maculata, 198 mandiocana, 191 marginata, 218 maris, 191, 194 martiana, 187 mexicana, 203

milaneziana,165, 167*,168,307* minor, 159 montana, 214 montevidensis,214 morifolia, 211

151, 191, 197 [multiformis], var.arifolia, 191 var.ceratosanthes, 197 var.ficifolia, 191 var.pinnatifida, 191 var.ramosa, 194 multisquamae,226 nervosa, 156, 183 var.angustifolia, 183 var. latifolia, 183 f. angustifolia, 183 f. latifolia, 183 nipensis, 226

nummularia, 223, 224*,309*

344

FLORANEOTROPICA

230 [ophidiana], [opifera],229 ovalis,203 210 pachecoleoneana, palmata,198 230 [palmata], panamensis,17,149,150,172,178, 180*,307* 230 [paradoxa], 211 paraguariensis, peltata, 191,223, 224*,225,309* 214 pernambucana, 151,175,176*,178,307* peruviana, petraea,220,221*, 222,307* picta,146, 152, 156 230 [placentoides], plumeriifolia,182 pseudo-opifera, 210

198 quadrangularis, var.integrifolia,198 var.pinnatifida,198 var.sinuata,198 quadrata,191 radiata,151 ramosa,144,146,147,149,150,151, 187,194, 195*, 196* subsp.dolichocaula,197,307* subsp.ramosa,152, 197, 195*, 196*,307* 230 [reniformis], riedeliana, 191

rocana,227*,228,309* roigii,222, 223, 224*, 225, 309* 230 [rotundifolia], sabanensis,214 schulzii,214,217,218 setosa,168, 169*,307* strangii,191 subdentata, 152, 156 sucrei,182 sychinium,194 tentaculata,191 tenuis,19, 149, 150, 153,213, 308* tomentosa,214 tricolor,229 tuberosa,227*,228,309* tubicina,145,214 var.[opifera],214 214 f. subexcentrica, f. major,214 tumerifolia,145,146,147,149,150, 152, 156, 157, 161, 163,307* umbricola, 151,172,174*,175,307* urceolata,146, 149, 150, 152, 156, 157,183, 184*,307* var.angustifolia,183, 185 var.latifolia,183 var.variegata,150, 183 uxpanapana,172,173*,307*

fagifolia, 32 Euartocarpeae, 4

Excoecaria,79 ilicifolia, 79, 88

Fatoua,4, 8, 9, 11, 14,25 Fatoueae,4 Ficeae,4, 9, 15,20, 21, 68, 289 Ficus,4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 20, 21, 22,25,289 sect.Adenosperma, 290 sect.Ficus,290 sect.Kalosyce,5, 290 subsect.Macrostyla,11, 15 sect.Neomorphe, 290 subsect.Palaeomorphe, 5 subg.Pharmacosycea, 16,290,292 sect.Oreosycea,290 sect.Pharmacosycea, 5, 6, 8, 13, 290,292 sect.Rhizocladus,5,290 sect.Sinosycidium, 290 sect.Sycidium,5, 8, 290 sect. Sycocarpus,290 subg.Sycomorus,290 subg.Urostigma,5, 290,291 sect.Galoglychia,15 sect.Americana, 6, 13,291 19 adhatodifolia, 17 albert-smithii, altissima,291 amazonica,291 americana,17, 19,20, 291 andicola,291 18 aripuanensis, aurea,17 berteroi,17 brittonii,16, 19,291 broadwayi,18,20 caballina,17 19 calyptroceras, carchiana,19 carica,13 19 carautana, tinctoria,56 casapiensis,20 20 zanthoxylon, 58 castellviana, christianii,18 Guatteria citrifolia,17, 19,291 cuspidata, 133 coerulescens,18 colubrinae,17 Helianthostylis, 8, 10, 12, 18, 144, 243 crassiuscula,16, 19 crocata,17,20 guggenheimii, 243 salzedoi, 243 cuatrecasana, 16, 19 20 dendrocida, schultesii, 243 20 donnell-smithii, sprucei, 18, 20, 243, 310* steyermarkii, 19, 244, 310* dugandii,291 eliadis,16, 19 suerpo, 243 enormis,16 Helicostylis, 7, 9, 10, 12, 266 eximia,291 bolivarensis, 233

varroniifolia, 159 velloziana, 197

faginea, 232

[vilella],229 [vitella],229

gameleira, 18

vitifolia, 210, 21 1 [volubilis], 230

zambesiaca,153 Dorstenieae,4, 9, 14, 15, 16, 17, 20, 68,143,149,231

32 Dumartroya,

killipii, 18 lauretana, 18, 19 leiophylla, 18 longifolia, 16 macbridei, 20 malacocarpa, 18 mariae, 16 maroma, 19 maroniensis, 18 mathewsii maxima, 20 membranacea,20 microcarpa,291 mollicula, 19 mutisii, 19 nymphaeifolia, 5, 19, 20 oblanceolata, 283 obtusifolia, 20 obtusiuscula, 19 organensis, 16, panurensis, 18 paraensis, 17, 20 pertusa, 17, 20 petiolaris, 16, 19 piresiana,20 pulchella, 20 rieberiana, 19 roraimensis, 18, 19 schippii, 5, 20 schultesii, 5, 18 subandina,291 subapiculata,291 tepuiensis, 16, 18 tequendamae, 19 tovarensis, 19 trigona, 17, 20 trigonata, 17 turrialbana,17 velutina, 16, 19 yoponensis, 20 ypsilophlebia, 20 Fusticus, 53, 56 [tataiba],59 [vera], 58

gigantosyce,19 gomelleira,5 greiffiana,291 guianensis,17, 19,20 hirsuta,16 insipida,17, 19,20, 21 jacobii,19 katherinae,283

elegans, 267 heterotricha,268 pedunculata, 17, 18, 20, 268 scabra,267 tomentosa, 5, 17, 20, 266 tovarensis, 16, 19, 268 turbinata,268 Hullettia, 67, 68 Inophloeum, 138

INDEXOFSCIENTIFIC NAMES armatum,138 Ioxilon,53 53 pomiferum, var.glaberrimum, 54 mora,66 Kosaria foetida, 145 radiata,151 Laurea,68 tiliufolia,68, 69 Lozanella, 8 Maclura, 5, 6, 7, 11, 20, 24, 25, 53 sect. Cardiogyne, 53, 54 sect. Chlorophora,54, 56 sect. Cudrania,53 sect. Maclura,53 sect. Plecospermum,54

affinis,59 afiicana, 53, 54

aurantiaca,53 brasiliensis, 20, 53, 54, 55*, 301*

59 chlorocarpa, cochinchinensis, 53

mora,53 plumeri,58 polyneura,59 pomifera, 25, 53, 54

59 sempervirens, sieberi,59 spinosa, 53

59 subintegerrima, tinctoria,16, 19, 20, 21, 22, 53, 54,56 subsp. tinctoria, 57*, 58,302* var.affinis, 59

var.chlorocarpa,59 subvar.sinuata,59 subsp.mora, 16, 19, 57*, 66, 302*

var.ovata,59 var.polyneura,59 var.subcuneata,59 var.subintegerrima, 59 subvar.quercina,59 var.zanthoxylon, 59 subvar.lobata,59 f.polyneura,59 triloba,66 velutina,59 58 zanthoxylon, Maillardia,33 borbonica,33 Malaisia,32,33 tortuosa,32 Maquira,7, 9, 12, 21, 263 calophylla, 16, 265 coriacea, 5, 16, 265

costaricana,263 [grandis],265 guianensis, 20,263 5,16,20,264,265 subsp.costaricana, subsp. guianensis, 263 sclerophylla, 265 Mesogyne, 15, 244 Milicia, 15, 25, 53 excelsa, 16 Monimiaceae, 10

345 Moraceae, 4,8,10,11,14,15,16,17,18,20 ulei, 20, 278, 282 Moreae,4, 7, 8, 9, 14, 15,17,20, 24,68 subsp. amara,283,287 subsp. puberula, 17, 284, 311 * 4 Moroideae, subsp. subandina,270, 283, 3 1 * Morus,6, 15, 16,25 subsp. ulei, 282 subg.Afromorus, 26 velutina, 272, 3 11* subg.Eumorus, 26 Noyera subg.Gomphomorus, 25,26 glabrifolia,263 alba,21, 26 rubra,262 albida,26 arbuscula,26 Ogcodeia australis,26 oblongifolia,275 betulifolia,26 Olmedia,32, 33, 138,246 canina,26 armata,138 celtidifolia,17, 19,22, 26, 27*,300* aspera,32 confinis,26 caucana,47 corylifolia,26 fa1cifolia,47 26 crataegifolia, poeppigiana,47 exelsa,53 virgata,133 goldmanii,26 40 urophylla, grisea,26 insignis,17, 19,25,26,29,30*,300* Olmedieae,4,231 Olmediopsis, 246 lactea,26 obliqua,246, 247 26 macroura, marmolii,29 Paraclarisia,92, 93, 131 mexicana,26 amazonica,92,131, 136 mesozygia,25, 26 Parartocarpus,67, 68 26 microphilyra, Perebea, 6, 7, 9, 21 microphylla, 26 sect. Noyera, 7. 253, 260 mollis,26 angustifolia, 5, 20, 258, 260, 310* nigra,25 glabrifolia,260,263 pandurata,26 guianensis, 20, 254 peruviana,29 subsp. acanthogyne, 255, 256 58 [plumeria], subsp.casfilloides,16, 17,254,256 radulina,26 subsp. guianensis, 254, 255 tinctoria,32, 53 subsp. hirsuta,255 trianae,27, 29 subsp. pseudopeltata,256 vernonii,26 humilis, 19,259 vitifolia,26 261 lecithogalacta, zanthoxylon, 32,58 longepedunculata,253,257,258,259 Musanga,4 mennegae, 19, 253, 259,310* 4 Myrianthus, mollis, 261 4 Myricaceae, Naucleopsis,7, 10, 12, 17,20, 22, 269 amara, 270, 278

caloneura, 272,274 capirensis,17,278,279* chiguila,17, 19,270, 278, 311* concinna,275,276 glabra,20, 273,274 guianensis,18,275 herrerensis, 276, 277*,311* humilis,19,285, 286*,311* imitans,274 inaequalis,16,276 jamariensis,18,22, 273 krukovii,286 272 macrophylla, mello-barretoi, 270,275 naga,16, 17,280 subsp.meridionalis, 270,281,311* subsp.naga,270,281 oblongifolia,17,20, 270, 275 pseudonaga, 274,287 riparia,272 stipularis,276 straminea,17,287,288*, 311* temstroemiiflora, 274,275, 276

subsp. lecithogalacta, 261 subsp. mollis, 260, 261, 31 1* subsp. rubra, 260 rubra,262 subsp. glabrifolia, 263, 311 * subsp. rubra,18, 262, 311* tessmannii, 253, 256 xanthochyma, 20, 257

290 Pharmacosycea, Pilea caulescens, 230 Piper

tiliifolium,68, 69 Plecospernum,53 spinosum,53 Poikilospermum,4 Poulsenia, 6, 67, 138

aculeata,138 arnata,10, 17,20,22,138, 139*,306* spec., 142, 306* Pourouma,4, 6, 14, 16 Prainea,67, 68 Procris

230 quinquenervia, Pseudolmedia, 6, 7, 9, 10, 12, 22, 246, 270

346

FLORANEOTROPICA

sect.Acomandra, 246 sect.Olmediopsis,246 sect.Pseudolmedia, 246,253 ferruginea,246 gentryi,16, 19,246, 252 glabrata,17,5 1, 246, 250 guaranitica,246,249 havanensis,246 hirtula,16,246,248 karstenii,40 laevigata,16, 17,20, 21, 246, 249, 311* laevis,12, 14, 17,21, 246, 247 macrophylla,16, 17, 21, 246, 248, 253, 310* malacocarpa,250 manabiensis, 17,22,246, 250,310* mollis,250 murure,246, 248 oxyphyllaria, 51, 246,250 rigida,16,20, 246, 251, 253 subsp.araguensis,19,252 subsp.eggersii,16, 19,20, 251 subsp.rigida,19,20, 251 simiarum,250 spuria,17,252 Pseudosorocea,92, 93 bonplandii,92, 100 poeppigii,88 sprucei,131 uaupensis,117 Sahagunia,79,80 colombiana,80 mexicana,79, 80 racemifera,88 strepitans,88 40 urophylla, [Sarcodiscus]

136 [amazonicus], Scyphosyce,144, 147 Sitodium altile, 143 Skutchia,33, 34 caudata,33,34 Soaresia,79, 80 nitida,79, 84, 138 Sorocea,6, 8, 10, 12, 17,67, 92, 93 subg.Sorocea,94, 96 subg.Paraclarisia, 94, 133 affinis,17,93, 123, 124*,125, 129, 305*

amazonica,114 arnoldoi,133 bonplandii, 16, 19,92, 93, 94, 100,

101*,304* briquetii,93, 118, 119*,305* castaneifolia,98 colombiana,40, 138 cufodontisii,105 dentata,114 duckei, 16, 92, 93, 94, 131, 136,

137*,306* faustiana,107 105 [graciliflora],

laurifolium, 53 grandifolia,98 guayanensis,117 Sychinium, 144, 151, 185 guilleminiana, 16, 17,20, 21,51, 92, ramosum,144,151, 185, 194 93, 94, 97*, 98, 100, 304*

hilarii,16,93,94,100,103,104*,304* Treculia, 7, 9, 12, 68 var.[graciliflora], 105 africana, 10 hilariana,98 Trilepisium, 7, 12, 144 hirtella, 110, 113

subsp.oligotricha,107 houlletiana,98 [ilicifolia],138 var.grandiflora,138 var.hilarii,103 f. laxiflora,100 jaramilloi, 17,19,94,129,130*,306* jureiana,98 98 klotzschiana, macrogyna, 98 103 macrophylla, martineziana, 120 mexicana,34, 138 muriculata, 17,93, 113,114, 120 subsp.muriculata,16, 21, 114, 115*, 305*

subsp.uaupensis,18, 115*,117, 305* nitida,138 opima,110 pileata,118 pubivena,20, 94, 105, 107, 113 subsp.hirtella,16,21, 108*,110, 305* subsp.oligotricha,16, 21, 107, 108*,113,305* subsp.pubivena,17,53, 92, 105, 106*,305* racemosa, 103 rhodorachis, 120

Trophis, 7, 9, 12, 24, 25, 32, 33, 246 sect. Calpidochlamys, 34 sect. Echinocarpa,34, 51 sect. Maillardia,34 sect. Malaisia, 34 sect. Olmedia, 34, 47 sect. Trophis, 34, 34

americana,32,40 var.meridionalis, 40 var.ramon, 40 aurantiaca, 47 borbonica, 33 caucana, 12, 20, 24, 32, 33, 47,48*, 301 *

chiapensis,34, 38 34 chlorizantha, cuspidata, 17, 38, 39*, 301* drupacea,33 glabrata, 51, 246,250

hilariana,51,98 involucrata,11, 24,33,51,52*, 301*

laurifolia,53 macrostachya, 53, 105 matudai,34 mexicana, 17, 20, 33, 34, 35*, 40,

138,300* nubium, 34 racemosa,17, 20,32,38,40,41*, 138 subsp. racemosa, 46

subsp.ramon,40,46 subsp. meridionalis, 40,46 ramon, 40 scandens,32 Trophisomia,92, 93, 131

ruminata,17, 19, 93, 94, 125, 127, 128*,306* sarcocarpa,17, 19, 94, 125, 126*, 305* edulis,92, 134 saxicola,134 Trymatococcus, 12, 18, 144, 231, 241 var.dentata,134 amazonicus, 14, 17,18, 241, 3 10* f. subrepanda,134 266 guanabarinus, sprucei,16, 19, 93, 94, 131 oligandrus, 243 subsp.saxicola,19,92, 132*,134, paraensis,241,242, 243 135,306* turbinatus,243 subsp.subumbellata,19, 132*, 135,306* subsp.sprucei, 132*, 133, 135, Uleodendron,269 amarum,284 306* ulei, 282 steinbachii,16,111, 112*,113,120, Ulmaceae, 4, 7, 15, 25 305*

Urera, 4, 138 Urostigma,290 sect. Americana, 291 trophoides, 16,17, 19,94,120, 121*, fagineum, 232 Urticaceae, 4, 7, 8, 10, 15, 20, 25, 138 269, 305* Urticales, 4, 10 subsp.rhodorachis,120 Utsetela, 144 uriamen,103

stenophylla, 90, 138 sylvicola, 100 var.caaguazuensis, 100

Sparattosyce, 9, 14,67, 68 Strebleae,4 Streblus,9, 11, 14,20, 25, 33 Styloceras

neglecta,144 Vaniera,53 cochinchinensis, 53