Confronting a morphological nightmare: revision of the Neotropical genus Guatteria (Annonaceae)
Abstract
Maas, P.J.M., Westra, L.Y.T., Guerrero, S. Arias, Lobão, A.Q., Scharf, U., Zamora, N.A., Erkens, R.H.J. (2015): Confronting a morphological nightmare: revision of the Neotropical genus Guatteria (Annonaceae). Blumea 60 (1): 1-219, DOI: 10.3767/000651915X690341, URL: https://doi.org/10.3767/000651915x690341
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Blumea 60, 2015: 1–219 www.ingentaconnect.com/content/nhn/blumea http://dx.doi.org/10.3767/000651915X690341 RESEARCH ARTICLE INTRODUCTION Guatteria is among the oldest still accepted genera of AnnonaceaeintheNewWorld.Annona waspublishedin1753, followed by Xylopia in1759, Asiminain1763,andPorcelia and Guatteriain1794(Maasetal.2011).Inthe220yearsofitsexistenceover425specieshavebeendescribedinthegenuswith someyearsagoeven307acceptednamespresent(Erkenset al.2008).Atthattimemanynewspecieswereforeseendueto thelargeamountsofunidentifiedmaterialinherbariaworldwide. Indeed,sincetheneightnewspecieshavebeendescribed (Lobãoetal.2010,Maas&Westra2010,2011,AriasG.etal. 2014,Lobão&Lopes2014),andsome25morearedescribed inthispublication.However,opposingtheincreaseofspecies standsadecreaseduetosynonymization.Forinstance,afew yearsago40namesweresynonymizedintoGuatteria australis, leading to a large reduction in species diversity in the genus (Lobãoetal.2011).Moresuchreductionstakeplaceinthis publication.Forinstance,34namesarenewlysynonymized under G. punctata (bringingthetotalnumberofsynonymsto 46)includingthetypespeciesofthegenus,G. glauca (fora furtherdiscussionaboutconservationandlectotypificationof GuatteriaseeVanSetten&Maas1990).Thecircumscription of G. punctata has always been problematic, but other species previously considered to be clearly circumscribed also disappear, such as for instance G. recurvisepala and G. boliviana that are now brought into synonymy with G. ucayalina (plussix morenewsynonyms). Thispatternisperhapsreminiscentofearlierdaysinthetaxonomic history of Guatteria (moreelaboratelydescribedby Erkensetal.2008).AfterthedescriptionofthegenusbyRuiz &Pavón(1794)thenumberofspeciessteadilyincreased viaforinstancetheworksofDunal(1817),DeSaint-Hilaire (1824–1825),VonSchlechtendahl(1834),VonMartius(1841), Triana&Planchon(1862),Diels(1905,1906,1924a,b,1927, 1931)andFries(1938).Atthatpointintime‘Sweden’sgrand oldmanofbotany’(Buchwald1970),RobertFries,hadtaken a keen interest in the 151 species of Guatteria described and heworkedontheonlyrevisionbeforethecurrentone.He synonymized and transferred species but also described 106 newones,andorganizedthesespeciesinto30sections(Fries 1939).Laterheerectedtwosubgenera(Fries1955a)andhe reducedthenumberofsectionsto22(Fries1959b).Afterhis revision, Fries continued to be the main describer of new speciesinthegenus(fordetailsagainseeErkensetal.2008). THE PIONEER OF ANNONACEAE SYSTEMATICS KlasRobertEliasFries(11July1876–29January1966;Fig.1) was the author of the only comprehensive revision of Guatteriatodate.Nexttothisrevisionhehasleftanextensiveand impressivebotanicalrecordofaccomplishment.Inhisworkhe coveredthreecontinents(Europe,SouthAmericaandAfrica) withworksontaxonomy,phytogeographyandmycology.He Confronting a morphological nightmare: revision of the Neotropical genus Guatteria (Annonaceae) P.J.M.Maas1,L.Y.T.Westra1,S.AriasGuerrero1,A.Q.Lobão2,U.Scharf3, N.A.Zamora4,R.H.J.Erkens1,5 1NaturalisBiodiversityCenter,sectionBotany,P.O.Box9517,2300RA Leiden,TheNetherlands. 2LaboratóriodeSistemáticaeBiogeografia,SetorBotânica,Departamento deBiologiaGeral,UniversidadeFederalFluminense,RiodeJaneiro,Brazil. 3Herbarium,InstitutfürBotanik,UniversitätLeipzig,Johannisallee21–23, D-04103Leipzig,Germany. 4 InstitutoNacionaldeBiodiversidad(INBIO),ApartadoAereo22-3100,Santo DomingodeHeredia,CostaRica. 5MaastrichtScienceProgramme,MaastrichtUniversity,Maastricht,The Netherlands. Key words Annonaceae chemistry descriptions flower biology and pollination Guatteria Guatteriella Guatteriopsis Heteropetalum history molecular studies morphology Neotropics palynology phylogeny taxonomy vernacular names Abstract A taxonomic revision of the genus Guatteria, including the former genera Guatteriella, Guatteriopsis and Heteropetalumisgiven.WithinthegenusGuatteria177speciesarerecognized,25ofwhicharenew.Includedare chapters on the history of the taxonomy of the genus, morphology, wood anatomy, karyology, palynology, chemistry, flower biology and pollination, dispersal, distribution and ecology, phylogeny and molecular studies, conservation, anduses.Asynopticalkeytoallspeciesisincluded,aswellastwodichotomouskeys,oneforthespeciesofCentral AmericaandMexico,andoneforthespeciesofNE,EandSEBrazil.Thespeciestreatmentsincludedescriptions, fullsynonymy,geographicalandecologicalnotes,vernacularnamesandtaxonomicnotes.Forallspeciesdistributionmapsaremade.Acompleteidentificationlistwithallexsiccataestudied,anindextovernacularnamesandan indexofscientificnamesisincludedattheend. Published on 22 December 2015
2Blumea–Volume60/1–3,2015 has written many publications on his trips to Africa and wrote ground-breaking treatments on the east African mountain vegetation(Jonselletal.1991).Nexttothisextensiveworkhecontributed greatly to the flora of Central America, South America andtheWestIndies,especiallytotheknowledgeofthefamily of Annonaceae.Hemonographedthefamilyinhis1959contributiontoEngler&Prantl’s‘DieNatürlichenPflanzenfamilien’ (Fries1959b).Hewasalsostronglyinterestedinmycologyand madeimportantcontributionstothatfield,suchashiscollectionsfromBolivia(aregionillcollectedatthattime).Todaywe canconcludethathismostimportantscientificcontributions have dealt mainly with Annonaceae and various groups of fungi(Unknown1966). IntermsofhisprofessionalcareerRobertFrieswaspartofabotanicaltradition.Hisfather,ThoreMagnusFries(1832–1913), was professor of Botany at Uppsala University and his grandfather,EliasMagnusFries(1794–1878),hadbeenScandinavia’s mostfamousmycologist(Buchwald1970).Hiscareerstartedat the age of 25 when he was part of an expedition to the northern partofArgentinaandtheSouthofBolivia(Chardon1947)which providedthebasisforhisdoctor’sdegreeatUppsalaUniversity in1905(Unknown1966).Afterthis,heobtainedateaching position at Uppsala University and in 1915 he was appointed ProfessorBergianus(directoroftheHortusBergianus)and director of the Bergius foundation at Stockholm, a position he retaineduntilhisretirementin1944(Chardon1947,Unknown 1966).Healsoheldmanyboardpositions.From1928–1939, hewasvice-secretaryoftheAcademyofSciencesofSweden; in1939–1940,hebecamepresidentofthelatterSocietyand in1942hewasvice-president.From1934–1939hewasalso president of the Botanical Society of Stockholm and member oftheDendrologicalSociety(1927–1946),theLinneanSociety ofSweden(from1924–1947;honorarypresidentfrom1947 untilhisdeath)andtheBotanicalSocietyofSweden(from 1943–unknown;Chardon1947). THE GUATTERIA PROJECT Around2000aprojectwasstartedattheNationaalHerbarium Nederland-UtrechtBranch(nowpartofNaturalisBiodiversity Center,Leiden;Erkens&Baas2008)toinvestigateGuatteria as part of the AnnonaceaeResearchProject(Maas1984). Theprojecthadthefollowingmainaims:1)torevisethespecies of Guatteria based on a geographic division among coworkers;2)toclarifythephylogeneticpositionofGuatteria within Annonaceae;3)toelucidatetherelationshipsbetweenthe genera in the Guatteria group;4)toproduceaphylogenetictree containingasignificantnumberofspeciesofGuatteria;5)to investigatethebiogeographichistoryofthegenus;and6)to provide insight into the evolution of key morphological and anatomicalcharacters. REVISIONARY WORK TherevisionofspeciesdiversityinGuatteria based on a geographicaldivisionprovedfruitful.ThesoutheasternBrazilian speciesweremainlyinvestigatedbyLobãoforherthesisand inpublications(Lobão&Mello-Silva2007,Lobãoetal.2010, 2011,2012,Lobão&Lopes2014).Scharfetal.focusedon speciesfromtheGuianashieldarea(2005,2006a,b,2008) andmadeacontributiontotheFloraoftheGuianas(inprep.). Erkensetal.focusedmoreonCentralAmerica,theCaribbean and non-Amazonian Colombia, as can be seen from the preliminarytreatmentforFloraMesoamericanainErkens’thesis (Erkens2007)andseveralnewlydescribedspeciesfromthat area(Erkensetal.2006,AriasGuerreroetal.2014,seealso thispublication).SusanaAriasG.revisedtheWestColombianAndeanspeciesforherMastersthesis(AriasGuerrero 2011,unpubl.data).AlsoZamoraetal.havebeenworkingon especiallytheCostaRicandiversity(Zamora&Maas2000, Erkensetal.2006,seealsothispublication).Lastly,Maaset al.focusedonAmazonianSouthAmerica(Maasetal.1988, Erkensetal.2008,Maas&Westra2010,2011)butMaasalso made contributions to many of the other publications mentioned above.ForamoreelaborateoverviewofpublicationsonGuatteria see Erkensetal.(2012b). Thisrevisionistheculminationofallthiswork.Itwillserveas anewtaxonomicbaselineforfuturework,finallyreplacing Fries’1939revision.Assuchthisrevisionfitsinalongtradition of systematic work on Annonaceae(Chatrouetal.2012a). Thecurrentrevisionrecognizes177speciesofGuatteria.This means that Guatteria is no longer the largest genus within Annonaceaecontrarytoasstatedbefore(Erkensetal.2008)and cannot be seen as one of the largest genera of woody Neotropicaltreesanymore. THE PHYLOGENETIC POSITION OF GUATTERIA WITHIN ANNONACEAE InmorphologicalanalysesofAnnonaceae the position of Guatteria wasunclear(e.g.,Doyle&LeThomas1996)because many of the character states for Guatteria are homoplastic or uniquewithinthefamily.Alsotheearlymolecularphylogenetic analyses placed Guatteria ambiguously with respect to its sister clades(Bygrave2000,Richardsonetal.2004,Couvreuretal. Fig. 1PotraitofRobertE.Friesatc.28yearsofage(originalintheRegnellianHerbarium,Stockholm,Sweden).
3 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria 2008).Itwasclear,however,thatthecladetowhichGuatteria belongs contains genera with African, Asian and South Americanorigins(Doyleetal.2004,Richardsonetal.2004, Couvreuretal.2008),nowcalledsubfamilyAnnonoideae.Areconstructionspecificallyofthephylogeneticrelationshipswithin Annonoideae was undertaken in order to address the issue of the phylogenetic position of Guatteria (Erkensetal.2009). Unfortunately, in that study there was still ambiguity with respect totheplacementoffourlineages(allclassifiedastribesbyChatrouetal.2012b):Guatterieae, Duguetieae, Xylopieae, and the remainderofthespeciesofthisclade(Annoneae, Monodoreae and Uvarieae).ThepositionofGuatteria at the end of a long branchmightbecauseforthesealternativeplacements(due tolong-branchattraction;Erkensetal.2009).Subsequent, family-wide analyses have not been able to resolve this issue either(Couvreuretal.2011,Chatrouetal.2012b). Thephylogenetically(longbranch)andmorphologically(unique characterstates)isolatedpositionofGuatteria might be explainedbyitsbiogeographicalhistory.Itwasshownthatthe most likely distribution of the stem lineage of Guatteria was the African-North American region and a scenario of migration fromAfricaviaEuropeintoCentralandSouthAmericahas beenhypothesisedforthisgenus(foranelaboratediscussion onthisandotherscenariosseeErkensetal.2009).Stepwise coolingfromtheMiddleEoceneonwardsremovedfrost-intoleranttaxafrommuchofNorthAmerica.Inthisscenariointhe Middle/EarlyMiocene(between18and11Ma)theancestors of Guatteria reached Central America, just before the northern climatebecametoohostile.Inthissituationanyintermediate NorthAmericanandEuropeantaxawentextinctandaphylogeneticallyisolatedgenusremainssittingonalongbranch. Also, all taxa with intermediate morphological character states were removed and therefore it is now impossible to connect extant Guatteria to other Annonaceae genera morphologically (Erkensetal.2009). THE FORMER GUATTERIA GROUP Until a few years ago four genera were thought, on morphological grounds, to be part of the so-called Guatteriagroup(Fries 1939,1943,1959b):Guatteria, Guatteriopsis R.E.Fr.,GuatteriellaR.E.Fr.andHeteropetalum Benth.Thecloseaffinitybetween these genera was established by many authors independentlyonmorphologicalgrounds(fordetailsseeErkensetal. 2007a,Erkens&Maas2008)andviamoreformal(molecularor morphological)phylogeneticanalyses(Heetal.1999,Bygrave 2000).However,noneofthelatterstudieshadsufficienttaxon sampling or enough data to adequately address the putative monophylyofthesefourgenera.Thisquestionwasspecifically addressed in a molecular phylogenetic study and it was shown that Guatteriopsis, Guatteriella and Heteropetalum were actually nested within Guatteria (Erkensetal.2007a).Asaresult the nested genera were transferred into Guatteria (Erkens& Maas2008). PHYLOGENETIC TREE OF GUATTERIA A phylogenetic tree based on four regions of the plastid genome andcontainingc.40%ofthespeciesofGuatteria was constructed(Erkensetal.2007a,b),butinternalresolutioninthis treeprovedtobeweak(Fig.2).Therefore,acompletelynew subgenericclassificationofGuatteria is still premature because of the lack of molecular and morphological synapomorphies to definenewsections(Erkensetal.2007a,2008).Currently,an improved version of this phylogenetic tree is being generated basedonfullplastomesequences(Erkens&Brandão,inprep.). all support values ≥ BS 85%/PP 0.95 one support value ≥ BS 85%/PP 0.95 both support values ≥ BS 50%/PP 0.5 Fig. 2MostparsimonioustreeaspublishedinErkensetal.2007based on plastid matK, trnLF, rbcL and psbA-trnH sequences but updated for the taxonomypresentedinthispaper.Supportvaluecatagoriesaresummaries ofbootstrappercentages(BS)andBayesianposteriorprobabilities(PP;for moredetailsontheanalysesseetheoriginalpaper).
4Blumea–Volume60/1–3,2015 INFRAGENERIC BIOGEOGRAPHY OF GUATTERIA Theinterestingquestioniswhatcausedthelackofresolutionin the phylogenetic tree of Guatteria.Threemajorbiogeographical events in the history of the genus were reconstructed and hypothesisedtoberesponsibleforthepatternfound(Erkens etal.2007b).TheoldestextantlineagesofGuatteria can be foundinCentralAmerica(Erkens etal.2007b),exceptfor the lineage leading to G. heteropetala (Erkens etal.2007a). TheCentralAmericanlineagehasdiversifiedandgivenrise to mainly endemic Central American species, which are morphologicallyclearlydistinguishable.Itisthoughtthatasecond trans-oceanicMiocenemigrationfromCentralintoSouthAmericahastakenplace(inadditiontotheoneleadingtoG. heteropetala)beforetheclosingoftheIsthmusofPanama,followed byamajordiversificationofthelineagewithinSouthAmerica (mainlyintheAmazon).Sucharapidandrecentburstof diversificationfromthemostrecentcommonancestorofthe extantspeciescanresultinpoorlyresolvedphylogenies(Richardsonetal.2001).Thethirdimportantbiogeographicalevent was the re-migration of several South American lineages into CentralAmericaviatheclosedPanamanianlandbridge.These speciesaremorphologicallydifficulttodistinguishfromeach other,asistobeexpectedofyoungspecies.Itcan,therefore, be concluded that Guatteria is not an Amazonian centred genus sensuGentry(Gentry1982)butagenuswithampleMiocene speciationfollowingitsdispersalintoSouthAmerica. Althoughthisrapiddiversificationofthe‘SouthAmerican Clade’withinGuatteria (Erkensetal.2007b)hasledtoalarge amount of species, Guatteria itself does not constitute a radiation(Erkensetal.2012a).Guatteria is in itself one of the larger annonaceous genera, but not larger than can be explained on thebasisofstochasticity(actually,incomparisontoitsputative sistergroupsitisrathersmall).Asamatteroffact,noneofthe largestgenerainthefamilyaretheresultofradiations(except perhaps Goniothalamus).ThelargercladeswithinAnnonaceae probably did not result from intrinsic key innovations since lineage-through-timeplotsarefairlylinear.Thisisgenerally associatedwithaconstantrateofdiversification.Annonaceae asawholediversifiedatanoverallsteadyrate(possiblydue tolowextinctionrates;Couvreuretal.2011). INSIGHT INTO THE EVOLUTION OF KEY MORPHOLOGICAL AND ANATOMICAL CHARACTERS Therecentdivergenceof[somespeciesof]Guatteria, the lack ofphylogeneticstructureandtheabsenceof(key-)characters associated with speciation severely hampers character mappingontoaphylogenetictree(Erkens2007).Somecharacters havebeentried(e.g.,woodanatomicalcharacters;Erkenset al.2007a)buttheevolutionofmostcharactersisstillpoorly understood.Forinstance,theindument(coveringthewhole plant in a species as G. tomentosa)isveryeasilyrecognizable andunidentifiedspecimenscanthereforeoftenbeassigned tospeciesbasedonthis.However,itisunclearwhetherornot indumenttypesshowhomoplasy.Thissituationiscomparable to the problematic use of the impressed secondary venation as a delimiting character for the Guatteria amplifolia-complex inCentralAmerica(Erkens2007).Thereisonenoteworthy exception(Maas&Westra2011):speciesthatarecharacterizedbyleaveswithtinywarts(verruculae)onbothsurfaces (incombinationwithelongateandshort-stipitatemonocarps) belongtooneparticularcladethatalignswithFries’sect.Mecocarpus.However,thefunctionofthesewartsisstillunclearand theirsynapomorphicstatusisamatterofdebate(Erkensetal. 2007a,Maas&Westra2011). GENERAL MORPHOLOGY Habit Species of Guatteria are trees, mostly medium-sized, or less oftenshrubs.Theonlylianescentspeciesarethecommonly collected G. scandens and the rarely collected G. fractiflexa, whichcanalsobeafacultativeshrub.Incidentalfieldnotesalso mention a liana habit in G. beckii, G. flexilis and G. synsepala. Theheightofthetreesorshrubsgenerallyvariesfrom1to 40m;amongtheexceptionsareG. grandiflora up to 60 m tall and G. modestaupto55mtall.Smalltreespeciesorshrubs generallynotexceeding4minheightareG. delicatula, G. maypurensis(bothinhabitingsavannas),G. rupestris and G. stenophylla.Somespeciescanbeveryvariable,e.g.,G. arenicola withindividualsincampinaranavegetationnomorethan2–3m high,butreachingupto30minforests.MostspeciesofGuatteria are quite slender with stems < 25 cm diam, but there are quiteafewexceptionshere,too,e.g.,G. cuatrecasasii(upto 85cmdiam),G. procera(upto90cmdiam),G. modesta(up to1mdiam),andthehugeMexican/CentralAmericanspecies G. grandiflorawhichcanhavestemsofupto170cmdiam. Leaves Theleavesaresimple,entire,petiolateandestipulate.Leaf disposition, as in the vast majority of Annonaceae, is in 2 rows (distichous),particularlyobviousinlateralbranches.Petioles mostly are comparatively short, normally varying from 1–10 mminlengthand0.5–5mmdiam.Thelongestpetioleswere found in G. chrysophylla(15–30by4–6mm),G. paludosa Fig. 3Ruguloseupperleafsurface,enlarged. a.Guatteria anteridiferaScharf&Maas;b.Guatteria paludosaR.E.Fr.(a:Sabatier & Prévost 4936,U;b:H.D. Clarke 11004,U).—Scalebar=1mm. ba
5 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria (15–30by3–4mm),G. castilloi(20–35by1–2mm)and G. megalophylla(15–50by3–10mm).Theshapeofthe lamina varies from elliptic, ovate, obovate, or narrowly so, ornarrowlyoblong-elliptic,obovate-oblong,etc.Thetexture varies from chartaceous to coriaceous, the surface varying from dull to shiny above, and is smooth in the majority, but verruculosebothaboveandbelowinabout1/4ofthespecies; infewofthespecies,e.g.,G. anteridi fera, G. ayangannae and G. paludosa, the epidermal surface shows minute more or less circular sculptures when viewed with a powerful lens, indicatedbythetermscabridulous(Fig.3).Thelaminamost oftenisbetween10–30cmlong,withalengthtowidthindex varyingfrom1.5–≥7,butmostlybetween2and5;thelargestleavesupto75by30cminsizewerefoundinG. megalophylla, while in about 10 species the lamina, as far as seen, does not exceed 10 cm in length, with G. odorata at the smallest extremewithleaves5–6by1.5–2.5cminsize.Theapexof the lamina generally is acute or mostly acuminate, the acumen between5–20mmlong(withfewextremesto≥30mmlong). A fairly small number of species has obtuse or even rounded apices,e.g.,G. castilloi, G. griseifolia, G. meliodora, or emarginateine.g.G. blainii and G. emarginata.Themostextreme are probably the leaves of G. clusiifoliawithatruncateapex. Thebaseofthelaminavariesfromacute,attenuate,obtuse tocordate.Themostcommonleafbaseisacute,attenuate orobtuse.Intheattenuateleafbasetheextremebaseis sometimesslightlydecurrentasawingalongthepetiole.Acordateleafbaseisencounteredinlessthan10species(e.g., G. candolleana, G. friesiana and G. pudica).Stem-clasping leavesareextremelyrareandseeninonlythreespecies(G. campinensis, G. longicuspis and G. tomentosa),butnotethatin these three species the base can vary from stem-clasping to notstem-clasping.Moregenerally,variationinshapeofthe leafbasewithinthesamespeciesisnotuncommon,e.g.,in G. amplifolia the base varies from obtuse, acute to cordate and in G. hirsutafromacute,obtusetorarelyattenuateorcordate. Inc.20speciesthebasalmarginsarerevolute,onespecies (G. revoluta)isevennamedafterthatfeature.Theprimaryvein isimpressedaboveinalmost50%ofthespecies.Inalittleover 10%ofthespeciestheprimaryveinisflattoimpressed,while inlessthan10%theprimaryveinisflat.Aflattoraisedprimary veinisfoundinalittlelessthan10%ofthespecies.Species withaprimaryveinraisedallalongarequiteexceptional(G.alticola, G. beniensis, G. crassivenia, G. gentryi and G. pachycarpa).Thevenationisbrochidodromous,i.e.secondaryveins joinedtogetherinaseriesofarches(loop-forming),mostly intheupper1/3to2/3,andmostlyeucamptodromousinthe basal part, with secondary veins gradually diminishing inside the margin and connected to superadjacent ones by a series ofcrossveinswithoutformingloops(Metcalfe&Chalk1979: 30).Inbrochidodromousvenationpatternstheloopscanbe relativelythickandbecomelined-upina(moreorlessundulating)marginalveininanumberofspecies(e.g.,G. guianensis, G. venosa).Thedistancebetweenloopsandmargininmost speciesvariesbetween1–5mm,to7mminfewspecies.A notable exception is G. sessilicarpa with smallest distance betweenloopsandmargin5–10mm.Thenumberofsecondary veins varies widely within the genus, ranging from only 5 to as manyas35oneachsideoftheprimaryvein,butmostspecies havingbetween10–20pairs.Specieswiththelowestnumber are G. acrantha and G. partangensis, both with 6–9 pairs of secondary veins, and G. eleganswith5–8pairs.Thehighest numbers occur in G. guianensis, G. insculpta(bothwith20–35 pairs),G. megalophylla(15–35pairs)andG. venosa(25–35 pairs).Thesecondaryveinsaredistinctincloseto85%of thespecies,andindistinct(hardlyvisible,sometimesdifficult tocount)intheremainder.Inroughly2/3ofthespeciesthe secondaryveinsareraisedontheupper(=adaxial)side,while in the remainder the secondary veins are flat to/or impressed above,infewspeciesstronglyimpressed(e.g.,G. megalophylla, G. punctatap.p.,G. venosa). Theangleofsecondaryveinswiththeprimaryveinranges between 60° and 90°, exceptions are G. elegans(30–40°)and G. campestris(40–55°). Thetertiaryvenationisconspicuousinalmostallspecies.Itis hardly visible or even seemingly absent in a handful of species, namely G. conspicua, G. decurrens, G. emarginata, G. friesiana and G. meliodora.Thetertiaryveinsareraisedontheupper sideofthelaminain3/4ofthespecies,intheremaining1/4 theyareflattoimpressed.Tertiaryvenationismostlyreticulate, butinlessthan1/5ofthespeciesitispercurrent(Metcalfe& Chalk1979).Agoodexampleofpercurrenttertiaryvenation is G. ucayalina. Indument.MostspeciesofGuatteria possess hairs on their leaves,youngbranchletsandfloweringparts(particularly sepalsandpetals),whereascompletelyglabrousspecies arerare.Theindumentiscomposedofsimplehairsonly.Ina few species, however, the hairs tend to be organized in tufts, almostsuggestingstellatehairs(e.g.,G. stipitata).Forspecific recognition it is important to know if the hairs are appressed or erect.Thisfeaturegenerallyisquitereliable,althoughthereare, unfortunately, some species where both erect and appressed hairsoccur. Inflorescences Annonaceaeshowavarietyofinflorescences.Fries(1919)was thefirsttopublishalargearticleonthesubject.Inthispaperhe investigated25genera(asrecognizedatthattime),mostlyfrom materialinspiritfromthebotanicalmuseuminUppsala(UPS) andtheBotaniskaMuséetinStockholm(S).Laterimportant papersoninflorescencestructurearebyFries(1959b:13–20) andWeberling&Hoppe(1996).Flowersinmostspeciesof Guatteria appear in the axils of leaves, and/or in the axils of fallenleaves.Inlittlelessthanhalfofthespeciestheflowers aresolitaryintheaxilswhilebetween35and40%have1–2 flowersintheaxils,andinaminority(about15%)leavescan alsoharbourmorethan2flowers.Thepedicelsvarygreatly within the genus, ranging from less than 5 mm to over 10 cm in length–butseealsonoteinthelastparagraphofthissection!In mostspeciesthepedicelsarebetween10–40mmlong,while inabout10%ofthespeciesthepedicellengthis10mmor less(e.g.,G. blepharophyllap.p.,G. campinensis, G. friesiana, G. schomburgkiana).Pedicelsover40mmlongarefoundin roughly15%ofthespecies(e.g.,G. alta, G. darienensis, G. dolichopoda, G. novogranatensis, G. pittieri),withthelongest exhibitedbyG.grandipes(to90mm)andG. macropus(to120 mm).Inanumberofspeciesthelengthincreasestoagreateror lesserextentbetweenfloweringandfruitingstage.Thereisalso variationindiameterfromc.1mminmanyspeciestoc.2mm inothers,andincreasingduringfruitingstageto5mm(oreven more).Thepedicel,orflowerstalk,isarticulatemostlyinthe basalpart.ThearticulationinGuatteria essentially is oblique, not transversal as in most Annonaceae(carefulexaminationis oftennecessary!).ThepedicelconformstoFries’stype1(Fries 1959b)inthatithasanumberofbractsinthepartbelowthe articulation with the two uppermost bracts placed immediately belowthearticulation,butnobractsabovethearticulation. Thereareusuallybetween5–8bracts,andtheyrepresent leavesreducedtosmallscales.Thelowermostbractisplaced on the adaxial side, and in fact is the addorsed prophyll already notedbyFries(1911).Thetwouppermostbractsareusuallythe largest, placed at the lower and at the upper end of the oblique articulation, and they protect the developing flower bud in the veryearlystage(Fig.4c).Bractsnormallyfallsoon,andtheir presence can only be inferred from the number of scars that
6Blumea–Volume60/1–3,2015 Fig. 4Inflorescencevariation.a.Guatteria terminalisR.E.Fr.:Fruitterminatingalateralbranchlet;b.idem,flowerterminalonasmalllateralbranchlet; c.Guatteria flabellataErkens&Maas:youngaxillaryinflorescencewithbractsbeforeshedding,thetwouppermostbractsenclosingtheflowerbud;d.Guatteria modestaDiels:axillaryinflorescencesonleaflessbranchletformedinpartbyaccessorybuds;e.Guatteria duodecimaMaas&Westra:axillaryinflorescences,partlysimple,partlywithonelateralflowerfromaxilofbract;f.Guatteria ayangannaeScharf&Maas:inflorescencewithtwofoliaceousupperbracts; g.Guatteria flagellifloraMaas&Westra:much-branchedinflorescence;h.Guatteria scalarinerviaD.R.Simpson:inflorescenceonsmalltrunkorlargebranch clearlyshowingbranchingpatterns;i.Guatteria ferrugineaA.St.-Hil.:oldinflorescenceontrunk;j.Guatteria notabilisMello-Silva&Pirani:terminal2-flowered inflorescencewithall(partlyfoliaceous)bractsinplace;k.same,3-floweredrhipidiateinflorescence(a,b:Farfán et al. 995,WAG;c:Croat 19183,WAG; d: Mutis 4486,US;e:Chatrou et al. 25,U;f:H.D. Clarke et el. 9819,U;g:Vásquez et al. 22500,WAG;h:Gentry et al.77361,U;i:Amorim et al. 818;j:Irwin et al. 22671,U;k:Hatschbach 64801, U). c b d a g h ij k e f
7 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria remainforashorterorlongertime.Amongthefewexceptions is G. ayangannae,anendemicofGuyana,wherethetwoupper bracts are foliaceous and mostly persist until flowering time (Fig.4f).Incidentallyabractenvelopedintoa(small)leafmay befoundatrandominmanyspecies. InflorescencesinAnnonaceaeareofthedeterminatetype(also termedmonotelicoranthotelic),i.e.,withtheaxisalwaysendinginaterminalflower(seealsoBriggs&Johnson1979and Weberling1981,1989).TheflowersinGuatteria actually are terminalonveryshortbranches(brachyblasts)fromtheaxilsof leaves.Onlyinfewspeciesdoflowersterminatenormalleafy branches, namely G. acrantha, G. grandiflora, G. notabilis and G. terminalis (Fig.4a,b).Guatteria blainii must be mentioned hereaswellbecause,superficially,itseemstohavebothaxillaryandterminalflowers.Carefulinspection,however,shows that most seemingly terminal flowers in this species are infactstillaxillaryduetoabortionofthemainshootapex; nevertheless,trueterminalflowersalsooccur,butremainrare. Following Fries, the transition is best understood – without pretending to be a phylogenetic sequence! – starting with G. terminalis(Fries1919:32etseq.,as‘G. glauca’).Herethesingle flowers stand on the top of normal leafy twigs, but also on somewhat smaller lateral twigs originating from axils, with leavesofnormalshape,butofsmallersize(Fig.4b).Amarkedlyobliquearticulationisfoundc.1–2cmbelowtheflower and showing the attachments of the two bracts continuing theleafdispositionintwoalternatingrows.Thelowerofthe twobractsoftenisstillinplaceandisfoliaceous,i.e.,with theshapeofanormalleaf,butof(moreorless)smallersize. Theupperbractalreadyhaddroppedinallmaterialthatwas available,andcouldnotbestudied.Lookingatabranchof G. terminalisinitsentiretyitisnotdifficulttoenvisagethesuppression of apical flower development and at the same time an extreme reduction of axillary twigs to minute short shoots, the leaves of which have become reduced to small scale-like bracts(Fig.4c).Guatteria acrantha, a recently discovered anddescribedendemicofsouthernPanama,isverysimilarto G. terminalis.Anotherrecentlydescribedspecies,G. notabilis, is remarkable in a number of aspects as shown in an illustration byitsauthors(Mello-Silva&Pirani1988).Heretheterminal inflorescencesbear1–3flowers(Fig.4j,k).Thefirstfloweris formed at the end of a leafy twig in a comparable way as in G. terminalis, and is subtended by two foliaceous bracts with theloweronebeingthelargest.Theupperoneusuallydrops early,whereastheloweroneisoftenpresentwiththeflower. Fromtheaxilofthelowerbractoftenasecondflowerdevelops. Thisflowerissubtendedbytwobracts.Thesebractshavethe characteristic shape for the genus: obovate and more or less narrowing towards the base, and the upper part with incurving sidestoprotectthedevelopingflowerbud.Boththesebracts areshedearly.Exceptionally3-floweredinflorescencesoccur, the third flower originating from the axil of the lower bract of the secondflower(Fig.4k).Theresultistherhipidiatearrangement common in many genera of Annonaceae, but not in Guatteria. Perhapstheoddestmemberofthegenustobementionedhere is G. grandiflora, occurring not only with terminal solitary flowers, but also quite often with 2–few-flowered terminal paniclelikeinflorescences.Inadditiontoterminalflowersthereare alsoflowersonsmallaxillarytwigs.Theinflorescencesseem to branch dichotomously, without showing the familiar pattern ofaxillaryshootsfromaxilsofleavesorbracts.Oncloserlook partialfusionofaxes(concaulescence)appearstoaccount forwhatseemedstrangeatfirst.Fries(1939)gaveanalyses based on several herbarium specimens which he illustrated withdrawings(1939:293–296,f.1;Fig.41).Thesmallaxillary twigs of G. grandiflora just mentioned are comparable to inflorescences in other Guatteria species, distinct mainly by theelongateinternodesbetweenthebracts.Pairedflowersin theaxilscanoriginateintwoways(Fries1919).Oneisthata serialaccessorybudsituatedbelowthefirstbudalsodevelops aflower(Fig.4d).Thereappeartwoflowersinamoreorless parallelstageofdevelopment(‘einandergleichwertig’,Fries 1919)(Fig.4e).Thesecondpossibilityisdevelopmentofa lateralshootfromtheaxilofabractofthefirst-orderstalk.The twoflowersthenareinsuccessivestagesofdevelopment(‘Die zweiteBlüte…dieTochterblütedererstenundnichtgleichwertigmitdieser’,Fries1919).Itislikelythatadditionalflowers that appear, often after the leaf has fallen, can originate via bothways.However,becauseofthehighlycompactinflorescence structure of many Guatteriaspeciesitisoftendifficult to determine whether a flower arose from an accessory bud orfromtheaxilofabractextremelyclosetothebase.Apeculiar species is the recently described G. flagelliflora where inflorescencebranchinghastakenplacemanytimes(Fig.4g). InthecauliflorousG. scalarinervia the inflorescences are much morecompact,butbranchingcanstillbeseen(Fig.4h).Inother cauliflorousspecies,e.g.,G. ferruginea, G. scandens, flowers appear as if implanted in the callous tissue that had been formingasinflorescencesbecameolder(Fig.4i).Moremorphological study is needed here, perhaps best by observing living plantsduringgrowthstages.Alsotheroleofaccessorybuds is important, particularly regarding the question of whether one or(many)moreaccessorybudsmightbeinvolved. Afinalnoteshouldbemaderegardingtheuseofthetermpedicel.Inourdescriptionsthisreferstotheflowerstalkaswe intuitively see it in the leaf axils and as sticking out from the compact inflorescences that can be observed on older leafless partsoftheplant.Strictlyspeaking,onlythepartabovethe articulation should be termed pedicel, the part below the articulationbeingashortshootasexplainedabove.Forpractical descriptive reasons we refrained from discriminating between pedicel proper and the supporting short shoot in measuring theflowerstalkoverthewholelength,exceptfore.g.,G. terminalis, where pedicel length is measured from the articulation upward.InG. grandiflora the free parts of the flower stalks are measuredaspedicels. Flowers Flower buds are generally depressed to broadly ovoid in outline,rarelyconical(e.g.,G. friesiana).Theirapexisgenerally obtuse,butinafewspeciestheapexisdistinctlypointed(Fig. 5).Thesepalsarealwaysmuchsmallerthanthepetals.In most species the sepals are free, but in a few species they arebasallyconnate.Theirshapevariesfromovate-triangular tobroadlyovate-triangular.Thesepalsareappressedtothe petalsorpartlyorcompletelyreflexed.Thelengthofthesepals rangesfrom2–20mm;thelengthismostlyaveragingfrom 2–10mm.Ahandfulofspecieshassepalslargerthan10mm. ExtremesareG. latifoliafromSEBrazilwithsepalsof10–20by 10 mm and G. talamancana from Central America with sepals of15–20by10–15mm.Petalsareintwo3-merouswhorls, muchsurpassingthesepalsinsize.Theaestivationisimbricate, exceptionallyvalvate(e.g.,G. blepharophylla, G. heteropetala), usuallyonlyvisibleinyoungbuds.Thepetalsgenerallysoon spreadwhilecontinuingtodevelopandbecomelarger.While the growth phase takes a certain amount of time, anthesis itself, as is generally the case in Annonaceae, lasts only a short time.Intheshortanthesisphasethepetalshavetheirmaximum sizebeforetheyareshed.Asmatureflowerscanbeconsiderably larger than flowers in the development stage immediately preceding it, and given the fact that good ripe flowers may still not have been collected in many species, the measurements in the descriptions in a number of species may have to be read withcaution.Thecolourmostoftenreported,apartfromgreen,
8Blumea–Volume60/1–3,2015 is(shadesof)yelloworcream;whitehasbeenreportedonvariousoccasionsasisalsothecasewithpinkorred(e.g.,G. stipitata).Thechocolate-brownpetalsofG. aberrans and the leadcoloured petals of G. terminalisdeservespecialmention.The petals of the two whorls generally are of comparable size and shape.ThemostnotableexceptionisG. heteropetala where theouterpetalsareonlyabouthalfthesizeoftheinnerpetals. Thisfeatureformedthemainreasonforplacingthisspecies in a separate genus Heteropetalum. Also G. vallensis must be mentioned where the inner petals, as far as seen in the type collection,areabouthalfaslongastheouterpetals(Fig.70f). Thegeneralshapevariesfromelliptic,ovate-elliptic,ovate, obovate,tonarrowlyso,oroblonginvariouscombinations. Morerareare,amongothers,ovate-trullate,obtrullateorflabellateshapes(G. flabellata).Inthedescriptionsusuallyone set of measurements is given for all six petals, except where outerandinnerwhorldiffervisibly.PetallengthinGuatteria variesfrom5–50mm,butgenerallyisover10mm.Species with the smallest petals are G. emarginata(5–9by6–7mm) and G. minutiflora(5–15by3–7mm).Thelargestpetalsare foundine.g.,G. grandiflora(15–40by5–13mm),G. ucayalina(20–45by7–25mm),G. punctata(10–50by5–20mm), G. flabellata(25–50by10–40mm).Thetorusvariesfrom broadly ovoid to conical, shortly cylindrical or cushion-shaped with a convex, flat or concave depression at the top, rarely with aflattop(G. heteropetala)(seeVanHeusden1992).Thereare many stamens in the genus Guatteria.Theyareprovidedwith averyshortfilament;theanthersareextrorselydehiscent.All species have a distinct apical and discoid prolongation of the connective,theso-calledconnectiveshield(Fig.6).Theconnective shield mostly is flat, but in a few species it is adorned withaswellinginthecentre,calledumbo.Thepresenceofthis umboisnotconstantwithinaspecies.Theconnectiveshieldis covered with minute papillae or less frequently with tiny simple, erecthairs,oritisglabrous.Thelengthofthestamensmostly variesfrom1–2mm,exceptionallymore.Thelargeststamens are found in G. grandiflora, G. guianensis, G. paludosa and G. pittieri,wherethestamenscanreachalengthof3mm. Carpelsarefree,generallynumerous,varyingfrom1.5–4mm long, hairy, narrowed into a style or not, the stigma obovoid, discoid, or obconical, sometimes cup-shaped and grooved, ovule1,basal(VanHeusden1992). Fig. 5Flowerbudshapes.a.Conical;b.pointed;c.rounded.—Scalebars=5mm.—Drawing:EsméeWinkel2014. cba Fig. 6Stamens,variationinconnectiveshield.a.Papillateorglabrous;b.hairy;c.umbonate.—Scalebar=1.5mm.—Drawing:EsméeWinkel2014. cba
9 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria Fruit ThefruitinGuatteria is apocarpous and composed of few to many,freefruitlets,ormonocarps.Theshapeofthemonocarps is generally ellipsoid, occasionally narrowly ellipsoid, and rarely globose.Guatteria maypurensis is the only species of Guatteria withnarrowlyoblongoidmonocarps.Inafewspeciesthemonocarpsarenarrowlyellipsoidandfusiform(e.g.,G. hispida and G. inundata).Inthelatterthenarrowmonocarpsarepointedat bothends,auniquefeaturewithinthegenus.Thelengthofa monocarpbodyvariesfrom5to35mm,withthesmallestmonocarpsof5–7mmlongseeninG. delicatula and G. mon ticola, andthelargestonesreachingalengthofupto30oreven35mm in G. hispida, G. inundata, G. talamancana and G. trichocarpa. Thelengthofthefruitingpedicelsvariesfromofafewmillimeters upto50mm,andexceptionallytoover100mmlong(G. macropus).Inafewspecies,however,thefruitmayevenbealmost sessile(e.g.,G. schomburgkiana).Themonocarpsarefleshy intexture.Theygenerallymaturetoblackorpurple-black. Monocarpsinherbariumspecimenshaveacolourvaryingfrom blacktobrown.Thenumberofmonocarpsperfruitgenerally variesfrom10–75.Inafewspecies,e.g.,G. ayang annae and G. anteridifera, the number of monocarps is lower than 10, whereasinseveralspecies,e.g.,G. dolichopoda, G. griseifolia, G. hirsuta, G. meliodora, G. punctata, G. saffordiana and G. scytophylla,upto100monocarpsareencountered.Each monocarp is generally placed on top of a fleshy stalk, often red-colouredinvivo,termedthestipe.Thestipesgenerally havealengthbetween5–25mm.InG. australis, G. latifolia, Fig. 7Variationinseedsculpture.a. Guatteria discolorR.E.Fr.:smooth,raphenotdistinct;b.G. ucayalinaHuber:smoothtopitted,rapheflat;c.G. amplifolia Triana&Planch.,d.G. ferrugineaA.St.-Hil.,e.G. beckiiMaas&Westra,andf.G. amplifoliaTriana&Planch.:pitted,rapheraised;g.G. australisA.St.-Hil.and h.G. pakaraimaeScharf&Maas:rugulose,raphenotdistinct;i.G. griseifoliaMaas&Westra:rugose,rapheraised;j.G. verrucosaR.E.Fr.:rugose,raphenot distinct;alsonotethickfruitwall;k.G.decurensR.E.Fr.andl.G. blepharophyllaMart.:longitudinallyandtransverselygrooved,raphenotdistinct(a:Grenand 554;b:A.L. Gentry 6745;c:Liesner et al. 15199;d:Mori & Benton 12993;e:Arroyo et al. 2638;f:Mori & Kallunki 3046;g:G. Hatschbach 40693;h:H.D. Clarke et al. 10861;i:Neill et al. 12948;j:Haber 3893;k:Croat 20759;l:Amaral et al. 674,allU).
16 Blumea–Volume60/1–3,2015 G. hirsuta, is also widely spread, occurring in the western part oftheAmazonBasin(withfewcollectionsfromtheeastern part),butalsoinNorthernVenezuelaandinthePacificcoastof Colombia.Thefourthspeciesisthemostcommonone,namely G. punctata occurring all over tropical South America except forSEandEBrazil. Geographical analysis for Guatteria (forabbreviationsseethelistonpage18) 1.GreaterAntilles(2species)—blai,cari 2.LesserAntilles(1species)—cari 3.CentralAmericaandMexico(33species)—aber,acra, aeru, alat, alle, ampl, chir, cost, cras, dari, doli, dota, gale, graf, herr, jefe, luce, oliv, pacc, pana, pudi, rein, rost, rotu, sess, slat, taca, tala, tene, tome, ucay, vero, zamo 4.CentralAmericatonorthwesternColombia(2species)— aber, dari 5.CentralAmericatowesternpartofSouthAmerica(1species)—ucay 6.CentralAmericaandlargepartoftropicalSouthAmerica (1species)—tome 7.NorthwesternandadjacentCentralColombiaandEcuador (16species)—alta,arge,brep,carc,cras,cuat,elet, esme, goud, micr, nari, paci, pich, pitt, vall, veno 8.CentralColombiaandNorthernVenezuela(9species)— anti, chas, novo, rufo, saff, spe2, syns, vene, veru 9.GuianasandadjacentBraziland/orVenezuela(23species)—alti,amap,ante,ayan,clus,cons,eles,flex,inte, leuc, minu, mont, obla, oure, paka, palu, pann, part, rich, rubr, scan, wach, woko 10.AmazonBasinandalsointheGuianas(15species)—aber, blep, citr, disc, foli, guia, lies, mayp, megp, meli, proc, punc, scho, scyt, subs 11.WesternAmazonBasin(Ecuador,Peru,Colombia,Venezuela,BrazilandBolivia)(49species)—alba,arar,aren, atab, auya, beck, beni, bern, cast, chry, conf, cusc, decu, deli, denu, duod, dura, elat, erio, euge, flab, flag, frac, gent, grap, gris, hete, hirs, insc, japu, magu, megc, mode, obli, pacp, past, peru, rami, revo, rubo, sanc, scal, spec, spe3,stip,stpe,term,tric,tris 12.CentralAmazonBasin(6species)—cami,duck,elon,frie, hisp, stph 13.AmazonBasin(3species)—inun,long,spe2 14.ThroughoutmostpartsoftropicalSouthAmerica(1species)—punc 15.EasternAmazonBasin(6species)—crya,myri,odor,orix, poly, sabu 16.Southeastern,easternandnortheasternBrazil(15species) —aust,came,cand,capi,emar,ferr,lati,macr,nota,olig, pogo, pohl, sell, stca, vill 17.CentralBrazil(1species)—rupe 18.CentralBrazilandadjacentAmazonianBrazil(1species) —rigi MostspeciesofGuatteria inhabit non-inundated rain forests, andonlyafewgrowinperiodicallyinundatedrainforests(e.g., G. heteropetala, G. inundata).Fewspeciesarerestrictedto savannas(amongothersG. campinensis, G. maypurensis). ThemainbulkofGuatteria species occur in lowland forest, with just a handful of species occurring or reaching into montane or submontaneforests(e.g.,G. goudotiana, G. oblongifolia).The main altitudinal range is from 0–1500 m, with only few species exceedingthis.TheextremesareG. oblongifolia to 2900 m, G. goudotiana,reaching3000m,andG. grandiflora with one specimencollectedat3150m!Manyspeciesarefoundonloamy, clayeyorlateriticsoils.Speciesgrowingonwhitesandsare uncommon(e.g.,G. arenicola, G. atabapensis, G. maypurensis). Brazil Colombia Peru Venezuela Panama Costa Rica Mexico Guatemala Nicaragua Honduras Bolivia Guyana Suriname French Guiana Puerto Rico Dominican republic Haiti Cuba Lesser Antilles Ecuador Map 1 Distribution of Guatteria basedonc.6850georeferencedherbariumcollectionsofalmostallspecies.
17 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria TERMINOLOGY Shape—Fortermsapplyingtoshapewehaveconsequently used the terminology recommended by the Systematics Association Committee for descriptive biological terminology (1962),Hickey(1979),Radfordetal.(1974)andStearn(1992). Measurements—Measurementsaremostlybasedonherbariummaterial.Incaseswhereweusedfreshorpickled material as well we indicated values obtained by including thembetween{},e.g.,‘fruitwall0.1–0.2{–1}mmthick’. Colour—Colourdescriptionsofleavesapplytoherbarium material.Coloursfromfreshmaterial(ofpetalsandfruit) arereferredtoas‘invivo’. Innerfloralparts—Measurementsofcarpelsandtorusarenot includedinthismonograph. TAXONOMIC TREATMENT Guatteria Guatteria Ruiz&Pav.(1794)85;R.E.Fr.(1939)291.—Lecto(selectedby Hutchinson 1923):Guatteria glaucaRuiz&Pav. (=Guatteria punctata (Aubl.)R.A.Howard). CanangaAubl.(1775)244,nom.rej.vsCananga(A.DC.)Hook.&Thomson(1855),nom.cons.—Type:Cananga ouregou(=Guatteria ouregou (Aubl.)Dunal). HeteropetalumBenth.(1861)69.—Type:Heteropetalum brasiliense Benth. (=Guatteria heteropetalaBenth.). GuatteriopsisR.E.Fr.(1934)108.—Lecto(selectedbyFries1959b):Guatteriopsis sessiliflora(Benth.)R.E.Fr.(=Guatteria blepharophylla Mart.). GuatteriaRuiz&Pav.sect.AcranthaR.E.Fr.(1939)303.—Type:Guatteria terminalisR.E.Fr. GuatteriaRuiz&Pav.sect.AustroguatteriaR.E.Fr.(1939)304.—Type: notdesignated. GuatteriaRuiz&Pav.sect.Asterantha R.E.Fr.(1939)318.—Type:not designated. GuatteriaRuiz&Pav.sect.MicrocalyxR.E.Fr.(1939)327.—Type:not designated. GuatteriaRuiz&Pav.sect.MicrophyllumR.E.Fr.(1939)332.—Type:Guatteria pohlianaSchltdl. GuatteriaRuiz&Pav.sect.Cordylocarpus R.E.Fr.(1939)333.—Type: Guatteria clavigeraR.E.Fr.(=Guatteria australisA.St.-Hil.). GuatteriaRuiz&Pav.sect.TrichocloniaR.E.Fr.(1939)335.—Type:not designated. GuatteriaRuiz&Pav.sect.StenophyllumR.E.Fr.(1939)354.—Type:not designated. GuatteriaRuiz&Pav.sect.LeptophyllumR.E.Fr.(1939)372.—Type:not designated. GuatteriaRuiz&Pav.sect.MacrophyllumR.E.Fr.(1939)375.—Type:not designated. GuatteriaRuiz&Pav.sect.Guatteria(1939)383.—Lecto(selectedby Hutchinson1923):Guatteria glaucaRuiz&Pav.(=Guatteria punctata (Aubl.)R.A.Howard). GuatteriaRuiz&Pav.sect.SclerophyllumR.E.Fr.(1939)390.—Type:not designated. GuatteriaRuiz&Pav.sect.MacroguatteriaR.E.Fr.(1939)395.—Lecto (selectedbyFries1959b):Guatteria ferruginea A.St.-Hil. GuatteriaRuiz&Pav.sect.OligocarpusR.E.Fr.(1939)405.—Type:not designated. GuatteriaRuiz&Pav.sect.StenocarpusR.E.Fr.(1939)410.—Type:notdesignated. GuatteriaRuiz&Pav.sect.PteropusR.E.Fr.(1939)416.—Type:notdesignated. GuatteriaRuiz&Pav.sect.PycnanthaR.E.Fr.(1939)431.—Type: Guatteria elongataBenth. GuatteriaRuiz&Pav.sect.TylodiscusR.E.Fr.(1939)432.—Lecto(selected byFries1959b):Guatteria chrysopetala (Steud.)Miq.(=Guatteria punctata (Aubl.)R.A.Howard). GuatteriaRuiz&Pav.sect.BrachystemonR.E.Fr.(1939)454.—Type:not designated. GuatteriaRuiz&Pav.sect.CephalocarpusR.E.Fr.(1939)460.—Type: notdesignated. GuatteriaRuiz&Pav.sect.TrichostemonR.E.Fr.(1939)470.—Type:not designated. GuatteriaRuiz&Pav.sect.DolichocarpusR.E.Fr.(1939)475.—Type:not designated. GuatteriaRuiz&Pav.sect.LeiophyllumR.E.Fr.(1939)483.—Type:not designated. GuatteriaRuiz&Pav.sect.MegalophyllumR.E.Fr.(1939)485.—Type: notdesignated. GuatteriaRuiz&Pav.sect.MecocarpusR.E.Fr.(1939)488.—Type:not designated. GuatteriaRuiz&Pav.sect.DichrophyllumR.E.Fr.(1939)509.—Type: Guatteria discolorR.E.Fr. GuatteriaRuiz&Pav.sect.StigmatophyllumR.E.Fr.(1939)511.—Type: Guatteria puncticulataR.E.Fr.(=Guatteria modestaDiels). GuatteriaRuiz&Pav.sect.ChasmanthaR.E.Fr.(1939)512.—Type:not designated. GuatteriaRuiz&Pav.sect.ChasmanthaR.E.Fr.subsect.Chasmantha(1939) 513.—Type:notdesignated. GuatteriaRuiz&Pav.sect.ChasmanthaR.E.Fr.subsect.VerrucosaeR.E.Fr. (1939)519.—Type:notdesignated. GuatteriaRuiz&Pav.sect.ChasmanthaR.E.Fr.subsect.GrandifloraeR.E.Fr. (1939)523.—Type:Guatteria grandifloraDonn.Sm. GuatteriaRuiz&Pav.sect.AnomalanthaR.E.Fr.(1939)524.—Guatteria Ruiz&Pav.subg.Anomalantha(R.E.Fr.)R.E.Fr.(1959b)86.—Type:Guatteria anomalaR.E.Fr.(=Guatteria grandifloraDonn.Sm.) GuatteriellaR.E.Fr.(1939)540.—Type:Guateriella tomentosaR.E.Fr.(= Guatteria trichocarpaErkens&Maas). Treesorshrubs,veryrarelylianas.Indumentcomposedofsimple, rarelytuftedhairsorabsent.Youngtwigsterete,rarelywinged, densely to sparsely covered with appressed to erect hairs, soonbecomingglabrousinmanyspecies.Leaves distichous, simple,entire,petiolate,estipulate;laminamostlyellipticto ovate, chartaceous to coriaceous, sometimes verruculose, rarely scabridulous, upper side glabrous or less often sparsely to densely covered with appressed to erect hairs, lower side glabrous to densely covered with appressed to erect hairs, base acute, attenuate, cordate, or obtuse, rarely stem-clasping, apex acuminate to acute, very rarely obtuse to rounded, primary vein impressed, flat, rarely raised above, secondary veins distinct,sometimesindistinct,between5and35oneither side of primary vein, angle of secondary veins with primary vein(30–)60–90°impressedtoraisedabove,smallestdistancebetweenloopsandmargin1–5(–10)mm,tertiaryveins impressedtoraisedabove,reticulatetopercurrent.Flowers bisexual,3-merous,commonlymedium-sized,solitaryor in 1–few-flowered inflorescences in axils of leaves or axils of fallen leaves, rarely terminal or terminal on lateral shoots, exceptionally inflorescences long-persisting and developing a varying number of successive flowers on older branches and incauliflorousspeciesandoneflagelliflorousspecies;pedicels with mostly oblique articulation below the middle, with 2 bracts justbelowthearticulationandmostly4–6bractsfurtherdown, without bracts above the articulation, bracts generally small, scale-like, soon falling, rarely one or more bracts persisting andfoliaceous;pedicelsandoutersideofbracts,sepalsand petals glabrous to densely covered with appressed to erect hairs;flowerbudsdepressedtobroadlyovoid,veryrarelyconical;sepals3,valvateorreduplicate-valvate,freeorconnateat thebase;petals6,free,generallysubequal,imbricate,much longer than the sepals, green, mostly maturing to cream or yellowinanthesis;stamensnumerous,withveryshortfilament, not septate, apex of connective discoid, papillate, glabrous, orrarelyhairy;staminodesabsent;carpelsnumerous,free, ovule 1, basal, stigma obovoid, discoid, or obconical, sometimescup-shapedandgrooved.Fruitapocarpous,consisting of numerous, indehiscent, stipitate, fleshy monocarps, stipes 5–50mmlong,sometimesabsentorstronglyreduced;seed1, a rudimentary aril present, brown, surface smooth, pitted, rugulose, rugose, or transversely and longitudinally grooved, endospermruminationsspiniformorlamellate. Chromosomenumber—2n=28.
18 Blumea–Volume60/1–3,2015 Distribution—About177species,allovertheNeotropics exceptforParaguay. Habitat&Ecology—Innon-inundatedorrarelyperiodically inundated, lowland rain forest or sometimes submontane to montaneforest,rarelyinsavannas.Generallyatlowelevations, butrarelyreachinguptoorover3000m. Note—ThegenusisnamedafterJ.B.Guatteri,professor ofBotanyinParma,Italy. Keys Guatteria, with few exceptions, is morphologically relatively homogeneouswhencomparedtootherannonaceoustaxa. Also,thereisgreatvariation(orplasticity)incharacterstates withmuchoverlapasaresult.Furthermore,Guatteria shows an extended period of flower development as contrasted with itsshortperiodofanthesis.Oftenripeflowersshowamarked increaseinsizeandachangeincolour(mostlylasting1–2days only).Unfortunatelyripeflowershavestillnotbeencollected inmanyspecies.Inevitably,anattempttoconstructadichotomous key given the three factors mentioned above is going to result in several leads leading to the same species in many instances.Suchakeyisnotonlyalmostimpossibletoconstruct butalsoextremelyfrustratingtouse.Inouropinion,therefore, a synoptical key is to be preferred, like we did in the past in revisions of Rollinia(Maasetal.1992)andDuguetia(Maaset al.2003).Fortwosmallerareas,however,viz.CentralAmerica andEandSEBrazil,inwhichonlyalimitednumberofspecies hadtobetreated,wedecidedtopresentadichotomouskey. Synoptical key—Theusermaystartwithanynumber,dependinguponthestateofthematerialathand.Ausefullead is,ofcourse,theentrydevotedtogeography.Thespeciesare representedbyfour-lettersymbols(seebelow).Speciesmentioned in more than 1 lead are printed in italics.Ifthereisa marked difference between the frequency ratio of character states, only the lead representing the one with the lowest valueisprintedinfull.Thealternateoneismentionedas‘vs’ betweenparentheses. aber = G. aberrans acra = G. acrantha aeru = G. aeruginosa alat = G. alata alba = G. alba alle = G. allenii alta = G. alta alti = G. alticola amap = G. amapaensis ampl = G. amplifolia ante = G. anteridifera anti = G. antioquensis arar = G. araracuarae aren = G. arenicola arge = G. argentea atab = G. atabapensis aust = G. australis auya = G. auyantepuiensis ayan = G. ayangannae beck = G. beckii beni = G. beniensis bern = G. bernardii blai = G. blainii blep = G. blepharophylla brev = G. brevipetiolata came = G. campestris cami = G. campinensis cand = G. candolleana capi = G. capixabae carc = G. carchiana cari = G. caribaea cast = G. castilloi chas = G. chasmantha chir = G. chiriquiensis chry = G. chrysophylla citr = G. citriodora clus = G. clusiifolia conf = G. confusa cons = G. conspicua cost = G. costaricensis crap = G. crassipes crav = G. crassivenia crya = G. cryandra cuat = G. cuatrecasasii cusc = G. cuscoensis dari = G. darienensis decu = G. decurrens deli = G. delicatula denu = G. denudata disc = G. discolor doli = G. dolichopoda dota = G. dotana duck = G. duckeana duod = G. duodecima dura = G. dura elat = G. elata eles = G. elegans elet = G. elegantissima elon = G. elongata emar = G. emarginata erio = G. eriopoda esme = G. esmeraldae euge = G. eugeniifolia ferr = G. ferruginea flab = G. flabellata flag = G. flagelliflora flex = G. flexilis foli = G. foliosa frac = G. fractiflexa frie = G. friesiana gale = G. galeottiana gent = G. gentryi goud = G. goudotiana graf = G. grandiflora grap = G. grandipes gris = G. griseifolia guia = G. guianensis herr = G. herrerana hete = G. heteropetala hirs = G. hirsuta hisp = G. hispida insc = G. insculpta inte = G. intermedia inun = G. inundata japu = G. japurensis jefe = G. jefensis lati = G. latifolia leuc = G. leucotricha lies = G. liesneri long = G. longicuspis luce = G. lucens macr = G. macropus magu = G. maguirei mayp = G. maypurensis megc = G. megalocarpa megp = G. megalophylla meli = G. meliodora micr = G. microcarpa minu = G. minutiflora mode = G. modesta mont = G. monticola myri = G. myriocarpa nari = G. narinensis nota = G. notabilis novo = G. novogranatensis obla = G. oblonga obli = G. oblongifolia odor = G. odorata olig = G. oligocarpa oliv = G. oliviformis orix = G. oriximinae oure = G. ouregou pacc = G. pachycarpa paci = G. pacifica pacp = G. pachyphylla paka = G. pakaraimae palu = G. paludosa pana = G. panamensis pann = G. pannosa part = G. partangensis past = G. pastazae peru = G. peruviana pich = G. pichinchae pitt = G. pittieri pogo = G. pogonopus pohl = G. pohliana poly = G. polyantha proc = G. procera pudi = G. pudica punc = G. punctata rami = G. ramiflora rein = G. reinaldii revo = G. revoluta rich = G. richardii rigi = G. rigida rost = G. rostrata rotu = G. rotundata rubo = G. ruboides rubr = G. rubrinervis rufo = G. rufotomentosa rupe = G. rupestris sabu = G. sabuletorum saff = G. saffordiana sanc = G. sanctae-crucis scal = G. scalarinervia scan = G. scandens scho = G. schomburgkiana scyt = G. scytophylla sell = G. sellowiana sess = G. sessilicarpa slat = G. slateri spe1 = G. sp. 1 spe2 = G. sp. 2 spec = G. spectabilis stca = G. stenocarpa stpe = G. stenopetala stph = G. stenophylla stip = G. stipitata subs = G. subsessilis syns = G. synsepala taca = G. tacarcunae tala = G. talamancana tene = G. tenera term = G. terminalis tome = G. tomentosa tric = G. trichocarpa tris = G. trichostemon ucay = G. ucayalina vall = G. vallensis vene = G. venezuelana veno = G. venosa vero = G. verrucosa veru = G. verruculosa vill = G. villosissima wach = G. wachenheimii woko = G. wokomungensis zamo = G. zamorae
19 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria KEY TO THE CENTRAL AMERICAN SPECIES 1. Plantcauliflorousorramiflorous.—CostaRica....... ................................78.G. herrerana 1. Plantnotcauliflorousorramiflorous ...............2 2. Flowersorinflorescenceterminal(someaxillaryflowers mayalsobepresent)...........................3 2. Flowersorinflorescenceaxillary..................4 3. Pedicels10–65mmlong;leaves10–22by3–8cm;petals15–40mmlong;flowersoftenin2–several-flowered inflorescences.—Mexico,Guatemala,Honduras...... ............................... 74.G. grandiflora 3. Pedicels5–12mmlong;leaves5–8by2–3.5cm;petals 7–9mmlong;flowersalwayssolitary.—Panama..... ..................................2.G. acrantha 4. Youngtwigsdenselycoveredwitherecthairs........5 4. Youngtwigscoveredwithappressedhairsorglabrous1 7 5. Leafbasecordate(sometimesobtuseinG. pudica)...6 5. Leafbaseacute,obtuseorattenuate(sometimesoblique in G. tomentosa)..............................7 6. Lowersideofleavescoveredwithreddishbrown,soft hairs,uppersideglabrousexceptforthehairyprimaryvein; pedicels10–35mmlong;flowerbudsobtuse;sepals5–8 mmlong.—CostaRica..............129.G. pudica 6. Bothsidesofleavescoveredwithbrown,stiffhairs;pedicels10–50(–70)mmlong;flowerbudsslightlypointed; sepals6–13mmlong.—CentralAmerica,tropicalSouth America....................... 163.G. tomentosa 7. Primaryveinraisedontheuppersideoftheleaves(but also slightly raised to flat above in G. zamorae and in G. tala-mancana);leavesnarrowlyoblong-elliptic.—Costa Rica,Panama..................34.G. chiriquiensis 7. Primaryveinimpressedtoflatontheuppersideofthe leaves(leavesnarrowlyovatetonarrowlyobovate,never oblong-elliptic)................................8 8. Leavesverruculose............................9 8. Leavesnotverruculose........................12 9. Sepals15–20mmlong;monocarps20–30mmlong;stipes ofmonocarps2–3mmlong.—CostaRica,Panama... .............................160.G. talamancana 9. Sepals4–10mmlong;monocarps5–16mmlong;stipes of monocarps 2–10 mm long....................10 10. Petioles3–9mmlong;youngtwigsfinallyglabrous.— Panama .........................175.G. zamorae 10. Petioles0–7mmlong;youngtwigscoveredwithlongpersisting hairs .............................. 11 11. Youngtwigsoftenzigzagging;monocarp2.3–3timeslongerthanthestipes;monocarps9–16mmlong,wallc. 0.5 mmthick.—CostaRica,Panama...110.G. oliviformis 11. Youngtwigsnotzigzagging;monocarp1.3–1.6timeslongerthanthestipes;monocarps8–12mmlong,wall0.2–0.3 mmthick.—Honduras,Nicaragua,CostaRica,Panama, Colombia........................ 3.G. aeruginosa 12. Flowerbudsdistinctlypointed(butseealsoG. ucayalina); petalsbrowntochocolate-coloured;leavescoriaceous.— Panama,Colombia,Brazil(Roraima)....1.G. aberrans 12. Flowerbudsobtuse;petalsgreentoyellow;leaveschartaceous(sometimescoriaceousinG. ucayalina)......13 13. Sepals2–5mmlong;petalscanaryyellow;leavesmostly greenishwhendried.—Panama.........6.G. allenii 13. Sepals5–13mmlong;petalsdifferentlycoloured;leaves never greenish when dried.....................14 14. Sepalsappressedtospreading,butfinallyreflexed;uppersideofleavesmostlycoveredwitherecthairs;flower budsslightlypointed.—CentralAmerica,tropicalSouth America ....................... 163.G. tomentosa 14. Sepalssoonreflexed;uppersideofleavesglabrousexcept forthehairyprimaryveinorcoveredwithappressedhairs; flower buds obtuse ...........................15 15. Tertiaryvenationofleavesmostlypercurrent;stipesofmonocarps8–50mmlong;sepals4–11mmlong.—Nicaragua, CostaRica,Panama,tropicalSouthAmerica......... ...............................166.G. ucayalina 15. Tertiaryvenationofleavesreticulate;stipesofmonocarps 3–20mmlong;sepals4–7mmlong.............16 16. Petioles2–6mmlong;pedicels30–60mmlong;monocarps7–11mmlong;stipes15–20mmlong.—Honduras, CostaRica,Panama.............51.G. dolichopoda 16. Petioles5–10mmlong;pedicels15–30mmlong;monocarps5–7mmlong;stipes3–10mmlong.—Panama. ..............................159.G. tacarcunae 17. Youngtwigsdistinctlywinged.—CostaRica,Panama.. .....................................4.G. alata 17. Youngtwigsterete ............................18 18. Leavesmostlylengthwisefoldedwhendried.—Panama ..................................86.G. jefensis 18. Leavesneverfoldedwhendried.................19 19. Leavesdistinctlyroundedattheapex,denselyverruculose. —Panama.....................137.G. rotundata 19. Leavesacuminateorsometimesacuteattheapex,sometimes verruculose ............................20 20. Leafvenationstronglyraisedonbothsides;primaryvein raisedontheupperside.—CostaRica............. ...............................42.G. crassivenia 20. Leafvenationneverraisedonbothsides;primaryveinflat to impressed on the upper side ..................21 21. Monocarpssessileorshortlystipitate(stipesupto3mm long).......................................22 21. Monocarpsdistinctlystipitate(stipes>2mmlong)...24 22. Leaves18–33by5–10cm,ratherdenselyverruculose. —Panama................... 150.G. sessilicarpa 22. Leaves7–20by3–6cm,notorsparselyverruculoseon the lower side ...............................23 23. Monocarpsglobose,20–25mmdiam,wall5–6mmthick; pedicels10–20mmlong.—CostaRica............ ..............................113.G. pachycarpa 23. Monocarpsnarrowlyellipsoid,10–14by4–8mm,wall 1–1.5mmthick;pedicels3–7mmlong.—CostaRica.. .................................132.G. reinaldii 24. Monocarpsdistinctlybeakedattheapex;petiole15–20 mmlong.—CostaRica.............136.G. rostrata 24. Monocarpsroundedtominutelyapiculateattheapex; petiolemostlymuchsmaller(exceptG. slateri 4–20mm long)......................................25 25. Leavesverruculose............... 170.G. verrucosa 25. Leavesnotverruculose........................26 26. Youngtwigsmostlyzigzagging;secondaryveinsraisedon the upper side of the leaves ....................27 26. Youngtwigsstraight;secondaryveinsimpressedtoraised on the upper side of the leaves..................29 27. Floweringpedicels0–5mmlong;monocarps3–5times longerthanthestipes;monocarps15–17mmlong.— CostaRica......................... 52.G. dotana 27. Floweringpedicels5–32mmlong;monocarps0.7–2.6 timeslongerthanthestipes;monocarps7–10mmlong ...........................................28
20 Blumea–Volume60/1–3,2015 28. Monocarp1.6–2.6timeslongerthanthestipes;stipesof monocarps3–10mmlong;uppersideofleavesglabrous. —Panama........................151.G. slateri 28. Monocarp0.7–0.8timeslongerthanthestipes;stipesof monocarps8–14mmlong;uppersideofleaveswitha hairyprimaryvein.—CostaRica,Panama.......... ............................. 40.G. costaricensis 29. Leavesverysmallandnarrow(7–12by2–3mm).—Costa Rica..............................161.G. tenera 29. Leavesgenerallymuchlarger...................30 30. Pedicels30–75mmlong.—Panama,Colombia...... ...............................46.G. darienensis 30. Pedicels7–50mmlong........................31 31. Basalmarginsofleavesoftenrevolute;leafvenationdistinctlyraisedontheupperside.—CostaRica,Panama ...................................91.G. lucens 31. Basalmarginsofleavesnotrevolute;leafvenationflatto impressed above, rarely slightly raised on the upper side ...........................................32 32. Seedspitted;leafvenationoftendistinctlyimpressedabove. —ThroughoutCentralAmericaandMexico.......... ................................10.G. amplifolia 32. Seedsrugose;leafvenationflattoslightlyraisedabove ...........................................33 33. Leaves10–15by3–5cm,secondaryveins8–15;monocarps10–13mmlong;stipesofmonocarps9–17mmlong. —Mexico.......................71.G. galeottiana 33. Leaves13–26by5.5–12cm,secondaryveins15–18; monocarps7–10mmlong;stipesofmonocarps6–10mm long.—Panama...............118.G. panamensis KEY TO THE SE AND E BRAzILIAN SPECIES 1. Plantcauliflorous.—Bahia,EspíritoSanto,MatoGrosso, MinasGerais,Pará,RiodeJaneiro....64.G. ferruginea 1. Plantnotcauliflorous ...........................2 2. Flowersorinflorescencesterminal.—MinasGerais... ................................104.G. notabilis 2. Flowersorinflorescencesinaxilsofleavesand/oronleafless branchlets................................3 3. Leafapexroundedoremarginate.................4 3. Leafapexacuminate,acute,rarelyobtusetoemarginate (seealsoG. rupestris, G. tomentosa)..............5 4. Leavesverruculose,apexemarginate,tertiaryveinsinconspicuous;pedicels15–30mmlong.—EspíritoSanto. . ...............................60.G. emarginata 4. Leavesnotverruculose,apexrounded,emarginate,or acute,tertiaryveinsconspicuous;pedicels10–15mm long.—MinasGerais..............141.G. rupestris 5. Stipesofmonocarps0–3mmlong ................6 5. Stipesofmonocarps4–30mmlong(seealsoG. campestris and G. villosissima).........................7 6. Youngtwigsdenselycoveredwitherecthairs;pedicels 2–10mmlong;monocarpsellipsoidtoglobose,6–13mm long;seedssmooth.—MatoGrosso,Paraíba,Pernambuco......................147.G. schomburgkiana 6. Youngtwigsglabrous;pedicels8–25mmlong;monocarps narrowlyellipsoid,13–17mmlong;seedsrugose.—Espírito Santo.....................29.G. capixabae 7. Monocarpsnarrowlyellipsoid ....................8 7. Monocarpsgloboseorellipsoid...................9 8. Monocarps18–25mmlong;stipesofmonocarps5–10 mmlong.—Bahia...............153.G. stenocarpa 8. Monocarps13–17mmlong;stipesofmonocarps2–4mm long.—EspíritoSanto.............29.G. capixabae 9. Monocarpsglobose;seedsnotattachedtothemonocarp wallinherbariummaterial.—Alagoas,Bahia.......... ...............................109.G. oligocarpa 9. Monocarpsellipsoid;seedsattachedtothemonocarpwall in herbarium material..........................10 10. Youngtwigsdenselycoveredwitherecthairs...... 11 10. Youngtwigsdenselycoveredwithappressedhairsorglabrous......................................16 11. Pedicels60–120mmlong.—Bahia,EspíritoSanto,Minas Gerais..........................92.G. macropus 11. Pedicels5–70mmlong........................12 12. Leafmarginsrevolute.........................13 12. Leafmarginsnotrevolute(seealsoG. tomentosa). . 14 13. Flowerbudsobtuse;leafmarginsrevoluteoverthewhole length;stipesofmonocarps3–4mmlong.—Ceará,EspíritoSanto,MinasGerais,RiodeJaneiro........... ..............................172.G. villosissima 13. Flowerbudsslightlypointed;leafmarginsrevoluteatthe baseonlyornotrevolute;stipesofmonocarps4–20mm long.—Bahia,EspíritoSanto,MinasGerais......... .............................. 163.G. tomentosa 14. Leaves17–40by5–15cm;stipesofmonocarps10–25 mmlong.—Bahia,EspíritoSanto,MatoGrosso,Minas Gerais,Pará,RiodeJaneiro .........64.G. ferruginea 14. Leaves5–15by1.5–4cm;stipesofmonocarps2–6mm long.......................................15 15. Leafbasecordate,rarelyobtuse.—Bahia,EspíritoSanto, MinasGerais,RiodeJaneiro......28.G. candolleana 15. Leafbaseacute.—Bahia,DistritoFederal,EspíritoSanto, Goiás,MatoGrosso,MinasGerais,Pernambuco,Riode Janeiro,SãoPaulo.............. 149.G. sellowiana 16. Leaves13–35by4–13cm.—Alagoas,Bahia,Ceará,EspíritoSanto,MatoGrosso,MinasGerais,Paraíba,Pernambuco..........................125.G. pogonopus 16. Leavesgenerallymuchsmaller(4–21by1.5–7cm). 17 17. Sepalsconnate,10–20mmlong.—MinasGerais,Riode Janeiro............................87.G. latifolia 17. Sepalsfreeorsometimesbasallyconnate,5–10mmlong ...........................................18 18. Stipesofmonocarps2–5mmlong.—Bahia,Espírito Santo,MinasGerais,RiodeJaneiro..26.G. campestris 18. Stipesofmonocarps5–30mmlong ..............19 19. Stipesofmonocarps5–9mmlong;sepalsfreeorinitially connate,5–7mmlong.—Bahia,EspíritoSanto,Minas Gerais..........................126.G. pohliana 19. Stipesofmonocarps10–30mmlong;sepalsfree,5–10 mmlong.—Bahia,EspíritoSanto,Goiás,MinasGerais, Paraná,RiodeJaneiro,RioGrandedoSul,SantaCatarina, SãoPaulo........................17.G. australis SYNOPTICAL KEY 1. Habit liana(vstreesorshrubs)—beck, flex, frac, scan, syns. 2. Young twigs zigzagging (vs straight)—alta,cost, dota, eles, frac, goud, lati, oliv, slat. 3. Young twigs coveredwitherecthairs—aber,aeru,alle,alta,alti,amap, anti, arar, ayan, beck, beni, bern, brev, cami, cand, carc, chir, chry, citr, cuat, decu, deli, disc, doli, duck, elet, erio, esme, euge, ferr, frie, grap, guia, herr, hirs, hisp, insc, inte, lies, macr, micr, minu, mont, nota, novo, obla, obli, oliv, oure, pann, part, peru, poly, proc, pseu, pudi, punc, revo, rich, rubo, rubr, rufo, rupe, saff, sanc, scho, sell,
21 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria stip, stpe, stph, taca, tala, tome, tric, tris, ucay, vene, vill, wach,zamo,spe2. coveredwithappressedhairs—acra,alat,alba,ampl, ante, aren, arge, atab, aust, auya, beck, blai, blep, brev, came, cari, cast, chas, clus, conf, cost, crap, crav, crya, cuat, cusc, dari, decu, deli, denu, dota, duod, dura, elat, eles, elon, emar, erio, esme, euge, flab, flag, flex, foli, frac, gale, gent, goud, graf, gris, hete, hirs, inun, jefe, leuc, lies, long, luce, magu, mayp, megc, megp, meli, mode, mont, myri, nari, odor, olig, orix, pacc, paci, palu, pana, past, peru, pich, pitt, pogo, pohl, punc, rami, rein, revo, rigi, rost, rotu, rupe, sabu, saff, scal, scan, scyt, sess, slat, spec, stpe, subs, syns, tene, term, tric, tris, vall, veno, vero,veru. glabrous—capi,cons,graf, japu, lati, mayp, orix, pacp, paka, pohl, rupe, stca, veno, vero,woko,spe1. 4. Petiole length 0–5mm—aber, acra, aeru, alba, alle, alta, alti, ampl, anti, arar, aren, atab, aust, auya, beck, beni, bern, blai, brev, came, cami, cand, capi, carc, cari, chir, chry, citr, cons, cost, crav, crya, cuat, cusc, dari, decu, deli, denu, disc, doli, dota, duck, duod, dura, elat, eles, elet, elon, emar, erio, esme, euge, foli, frac, frie, gale, goud, graf, grap, gris, guia, hirs, hisp, insc, inte, inun, jefe, lati, leuc, lies, long, luce, macr, magu, mayp, meli, micr, minu, mode, mont, myri, nari, nota, novo, obla, obli, odor, olig, oliv, orix, oure, paka, pana, part, past, peru, pitt, pogo, pohl, poly, proc, pseu, pudi, punc, rein, rich, rigi, rubo, rubr, rufo, rupe, sabu, saff, sanc, scho, scyt, sell, sess, slat, spec, stip, stpe, stph, subs, syns, taca, tala, tene, term, tome, tric, tris, ucay, vall, vene, vero, veru, vill, wach, woko, zamo, spe1, spe2. 6–10mm—aber, alat, alba, alle, ampl, anti, aren, arge, atab, aust, ayan, beck, blai, blep, came, capi, chas, chry, citr, clus, conf, cons, cost, crap, cusc, dari, decu, denu, disc, doli, dota, duck, duod, dura, elat, elon, erio, euge, ferr, flab, flag, flex, foli, frie, gale, gent, goud, graf, gris, guia, herr, hete, insc, inte, inun, japu, jefe, lati, lies, luce, macr, mayp, meli, micr, minu, mode, mont, obla, obli, olig, oliv, orix, oure, pacc, paci, pacp, paka, pana, pann, past, peru, pich, pitt, pogo, proc, pseu, pudi, punc, rami, revo, rich, rigi, rotu, rufo, sabu, saff, sanc, scan, scho, scyt, sell, sess, slat, spec, stca, stip, stpe, stph, subs, syns, taca, term, tric, tris, ucay, vall, vene, veno, veru, vill, wach, woko, zamo, spe1, spe2. >10mm—alat, amap, ampl, ante, arge, ayan, blep, cast, clus, crap, denu, disc, elon, ferr, flab, flex, foli, gent, hete, inun, luce, mayp, megc, megp, olig, pacp, paci, palu, past, pitt, pogo, proc, punc, rami, revo, rost, saff, scal, scan, scyt, sess, slat, spec, stca, ucay, veno. 5. Leaf surface verruculose—aeru,alat,arar,arge,atab, auya, blep, cand, capi, chas, crap, crav, crya, cuat, decu, disc, duod, dura, elet, emar, euge, gent, goud, graf, grap, gris, guia, hisp, insc, japu, long, luce, megc, meli, mode, nari, nota, novo, obli, oliv, paci, paka, past, peru, pitt, rein, rigi, rotu, sess, tala, tene, tric, vall, veno, vero, zamo, spe1. scabridulous—ante,ayan,clus,cons,flex,frie,inun, japu, palu,scan,veru,wach. smooth—allotherspecies. 6. Leaves folded lengthwise (vs not folded)—jefe, mayp. 7. Leaf base cordate—ampl, cami, cand, flab, frie, hirs, pudi, punc, scan, tome, vill. stem-clasping—cami, long, tome. different—allotherspecies. 8. Basal margins of leaves revolute—ampl, anti, blai, carc, cast, duck, emar, goud, luce, mode, nota, pacp, past, pudi, revo, rufo, rupe, sess, stip, tala, tome,vill. 9. Leaf apex rounded,truncateorobtuse—acra,aren, cast, crap, dari, flag, gris, hete, meli, nota, pacp, rigi, rotu, rupe, term, tome. emarginate—blai, clus, emar, rigi, rupe, tome. different—allotherspecies. 10. Hairs on lower side of lamina (sometimes on large veins only): appressed(toerectonlargeveinsonly)—aber,acra,alat, alba, alta, ampl, ante, aren, arge, atab, aust, auya, ayan, beck, beni, bern, blai, blep, brev, came, capi, cari, cast, chas, citr, clus, conf, cons, cost, crap, crav, crya, cuat, cusc, dari, decu, deli, doli, dota, duod, dura, elat, eles, elon, emar, erio, esme, euge, flab, flag, flex, foli, frac, frie, gent, goud, graf, gris, guia, hete, hirs, inte, inun, jefe, lati, leuc, lies, long, luce, magu, mayp, mega, megc, meli, mode, mont, myri, nari, novo, obla, obli, odor, olig, orix, pacc, paci, paka, pana, pann, past, peru, pich, pitt, pohl, proc, pseu, punc, rami, rein, revo, rigi, rost, rotu, rubr, rupe, sabu, saff, sanc, scal, scho, scyt, sell, sess, slat, spec, stip, stpe, stph, subs, syns, taca, tene, term, tric, tris, ucay, vall, veno, vero, veru, spe1, spe2. erectallover—aeru,alle,alti,amap,anti,arar,beni, bern, brev, cami, cand, carc, chir, chry, citr, crav, cuat, decu, deli, disc, doli, duck, elet, erio, ferr, graf, grap, guia, herr, hirs, hisp, insc, macr, micr, minu, nota, novo, obli, oliv, oure, part, poly, proc, pudi, punc, revo, rich, rubo, rubr, rufo, saff, sanc, scho, sell, stip, tala, tome, tric, tris, ucay, vene,vill,zamo. glabrous—blai, came, cand, capi, chas, conf, cons, crap, dari, denu, euge, ferr, flex, gale, gent, goud, graf, hete, hirs, japu, lies, macr, magu, mayp, mega, meli, mode, nari, olig, orix, oure, pacc, paci, pacp, palu, pich, pitt, pogo, pohl, pseu, sabu, scan, sess, slat, stca, stph, subs, vene, veno, veru, wach, woko, spe1, spe2. 11. Primary vein raised on upper side of lamina (vs impressed to flat) — aeru, alat, alle, alti, aren, beni, chas, chir, citr, conf, crav, crya, deli, disc, dota, gent, gris, hete, hirs, nota, oliv, pacc, proc, slat, stpe, subs, tala, vene, vero, zamo. 12. Marginal vein present (vs absent)—arar,cami, cuat, decu, elet, grap, guia, insc, long, megp, novo, veno, wach,woko. 13. Secondary venation indistinct (vs distinct)—aren, atab, clus, cons, crya, denu, doli, duck, elat, elet, euge, frac, frie, japu, leuc, macr, mayp, minu, mode, nari, nota, obla,olig,pacp,paka,past,pogo,pohl,rubo,spe1. 14. Number of secondary veins 5–9(vs≥10)—acra,alle, alti, aren, arge, aust, auya, ayan, blai, cand, capi, cari, clus, conf, cost, cusc, duck, eles, emar, euge, flex, foli, gale, graf, herr, hirs, leuc, lies, macr, magu, minu, odor, oliv, orix, paka, part, pohl, pudi, rein, rigi, rotu, rubr, rupe, scan, scho, slat, stpe, syns, taca, tala, term, tome, vene, vero, vill, woko. >20(vs≤20)—aber, aeru, alata, ampl, arar, aust, blep, cami, cons, crav, crya, cuat, decu, disc, ferr, flab, frie, grap, guia, insc, inun, lies, long, mega, nari, novo, paci, palu, punc, rami, revo, rufo, sabu, sanc, stca, stip, tric, ucay, veno. 15. Distance between loops/marginal vein and margin ≥ 5 mm (vs ≤ 4)—atab, cari, citr, conf, dari, decu, denu, duck, dura, elat, ferr, flab, flex, frie, goud, gris, guia, hisp, inun, lies, micr, novo, obli, orix, oure, past, pich,
22 Blumea–Volume60/1–3,2015 pitt, pogo, proc, pudi, punc, rein, revo, rost, rubri, sabu, sanc, scan, sess, stip, subs, ucay, veno, veru, wach, spe1. 16. Tertiary venation percurrent(vsreticulate)—alat,amap,cami,chrys, crap, disc, dura, elat, eles, graf, grap, guia, herr, insc, inte, megp, meli, novo, obla, orix, oure, palu, pana, punc, rami, revo, rich, sabu, syns, taca, tene, tric, ucay, veno. indistinct(vsdistinct)—alat,ante,arar,cons,cuat, dari, decu, dura, elat, elon, emar, frie, long, megp, mode, nota, novo, obla, oliv, orix, palu, pana, past, pohl, punc, scal, scan, stca, subs, tala, wach. 17. Flower position terminal—acra,blai, graf,nota,term. inleafaxils—aber,aeru, alba, alle, alta, alti, amap, ampl, ante, anti, arar, aren, arge, atab, aust, auya, ayan, beck, beni, bern, blai, blep, brev, came, cami, cand, capi, carc, cari, cast, chas, chir, chry, citr, clus, conf, cons, cost, crap, crav, crya, cuat, cusc, dari, decu, deli, denu, disc, doli, dota, duck, duod, dura, elat, elet, elon, emar, erio, esme, euge, ferr, flab, flex, foli, frac, frie, gale, gent, goud, graf, grap, gris, guia, hete, hirs, hisp, insc, inte, inun, japu, jefe, lati, lies, long, luce, macr, magu, mayp, megc, megp, meli, micr, minu, mode, mont, myri, nari, obla, obli, odor, olig, oliv, orix, oure, pacc, paci, pacp, paka, palu, pana, pann, part, past, peru, pich, pitt, pogo, pohl, poly, proc, pseu, pudi, punc, rami, rein, revo, rich, rigi, rost, rotu, rubo, rubr, rufo, rupe, sabu, saff, sanc, scho, scyt, sell, sess, slat, spec, stca, stpe, stph, stip, subs, syns, taca, tala, tene, tome, tric, tris, ucay, vall, vene, veno, vero, veru, vill, wach, woko, zamo, spe1, spe2. onleaflessbranchlets—aeru, alat, alba, amap, ampl, arar, aust, blai, brev, cami, cari, chas, chry, citr, crap, crya, cusc, denu, dota, duod, dura, elat, eles, elon, erio, ferr, flab, flex, gent, gris, guia, herr, hirs, insc, inun, japu, jefe, leuc, lies, long, mayp, mega, meli, micr, mode, mont, myri, obla, orix, oure, paci, pacp, paka, palu, pana, pann, part, past, poly, proc, punc, rami, revo, rich, rost, rubo, rufo, saff, sanc, scan, scho, scyt, stip, subs, syns, tome, ucay, vall, veno, wach. plantcauliflorous—cons, ferr, herr, long, novo, scan,scal. plantflagelliflorous—flag. 18. Flowers solitary(exceptionally≥2)—acra,aeru,alat,alba,alta, alti, ampl, ante, arar, aren, arge, aust, auya, beck, beni, bern, blai, cand, capi, cast, chry, clus, conf, cons, cost, crav, cuat, cusc, decu, deli, denu, doli, dota, dura, elat, eles, emar, erio, esme, euge, ferr, flex, foli, frac, gale, graf, grap, guia, hete, hirs, hisp, insc, inte, inun, lati, leuc, long, macr, magu, mayp, megp, meli, micr, minu, mont, myrio, nari, obla, obli, odor, olig, oliv, oure, pacc, paci, paka, palu, part, past, peru, pich, pohl, proc, pudi, rigi, rotu, rubo, rubr, rupe, scho, sell, sess, slat, stca, stpe, stph, taca, tala, tene, term, tome, tric, tris, vall, vene, vero,veru,vill,woko,zamo. 1– ≥2-floweredinflorescences—aber,alle,amap,ampl, anti, atab, ayan, blep, brev, came, cami, carc, cari, chas, chir, citr, crap, crya, dari, disc, duck, duod, elet, erio, ferr, flab, frie, gent, goud, gris, herr, japu, jefe, lies, long, luce, mode, nota, orix, pacp, pana, pann, pitt, pogo, poly, pseu, punc, rami, rein, revo, rich, rost, rufo, sabu, saff, sanc, scyt,spec,stip,subs,ucay,veno,wach,spe1,spe2. 2–∞ —disc, elon, erio, ferr, flag, graf, long, novo, scal, scan. 19. Length of pedicels (flowering, fruiting) ≤10(–15)mm—acra,alle, amap, ampl, ante, arar, aren, arge, atab, auya, ayan, blep, came, cami, capi, cari, chas, citr, conf, cons, cost, crav, crya, denu, disc, dota, duck, duod, dura, elat, elon, flex, frie, gale, gent, goud, graf, hete, hirs, hisp, inte, jefe, leuc, lies, luce, magu, megp, meli, minu, mode, mont, nota, obla, obli, olig, oliv, orix, oure, pacc, palu, pann, past, peru, pogo, pohl, poly, pseu, pudi, punc, rein, revo, rich, rotu, rufo, rupe, sabu, saff, scho, scyt, sell, sess, stip, stph, subs, syns, tene, term, tome, tric, ucay, veno, vero, vill, wach, zamo, spe1, spe2. ≥30mm—alta,ampl, aust, beni, blai, blep, brev, cand, chir, chry, crap, cuat, cusc, dari, decu, doli, dura, elet, emar, ferr, flag, flex, foli, gale, goud, graf, grap, gris, guia, hirs, insc, inun, japu, jefe, lati, lies, long, luce, macr, magu, mayp, megc, micr, nari, novo, obli, oliv, paci, paka, pana, part, pich, pitt, pogo, pohl, pudi, punc, rami, rigi, rost, rubo, rubr, rufo, saff, sanc, scal, scan, slat, spec, stca, stpe, taca, tala, tome, tris, ucay, vall, vene, vill, woko, zamo. 10–30mm—allotherspecies. 20. Flower buds pointedoracute(vsobtuse;unknowninsomespecies)— aber, amap, aren, ayan, blai, chry, disc, doli, guia, hirs, megp, palu, pann, punc, rami, rubo, rubr, sanc, scyt, tome, ucay. conical(vsdepressedtobroadlyovoidortriangular-ovoid; unknowninsomespecies)—aber,amap,frie,hete. 21. Sepal length >10mmlong(vs≤10mmlong)—aber, alat, ampl, arar, areu, chry, cuat, decu, disc, ferr, frie, graf, grap, guia, hirs, hisp, insc, lati, macr, megp, nota, novo, pann, past, pitt, punc, rost, sanc, spec, tala, tome, tric, ucay, vill. 22. Petals unequal (vs subequal)—frie,hete,rost. 23. Petal colour (but notice that the flower colour of many species is still unknown) browntoblack—aber,cami,foli, mayp, sanc, syns, veno. reddishtopurple—cuat, flab, hete, luce, megp, nota, obli, pann, past, peru, revo, sanc, scho, stip, vill. greytoblack—goud,term. orange—carc, luce, megp, oure, rubr, vill. differentcolours—allothers,includingthespeciesfor whichtheflowercolourisstillunknown). 24. Stamens with connective shield densely hairy (vs glabrous, papillate or [thinly] hairy)—amap,anti, citr,duck,poly,scho,stip. 25. Monocarp size <10mm(vs≥10mm)—acre, aeru, alle, alta, aren, atab, aust, beck, beni, blai, cand, cast, chir, cons, cost, crya, cuat, dari, deli, denu, doli, duck, eles, elet, esme, ferr, flab, flag, foli, hirs, inte, lati, leuc, lies, luce, magu, mayp, micr, minu, mode, mont, myri, nari, obli, oliv, oure, pana, part, pitt, pohl, proc, pudi, punc, rich, rigi, rubo, rubr, rufo, rupe, sabu, saff, scho, scyt, sell, slat, stip, stpe, syns, taca, tome, ucay, vall, vene, vill, wach, zamo. >20mmlong—alat, ante, ayan, blep, chry, clus, conf, decu, disc, frie, graf, guia, hete, hisp, inun, long, megc, megp, meli, pacc, peru, rami, scal, sess, stca, tala, tric. differentlength—allotherspecies. unknown—alba,alti,auya,brev,crav,elon,emar,erio, frac,odor,poly,rufo,spec,stph,tene,tris. 26. Monocarps < 10—acra,alat, ante, arar, ayan, cand, capi, cari, cast, chry, citr, clus, frie, graf, hete, leuco, orix, paka, part, pseu, punc, rein, rost, scho, sess, stca, stip, subs, woko, spe2. 27. Monocarp shape fusiform(vsdifferentlyshaped)—hisp,inun. narrowlyoblongoid(vsdifferentlyshaped)—mayp.
23 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria 28. Thickness of monocarp wall 1–2mm—alat,ante,arge,cami, cast, chry, clus, crap, decu, disc, duck, duod, euge, gent, guia, insc, mega, megp, meli, mode, nota, olig, pann, past, pich, pogo, punc, rein, scal, sess, stca, tric, vero. >3mm—pacc,vero. unknown—alba,alti,auya,beni,brev,crav,elon,emar, erio, euge, frac, graf, odor, pana, poly, spec, stph, tene, tris, vero,woko,spe2. <1mm—allotherspecies. 29. Monocarp indument denselycoveredwithappressedhairs—amap,lies(but soonglabrous!),tric. denselytoratherdenselycoveredwitherecthairs(‘hirsute’) —arar, decu, hisp, rubo. denselycoveredwitherecthairs(‘velutinous’)—cami, nota, pann,stip. rather densely covered with appressed and erect hairs — beni. 30. Stipe length 0–4mm—acra,alat, alle, aren, arge, atab, beni, blai, came, cami, capi, cari, chas, citr, clus, conf, cons, crap, crya, cuat, cusc, denu, disc, dota, duck, duod, elet, esme, frie, graf, gris, guia, hete, inun, jefe, magu, mayp, megp, nota, obli, oliv, pacc, paci, pacp, paka, past, rein, revo, rigi, rost, rotu, rubo, rufo, rupe, sabu, sanc, scho, sess, slat, subs, syns, taca, tala, term, tome, tric, vene, vero, veru, vill, wach. >20mm—alat, ampl, aust, carc, elat, ferr, foli, herr, hirs, inte, lati, lies, micr, myri, olig, our, pann, proc, punc, rubr, saff, scal, scyt, ucay, woko. differentlength—allotherspecies. unknown—alba,alti,auya,brev,crav,elon,emar,erio, frac,odor,poly,rufo,spec,stph,tene,tris. 31. Seed surface smooth—aber, aren, ayan, bern, blai, chir, citr, disc, duck, euge, flex, foli, minu, mode, olig, pann, pogo, punc, revo, rubr, rupe, scho, scyt, stip, syns, term, ucay. pitted—acra, aber, ampl, anti, arge, atab, ayan, beck, came, cand, cari, chir, cons, crya, cusc, dari, denu, doli, dota, elat, eles, euge, ferr, flab, goud, herr, hete, hirs, inte, inun, jefe, lati, lies, luce, macr, magu, mayp, micr, minu, mode, mont, myri, nari, obla, obli, olig, oure, paci, pann, part, past, pich, pogo, pohl, proc, pudi, punc, rein, revo, rich, rami, rigi, rost, rubr, rupe, sabu, saff, sanc, scyt, sell, sess, slat, stpe, syns, tala, term, tome, ucay, vall, vill, wach,woko,spe1. transverselygrooved—ante, atab, camp, cons, cost, deli, dota, eles, esme, flab, flex, frie, hete, hirs, leuc, lies, mont, oure, paci, past, pich, pohl, proc, punc, rich, rost, scyt, sess, slat, stpe, zamo. longitudinally grooved — acra, blai, camp, dura, inun, pacp, wach. rugulose—alat,ante, aust, blep, came, camp, carc, clus, cons, cost, duod, dura, elet, goud, graf, guia, hisp, leuc, long, magu, minu, nota, obli, pacp, paka, palu, rubo, saff, sanc, tala, vall,vene,veno. rugose—acra, aeru, alle, alta, amap, blep, capi, chry, crap, cuat, denu, duod, esme, flag, gale, gent, graf, grap, gris, megc, megp, oliv, orix, pacc, pana, pitt, punc, rotu, rufo, subs, vero, zamo. longitudinallyandtransverselygrooved—arar,blai, blep, cami, cari, conf, decu, gent, guia, hisp, insc, japu, long, megp, novo, peru, scal, scan, stca, subs,tric,veru. unknown—alba,alti,brev,cast,chas,crav,elon,emar, erio, frac, meli, odor, pana, poly, spec, stph, taca, tene, tris,spe2. 32. Distribution Antilles—blai,cari. CentralAmericaandMexico—aber, acra, aeru, alat, alle, ampl, chir, cost, crav, dari, doli, dota, gale, graf, herr, jefe, luce, oliv, pacc, pana, pudi, rein, rost, rotu, sess, slat, taca, tala, tene, tome, ucay,vero,zamo. PacificColombiaandPacificEcuador—aber, alta, anti, arge, brev, carc, citr, crap, cuat, dari, elet, esme, goud, hirs, micr, nari, paci, pich, pitt, punc, taca, tome, ucay, vall,veno. CentralColombiaandCentralEcuador—arge, chas, crap, cuat, dari, goud, guia, hirs, lies, megp, novo, punc, rufo, stip, subs, tome, ucay, veru. NorthernVenezuela—bern,foli, hirs, mayp, punc, saff, scho, subs, ucay, vene, veru. Guianas(includingadjacentregionsinBrazilandVenezuela)—alti,ante, ayan, blep, citr, clus, cons, disc, eles, flex, foli, guia, inte, leuc, lies, mayp, megp, meli, minu, mont, obla, oure, paka, palu, pann, part, proc, punc, rich, rubr, scan, scho, scyt, subs, ucay, wach,woko. WesternAmazonBasin(WofManaus)—alba,arar,aren, atab, auya, beck, beni, blep, brev, cast, chry, citr, conf, cusc, decu, deli, denu, disc, duod, dura, elat, erio, euge, flab, flag, flex, foli, frac, gent, goud, grap, gris, guia, hete, hirs, insc, inun, japu, lies, long, magu, mayp, megc, megp, meli, mode, mont, obli, pacp, past, peru, proc, pseu, punc, rami, revo, rigi, rubo, rubr, sanc, scal, scan, scho, scyt, spec, stip, stpe, stph, subs, term, tome, tric, tris, ucay, spe1,spe2. CentralAmazonBasin—blep, cami, citr, duck, elon, frie, hisp, inun, lies, long, past, scyt, subs. NorthernBrazil(Amapá,Roraima)—aber, amap, ante, blep, crya, guia, lies, mayp, megp, pann, punc, rich, scan, scho, tric, wach. EasternAmazonBasin(EofManaus)—blep, citr, crya, ferr, foli, guia, hirs, inun, lies, long, megp, myri, odor, orix, poly, proc, punc, rigi, sabu, scan, scho, scyt, subs, spe1. NortheasternBrazil(Alagoas,Ceará,Maranhão,Paraiba, Pernambuco,Piauí)—citr, mode, olig, pogo, punc, scan, scho. EasternBrazil(Bahia,EspiritoSanto)—aust, came, cand, capi, emar, ferr, macr, nota, olig, pogo, pohl, sell, stca, tome, vill. CentralBrazil(Goiás,MatoGrosso,MinasGerais)—aust, blep, came, cand, dura, ferr, foli, lati, macr, mayp, nota, pogo, pohl, punc, rigi, rupe, scyt, sell, tome, vill. SoutheasternBrazil(Paraná,RiodeJaneiro,RioGrande doSul,SantaCatarina,SãoPaulo)—aust, came, cand, ferr, lati, sell, vill. 33. Elevation above2000m(vs0–2000m)—aren, carc, crap, frac, goud, gris, obli, oliv, past, punc, slat, syns, term. 1. Guatteria aberransErkens&Maas—Map2 Guatteria aberransErkens&MaasinErkensetal.(2006)201,t.1,f. 1.— Type:Maas et al. 9570(holoU;isoINB,K,MO,PMA,SCZ),Panama, Colón,SantaRitaRidge,Parcela31ofCTFS,250m,9June2004. Tree8–25mtall,10–30cmdiam;youngtwigsdenselycovered witherect,curly,brownhairs,soonglabrous.Leaves: petiole 5–10mmlong,2–3mmdiam,decurrentasprominentridge inyoungtwigs;laminanarrowlyelliptic,sometimesnarrowly ovate,15–32by4–9.5cm(leafindex2.9–3.7),coriaceous,not verruculose, shiny, dark blackish brown to grey above, brown to pale brown below, sparsely covered with appressed hairs above, soon glabrous, sparsely covered with appressed hairs
24 Blumea–Volume60/1–3,2015 below, base obtuse, extreme base sometimes slightly attenuate, apexacuminate(acumen10–20mmlong),primaryveinflat to slightly impressed above, secondary veins distinct, 15–22 on either side of primary vein, raised above, smallest distance betweenloopsandmargin3–4mm,tertiaryveinsraisedabove, reticulate.Flowersin1–2(–3)-floweredinflorescencesinaxils ofleavesoronleaflessbranchlets;pedicels7–20mmlong, 1–2mmdiam,fruitingpedicels2–3mmdiam,denselycovered witherect,curly,brownhairs(toratherdenselycoveredwith appressedhairsintheRoraimaspecimen),articulatedatc.0.2 fromthebase,bracts4–6,soonfalling,notseen;flowerbuds conicaltoovoid,distinctlypointed;sepalsfree,triangularto broadlytriangular,7–12by5–8mm,spreadingbutsoonbecoming completely reflexed, outer side densely covered with erect,curly,brownhairs;petalsbrowntochocolate-colouredin vivo,narrowlyelliptictoelliptic,13–27{–32}by5–15mm,outer sidedenselycoveredwitherect,curly,brownhairs;stamens 1–1.5mmlong,connectiveshieldglabrous.Monocarps10–40, green, maturing blue-black in vivo, blackish in sicco, ellipsoid, 10–16by7–9mm,sparselycoveredwitherect,curly,brown hairs,soonglabrous,apexapiculate(apiculum<0.5mmlong), wallc.1mmthick,wrinkledinsicco,stipes9–15by1–2mm. Seed ellipsoid, 10–12 by 5–6 mm, dark brown, smooth to pitted,raphenotdistinctfromrestofseed. Distribution—Panama,Colombia(Chocó),Brazil(Roraima). Habitat&Ecology—Inforest,sometimesalongroadsides. Atelevationsof0–300(–540)m.Flowering:January,February, June,July;fruiting:February,June. Vernacularname—Panama:Yayo(Howell 20). Notes—Guatteria aberrans is remarkable by its pointed flowerbuds(afeatureonlysometimesseeninG. ucayalina and G. pannosa).Anothernoteworthyaspectincollectionsfrom Panamaareitschocolate-brownpetals.Afurtherpeculiarityof this species is that the inner petals seem to stay closed during thewholelifecycle(noopenbuddevelopmentseen). Cogollo & Brand 373(JAUM,MO)fromColombia,Antioquia, Mun.SanLuis,HighwayMedellín-Bogotá,2kmfromRíoClaro, nearRíoSamaná,alt.790m,probablybelongstothisspecies. ItmatchesG. aberransinalmostallaspects(pointedflower buds,leafsizeandflowersize).Itsflowersareannotatedas pink(‘rosada’)anditsfruitsasblack. Sá et al. 3(K)fromBrazil,Roraima,Rorianópolis,RioXixuaú, alt.36misplacedherealthoughitisgeographicallyquitefar fromthelocalitiesinPanamaandthePacificcoastofColombia. Itsharesalmostallflowerandleaffeatures,butthebranchlets are glabrous or sparsely covered with minute hairs at most, and Brazil Colombia Peru Venezuela Panama Costa Rica m ala Nicaragua H o n d u r a s Lesser Antilles Ecuador Map 2 Distribution of Guatteria aberrans (●),G. acrantha (£ ),G. aeruginosa (■)andG. allennii(p). the pedicels and leaves are covered with appressed instead oferecthairs. 2. Guatteria acranthaErkens&Maas—Map2 Guatteria acrantha Erkens&MaasinErkensetal.(2006)202, t.1,f.2,3. —Type:Rivera 355(holoSTRI;isoMO,PMA,U),Panama,LosSantos, Distr.Tonosí,CerroLosPiraguales,ElCortezo,900m,20April1994. Treeorshrub4–20mtall;youngtwigssparselycoveredwith appressedhairs,soonglabrous.Leaves:petiole1–3mm long,c.1mmdiam;laminanarrowlyobovatetonarrowlyelliptic,5–8by2–3.5cm(leafindex2.2–2.8),chartaceous,not verruculose, dull, dark brown above, brown to brownish green below, glabrous above, rather densely covered with appressed hairs below, base attenuate, apex rounded or sometimes very shortlyandbluntlyacuminate(acumen<2mmlong),primary vein flat to slightly impressed above, secondary veins distinct, 6–9 on either side of primary vein, flat to slightly raised above, smallest distance between loops and margin 1–2 mm, tertiary veinsflattoslightlyraisedabove,reticulate.Flowers solitary, terminaloralsoinaxilsofleaves;floweringandfruitingpedicels 5–12mmlong,c.1mmdiam,denselycoveredwithappressed, brownhairs,articulatedat0.2–0.3fromthebase,bractsof terminalflowers1–2,foliaceous(?),soonfalling,notseen, bractsofaxillaryflowersc.5,soonfalling,notseen;flower budsbroadlyovoid;sepalsfree,broadlyovate-triangular,3–5 by3–4mm,appressedtoreflexed,outersidedenselycovered withappressed,brownhairs;petalsgreenishinvivo,broadly ovate-triangular,7–9by5–7mm,outersidedenselycovered withappressed,brownhairs;stamens1–1.5mmlong,connectiveshieldpapillate.Monocarpsc.10,greeninvivo,blackin sicco,narrowlyellipsoid,7–10by3–4mm,subglabrous,apex apiculate(apiculumc.1mmlong),wallc.0.1mmthick,stipes 1–2by1mm.Seednarrowlyellipsoid,c.8by3mm,brown,pitted to rugose and more or less distinctly longitudinally grooved, raphedistinct,flattoslightlyraised. Distribution—Panama(Chiriquí,LosSantos,Veraguas). Habitat&Ecology—Incloudforest.Atelevationsof900– 1500m.Flowering:FebruarytoApril,July;fruiting:April. Note—Guatteria acrantha is unique by its solitary terminal, minute flowers and by its very tiny, mostly roundish tipped leaves.Superficially,itlookssimilartoG. rotundata by the shape ofitsleaves,butitisquitedistinctbyhavingterminalflowers. 3. Guatteria aeruginosaStandl.—Plate1a,b;Map2 Guatteria aeruginosa Standl.(1929)206;R.E.Fr.(1939)522,f.35g.—Type: Cooper 526 (holoF;isoBM,G,K,NY,US),Panama,BocasdelToro, RegionofAlmirante,Jan.1928. Tree5–25mtall,10–55cmdiam;youngtwigsdenselycoveredwithlong-persisting,erect,brownhairs.Leaves:petiole1–8 mmlong,2–4mmdiam;laminanarrowlyellipticornarrowly oblong-elliptictonarrowlyobovate,18–42by5–16cm(leaf index2.2–4.2),chartaceous,denselyverruculosetolessoften not verruculose, dull, greenish grey, grey, to dark brown above, brown below, sparsely covered with appressed hairs to glabrous above, but primary vein densely covered with erect, brown hairs, densely to sparsely covered with erect, brown hairs below, base acute to obtuse, often slightly decurrent along petiole, apexacuminate(acumen5–20mmlong),primaryveinflat to impressed or rarely raised above, secondary veins distinct, 12–24oneithersideofprimaryvein,flattoraisedabove,smallest distance between loops and margin 2–5 mm, tertiary veins slightlyraisedabove,reticulate.Flowers solitary on leafless branchletsorinaxilsofleaves;floweringandfruitingpedicels 13–30mmlong,1–2mmdiam,fruitingpedicelstoc.4mm diam, densely covered with erect brown hairs, articulated at
25 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria Plate 1a,b.Guatteria aeruginosaStandl.a.Floweringbranch;b.flowerandfruit.–c.Guatteria alataMaas&Setten.Partoftwig.–d.Guatteria allenii R.E.Fr.Flowers.–e–g.Guatteria altaR.E.Fr.e.Floweringbranch;f.flower;g.fruit.–h.Guatteria australis A.St.-Hil.Ripeflower(a:Chatrou et al. 720; b: Chatrou et al. 741;c:McPherson 20133;d:Maas et al. 9543;e–g:Gal Tadri 560;h:Maas et al. 8816).—Photos:a,b:L.W.Chatrou;c:G.McPherson; d:P.J.M.Maas;e–g:GalTadri;h.P.J.M.Maas. c b d a g h e f
32 Blumea–Volume60/1–3,2015 Fig. 13 a.Guatteria amapaensisMaas&Westra.Fruit.–b.Guatteria antioquensisMaas&Westra.Fruit.–c.Guatteria atabapensisAristeg.exD.M.Johnson &N.A.Murray.Fruitingbranch,detail.–d.Guatteria auyantepuiensisMaas&Westra.Detailofflower.–e,f.Guatteria beckiiMaas&Westra.e.Flower; f.fruit.–g.Guatteria beniensisMaas&Westra.Youngfruit,alsonotedenselyerecthairyjuvenilepartsinupperleft.–h.Guatteria blainii(Griseb.)Urb.Detail offlower(a:Daly et al. 4028,U;b:E. Correa et al. 93,U;c:Wurdack & Adderley 42759,isotypeUS;d:F. Cardona 2619,holotypeUS;e,f:Beck & Haase 9895,holotypeU;g:Nee 34593,holotypeNY;h:Ekman 5920, S). c b d a gh ef
33 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria aryveinsdistinctlyimpressedontheupperside;furthermorethe leaf base is obtuse to cordate with the basal margins somewhat revolute.TowardstheWest,inCostaRica,Nicaragua,andall other western Central American countries, leaves and petioles tend to be smaller, and the secondary veins are less impressed toevenflatontheupperside.Moreover,theleafbaseismostly acute,althoughobtuseleafbasesarealsosometimesfound. We consider all of the variation to be encompassed within a singlespecies.Foranillustrationofthisvariation,seeErkens 2007:104,f.1.However,itisclearthatthiscomplexshouldbe analysedfurthermorphologicallyaswellaswithmoleculardata. Ithasbeennoted,inthefieldaswellasinherbariummaterial, thatthesepalsofthePanamanianmaterialaresometimes persistentinfruit. 11. Guatteria anteridiferaScharf&Maas—Map3 Guatteria anteridifera Scharf&MaasinScharfetal.(2008)517,f.1–3.— Type:Sabatier & Prévost 5047(holoU2sheets;isoCAY),FrenchGuiana, km122.5ofroadfromCayennetoSaintGeorges,near‘SavaneRoche Virginie’,27July2006. Treeuptoc.40mtall,50–80cmdiam,withbuttressesupto c.2mhighandc.80cmatthebase,limbsdecurrent;young twigs densely covered with appressed, silvery hairs, soon glabrous.Leaves:petiole15–20mmlong,2–3mmdiam, winged;laminaobovatetonarrowlyoblong-elliptic,9–21by 5–8cm(leafindex1.8–2.7),coriaceous,scabridulous,dull, greyish to greenish brown above, dark brown below, glabrous above, sparsely covered with appressed hairs below, base longattenuate,apexshortlyacuminate(acumen3–10mmlong), primary vein impressed above, secondary veins distinct, 12–16 on either side of primary vein, slightly impressed above, smallestdistancebetweenloopsandmargin2–3mm,tertiaryveins inconspicuous,flatabove,reticulate.Flowersin1(–2)-flowered inflorescencesinaxilsofleaves;floweringandfruitingpedicels 10–15 mm long, 1–2 mm diam, densely to rather densely coveredwithappressed,silveryhairs,articulatedat0.2–0.3from the base, bracts 5–6, soon falling, broadly to very broadly ovate atthebase,1–2mmlong,otherbractsnotseen;flowerbuds notseen;sepalsfree,broadlyovate-triangular,5–6by5–6mm, reflexed, outer side densely covered with appressed, silvery hairs;petalsyellowishgreeninvivo,elliptic,oblong-elliptic,to obovate,21–23by10–13mm,outersidedenselycoveredwith appressed,short,silveryhairs;stamensc.1mmlong,connectiveshieldhairy,umbonate.Monocarpsc.4,greeninvivo (immatureones),greyishtoblackishbrowninsicco,ellipsoid, 20–25by8–11mm,apexsparselycoveredwithappressed hairs,apexapiculate(apiculum1–1.5mmlong),wallc.1mm thick,stipes5–8by1–1.5mm.Seedellipsoid,16–19by7–9 mm, pale to dark brown, rugulose to transversely grooved, rapheimpressed. Distribution—FrenchGuiana,AmazonianBrazil(Amapá). Habitat&Ecology—Innon-inundated,primaryforest.At aboutsealevel.Flowering:July,August;fruiting:November. Vernacularnames—Notrecorded. Note—Guatteria anteridifera is well marked by large buttressesofupto2mhigh.Itseemssimilartosomespeciesof sect.Mecocarpus(e.g.,G. blepharophylla),butitcompletely lackstheverruculosestructuressotypicalforthatsection. Instead,ithasscabridulousleaveswhicharecoveredwith minute,circularstructures.Itperhapscomesclosesttothe GuyananendemicG. paludosa, but that species lacks the buttresses,hasmuchlargerleaves(30–55cmvs9–21cmlong in G. anteridifera),long-acuminatevsshortlyacuminateleaves (acumen10–45mmvs3–10mmlonginG. anteridifera)and largerstamens(2–3mmvs1mmlonginG. anteridifera). 12. Guatteria antioquensisMaas&Westra,sp. nov.—Fig. 13b,14;Map5 Guatteria citriodoraaffinis,sedpedicellislongioribusseminibusfoveatis benediffert.—Typus:Cogollo et al. 3796 (holoJAUM2sheets;isoCOL, MO),Colombia,Antioquia,Mun.SanLuis,nearAquitania,8kmfromHwy. Medellín-Bogotá,830m,25Nov.1988. Tree5–23mtall,c.6cmdiam(oncerecorded);youngtwigs denselycoveredwitherect,brownlong-persistinghairs.Leaves: petiole3–6mmlong,c.1mmdiam;laminanarrowlyovate tonarrowlyelliptic,10–19by3–5cm(leafindex2.4–3.6), chartaceous, not verruculose, shiny, dark brown to greyish black above, brown below, sparsely covered with erect hairs mainly along primary vein above, soon glabrous, densely covered with erect, brown hairs, hairs simple or in bundles of 2–4,below,baseacutetoobtuse,basalmarginsoftenrevolute,apexacuminate(acumen5–10mmlong),primaryvein impressed above, secondary veins distinct, 10–15 on either side of primary vein, slightly raised above, smallest distance between loops and margin 1–2 mm, tertiary veins slightly raised above,reticulate.Flowers in 1–2-flowered inflorescences, in axilsofleaves;pedicels15–20mmlong,c.1mmdiam,fruitingpedicels20–35mmlong,1–2mmdiam,denselycovered witherect,brownhairs,articulatedat0.2–0.5fromthebase, bracts5–7,soonfalling,notseen;flowerbudsdepressed ovoid;sepalsfree,broadlyovate-triangular,3–6by3–6mm, apex and margins often reflexed, outer side densely covered witherect,brownhairs;petalsgreeninvivo,narrowlyelliptic tonarrowlyobovate,8–20by4–9mm,outersidedensely coveredwitherecttoappressed,brownhairs;stamensc.1.5 mmlong,connectiveshielddenselyhairy(hairsc.1mmlong). Monocarpsc.50,green,maturingpurple-blacktoredinvivo, blackinsicco,ellipsoidtonarrowlyellipsoid,7–10by3–4mm, rather densely to sparsely covered with erect to appressed, brownhairs,toglabrous,apexapiculate(apiculumc.1mm long)torounded,wallc.0.1mmthick,stipes5–10by0.5mm, rather densely to sparsely covered with erect to appressed, brownhairstoglabrous.Seedellipsoid,7–8by4mm,brown, deeplypitted,rapheraised. Distribution—Colombia(Antioquia). Habitat&Ecology—Innon-inundatedforest.Atelevations of790–1550m.Flowering:NovembertoFebruary;fruiting: February,June,November,December. Vernacularnames—Colombia:Frisolo(Cárdenas L. & Ramírez 2746), Garrapato(Fonnegra et al. 3109),Garrapato lanudo(Giraldo 190). Brazil Colombia Peru Venezuela Panama Costa Rica a temala Nicaragua Honduras Bolivia Guyana Suriname French Gui Haiti Lesser Antilles Ecuador Map 5 Distribution of Guatteria antioquensis (●),G. araracuarae (£ ),G. arenicola (■),G. argentea (p),G. atabapensis (u)andG. bernardii (✸).
34 Blumea–Volume60/1–3,2015 Fig. 14 Guatteria antioquensisMaas&Westra.Floweringbranch(Cogollo & Estrada 202; MO).
35 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria Other specimens examined. ColomBia, Antioquia,Mun.SanLuis,margen derechadelacarreteraQuebradas-NaranjalesylaCristalina,420–460m, Callejas et al. 4151(HUA);Mun.Amalfí,VeredasChorritos,LaMaría,Montebello,km15–35oftheroadfromChorritostoLosMonos,SNEofAmalfí, 1180–1550m,Callejas et al. 9022(HUA,NY,U);Mun.SanLuis,VeredaLa Josefina,QuebradalaMariola,700–925m,Cárdenas L. & Ramírez 2746 (JAUM);Mun.SanLuis,sectorRíoSamaná-RíoClaro,nearVeredaTulipán, 935m,Cogollo & Estrada 202(MO);Mun.SanLuis,sectorRíoSamaná-Río Claro,nearVeredaLaJosefina,790m,Cogollo & Estrada 299(MO);Mun. SanLuis,ElReposo,km152ofHwy.Medellin-Bogotá,sectorRíoSamanáRíoClaro,790m,Cogollo & Brand 427(JAUM,MO);Mun.SanLuis,near Aquitania,8kmfromHwy.Medellín-Bogotá,830m,Cogollo et al. 3796 (COL, JAUM,MO);Mun.SanCarlos,VeredaPatioBonito,AltoElCerrón,linea detransmisiónSanCarlos-SanMarcos(entrelastorres22–23),1100m, E. Correa et al. 93 (U);Mun.SanCarlos,roadofVeredaMiraflorestothe CorregimientoAltodeSamanáNorte,800–900m,Fonnegra et al. 3109 (HUA,MO,U);Mun.Cáceres,VeredaMadreseca,ReservaRegionalNatural BajoCauca-Nechí,830–850m,Giraldo 190(JAUM);Mun.SanLuis,Vereda LaJosefina,Hwy.Medellin-Bogotá,800m,Hoyos M. & Hernández 261 (JAUM,MO);Mun.SanRafael,VeredaElCharco,1010–1200m,Montes Guarín 200(COL). Notes—Guatteria antioquensis, restricted to the Colombian department of Antioquia, is easily recognizable by a combination ofitsdenseandlong-persistingindumentoferecthairs(hairs simpleorinbundlesof2–4)onthelowersideoftheleaves and a densely hairy connective shield, which is so typical for all species related to G. schomburgkiana.Anotherpeculiarityis the hairy stipes, which in almost all other species in the genus aredevoidofanyindument. Guatteria antioquensis looks somewhat like G. citriodora because of its indument, but from that species it clearly differs bymuchlongerpedicels,bothfloweringandfruiting(15–35 vs5–17mm),amuchhighernumberofmonocarps(c.50 vs2–10),athinnerfruitwall(0.1vs0.5mmthick)andpitted insteadofsmoothseeds. 13. Guatteria araracuaraeMaas&Westra,sp. nov.—Map5 Guatteria hispida affinissedfoliisvenamarginalibenedistincta(necnonvenis secundariisarctetantumconjunctisdiffert.—Typus:Ronderos & Rosselli 58 (holoCOAH),Colombia,Amazonas,cuencadelmedioCaquetá,región deAraracuara,ComunidaddePeñaRoja,bosquemadurodelasterrazas altasdelRíoCaquetá,160m,17July1996. Guatteria sp. B MurilloA.&Restrepo(2000)119,f.36;Maas&Westra (2011)141,f.23. Tree3.5–16mtall,2.5–7cmdiam;youngtwigsdenselyto rather densely covered with long-persistent, erect, stiff hairs toc.3mmlong.Leaves:petiole3–5mmlong,2–4mmdiam; laminanarrowlyelliptictonarrowlyobovate,15–28by4–7cm (leafindex3–5),chartaceous,verydenselyverruculose,dull, brown to greyish brown above, brown below, sparsely covered with erect hairs mainly along primary vein above, densely coveredwitherect,stiffhairstoc.3mmlongbelow,baseobtuse, apexacuminate(acumen10–20mmlong),primaryveinimpressed above, secondary veins distinct, 16–22 on either side of primary vein, impressed above, forming a distinct marginal vein,atasmallestdistanceof2–4mmfromthemargin,tertiary veins inconspicuous and hardly visible above, percurrent toreticulate.Flowers solitary in axils of leaves or on leafless branchlets;floweringandfruitingpedicels(3–)11mmlong, 1–2 mm diam, densely covered with long-persistent, erect, stiffhairstoc.3mmlong,articulatedatc.0.1fromthebase, bracts not countable in available material, uppermost bract ovate-elliptic,toc.11mmlong;flowerbudsovoid;sepalsfree, broadlyovate-triangular,8–10by5–8mm,appressed,apex acuminate,outersidedenselycoveredwitherect,brownhairs; petalsgreenorwhiteinvivo,oblong-ovatetoelliptic,8–15by 4–10mm,outersidedenselycoveredwithappressed,wooly hairs;stamens1.5–2mmlong,connectiveshieldpapillate. Monocarps 5–10, purplish black in vivo, black in sicco, ellipsoid, 15–20by8–9mm,denselytosparselycoveredwitherect, stiffhairstoc.2mmlong,apexroundedtoapiculate(apicule <1mmlong),wall0.5–0.7mmthick,stipes10–20by1mm, denselycoveredwitherecthairs.Seedellipsoid,c.15by7mm, dark brown, horizontally and transversely grooved, raphe not distinctfromrestofseed. Distribution—AmazonianColombia(Amazonas,Caquetá). Habitat&Ecology—Innon-inundatedforest,onclayey tosandysoil.Atelevationsofupto300m.Flowering:July, November;fruiting:January,July. Vernacularnames—Colombia:Buruchicu(Muinanename) (Van Andel et al. 160),Carguero(Muinanename)(Cárdenas L. et al. 4064),Dujïku(Huitotoname)(Cárdenas L. et al. 4064), Jïdïra(Huitotoname)(Cárdenas L. et al. 4064),Ñaajeku(Muinanename)(Murillo A. et al. 510). Other specimens examined. ColomBia, Amazonas,PuertoSantander, Monochoa,Cárdenas L. et al.4064(COAH,U);RíoCaquetá,Leticia,Villa Azul,200–270m,Duque & Posada 4189(U);Araracuara,AeropuertoRestrepo,rightmarginofRíoCaquetá,QuebradaBocaduche,Murillo A. et al. 510(COAH,COL,U);VillaAzul,RíoCaquetá,Van Andel et al. 161(COAH, U).Caquetá,Mun.Solano,RíoMesay,RaudalMasaca,300m,Cárdenas L. et al. 6772(COAH,U). Notes—Guatteria araracuarae looks similar to G. hispida astotheleafshapeandthedenseindumentofstifferecthairs. Itclearlydiffersfromthelatter,however,bythepresenceofa distinct marginal leaf vein, while in G. hispida the leaves do not showamarginalvein. MurilloA.&Restrepo(2000)attributedthecollectionStein et al. 4002(MO,U)fromLoreto,Peru,tothisspeciesaswell.Actually, it is the type collection of G. grandipesMaas&Westra(2011). 14. Guatteria arenicolaMaas&Erkens—Plate2a,b;Map5 Guatteria arenicolaMaas&ErkensinErkensetal.(2008)471,f.1,pl.1. —Type:Maas et al. 8980(holoU;isoNY),Brazil,Acre,km6ofroadfrom CruzeirodoSultoBoaFé(BR-307),RamaldosCarobas,c.200m,16 Oct.2001. Shrubortree2–30mtall,5–40cmdiam,trunkfissured,slash orange(inmaterialfromAcre);youngtwigsdenselytosparsely coveredwithappressedhairs,soonglabrous.Leaves: petiole 2–7mmlong,c.1mmdiam;laminaelliptictonarrowlyelliptic, 5–9by1–3cm(leafindex2–3.6),coriaceoustochartaceous, not verruculose, dull, dark brown, brownish black to greyish black above, somewhat paler brown below, glabrous, except for the hairy primary vein above, rather densely to sparsely covered with appressed hairs below, base attenuate to acute, apexshortlyandbluntlyacuminate(acumen1–3mmlong)to rounded, primary vein slightly impressed to flat above, secondaryveinsindistincttodistinct,6–13oneithersideofprimary vein, flat to raised above, smallest distance between loops and margin2–3mm,tertiaryveinsraisedabove,reticulate.Flowers solitaryinaxilsofleaves;pedicelsstronglyrecurved(inmaterialfromAcre)toerect,10–13mmlong,c.1mmdiam,fruiting pedicels15–20mmlong,c.2mmdiam,articulatedat0.2–0.5 fromthebase,bracts4–5,the2upperonesnarrowlyobovate, 4–6by1.5–2mm,the3lowerones1–1.5mmlong,denselyto sparselycoveredwithappressedhairs;flowerbudsdepressed ovoid,slightlypointed;sepalsfreeorbasallyconnate,broadly ovate-triangular,3–6by3–5mm,initiallyappressed,soon spreading and apically reflexed, outer side densely to rather denselycoveredwithappressedhairs;petalsgreen,maturing yellowinvivo,ovate,5–18by3–12mm,outersidedensely coveredwithappressedtoerecthairs;stamens1–1.5mmlong, connectiveshieldpapillatetohairy.Monocarps10–40,green, maturingblackinvivo,brownishblackinsicco,ellipsoid,7–13by 3–7mm,sparselycoveredwithappressedhairs,soonglabrous,
36 Blumea–Volume60/1–3,2015 Plate 2a,b.Guatteria arenicolaMaas&Erkens.a.Fruitingbranches;b.fruits.–c.Guatteria beckiiMaas&Westra.Fruitingbranch.–d.Guatteria argenteaErkens&Maas.Topofbranchletshowingsilveryindument;e.fruits.–f,g.Guatteria blainii(Griseb.)Urb.f.Flower;g.fruit(a,b:Rodriguez R.3095; c: Solomon 16789;d,e:Maas et al. 10525;f,g:Maas & Mejia 6443).—Photos:a,b:E.RodriguezR.;c:J.Solomon;d–g:P.J.M.Maas. c b d a g e f
37 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria apexroundedtobluntlyapiculate(apiculum<0.5mmlong), wall0.3–0.5mmthick,stipes0–2by1.5–2mm.Seed ellipsoid,10–11by3–5mm,palebrown,smooth,raphenotdistinct fromrestofseed. Distribution—AmazonianEcuador(Zamora-Chinchipe), Peru(Cajamarca,SanMartín),Brazil(Acre). Habitat&Ecology—Incampinaorcampinaranavegetation, orinprimaryforest,onwhite,oftenhumoussand.Atelevations of200–2170m.Flowering:June,SeptembertoNovember; fruiting:May,December. Vernacularnames—Notrecorded. Notes—Guatteria arenicola is a small-leaved species from the white sand campina or campinarana vegetations of Acre, Brazil,andforestsonwhitesandsinPeru(SanMartín).Itis quite noteworthy by its tiny, prominently veined, coriaceous leaves, its very small flowers, which often have strongly recurvedpedicels,andstronglyrugoseseeds.Anothercharacter of this species is the presence of orange slash, a feature not seen so far in any other representative of the genus Guatteria. Guatteria arenicola has been compared with G. rupestris, a specieswhichisknownfromthecerradosinMinasGerais,Brazil. Theybothsharesomeleafcharacteristics,butinG. rupestris, the leaves mostly have a rounded to even emarginate apex, whereasthebasalmarginsarerevolute. 15. Guatteria argenteaErkens&Maas—Plate2d;Map5 Guatteria argentea Erkens&MaasinErkensetal.(2008)473,f.2.—Type: Monsalve B.1262 (holoU;isoCUVC,MO),Colombia,ValledelCauca, BajoCalima,ConcesiónPulpapel/Buenaventura,100m,13Nov.1986. Tree6–12mtall,diamnotrecorded;youngtwigsdensely coveredwithappressed,silveryhairs,soonglabrous.Leaves: petiole8–15mmlong,2–3mmdiam;laminaobovate,rarely narrowlyso,9–12by3.5–6cm(leafindex2–2.5),coriaceous, densely verruculose, slightly shiny, dark brown to greyish black above, brown below, glabrous above, densely covered with appressed, silvery hairs below, base attenuate, apex shortly acuminate(acumen1–5mmlong),primaryveinimpressed above,keeledbelow,secondaryveinsdistinct,9–13oneither side of primary vein, raised to flat above, smallest distance betweenloopsandmargin3–4mm,tertiaryveinsslightlyraised toflatabove,reticulate.Flowerssolitaryinaxilsofleaves; pedicels 5–9 mm long, 1–2 mm diam, fruiting pedicels 10–15 mmlong,c.2mmdiam,denselytoratherdenselycoveredwith appressed,silveryhairs,articulatedat0.3–0.5fromthebase, bracts3–4,soonfalling,theupperonec.4by3mm;flower budsdepressedovoidtoovoid;sepalsfree,broadlyovatetriangular,3–5by4–6mm,appressed,outersidedensely coveredwithappressed,silveryhairs;petalsyellowishgreenin vivo,elliptictoobovate,9–13by5–8mm,outersidedensely coveredwithappressed,silveryhairs;stamens1–1.5mmlong, connectiveshieldhairy.Monocarps 10–15, green to yellow invivo,browntoblackishinsicco,ellipsoid,12–16by5–8 mm, sparsely covered with appressed hairs, apex apiculate (apiculum<0.5mmlong),wall1–1.5mmthick,stipes2–6by c.2mm.Seedellipsoid,10–12byc.5mm,shiny,darkbrown, pitted,raphenotdistinctfromrestofseed. Distribution—Colombia(Antioquia,ValledelCauca). Habitat&Ecology—Inforest(‘bosquepluvialtropical’).At elevationsof50–200m.Flowering:February,August,October toDecember;fruiting:February,August,November,December. Vernacularnames—Notrecorded. Notes—Guatteria argentea is very easily recognizable by its silvery indument on many parts of the plant and by its obovate, denselyverruculoseleaveswithanattenuatebase. Thetypeofthepresentspecies(Monsalve B.1262)hasbeen usedbyErkensetal.(2007a,b)foraphylogeneticstudyon Guatteria.However,inthesestudiesthisspecimenwasassigned to G. heterotricha R.E.Fr.ItwasplacedsistertoG. rotundata to whichitlooksquitesimilar.Guatteria rotundataisfromPanama (PanamáandSanBlas)andthetwospeciessharetheobovate leaves.Guatteria argentea differs by its dense silvery indument of the young twigs and lower side of the lamina and by its more denselyverruculoseleaves. 16. Guatteria atabapensisAristeg.exD.M.Johnson&N.A. Murray—Fig.13c,15;Map5 Guatteria atabapensis Aristeg.exD.M.Johnson&N.A.Murray(1990)599; Steyerm.etal.(1995)444;MurilloA.&Restrepo(2000)92.—Type:Wurdack & Adderley 42759(holoNY;isoF,G,RB,US,VEN),Venezuela,Amazonas,Dep.Atabapo,RíoAtabapo,marginofSabanaCumare,rightbank ofCañoCumare(20kmaboveSanFernandodeAtabapo),125–140m, 3June1959. Treeorshrub,3–6(–28)mtall,to23cmdiam;youngtwigs sparselycoveredwithappressedhairs,soonglabrous.Leaves: petiole5–10mmlong,1–3mmdiam;laminaelliptictoovateor narrowlyso,8–16by4–6cm(leafindex1.8–2.8),coriaceous, not or less frequently verruculose, dark brown to greyish green above, brown to pale brown below, glabrous above, sparsely covered with appressed hairs to almost glabrous below, base obtuse,sometimesacute,apexacuminate(acumen5–15mm long),primaryveinimpressedabove,secondaryveinsindistinct, 10–14oneithersideofprimaryvein,raisedabove,smallest distancebetweenloopsandmargin4–7mm,tertiaryveins raisedabove,reticulate.Flowers in 1–2-flowered inflorescences inaxilsofleaves;floweringandfruitingpedicels4–8mmlong, 1–2 mm diam, rather densely covered with appressed hairs, articulatedatc.0.3fromthebase,bracts4–6,soonfalling, broadlyovate,1–3mmlong;flowerbudsdepressedovoid; sepalsfree,broadlyovate-triangular,4–5by4–5mm,spreading,outersidedenselycoveredwithappressedhairs;petals greenishyelloworyellowinvivo,ovatetoobovate,8–23by 4–12mm,outersidedenselytoratherdenselycoveredwith appressedhairs;stamensc.1mmlong,connectiveshieldpapillate.Monocarps 10–25, green, maturing black in vivo, black to brownishinsicco,ellipsoid,7–10by4–6mm,sparselycovered withappressedhairs,soonglabrous,apexapiculate(apiculum 0.1–0.2mmlong),wall0.2–0.3mmthick,stipes1–5by1mm. Seedellipsoid-fusiform,7–9by4–5mm,darkbrown,pittedto transverselygrooved,rapheimpressed. Distribution—AmazonianColombia(Vaupés),Venezuela (Amazonas,Bolívar). Habitat&Ecology—Inshrubislandsinsavannasandin forest,onwhitesands.Atelevationsof0–150(–560)m.Flowering:JanuarytoMay;fruiting:January,May,July,September. Vernacularnames—Colombia:Carguerodehojanegra (M. Sánchez S. et al. 802, 879),Quïbojïdujecu(Muinanename) (M. Sánchez S. et al. 802, 879). Notes—Guatteria atabapensis is well recognizable by its coriaceous, obscurely veined leaves, with submarginal loops quitefarremovedfromthemargins(4–7mm),incombination withveryshortpedicelsandshortstipes.Itlooksquitesimilar to G. maguirei,whichisalsoconfinedtosavannas,butitdiffers fromthelatterbymuchshorterpedicelsandlargerleaves. SomesterileColombiancollectionsidentifiedbyMurilloA.& Restrepo(2000)asG. atabapensisdeservefurtherattention. Theydifferfromtheremainderbyverruculoseleaves,butare otherwisequitesimilar.Itconcernsthefollowing: ColomBia, Amazonas,RíoCaquetá,IslaMariñame,M. Sánchez S. et al. 802(COAH,U);RíoCaquetá,IslaMorocoy,M. Sánchez S. et al. 879(COAH, U).Caquetá, Sierra de Chiribiquete, 560 m, P. A. Palacios et al. 2627(U).
38 Blumea–Volume60/1–3,2015 Fig. 15 Guatteria atabapensisAristeg.exD.M.Johnson&N.A.Murray.Floweringbranch(Wurdack & Adderley 42759;isotypeUS).
39 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria 17. Guatteria australis A.St.-Hil.—Fig.16;Plate1h;Map6 Guatteria australis A.St.-Hil.(1825)37;R.E.Fr.(1939)306,t.1.—Guatteria australisvar.glabrataMart.(1841)26.—Type:A.C.F.P. de Saint-Hilaire 1452 (holoP3sheets),Brazil,SãoPaulo,Itararé(‘bordsofTararé’),Jan. 1817. Guatteria gomeziana A.St.-Hil.(1825)36;R.E.Fr.(1939)319.—Type: A.F.C.P. de Saint-Hilaire 560 (holo P2sheets;isoK),Brazil,MinasGerais, RioPiracicaba(‘ItajurudeSãoMigueldeMatoDentro’),Jan.–Feb.1817. Guatteria lutea A.St.-Hil.(1825)37;R.E.Fr.(1939)322.—Guatteria flava A.St.-Hil.exMart.(1841)27,nom.superfl.—Type:A.F.C.P. de Saint-Hilaire 304 (holoP),Brazil,MinasGerais,OuroPreto(‘entreVilaRicaetAntônio Pereira’),Jan.1817. Uvaria hirsuta Vell.(1829)224;(1831)t.124,nom.illeg.,nonUvaria hirsuta Jack(1820)46.—Type:Velloso’splate124(lecto,selectedbyMaaset al.2011). Uvaria monosperma Vell.(1829)224;(1831)t.123,nom.illeg.,nonUvaria monosperma Lam.(1785)596.—Type:Velloso’splate123(lecto,selected byMaasetal.2011). Guatteria hilarianaSchltdl.(1834)324;R.E.Fr.(1939)396. — Guatteria hilariana f. latifolia Schltdl.(1834)324.—Guatteria hilariana var. latifolia (Schltdl.)Mart. (1841)35.—Type:Sellow s.n. (holoB;iso HAL),Brazil, withoutlocalityanddate. Guatteria hilarianaSchltdl. f. angustifolia Schltdl.(1834)324;R.E.Fr.(1939) 397.—Guatteria hilarianavar. angustifolia (Schltdl.)Mart.(1841)35.— Type:Sellow 5943 p.p. (holoB2sheets;iso HAL),Brazil,SãoPaulo, Santos,Apr.–July1829. Guatteria psilopus Mart.(1841)27,t.7,f.1;R.E.Fr.(1939)339.—Type: Luschnath s.n.(holoBR),Brazil,RiodeJaneiro,Teresópolis,Serrados Orgãos,June1834. Guatteria acutifloraMart.(1841)29.—Type: Wied zu Neuwied s.n.(lectoBR, selectedbyMoraes2013;isolectoBR,MEL),Brazil,Bahia,NovaAlmeida (‘crescitinsilvisadVillamNovamdeAlmeida’),Mar.1816. Guatteria nigrescens Mart.(1841)31;R.E.Fr.(1939)319.—Guatteria nigrescens Mart. var. oblongifolia Mart.(1841)31.—Type:Martius s.n. (lecto M,selectedbyFries1939),Brazil,SãoPaulo,Guaratinguetá,Dec.1817. Guatteria densicoma Mart.(1841)32;R.E.Fr.(1939)309,f.3b,t.3.—Type: Martius 711 (holoM; isoB,BR4sheets,F,G,HAL,K,NY,P),Brazil,Bahia, Ilhéus,FazendaAlmada,Dec.1818. Guatteria klotzschiana Mart. (1841)32;R.E.Fr.(1939)327,f.5d–f.—Cananga klotzschiana (Mart.)Warm.(1873)145.—Type:Schott s.n. (holoBR 2sheets;isoB,F,US),Brazil,RiodeJaneiro,‘propeTocaia’. Guatteria odontopetala Mart.(1841)33,t.11;R.E.Fr.(1939)407.—Type: Pohl 2964 (lectoM,selectedbyFries1939;isolectoB,BR,F,G),Brazil, MinasGerais,ManoelPereira. Guatteria hookeri A.St.-Hil.&Tul.(1842)132;R.E.Fr.(1939)342,f.6c,d.— Type:Gardner 306 p.p. (holoP3sheets;isoBM,E,F,G,K3sheets,LZ, NY,OXF,S),Brazil,RiodeJaneiro,Teresópolis,OrganMountains,banks oftheRiverPaquequer,Mar.1837. Guatteria neglecta R.E.Fr.(1939)308,f.3a,t.2.—Type:Dusén 6780 p.p. (holoS2sheets; iso F,K,L,MO,NY2sheets,P,U),Brazil,Paraná,Morretes,Jacareí(‘Jacarehý’),30Sept.1908. Guatteria polycarpa R.E.Fr.[subsp.polycarpa](1939)310,t.4.—Type: Dusén 7414 (holoS; iso F,K,L,MO,NY,P),Brazil,Paraná,Piraquara, RoçaNova-Banhado,30Dec.1908. Guatteria polycarpa R.E.Fr.subsp.drupacea R.E.Fr.(1939)311,f.3c. — Type:J.G. Kuhlmann RB 19677 (holoS;isoRB),Brazil,SãoPaulo,without locality,26Sept.1922. Guatteria glabrescens R.E.Fr.(1939)311,t.5. — Type:J.G. Kuhlmann RB 4483 (holoS;isoF,G,RB2sheets),Brazil,RiodeJaneiro,Reserva FlorestaldafábricaAlliança,11Jan.1922. Guatteria fruticosa R.E.Fr.(1939)312,t.6. — Type:Löfgren 531 (holoS), Brazil,SãoPaulo,SãoJosédosCampos,14Dec.1909. Guatteria dusenii R.E.Fr.(1939)313,t.7. — Type:Dusén 13752 (holoS; iso L,MO),Brazil,Paraná,Guaratuba,July1911. Guatteria dusenii R.E.Fr. var. subglabra R.E.Fr.(1939)313.—Type:Dusén 4497 (holoS),Brazil,Paraná,Paranaguá(‘PortoD.PedroII’),19Apr.1904. Guatteria paranensis R.E.Fr.(1939)314,t.8.—Type:Dusén 6780 p.p. (holoS),Brazil,Paraná,Morretes,Jacareí(‘Jacarehý’),30Sept.1908. Guatteria parvifolia R.E.Fr.(1939)314,t.9. — Type:F.C. Hoehne SP 28405 (holoS;isoB,F,HB,K,MO,SP,SPF,US),Brazil,SãoPaulo,SãoPaulo, ParquedoEstado,28Oct.1931. Guatteria parvifolia R.E.Fr. var.vestita R.E.Fr.(1939)315. — Type:Sellow 211 (holoB;isoK),Brazil,SãoPaulo,withoutlocalityanddate. Guatteria salicifolia R.E.Fr.[var.salicifolia](1939)316,t.10,f.3d.—Type: Glaziou 7508(holoB;isoC,K,MG,P),Brazil,RiodeJaneiro,Riode Janeiro,Tijuca,21Dec.1874. Guatteria salicifolia R.E.Fr. var. erosa R.E.Fr.(1939)316.—Type:Dusén s.n. (holoS; iso F),Brazil,Paraná,Morretes,Jacareí(‘Jacareý’),1914. Guatteria curvinervia R.E.Fr.(1939) 317,t.11.—Type:Frazão 8667 (holo S;iso RB),Brazil,SãoPaulo,Barretos,Dec.1917. Guatteria mosenii R.E.Fr.(1939)321. — Type:Mosén 4002 (holoS),Brazil, SãoPaulo(‘SerradoCaracol’),15Dec.1875. Guatteria asterantha R.E.Fr.(1939)323,t.12.—Type:Mexia 5485 (holoS; isoBM,F2sheets,G,K,NY,U,US),Brazil,MinasGerais,Viçosa(‘road toCajurý,Canella’),750m,7Jan.1931. Guatteria acutipetala R.E.Fr.(1939)324,t.13.—Type:Ule 3962 (holoB), Brazil,RiodeJaneiro,Teresópolis,Jan.1897. Guatteria reflexa R.E.Fr.(1939)329.—Type:Glaziou 5725 (holoS;isoB, C,F,K,P,R,RB),Brazil,RiodeJaneiro,Niterói,MorrodaViração,28 July1870. Guatteria silvatica R.E.Fr.(1939)330.—Type:Peckolt 151 (holoBR),Brazil, RiodeJaneiro,Cantagalo(‘CantaGallo’),1857. Guatteria blanchetiana R.E.Fr.(1939)331,t.14.—Type:Blanchet 2114 (holoG6sheets;isoB,BM,F,FI,K2sheets,MO,NY,P2sheets,W), Brazil,Bahia,Ilhéus,FazendaAlmada,1835. Guatteria dimorphopetala R.E.Fr.(1939)333,f.5c.—Type:Sellow 175 (holo B),Brazil,betweenVitoria(EspíritoSanto)andBahia. Guatteria clavigera R.E.Fr.(1939)334,f.5g,h.—Type:Koscinsky 214 (holo S; isoSP),Brazil,SãoPaulo,SãoPaulo,SerradaCantareira,15Dec.1933. Guatteria riedelianaR.E.Fr.(1939)339,f.6a,b. —Type:Riedel 424 (holo S;isoLE,NY,U),Brazil,Bahia,Castelnovo,Nov.1821. Guatteria penduliflora R.E.Fr.(1939)341,t.15.—Type:Allemão e Cysneiro s.n. (holoG2sheets),Brazil,RiodeJaneiro,withoutlocality,1860. Guatteria minarum R.E.Fr.(1939)346,t.16.—Type:Mexia 5130 (holo S;isoBM,C,F2sheets,G,K,NA,NY,U),Brazil,MinasGerais,Viçosa (‘FazendaParaiso’),1Oct.1930. Guatteria hilarianaSchltdl. var.cuneata R.E.Fr.(1939)397.—Type:Burchell 3366 (holoK;isoK,P),Brazil,SãoPaulo,Santos,Montserrat,NEpartof islandSãoVicente,3Nov.1826. Guatteria hilarianaSchltdl. var.verruculosa R.E.Fr.(1939)397.—Type: Mosén 3337 (holoS;isoB,P,US),Brazil,SãoPaulo,Santos,Baturoca, 25Jan.1875. Guatteria sordida R.E.Fr.[var.sordida](1939)532.—Guatteria sordida var.stenopetala R.E.Fr.(1939)533,t.39p.p.—Type:Lund 959 (synC), Brazil,SãoPaulo,Mogi(‘Mugy’),Nov.1833andLund s.n. (synC),Brazil, SãoPaulo,withoutlocalityanddate. Guatteria sordida R.E.Fr. var.lancifolia R.E.Fr.(1939)533,t.39p.p.—Type: Riedel 1689 p.p. (holoS;isoK),Brazil,SãoPaulo,Mogi(‘propeMuji’), Nov.1833. Guatteria sordida R.E.Fr. var.ovalis R.E.Fr.(1939)533,t.39p.p.—Type: Riedel 1689 p.p. (holoS;isoB,K,P),Brazil,SãoPaulo,Mogi(‘prope Muji’),Nov.1833. Guatteria tenuis R.E.Fr.(1939)535,t.40.—Type:Mexia 5482 (holoS;iso BM,F,G,K,U),Brazil,MinasGerais,Viçosa,FazendadoDeserto,690m, 5Jan.1931. Tree1–26mtall,6–20cmdiam;youngtwigsdenselycovered withappressed,rarelyerecthairs,soonglabrous.Leaves: petiole2–8mmlong,1–3mmdiam;laminanarrowlyelliptic toelliptic,5–20by1.5–7cm(leafindex2–6.7(–9.5)),chartaceous, not verruculose, shiny, blackish green to blue above, pale brown to brown below, sparsely covered with appressed hairs to glabrous above, densely covered with appressed hairs to glabrous below, base acute to attenuate, apex acute to acuminate(acumen5–15mmlong),primaryveinimpressed above,secondaryveinsdistinct,8–20oneithersideofprimary vein, raised above, smallest distance between loops and margin2–4mm,tertiaryveinsraisedabove,reticulate.Flowers solitaryinaxilsofleavesoronleaflessbranchlets;flowering andfruitingpedicels15–60mmlong,1–3mmdiam,densely covered with appressed, brown hairs to glabrous, articulated at0.2–0.3fromthebase,bracts3–5,soonfalling,ellipticto narrowlyelliptic,4–7(–20)mmlong;flowerbudsbroadlyovoid; sepalsfree,broadlytriangular,5–10by5–7mm,reflexed, outersideglabrous;petalscreaminvivo,narrowlyellipticto elliptic,10–38by5–20mm,outersidedenselycoveredwith appressed,brownhairstoglabrous;stamensc.2mmlong,connectiveshieldhairy.Monocarps15–45,green,maturingredto purple-blackinvivo,blackinsicco,ellipsoid,6–15by5–7mm, glabrous,apexapiculate(apiculumc.0.5mmlong),wallc.0.5
40 Blumea–Volume60/1–3,2015 mmthick,stipes10–30by1mm.Seedellipsoid,6–10by5–7 mm,darkbrown,rugulose,raphenotdistinctfromrestofseed. Distribution—Brazil(Bahia,EspíritoSanto,Goiás,Minas Gerais,Paraná,RiodeJaneiro,RioGrandedoSul,SantaCatarina,SãoPaulo). Habitat&Ecology—Innon-inundatedandperiodicallyinundated, often Atlantic forest, or cerrado, on clayey to sandy soil.Atelevationsof0–2000m.Flowering:throughouttheyear; fruiting:throughouttheyear. Vernacularnames—Brazil:Cambuí(P. Rosa RB 81409), Imbira(H.C. Lima 3405),Pindaíba. Note—Guatteria australis is very variable, generally being characterizedbymedium-sizedleaves(5–20by1.5–7cm)with Fig. 16 Guatteria australis A.St.-Hil.a.Floweringhabit;b–d,f,g,i,k,l.leaves;e.leafapex;h,j.leafbase;m.flower;n.fruit(allfromLobãoetal.(2011)f.1). cb d a g ih j k m n l e f 2 cm 2 cm 5 mm 2 cm 5 mm 5 mm 1 mm
41 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria acute to acuminate apex and flowering pedicels often around 50 mmlong.Theindumentisalsohighlyvariable,individualswith glabrousbranchesandleaves(e.g.,Dusén 13746 fromGuaratuba,Paraná)occurringinthesameareaasindividualswith a dense hair cover on branches and the lower side of leaves (e.g.,Barbosa 556,alsofromGuaratuba,Paraná).Guatteria acutiflora was recently added to the synonymy of G. aus tralis (Lobãoetal.2012).Guatteria acutiflora supposedly differs fromthelatterbyroundedleafbases(VonMartius1841),but, asLobãofoundaroundedleafbaseinseveralspecimensof G. australisaswell,sheseesnojustificationforkeepingthe twospeciesapart.ThenameG. acutifloraDC(DeCandolle 1817)iscitedinreferencetoG. acutifolia Dunal and is an orthographicvariant(art.60.1ICBN). 18. Guatteria auyantepuiensisMaas&Westra,sp. nov. — Fig.17,13d;Map7 Foliis sparse verruculosis, coriaceis, apice obtusis vel breviter acuminatis, floribusquebreviterpedicellatisdistincta.—Typus:F. Cardona 2619(holo US;isoVEN,notseen),Venezuela,Bolívar,Auyantepui(‘CerroAuyan’), Alto Caroni, en pequeños bosques abrigados, lugares rocosos, 1600 m, Jan.1949. Treec.6mtall,diamnotrecorded;youngtwigsdenselycovered withappressedhairs,soonglabrous.Leaves:petiolec.5mm long,2mmdiam;laminaelliptic,9–12by3–4cm(leafindex 2.5–3),coriaceous,sparselyverruculose,shiny,darkbrown above, brown below, glabrous above, sparsely covered with appressed hairs below, base acute to attenuate, apex obtuse toshortlyacuminate(acumentoc.5mmlong),primaryvein impressedabove,secondaryveinsdistinct,8–10oneitherside of primary vein, raised above, smallest distance between loops andmargin2–4mm,tertiaryveinsraisedabove,reticulate. Flowerssolitaryinaxilsofleaves;pedicelsc.5mmlong,2 mm diam, densely covered with appressed hairs, articulated at c.0.3.fromthebase,bractsc.6,soonfalling,notseen;flower budsnotseen;sepalsbasallyconnate,broadlyovate-triangular, 4–5by5–6mm,appressed,outersidedenselycoveredwith appressedhairs;petalscolourinvivounknown,oblong-elliptic, 10–12 by 5–6 mm, outer side densely covered with erect and appressedhairs;stamensc.2mmlong,connectiveshield papillate.Monocarps and seednotseen. Distribution—Venezuela(Bolívar). Habitat&Ecology—Innon-inundatedforest,onrockyoutcrops.Atanelevationofc.1600m.Flowering:January;fruiting: notknown. Vernacularnames—Notrecorded. Note—Guatteria auyantepuiensis has been recorded by JohnsonandMurray(seeSteyermarketal.1995)asGuatteria sp. A.Itbelongstosect.Mecocarpus and shares some features with G. meliodora(e.g.,theshinyandsparselyverruculoseand obtusetoshortlyacuminateleaves),buttheleavesaremuch smaller(9–12vs22–35cmlong),havefewersecondaryveins (8–12vs18–20)andtheflowershaveshorterpedicels(5vs 10–20mmlong).Moreover,theleavesarequitedistinctbythe stronglyreticulatetertiaryvenation. 19. Guatteria ayangannaeScharf&Maas—Map7 Guatteria ayangannae Scharf&MaasinScharfetal.(2005)565,f.2.—Type: Clarke et al. 9819(holoU2sheets;isoBRG,NY4sheets),Guyana, Potaro-SiparuniRegion,UpperPotaroRiver,nearcamp,MtAyanganna, 650m,3July2001. Tree5–7mtall,c.10cmdiam;youngtwigsdenselycovered witherect,orange-brownhairstoc.1.5mmlong,soonglabrous. Leaves:petiole7–14mmlong,1–2mmdiam;laminaellipticto obovate,10–17by4–6cm(leafindex2.1–2.9),coriaceous, scabridulous particularly below, pale to creamy brown above, brown below, glabrous above, densely covered with appressed, orange-browntosilveryhairstoc.1.5mmlongbelowwhen young, soon becoming sparsely covered with appressed hairs toc.0.5mmlong,thenbecomingglabrous,baseacute,apex acuminate(acumen5–10mmlong),primaryveinimpressed above,secondaryveinsdistinct,9–14oneithersideofprimary vein, flat above, smallest distance between loops and margin 2–4mm,tertiaryveinsflatabove,reticulate.Flowers in 1–2-floweredinflorescencesinaxilsofleaves;pedicels5–15mmlong, c.2mmdiam,fruitingpedicelstoc.15mmlong,c.3mm diam, densely covered with appressed, orange-brown hairs, articulatedat0.3–0.5fromthebase,bracts5–6,thelowerbracts soonfalling,elliptic-ovate,c.2mmlong,the2upperonesoften persisting till flowering time, often foliaceous, elliptic to narrowly elliptic,7–40mmlong,to15mmwide;flowerbudsbroadly ovoid,slightlypointed;sepalsbasallyconnate,triangular,9–10 by 5–6 mm, often reflexed, apex strongly acumi nate, outer side denselycoveredwithappressed,orange-brownhairs;petals greenishyelloworyellowinvivo,elliptictoovate,14–18by 6–10 mm, outer side densely covered with appressed, orangebrownhairs;stamensc.2mmlong,connectiveshieldhairy. Monocarps2–8,greentoreddishbrowninvivo,blackinsicco, narrowlyellipsoid,15–23by4–6mm,sparselycoveredwith appressedhairstoc.0.5mmlongtoglabrous,apexnipple- Brazil Colombia Peru Venezuela s ta Rica Bolivia G u y a n a Suriname French Guiana Ecuador Map 6 Distribution of Guatteria australis (p),G. beckii (£ ),G. beniensis (n), G. brevipetiolata (●)andG. campestris (u). Brazil Colombia Peru Venezuela Panama Costa Rica M e x i c o Guatemala Nicaragua Honduras Bolivia Guyana Suriname French Guiana P u e r t o R i c o Haiti Lesser Antilles Ecuador Map 7 Distribution of Guatteria auyantepuiensis (●),G. ayangannae (£), G. blepharophylla (■),G. campinensis (p)andG. candolleana (u).
48 Blumea–Volume60/1–3,2015 Flowering:throughouttheyear;fruiting:January,ApriltoJuly, November,December. Vernacularnames—Cuba:Pimientamalagueta.Dominican Republic:Yaya(Mera 2054),Yayafrancesa(J. Jiménez 4627). Fieldobservations—InZanoni et al. 30964 from the DominicanRepublictheflowersare‘bananascented,withascent of Cananga odorata’. Notes—Guatteria blainii, the only species of Guatteria occurringintheGreaterAntilles,isaquitevariablespeciestobe recognized by relatively small and shiny leaves with the secondary veins strongly raised on the upper side, and by sessile or shortlystipitatemonocarps. ThereisalotofvariationintheseedstructureofG. blainii, much morethaninotherspeciesofthegenus.Theseedsvaryfrom smooth,longitudinallygrooved,totransverselygrooved. Leavesofsterilematerialaredifferentfromleavesoffertile material, they are larger and long-acuminate, with a petiole rangingfrom7–12mmby2mmandthelaminafrom8–11cm by4–5cm. 24. Guatteria blepharophyllaMart.—Map7 Guatteria blepharophylla Mart.(1841)38;Maas&Westra(2011)115,f.1a, 2,3a,pl.1a,b.—Guatteriopsis blepharophylla(Mart.)R.E.Fr.(1934)110, t.6;Maasetal.(2007)644.—Type:Martius s.n.(lectoM,selectedby Erkens&Maas2008),Brazil,Amazonas,RioNegro,Coarí,Nov.1819. Annona sessiliflora Benth.(1853)8.—Guatteria sessiliflora(Benth.)Saff. (1914)6.—Guatteriopsis sessiliflora(Benth.)R.E.Fr.(1934)109.—Type: Spruce 1668(holoK;isoB,BM,FI,G2sheets,LE,M,NY,P,W),Brazil, Amazonas,RioNegro,towardsconfluencewithRioSolimões,May1851. Guatteria ucayalianaDiels(1924b)138.—Guatteria dielsianaR.E.Fr.(1938) 719.—Type:Tessmann 3212(holoB;isoG,S),Peru,Loreto,Yarina Cocha,MiddleRíoUcayali,155m,24Sept.1923. Guatteria brevicuspisR.E.Fr.(1939)491,f.28e,f.—Type:Krukoff 5589 (holoS;isoBM,F,G,K,M,MO,NY2sheets,RB,S,U2sheets,US), Brazil,Acre,RioPurus,nearmouthofRioMacauã(‘Macauhan’),tributary ofRioIaco(‘Yacu’),21Aug.1933. Guatteria cylindrocarpa R.E.Fr.(1957b)601,t.2.—Type:Schultes & López 8949(holoUS),Brazil,Amazonas,RioNegro,Tapurucuara,11Sept.1947. Guatteria sp. 1Chatrouetal. (1997)109. Shrubortree,2–13(–25)mtall,toc.10(–30)cmdiam;young twigsdenselycoveredwithappressedhairs,soonglabrous. Leaves:petiole8–13mmlong,2–4mmdiam;laminanarrowly obovate-oblongtonarrowlyovate-oblong,rarelyelliptic,15–35 by3.5–8.5(–12)cm(leafindex3.5–5.5),chartaceous,densely to rather densely verruculose, dull greenish brown to brown above, pale brown to brown below, glabrous above, densely to sparselycoveredwithappressedhairstoc.1mmlongbelow, base long-attenuate, sometimes acute to obtuse, apex acuminate(acumen10–30mmlong),primaryveinimpressedto flat above, secondary veins distinct, 16–22 on either side of primary vein, flat or slightly impressed above, smallest distance betweenloopsandmargin2–3mm,tertiaryveinsslightlyraised above,reticulate.Flowers in 1–2-flowered inflorescences in axilsofleaves;pedicels4–10(–20)mmlong,1–3mmdiam, fruitingpedicelsrarelytoc.40mmlong,denselycoveredwith appressedhairs,articulatedat0.2–0.5fromthebase,bracts c.5,soonfalling;flowerbudsovoidtobroadlyovoid;sepals free or almost so, broadly ovate to broadly ovate-triangular, 4–9by3–7mm,appressed,outersidedenselycoveredwith appressedhairs;petalsgreenishyellow,yellow,yellow-orange orcreaminvivo,rhombic-ovatetoelliptic,8–20by5–10mm, outersidedenselycoveredwithappressed,silveryhairs;stamens1.5–2mmlong,connectiveshieldpapillatetoglabrous. Monocarps 10–25, green, maturing red to black in vivo, brown todarkbrowninsicco,narrowlyellipsoidtoellipsoid,17–25by 7–9mm,sparselycoveredwithappressedhairs,apexacute toapiculate(apiculumc.0.5mmlong),wall0.2–0.5mmthick, stipes5–13by1.5–2mm.Seed narrowlyellipsoid,18–22by 6–9 mm, brown, rugulose to rugose, longitudinally and transverselygrooved,raphenotdistinctfromrestofseed. Distribution—AmazonianVenezuela(Amazonas,Bolívar), Guyana,FrenchGuiana,AmazonianEcuador(Napo,Sucumbios),Peru(Huánuco,Loreto,MadredeDios,SanMartín, Ucayali),Brazil(Acre,Amapá,Amazonas,Maranhão,Mato Grosso,Pará,Rondônia,Roraima),Bolivia(Beni). Habitat&Ecology—Innon-inundatedorperiodicallyinundated(restinga,tahuampa)forest,onclayeyorrarelyonsandy soil.Atelevationsof0–800m.Flowering:throughouttheyear; fruiting:throughouttheyear. Vernacularnames—Brazil:Envira-preta(H.C. & R.P. Lima 2718),Envireira(Rodrigues 534),Pindaíba-preta(Marimon 51). Peru:Anonilla(Freitas 7, Ruiz & Melendez 1344),Carahuasca amarilla(Tello 2999),Hicoja(Schunke V. 7688),Hicojanegra (Schunke V. 4294, 8674),Huascaanonilla(Vásquez 12285), Huascahicoja(Schunke V. 7844),Icoja(Rimachi Y. 4175), Tortuga(Plowman 2537).Venezuela:Kunguate(Yekuana) (Aymard C.& Fernández 7324),Kunwatö(Yekuana)(Ang. Fernández 5266). Notes—Guatteria blepharophylla is recognizable by shortly pedicellateflowers(pedicelsgenerallyuptoc.10mmlong) and by petals which are densely covered with apressed, silvery hairs. Guatteria blepharophyllaisavariablespecies.Characteristically it has densely to rather densely verruculose leaves with a long-attenuate base, and it is found as such particularly in AmazonianBrazilandVenezuela.InEcuadorandPeruthe leaves tend to have a shortly attenuate, acute or even obtuse base.SpecimensfromJenaroHerrera,Loreto,Peruarevery variable in leaf shape, particularly the leaf base which shows thewholevariationrangebetweenlong-attenuateandobtuse. 25. Guatteria brevipetiolataMaas&Westra—Map6 Guatteria brevipetiolata Maas&WestrainErkensetal.(2008)478,f.4. —Type:T.D. Pennington et al. 14941(holoU2sheets;isoK2sheets), Ecuador,Esmeraldas,Anchayacu,EloyAlfaro,Mayronga,100–250m, 14Apr.1994. Guatteria sp. 5Chatrouetal. (1997)110. Treeorshrub3–20mtall,toc.30cmdiam,slashorangebrown;youngtwigsdenselycoveredwitherectandappressed, palebrown,somewhatcurlyhairs.Leaves: petiole 2–5 mm long,1–3mmdiam;laminanarrowlyoblong-elliptictonarrowly elliptic,16–28by7.5–10cm(leafindex2.6–3.2),chartaceous, not verruculose, shiny, green above, green below, sparsely covered with erect and appressed, pale brown, somewhat curly hairs above, rather densely covered with erect and appressed, pale brown, somewhat curly hairs below, base obtuse, sometimes oblique, apex acute to shortly and very gradually, bluntlyacuminate(acumen5–10mmlong),primaryveinimpressed above, secondary veins distinct, 10–15 on either side of primary vein, raised above, smallest distance between loops andmargin2–3mm,tertiaryveinsraisedabove,reticulate. Flowers 1–2-flowered inflorescences in axils of or on leafless branchlets;pedicels30–55mmlong,1–2mmdiam,densely to rather densely covered with erect and appressed, pale brown,somewhatcurlyhairs,articulatedat0.1–0.2fromthe base,bracts4–6,soonfalling;flowerbudsnotseen;sepals free,ovate,6–7by4–5mm,spreadingtoreflexed,outerside denselycoveredwithappressedhairs;petalspalegreen, greenish brown or greenish red in vivo, oblong-elliptic to narrowlyoblong-elliptic,13–20by5–11mm,outersidedensely coveredwithmostlyappressedhairs;stamens1.5–2mmlong, connectiveshieldpapillate,hairyorglabrous.Monocarps and seednotseen(butonlabelofAulestia & Gonti 1736:‘frutos jovenescremas’).
49 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria Distribution—Ecuador(Esmeraldas,Napo). Habitat&Ecology—Inprimaryforest,onred,clayeysoil.At elevationsof60–250m.Flowering:FebruarytoApril;fruiting: February(?). Vernacularname—Ecuador:Oitahumo(Aulestia & Gonti 1736). Note—Guatteria brevipetiolata can be recognized by a short petiole, an oblique and obtuse leaf base, a bluntly acuminate leafapexandlong-pedicellateflowers. 26. Guatteria campestris R.E.Fr.—Fig.21;Map6 Guatteria campestris R.E.Fr.(1939)402.—Type:Glaziou 14466 (holoB;iso BR,C,G,K,P),Brazil,MinasGerais,Araçuaí(‘Arassuahy’). Guatteria reticulata R.E.Fr.(1939)324,f.4b–d.—Type:Schwacke 9155 (holoB),Brazil,MinasGerais,OuroPreto,7Feb.1893. Guatteria xylopioidesR.E.Fr.(1939)459,f.21g.—Type:Glaziou 13401(holo B;isoK,P),Brazil,RiodeJaneiro,CaboFrio,Feb.1882. Tree3–18(–22)mtall,13–35cmdiam;youngtwigsdenselyto sparselycoveredwithappressedhairs,soonglabrous.Leaves: petiole5–10mmlong,1.5–2mmdiam;laminanarrowlyelliptic,sometimeselliptic,7–21by2–6(–9)cm(leafindex 1.8–6),chartaceous,notverruculose,palebrowntobrownon both sides, sparsely covered with appressed hairs to glabrous above, densely covered with appressed hairs to glabrous below, base obtuse, rarely acute, apex acute, primary vein impressed above, secondary veins distinct, 12–16 on either side of primary vein, raised above, angle of secondary vein with primary vein 40–55°,smallestdistancebetweenloopsandmargin1–2mm, tertiaryveinsraisedabove,reticulate.Flowers in 1–2-flowered inflorescencesinaxilsofleaves;pedicels5–25mmlong,1–2 mmdiam,fruitingpedicels2–3mmdiam,denselytosparsely coveredwithappressed,brownhairs,finallyglabrous,articulatedat0.2–0.3fromthebase,bracts2–4,soonfalling,2–4 mmlong;flowerbudsdepressedovoid;sepalsfree,broadly ovate-triangular,5–6by5–6mm,reflexed,outersideglabrous; petals cream in vivo, ovate, 10–25 by 6–10 mm, outer side denselycoveredwithappressed,brownhairs;stamensc.1mm long,connectiveshieldhairy.Monocarpsc.15,greenwhen younginvivo,blackinsicco,ellipsoid,12–14by8mm,glabrous,apexapiculate(apiculum<1mmlong),wallc.0.8mm thick,stipes2–5by1mm.Seedellipsoid,12–14by5mm, orange-brown,pittedtorugulose,rapheraised. Distribution—Brazil(Bahia,EspíritoSanto,MinasGerais, RiodeJaneiro). Habitat&Ecology—Innon-inundated,Atlanticrainforest orcerrado,onclayeysoil.Atelevationsof0–1050m.Flowering:throughouttheyear;fruiting:January,April,May,October, November. Vernacularnames—Brazil:Araticum(Ramalho & Rodrigues 1594),Conde(Piratininga 14),Imbuí(R. Guedes 2280, Piratininga 14). Note—Guatteria campestris is distinct by mostly narrowly elliptic leaves with an obtuse base, with secondary veins forminganangleof40–55°totheprimaryvein,whichisanarrow angleforthegenus.ItresemblesG. australis in the size and theshapeoftheleaves(narrowlyelliptictoelliptic).However, G. australis has secondary leaf veins forming an angle of 70–90°withtheprimaryvein. 27. Guatteria campinensis(Morawetz&Maas)Erkens&Maas —Map7 Guatteria campinensis (Morawetz&Maas)Erkens&Maas(2008)405. —Guatteriella campinensis Morawetz&Maas(1984)20,f.1,2.—Type: Morawetz & D. Coêlho 31-24883(holoINPA;isoINPA2sheets,U,WU), Brazil,Amazonas,km165ofnewroadfromManaustoItacoatiara,24 Aug.1983. Tree12–15mtall,toc.15cmdiam;youngtwigsdenselycovered with a velutinous indument of erect, brown, long-persistent Fig. 21 Guatteria campestrisR.E.Fr.a.Floweringbranch;b.leaf;c.flower; d.fruit(a:Kollmann 4276;b:Sucre 7775;c:Ramalho & Rodrigues 1594; d: Demuner 142;fromLobão(2009)f.4). c b d a 2 cm 1 cm 2 cm 1 cm
50 Blumea–Volume60/1–3,2015 Fig. 22 Guatteria candolleanaSchltdl.a.Floweringbranch;b:leafbase; c:flower;d:fruit(a,d:Mori 10901;b,c:Farias 263;fromLobão(2009)f.4). hairs.Leaves:petiole2–5mmlong,3–5mmdiam;lamina narrowlyelliptictonarrowlyobovate,15–32by7–12cm(leaf index2.2–2.7),chartaceous,notverruculose,shiny,dark brown above, brown below, sparsely covered with erect, brown hairs above, the primary vein densely so, rather densely covered with a velutinous indument of erect, brown hairs below, base obtuse to subcordate and almost stem-clasping, apex acutetoacuminate(acumentoc.10mmlong),primaryvein impressed above, secondary veins distinct, 20–26 on either side of primary vein, impressed above, more or less forming amarginalvein,atasmallestdistanceof2–3mmfromthe margin,tertiaryveinsimpressedabove,percurrent.Flowers in 1–2-flowered inflorescences in axils of leaves or on leaflessbranchlets;pedicels5–7mmlong,c.3mmdiam,fruiting pedicels5–11mmlong,4–5mmdiam,denselycoveredwith a velutinous indument of erect, brown hairs, articulated at c.0.5fromthebase,bractsnotseen,notcountable;flower budsnotseen;sepalsfree,broadlyovate-triangular,c.10by10 mm, appressed, outer side densely covered with a velutinous indumentoferect,brownhairs;petalsbrownorgoldenyellow invivo,ovate,c.20by13mm,outersidedenselycoveredwith avelutinousindumentoferect,brownhairs;stamens1–2mm long,connectiveshieldhairy.Monocarps 25–50, brown when ripeinvivo,browninsicco,ellipsoid,15–20by6–13mm, densely covered with a velutinous indument of erect, brown hairs,apexrounded,wall0.5–1{–2}mmthick,stipes2–5by 2–5mm.Seedellipsoid,14–17by4–7mm,palegreyishbown, rugulose and longitudinally and transversely grooved, raphe notdistinctfromrestofseed. Distribution—AmazonianBrazil(CentralAmazonas). Habitat&Ecology—Incampinaforest,onwhitesand.At elevationsbelow100m.Flowering:August;fruiting:August, December. Vernacularnames—Notrecorded. Note—Guatteria campinensis, formerly put in the genus Guatteriella, shares the velutinous indument of most parts and the leaf venation with G. trichocarpa.Itdiffersfromthatspecies, however,byitsmuchshorterpetioles(2–5vs5–10mm)and thelackofverruculosestructuresintheleaves. 28. Guatteria candolleana Schltdl.—Fig.22;Map7 Guatteria candolleana Schltdl.(1834)325;R.E.Fr.(1939)342.—Cananga candolleana (Schltdl.)Warm.(1873)145.—Type:Sellow 5442 p.p. (holo B;isoBR, G,K),Brazil,RiodeJaneiro,Sumidouro. Guatteria hilarianaSchltdl.var.pallescens R.E.Fr.(1900)15.—Type:Glaziou 3855(lectoB,selectedbyLobãoinMaasetal.2011;isolectoC,K, P),Brazil,RiodeJaneiro,RiodeJaneiro,Jacarepaguá,29Sept.1869. Treeletorshrub0.7–4mtall,5–17cmdiam;youngtwigs denselycoveredwitherect,brownhairstoc.2mmlong,soon glabrous.Leaves:petiole2–4mmlong,c.1mmdiam;lamina elliptic,5–15by2–4cm(leafindex2–4.3),chartaceous,sparsely verruculose, dark brown, brown below, glabrous above, denselycoveredwitherect,brownhairstoc.2mmlongto glabrous below, base cordate, sometimes obtuse, apex acute toshortlyacuminate(acumenc.5mmlong),primaryvein impressedabove,secondaryveinsdistinct,9–13oneither side of primary vein, raised above, smallest distance between loopsandmarginc.1mm,tertiaryveinsraisedabove,reticulate.Flowerssolitaryinaxilsofleaves;floweringandfruiting pedicels12–60mmlong,0.5–1mmdiam,denselytosparsely coveredwitherect,brownhairs,articulatedat0.2–0.3from thebase,bracts2–4,soonfalling,5–8mmlong;flowerbuds broadlyovoid;sepalsfree,broadlyovate-triangular,2–6by 2–5 mm, reflexed, outer side densely to sparsely covered witherect,brownhairs;petalscreamoryellowinvivo,elliptic, 10–30by8–12mm,outersidedenselycoveredwitherectto appressed,brownhairs;stamensc.2mmlong,connective c b d a 5 mm 2 cm 1 cm 5 mm
51 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria shieldpapillate.Monocarpsc.10,blackishpurpleinvivo,black insicco,ellipsoid,7–10by4–5mm,glabrous,apexapiculate (apiculumc.1mmlong),wall0.1–0.3mmthick,stipesc.5by 0.5–1mm.Seedellipsoid,7–10by4–5mm,orange-brown, pitted,rapheslightlyraised. Distribution—Brazil(Bahia,EspíritoSanto,MinasGerais, RiodeJaneiro). Habitat&Ecology—Innon-inundated,Atlantic,sometimes disturbedrainforestorcerrado,onclayeytosandysoil.Atelevationsof0–380m.Flowering:JanuarytoAugust;fruiting: August,October. Vernacularnames—Brazil:Pindaíba(Eupunino 100, Folli 698),Pindaíba-candoleana(L.A.M. Silva et al. 928),Pindaíbada/de-capoeira(L.A.M. Silva et al. 860, 928),Pindaíba-mole (L.A.M. Silva & Brito 860),Puruna-do-nativo(V.C. Souza 166). Notes—Guatteria candolleana is characterized by leaves with a cordate or sometimes obtuse base, densely covered with erecthairsonthelowerside.ItissimilartoG. australis but that specieshasleaveswithanacutetodecurrentbase. For differences with G. macropusseeunderthatspecies. 29. Guatteria capixabaeLobão&J.C.Lopes—Map9 Guatteria capixabaeLobão&J.C.Lopes(2014)205,f.1.—Type:Kollmann et al. 1948 (holoRB;isoMBML),Brazil,EspíritoSanto,SantaTeresa, ValsuganaVelha,EstaçãoBiológicadeSantaLúcia,500m,11Feb.1999. Treeorshrub2–6mtall,4–6cmdiam; youngtwigsglabrous. Leaves:petiole4–6mmlong,1–1.5mmdiam;laminanarrowly elliptic,rarelynarrowlyobovate,7–15by1.5–5cm(leafindex 2.3–5.3),chartaceous,notverruculosetodenselyverruculose, greyish above, dark grey below, glabrous above, glabrous or rarely sparsely covered with appressed hairs below, base attenuate,apexacuminate(1–10mmlong),primaryveinflatabove, secondaryveinsdistinct,9–18oneithersideofprimaryvein, smallestdistancebetweenloopsandmargin2–3mm,tertiary veinsreticulate,raisedabove.Flowerssolitaryinaxilsofleaves; bracts3–4,soonfalling,notseen;pedicels8–10(–30)mm long,c.1mmdiam,fruitingpedicelstoc.25mmlong,1–1.5mm diam, sparsely covered with appressed hairs to glabrous, articulatedat0.2–0.4fromthebase;flowerbudsnotseen;sepals free,deltate,c.3by3–4mm,appressed,outersideglabrous orrarelysparselycoveredwithappressedhairs;petalsyellow invivo,elliptictorhombic-ovate,10–15by4–8mm,outer sidedenselycoveredwitherect,greyhairs;stamensc.2mm long,connectiveshieldpapillate.Monocarps 5–7,reddishin vivo,blackishinsicco,narrowlyellipsoid,13–17by4–7mm, glabrous or rarely sparsely covered with appressed hairs, apex apiculate(apiculumc.0.5mmlong),wallc.0.5mmthick;stipes 2–4by1–2mm.Seednarrowlyellipsoid,12–15by4–5mm brownreddish,rugose. Distribution—Brazil(EspíritoSanto). Habitat&Ecology—Innon-inundated,Atlanticrainforest.At elevationsof500–850m.Flowering:August,February;fruiting: November,March. Vernacularnames—Notrecorded. Note—Guatteria capixabae, endemic to the Brazilian state ofEspíritoSanto,isrecognizablebyveryshortlystipitatemonocarpsandanattenuateleafbase.Intheoriginaldescriptionitis mentioned that the leaves are verruculose and that this species shouldbecomparedwithspeciesofsect.Mecocarpus(like G. duodecima),butthespecimenwehadathandinLeidenhad only very vague verruci and is in that respect completely differentfromthestructuressotypicalforsect.Mecocarpus.Inour opinion G. capixabae is very close to G. stenocarpa, endemic totheBrazilianstateofBahia,differinginshorterstipes(2–4 vs5–10mm)andsmallermonocarps(13–17vs18–25mm). 30. Guatteria carchianaMaas&Westra,sp. nov.—Fig.23, 24a;Map9 Foliis coriaceis, pilis erectis subtus obtectis, apice acutis, summo obtuso, monocarpiisseminibusqueapicepunctatisbenedistincta.—Typus:Vargas L. et al. 4242(holoMO;isoQCNE,U),Ecuador,Carchi,Espejo,ReservaGolondrinas,pathbetweenEstaciónSantaRosaandElCorazón,1700–2050m, 21Jan.2004. Tree10–30mtall,diamnotrecorded;youngtwigsdenselycoveredwitherect,brownhairstoc.1mmlong,soonglabrous. Leaves:petiole3–5mmlong,c.2mmdiam;laminaellipticto obovateornarrowlyso,15–18by6–8cm(leafindex2.1–2.8), coriaceous, not verruculose, dull, grey above, brown below, glabrous above, densely covered with erect, brown hairs to c.1mmlongbelow,baseacutetoobtuse,marginsslightly revolute, apex bluntly acute, primary vein impressed above, keeled below, secondary veins distinct, 12–15 on either side of primary vein, raised above, smallest distance between loops andmargin2–4mm,tertiaryveinsraisedabove,reticulate. Flowersin1–2-floweredinflorescencesinaxilsofleaves; pedicels15–20mmlong,1–1.5mmdiam,fruitingpedicels toc.25mmlong,2–4mmdiam,denselytosparselycovered witherect,brownhairs,finallyglabrous,articulatedat0.2–0.3 fromthebase,bracts5–6,soonfalling,notseen;flowerbuds depressedovoid;sepalsfree,broadlyovate-triangular,5–6by 5–6 mm, reflexed, outer side densely covered with erect, brown hairs;petalsgreenishyellowtodullorangeinvivo,ovatetoelliptic,c.15by8–10mm,outersidedenselycoveredwitherect, brownhairs;stamensc.2mmlong,connectiveshieldpapillate. Monocarpsc.75,darkred-purpletoblackinvivo,blackinsicco, narrowlyellipsoid,14–16by5–6mm,sparselycoveredwith appressedhairs,soonglabrous,apexhard-pointed(apiculum c.1mmlong),wall0.2–0.3mmthick,stipes15–25by1mm. Seednarrowlyellipsoid,c.15by6mm,brown,apexpointed, rugulose. Distribution—Ecuador(Carchi). Habitat&Ecology—Inwet,montaneforest.Atelevationsof 1450–2050m.Flowering:January,May;fruiting:January,May. Vernacularnames—Ecuador:Cafesillo(Vargas L. et al. 4242). Fieldobservations—Flowerswithcherimoya-likescent (Madison et al. 4442). Uses—Ecuador:Woodusedforformwork(‘Maderapara encofrados’). Other specimens examined. eCuador, Carchi,nearMaldonado,1450– 1650m,24May1978,Madison et al. 4442(AAU,F,QCA,SEL,U). Brazil Colombia Peru Venezuela Panama a Rica a Nicaragua Bolivia Guyana Suriname French Guiana L e s s e r A n t i l l e s Ecuador Map 9 Distribution of Guatteria capixabae (●),G. carchiana (£ ),G. castilloi (■),G. chasmantha (p)andG. chrysophylla (u).
52 Blumea–Volume60/1–3,2015 Fig. 23 Guatteria carchianaMaas&Westra.Fruitingbranch(Vargas L. et al. 4242,holotypeMO).
53 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria Fig. 24 a.Guatteria carchianaMaas&Westra.Flowers.–b.Guatteria chiriquiensisR.E.Fr.Fruit.–c.Guatteria citriodoraDucke.Fruit.–d,e.Guatteria conspicuaR.E.Fr.d.Flowers;e.fruit.–f.Guatteria crassiveniaN.Zamora&Maas.Detailoflowerleafsurface.–g.Guatteria delicatulaMaas&Westra. Flowerpedicelwithperianthandstamensgone.–h.Guatteria dotanaN.Zamora&Erkens.Fruit(a:Vargas L. et al. 4242,isotypeU;b: Zamora et al. 1461,U; c: Mori et al. 23673,U;d,e:Jonker-Verhoef & Jonker 494, U;f:Herrera C. & Chacón 2746,U;g:Solomon 9282, holotypeU;h:Estrada & Solano 1817,MO). c b d a gh e f
54 Blumea–Volume60/1–3,2015 Note—Guatteria carchiana is well recognizable by coriaceous leaves with a dense indument of erect, brown hairs on the lower side, an acute leaf apex, hard-pointed monocarps, andpointedseeds.Itisprobablyclosesttothepoorlyknown Colombian G. rufotomentosa, sharing most of its indument and leaf features, but from which it differs, however, by a shorter petiole(3–5vs5–15mmlong),anobtusetoacuteinstead ofattenuateleafbase,andlesssecondaryveins(12–15vs 15–25). 31. Guatteria caribaeaUrb.—Plate3a,b;Map8 Guatteria caribaea Urb.(1905)240;R.E.Fr.(1939)480,f.25b,c.—Cananga caribaea(Urb.)BrittoninBritton&Wilson(1924)311. —Type:Sintenis 1535 (lectoB,selectedbyFries1939;isolectoBM,BP,G2sheets,GH, JE,K,L,M,NY,P,PR,S,US,WU),PuertoRico,SierradeLuquillo,‘in silvismontisJiménes’,June1885. Tree4–40mtall,5–60cmdiam;youngtwigsratherdensely coveredwithappressedhairs,soonglabrous.Leaves: petiole 4–5mmlong,1–3mmdiam;laminanarrowlyelliptic,8–21by 3–8cm(leafindex2.2–4),chartaceous,notverruculose,dull, green to brown above and below, glabrous above, sparsely to rather densely covered with appressed hairs below but primary vein densely covered with appressed hairs, base acute, apex acuminate(acumen10–20mmlong),primaryveinimpressed to flat above, secondary veins distinct, 6–12 on either side of primary vein, raised above, smallest distance between loops andmargin5–7mm,tertiaryveinsraisedabove,reticulate. Flowers in 1–2-flowered inflorescences in axils of leaves or on leaflessbranchlets;floweringandfruitingpedicels10–25mm long,c.1mmdiam,sparselycoveredwithappressedhairsto glabrous,articulatedatc.0.2fromthebase,bracts4–5,soon falling,lowerbractsverybroadlyovate,c.1mmlong,uppermostbractsoneseen,broadlyovate,c.4mmlong;flowerbuds depressedovoid;sepalsfree,broadlytoshallowlygrooved, ovate-triangular,2–4by3–4mm,appressedtospreading, sparselycoveredwithappressedhairs;petalswhiteorcreamy whiteinvivo,narrowlyelliptic,10–15by3–5mm,outerside denselycoveredwithappressedhairs;stamensc.1mmlong, connectiveshieldpapillate.Monocarps 5–25, green, maturing black to purple-black in vivo, black in sicco, narrowly ellipsoid, 11–20by5–7mm,sparselycoveredwithappressedhairs, soonglabrous,apexobtuselyapiculate(apiculum<1mmlong), wall0.3–0.5mmthick,stipes2–5by1mm.Seed narrowly ellipsoid,11–18by4–6mm,darkbrown,longitudinallyand transversely grooved, sometimes also pitted, raphe not distinct fromrestofseed. Distribution—LesserandGreaterAntilles:Dominica,Grenada,Guadeloupe,Martinique,Montserrat,PuertoRico,Saint KittsandNevis,St.Lucia. Habitat&Ecology—Indenserainforest,onclayeyloam soil.Atelevationsof200–700m.Flowering:January,March toAugust,November,December;fruiting:January,Marchto May,September,November. Vernacularnames—Boisnouè(Jérémie 1115),Boisviolin (Hill 25840, Jérémie 1115, Shillingford 450),Corchoblanco (Little Jr. 13764),Corossolmontagne(Jérémie 1115),Mahaut noir(Hill 25840),Ti-cachiman-bois(Slane 222),Wildsoursop (Slane 222). Fieldobservations—AccordingtoStern & Washaussen 2595(US)fromDominicatheflowersare‘fragrant,highlyaromatic’. Notes—Guatteria caribaea, the only species of the genus occurringintheLesserAntilles,isrecognizablebythesecondary veins which are raised on the upper side of the leaves, slender pedicels, narrowly ellipsoid monocarps on short stipes, and seeds which are mostly longitudinally and transversely grooved. InStern & Wasshausen 2595(US)thethirdbractfromthetop is more or less persisting, foliaceous, narrowly elliptic and 15– 25mmlong. Sterilematerialcanhaveleavesofupto28by8cm. 32. Guatteria castilloiMaas&Westra—Map9 Guatteria castilloiMaas&Westra(2010)264,f.5.—Type:Castillo S. 5444 (holoVEN;isoMO),Venezuela,Amazonas,Mun.Autana,RíoSipapo, betweencommunityMonteNegroandCaldero,250m,18Aug.1997. Tree8–20mtall,diamnotrecorded;youngtwigsratherdensely coveredwithappressed,whitehairs,soonglabrous.Leaves: petiole20–35mmlong,1–2mmdiam;laminanarrowlyoblongelliptictonarrowlyovate,12–19by4–8cm(leafindex2.4–3.7), coriaceous, not verruculose, shiny, greyish green to brown above, brown below, glabrous above, sparsely covered with appressed, white hairs below, base acute to obtuse, extreme base slightly attenuate, apex obtuse, sometimes shortly acuminate(acumentoc.5mmlong),basalmarginssomewhatrevolute, primary vein impressed above, secondary veins distinct, 13–15oneithersideofprimaryvein,stronglyraisedabove, smallest distance between loops and margin 1–2 mm, tertiary veinsraisedabove,stronglyreticulate.Flowers solitary in axils ofleaves;pedicels15–20mmlong,c.1mmdiam,fruiting pedicelsc.22mmlong,1–2mmdiam,sparselycoveredwith appressed,whitehairs,articulatedat0.2–0.3fromthebase, bracts3–5,soonfalling,upperbractc.5mmlong;flowerbuds notseen;sepalsfree,broadlyovate-triangular,5–6by5–6 mm, appressed, rigid, margins somewhat revolute, outer side sparselycoveredwithappressedhairs;petalsgreeninvivo, narrowlyoblong-elliptic,13–20by5–8mm,outersiderather denselytosparselycoveredwithappressedhairs;stamens c.2mmlong,connectiveshieldpapillate.Monocarpsc.10, greeninvivo,browninsicco,ellipsoid,13–15by7–10mm, sparselycoveredwithappressedhairs,apexapiculate(apiculum <0.2mmlong),wallc.1.5mmthick,stipes10–12byc.1mm. Seednotstudied(abortive?). Distribution—AmazonianVenezuela(Amazonas). Habitat&Ecology—Inblackwater-floodedforest(‘bosque ribereño,influenciadoporaguasnegras’).Atelevationsof c.250m.Flowering:AugusttoOctober;fruiting:October. Vernacularnames—Notrecorded. Note—Guatteria castilloi is unique by the combination of longpetiolesforthegenus(20–35mm),araised,reticulate leafvenation,andamostlyobtuseleafapex. 33. Guatteria chasmanthaR.E.Fr.—Map9 Guatteria chasmanthaR.E.Fr.(1939)513,f.34a.—Type:Lawrance 422 (holoS;isoA,F,G,MO,US),Colombia,Boyacá,AltoChapón,100km NWofBogotá,highforestfront,6000ft,9Aug.1932. Tree9–25mtall,toc.1.2mdiam;youngtwigsdenselyor rather densely covered with small, appressed, more or less curlyhairs,soonglabrous.Leaves:petiole8–15mmlong, 1.5–2mmdiam;laminaelliptictomostlynarrowlyellipticor narrowlyobovate-elliptic,12–20by4.5–7.5cm(leafindex 2.4–2.9),chartaceous,sparselyornotverruculose,greyish black and more or less shiny above, brownish black to dark brown below, glabrous above, sparsely covered with small, appressed hairs to glabrous below, base acute to rounded, extremebaseattenuate,apexacutetoacuminate(acumento c.5mmlong),primaryveinflattoslightlyraisedabove,secondary veins distinct, 10–15 on either side of the primary vein, shortestdistancebetweenloopsandmargin2–3mm,tertiary
55 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria Plate 3a,b.Guatteria caribaeaUrb.a.Floweringbranch;b.matureflower.–c.Guatteria costaricensisR.E.Fr.Leavesandfruit.–d.Guatteria cuatrecasasii SánchezS.Floweringbranch.–e.Guatteria dolichopodaDonn.Sm.Floweringbranchandleaves.–f.Guatteria foliosaBenth.Floweringbranch.–g,h.Guatteria talamancanaN.Zamora&Maas.g.Floweringbranch;h.flower(a,b:Graveson et al. 3093;c:Maas et al. 9392;d:R. Cámara Leret & Copete Maturana 1743; e: Maas et al. 9462;f: Mori & Gracie 23929;g,h:Monro 6108).—Photos:a,b:R.H.J.Erkens;c,e:P.J.M.Maas;d:R.CámaraLeret;f:C.Gracie;g,h:A.K.Monro. c b d a g h e f
56 Blumea–Volume60/1–3,2015 Fig. 25 Guatteria chiriquiensisR.E.Fr.Floweringbranch(Biesmeijer et al. 321,U).
57 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria veinsraisedabove,reticulate.Flowers in 1–2-flowered inflorescences in axils of leaves or on leafless branchlets, pedicels 12–20by1–1.5mm,fruitingpedicelstoc.2mmdiam,rather densely to sparsely covered with small, appressed hairs, becomingglabrous,articulatedat0.2–0.4frombase,bracts5–6, soonfalling,notseen;flowerbudsnotseen;sepalsfree,broadly ovate,3–4by4–5mm,appressed,ratherdenselycovered with small, appressed hairs, petals yellowish green or peachcolouredinvivo,elliptic-ovate,7–13by7–10mm,innerpetals distinctlysmallerthanouterpetals,densely(base)torather denselycoveredwithwhitishhairs,stamensc.1.5mmlong, connectiveshieldpapillate.Monocarps 10–15, green, maturing blackinvivo,blackinsicco,glabrous,ellipsoid,12–15by7–10 mm,apexacutetoobtuse,wall0.5–0.7mmthick,stipes1–10 by2mm.Seed:nogoodseedsseeninsidemonocarps(eaten byinsectsorattackedbyfungus?). Distribution—Colombia(Antioquia,Boyacá). Habitat&Ecology—Inforestandmontaneforest.Atelevationsof1500–1900m.Flowering:MayandAugust;fruiting: August. Vernacularnames—Notrecorded. Note—Guatteria chasmantha is noteworthy by shiny almost blackishleavesincombinationwithshortpedicels(12–20mm long)andsmallsepals(3–4mmlong). 34. Guatteria chiriquiensisR.E.Fr.—Fig.24b,25;Map8 Guatteria chiriquiensisR.E.Fr.(1948b)11.—Type:Pittier 5748 (holoUS 4sheets),Panama,Chiriquí,vicinityofSanFelix,easternChiriquí,0–120m, Sept.1912. Tree5–25mtall,5–25cmdiam;youngtwigsslightlywinged, densely covered with a velutinous indument of long-persisting, erect,brownhairs.Leaves:petiole1–5mmlong,1–2mmdiam; laminanarrowlyoblong-elliptic,13–32by3–7cm(leafindex 3.5–4.2),chartaceous,notverruculose,dull,greyishgreen above, brown to greenish brown below, rather densely covered with erect, brown hairs above, densely covered with a velutinous indument of erect, brown hairs below, base acute to obtuse, sometimesslightlyoblique,apexacuminate(acumen5–25 mmlong),primaryveinflatapically,becomingdistinctlyraised basallyabove,secondaryveinsdistinct,13–15oneitherside of primary vein, raised above, smallest distance between loops andmargin2–3mm,tertiaryveinsraisedabove,reticulate. Flowersin1–2(–3)-floweredinflorescencesinaxilsofleavesor onleaflessbranchlets;pedicels15–20mmlong,1–2mmdiam, fruitingpedicelstoc.35mmlong,toc.2mmdiam,densely covered with appressed and erect, brown hairs, articulated at 0.3–0.4fromthebase,bracts5–7,soonfalling;flowerbuds depressedovoid;sepalsfree,broadlyovate-triangular,5–7 by5–7{–9}mm,appressed,outersidedenselycoveredwith appressed,brownhairs;petalsyelloworcreaminvivo,broadly oblong-ovate to oblong-ovate, 10–15{–25} by 10–12{–15} mm, outersidedenselycoveredwithappressedhairs;stamens1–2 mmlong,connectiveshieldglabrous.Monocarps15–40,dark purpletoblackinvivo,blackinsicco,ellipsoid,7–10by3–5 mm, sparsely covered with appressed hairs, particularly near theapicalpart,apexapiculate(apiculum<0.5mmlong),wall 0.1–0.2mmthick,stipes5–8byc.1mm.Seedellipsoid,6–8by 3–4mm,dark,shinybrown,smoothtopitted,rapheimpressed. Distribution—CostaRica,Panama. Habitat&Ecology—Inforest.Atelevationsof0–500m. Flowering:January,March,May,June,December;fruiting: AugusttoOctober. Vernacularname—Panama:Burillo(P.H. Allen 5217). Note—Guatteria chiriquiensis is easily recognizable by its almost oblong leaves, its short petioles, and particularly by its primary vein which is raised on the upper side, the last feature veryrarelyseeninthegenus.Thebasalleafmarginsaremostly revolute. 35. Guatteria chrysophyllaMaas&Setten—Map9 Guatteria chrysophylla Maas&Setten(1988)252,f.10.—Type:Davis & Yost 1011(holoNY;isoF,U2sheets),Ecuador,Napo,confluenceofQuiwado andTiwaenoRivers,22Apr.1981. Tree20–30mtall,>25cmdiam;youngtwigsdenselycovered witherect,goldenbrown,curlyhairs.Leaves:petiole15–30mm long,4–6mmdiam;laminaelliptictonarrowlyelliptic,20–30 by8–15cm(leafindex2–2.6),coriaceous,notverruculose, dull, greyish green above, golden brown below, densely to rather densely covered with hairs on primary vein, otherwise glabrous above, densely covered with erect, golden brown, curly hairs below, base acute to attenuate, extreme base decurrentalongpetiole,apexacuminate(acumenc.15mmlong), primary vein impressed to flat above, secondary veins distinct, 15–18oneithersideofprimaryvein,raisedabove,smallest distancebetweenloopsandmargin0.5–1.5mm,tertiaryveins raisedabove,mostlypercurrent.Flowersin1(–2)-flowered inflorescencesinaxilsofleavesorleaflessbranchlets;pedicels 15–20mmlong,c.2mmdiam,fruitingpedicelstoc.30mm long,3–6mmdiam,denselycoveredwitherect,goldenbrown, curlyhairs,becomingglabrous,articulatedat0.2–0.3fromthe base,bractsc.6,soonfalling,notseen;flowerbudsovoid, pointed;sepalsfree,broadlyovate-triangular,10–13by7–9 mm, appressed, outer side densely covered with erect, golden brown,curly,hairs;petalscolournotrecorded,onlyseeninbud stage, outer side densely covered with erect, golden brown, curly,hairs;stamensnotseen.Monocarps3–20,greenin vivo,blackinsicco,ellipsoidtoobovoid,17–25by12–18mm, surface smooth to irregularly wrinkled in sicco, subglabrous, apexroundedtoacute,wall1.5–2mmthick,stipes8–18by 1.5–3mm.Seedellipsoid,13–21by8–12mm,reddishbrown, rugose,raphenotdistinctfromrestofseed. Distribution—AmazonianEcuador(Napo,Pastaza),Peru (Loreto). Habitat&Ecology—Innon-inundatedforest,onredsoil.At elevationsof0–400m.Flowering:May;fruiting:April,October. Vernacularname—Ecuador:Menedowe(TomoUpriver dialect),whichmeansjaguar’stree(Davis & Yost 1011). Uses—Ecuador:‘Barkcrushedandmixedwithwater,rubbedoverheadandshouldersandfeverwillflee’(Davis & Yost 1011). Note—Guatteria chrysophylla shares the indument of curly hairs, the pointed flower buds, long sepals and the large, wrinkled monocarps with G. discolor, a species occurring in AmazonianBolivia,Brazil,PeruandFrenchGuiana.Itdiffers, however, by rugose instead of smooth seeds, the longer petioles(15–30vs5–10(–15)mm),amostlypercurrenttertiary venation,andthelongerstipes(8–18vs2–10mm). 36. Guatteria citriodoraDucke—Fig.24c,26;Map10 Guatteria citriodora Ducke(1930)104,t.2,f.3a–c;R.E.Fr.(1939)467, f.23a,b;Maasetal.(2007)638.—Type:Ducke RB 19609(holoRB;iso B,F,G,K,NY,P,S,U,US),Brazil,Amazonas,Maués,nearRioCurucá, 4Juneand17Dec.1927. Guatteria paraensisR.E.Fr.(1939)464,f.22c–e,syn.nov.—Type:Ducke RB 17866(holoS),Brazil,Pará,Belém(‘BelémdoPará’),7Nov.1922. Guatteria duckeanaR.E.Fr.var.subcordataR.E.Fr.(1948b)10,pl.4a.— Type:Ll. Williams 15777(holoUS3sheets;isoG,NY),Venezuela, Amazonas,Capibara(‘Capihuara’),AltoCasiquiare,120m,3June1942. Tree2–33mtall,5–50cmdiam;youngtwigsdenselycovered witherect,brownhairs,finallyglabrous.Leaves:petiole2–7 mmlong,1.5–2mmdiam;laminaelliptictoovate,mostlynar-
64 Blumea–Volume60/1–3,2015 Vernacularnames—Colombia:Cargamarillo(Mondragón 87, Restrepo & Mondragón 261).Ecuador:Cargaderanegra (Tipaz et al. 1725). Notes—Guatteria crassipes is well recognizable by a combination of coriaceous narrowly obovate to obovate leaves with along-attenuatebase. SomematerialcollectedintheEcuadorianstateofCarchi (Freire-Fierro 2614, Tipaz et al. 1725, Rubio et al. 2199) is aberrant in having verruculae all over the lower side of the lamina, thickerpetioles(uptoc.10mmdiam)andfruitingpedicels(up toc.8mmdiam). 42. Guatteria crassiveniaN.Zamora&Maas,sp. nov.—Fig. 24f,30;Map8 Foliis nitidis, verruculosis, venulis omnino distincte prominentibus bene distincta. — Typus:Herrera C. & Chacón 2746(holoINB;isoU),CostaRica, Limón,CantóndeLimón,ElProgreso,áreadesuelosinundadosentre 1500–1700m,FilaMatama,1600m,24Abril1989. Treec.6mtall,diamnotrecorded;youngtwigsdenselycovered withappressed,brownhairs,soonglabrous.Leaves: petiole 0–3mmlong,1–2mmdiam;laminaelliptic,6–12by3.5–5 cm(leafindex1.5–2.5),chartaceous,denselyverruculose along primary vein to sparsely so elsewhere, shiny, dark brown Fig. 29 Guatteria crassipesR.E.Fr.a.Branchwithflowersandfruit;b.flower; c.fruit;d.seed(all:D.A. Sánchez S. et al. 727,MEDEL;fromSánchezS. (1986)f.18. 1 cm c b d a 0.5 cm 1 cm 1 cm
65 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria Fig. 30 Guatteria crassiveniaN.Zamora&Maas.Floweringbranch(Herrera C. & Chacón 2746,isotypeU).
66 Blumea–Volume60/1–3,2015 above, brown below, densely to rather densely covered with erect, brown hairs on primary vein above, glabrous elsewhere, sparsely covered with appressed to more or less erect hairs below, base acute to slightly attenuate, apex acute to acuminate(acumen5–15mmlong),primaryveinraisedabove, secondary veins distinct, 12–22 on either side of primary vein, strongly raised above, smallest distance between loops and margin1–2mm,tertiaryveinsstronglyraisedabove,reticulate. Flowerssolitary,inaxilsofleaves;pedicels10–25mmlong, c.1mmdiam,denselytoratherdenselycoveredwithappressed hairs,articulatedat0.3–0.5fromthebase,bractsc.5,soon falling,notseen;flowerbudsdepressedovoid;sepalsfree, broadlyovate-triangular,3–6by4–5mm,appressed,finally reflexed, outer side sparsely covered with appressed to more orlesserecthairs;petalsyellowinvivo,ovate,10–15by6–11 mm,outersidedenselycoveredwithappressedhairs;stamens c.1.5mmlong,connectiveshieldpapillate.Monocarps and seednotseen. Distribution—CostaRica(Limón). Habitat&Ecology—Inwetforest.Atanelevationofc.1600 m.Flowering:April;fruiting:unknown. Vernacularnames—Notrecorded. Note—Guatteria crassivenia,endemictoCostaRicaand only once collected, is very easily recognized by its shiny leaves which are densely verruculose, particularly near the primary vein, by their primary vein which is raised on the upper side, andbytheveryprominentvenationonbothsides. 43. Guatteria cryandraErkens&Maas—Map11 Guatteria cryandra Erkens&Maas(2008)404;Maas&Westra(2011)118, f.3b.—Guatteriopsis kuhlmannii R.E.Fr.(1937)275,t.8,notGuatteria kuhlmanniiR.E.Fr.(1939).— Type:J.G. Kuhlmann 2015 = RB 24361 (holoS;isoRB4sheets),Brazil,Pará,RioTucuruí,affluentofRioXingu, Vitória,17Apr.1924. Tree3–10mtall,c.10cmdiam;youngtwigsdenselycovered withappressedhairs,soonglabrous.Leaves: petiole 2–5 mm long,1–2mmdiam;laminanarrowlyelliptictonarrowlyoblongelliptic,13–25by4–8cm(leafindex2.6–4.1),chartaceous, rather densely verruculose, pale greenish brown above, pale brown below, glabrous above, sparsely covered with appressed hairs below, the primary vein rather densely so, base obtuse to rounded,apexacuminate(acumen15–25mmlong),primary vein flat to slightly raised above, secondary veins indistinct, 17–25oneithersideofprimaryvein,slightlyraisedabove,not or indistinctly loop-forming, smallest distance between loops andmargin1–4mm,tertiaryveinsslightlyraisedabove,reticulate.Flowers in 1–2-flowered inflorescences in axils of leaves oronleaflessbranchlets;pedicels1–5mmlong,1–2mmdiam, fruitingpedicelstoc.5mmlong,c.3mmdiam,denselycovered withappressed,brownhairs,articulatedatc.0.2fromthebase, bracts not countable with certainty, one bract seen: broadly ovate-triangular;flowerbudsbroadlyovoid;sepalsbasally connate, broadly ovate-triangular, 5–6 by 5–6 mm, appressed, outersidedenselycoveredwithappressedhairs;petalsyellow orgoldenyellowinvivo,broadlyovatetoovate,10–15by8–10 mm,outersidedenselycoveredwithappressed,brownhairs; stamensc.2mmlong,connectiveshieldpapillate.Monocarps 10–30,redtored-orangewhenripeinvivo,blackishinsicco, ellipsoid,9–12by4–6mm,sparselycoveredwithappressed hairs,apexapiculate(apiculum0.5–1mmlong),wallc.0.2mm thick,stipes1–3byc.1mm.Seedellipsoid,9–10by4mm, pitted,raphenotdistinctfromrestofseed. Distribution—AmazonianBrazil(Amapá,Pará). Habitat&Ecology—Innon-inundatedforest,onespecimen growingalongriver,oftenonclayeysoil.Atelevationsofupto 250m.Flowering:March,June,November;fruiting:August, September. Vernacularnames—Notrecorded. Note—Guatteria cryandra is characterized by shortly petiolate leaves with a rounded to obtuse base, in combination with the stiffly appressed hairs on the leaves and young branchlets, aswellaswiththeshortlystipitatemonocarpswithstipes1–3 mm long, and very short pedicels making the flowers appear almostsessile. 44. Guatteria cuatrecasasiiD.Sánchez—Plate3d;Map12 Guatteria cuatrecasasii D.SánchezinMaasetal.(1988)253,f.12.—Type: D.A. Sánchez S. et al.907(holoMEDEL;isoCOL,CUVC,HUA,U),Colombia,Chocó,Quibdó,Mun.Lloró,roadfromYutotoLloró,2.8kmtothe ferry,65m,15Sept.1988. Treeorshrub3–35mtall,5–85cmdiam,oftenwithbuttresses orstiltroots;youngtwigswithnarrowwingsdecurrentfrompetioles, densely covered with erect to appressed, brown to greyish hairs2–3mmlong.Leaves:petiole1–5mmlong,2–4mm diam;laminanarrowlyoblong-elliptictonarrowlyovate,10–35 by2–7cm(leafindex3.5–5(–8)),subcoriaceous,denselyor rarely sparsely verruculose, blackish brown to greyish above, brown below, glabrous or rarely sparsely covered with erect to appressed, brown hairs above, primary vein permanently densely to sparsely covered with erect to appressed, brown hairs above, densely covered with erect to appressed, brown hairs2–3mmlongbelow,baseacutetoobtuse,oftenslightly oblique,apexacuminate(acumen5–30mmlong),rarelyacute, primary vein impressed to flat above, secondary veins distinct, 15–30oneithersideofprimaryvein,flatandofteninconspicuous above, more or less forming a marginal vein, at a smallest distance of 1–2 mm from the margin, tertiary veins often inconspicuous,flatabove,reticulate.Flowersin1(–2)-flowered inflorescencesinaxilsofleaves;pedicels20–35mmlong,2–4 mm diam, fruiting pedicels 25–50 mm long, 2–5 mm diam, densely to rather densely covered with erect to appressed, brownhairs,articulatedat0.3–0.5fromthebase,bracts2–4, soonfalling,theupperoneconcave,15–25mmlong;flower budsbroadlyovoid;sepalsfree,ovate-triangulartobroadly ovate-triangular,4–15(–22)by5–15(–22)mm,appressed, denselytosparselycoveredwithappressed,brownhairs;petals green, maturing yellow or purple in vivo, broadly ovate to obovate, 10–20 by 10–15 mm, outer side densely to sparsely coveredwithappressed,brownhairs0.5–1.5mmlong;stamens 1–1.5mmlong,connectiveshieldpapillate.Monocarps 25–65, green, maturing violet to black in vivo, black in sicco, ellipsoid, 8–12by4–6mm,glabrous,apexapiculate(apiculum0.3–0.5 mmlong),wallc.0.5mmthick;stipes3–10by1–1.5mm. Brazil Colombia Peru Venezuela Panama Costa Rica M e x i c o Guatemala Nicaragua Honduras Bolivia Guyana Suriname French Guiana P ue r t o R i co Haiti C u b a Lesser Antilles Ecuador Map 12 Distribution of Guatteria cuatrecasasii (●),G. denudata (£),G. discolor (■)andG. dolichopoda (p).
67 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria Seedellipsoid,7–10by3–4mm,darkbrowntoblack,rugose, raphenotdistinctfromrestofseed. Distribution—Colombia(Antioquia,Chocó,Riseralda),Ecuador(Esmeraldas). Habitat&Ecology—Inwet,pluvial,orpremontane,often secondaryforest.Atelevationsof80–1500m.Flowering:April toMay;fruiting:AugusttoNovember. Vernacularnames—Colombia:Cargaderoblanco,Cargaderomachetico(D.A. Sánchez S. et al. 885), Cargadero negro (Forero et al. 9499),Cargaderopuntadelanza(D.A. Sánchez S. & Velasquez 878, 894, Hoyos M. & D.A. Sánchez S. 1175), Cedrillo,Guasco(Cogollo et al. 2844),Nuanamú(D.A. Sánchez S. et al. 307).Ecuador:Chalviande(Rubio & Quelal 1323). Uses—Colombia:Woodusedasfirewoodandforconstructionofhouses(‘Lamaderaseusacomoleñayparaconstruccióndeviviendas’)(D.A. Sánchez S. & Velasquez 894, D.A. Sánchez S. et al. 907);thebarkisusedforbagsofguasca (fibers)(‘lacortezaseutilizaparasacarguasca(fibra)’);thebark isusedtotieupdifferenttools(‘paraamarraroatardiferentes implementos’)(D.A. Sánchez S. et al. 907).Ecuador:Usedfor construction(‘Usadoenconstrucción’)(Rubio & Quelal 1323). Note—Guatteria cuatrecasasii may be confused with G. elegantissima. For the differences between the species see under the latter. 45. Guatteria cuscoensis Maas&Westra, sp. nov.—Fig.31; Map11 Abomnibusspeciebusperuvianisandicolispedicellislongioribusdiffert.— Typus:Garcia C. et al. 984(holoWAG;isoCUZ,USM),Peru,Cusco,Paucartambo,Kosñipata,SanPedro,1650m,6Sept.2006. Tree4–20mtall,10–15cmdiam;youngtwigsratherdensely coveredwithappressedhairs,soonglabrous.Leaves: petiole 4–7mmlong,1–2mmdiam;laminanarrowlyelliptic,9–15by 3–5cm(leafindex2.4–3.5),chartaceous,notverruculose,dull, brown above and below, glabrous above, sparsely covered with appressed hairs below, primary vein rather densely so, base acutetoslightlyattenuate,apexacuminate(acumen5–15 mmlong),primaryveinimpressedabove,secondaryveins distinct,8–12oneithersideofprimaryvein,slightlyraised above,smallestdistancebetweenloopsandmargin2–3mm, tertiaryveinsslightlyraisedabove,reticulate.Flowers solitary inaxilsofleavesoronleaflessbranchlets;pedicels30–60mm long,1–1.5mmdiam,fruitingpedicelstoc.2mmdiam,rather densely to sparsely covered with appressed hairs, articulated at0.1–0.3fromthebase,bracts5–6,soonfalling,exceptionallytheupperonefoliaceous,narrowlyelliptic,c.30mmlong; flowerbudsdepressedovoid;sepalsbasallyconnateorfree, broadlyovate-triangular,3–10by4–10mm,appressed,outer sidedenselytoratherdenselycoveredwithappressedhairs; petals pale green, maturing cream, creamy yellow, or beige in vivo,ovate,15–20by8–9mm,outersidedenselycovered withappressed,brownhairs;stamensc.2mmlong,connective shieldpapillate.Monocarps20–40,green,maturingreddishin vivo,browninsicco,ellipsoid,10–11by5.5–6mm,glabrous, apexapiculate(apiculum0.5–1mmlong),wallc.0.2mmthick, stipesc.4by1mm.Seedellipsoid,c.7by4mm,brown,pitted, raphenotdistinctfromrestofseed. Distribution—Peru(Cusco). Habitat&Ecology—Inmontaneprimaryorsecondary,cloud forest,onrichsoil.Atelevationsof800–1750m.Flowering: January,March,June,September;fruiting:July,September, October. Vernacularnames—Notrecorded. Other specimens examined. Peru, Cusco,Paucartambo,Kosñipata,San Pedro,1650m,2Sept.2006,Farfán R. et al. 2558(CUZ,USM,WAG);Paucartambo,Kosñipata,Callanga,1250m,23–30June2008,Farfán R. et al. 4088(CUZ,USM,WAG);Paucartambo,Kosñipata,SanPedro,1650m,1Sept. 2006, Garcia C. et al. 946(CUZ,USM,WAG);Paucartambo,Kosñipata,San Pedro,1650m,6Sept.2006,Garcia C. et al. 990(CUZ,USM,WAG);La Convención,Distr.Echarate,Kepashiato,Puguientimari-Pomoreni,800m, 24Mar.2007,Valenzuela et al. 9413(WAG);LaConvención,Distr.Echarate,Puyentimari,831m,23Jan.2008,Valenzuela et al. 10766(WAG);La Convención,Distr.Echarate,Cigakiato,1000m,29Jan.2008,Valenzuela et al. 10962(WAG). Note—Guatteria cuscoensisoccursinthePeruvianDepartment of Cusco high in the tropical montane cloud forests of the ManuNationalPark,andnearKepashiato.Itisrecognizable from other Andean species of Guatteria by its relatively long pedicelsandshortstipes. 46. Guatteria darienensisSusanaArias&Maas—Map8 Guatteria darienensisSusanaArias&MaasinAriasG.etal.(2014)150,f.1. —Type:Brand & Ascanio 439 (holoHUA;isoJAUM,MO),Colombia, Antioquia,Turbo,CarreteraTapóndelDarién,sectorRíoLeón-Lomas Aisladas,km36,20m,27Aug.1983. Guatteria sp. 2Erkens(2007)208. Treeorrarelyashrub3–16mtall,4–18cmdiam;youngtwigs rather densely to sparsely covered with appressed hairs, soon glabrous.Leaves:petiole2–7mmlong,1–3mmdiam;lamina narrowlyelliptictonarrowlyobovate,rarelyelliptic,8–21by 2.5–8cm(leafindex2.5–3.6),chartaceous,notverruculose, dull to shiny, greyish, dark green or brown above, dull, pale brown or brown below, glabrous above, sparsely covered with appressed hairs to almost glabrous below, base acute to attenuate,rarelyobtuse,apexacuminate(acumen5–15mm long),primaryveinimpressedabove,secondaryveinsdistinct, 10–14oneithersideofprimaryvein,impressedtoslightlyraised above,smallestdistancebetweenloopsandmargin3–5mm, tertiary veins inconspicuous or conspicuous, slightly raised above,reticulate.Flowers in 1–2-flowered inflorescences in axilsofleaves;pedicels30–75mmlong,c.1mmdiam,fruiting pedicels1–1.5mmdiam,ratherdenselytosparselycovered withappressedandsomeerecthairs,articulatedat0.2–0.3 fromthebase,bracts3–7,soonfalling,onlyoneuppermost bractseen,elliptic,c.7mmlong;flowerbudsdepressedovoid; sepalsbasallyconnateorfree,broadlyovate-triangular,3–7by 4–6mm,reflexed,outersidedenselycoveredwithappressed hairs;petalsgreen,maturingyellowinvivo,narrowlyelliptic, 12–26 by 5–10 mm, outer side densely covered with appressed hairs;stamens1.5–2mmlong,connectiveshieldpapillate. Monocarps35–50,greenorpinkish,maturingredtopurplein vivo,browninsicco,ellipsoidorobovoid,7–10by4–5mm, sparselycoveredwithappressedhairs,apexapiculate(apiculumc.0.5mmlong),wallc.0.1mmthick,stipes7–15(–20)by 0.5–1mm.Seedellipsoid,6–9by3.5–4mm,shiny,reddish brown,pitted,raphenotdistinctfromrestofseed. Distribution—Panama,Colombia(Antioquia,Chocó,Córdoba). Habitat&Ecology—Inlowlandorpremontaneforest,along rivers,creeks,onsteepslopesandswampysoils.Atelevations of20–800(–1200)m.Flowering:throughouttheyear;fruiting: throughouttheyear. Vernacularnames—Colombia:Guanabanillo(Zuluaga R. 1226),Yayablanca(D.A. Sánchez S. & Hoyos M. 536). Uses—Colombia:Woodisusedforbuildinghousesand thebarkforlashingmaterialandheadstraps(D.A. Sánchez S. & Hoyos M. 536).Thedecoctionofanypartoftheplantis usedagainstmalaria(Fonnegra et al. 8877). Notes—Guatteria darienensis is well recognizable by relativatelylongandslenderpedicels(upto75mmlonginfruit), relativelysmallmonocarpswithaverythinwall(c.0.1mmthick) andlongandslenderstipes.
68 Blumea–Volume60/1–3,2015 Fig. 31 Guatteria cuscoensisMaas&Westra.Floweringbranch(Farfán et al. 4088, WAG).
69 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria SomePanamanianmaterialcollectedintheprovinceofPanama from[“?]ParqueNacionalAltosCampana(Espinosa et al.720, FLORPAN 2962, Galdames et al. 3081, 4136, Van der Werff et al. 6198, 6933),andonecollectionfromtheprovinceofVeraguas(McPherson 12812),isaberrantbyrelativelysmallerand thickerleaves(8–12by2.5–4.5cm)withanacutetoobtuse base.Asallflower,fruitandseedcharactersmatchthespecies concept of G. darienensis very well, we refrained from giving thismaterialastatusofitsown. 47. Guatteria decurrensR.E.Fr.—Map11 Guatteria decurrensR.E.Fr.(1938)720;Maas&Westra(2011)119,f.3c,5. —Type:Killip & Smith 29585(holoUS;isoF,NY),Peru,Loreto,Soledad, onRíoItaya,110m,20–22Sept.1929. Guatteria rugosaR.E.Fr.(1939)501.—Type:Krukoff 4664(holoS;isoBM, F,G,K,M,MO,NY,U,US),Brazil,Amazonas,BasinofRioJuruá,near mouthofRioEmbira,tributaryofRioTarauacá,4June1933. Tree2–35mtall,2–60cmdiam;youngtwigsdenselytorather densely covered with rough, brown, erect to half-appressed hairs(‘hirsute’)toc.3mmlong,finallyglabrous.Leaves: petioles1–7mmlong,2–3mmdiam;laminanarrowlyellipticobovateornarrowlyelliptic,rarelyelliptic,11–34by4–11cm (leafindex2.2–5.5),chartaceous,brownishgreenabove, brownish green to pale brown below, very densely to densely verruculose on both sides, glabrous above, densely to rather densely, sometimes sparsely covered with rough, erect to appressedhairstoc.2mmlongbelow,baseattenuatetoacute, apexacuminate(acumen10–40mmlong),primaryveinimpressedabove,secondaryveinsdistinct,13–30oneachside, flat to slightly impressed above, forming a marginal vein, at a smallest distance of 1–5 mm from the margin, tertiary veins inconspicuousandhardlyvisibleabove,reticulate.Flowers in1(–2)-floweredinflorescencesinaxilsofleaves;pedicels 12–30mmlong,1.5–2mmdiam,fruitingpedicels25–40(–50) mm long, rather densely to sparsely covered with erect, halfappressed, or sometimes appressed hairs to 2 mm long, articulatedat0.1–0.3fromthebase,bractsc.5,soonfalling,not seen;flowerbudsbroadlyovoidtobroadlyellipsoid;sepalsfree or basally connate, ovate-triangular to broadly ovate-triangular, 5–12by5–7mm,appressedoratlastreflexed,outerside densely to rather densely, sometimes sparsely covered with erect,half-appressed,orsometimesappressedhairstoc.2mm long;petalsgreen,yellowishgreen,maturingyelloworcreamy yellow in vivo, elliptic to ovate-elliptic or narrowly so, 15–25 by 6–13mm,outersidedensely(particularlythebase)torather denselycoveredwithappressed,palebrownhairs;stamens 1.5–2.5mmlong,connectiveshieldpapillate.Monocarps 15–60, green, maturing purplish black to black in vivo, brown in sicco,ellipsoid,15–25by7–12mm,sparselytoratherdensely covered with rough, appressed hairs, apex rounded to apiculate (apiculum<0.2mmlong),wall0.3–1(–1.5)mmthick,stipes 5–15by1–2mm.Seedellipsoid,15–25by7–12mm,dark brown to brown, shiny, longitudinally and transversely grooved, raphenotdistinctfromrestofseed. Distribution—AmazonianColombia(Amazonas),Ecuador(Carchi,Morona-Santiago,Napo,Pastaza,Sucumbios, Zamora-Chinchipe),Peru(Amazonas,Loreto,Pasco),Brazil (Acre,Amazonas). Habitat&Ecology—Inprimary,non-inundated,lowlandor rarely premontane forest, sometimes in periodically inundated forest,onlateritictoclayey,orrarelywhitesandysoil.Atelevationsof100–500m,rarely(inEcuador)to1300m.Flowering: AugusttoApril;fruiting:ApriltoJanuary. Vernacularnames—Ecuador:Caracaspi(Alvarez et al. 2407, Zuleta 212).Peru:Carahuasca(Ellenberg 2852, Tello 1828),Espintana(Ayala et al. 2546),Yais(Shuarname)(R. Rojas et al. 36). Note—Guatteria decurrens can be confused with G. guianensis,butitdiffersbytheindument.Forfurtherinformation seeunderthelatterspecies. 48. Guatteria delicatulaMaas&Westra,sp. nov. —Fig.24g, 32;Map11 Frutex parva pro genere, foliis parvis supra conspicue venosis venulis omnino prominentibusmonocarpiisinstipitibusarticulatisbenedistincta.—Typus: Solomon 9282(holoU;isoLPB,MO),Bolivia,LaPaz,Prov.NorYungas, 13.7kmNWofSanPedroonroadthroughIncahuara-Mejillonesandalong trailto12deOctubre,1500m,15–16Jan.1983. Shrubc.1.5mtall;youngtwigsratherdenselycoveredwith appressedtosemi-erectwhitishbrownhairstoc.1mmlong, eventuallybecomingglabrous.Leaves:petiole2–3mmlong, c.1mmdiam;laminanarrowlyelliptic,5–7.5by1.5–2cm (leafindex2.7–3.8),subcoriaceous,notverruculose,dark brown and shiny above, brown and dull below, glabrous above except for rather densely to sparsely hairy primary vein, rather denselycoveredwithappressedtosemi-erecthairstoc.1mm long on primary vein, elsewhere rather densely to sparsely so below,baseacutetoobtuse,apexacuminate(acumen5–10 mmlong),primaryveinflattoslightlyraisedabove,secondary veins distinct, 10–12 on either side of primary vein, raised above, smallest distance between loops and margin 1–2 mm, tertiaryveinsraisedabove,reticulate.Flowers solitary in leaf axils,onlyseeninbudandfruitingstage;fruitingpedicels c.25mmlong,c.1mmdiam,ratherdenselycoveredwith appressedhairs,ultimatelyglabrous,articulatedatc.0.1from thebase,bractssoonfalling,notseen,notcountable;flower budsgreeninvivo,onlyonelooseyoungbudseen,ovoid,c.3 by3mm,pointed;flowersnotseen.Monocarps≤20,purplish invivo,blackinsicco,ellipsoid,5–7by4–4.5mm,sparsely covered with appressed hairs, soon glabrous, apex apiculate (apiculumc.0.5mmlong),wallc.0.3mmthick,stipes8–10 by1–1.5mm,articulatedtothemonocarpbody.Seed ellipsoid, c.6by4mm,brown,transverselygrooved,raphenotdistinct fromrestofseed. Distribution—Bolivia(LaPaz). Habitat&Ecology—In‘mixtureofmoistforestwithlittle disturbanceandsmallfields’,insecondarygrowth.Atanelevationofc.1500m.Flowering:unknown;fruiting:January. Vernacularnames—Notrecorded. Note—Guatteria delicatula, only known from the type col lection, is oneofthesmallest(c.1.5mtall!)speciesinthegenus, and is recognizable by small leaves with a distinctly raised venation on the upper side and by the stipes which are articulate tothemonocarpbody.Thearticulationisclearlyvisibleasa constrictionbetweenthestipesandthemonocarpbody.This isnotoftenseeninthegenusasawhole. 49. Guatteria denudataR.E.Fr.—Map12 Guatteria denudataR.E.Fr.(1957b)601;Maas&Westra(2010)264,f.6. —Type:Schultes & Cabrera R. 17374(holoS;isoCOAH,GH,NY,US), Colombia,Vaupés,RíoPiraparaná,tributaryofRíoApaporis,headwaters ofCañoTeemeeña,10Sept.1952. Treesmall(1.5m)tomedium-sized,diamnotrecorded;young twigs sparsely covered with appressed hairs or very soon glabrous.Leaves:petiole5–15mmlong,1–2mmdiam;lamina elliptictonarrowlyelliptic,ornarrowlyoblong-elliptic,7–15 by3–6cm(leafindex1.7–2.8),coriaceous,notverruculose, shiny and dark brown above, brown below, glabrous above and below, base obtuse, extreme base slightly attenuate, apex acuminate(acumen10–20mmlong),primaryveinimpressed above, secondary veins indistinct, 12–15 on either side of primaryvein(hardlycountable),slightlyraisedabove,smallest
70 Blumea–Volume60/1–3,2015 Fig. 32 Guatteria delicatulaMaas&Westra.Floweringandfruitingbranch (Solomon 9282,holotypeU).
71 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria distance between loops and margin 2–5 mm, tertiary veins flat toslightlyraisedabove,reticulate.Flowersin1(–2)-flowered inflorescencesinaxilsofleavesoronleaflessbranchlets;pedicels6–22mmlong,1–1.5mmdiam,fruitingpedicelstoc.2mm diam,articulatedat0.1–0.2fromthebase,ratherdensely covered with appressed, brownish hairs to glabrous, bracts 4–5,soonfalling,notseen;flowerbudsnotseen;sepalsfree, broadlyovate-triangular,3–4by4–6mm,appressed,outer sidesparselycoveredwithappressedhairs;petalsgreenin vivo,narrowlyoblong-elliptictooblong-elliptic,10–22by4–13 mm, outer base densely covered with appressed hairs, otherwisesparselyso;stamens1.5–2mmlong,connectiveshield papillate.Monocarps25–30,colourinvivonotrecorded,dark browntobrowninsicco,ellipsoid,8–9by3–4mm,glabrous, apexrounded,wallc.0.2mmthick,stipes2–3by1–1.5mm. Seedellipsoid,8–9by4mm,brown,rugosetopitted,raphenot distinctfromrestofseed. Distribution—AmazonianColombia(Vaupés). Habitat&Ecology—Vegetationtypenotrecorded.Atabout sealevel.Flowering:September;fruiting:September. Vernacularname—Colombia:Feé-ka-no(Barasname) (Schultes & Cabrera R. 17374). Note—Guatteria denudata(includedbyR.E.Friesinsect. Pteropus)isapoorlyknownspeciesfrom2gatheringsonly.It is quite noteworthy by its very thick, shiny, glabrous leaves with anobtusebase,theextremebasebeingslightlyattenuate. 50. Guatteria discolorR.E.Fr.—Map12 Guatteria discolor R.E.Fr.(1939)509,f.33a;Maasetal.(2007)638;Erkens etal.(2008)483,pl.2.—Type: Krukoff 7047(holoS;isoB,BR,F2sheets, G,K,MO,NY,U,US2sheets),Brazil,Amazonas,BasinofRioMadeira, Mun.Humaitá,onplateaubetweenRioLivramentoandRioIpixuna,7–18 Nov.1934. Tree13–30mtall,25–80cmdiam,oftenwithplankbuttresses; young twigs densely to rather densely covered with erect, reddishbrown,curlyhairs,soonglabrous.Leaves: petiole 5– 10(–15)mmlong,2–4mmdiam;laminanarrowlyelliptic,15–30 by4–10cm(leafindex2.8–3.7),coriaceous,sparselyto densely verruculose or not verruculose, dull, grey to greyish black above, dark brown below, rather densely covered with erect and appressed, brown hairs above, soon glabrous, densely covered with erect, reddish brown, curly hairs below, base acute, extreme base attenuate, shortly decurrent along petiole,apexacutetoacuminate(acumen10–15mmlong), primary vein impressed to raised above, secondary veins distinct, 15–25 on either side of primary vein, impressed above, smallest distance between loops and margin 1–2 mm, tertiary veinsflattoslightlyraisedabove,reticulatetopercurrent.Flowersin1–3-floweredinflorescencesinaxilsofleaves;pedicels 10–20mmlong,1–2mmdiam,fruitingpedicelstoc.25mm long,3–5mmdiam,denselycoveredwitherect,reddish brown,curlyhairs,articulatedat0.3–0.5fromthebase,bracts 5–6, soon falling, broadly elliptic to elliptic, basal bracts 1–2 mmlong,upperones8–11mm;flowerbudsovoid,pointed; sepalsfree,broadlyovate-triangulartoovate-triangular,6–13 by 5–6 mm, spreading to reflexed, outer side densely covered witherect,reddishbrown,curlyhairs;petalsgreen,maturing yellowishinvivo,narrowlyovatetonarrowlyobovate,15–40 by 6–16 mm, outer side densely covered with erect, reddish brown,curlyhairs;stamensc.2mmlong,connectiveshield papillate,umbonateornot.Monocarps10–30,darkgreen, maturing blueish black to black in vivo, dark brown to black insicco,ellipsoidtonarrowlyso,15–30by7–15mm,surface irregularly wrinkled in sicco, sparsely covered with erect, reddishbrown,curlyhairs,apexroundedtoapiculate(apiculum <0.5mmlong),wall0.5–1mmthick,stipes2–10by1–2mm. Seednarrowlyellipsoidtoellipsoid,13–17by6–9mm,brown, smooth,raphenotdistinctfromrestofseed. Distribution—Colombia(Amazonas,Vaupés),FrenchGuiana,AmazonianPeru(Loreto,MadredeDios),Brazil(Acre, Amazonas,Rondônia),Bolivia(LaPaz,Pando). Habitat&Ecology—Innon-inundatedforest,sometimesin campinaranavegetation(Brazil,Acre),onclayey,sometimes onsandysoil.Atelevationsof0–600m.Flowering:Octoberto January,April;fruiting:January,ApriltoJuly,October,November. Vernacularnames—Bolivia:Piraquinablanca(Terceros 1398).Brazil:Envira(Krukoff 7047, Unknown collector 17, 21), Envira-bobo(Boom et al. 8635),Envira-fofa,Envira-mole-dafolha-grande(Daly et al. 7794),Envira-preta,Envira-rolinha, Envireira(Daly et al. 7037).FrenchGuiana:Matau’i,(Wayãpi name)(Grenand 554),Miret(Palikurname). Notes—Guatteria discolor can be distinguished fairly easily by its 2-coloured leaves which are covered with an indument oferect,reddishbrown,curlyhairsonthelowerside.Furthermore, the monocarps are fairly large and wrinkled when dry, the latterfeaturenotoftenseeninthegenus.Athirdremarkable aspectarethedistinctlypointedflowerbuds.Seealsounder G. chrysophylla. Guatteria discolorwasplacedbyFries(1939:509)intoasectionofitsown,namelysect.Dichrophyllum, characterized by 2-coloured leaves, a persistent indument of brownish hairs, and byleaveswhichareverruculoseontheupperside.Thelatter feature appears to be quite variable, the density of verruculae ranging from densely to sparsely, and we even noticed leaves withoutanywarts. AvegetativecollectionfromBolivia(Chatrou et al. 438(MO, U))hasverylargeleavesmeasuring40by15cm! 51. Guatteria dolichopodaDonn.Sm.—Plate3e;Map12 Guatteria dolichopoda Donn.Sm.(1897)2;R.E.Fr.(1939)357.—Type: Donnell Smith 6429(lectoUS,selectedbyErkens2007;isolectoB,C, G,GH,K2sheets,M,MO,US),CostaRica,SanJosé,LaConcepción, LlanurasdeSantaClara,250m,Feb.1896. Guatteria tonduziiDiels[var.tonduzii ](1931)75;R.E.Fr.(1939)356.—Type: Tonduz 17680(holoB;isoBM,F,G3sheets,K,US2sheets,Z),Costa Rica,Heredia,‘CollinesdeTremedal,prėsSanRamon’,1300–1400m, 10Apr.1913. Guatteria tonduziiDielsvar.leptopusR.E.Fr.(1939)357.—Type:Pittier 10958(holoM;isoBR2sheets,C,G3sheets,US),CostaRica,Puntarenas,CañasGordas,1100m,Mar.1897. Guatteria dolichopodaDonn.Sm.var.microspermaR.E.Fr.(1939)358,f.8a. —Type:Tonduz 9166(holoS;isoBM,BR3sheets,C,K2sheets,M, P,S,US7sheets,W),CostaRica,Limón,Shiroles(‘ForêtsaShirores, Talamanca’),100m,Feb.1895. Guatteria rigidipesR.E.Fr.(1939)358,t.21.—Type:Skutch 2553(holoS; isoK,MO,NY,US),CostaRica,SanJosé,vicinityofElGeneral,1040m, Feb.1936. Treeorshrub3–20mtall,10–20(–100)cmdiam;ratherdenselytosparselycoveredwitherect,brownhairsc.0.5mmlong, finallyglabrous.Leaves:petiole2–6mmlong,1mmdiam; laminanarrowlyelliptic,sometimesnarrowlyovate,11–14by 2–5cm(leafindex2.8–5.5),chartaceous,notverruculose, dull, greyish black to greyish brown above, greyish to brown below, glabrous above, but primary vein and sometimes the secondary veins covered with erect, brown hairs, sparsely coveredwitherect(andappressed),brownhairsbelow,base acutetoattenuate,apexacuminate(acumen10–20mmlong), primary vein impressed to flat above, secondary veins indistinct, 10–15 on either side of primary vein, flat to slightly raised above,smallestdistancebetweenloopsandmargin2–3mm, tertiaryveinsraisedabove,reticulate.Flowersin1(–2)-flowered inflorescencesinaxilsofleaves;floweringandfruitingpedicels
72 Blumea–Volume60/1–3,2015 Fig. 33 Guatteria dotanaN.Zamora&Erkens.Fruitingbranch(Estrada & Valverde 1084,isotypeMO).
73 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria 30–60mmlong,c.1mmdiam,ratherdenselytosparsely covered with erect hairs, becoming almost glabrous in fruit, articulatedat0.1–0.2fromthebase,bracts4–6,soonfalling, rarelypresent,uppermostbractnarrowlyelliptic,toc.8mm long;flowerbudsdepressedovoid,slightlypointed;sepalsfree, ovate-triangulartobroadlyovate-triangular,5–7by3–6mm, reflexed, outer margins revolute, outer side rather densely to sparselycoveredwithappressedhairs;petalsgreen,maturing yellowinvivo,narrowlyoblong-elliptictonarrowlyovate,14–30 by4–11mm,outersidedenselytoratherdenselycovered withappressed,whitehairs,particularlytowardsthebase;stamens1.5–2mmlong,connectiveshieldpapillate.Monocarps 75–100,blackinvivo,blackinsicco,ellipsoid,7–11by4–5 mm, sparsely covered with appressed hairs, soon glabrous, apexapiculate(apiculum<1mmlong),wall0.1–0.2mmthick, stipes15–20by1mm.Seedellipsoid,6–9by4–5mm,dark brown,pitted,rapheraised. Distribution—Honduras,CostaRica,Panama. Habitat&Ecology—Inlowlandorlowermontaneforest.At elevationsof0–1600m.Flowering:JanuarytoJuly;fruiting: DecembertoJuly. Vernacularnames—Notrecorded. Note—Guatteria dolichopoda is recognized by its quite long pedicelsandyoungtwigscoveredwitherecthairsc.0.5mm long.IthasoftenbeenconfusedwithG. tomentosa, see under thatspeciesforadiscussion. 52. Guatteria dotanaN.Zamora&Erkens,sp. nov.—Fig.24h, 33;Map11 Ramulisnovellisplusminusvefractiflexispilisappressisdenseobtectis, floribus breviter pedicellatis monocarpiis breviter stipitatis facile recognoscenda.—Typus:Estrada & Valverde 1084(holoCR;isoMO),CostaRica, SanJosé,Tarrazu,LlanosdeSantaMaríadeDota,CerroNene,Fincade HenryMarín,900–1300m,19Aug.1997. Tree4–7mtall,diamnotrecorded;youngtwigsdenselyto rather densely covered with brown, appressed hairs, slightly zigzagging.Leaves:petiole5–8mmlong,c.2mmdiam; laminanarrowlyoblong-elliptic,10–21by3–8cm(leafindex 2.6–3.3),coriaceous,notverruculose,shinytodull,greyish green above, pale brown below, glabrous above, densely to sparsely covered with appressed, reddish hairs mainly along the primary vein below, base acute to obtuse, apex acuminate (acumen5–15mmlong),primaryveinflattoslightlyraised above,secondaryveinsdistinct,11–17oneithersideofprimary vein, slightly raised above, smallest distance between loops and margin2–3mm,tertiaryveinsslightlyraisedabove,reticulate. Flowerssolitaryinaxilsofleavesoronleaflessbranchlets; pedicels0–7mmlong,c.1mmdiam,fruitingpedicels5–15 mmlong,2–4mmdiam,denselycoveredwithappressed,pale brownhairs,articulatedat0.3–0.5fromthebase,bracts5–6, soonfalling,notseen;flowerbudsovoid;sepalsfree,broadly ovate-triangular,6–8by5–7mm,reflexed,outersidedensely coveredwithappressedhairs;petalsgreenishtocreamin vivo,ovate-elliptic,7–12(–14)by6–8mm,outersidedensely coveredwithappressedhairs;stamensc.2mmlong,connectiveshieldhairy.Monocarps 15–25, green, maturing purplish blackinvivo,blackinsicco,ellipsoid,15–17by6–10mm, glabrous,apexrounded,wall0.1–0.2mmthick,stipes4–6by 1mm.Seedellipsoid,12–14by6–7mm,dark,shinybrown, transverselygrooved(topitted),rapheraised. Distribution—CostaRica. Habitat&Ecology—Wetforest(‘bosquemuyhúmedo’).At elevationsof500–1300m.Flowering:February,May;fruiting: August,October. Vernacularnames—Notrecorded. Other specimens examined. Costa riCa, Puntarenas, Buenos Aires, Colinas,FilaRetinto,partealtadelaFila,1017m,22May2012(fl),Lobo 3343(CR).San José,Dota,SanIsidro,CerroTijerilla,900–1000m,21Oct. 1998(fr),Estrada & Solano 1817(CR,MO);Tarrazú,ZPCerroNara,Cuenca delNaranjoyPaquita,Dota,2kmdelcruceàRíoBlanco,caminoaSan Isidro,300–600m,27Feb.2005,Santamaría & Morales 682 (CR,L);Dota, ReservaForestalLosSantos,CuencadelSavegre,QuebradaBomba,cruce aFilaMonayLaBomba,500m,28Feb.2005(fl),Santamaría & Morales 757(INB);Dota,ReservaForestalLosSantos,CuencadelSavegre,San IsidrodeDota,albergueTimaná,800–900m,1Mar.2005,Santamaría & Morales 809 (CR,L). Note—Guatteria dotana,endemictoCostaRica,canbe distinguished by zigzagging young twigs which are densely covered with appressed, brownish to reddish hairs, very shortly pedicellateflowersandshortlystipitatemonocarps. 53. Guatteria duckeana R.E.Fr.—Fig.34;Map13 Guatteria duckeana R.E.Fr.(1939)468,f.22f,g. —Type:Ducke RB 29019 (holoS;isoRB),Brazil,Amazonas,Manaus,PontedoMindu(‘locoPonte deMindú,adripamrivuli’),Nov.1935. Tree6–15mtall,c.7.5cmdiam;youngtwigsdenselytorather denselycoveredwitherecthairs,soonglabrous.Leaves: petiole 3–10mmlong,1–2mmdiam;laminanarrowlyelliptictonarrowlyovate,10–22by4–6cm(leafindex2.5–4),coriaceous, not verruculose, shiny, dark brown above, brown below, rather densely to sparsely covered with erect hairs above, soon glabrous, densely to rather densely covered with erect hairs below (hairssimple,intermixedornotwithfewbundlesof2hairs), base acute, obtuse, or attenuate, basal margins mostly revolute,apexacuminate(acumen5–15mmlong),primaryvein flatabove,secondaryveinsindistinct,8–15oneithersideof primary vein, flat above, smallest distance between loops and margin2–4(–8)mm,tertiaryveinsflatabove,reticulate.Flowersin1–3-floweredinflorescencesinaxilsofleaves;flowering andfruitingpedicels5–8mmlong,1–2mmdiam,denselycovered with erect hairs, soon becoming subglabrous, articulated at0.3–0.6fromthebase,bractsc.5,soonfalling,onebasal bractseen,broadlyelliptic,c.2mmlong;flowerbudsdepressed ovoid;sepalsfree,broadlyovate-triangular,3–5by3–5mm, reflexed, outer side densely covered with appressed hairs, marginsrevolute;petalsgreenwhenyounginvivo,oblongelliptic,8–20by5–8mm,outersidedenselycoveredwitherect hairs,soonbecomingsubglabrous,marginsrevolute;stamens 1–1.5mmlong,connectiveshielddenselyhairy.Monocarps 10–20, green in vivo, brown in sicco, ellipsoid to fusiform, 9–18by6–10mm,longitudinallywrinkledinsicco,sparsely covered with erect hairs, soon becoming glabrous, apex roundedtoapiculate(apiculumto.1mmlong),wallc.1mm thick,stipes0–4by2mm.Seedellipsoid,c.15by7mm,pale brown,smooth,raphenotdistinctfromrestofseed. Distribution—Brazil(Amazonas,ManausRegion). Habitat&Ecology—Innon-inundated,primaryorsecondary terrafirmeforest,onecollectionfromcampinaranavegetation, onsandysoil.Atelevationsofaboutsealevel.Flowering:January,October;fruiting:January,AugusttoOctober. Vernacularnames—Brazil:Envira(Krukoff 7923),Envireira (L. Coêlho INPA 5805). Notes— Guatteria duckeana together with G. citriodora, G. schomburgkiana and G. stipitata forms part of the G. schomburgkianacomplex,seeTable4(p.59).Thisgroup,placedby Fries(1939)insect.Cephalocarpus which originally consisted of eight species, has as its main features short-pedicelled flowersandsessiletoveryshortlystipitatemonocarps.Thedensely hairy connective shield also is a distinctive feature of this group (vspapillateorglabrousinmostGuatteriaspecies).
80 Blumea–Volume60/1–3,2015 index3.8–5.8),chartaceous,denselyverruculose,dull,dark brown or dark green above, paler brown or green below, glabrous above, but the primary vein densely covered with erect, brown hairs, densely covered with erect, brown hairs below, baseacute,rarelyobtuse,apexacuminate(acumen5–20mm long),primaryveinimpressedabove,secondaryveinsindistinct to distinct, 12–20 on either side of primary vein, flat to slightly impressed above, more or less forming a marginal vein, at a smallestdistanceof1–3mmfomthemargin,tertiaryveins inconspicuous,flattoslightlyraisedabove,reticulate.Flowersin1–2-floweredinflorescencesinaxilsofleaves;pedicels 20–40mmlong,1–1.5mmdiam,fruitingpedicelstoc.3mm diam, rather densely to densely covered with erect, brown hairs, articulatedatc.0.3fromthebase,bracts3–6,soonfalling, thebasalones10–14mmlong,the2upperones13–16mm long;flowerbudsdepressedovoid;sepalsfree,broadlyovatetriangular,5–7by5–6mm,appressed,outersidesparsely coveredwithappressed,brownhairs;petalsgreenoryellow invivo,broadlyovatetoovate,9–13by7–10mm,outerside densely covered with erect, brown hairs to almost glabrous, exceptforthehairymargins;stamensc.1.5mmlong,connective shieldpapillate.Monocarps40–50,darkgreen,maturingblack invivo,blackishinsicco,narrowlyellipsoidtoellipsoid,5–8 by2–4mm,glabrous,exceptforafewscattered,appressed hairs,apexacutetoapiculate(apiculum0.1–0.2mmlong), wall0.1–0.5mmthick,stipes2–5byc.1mm.Seed narrowly ellipsoid,6–8by2–4mm,dull,darkbrown,rugulose,raphe notdistinctfromrestofseed. Distribution—PacificcoastofColombia(ValledelCauca). Habitat&Ecology—Inwet,tropicalrainforest,alongthe marginofriversandalongroads,onloamytoclayeysoil.At elevationsof0–350m.Flowering:FebruarytoAugust;fruiting: February,April,May,July,August,October. Vernacularnames—Colombia:Cargadero(Gentry et al. 48326),Cargaderonegro(D.A. Sánchez S. 676). Notes—Guatteria elegantissima can very easily be recognized by the very densely verruculose and shortly petiolate leaves which are densely covered with erect, brown hairs on thelowerside.Moreover,themonocarpsarequitecharacteristic inoftenbeingnarrowlyellipsoidandshortlystipitate.Guatteria elegantissima looks somewhat similar to G. cuatrecasasii, differing by smaller leaves and a less dense indument of shorter hairs than in G. cuatrecasasii. Wisum et al. 24(MO)fromEcuador,Morona-Santiagooccurs at quite some distance from Valle del Cauca in Colombia were allothergatheringsweredone.Itisalsoremarkablebecauseit occursatanelevationof>1200mwhileinColombiacollections werefoundatelevationsbelow400m.Apartfromthisitmatches welltheothermaterialalbeitwithslightlylargermonocarps. 59. Guatteria elongataBenth.—Map14 Guatteria elongata Benth.(1843)359;R.E.Fr.(1939)431,t.30.—Type:R.H. Schomburgk I 962(holoK;isoB,BM,E,F,G,K,L,P,U,US,W),Brazil, RioNegro,withoutfurtherexactlocation,anno1839. Treeofunknownheightanddiam;youngtwigsdenselycovered withappressed,brownhairstoc.1mmlong,soonglabrous. Leaves:petiole5–15mmlong,1–2mmdiam;laminanarrowlyoblong-elliptic,12–23by5–6.5cm(leafindex3.8–5), chartaceous, not verruculose, shiny above, brown on both sides, glabrous above, sparsely covered with appressed, brown hairstoc.1mmlongbelow,baseacute,extremebaseattenuate,apexacutetoacuminate(acumentoc.10mmlong),primary vein impressed above, secondary veins distinct, 12–15 on either side of primary vein, slightly raised above, smallest distancebetweenloopsandmargin3–4mm,tertiaryveins inconspicuous,slightlyraisedabove,reticulate.Flowers in (1–)2–4-floweredinflorescencesinaxilsofleavesoronleaflessbranchlets;pedicels2–8mmlong,1.5–2mmdiam, denselycoveredwithappressed,brownhairstoc.1mmlong, articulatedat0.5–0.6fromthebase,bractsc.6,soonfalling, basalbract(oneseen)verybroadlyovate,c.1mmlong;flower budsdepressedovoid;sepalsfree,broadlyovate-triangular, 4–5by5–6mm,appressed,outersidedenselycoveredwith appressed,brownhairstoc.1mmlong;petalscolournot recorded,obovate,12–18by6–8mm,outersidedensely coveredwithappressed,brownhairstoc.1mmlongatthe base,towardstheapexthehairswhitish;stamensc.1.5mm long,connectiveshieldpapillate.Monocarps and seednotseen. Distribution—AmazonianBrazil(Amazonas). Habitat&Ecology—Vegetationtypenotrecorded.Atan elevationofaboutsealevel.Flowering:unknown;fruiting:unknown. Vernacularnames—Notrecorded. Notes—Guatteria elongatawasplacedbyFries(1939)into asectionofitsown,sect.Pycnantha(=‘flowerscrowdedtogether’),becauseofitsmany-floweredinflorescencesandvery shortpedicels.Asthespecieshasneverbeencollectedsince and as the material available is incomplete we cannot place it yetanywhereinthegenus.Itshowssuperficiallysomefeatures of G. subsessilis(theshortpedicels)butfromthatspeciesitdiffersbyacompletelydifferentindument(long,brown,appressed hairsinleaves,flowers,pedicels,sepals,etc.inG. elongata vs almost completely glabrous in all parts of G. subsessilis). Several collections of this species have been labelled as ‘BritishGuiana’,butaccordingtoVanDam(2002:88,89)the RobertSchomburgktypecollectioncomesfromtheRioNegro inAmazonianBrazil.Theexactlocationcouldnotbetraced, butitisprobablysituatedbetweenBarcelosandRioPadauiri, accordingtothedatabyVanDam.Thepositiononthemap hasbeenindicatedaccordingly. 60. Guatteria emarginataLobão,Maas&Mello-Silva—Map 14 Guatteria emarginata Lobão,Maas&Mello-Silva(2010)120,f.1a–f.—Type: Vervloet et al. 2316(holoSPF;isoMBML),Brazil, EspíritoSanto,Nova Lombardia,ReservaBiológicaAugustoRuschi,GoipaboAçu,Boeirão, linhadedivisa(marco53e52)600–800m,29Apr.2003. Treec.17mtall,diamnotrecorded;youngtwigssparselycoveredwithappressed,greyhairs,soonglabrous.Leaves: petiole3–4mmlong,c.1mmdiam;laminanarrowlyobovate,5–9.5 by1.5–3cm(leafindex3.1–3.3),slightlycoriaceous,rather Brazil Colombia Peru Venezuela Panama C osta Rica e mala N i ca r agua Bolivia Guyana Suriname French Guiana Ecuador Map 14 Distribution of Guatteria elegantissima (●),G. elongata (£ ),G. emarginata (■),G. eriopoda(p),G. esmeraldae (u)andG. eugeniifolia (✸).
81 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria densely verruculose, shiny, brown above, pale brown below, glabrous above, densely to sparsely covered with appressed hairs below, base attenuate, basal margins revolute, apex emarginate, primary vein impressed above, secondary veins distinct,9–13oneithersideofprimaryvein,raisedabove, smallestdistancebetweenloopsandmarginc.1mm,tertiary veinsinconspicuous,raisedabove,reticulate.Flowers solitary inaxilsofleaves;pedicels15–30mmlong,0.5–1mmdiam, densely to sparsely covered with appressed, brown hairs, articulatedat0.2–0.3fromthebase,bracts2–4,soonfalling, 5–8mmlong;flowerbudsbroadlyovoid;sepalsfree,broadly triangular,4–6by6–7mm,reflexed,outersidedenselycoveredwithappressed,brownhairs;petalsgreenishpinkinvivo, broadlyelliptic,5–9by6–7mm,outersidedenselycovered withappressed,brownhairs;stamensc.1.5mmlong,connectiveshieldpapillate.Monocarps and seednotseen. Distribution—Brazil(EspíritoSanto). Habitat&Ecology—Inmontane,Atlanticforest.Atelevationsof600–850m.Flowering:April;fruiting:unknown. Vernacularnames—Notrecorded. Note—Guatteria emarginata is characterized by young twigs covered with grey hairs and narrowly obovate, verruculose leaveswithanemarginateapex. 61. Guatteria eriopodaDC.—Map14 Guatteria eriopodaDC.inDunal(1817)130;R.E.Fr.(1939)351,t.19.— Type:Dombey s.n. (holoP;isoF,G,P),Peru,Huánuco,Cuchero(‘circa Cochera’). Tree‘3-orgyalis’;youngtwigsdenselycoveredwitherectto appressed,curly,whitehairstoc.1mmlong,soonglabrous. Leaves:petiole4–6mmlong,1–2mmdiam;laminanarrowly ellipticorelliptic-oblong,12–25by4–8cm(leafindex3.1–3.5), chartaceous, not verruculose, dull, greyish or brownish black above, brown below, sparsely covered, to densely covered on primary vein, with mostly appressed, curly, white hairs to glabrous at last above, densely to rather densely covered with erect to appressed, curly, white hairs below, base obtuse, apex acuminate(acumen10–20mmlong),primaryveinimpressed above,secondaryveinsdistinct,13–17oneithersideofprimary vein, impressed above, smallest distance between loops and margin1–2mm,tertiaryveinsflatabove,reticulate.Flowers in 1–2-flowered inflorescences in axils of leaves or on leaflessbranchlets;pedicels20–25mmlong,1–1.5mmdiam, densely covered with erect, more or less curly, whitish to light brownhairs,articulatedat0.2–0.4fromthebase,bracts4–5?, soonfalling,onlybasalonesseen,broadlyelliptic(-ovate), 1.5–3mmlong;flowerbudsbroadlyovoid;sepalsfree,broadly ovate-triangular,7–8by7–8mm,appressed(asfarasseen inyoungstage),outersidedenselycoveredwithappressedto erecthairs;petalscolournotrecorded,elliptictoobovate,c.16 by 9 mm, outer side densely covered with appressed to erect hairs;stamenstoc.1.5mmlong,connectiveshieldglabrous. Monocarps and seednotseen. Distribution—AmazonianPeru(Huánuco,Loreto). Habitat&Ecology—Notrecorded. Vernacularnames—Notrecorded. Notes—Guatteria eriopodawasplacedbyFriesinsect. Trichoclonia. An incompletely known and perhaps rare species, G. eriopoda somewhat resembles the widely distributed G. ucayalina, but isdistinctfromitbythestrikinglycurlyhairs.Notethatthelong hairs(toc.1mm)ontheleavestendtocomeoffearlierthan the shorter hairs thus giving the false impression of small-haired leaves(seethePoeppigspecimens). 62. Guatteria esmeraldae Maas&Westra,sp. nov.—Fig.38; Map14 Foliis pro genere parvis 15 cm non superantibus atque pedicellis longitudine mediocrismonocarpiisbrevistipitatiscognita.—Typus:Neill & QCNE botany interns 12741(holoU;isoMO,QCNE),Ecuador,Esmeraldas,Muisne,RecintoPalmaJunta,10kmEofMuisne,nearroadtoTonchique,ridgeabove EsteroLaPedorrera,1kmNofhighway,100m,19May2000. Treec.8mtall,diamnotrecorded;youngtwigssparselycoveredwithappressedtoerect,whitehairstoc.0.2mmlong,soon glabrous.Leaves:petiole2–4mmlong,1mmdiam;laminanarrowlyelliptic,10–15by3–5cm(leafindex3–3.3),chartaceous, not verruculose, dull, greyish above, greyish brown below, glabrous above, sparsely covered with appressed hairs below, mainlyalongprimaryveinandtoc.0.5mmlong,baseacute, apexacuminate(acumen10–15mmlong),primaryveinimpressed above, secondary veins distinct, 10–12 on either side of primary vein, impressed above, smallest distance between loopsandmargin3–4mm,tertiaryveinsraisedabove,reticulate.Flowerssolitaryinaxilsofleaves;floweringandfruiting pedicelsc.20mmlong,c.1mmdiam,sparselycoveredwith appressedhairstoc.0.3mmlong,articulatedatc.0.2fromthe base,bracts5–6,soonfalling,notseen;flowerbudsnotseen; sepalsfree,broadlyovate-triangular,c.3by3mm,appressed, outersidedenselycoveredwithappressedhairs;petalspale greeninvivo,ovate-triangular,7–9by4–5mm,outerside denselycoveredwithappressed,whitehairs;stamensc.1.5 mmlong,connectiveshieldpapillate.Monocarpsc.40,bright redinvivo,blackinsicco,ellipsoid,8–9by4mm,sparsely covered with appressed hairs to glabrous, apex rounded, wall c.0.1mmthick,stipes4–6by1mm.Seedellipsoid,c.8by4 mm,brown,transverselygrooved,rugose,rapheslightlyraised. Distribution—Ecuador(Esmeraldas). Habitat&Ecology—Innon-inundated,tropicalwetforest. Atanelevationofc.100m.Flowering:May;fruiting:May. Vernacularnames—Notrecorded. Note—Guatteria esmeraldae, only known from the type collection, is the smallest-leaved species in the area as far as known.Itisbestcharacterizedbythecombinationofrelatively smallleaves(toc.15cmlong),fairlylongpedicels(toc.20 mmlong),monocarpswithstipeslessthanthemonocarpbody inlength,andthetransverselygroovedtorugoseseeds. 63. Guatteria eugeniifoliaA.DC.exR.E.Fr.—Map14 Guatteria eugeniifoliaA.DC.exR.E.Fr.(1938)714;(1939)453.—Type: Poeppig 1859(holoG;isoB,BR,F,G,P2sheets,W),Peru,SanMartín, TocacheNuevo(‘Loreto,Tocache,Maynas’),June1831. Treetoc.20mtall,toc.40cmdiam;youngtwigsdenselycovered with appressed to more or less erect, curly or not, brown hairstoglabrous,becomingglabrousallover.Leaves: petiole 3–10mmlong,1–2mmdiam;laminanarrowlyelliptic,8–20by 3–7cm(leafindex2.5–3),chartaceoustocoriaceous,rather densely to sparsely verruculose, shiny, greyish brown or dark brown above, brown below, glabrous or with some scattered, appressed hairs above, sparsely covered with appressed hairs toc.1mmlongtoglabrousbelow,baseacutetoobtuse,the extremebaseslightlyattenuate,apexacuminate(acumen5–10 mmlong),primaryveinflattoimpressedabove,secondary veinsdistincttoindistinct,8–15oneithersideofprimaryvein, raisedabove,smallestdistancebetweenloopsandmargin2–4 mm,tertiaryveinsraisedabove,reticulate.Flowers in 1–2-flowered inflorescences in axils of leaves or paniculately grouped attheendofatwigbyreductionofleavestobracts;flowering pedicels12–15mmlong,c.1mmdiam,fruitingpedicelsto c.25mmlong,1–2mmdiam,denselycoveredwitherectto more or less appressed, curly, brown hairs to glabrous, articulatedat0.2–0.4fromthebase,bracts5–6,soonfalling,not
82 Blumea–Volume60/1–3,2015 Fig. 38 Guatteria esmeraldaeMaas&Westra.Fruitingbranch(Neill et al. 12741, holotypeU).
83 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria seen;flowerbudsnotseen;sepalsfree,broadlyovate-triangular,4–5by4–5mm,appressedtospreading,outerside denselycoveredwitherect,curly,brownhairs;petalscolour notrecorded,ovatetoelliptic,12–17by5–7mm,outerside denselycoveredwitherect,curly,brownhairs;stamensc.1mm long,connectiveshieldpapillate,umbonate.Monocarps10–75, yellowish green when young in vivo, brown in sicco, ellipsoid, 10–13by8–11mm,sparselycoveredwithappressedhairsto glabrous,apexroundedtoapiculate(apiculum<0.5mmlong), wall1–1.5mmthick,stipes5–7by1–1.5mm.Seed ellipsoid, 10–11by6–7mm,shinybrown,smoothtoslightlypitted. Distribution—AmazonianPeru(SanMartín). Habitat&Ecology—Innon-inundatedforest.Atanelevation of400–500m.Flowering:June;fruiting:June,November. Vernacularnames—Notrecorded. Note—ThetaxonomicpositionofG. eugeniifolia, only knownfromtwocollectionsfromthePeruviantypelocality, needstobeinvestigatedfurther.Theverruculoseleavescould possiblypointtosect.Mecocarpus(Maas&Westra2011).However, the raised secondary and tertiary venation on the upper sideoftheleavesandthesmoothtoslightlypitted(ratherthan muchmoresculptured)seedsdonotfitatallinthissection makingaplacewithinthatsectionunlikely.Fries(1939)placed G. eugeniifoliainsect.Tylodiscus, based on the umbonate staminal shield, setting it apart in that section because of the lack of a sharply delimited glabrous area at the inner base of theinnerpetals. 64. Guatteria ferruginea A.St.-Hil.—Plate4a;Map15 Guatteria ferruginea A.St.-Hil.(1825)38;R.E.Fr.(1939)398.—Type: A.F.C.P. de Saint-Hilaire 2 (holoP;isoS),Brazil,RiodeJaneiro,Nova Iguaçu(‘propevicumAguassu’),July1816. Guatteria glazioviana R.E.Fr.(1900)19,t.2,f.1,2.—Type:Glaziou 6856 (lectoBselectedbyLobãoinMaasetal.2011;isolectoBR,C,G,K,P, S),Brazil,RiodeJaneiro,NovoFriburgo(‘AltodaBoaVistadeNova Friburgo’),23Jan.1874. Guatteria burchellii R.E.Fr.(1939)398.—Type:Burchell 2698(holoK;iso P),Brazil,RiodeJaneiro,Magé(‘FrechaltoMagé’),25Feb.1826. Anoftencaulifloroustreeorshrub3–10(–18)mtall,6–18cm diam;youngtwigsdenselycoveredwitherect,brownhairs. Leaves:petiole6–12mmlong,2–5mmdiam;laminanarrowly elliptictonarrowlyobovate,17–40by5–15cm(leafindex 1.5–3.8),chartaceoustocoriaceous,notverruculose,blackish green above, brown below, sparsely covered with erect hairs to glabrous above, densely covered with erect hairs to sometimes glabrous below, base obtuse to attenuate, apex acuminate (acumen10–20mmlong),primaryveinimpressedabove, secondary veins distinct, 10–20 on either side of primary vein, raised or impressed above, smallest distance between loops andmargin2–4mm,tertiaryveinsraisedabove,reticulate. Flowerssolitaryinaxilsofleavesormostlyin1–2(–more?)- flowered,denselyclusteredinflorescencesonthestem;pedicels15–60mmlong,1–3mmdiam,fruitingpedicelstoc.70 mm long, densely to sparsely covered with erect, brown hairs, articulatedat0.1–0.2fromthebase,bracts2–3,soonfalling, 5–6mmlong;flowerbudsbroadlyovoid;sepalsfree,broadlyto shallowlyovate-triangular,5–10by7mm,reflexed,outerside denselycoveredwitherecthairstoglabrous;petalscreamor yellowinvivo,elliptictoovate,13–20by6–12mm,outerside sparselycoveredwitherect,brownhairs;stamensc.2mmlong, connectiveshieldpapillate.Monocarps 25–55, green, maturing reddishtopurple-blackinvivo,blackinsicco,ellipsoid,8–12by 5–7mm,glabrous,apexapiculate(apiculum0.5–1mmlong), wall0.1–0.3mmthick,stipes10–25by1mm.Seed ellipsoid, 8–12by5–7mm,brown,pitted,rapheraised. Distribution—Brazil(Bahia,EspíritoSanto,MatoGrosso, MinasGerais,Pará,RiodeJaneiro). Habitat&Ecology—Inperiodicallyinundatedornon-inundatedforest,onclayeytosandysoil.Atelevationsof0–1000m. Flowering:throughouttheyear;fruiting:throughouttheyear. Vernacularnames—Notrecorded. Fieldobservations—A. Rylands 30/1980(U)fromUna, Bahia,Brazil:‘frutosmadurosverdeamareladoscomidospor Callithrix e Leontopithecus’. Note—Guatteria ferruginea is the only cauliflorous representativeofthisgenusintheAtlanticforestsofBrazil.Itis characterizedbylargeleaves(17–40by5–15cm),usually densely covered with erect, brown hairs on the lower side, and outer side of sepals and petals also often covered with erect, brown hairs.Guatteria ferruginea is similar to G. pogonopus. Itsharesthelargeleaves,butthepedicelsinthelatterare muchshorter.However,G. pogonopus has glabrous leaves and,moreover,itisnotcauliflorous. 65. Guatteria flabellataErkens&Maas—Map15 Guatteria flabellataErkens&MaasinErkensetal.(2008)484,f.7,pl.2. —Type:Maas et al. 6297(holoU2sheets;isoAMAZ,F,K,MO,WIS), Peru,Loreto,Prov.Maynas,RíoMomón,0–5kmaboveconfluencewith RíoNanay,100m,15Nov.1984. Guatteria sp. AVásquez(1997)100. Tree(3–)5–28mtall,to60cmdiam;youngtwigssparsely coveredwithappressedhairs,soonglabrous.Leaves: petiole 8–12(–15)mmlong,3–5mmdiam;laminanarrowlyelliptic, 20–38by7–16cm(leafindex1.6–3.3),coriaceous,notverruculose, dull, brown above and below, glabrous above, sparsely to rather densely covered with appressed, very short hairs below,basecordatetoobtuse,apexacuminate(acumen10– 25(–35)mmlong),primaryveinimpressedabove,secondary veinsdistinct,14–22oneithersideofprimaryvein,impressed above,smallestdistancebetweenloopsandmargin3–5 mm,tertiaryveinsslightlyraisedabove,reticulate.Flowers in 1–3-floweredinflorescencesinaxilsofleavesoronleafless branchlets;floweringandfruitingpedicels15–25mmlong,1–2 mm diam, densely to sparsely covered with appressed, brown hairs,articulatedatc.0.2fromthebase,bracts5–7,soonfalling, uppermost bract elliptic, 5–6 mm long, lower bracts broadly elliptic,1–3mmlong;flowerbudsdepressedovoid;sepalsfree, broadlyovate-triangular,5–9by4–7mm,stronglyreflexed, outersidedenselycoveredwithappressed,brownhairs;petals green, maturing cream or reddish in vivo, obovate to broadly Brazil Colombia Peru Venezuela Panama Costa Rica M e x i c o Guatemala Nicaragua Honduras Bolivia Guyana Suriname French Guiana P uer t o R i co Haiti Lesser Antilles Ecuador Map 15 Distribution of Guatteria ferruginea (●),G. flabellata (£),G. flagelliflora (■),G. flexilis (p)andG. foliosa (u).
84 Blumea–Volume60/1–3,2015 Plate 4a.Guatteria ferrugineaA.St.-Hil.Stemwithflowersandfruit.–b,d,e.Guatteria goudotianaTriana&Planch.b.Fruit;d.floweringbranch;e.flower. –c,f.Guatteria heteropetalaBenth.c.Fruitingbranch;f.floweringbranch.–g.Guatteria hirsutaRuiz&Pav.Fruitingbranch.–h,i.Guatteria inundataMart. h.Floweringbranch;i.fruitingbranch(a:Amorim et al. 818;b,d,e:Sánchez S. et al. 4776;c:Stevenson et al. 1115;f:Berry s.n.;g:Clark et al. 5291;h:Croat 19234;i:Maas et al. 6288).—Photos:a,i:P.J.M.Maas;b,d,e:D.SánchezS.;c:D.W.Stevenson;f:P.E.Berry;g:J.L.Clark;h:T.B.Croat. cb d a g h i e f
85 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria obovate,25–40{–50}by15–25{–40}mm,basedistinctlynarrowed(unguiculate)andmarginsrolledoutwards,apexoften emarginate, outer base densely covered with appressed hairs, middle and apical part sparsely so to glabrous, outer side with distinct,impressedveins;stamens1–2mmlong,connective shieldpapillatetohairy.Monocarps50–75,green,maturing redtoblackinvivo,blackinsicco,ellipsoid,9–15by5–8mm, sparselycoveredwithappressedhairs,apexapiculate(apiculum<0.5mmlong),wall0.2–0.4{–1}mmthick,stipes8–16by 0.5–1{–1.5}mm.Seedellipsoid,8–14by5–7mm,paletodark brown,pittedtotransverselygrooved,rapheraised. Distribution—AmazonianPeru(Loreto),Brazil(Amazonas, Rondônia). Habitat&Ecology—Inperiodicallyinundatedforest(igapó, restingabaja,tahuampa,várzea),rarelyinnon-inundatedforest.Atelevationsof0–300m.Flowering:ApriltoDecember; fruiting:MarchtoNovember. Vernacularnames—Peru:Anona(Rimachi Y. 6713),Anonilla(Vásquez & Jaramillo 7998),Anonilla-carahuasca(Vás quez & Jaramillo 9690),Baracaspi(Rimachi Y. 1864, 6454),Carahuasca(Rimachi Y. 3437, J. Ruíz & Jaramillo 1134, Vás quez & Jaramillo 9434),Carahuascablanca(J. Ruíz 1078),Carahuascadebajial,Carahuascadehojaancha. Note—Guatteria flabellata is recognizable by petals which are very well marked by a distinct impressed venation on theirouterside;furthermorethepetalsaredistinctlynarrowed towardsthebase(almostclawed),whereastheapexisoften somewhatemarginate. 66. Guatteria flagellifloraMaas&Westra—Map15 Guatteria flagellifloraMaas&WestrainErkensetal.(2008)486,f.8.—Type: Vásquez et al. 22500(holoU2sheets;isoF,MO,WAG2sheets),Peru, Amazonas,Prov.Condorcanqui,Distr.ElCenepa,Mamayaque,RíoCenepa,CerroSakeegaig,900–1000m,12Feb.1997. Treec.25mtall,diamnotrecorded;youngtwigssparsely coveredwithappressedhairs,soonglabrous,black.Leaves: petiolec.10mmlong,2mmdiam;laminanarrowlyellipticto narrowlyobovate,23–32by9–10cm(leafindex2.6–3.6), chartaceous, not verruculose, blackish brown above, brown below, glabrous above, sparsely covered with appressed hairs below,baseacute,apexobtusetoacuminate(acumenc.15mm long),primaryveinflatabove,secondaryveinsdistinct,12–16 on either side of primary vein, slightly raised above, smallest distancebetweenloopsandmargin2–4mm,tertiaryveins raisedabove,reticulate.Flowers in condensed panicle-like inflorescences(lengthnotmentioned)onleaflesssproutsfrom thestem;pedicels15–17mmlong,c.1.5mmdiam,youngfruitingpedicels15–30mmlong,2–3mmdiam,denselycovered withappressedhairs,articulatedatc.0.2fromthebase,bracts 5–6,soonfalling,notseen;flowerbudsdepessedovoid;sepals free,broadlyovate,5–7by4–6mm,appressed,outerside denselycoveredwithappressedhairs;petalsgreeninvivo, outer ones elliptic to elliptic-ovate, 15–20 by 10–15 mm, inner ones broadly to very broadly elliptic-ovate, 10–15 by 10–12 mm,outersidedenselycoveredwithappressedhairs;stamens 1.5–2mmlong,connectiveshieldpapillatetohairy.Monocarps 25–50,blackinvivo,blackinsicco,ellipsoid,8–10by4–5mm, wrinkledinsicco,glabrous,apexapiculate(apiculum<1mm long),wall0.1–0.2mmthick,stipes5–8byc.1mm.Seed ellipsoid,9–10by4–6mm,darkbrown,rugose,raphenotdistinct fromrestofseed. Distribution—AmazonianPeru(Amazonas). Habitat&Ecology—Inprimaryforest.Atelevationsof 900–1000m.Flowering:February;fruiting:February. Vernacularnames—Notrecorded. Note—Guatteria flagelliflora is unique by its flagelliform inflorescence, a feature not reported so far in Guatteria. 67. Guatteria flexilisR.E.Fr.—Fig.39;Map15 Guatteria flexilisR.E.Fr.(1952b)255.—Type:Forest Department British Guiana 5603=Fanshawe 2804(holoK2sheets;isoFDG,K,NY,U), Guyana,EsequiboRiver,WiniperuCreek,16Mar.1948. Treeorrarelyliana6–9mtall,2.5–12cmdiam;youngtwigs sparselycoveredwithappressedhairs,soonglabrous.Leaves: petiole4–6mmlong,c.1mmdiam;laminanarrowlyelliptic, 8–17by3–6cm(leafindex2.2–3.2),chartaceous,scabridulous, dull, grey to greyish brown above, greyish brown below, youngest leaves densely covered with appressed hairs, soon glabrous above, glabrous below, except for some scattered appressed hairs along primary vein, base attenuate, rarely obtuse,apexacuminate(acumen5–15mmlong),primaryvein impressedabove,secondaryveinsdistinct,9–14oneitherside of primary vein, impressed to slightly raised above, smallest distancebetweenloopsandmargin3–5mm,tertiaryveinsflatto slightlyraisedabove,reticulate.Flowers solitary in axils of leaves oronleaflessbranchlets;pedicels25–35mmlong,c.1mm diam,fruitingpedicelstoc.50mmlong,c.2mmdiam,sparsely coveredwithappressedhairstoglabrous,articulatedat0.1–0.2 fromthebase,bracts5–7,soonfalling,elliptic,basalones c.1mmlong,theupperonestoc.3mmlong,rarelyfoliaceous andpersisting,toc.20mmlong;flowerbudsdepressedovoid; sepalsfree,broadlytoshallowlyovate-triangular,3–4by3–5 mm, appressed, outer side rather densely covered with appressed hairs when young, later becoming almost glabrous, exceptforthehairymargin;petalspalegreyishgreenoryellow in vivo, oblong to elliptic to narrowly so, 10–26 by 5–10 mm, outersidedenselycoveredwithappressedhairs;stamens 1.5–2mmlong,connectiveshieldhairy.Monocarps 10–50, green,maturingred,tofinallyblacktopurple-blackinvivo,black tobrowninsicco,ellipsoid,10–14by5–6mm,glabrous,but apex sparsely covered with appressed hairs, apex rounded to apiculate(apiculum<0.5mmlong),wall0.1–0.2mmthick, stipes5–10byc.1mm.Seedellipsoid,9–11byc.5mm,shiny, reddish brown to dark brown, smooth to transversely grooved, raphenotdistinctfromrestofseed. Distribution—AmazonianVenezuela(Bolívar,DeltaAmacuro),Guyana. Habitat&Ecology—Innon-inundated,primaryorsecondaryforest,thetypecollectionfromGreenheart(Chlorocardium rodiaei)forest,onsandyorlateriticsoil.Atelevationsof0– 120m.Flowering:MarchtoMay,August,September;fruiting: February,March,May,September. Vernacularnames—Guyana:Blackyariyari(Creolename) (Van Andel et al. 1813),Karishiri(Arawakname)(Forest Department British Guiana 6336=Fanshawe 3006).Venezuela:Majagua(Ang. Fernández 3949),Yarayaramorada(Marcano-Berti 755). Notes—Guatteria flexilis is well recognizable by its scabridulous, almost completely glabrous leaves combined with relativelylongpedicelsofuptoc.50mmlonginfruit. Fries(1939)placedG. flexilisinsect.Dolichocarpusand‘most nearly connected with Guatteria subsessilis’. 68. Guatteria foliosaBenth.—Plate3f;Map15 Guatteria foliosaBenth.(1843)360;MurilloA.&Restrepo(2000)94;S.A.Mori etal.(2002)62,pl.12e;Maasetal.(2007)639.—Type:R.H. Schomburgk I 995(holoK3sheets;isoB3sheets,BM,E,F2sheets,G3sheets,K, L,NY,P,U,US),Brazil,Amazonas,RioNegro,anno1837. Tree4–25mtall,5–50cmdiam;youngtwigsratherdenselyto sparselycoveredwithappressedhairs,soonglabrous.Leaves:
86 Blumea–Volume60/1–3,2015 Fig. 39 Guatteria flexilisR.E.Fr.Floweringbranch(Van Andel et al. 1813,U).
87 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria petiole5–15mmlong,0.5–1mmdiam;laminanarrowlyelliptic tonarrowlyovate,orelliptic,6–16by2–6cm(leafindex1.9–4) chartaceous, not verruculose, shiny above, dark brown above, pale brown below, glabrous or sometimes sparsely covered with appressed hairs above, but often with appressed to erect, brown hairs along the primary vein above, rather densely to sparsely covered with appressed, brown hairs below, base obtuse, rarely acute, the extreme base long-attenuate, apex acuminate (acumen5–20mmlong),primaryveinflattoimpressedabove, secondaryveinsdistinct,6–13oneithersideofprimaryvein, often strongly raised above, smallest distance between loops andmargin2–3mm,tertiaryveinsraisedabove,reticulate. Flowersin1(–3)-floweredinflorescencesinaxilsofleaves; pedicels10–30mmlong,1–2mmdiam,fruitingpedicelsto c.3mmdiam,sparselycoveredwithappressedhairs,articulatedat0.2–0.3fromthebase,bracts5–6,notseen;flowerbuds depressedovoid;sepalsfree,broadlyovate-triangular,3–5by 3–5mm,reflexed,outersidedenselycoveredwithappressed, palebrownhairs;petalsgreen,maturingyellowishgreenor brown in vivo, narrowly oblong to narrowly obovate, 10–25 by4–10mm,outersidedenselycoveredwithappressed, brownhairs;stamens1–1.5mmlong,connectiveshieldhairy topapillate.Monocarps25–40,green,maturingblackinvivo, blacktobrownishblackinsicco,ellipsoid,5–10by3–4mm, sparsely covered with appressed hairs, soon glabrous, apex apiculate(apiculum<0.5mmlong),wall0.1–0.2mmthick, stipes10–25by0.5–1mm.Seedellipsoid,7–8by3–4mm, dark,shinybrown,smooth,raphenotdistinctfromrestofseed. Distribution—AmazonianColombia(Caquetá,Casanare, Vichada),Venezuela(Bolívar,Tachira),FrenchGuiana,Peru (Loreto),Brazil(Amazonas,MatoGrosso,Pará,Rondônia),Bolivia(Beni,SantaCruz). Habitat&Ecology—Innon-inundatedforest,sometimesin campinaraforest,orinperiodicallyinundatedforest;onsandy tosandy-clayeysoil.Atelevationsof0–500(–1300)m.Flowering:mainlyinSeptemberandOctober;fruiting:February,April, AugusttoOctober. Vernacularnames—Brazil:Envireira-da-casca-verde(D. Coêlho INPA 3872).Colombia:Cïbodujecu(Muinanename), Cargueronegro.Peru:Anonilla(J. Rodriguez 224),Carahuasca negra(J. Ruiz 1073).Venezuela:Aceitunonegro(J.O. Ramirez & Paredes 46),Karaurinyek(Pemónlanguage)(Ang. Fernández 4487),Majagua(Breteler 4752),Majaguaverde(H.L. Clark 7244, Liesner 6067). Notes—Guatteria foliosa is one of the species of this genus most easy to recognize by its long and slender petioles, a mostly obtuse leaf base that is abruptly and long-attenuate at the extreme base, and secondary veins which are strongly raisedontheuppersideofthelamina. For differences with G. liesneri and G. maypurensis see under thosespecies. Castillo 4328(MO)fromAmazonianVenezuelaisaberrantin havingpedicelsuptoc.45mmlong. Cid et al. 8344(U)fromAmazonianBrazildeviatesinhaving someoftheyoungesttwigscoveredwitherect(insteadofappressed)tocurlyhairs. Farney et al. 1838(U)fromAmazonianBrazilissomewhat aberrant in having sparsely verruculose leaves, a feature not met with in any other specimen of G. foliosa. Onecollection,Glaziou 13504 (P:‘Brazil,MinasGerais,Mar d’Espanha,8May1870’)fallsquitefaroutofthedistributionof thisspecies.AsimilarcaseisthatofGlaziou 13505(seeunder G. inundata).AsGlaziouquiteoftenusederroneousdata(see Wurdack1970),wehavethesuspicionthatthelocalitydata areincorrect. 69. Guatteria fractiflexaMaas&Westra—Fig.36b,40;Map 16 Guatteria fractiflexaMaas&WestrainErkensetal.(2008)491,f.9.—Type: Christenhusz & Cárdenas-Ramires 2110(holoU;isoLZ,TUR,USM),Peru, SanMartín,CerroGuayrapurina,topoftheridge,steepsouthfacingslope, 1300m,14Aug.2002. Liana,shrub,orsmalltreetoc.4mtall,diamnotrecorded; young twigs mostly with a zigzag pattern, sparsely covered withappressedhairs,soonglabrous.Leaves:petiole3–5mm long,1–2mmdiam;laminanarrowlyelliptic,6–15by1.5–5 cm(leafindex3–6),coriaceous,notverruculose,dull,greyish green above, pale brown below, glabrous above, except for some hairs along primary vein, sparsely to rather densely covered with appressed hairs below, base acute to obtuse, slightlyoblique,apexlong-acutetoacuminate(acumentoc. 10mmlong),primaryveinimpressedabove,secondaryveins indistinct,13–20oneithersideofprimaryvein,flatabove, smallest distance between loops and margin 1–2 mm, tertiary veinsflatabove,reticulate.Flowerssolitaryinaxilsofleaves; pedicels10–25mmlong,1–1.5mmdiam,denselycoveredwith erecttoappressed,brownhairs,articulatedat0.2–0.6fromthe base,bracts5–6,soonfalling,notseen;flowerbudsbroadly ovoid;sepalsfreetobasallyconnate,broadlyovate-triangular, 5–6by5–5.5mm,appressed,outersidedenselycoveredwith erecttoappressed,brownhairs;petalspaleyellowtoyellowin vivo,narrowlyelliptictonarrowlyelliptic-obovate,13–15by5–6 mm, outer side densely covered with erect to appressed, brown hairs;stamensc.1.5mmlong,connectiveshieldpapillateto glabrous.Monocarpsblackinvivo(accordingtothelabelof Rodriguez et al. 2740),notseen.Seednotseen. Distribution—AmazonianEcuador(Zamora-Chinchipe),Peru (Amazonas,Huánuco,SanMartín). Habitat&Ecology—Inlow,primaryforest,withmanyepiphytes, and with a thick layer of moss, roots and leaves, on sandstonesoil.Atelevationsof320–2190m.Flowering:February,May,June,August,November;fruiting:unknown. Vernacularnames—Notrecorded. Notes—Guatteria fractiflexa is with G. scandens(andsometimes G. beckii, G. flexilis and G. synsepala)oneofthefive lianaspeciesinthegenus.Anoteworthyfeatureisthezigzag patternofthelateral(plagiotropic)branchlets,mostextreme inAmazonianPeru(AmazonasandSanMartín),butlesspronouncedincollectionsfromPeru,Huánuco(Schunke V. 10299) andEcuador,Zamora-Chinchipe(Jørgensen et al. 2304). Pirie et al. 130(Peru,Amazonas;sterileliana)shouldbementioned here: it resembles collections of G. fractiflexa, but it differs by branchlets, petioles and lower leaf sides all densely covered witherecthairstoc.1mmlong.Wehavedecidedtoleaveit out as the number of collections of G. fractiflexa is too low to haveagoodpictureofthevariabilityofthiscuriousspecies. 70. Guatteria friesiana(W.A.Rodrigues)Erkens&Maas— Map16 Guatteria friesiana(W.A.Rodrigues)Erkens&Maas(2008)404;Maas& Westra(2011)123,f.8a,9,pl.1c.—Guatteriopsis friesianaW.A.Rodrigues (1981)49,f.1;Maasetal.(2007)644.—Type:D.F. Coêlho INPA 3609 (holoINPA;isoS),Brazil,Amazonas,Manaus,IgarapédoPassarinho, 14Mar.1956. Tree3–10mtall,4–10cmdiam;youngtwigsandpetioles densely covered with a velutinous indument of erect, dark browntoblackishbrown,long-persistenthairstoc.0.5mm long.Leaves:petiole2–8mmlong,2–3mmdiam;laminanarrowlyelliptictonarrowlyobovate,12–28by4–9cm(leafindex 2.7–3.5),chartaceoustothinlycoriaceous,scabridulous,pale green to greyish green on both sides, glabrous above, sparsely
88 Blumea–Volume60/1–3,2015 Fig. 40 Guatteria fractiflexaMaas&Westra.Flowering branch (Vásquez 27596,WAG).
89 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria covered with appressed brown hairs below, but rather densely to sparsely covered on primary vein, base cordate, apex acuminate(acumen10–20mmlong),primaryveinflatoralmostso above,secondaryveinsindistinct,10–24oneitherside,flator slightly impressed above, indistinctly loop-forming over most of the length or the entire length, smallest distance between loops and margin 2–5 mm, tertiary veins inconspicuous and hardly visible,reticulate.Flowers in 1–2-flowered inflorescences in axilsofleaves;floweringandfruitingpedicels5–10mmlong, 2–3mmdiam,denselycoveredwithdarkbrown,appressed hairs,articulatedat0.3–0.5fromthebase,bractsc.5,soon falling,theupperbractbroadlytodepressedovate,c.5mm long;flowerbudsovoidtoconical,acute;sepalsfree,triangular toovate-triangular,10–16by6–7mm,appressed,soonbecoming spreading to reflexed, outer side densely covered with darkbrown,appressedhairs;petalscreaminvivo,unequal, narrowly ovate-triangular to narrowly oblong-ovate, outer ones 20–25by8–12mm,innerones15–19by6–8mm,outerside denselycoveredwithdarkbrown,appressedhairs;stamens c.2mmlong,connectiveshieldhairy.Monocarps7–10,green, maturingredinvivo,browninsicco,ellipsoid,19–23by6–7 mm, sparsely covered with appressed and erect hairs, apex apiculate(apiculumc.2mmlong),wall0.1–0.2mmthick, stipes1–3by2mm.Seedellipsoid,13–19by6–8mm,dark, shiny brown, slightly transversally grooved, slightly pointed at theapex,raphestronglyimpressed. Distribution—AmazonianBrazil(Amazonas). Habitat&Ecology—Innon-inundatedforest,onclayeyto sandysoil.Atelevationsof50–125m.Flowering:Februaryto May;fruiting:MarchtoJune. Vernacularname—Brazil:Envireira(Rodrigues & Loureiro 5908). Note—Guatteria friesiana can be recognized by a cordate leaf base, obscure secondary veins, a velutinous indument on youngtwigs,andshortlypedicellateflowers. 71. Guatteria galeottianaBaill.—Map16 Guatteria galeottianaBaill.(1868b)268;R.E.Fr.(1939)373,f.10b–d.— Type:Liebmann 16(holoC;isoP),Mexico,Oaxaca,Comaltepec,Aug. 1842. Tree2–10mtall,10–20cmdiam;youngtwigssparselycovered withappressedhairs,soonglabrous.Leaves:petiole4–8mm long,1–2mmdiam;laminanarrowlyelliptic,rarelyobovate, 10–15by3–5cm(leafindex3–3.3),chartaceous,notverruculose, shiny, often bright green when dry above and below, glabrous above, glabrous below, except for some hairs along theprimaryvein,baseattenuate,apexacuminate(acumen 10–20mmlong),primaryveinflatabove,secondaryveinsdistinct,8–15oneithersideofprimaryveinatalmostrightangles, slightly raised above, smallest distance between loops and margin2–4mm,tertiaryveinsraisedabove,reticulate.Flowers solitaryinaxilsofleaves;pedicels10–30mmlong,c.1mm diam,fruitingpedicels35–50mmlong,c.2mmdiam,sparsely coveredwithappressedhairstoglabrous,articulatedatc.0.2 fromthebase,bracts5–7,soonfalling;flowerbudsdepressed ovoid;sepalsfree,broadlyovate-triangular,4–5by4–5mm, reflexed, outer side rather densely covered with appressed hairs;petalsyellowishgreentoyellowinvivo,ovate,12–15 by6–7mm,outersideratherdenselycoveredwithappressed hairs;stamens1–2mmlong,connectiveshieldpapillate. Monocarps 10–20, green, maturing blackish purple in vivo, blackinsicco,narrowlyellipsoid,10–13by5–6mm,sparsely covered with appressed hairs, soon glabrous, apex apiculate (apiculum<0.5mmlong),wallc.0.2mmthick,stipes9–17by 1mm.Seedellipsoid,9–12by4–5mm,darkbrown,rugose, raphenotdistinctfromrestofseed. Distribution—Mexico(Oaxaca,Veracruz). Habitat&Ecology—Inforest(‘selvaaltaperennifolia’).At elevationsof0–1400m.Flowering:June,October,November; fruiting:February,April,November. Vernacularnames—Mexico:Peseʉ(Zoquelanguage), Zapotedemonte(Osorio et al. 17, Rincón G. et al. 700). Note—Guatteria galeottiana is an endemic from the VeracruzandOaxacaregionsinMexico.Itcanberecognizedbyits leaves that often dry bright green, its long pedicels, the narrowly ellipsoid and apiculate monocarps, and its almost perpendicular secondaryvenation.Allthesecharactersfitthetypeverywell. However,thecollectionsfromOaxaca(e.g.,H. Hernández G. 1695)differsomewhatbyhavingroundermonocarpsanda somewhat more ascending secondary venation, but otherwise fallwithinthedescriptionofthisspecies. 72. Guatteria gentryiMaas&Erkens—Map16 Guatteria gentryiMaas&ErkensinErkensetal.(2008)495,f.10,pl.3.— Type:Gentry 12444(holoMO2sheets;isoU),Ecuador,Sucumbíos,17 kmWofLumbaque(70kmWofLagoAgrio),1130m,4Nov.1974. Guatteria sp. 8Chatrouetal.(1997)110. Tree5–40mtall,10–40cmdiam;youngtwigssparselycoveredwithappressedhairs,verysoonglabrous.Leaves: petiole 6–12mmlong,2–3mmdiam;laminanarrowlyelliptic,14–25 by4–8cm(leafindex2.8–4),chartaceous,notverruculose, dull, greenish brown above, not or sometimes slightly verruculose below, brownish below, glabrous above, rather densely covered with appressed hairs in the youngest stages but very soon becoming almost completely glabrous below, base acute toobtuse,apexacuminate(acumen5–15mmlong,itsapex itselfobtuse),primaryveinslightlyraisedabove,secondary veinsdistinct,10–13oneithersideofprimaryvein,raised above,smallestdistancebetweenloopsandmargin2–3mm, tertiaryveinsflattoslightlyraisedabove,reticulate.Flowers in 1–2-flowered inflorescences in axils of leaves or on leafless branchlets;pedicels4–8mmlong,1.5–2mmdiam,fruiting pedicels5–10mmlong,3–4mmdiam,denselycoveredwith appressedhairs,articulatedat0.5–0.6fromthebase,bracts 5–7,soonfalling,thetwouppermostonesbroadlyovate,toc.8 mmlong;flowerbudsbroadlyovoid;sepalsfreeorbasallyconnate,broadlyovate,5–6byc.5mm,appressedtospreading, outersidedenselycoveredwithappressedhairs;petalsgreen, maturing dull yellow, yellow or cream in vivo, ovate, elliptic to oblong-ovate, 12–20 by 6–9 mm, outer side densely covered withappressedhairs;stamens1.5–2mmlong,connective shieldpapillatetohairy.Monocarps10–30,green,maturing black in vivo, black, rarely brown in sicco, ellipsoid, 15–20 by Brazil Peru Ecuador Colombia Venezuela Guyana Surina m Fr e Panama Costa Rica Lesser Antilles Nicaragua Guatemala Cuba Haiti Honduras Dominican Republic Puerto Rico Mexico Map 16 Distribution of Guatteria fractiflexa (●),G. friesiana (£),G. galeottiana (■),G. gentryi (p),G. goudotiana (u)andG. herrerana (✸).
96 Blumea–Volume60/1–3,2015 Fig. 43 Guatteria hirsutaRuiz&Pav.Floweringbranch(Nee 37977, U).
97 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria WedecidedtouniteherefourspeciesdescribedbyTrianaand Planchoninthesamepaper(1862).Thesespeciesdifferfrom eachotheronlyinminoraspects(amongothers,leafbase,leaf indument,shapeofpetals),butexaminationofmorespecimens thanmusthavebeenatthedisposalofTrianaandPlanchon makesobviousthattherearemanyintergradations. ItiswithsomereluctancethatweunitedG. latisepala with G. hirsuta.Guatteria latisepala is found mostly at higher elevations(c.800mandup),andmanyspecimenslookquitedistinct by a more dense indument and by markedly thickened pedicels abovethearticulationpoint.Therearetoomanytransitional forms,though,tojustifykeepingthetwospeciesapart. Guatteria alutaceaasdescribedatfirstseemswelldistinct by the greyish to greyish black colour of the upper side of the leaves, the mostly obtuse to acute leaf base, and often the presence of rather long appressed and forward-pointing hairs onthelowersideofthelamina.However,therearealsomany intergradations here that make it impossible to uphold G. alu taceaasaseparatespecies. Forms described as G. alutaceavar.angustifolia are noticeable because of very narrow and attenuate leaves with a leaf indexupto8andgenerallylongpedicels,butotherwiseare notclearlydistinct. Guatteria longipes, known only from the type collected in the ColombianstateofCundinamarca(‘Prov.deBogotá,Biota, AndesdeBogotá’),atanelevationof2000m,differsbypedicels 50–90(–120)mmlong! Someforms,allfromtheColombianstateofChocó,arenoteworthy because of their tendency towards longer-than-average pedicels: Cuatrecasas & Llano 24070(to80mmlong!),Gentry et al. 17844 and 30242. Apart from that, they match well with theoveralldescription. 81. Guatteria hispida(R.E.Fr.)Erkens&Maas—Map18 Guatteria hispida(R.E.Fr.)Erkens&Maas(2008)404;Maas&Westra(2011) 129,f.13.—Guatteriopsis hispidaR.E.Fr.(1934)111,t.7,8;Maasetal. (2007)645.—Type:Ducke 14 = RB 23903(holoS;isoK,RB,US),Brazil, Amazonas,Manaus,EstradadoAleixo,km12,9Dec.1932. Treeorshrub2–6mtall,c.5cmdiam;youngtwigsdensely coveredwithbrown,rough,erect,long-persistenthairs(‘hirsute’)toc.3mmlong.Leaves:petiole3–5mmlong,1–1.5mm diam;laminanarrowlyoblong-elliptic,10–27by3–6cm(leaf index3.2–4.5),chartaceous,denselyverruculose,dull,greyish green to brownish green above, brown below, glabrous above, rather densely to sparsely covered with brown, rough, erect hairs (‘hirsute’)below,baseacutetoobtuse,apexabruptlyandlongacuminate(acumen10–25mmlong),primaryveinimpressed above, secondary veins distinct, 10–15 on either side of primary vein, slightly raised above, smallest distance between loops and margin3–7mm,tertiaryveinsslightlyraisedabove,reticulate. Flowerssolitaryinaxilsofleaves;floweringandfruitingpedicels5–8mmlong,3–5mmdiam,denselytofinallysparsely coveredwitherect,brownhairs,articulatedatc.0.7fromthe base,bracts5–6,soonfalling,theuppermostbracttoc.7mm long;flowerbudssubglobose;sepalsfree,broadlyovatetriangular,8–10by8–10mm,appressed,outersidedensely coveredwithappressed,brown,longhairs;petalsyellowin vivo,ovate-triangular,15–20by8–14mm,outersidedensely coveredwithappressed,brown,hairstoc.3mmlong;stamens c.2mmlong,connectiveshieldhairy.Monocarps 10–50, greenish red to wine-red in vivo, brown in sicco, narrowly ellipsoidandfusiform,20–35by7–8mm,denselycoveredwith erect,rough,brownhairs(‘hirsute’),apexdistinctlyandbluntly apiculate(apiculum<2mmlong),wall0.5–0.7mmthick,stipes 5–10by1–1.5mm.Seednarrowlyellipsoid,17–20by6–7mm, apex pointed, brown, longitudinally and transversely grooved torugulose,raphenotdistinctfromrestofseed. Distribution—AmazonianBrazil(Amazonas). Habitat&Ecology—Mostlyincampinaranaforest,onsandysoil.Atelevationsbelow200m.Flowering:February,December;fruiting:MaytoJuly. Vernacularname—Brazil:Envireira. Note—Guatteria hispida is one of the very few species in the genus with a long-persistent indument of erect, rough, brown hairs(‘hirsute’)ontheleafytwigs,leavesandalsoonthemonocarps. 82. Guatteria insculptaR.E.Fr.—Map18 Guatteria insculptaR.E.Fr.(1939)504,f.28a,b;MurilloA.&Restrepo(2000) 95,f.25;Maas&Westra(2011)131,f.1c,14,15.—Type:Spruce 2896 (holoK2sheets;isoBM,BR,P),Brazil,Amazonas,Ipanoré(‘Panuré’), RioUaupés,Mar.1853. Treeorshrub5–25mtall,5–30cmdiam;youngtwigsdensely covered with a velutinous indument of erect, red-brown hairs, finallyglabrous.Leaves:petiole5–10mmlong,4–7mmdiam; lamina narrowly elliptic to narrowly obovate, 20–50 by 5–16 cm (leafindex2.3–4.4),chartaceous,verydenselyandminutely verruculose, dull, greyish green above, brown below, glabrous above, densely covered with erect, red-brown hairs below, base acute,extremebaseobtuse,apexacuminate(acumen5–15mm long),primaryveinimpressedabove,secondaryveinsdistinct, 20–35oneithersideofprimaryvein,impressedabove,sometimes forming a more or less distinct marginal vein, at a smallest distanceof3–4mmfromthemargin,tertiaryveinsimpressed above,percurrent.Flowers solitary in axils of leaves or on leaflessbranchlets;pedicels18–40mmlong,2–4mmdiam, fruitingpedicelstoc.50mmlong,c.5mmdiam,denselycoveredwitherect,red-brownhairs,articulatedat0.1–0.2from thebase,bractssoonfalling,theupperbracttoc.7mmlong; flowerbudsnotseen;sepalsfree,triangulartobroadlyovatetriangular,10–12by7–8mm,reflexed,outersidedenselycoveredwitherect,red-brownhairs;petalsgreeninvivo,obovate toelliptic,16–20by8–12mm,outersidesparselycovered withappressed,whitehairs,thebasedenselyso;stamens 1.5–2mmlong,connectiveshieldpapillate.Monocarps 10–50, green, maturing brown in vivo, pale brown in sicco, ellipsoid, 10–18by5–12mm,sparselycoveredwithappressedhairs, apexroundedtoapiculate(apiculum<0.5mmlong),wall0.5–1 mmthick,stipes5–10by1–2mm.Seed ellipsoid, 12–19 by 6–8mm,shinybrown,longitudinallyandtransverselygrooved, raphenotdistinctfromrestofseed. Brazil Colombia Peru Venezuela Panama Costa Rica M e x i c o Guatemala Nicaragua Honduras Bolivia Guyana Suriname French Guiana P u e r t o R i c o Haiti Lesser Antilles Ecuador Map 18 Distribution of Guatteria hirsuta (●),G. hispida (£ ),G. insculpta (■) and G. intermedia (p).
98 Blumea–Volume60/1–3,2015 Distribution—AmazonianColombia(Amazonas,Caquetá), Venezuela(Amazonas),Brazil(Amazonas). Habitat&Ecology—Innon-inundated,oftencaatingaforest, onsandysoil.Atelevationsofupto270m.Flowering:March, April,July,October;fruiting:January,February,April,October, November. Vernacularnames—Colombia:Jakuo(Muinanename)(Murillo A. & Rodríguez 544),Jimokai(Huitotoname),Palodeperfume. Note—Guatteria insculpta can be recognized by very densely and minutely verruculose leaves, and by young twigs coveredwithavelutinousindumentoferectred-brownhairs. Furthermore, this species stands out by the dense cover of erectbrownhairsonthelowersideoftheleaves. 83. Guatteria intermediaScharf—Map18 Guatteria intermediaScharfinScharfetal.(2006b)545,f.2.—Type:Oldeman B.4125(holoCAY;isoNY),FrenchGuiana,Saül,CriqueCochon, 21Oct.1971. Tree2–10mtall,5–15cmdiam;youngtwigsdenselycovered withsemi-erect,brown,long-persistinghairs.Leaves: petiole 3–10mmlong,1–2mmdiam;laminanarrowlyelliptictonarrowly obovate,10–23by4–7.5cm(leafindex2.6–3.4),chartaceous, not verruculose, dull greyish above, greyish brown to reddish brown below, sparsely covered with appressed, long hairs above, soon glabrous, primary vein with a row of semi-erect hairs, densely covered with appressed, pale brown hairs below, baseacute,apexacuminate(acumen10–20mmlong),primary vein impressed above, secondary veins distinct, 10–12 on either side of primary vein, flat to slightly raised above, smallestdistancebetweenloopsandmargin1.5–3mm,tertiary veinsslightlyraisedabove,percurrenttoreticulate.Flowers in1(–2)-floweredinflorescencesinaxilsofleaves;flowering and fruiting pedicels 5–20 mm long, 1–2 mm diam, densely coveredwithsemi-erect,brownhairs,articulatedat0.3–0.4 fromthebase,bracts5–6,soonfalling,notseen;flowerbuds depressedovoid;sepalsfreeorslightlyconnateatthebase, broadlytoshallowlyovate-triangular,4–5by5–7mm,apex reflexed,outersidedenselycoveredwithappressedhairs(apex glabrous);petalsgreenishyelloworyellowinvivo,narrowly oblong-elliptic to oblong-elliptic,10–20 by 5–10 mm, outer side densely covered with appressed to erect, rusty brown, curly hairs;stamens1.5–2mmlong,connectiveshieldpapillate, rarelyhairy,mostlyumbonate.Monocarps 20–50, green, maturing black in vivo, shiny black or blackish brown in sicco, ellipsoid,5–9by3–5mm,sparselycoveredwithappressedhairs, soonglabrous,apexapiculate(apiculum<1mmlong),wall 0.1–0.2mmthick,stipes10–25by0.5–1mm.Seed ellipsoid, 6–8by4–5mm,shiny,darkbrown,pitted,raphenotdistinct fromrestofseed. Distribution—NWSuriname,FrenchGuiana,Brazil(Amapá). Habitat&Ecology—Innon-inundatedforest,onlateriticor sometimesongraniticsoil.Atelevationsof0–400m.Flowering:JunetoMarch;fruiting:February,OctobertoDecember. Vernacularnames—FrenchGuiana:Iwi,I-wi(Wayãpiname) (de Granville T.1151, Grenand 632, Prévost & Grenand 967, Prévost & Sabatier 2781),Mamayavé(Creolename)(de Granville B.4561). Notes—TherecentlydescribedG. intermedia differs from G. ouregou and G. richardii, by having semi-erect to appressed insteadoferecthairsonyoungtwigsandpedicels. Exceptfortheindument,G. intermedia shares many features with G. punctata. Tocomplicatematters,though, there are somespecimenscitedbyScharf(Scharfetal.2006b)under G. intermedia which lack the indument of semi-erect hairs, havinganindumentofappressedhairsinstead.ThisinvolvesMori et al. 25705fromFrenchGuiana,and Van Andel et al. 4657 fromSuriname.InalmostallmaterialofG. punctata studied by us we observed a very distinct indument of appressed hairs on mostoftheparts.IntheBrazilianstateofMatoGrosso,however, we came across some material with erect or almost erect hairs.ThesespecimenshadbeenidentifiedasG. sylvicola.In that case, too, we found various specimens with an indument intermediate between that of G. sylvicolaandthatof‘typical’ G. punctata making us conclude that all material should be referred to G. punctata.PossiblyG. intermedia represents a similar case of a slightly deviating population within G. punctata inthewidesense. 84. Guatteria inundataMart.—Plate4h,i;Map19 Guatteria inundataMart.(1841)36;R.E.Fr.(1939)411,f.14f,g,t.28.—Type: Martius s.n.(lectoM,selectedbyFries1939),Brazil,Amazonas,RioSolimõesandRioJapurá(‘ininundatisripaefluv.SolimõesetJapurae’),14 Dec.1819. Guatteria inundataMart.var.longifoliaPoepp.exMart.(1841)36.—Guatteria dolichophyllaR.E.Fr.(1939)414,syn.nov.—Type:Poeppig 2693 (holoBR;isoB,BM,F,G,HAL,P,W),Brazil,Amazonas,alongalakenear Tefé(‘insylvislitoreislacusEgensis’),Oct.1831. Guatteria phanerocamptaDiels(1931)76;R.E.Fr.(1939)415.—Type: Ll. Williams 3436(synF,G,S,US),Peru,Loreto,UpperRíoItaya,San Antonio,145m,5Oct.1939. Guatteria ripariaR.E.Fr.(1939)410;Steyerm.etal.(1995)446,f.377,syn. nov.—Type:Spruce 3105(holoK;isoBM,BR,C,G3sheets,NY,P), Venezuela,Amazonas,banksofRíoNegroandRíoCasiquiare,Oct.1853. Guatteria obovataR.E.Fr.(1939)412,f.14h;Morawetz(1984)59,syn.nov. —Type:R.H. Schomburgk I 922(holoK;isoB,BM,E,F,FI2sheets,G3 sheets,L,P,U,US,W),Brazil,Amazonas,RioNegro,LagoPedrero,1840. Guatteria oblanceolataR.E.Fr.(1939)415.—Type:Krukoff 6721(holoS; isoBM,BR,F,G,K,MO,NY,RB,U,US2sheets),Brazil,Amazonas,Mun. Humaitá,nearLivramento,onRioLivramento,12Oct.to6Nov.1934. Many-stemmedshruborsometimesatree1–10(–20)mtall,up toc.10cmdiam;youngtwigssparselycoveredwithappressed hairs,soonglabrous.Leaves: petiole 5–15 mm long, 1–2 mm diam;laminanarrowlyelliptic,15–27by4–8cm(leafindex 2.3–5.6),chartaceous,sometimesrugulose,brownishgreen, brown, to greyish green above, brown to greenish brown below, glabrous above, sparsely covered with appressed hairs below, baseattenuate,apexacuminate(acumen10–20mmlong)to long-acute, primary vein impressed above, secondary veins distinct, 12–20 on either side of primary vein, impressed to slightly raised above, smallest distance between loops and margin 2–6 mm,tertiaryveinsflattoslightlyraisedabove,reticulate.Flowerssolitaryinleafaxils,rarelyonleaflessbranchlets;pedicels 12–40mmlong,1–1.5mmdiam,fruitingpedicelstoc.50mm long,c.2mmdiam,sparselycoveredwithappressedhairs, articulatedat0.2–0.6fromthebase,bracts4–6,soonfalling, broadlytriangular-elliptic,broadlyovate,orelliptic,1.5–6mm long;flowerbudsdepressedovoid;sepalsfree,broadlyovatetriangular,4–6by4–6mm,appressed,outersidedenselyto ratherdenselycoveredwithappressedhairs;petalsgreen, maturingcreamoryellowinvivo,ovatetoobovate,15–35 by8–20mm,outerbasedenselycoveredwithappressed hairs,apicalpartratherdenselytosparselyso;stamensc.2 mmlong,connectiveshieldpapillatetoglabrous.Monocarps 25–50, green, maturing reddish in vivo, black, rarely brownish insicco,narrowlyellipsoidandmostlyfusiform,17–30by5–7 mm, sparsely covered with appressed hairs, soon glabrous, apexdistinctlyapiculateandhard-pointed(apiculum1–3mm long),wall0.1–0.3mmthick,stipes2–5by1–2mm.Seed narrowlyellipsoid,15–25by5–8mm,palebrown,pitted, longitudinally grooved, with spongy appearance, raphe not distinctfromrestofseed.
99 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria Distribution—AmazonianColombia(Vaupés),Venezuela (Amazonas),Peru(Loreto),Brazil(Amazonas,Pará,Roraima). Habitat&Ecology—Mostlyinperiodicallyinundatedforest (igapó,várzeaortahuampa),onclayeytosandysoil.Atelevationsof0–200m.Flowering:throughouttheyear,butmostly betweenSeptemberandNovember;fruiting:NovembertoJune. Vernacularnames—Brazil:Cacaia-velha(Zappi et al. 2967),Envira-do-igapó(Albuquerque & Lima 227 ).Peru: Anona(McDaniel & Rimachi Y.17539, Rimachi Y. 2443, 2557), Anonilla(Rimachi Y. 12305),Bara(Rimachi Y. 907, 2706, 6447), Bar(r)acaspi(McDaniel & Rimachi Y.20386, Rimachi Y. 1840), Carahuasca(Del Carpio & Ruiz 1606),Espintana(Peters 22, Rimachi Y. 1809, 10719),Motelocaspi(Rimachi Y. 3348).Venezuela:Kunwata(L. Delgado et al. 953),Majagua(Breteler 4844, Stergios & Aymard C. 9010),Majaguanegra(Liesner 6233), Majaguaorillera(Liesner 6501),Majaguarebalsera(Delgado 593),Palodeboyanegro(Ll. Williams 14727). Notes—Guatteria inundata is a species quite often collected inperiodicallyinundatedvárzea,tahuampaandigapóforests ofAmazonianBrazil,PeruandVenezuela.Itismosteasyto recognize by the long-apiculate and almost pointed monocarps, veryrarelymetwithinthegenus.Theseedsofthisspecieshave a spongy outer layer which may be an adaptaton to hydrochory (thespeciesisrestrictedtoriverbanks!).Fries(1939)placed G. inundata, G. dolichophylla, G. oblanceolata, G. obovata, G. phanerocampta and G. ripariainsect.Stenocarpus.Inhiskey to the 6 species he used characters such as leaf shape, secondaryveinsandleafcolour.Intheinitialphaseofourstudyweonly recognizedameretwospecies,viz.G. inundata and G. riparia, with as main differentiating characters the leaf shape, leaf apex anddistancebetweenloopsandthemargin.Withthesteady increase of many new collections these differences gradually became blurred, leaving us no other option but to unite all in G. inundata. Inmostcollectionsstudiedthemonocarpsareverynarrow and almost fusiform, but in a few collections the monocarps aremuchbroaderandtheapexevenobtuse. Onecollection,Glaziou 13505 (P:‘Brazil,RiodeJaneiro,Morro doCocco,9February1882’),fallsquitefaroutofthedistributionrangeofthistypicallyAmazonianspecies.Asimilarcaseis that of Glaziou 13504(seeunderG. foliosa).AsGlaziouquite oftenusederroneousdata(seeWurdack1970)wehavethe suspicionthatthelocalitydataareincorrect. 85. Guatteria japurensisMaas&Westra—Map20 Guatteria japurensis Maas&WestrainErkensetal.(2008)497,f.12;Maas &Westra(2011)131.—Type:Amaral et al. 518(holoU;isoINPA,NY, RB),Brazil,Amazonas,rightbankofRioJapurá,VillaBittencourt,Serrinha, 100m,16Nov.1982. Smalltreec.6.5mtall,diamnotrecorded;twigs(nogrowthtip seen)glabrous.Leaves:petiolec.10mmlong,3–4mmdiam; laminanarrowlyelliptic-ovate,16–27by5–8.5cm(index3–4), chartaceous to thinly coriaceous, scabridulous above, not or sparsely verruculose along primary vein only, slightly shiny and brownish green above, dull brownish green below, glabrous on both sides, base obtuse, extreme base shortly attenuate, apex acuminate(acumenc.10mmlong),primaryveinflatabove, secondaryveinsindistinct,15–18oneithersideofprimaryvein, raised above, smallest distance between loops and margin 1–2mm,tertiaryveinsflattoslightlyraisedabove,reticulate. Flowers in 1–several-flowered inflorescences in axils of leaves or on leafless branchlets, only seen in fruiting stage: fruiting pedicels50–60mmlong,1–1.5mmdiam,sparselycovered withappressedhairstoglabrous,articulatedatc.0.1fromthe base,bracts5–6,soonfalling,notseen;flowerbuds,sepals, petalsandstamensnotseen.Monocarps 10–15, immature, colour not recorded in vivo, blackish brown in sicco, ellipsoid to narrowlyellipsoid,17–20by7–8mm,glabrous,apexapiculate (apiculum0.2–0.4mmlong),wallc.0.2mmthick,stipes11–17 by1–1.5mm.Seednarrowlyellipsoid,c.18by7mm,brown, shiny, longitudinally and transversely grooved, raphe not distinct fromrestofseed. Distribution—WesternAmazonianBrazil(Amazonas). Habitat&Ecology—Innon-inundatedforest,onstonysoil coveredwithlichensandmosses.Atanelevationof100m. Flowering:unknown;fruiting:November. Vernacularnames—Notrecorded. Note—Guatteria japurensis is most distinctive by the long and slender fruiting pedicels bearing rather long monocarps, a combination very uncommon in Guatteria.Anothernoteworthy feature is found in the leaves which are indistinctly veined and arescabridulousontheuppersurface. 86. Guatteria jefensisBarringer—Plate5a,b;Map19 Guatteria jefensis Barringer(1984)1186.—Type:Hammel 6302 (holoMO), Panama,Panamá,CerroJefenearradiotower,inClusiaforest,1000m, 3Mar.1979. Treeorshrub1–6mtall,2–5cmdiam;youngtwigssparselycoveredwithappressedhairs,soonglabrous.Leaves:petiole3–8 mmlong,1–3mmdiam;laminanarrowlyovatetoelliptic,7–20 by3–7cm(leafindex2.3–3),coriaceous,oftenfoldedlengthwise when dry, not verruculose, dull, greyish green above, brown below, sparsely covered with erect hairs along primary vein and secondary veins above, sparsely covered with appressedhairsbelow,baseobtuse,apexacuminate(acumen toc.10mmlong),primaryveinimpressedabove,secondary veinsdistinct,10–13oneithersideofprimaryvein,impressed to flat above, smallest distance between loops and margin 3–4mm,tertiaryveinsflatabove,reticulate.Flowers in 1–2-flowered inflorescences in axils of leaves or on leafless branchlets;pedicels10–50mmlong,1–3mmdiam,fruiting pedicels30–50mmlong,3–4mmdiam,denselycoveredwith appressedhairs,articulatedatc.0.2fromthebase,bracts5–7, graduallyincreasinginsizefrombasaltoc.2mmlongtouppermosttoc.5mmlong,occasionallyfoliaceous,elliptic,20–30 mmlong;flowerbudsdepressedovoid;sepalsfree,broadly ovate-triangular,5–8by5–7mm,appressed,finallybecoming spreading to slightly reflexed, margins revolute, outer side denselycoveredwithappressedhairs;petalsyellowishgreen Brazil Colombia Peru Venezuela Panama Costa Rica M e x i c o Guatemala Nicaragua Honduras Bolivia Guyana Suriname French Guiana P ue r to R i co Haiti C u b a Lesser Antilles Ecuador Map 19 Distribution of Guatteria inundata (●),G. jefensis (£ ),G. latifolia (■), G. leucotricha (p)andG. pachycarpa (u).
100 Blumea–Volume60/1–3,2015 Plate 5a,b.Guatteria jefensisBarringer.a.Floweringbranchofdwarftree;b.ripeflower(top),fruit(bottom).–c.Guatteria latifolia(Mart.)R.E.Fr.Flower bud.–d–f.Guatteria lucensStandl.d.Flowers;e.leaves;f.fruit.–g,h.Guatteria megalophyllaDiels.–g.Flower;h.fruit.–i.Guatteria oblongifoliaRusby. Floweringbranch(a:Maas et al. 9553;b:CerroJefe,Panama;c:Maas & Martinelli 3222;d;Maas et al. 9491;e:Maas et al. 9519;f:Chatrou et al. 705;g,h: Chatrou et al. 387;i:Homeier 4504).—Photos:a,c–e:P.J.M.Maas;b:J.Harrison;f–h:L.W.Chatrou;i:J.Homeier. cb d a g h i e f
101 P.J.M.Maasetal.:RevisionoftheNeotropicalgenusGuatteria oryellowinvivo,broadlyovatetoobovate,10–15{–20}by7–15 mm,outersidedenselycoveredwithappressedhairs;stamens 1–2 mm long, connective shield glabrous, sometimes slightly umbonate.Monocarps 25–50, green, maturing blackish purple invivo,blackinsicco,ellipsoid,10–13by5–7mm,glabrous, apexroundedtoapiculate(apiculumnipple-shaped,toc.1mm long),wall0.2–0.3mmthick,stipes4–8by1–2mm,distinctly constrictedattheapex.Seedellipsoid,8–11by5–6mm,dark brown,pitted,raphenotdistinctfromrestofseed. Distribution—Panama. Habitat&Ecology—Indwarfcloudforest.Atelevationsof 800–1000m.Flowering:JanuarytoJune,September,November;fruiting:JanuarytoApril,October,December. Vernacularnames—Notrecorded. Note—Guatteria jefensis, one of the many narrow endemics ofCerroJefe,iseasilydistinguishedbyitsthick,oftenlengthwisefoldedleavesandbyitsconstrictedstipes. 87. Guatteria latifolia (Mart.)R.E.Fr.—Fig.44;Plate5c;Map 19 Guatteria latifolia (Mart.)R.E.Fr.(1939)326,f.4a.—Guatteria nigrescens Mart.var. latifolia Mart.(1841)31. — Type: Schott s.n. (holoBR),Brazil, RiodeJaneiro,SerradoTinguá. Treeorshrub4–10mtall,6–20cmdiam;youngtwigsglabrous. Leaves:petiole5–8mmlong,c.2mmdiam;laminanarrowly elliptictoelliptic,10–20by3–7cm(leafindex2–4),chartaceous, not verruculose, shiny, blackish green to brown above, pale brown below, glabrous above, sparsely covered with appressed, brown hairs to glabrous below, base acute, apex acute toacuminate(acumenc.10mmlong),primaryveinimpressed above, secondary veins distinct, 10–12 on either side of primary vein, raised to impressed above, smallest distance between Brazil Colombia Peru Venezuela Panama s ta Rica a la Nicaragua H o n d u r a s Guyana Suriname French G u H a i t i Lesser Antilles Ecuador Map 20 Distribution of Guatteria japurensis (●),G. megalocarpa (£ ),G. meliodora (■),G. microcarpa (p)andG. minutiflora (u). Fig. 44 Guatteria latifolia (Mart.)R.E.Fr.a.Fruitingbranch;b.flowerbuds; c,d.flowers(a,b:Maas & Martinelli 3222;c,d: Marquete 3416;fromLobão (2009)f.7A–D). cb d a 2 cm 5 cm 2 cm
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