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 an...

Author: Cornelis C. Berg

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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

Issued 31 October 2001

C 2001 by TheNewYorkBotanicalGarden

All rightsreserved. Publishedby

TheNewYorkBotanicalGarden Bronx,NY 10458

InternationalStandardSerial Number0071-5794

of Thepaperusedin thispublication meetstherequirements forInformation Sciences- Permanence of Paperfor theAmericanNationalStandard Publications andDocumentsin LibrariesandArchives,ANSI/NISO(Z39.48-1992).

Printedin theUnitedStatesof Americausingsoy-basedinkon recycledpaper. is a leadershipfunderof TheNewYorkBotanicalGardenPress. LifeFoundation Metropolitan

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

581 .98'012-dcl9

85-647083

[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-

2

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

FLORANEOTROPICA

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


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.


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.


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,


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


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


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 oblong 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)

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