Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Taxonomists. All rights reserved. © Copyright2010bythe American SocietyofPlantTaxonomists Systematic Botany were lifted. Thus,botanicalinventories intheregion have location ofthe border andrestrictions inaccesstotheregion tries in1995. A treaty signedin1998establishedtheprecise der, culminating in anarmedconflictbetweenthetwocoun- Ecuador andPeru overthelocationof internationalbor- ing the1990swaslimiteddue toanongoingdisputebetween in Ecuadorwasinitiated1990, butfurtherfieldworkdur- and theothersub-Andeancordilleras. gold, copper, andothermineralsalsoformpart oftheCóndor years. Intrusive igneousformations,oftenbearingdepositsof concurrently withthemain Andes, largely inthepastmillion of South America priortotheriseof Andes anduplifted sandstone andlimestone,depositedatthewesternmargin up principallyofMesozoicandearlyTertiary sediments, phic andvolcanicrocks, thesub-Andeancordilleras are made the mainEasternCordillera iscomprisedmostlyofmetamor- the Cordillera Azul and Yanachaga rangesinPeru. Whereas ras includetheGalerasandCutucúrangesinEcuador valleys. BesidestheCóndor, someothersub-Andean cordille- separated from themainEastern Cordillera bylowerelevation to themain Andean chainbutattaininglowerelevationsand Cordillera ofthe Andes andthe Amazon lowlands,parallel Andean cordilleras that are situatedbetweentheEastern The Cóndorformspartofthediscontinuouschainsub- south insoutheasternEcuadorandadjacentnorthernPeru. Andean mountainsthatextendsabout150kmfrom northto Botanical exploration oftheCordillera delCóndorregion The Cordillera delCóndorisaneastern outlierofthe Shuaria bosque nubladomontanobajoenlasvertientesoccidentalesdelaCordillera delCóndor. Sepresenta unaclaveytablapara provincias ecuatorianasdePastaza,Morona-Santiago, yZamora-Chinchipeenelbosquehúmedotropical delastierrasbajasamazó conjunto decaracteres inusualesinicialmentehizodifícilcolocar etativas yflorales;pequeñasflores blancasconunagibosidaddorsalenlabasedeltubodecorola; yunacápsulasepticida : elhábitoarborescente; hojasopuestasalgunasvecesintercaladas conhojasalternas;tricomaslepidotosenlase tribus BesleriaceaeyNapeantheae. trnL Beslerieae sebasaenlassecuenciasde ADN generadasporlosanálisisdelaregión ITSdel ADN ribosomalnuclear, ydelaregi con unasolaespecie, forest onthewesternslopesofCordillera delCóndor. A keyandatable are presented fordifferentiating Ecuadorian provinces ofPastaza,Morona-Santiago, andZamora-Chinchipeinlowland Amazonian tropical wetforest and inlowermo suite ofuncommoncharactersinitiallymade and floralstructures; smallwhiteflowerswithadorsalgibbosityatthebaseofcorolla tube;andabivalvedsepticidally acters intheGesneriaceae:arborescent habit;oppositeleafarrangementsometimesinterrupted byalternateleaves;lepidotetr diversity from thetribesBeslerieaeandNapeantheae. the chloroplast DNA strongly supportedbymolecularsequencedatageneratedfrom analysesofnuclearribosomalDNA internaltranscribedspacerregi regions ofsoutheasternEcuadorandisplacedinthefamilyGesneriaceae,tribeBeslerieae.Theplacement Shuaria Keywords— Resumen— Abstract— intron,

3 University ofVienna, DepartmentofPalynologyandStructural Botany, Rennweg14, A-1030 Vienna, Austria yotros géneros estrechamente vinculados. (2010), 35(3):pp.662–674 4

on . Clark, L. John Universidad SanFranciscodeQuito,Ecuador;FederaciónInterprovincial deCentros Shuar, trnL-trnF A newneotropical , 1 e ecie describe Se (Gesneriaceae),anArborescentNewGenusfromtheCordilleradelCóndor University of Alabama, DepartmentofBiologicalSciences,Tuscaloosa, Alabama 35487U.S. A. elree, Eudr, Gseica aenha hlgntc . phylogenetics, Napeantheae, Gesneriaceae, Ecuador , Beslerieae, trnL del ADN delcloroplasto. Sesecuenciaron 33especies,representando lamayorpartedediversidadgenéricalas S.ecuadorica intron, Shuaria 2

Missouri BotanicalGarden, P.O. Box299,St.LouisMissouri63166U.S. A. 1 Dvd . Neill, A. David trnL-trnF , unnuevogénero neotropical delaCordillera delCóndorylaregión amazónicadelsuroriente delEcuador, Shuaria , yseleincluyeenlatribuBeslerieaedefamliaGesneriaceae.Laubicación Shuaria intergenic spacerregion. Sequencedatawere generatedfor33speciesrepresenting mostofthegeneric secaracterizaporlasiguientecombinación únicadecaracteres quenosoncomunesenlafamilia Shuaria

1 , withasinglespecies, Author forcorrespondence ( [email protected] ) omnctn Eio: itra Sosa Victoria Editor: Communicating Sucúa, Morona-Santiago, Ecuador and AmazonianEcuador difficult toplaceintheGesneriaceaeandwithinaknowntribe. Shuaria 2 Atn Weber , Anton ischaracterizedbythefollowinguniquecombinationofrelatively uncommonchar- Shuaria S.ecuadorica 662 enGesneriaceaeysustribusconocidas. to Gesneriaceae by Wiehler (1994) followingmorphological been placedin Buddlejaceae,but G. S.Bunting&J. A. Duke and neously determinedas Peltantherafloribunda ern Ecuador: monotypic generathatoccurin thesamegeneralregion of east- examined thematerialtobesimilar inoverallappearancetotwo matic taxonseemedtothecollectorsandotherbotanistswho be assignedtoanextantgenusorfamilyofthatorder. Thisenig- recognized asamemberoftheLamiales,butcouldnotreadily ynous flowers,anddrycapsularfruit. Thistaxonwasinitially inflorescences withrathersmall,white,gamopetalous,hypog- opposite (occasionallyalternate),exstipulateleaves,branched region onseveraloccasionsisasmallunderstorytree with Melastomataceae ( Ulloa andNeill2006 ). 1998 ) and include Shield insouthernVenezuela andadjacent areas. Examples endemic tothesandstoneplateausortepuisofGuayana eral specieswithingenerathatwere previously considered flowering planttaxafrom theCóndorregion includesev- areas oftheCóndorregion ( Neill 2005 ). Therecently described restricted totheoligotrophic sandstoneplateausandadjacent the newlydescribedtaxaare locallyendemicandedaphically have revealed anumberof taxonomicnovelties,andmostof continued duringthepastdecade.Thesefloristicinventories 3 Jnie A Gruhn, A. Jennifer Among thecollectionsmadeinCordillera delCóndor , isdescribedfrom theCordillera delCóndorand Amazonian Stenopadus andicola Phainantha shuariorum Sanango racemosum Benth.Infact,somecollectionswere erro- Peltanthera 2 n uta Katan Tuntiak and Pruski inthe ( Pruski Shuaria Shuaria C.Ulloa&D. A. Neillinthe (Ruiz&Pav.) Barringerand Peltanthera Benth&Hook.Both from otherrelated genera. Shuaria Shuaria Sanango inthetribeBeslerieaeis dehiscent . The dehiscentcapsule.The ichomes on vegetative ichomes onvegetative Shuaria isrestricted tothe serestringe alas have traditionally traditionally have bivalvada.Este distinguir entre a transferred was structuras veg- ón intergénica on (ITS), and on (ITS),and 4 enlatribu

ntane cloud ntane cloud nicas, yen Sanango

Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. described inJohnson andSoltis(1995).Direct cyclesequencing ofpurified using PEG8000(polyethylene glycol)in2.5MNaCl under theconditions stained withethidium bromide toconfirmasingleproduct, andpurified were electrophoresed usinga1.0%agarose gelin1 to use5%DMSOandBSA inPCRreactions forITS.ThePCRproducts can result ininterference withTaq polymeraseactivity, we founditessential (Promega, Madison,Wisconsin). To reduce within-strandbasepairingthat dures describedby Baldwin etal.(1995) utilizingTaq DNA polymerase reads fortheentire ITSregion. used incyclesequencingtoobtainindependentandoverlappingsequence and forward oftheinternal primersITS2andITS3( White etal.1990 ) were fied usingtheprimerstrnLcandtrnLf( Taberlet etal.1991 ). Thereverse 1990 ). ThecpDNA prepared usingtheprimersITS5HP ( Suh etal.1993 ) andITS4( White etal. well asthe region of18S-26SnuclearribosomalDNA (nrDNA)andthe twenty-six specieswere sequencedfortheinternaltranscribedspacer(ITS) included intheanalyses(Appendix1).Twenty-seven taxa representing phylogenetic placement,ecology, anddistribution. Gesneriaceae, tribeBeslerieae,andpresent adiscussionofits species, placement. We describethistaxonherein asanewgenusand material forDNA extractionandsequencingtohelpresolve its 663 Recent collectionsofthissmallunderstorytree includedleaf Garden (MO)andtheNationalHerbariumofEcuador(QCNE). up inthe“mysteryplantcabinets”atMissouriBotanical ities farthernorthinlowland Amazonian Ecuadoralsoturned Peltanthera CLARKET AL.: SHUARIA (GESNERIACEAE), A NEWGENUSFROMECUADOR 1999 ; Bremer etal.2002 ; ( WangNorman 2000 etal.2004 ). Molecularsystematicstudies( ) strongly support Oxelman etal. (1994) ; whereas and anatomicalstudiesby Norman (1994) and Dickison 2010] herbarium collectionfrom 1966( ( collected leafsampledriedinsilicagelof of thesetwotribes.Taxon samplingforthisstudyincludesarecent field- Roalson etal.2008 ) andtaxonsamplingtherefore emphasizedmembers Napeantheae ( Smith 2000 ; Zimmer etal.2002 ; Roalson andClark2006 ; ses. All otherspecies intheanalysesare represented byonlyoneaccession. resenting twopopulationsof Alabama MuseumofNaturalHistory(UNA).Two separatecollectionsrep- U. S.NationalHerbarium(US),MarieSelbyBotanicalGardens (SEL),orthe have fertilevoucherspecimensarchived attheSmithsonianInstitution’s herbarium voucherspecimens,photographs,andtheliterature. All taxa cies were photographedinthefieldanddeterminationswere verifiedwith species from theOldWorld were represented intheanalyses.Mostspe- Zimmer etal.(2002) sothatallmajortribesfrom theNewWorld andtwo Sequence dataobtainedfrom GenBankforsixspecieswere selectedfrom et al.1999 ; Bremer etal.2002 ) strongly support groups. Molecularphylogeneticstudies( Smith etal.1997 ; Oxelman Qiagen DNeasy disrupter (Qbiogene,Carlsbad,California).DNA was isolatedusingthe gel. Leafsampleswere ground usingaThermSavantFastPrep FP120cell included intheanalysiswere collectedinthefieldanddriedsilica tionship of on Wang et al.(2004) , whichunequivocallysupportsthesister-group rela- Loganiaceae. Thephylogeneticanalysesare rooted with Gesneriaceae, buttheyare clearlynotcloselyrelated toBuddlejaceaeor are recognized assister-taxa totheGesneriaceaeorbasalmembersof bution ( Norman 1994 ). Itisequivocalwhether ( Dickison 1994 ), chemotaxonomy( Jensen 1994 and Gesneriaceraeissupportedbyvegetativefloralanatomy ), morphology, and distri- as closelyrelated totheGesneriaceae.Thecloserelationship of J. L.Clark10003 Additional specimensofthisundeterminedtaxonfrom local- Polymerase chainreaction (PCR)amplificationsfollowedtheproce- Templates ofthenrDNA internaltranscribedspacerregion (ITS)were Taxon SamplingandOutgroup Selection— Peltantherafloribunda Previous studiesstrongly supportmonophylyofBeslerieaeand DNA Extraction,Amplification,andSequencing— Shuaria ecuadorica trnL-trnF Peltanthera ascloselyrelated totheGesneriaceae. TM ) whereas Smith (2000) utilizedDNA extracted from a DNA isolationkit(Qiagen,Valencia, California). trnL Peltanthera intergenic spacerregion ofchloroplast DNA (cpDNA). aeil ad Methods and Materials totherest ofGesneriaceae. intron and and Shuaria ecuadorica Sanango racemosum D. A. Neill&J.L.Clark,placedin hasbeenretained inBuddlejaceae Irwin etal.14055 trnL-trnF intergenic spacerwere ampli- Anetanthus were included intheanaly- hrytre pce are species Thirty-three ). Sanango Peltanthera were selectedasout- × TBE(pH8.3)buffer, l tsu samples tissue All HiernexBenth. Peltanthera n and trnL n and Peltanthera intron as Sanango Sanango based

for 1,000,000generations withatree sampled every50generations.The version 3.1.2( Ronquist and Huelsenbeck2003 ). Fourchainswere run significantly different from randompartitions ofthecombineddata. sequences are depositedinGenBank(Appendix1). parisons withotherGesneriaceaesequences( Zimmer etal.2002 ). All Identification oftheendsITS1andITS2were determinedbycom- sequences were truncated toincludeonlyITS1,5.8S,andITS2regions. Sequencher 3.0(GeneCodesCorporation, Ann Arbor, Michigan).The Model 377 Automated DNA SequencingSystem(PEBiosystems). et al.1990 ). Sequencingwasperformedusingan Applied Biosystems the twoinitialPCRprimersandinternalprimers,ITS3ITS2( White Biosystems, FosterCity, California).Cyclesequencingwascarriedoutwith Prism® BigDye™Terminator CycleSequencingReadyReactionKit(PE specifications,usingthe ABI template DNAsfollowedthemanufacturer’s ( Swofford 2002 ) todeterminewhethertheITSand ence test(ILD; Farris etal.1994 ) wasperformedintheprogram PAUP* corresponding modelsforeachregion. Theincongruence lengthdiffer- best fitthedata.ThefinalBayesian analysishadtwodatapartitionswith evaluated separatelytodetermine the modelofsequenceevolutionthat 3.1 ( Ronquist andHuelsenbeck 2003 ). TheITSand the 24different modelsofsequenceevolutionthatcanbeusedinMrBayes Information Criteriontoassessthesignificanceofaddingparameters to sion ofModeltest3.6( Posada andCrandall1998 ) thatusesthe Aikaike vidual nrDNA andcpDNA datasets.MrModeltest2.2isamodifiedver- used todeterminethemodelofsequenceevolutionthatbestfitindi- genic regions were therefore combined inatotalevidenceanalysis. values from individualdatasets. There were nowell-supportedclades(bootstrap was evaluatedbycomparingincongruence ofstrongly supportedclades vidual ITSand The parsimonyanalysesandcladesupportwere evaluatedfortheindi- sensus tree wasobtained for eachsetoftrees untilallbranchescollapsed. length resulted inanincreased numberofcorresponding trees. A strictcon- with theheuristicsearch optioninPAUP*. Eachsuccessiveincrease intree that collapsedastree lengthwasincreased byone stepinconjunction trees for1,000replicates. Bremer supportwasusedtoexaminebranches bisection-reconstruction (TBR)branchswapping,savingamaximumof10 each search: 10randomadditioncycles,holdingtrees ateachstep;tree with thefullheuristicsearch optionineffect withthe followingsettingsfor Donoghue etal.1992 ). Thebootstrapanalysiswasbasedon10,000replicates using thebootstrap( Felsenstein 1985 ) andBremer support( Bremer 1988 ; tions thanonelarger search withmore iterations. it istherefore bettertoperformmanyseparatesearches withfeweritera- the ratchetoptioncansometimesgetstuckonsuboptimal“islands”and searches were performedinWinClada assuggestedby Nixon (1999) since ous tosupportaclade),andrandomconstraintlevelof10.Separatetree total), amb=poly-(onlyusescharactersthatcanbedefinedasunambigu- search, onetree heldforeachiteration,160characters sampled(10%ofthe analysis inWinClada usingthefollowingsettings:400iterationsper editor. Ten separatetree searches were doneusingtheparsimonyratchet acting astheparsimonysearch engineandWinClada asthetree anddata ysis ofWinClada ( Nixon 2002 ) andNONA ( Goloboff 1994 ) withNONA of equallengthforeachthereplicates. for eachofthereplicates; 1,000randomadditioncycleslimitedto100trees changes: 10randomadditioncycleslimitedto1,000trees ofequallength but didnotfindshortertrees usingthesettingsabovewithfollowing but withtheMULTREES optionineffect. Othersearches were conducted, formed onallmostparsimonioustrees inmemorywiththesamesettings, MULTREES optionnotineffect. Thesecondstageoftheanalysiswasper- (TBR) branchswappingwithnomore than10trees savedforeachrep; additions, holding10trees ateachstep;tree bisection-reconstruction of theanalysiswasdoneusingfollowingsettings:1,000random stage heuristicsearch inPAUP* 4.0b10( Swofford 2002 ). Thefirststage (study numberS2572). Phylogenetic datamatricesare availableinNexusformatfrom TreeBASE to makeminoradjustmentssothatoverlappinggapswere minimized. ment. Becausethesequenceswere nothighlydivergent, itwaspossible version 1.0a1( Rambaut 1996 ) multiplesequenceeditorforthefinalalign- mode. Theresulting sequenceswere thenimportedintotheprogram SeAl aligned usingClustalW( Larkin etal.2007 ) underthemultiplealignment The Bayesianphylogeneticanalysiswas implementedusingMrBayes AlignmentandPhylogeneticAnalyses— DNA chromatograms were editedandcontigswere assembledusing For theBayesiananalysisMrModeltest2.2( Nylander 2004 ) was Clade robustness fortheparsimonyanalysiswasevaluatedinPAUP* Additional tree searches were doneusingtheparsimonyratchetanal- The parsimonyanalysiswasperformedtocompletionusingatwo ≥ 70%)betweenthetwotopologiesthatwere inconflict.Thetwo trnL-F regions (data notshown).Conflictbetweendatasets All sequenceswere initially trnL-F trnL-F matriceswere datasetsare Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. probability (PP)valuesandmeanbranch lengths. analysis ofthe combinedITSand rule consensustree resulting from thepartitionedBayesian tionarity after5,550generations. Figure 2 showsthemajority trees were excludedas burn-inandtheanalysisachievedsta- modeled underdifferent parametervalues.Thefirst20%of tioned thetwodatasets,which allowedforeachmarkertobe G +IforITS.TheBayesian analysispresented here parti- and Crandall1998 ) resulted inGTR+Gfor selection implementedintheprogram Modeltest2.2( Posada trnL-F et al.1995 ; Nixon andCarpenter1996 ; Graham etal.1998 a totalevidenceapproach wasfollowed( ). Kluge 1989 ; Bruneau more resolution overtheanalysisofindividualdatasets.Thus, combined analysisofthetwodatasetsprovided significantly a well-resolved phylogeny(results notpresented here). The tribe Beslerieae( Fig. 1 ). ported (BS=99%;Bremer support=14)asamemberofthe the lowestof87%( also strongly supportedwith bootstrapvaluesrangingfrom ( Fig. 1 ). Monophylyforallgenerawithtwoormore speciesis ported withrespective bootstrap valuesof99%and100% 0.49). Figure 1 showsthe strictconsensusofthesetrees. retention index[RI]=0.72, rescaled consistencyindex[RC]= ous trees (length=1,579 steps, consistencyindex[CI]=0.68, bined ITSand and the lations. TheITSsequencesofthetwosamplesdiffered by7bp trnL-F from theanalysis.Themean pairwisedivergence fortheentire characters. There were noambiguouslyalignedsitesexcluded the analysiscontributed11 ofthe100parsimonyinformative informative characterswas100(11%). Thetwooutgroups of sequences variedfrom 745–890 bp.Thenumberofparsimony variable sites(28%).Thelengthoftheunalignedcomplete The aligned 6 SSEAI OAY [Volume 35 the fragments thatcollectivelycovered theentire region was18.8%(witharangeof8–37.8%). from theanalysis.Themeanpairwisedivergence for theITS characters. There were noambiguouslyalignedsites excluded SYSTEMATICthe analysiscontributed15of319parsimonyinformative BOTANY sequences variedfrom 607–647bp.Thetwooutgroups of were parsimonyinformative.Thelengthoftheunaligned (bp) longwith430(62%)variablesites,ofwhich319(46%) both strands.ThealignedITSdatamatrixwas699basepairs tively covered theentire spacerand5.8SrDNA regions along ing primersproduced overlappingfragmentsthatcollec- sampled duringthe“burn-inperiod”. 50% majority-rule consensustree ofallcompatibletrees, excludingtrees began. Posteriorprobability valueswere determinedbycomputingthe the pointinwhichsamplingwithinposteriorprobability distribution these valuesstabilizeandrandomlyhoveraround ameanwastakenas maximum likelihoodvaluesinaMS-Excelspreadsheet. Thepointwhere burn-in periodwasdeterminedbyconstructing aXY scatterplotoftree 664 Bysa Analysis— Bayesian The individualanalysisofthe The tribesBeslerieaeandNapeantheaeare strongly sup- Analysis— Parsimony The twosamplesof The two DNA SequencingandAlignment— trnL-trnF data setsare significantlydifferent ( region was3.7%.(witha rangeof0.20–10%). trnL-F trnL-F intergenic spacerregions alongbothstrands. trnL-F sequencesdiffered by8bp. trnL-F sequencingprimersproduced overlapping datamatrixwas912bplongwith252 Besleria dataresulted inninemost parsimoni- Shuaria The ILDtestsuggeststhat the ITSand The parsimonyanalysisofthecom- Results ) to100%. represented twodistinctpopu- trnL-F trnL-F Shuaria The fourITSsequenc- datadidnotresult in datawithposterior p = 0.01).Themodel trnL-F isstrongly sup- trnL andGTR+ intron and ance to in theorder ,despiteitssimilarityinoverallappear- make itreadily identifiabletocurrently recognized families and genusbecauseitdoesnotpresent charactersthatwould of tribe Beslerieaeandasthesistergenus(PP =0.99)toaclade Shuaria strongly supportedwithposteriorprobability values of1.00. Monophyly forallgenerawithtwoormore speciesisalso ported withrespective PP valuesof1.00and0.99 ( Fig. 1 ). prostrate herbwithelongate stemsand ing because Anetanthus ses suggeststhat ular data( Figs. 1 , supported asbelongingtothetribeBeslerieaebyourmolec- 2 ). TheparsimonyandBayesiananaly- logical charactersalone. it wasnoteasytoplaceinaknowntribebasedonmorpho- porting theplacementof ment inthatfamily. Evenwithmolecularresults strongly sup- R. Liesner, andthepresent authors)were suspectofitsplace- Other specialistswhostudiedthematerial(e.g.L.E.Skog, seemed tobelongintheGesneriaceae(pers.comm.2007). examined someofthecollectionsandremarked that who included and nooppositeleaves(e.g. herbarium collectionsof arrangement interrupted byalternateleaves( Fig. 3E ). Some opposite butitiscommontofindshootswithleaf acters are autapomorphies atthefamilylevel. on vegetativeandfloralparts. At theleast,lattertwochar- opposite andalternateleaves,lepidotetrichomesfound mon characterssuchasarborescent habit,presence ofboth the Gesneriaceae.However, there are anumberofuncom- flowered cyme) in theBeslerieaeare small herbstosubshrubs ( Table 1 ). herb withabasalrosette of leaves.Theremaining fourgenera The peltateshield isca.100–120 a shallowersinus betweenthelobesare “dentate-lepidote”. lobes thatgiveitastar-like appearance( Fig. 3 ) andthose with mis onashortstalkandisstellate bythepresence of 15–20 et al.(1996) thestellate-lepidoteshieldliesclosetoepider- Webster etal.(1996) . According totheterminologyof Webster of among allknownGesneriaceae, butstrikinglysimilartothose lar stellate-lepidoteordentate-lepidotetrichomesare unique and (3)unicellular–unbranchedtrichomes.Themulticellu- dular (restricted totheinnercorolla surfaceneartheanthers), composed ofthree typesoftrichomes:(1)lepidote,(2)glan- for theGesneriaceae. ( Fig. 3E ) in viduals. Thepresence ofbothoppositeandalternateleaves have confirmedthatalternateleavesare present inmostindi- L. Santi3352 Shuaria The tribesBeslerieaeandNapeantheaeare strongly sup- Phylogenetic PlacementofShuaria— Phyllotaxy— In theflowercharactersandtypeofinflorescence (pair- Indumentum— Anetanthus Croton isstrongly supported(PP =0.99)asamemberofthe Peltanthera L.(Euphorbiaceae) asdescribedandillustratedby wasinitiallydifficult toplaceinaknownfamily is 1, 2) Ti rltosi ws ntal surpris- initially was relationship This 2). 1, (Figs. Shuaria Shuaria ). Fieldobservationsmadebytwooftheauthors sp.+ Peltanthera The leafarrangementin Shuaria The indumentumof Shuaria represents auniquepatternofphyllotaxy n and Tylopsacas cuneatum isatree, 3–5mtall,while canbeunambiguouslyreferred to Discussion isthesistertaxonto inhermonographofBuddlejaceae, Sanango Shuaria Shuaria W. Palacios12109 μ inthefamilyGesneriaceae, haveonlyalternate leaves al 1 ) omn (2000), Norman 1). (Table m indiameter and thestalk (Fg 2). (Fig. Shuaria ecuadorica Tylopsacas Shuaria Shuaria and Anetanthus Tylopsacas s strongly is s usually is isasmall E. Freire & Shuaria and s a is is

Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. rmr upr values. support Bremer (CI =0.68,RI0.72). Tree rooted with 665 CLARKET AL.: SHUARIA (GESNERIACEAE), A NEWGENUSFROMECUADOR 2010] Fig. 1 . . Strict consensusofninemostparsimonioustrees (length=1,579steps)from parsimonyanalysisofnrDNA ITSandcpDNA Peltanthera floribunda . Numbersabovebranches are bootstrapvalues ( > 50%) andnumbers belowbranchesare trnL-F datasets Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. Bayesian posterior probability (PP)values. and ITSdata.Branch lengthsare proportional tothemeannumberof substitutionspersiteasmeasured bythescalebar. Number [Volume 35 SYSTEMATIC BOTANY 666 Fig. 2 . . Majority rule consensustree (excludingburn-intrees) withmeanbranchlengthsfrom thepartitionedBayesiananalysis ofth s onbranchesindicate e combined trnL-F

Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 00 CAKE L:SURA(ENRAEE,ANWGNSFO CAO 667 CLARKET AL.: SHUARIA (GESNERIACEAE), A NEWGENUSFROMECUADOR 2010] Table 1. Morphological characters of Shuaria compared to other genera in the tribes Beslerieae and Napeantheae as well as genera in unassigned tribes (Resia , Cremospermopsis , Peltanthera and Sanango ).

Character Shuaria Besleria Anetanthus Cremosperma Reldia Tylopsacas Cremospermopsis Napeanthus Resia Peltanthera Sanango Habit Tree or Herb or Herb Herb Herb Herb Herb Herb or Herb Herb Tree Tree shrub Leaf arrangement Opposite & Opposite Opposite Opposite Opposite Alternate Congested Opposite Congested Congested Opposite Opposite alternate Fruit type Capsule Capsule Capsule Capsule Capsule Capsule Capsule Capsule Capsule Capsule Capsule Placentation Parietal Parietal Parietal Parietal Parietal Parietal Parietal Parietal Parietal Parietal Parietal Parietal Style persistence Persistent Caducous Persistent Caducous Caducous Persistent Not observed Not observed Not observed Not observed Not observed Not observed in fruit Inflorescence Absent Absent Present or Absent Absent Absent Absent Present Present Present or Absent Absent bracts absent absent Peduncle Present Absent Absent Present Usually present, Present Present Present Present Present Present Present (=hypopodium) rarely absent Calyx lobe Basal only Basal only Basal only 2/3 to full Basal only Basal only Basal only 1/3–2/3 Basal only Basal to 1/3 Basal only Basal only connation Calyx lobe Equal Equal Equal Equal Equal Equal Equal Unequal to Equal Equal Equal Equal equality equal Corolla symmetry Zygomorphic Zygomorphic Zygomorphic Zygomorphic Zygomorphic Zygomorphic Zygomorphic Zygomorphic Slightly Zygomorphic Actinomorphic Slightly zygo- zygomorphic morphic Gibbosity [“spur”] Present Present Present Present Present Present Absent Absent Absent Absent Absent Absent Nectary Absent Present Present Present Present Present Present Present Absent Present Present Present Stamen number Four Four Four Four Four Four Four Four Four Four Five Four Papillate or Reticulate/ Reticulate- Seed surface Reticulate Striate Striate Striate striate Striate Not observed Papillate Striate Striate honey-combed striate lutea ening ofthepeduncle,besessileinleafaxils(e.g. of 4–6(-16)flowersonalongpeduncle)or, byextreme short- (e.g. dense flowerclustersmaybepositionedonelongatepeduncles inflorescences inwhichtheflowersare strongly congested.The branching ofthecyme.MostspeciesinBeslerieaehave ing awell-developedpeduncleandnoncondenseddichasial ered cymes),butuncommoninthetribeBeslerieaeforhav- Shuaria form amattedindumentum. Their densityishighestontheabaxialleafsurfacewhere they can beobservedonmostfloralandvegetativepartsof and mayfalloff withminorabrasion.Thepeltatetrichomes trichome stalksare persistent,buttheshieldisoftenabsent secting scope,duetotheirflatandglisteningsurface.The width is10–15 perhaps with ers are derivedfrom pair-flowered cymesbyreduction). axillary flowers(inaphylogeneticcontext,thesesingleflow- cies of to thecompounddichasiaof inflorescences thusrepresenting “doublecincinni”incontrast be dichasial,whilethesubsequentonesare monochasial(the condensed. However, here onlythefirstbranchingseemsto inflorescences havealso an elongatepeduncleandare not However, in similar becausethecymeunitsare notstrongly condensed. gle axillaryflower(Clark,inprep.). in eral (manyflowered cincinni, incontrasttodichasialcymes of symplesiomorphic. elongate peduncleandnoncondensedlateralcymeunitsis more typicalofotherGesneriaceae andthepresence ofan alternate manner. Theinflorescence structure of ing unitprimordia (from whichflowerpairsdevelop)inan 1975 , 1988 , 1995 Monophyllaea ), withapermanentapicalmeristemproduc- between cences asasynapomorphyfor theBeslerieae. logeny presented here strongly supportsebracteate inflores- of thetribeNapeantheae( Skog andBoggan2006 ). Thephy- classifications haveincluded the absenceorpresence ofbracteoleswasconjectural.Recent sized thattribalboundariesforBeslerieaebasedprimarily on would betheonlygenuswithbracteoles,buttheyalsoempha- from assigning within thecyme. Skog andKvist(2002) , therefore, refrained This genusappearsmuchlike Skog andKvist(2002) in theirdescriptionof 1995 , 2004 ), butitsimportanceattriballevelwasquestionedby defining characterforthetribeBeslerieae( Wiehler 1983 ; Weber the cyme.Theabsenceofbracteoleshasbeenregarded asa bracteoles (prophylls, oftensimplyreferred toasbracts)within fruit withsepticidaldehiscence ( Fig. 4F bers oftheGesneriaceae. All three generahaveadrycapsular ). Itisambiguous era share afruit characterthatisrare intheNewWorld mem- Ifoecne Structure— Inflorescence The greatest resemblance oftheinflorescences of Shuaria A conspicuouscharacterthatdifferentiates theinflorescence Fut Morphology— Fruit Shuaria Shuaria L.).Otherspeciesof Gasteranthus corallinus Anetanthus isingeneralagreement withGesneriaceae(pair-flow- Shuaria n and from thatof ) andtheirdevelopmentmaybesimilarasin R.Br. and Tylopsacas Anetanthus gracilis μ Cremospermopsis , Tylopsacas m. Thesetrichomesare visibleunderadis- Tylopsacas from Ecuadorthecymeisreduced toasin- Sanango Despite extreme differences inhabit thepair-flowered cymesare unilat- Epithema Gasteranthus inflorescences are superficially (Fritsch)Wiehler, bearingacluster ad and , The inflorescence structure in Shuaria n and Cremosperma Cremospermopsis totheBeslerieaebecauseit Hiern,inwhichtheaxillary Blume(describedby Weber Anetanthus, Peltanthera . n n udsrbd spe- undescribed one In ). n and Besleria , buthasbracteoles thesethree gen- , istheabsenceof Cremospermopsis asamember ae single have Shuaria Shuaria Shuaria Besleria is is . . Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. Scale barsfor A &B=50 leafy branchingshoot showingphyllotaxychanging from alternateatbasethen oppositethenalternate nearapex( leaf surface( 6 SSEAI OAY [Volume 35 SYSTEMATIC BOTANY 668 Fig. 3 . . SEM photographsoftrichomes and phyllotaxyof H. vanderWerff 21704 μ m, scalebarforC = 100 ); C.Outersurfaceoflowersepal ( μ m, scalebarforD =150 Shuaria . A. Trichome onpedicelofimmature fruit ( D. A.Neill15912 μ m. ); D.Outersurfaceofmedianpetal( D. A.Neill15912 D. A.Neill15912 ); B.Trichome onadaxial

J. L.Clarketal.10825 ); E. Apex of ). Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. D &F: branching withpaired flowers;F. Mature capsule(lateral view)showingsepticidaldehiscence andpersistentstyle(A: C. Openedflower showingstamensandpistil;D.Portion ofcymewithopenflowers;E.Portion ofcymeshowingdichasialand sub 669 CLARKET AL.: SHUARIA (GESNERIACEAE), A NEWGENUSFROMECUADOR 2010] Fig. Fuentes 1171 4 . .

Shuaria ecuadorica. ; E: Palacios 12109 A. Floweringbranchshowingaxillaryinflorescences withrepeated dichasialbranching;B.Seed withreticulate surface; ). Morales 1922 sequently monochasial ; B& C: Neill 13180 ; Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 7 SSEAI OAY [Volume 35 of L. P. Kvist &L.E.Skog).Fieldobservationsandphotographs SYSTEMATICis acapsulethatsplitsirregularly (e.g. BOTANY Kvist 2000 ). A third type offruit dehiscenceintheBeslerieae subsp. cidally withfourvalvesasseenin cases, fruits splitloculicidallyandthensecondarilysepti- dehisces loculicidallywithtworeflexing valves.Insome in thetribeBeslerieaeisalaterallycompressed capsulethat indehiscent fleshyberriesin fleshy tocapsulesthatsplitloculicidally. Apart from have eitherindehiscentfleshyberries( be plesiomorphic( Weber 2004 ). OthergeneraintheBeslerieae Batalin, of the“Basal Asiatic andEuropean genera”( septicidally dehiscentcapsulesare onlyfoundinsomeclades derived inthisgroup. IntheDidymocarpoidGesneriaceae whether septicidaldehiscenceisplesiomorphic(relictual) or 670 subg. especially inthebasalclades, butalsoin surfaces frequently occurinthepaleotropical Gesneriaceae, row, slightlyortightlyspirallyarrangedcells.Reticulateseed in theNewWorld Gesneriaceaeare striate,withlongandnar- and form.Thesurfaceisreticulate whereas mostseedsurfaces A.rubra cilis Beaufort-Murphy (1983) . Thewingedseedsof trated by Ivanina (1965) andSEM imageswere presented by seed withpustulatesurfaceof close relationship ofthetwogenera.Thedistinctivewinged in theGesneriaceae. Wiehler (1976) , therefore, suggesteda The round andflatseedwithdevelopedwingsisunique Anetanthus studies. dehiscence, willneedtobecarefully evaluatedinfuture which are saidtohavecapsules withirregular orfour-valved in speciesof ance ofdehiscingloculicidally. Whetherornotthisoccurs occasionally furtherdevelopandgivethecapsuleappear- the capsule.Thesesmalltearsinapexofcapsulewill but occasionallyasmalltearorslitisproduced attheapexof that thefruit dehiscencein capsules wasgivenby Wiehler (1976) where hedocumented 1958 ). An interesting noteregarding four- andtwo-valved four-valved itisloculicidal andsepticidal( Leeuwenberg Napeanthus loculicidally ( Kvist andSkog1989 ). Thefruit dehiscencein cence in ondarily septicidally( Skog andKvist2000 ). Thefruit dehis- species where thefruits firstsplitloculicidally andthensec- fruits of Benth. alsohavefour-valved capsules.Itislikelythatthe by photographssuggestthatotherspeciesof Kvist andSkog(1988) . Careful fieldobservationsdocumented ally four-valved insteadofsplittingirregularly asreported by Ecuador suggestthatinsomeindividualsthecapsuleisactu- that, upondehiscence, appearspeltate(from whencethe superficially similar in some speciesof a reticulate seedsurfaceisuncommon,butdocumented in The seedsof Morphology— Seed Sura Platea ad Sanango— and Peltanthera, Shuaria, Cremosperma muscicola were illustrated by Wiehler (1976) andsimilarseedsof Streptocarpus oncogastrus L.E.Skogwere illustratedby Skog (1982) . Haberlea Cremosperma Reldia n and Gardn. istwo-valvedandloculicidalwhen Shuaria Wiehler isadrybivalvedcapsulethatsplits Napeanthus, Gasteranthus Cremosperma Friv., Tylopsacas (Hanst.)L.E.Skog&P. Kvist( Skog and and are similartothoseofsome are notwinged andoftheusualsmallsize Ramonda An interesting charactershared by Didymocarpus Shuaria ( ( isthepresence ofwingedseeds. J. L.Clark6328 Anetanthus euotMrh 1983). (Beaufort-Murphy Besleria Rich.,etc.)andare thoughtto Tylopsacas cuneatum n and , thestandard fruit type Peltanthera Wall. IntheNewWorld isdefinitelysepticidal, Gasteranthus calcaratus Cremosperma muscicola Besleria , and ) from northwestern One feature thatis Streptocarpus Anetanthus gra- is theanther L.)orsemi- Cremosperma Cremosperma Corallodiscus Gasteranthus wasillus- Lindl. ,

Shuariaecuadorica 2000). ( Smith or theywere basedongenes(e.g. taxon sampling( Zimmer etal.2002 ; Roalson etal.2005 , assessed more carefully asmanyofthesestudieslacksufficient 2008 ) tionship oftheNapeantheaeandBeslerieaewillneedto be logeny usingthefloralsymmetrygene relationship usingtwospeciesforeachtribebasedonaphy- from 68–70. Smith etal.(2004) didnotrecover asister-group Napeantheae andBeslerieaewithbootstrapvaluesranging et al.2005 , 2008 ) suggestasister-group relationship between a singlespeciesofNapeantheae( Zimmer etal.2002 ; Roalson genetic studiesusingthesamethree samplesofBeslerieaeand (Bremer supportof10and bootstrapvalueof95).Otherphylo- sister-group relationship betweenNapeantheaeeandBeslerieae the cpDNA marker representatives from alltribeswithintheGesnerioideaeusing from theBeslerieae,fourspecies from theNapeantheae,and them assister-groups. Smith (2000) includedfourteenspecies that includedtaxafrom thesetwo tribesconsistentlysupported the tribesNapeantheaeandBeslerieae.Phylogeneticstudies ship ( Fig. 1 ; parsimony)forasistergroup relationship between of support( Fig. 2 ; Bayesianinference) orunresolved relation- One importantresult ofthephylogenypresented here isthelack (e.g. cies havebracteoles(e.g. taxon samplingisessentialfor accurate assessmentoftheBeslerieaeandNapeantheae.More and includingtheminfuture studies willallowforamore of thesegenerahavespecieswithbracteateinflorescences the lowlandsofnorthernPeru (I.Salinas,Pers.Comm.).Both A third speciesof Colombian departmentsof Antioquia andadjacentBolívar. a genusoftwospeciesrestricted tothehumidforests inthe and deJesús1997 ). genus oftwospeciesfrom ColombiaandVenezuela ( Skog Skog &L.P. Kvsit. lar analysesare Bremer etal.2002 ; Wang etal.2004 ). close relationship withtheGesneriaceae( Oxelman etal.1999 ; though thisisnotconsistentwithotherstudiesthatsupporta transferred toGesneriaceae,orremain inBuddlejaceae even equivocal astowhether Norman (2000) suggestedthatmorphologicalevidenceis mens andaloculicidally, notsepticidallydehiscentcapsule. Shuaria distinct from thethrysemadeupofpair-flowered cymesin rescence of structures ofthesetwogeneraare ratherdifferent: theinflo- generic name Phylogenetic RelationshipofNapeantheaeandBeslerieae— Sampling— Taxon Future Sep 2007(fl,fr), sandstone plateau, 1,130m,3°34 ridge, onsubstrateofmixed sandandclaysoil,below forest inventory“Wawaime Plot”,tallcloudforest on posed EcuaCorrientecopper mine,onridgenear1-ha watershed, tributaryofRío Quimi, belowsiteofpro- El Pangui,Cordillera delCóndorregion, RíoWawaime nov.—TYPE: ECUADOR.Zamora-Chinchipe,Cantón Anetanthus Mroe, Moreover, . Peltanthera Peltanthera sp.usedforthisstudy). Resia Cremospermopsis Resia aooi Treatment Taxonomic Peltanthera Cremospermopsis D. A. Neill&J.L.Clark,gen.etsp. ndhF H.E.Moore and D. A.Neill,C. Davidson &S.Christoph isamany-flowered compoundpanicle, isapoorlyknownandrarely collected A. gracilis . Smith (2000) strongly supporteda Peltanthera ). Butotherwisethereproductive Absent from thecurrent molecu- hasfiveratherthanfoursta- Anetanthus ndhF wasrecently discovered in ) andotherslackbracteoles lk like , ( Skog andKvist2002 ) is ) withlimitedvariability CYCLOIDEA ′ Cremospermopsis 18”S 78°26 Sanango becausesomespe- sol be should , ′ . Therela- 46”W, 22 . E. L. Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 3°20 Río Tsurim entre loscentros Shuarde NumpatkainyBanderas,1,100m, Cantón SanJuanBosco,Cordillera delCóndor, valledel Río Coangos, minute, elliptic,surfacereticulate, lessthan1mmlong. from thebasebysplittingofvalves.Seedsnumerous, cidally dehiscent,withalongpersistentstyleandoftentearing the anthers,stigmabilobed.Fruit abivalveddrycapsule,septi- style extendingtonearthemouthofcorolla tube,evenwith broad baseandatapering apex,transgressing intothestyle, anthesis. Nectaryabsent.Gynoeciumunilocular, ovoidwitha to anthesisandseparatingatanthesis,appearingpeltateafter fertile filaments;anthersincluded,the4connateprior corolla tubefor1–2mm,staminode present, halfaslongthe broad andflattenednear thebase,adnatetobaseof with fourfertilesubequaltodidynamousstamens,filaments half withgland-tippedunbranchedtrichomes. Androecium 1 mmwide,reflexed atanthesis, innercorolla tubeonupper limb offiveequal,rounded lobes,lobesto1mmlongand ative tothecalyxatanthesis,tube4–7mmlong,2wide, pouched ontheventralside,corolla posture perpendicularrel- uniformly white,tubular, gibbousatthebaseandshallowly slightly connateatthebaseandattenuateapex.Corolla 2.5 mmlong,0.4–0.5wideatbase,broadly triangular, long. Calyxuniformlylightgreen, offiveequallobes,1.5– axes oflateralcymeunits1.0–2.5cmlong;pedicels3–5mm number perinflorescence from 6–16;peduncle3–6cmlong; branches, bracteoles(bractswithinthecyme)lacking,flower age leaves,pair-flowered dichasialcymeswith2–4orders of 9–14 perside.Inflorescences emerging from theupperfoli- nected bywebbingformingashield-likescale,lateralveins and adaxiallywithstellatetrichomesradiithatare con- with sparse,unbranchedsingle-celledtrichomes;abaxially glandular teethseparatedatintervalsof±4–5mm;abaxially oblique, apexacuminate,margins serrulate toserratewith to elliptic,sometimesslightlyfalcate,baseacuteandoften 9.5–19.0 cmlong,(1.0-)1.5–5.0wide,narrowly oblong iole 5–15mmlong,shallowlysulcate,glabrous; blade(6.0-) with alternateleaves,nearlyisophyllouswhenopposite;pet- puberulent. Leavesoppositeanddecussate,sometimesmixed 671 with multipletrunks arisingfrom thetree base;youngstems dichasialibus. capsulis septicidedehiscentibusatquecymispedunculatiset floralibus vegetativisquetrichomatibuslepidotisvestitis, foliis oppositisinterdum cumfoliisalternismixtis,partibus CLARKET AL.: SHUARIAcharacterum sequentiumdistinguitur:habituarborescente, (GESNERIACEAE), A NEWGENUSFROMECUADOR 2010] m, 1°30’S,77°56’W, 1Sep1995(fl), US); LosVencedores, Estación ExperimentalPastazaESPOCH, 800–1000 600 m,1°31’S77°30’W, 25 Feb1994(fr), QCNE, US);km17delpropuesto oleoducto ARCO Villano-El Triunfo, Lliquino, 420m,1°29’S77°22’W, 7–10May1992(fl), E.Freire &L.Santi 3352 noroeste delpozoVillano 2,700 m, 1°27’S77°36’W, 3–14Sep1988(fl,fr), 9, 22y25,líneapropuesta deloleoductoVillano-CPF por ARCO, km25 (MO, QCNE,UNA).Pastaza:Cantón Arajuno, campamentos temporales lage ofTinkimints, 1,600m,3°15 del CóndorandRíoChangos,forest onsandstoneridge,eastofShuarvil- UNA); CantónLimonIndanza,Cordillera deHuaracayo,eastofCordillera Zamora-Chinchipe: CantónElPangui,parroquia Tundayme, Cordillera Shuar Warints, 1220m,3°10 Cóndor, cumbre delCerro ChikichikiNainta4kmalsureste delcentro D.A.Neill&J.Manzanares 13180 Small tree 3–5(-8)mtall,withtrunk to6cmDBH,frequently Hoc genusetspeciesaceterisGesneriaceiscombinatione AdditionalSpecimensExamined— ′ 44”S 78°14 K!, LOJA!,NY!,MO!,P!,QCA!,UNA!,US!,USM!). 15912 (holotype:QCNE!;isotypes: AAU!, F!,GB!,HA!, ′ 08”W, 21Oct1999(fl,fr), (MO,QCNE,US);CantónPastaza,Pambayacu, Río ′ 13”S 78°14 (MO,QCNE,UNA,US);Cordillera del ′ R. Pinusetal.RBS-0017 51”S 78°11 ′ 33”W, 9Oct2002(fr), CAO. Morona-Santiago: ECUADOR. P. Fuente etal.1171 W. Palacios 12109 ′ 30”W, 24Mar2001(fl,fr), W. Palacios 10144 (CHEP, QCNE). (MO,QCNE, (MO,QCNE, G. Toasa 8955 (MO,

C.Cerón etal.16759 desde elcampamentomilitar, 1,050–1,100m,4°07’S78°37’W, 17Oct1991, Parroquia Guayzimi,Cordillera delCóndor, caminoalhitodePachicutza D.Neilletal.16363 viduals inanarea ofabout0.1ha(1,000m type localityintheRíoQuimi watershed,of30ormore indi- times occursinratherdenseunderstory populations,asatthe small tree intheshadedunderstoryofprimaryforest. Itsome- pure crystallinesandstonesubstrate.Thespeciesgrows asa plateau where thesandiseroded from above,ratherthanon stone formation,butprimarilyonslopesbelowthesandstone clay soil,derivedinpartfrom thenutrient-poorHollínsand- inces, itoccursat900–1,600melevationonmixedsandand del CóndorinMorona-Santiago andZamora-Chinchipeprov- 700 minPastazaprovince. OntheslopesofCordillera labels onthespecimensfrom lowland Amazonia at420– ridge tops.Soiltypeswere notincludedinthecollection occurs ondissected,hillyterrain,mostlyalongwell-drained Amazonian peneplainofPastazaprovince, known populationsfrom 420mto1,600m.Inthewestern of theCordillera delCóndor, withanelevationrangeforall est, andinlowermontanecloudforest onthewesternslopes Chinchipe, Ecuador, inlowland Amazonian tropical wetfor- from theprovinces ofPastaza,Morona-Santiago, andZamora- per mineconcession,1,230m,3°34 Plot”, 1-hectare forest inventoryplotonsameridge,EcuaCorrientecop- Quimi, tallforest onridge,withmixedsand/claysoil,above“Wawaime (LOJA, MO,QCNE,UNA);RíoWawaime watershed,tributaryofRío is presently known. the speciesisendemic,asfaritsgeographicaldistribution cific epithetrefers tothecountry oforigin,Ecuador, towhich made inforests ownedby rural Shuar communities.Thespe- m, 3°34 tributary ofRíoQuimi,alongroad aboveEcuaCorrienteminecamp,1,300 D.Reyes&C.Morales1197 78°26 Río Wawaime, propiedad delaempresa EcuaCorriente,970m,3°33 900 m,4°18 por launióndelosRíosNangaritzayNumpatakaime,sitioLaWantza, 13947 US); Miazi,sendero haciaelhito,900–1,200m,16Oct1991, W. Palacios&Exp.Tratado CooperaciónAmazónica8192 camino alhitodePachicutza,900–1,000m,4°07’S78°37’W, 18Oct1991(fr), 970 m,3°33 96979 range of Chinchipe provinces corresponds quitecloselytotheknown Shuar territoryinPastaza,Morona-Santiago andZamora- now compriseapopulationofabout110,000. Theancestral Shuar indigenousethnicgroup of Amazonian Ecuador, who 1922 eral oftheEcuadorian localitiesare less than 10kmfrom the del Cóndorin the departmentof Amazonas, Peru, butsev- yet beenrecorded from theeasternslopesof Cordillera trees inthepopulationflowersimultaneously. Ithasnot MO, QCNE,UNA). headquarters, 1,135m,3°26 tion inroad toTundayme, SanMarcos andEcuaCorrientecoppermine military posttoCóndorMiradoroutpost,ca.3.5kmSofjunc- UNA); vicinityofRíoZamoraandvillageQuimi,alongroad from 78°25 opment, trailPiloto,6–8kmSofmineheadquarters,1,291m,3°35 del Cóndorregion, Terreno deEcuacorrienteS. A. copperminedevel- 3°35 Wawaime, 8.1kmSofmineheadquartersonroad tominesite,1,291m, region, vicinityofEcuaCorrientecopperminedevelopment,valleyRío QCNE, SEL,UNA,US,W);CantónElPangui,Cordillera delCóndor Dsrbto ad Habitat— and Distribution Etymology— ′ (MO,QCNE); 51”S 78°25 ′ ′ (MO,QCA);cuencaaltadelRíoNangaritza,Centro ShuarShaime, (MO,QCNE,UNA);Parroquia Tundayme, valledelRíoQuimi, 59”W, 6Oct2006(fr), 57”W, 15May2009(fl), ′ 43”S 78°26 Shuaria ecuadorica ′ 49”S 78°40 ′ 07”S 78°26 ′ 57”W, 7 Apr 2006(fr), The generic nameisbasedontheof ′ 07”W, 26Oct2006(fl), C. Morales&D.Reyes1951 (MO,QCNE,UNA);RíoNangaritza,Pachicutza, (LOJA,MO,QCNE,UNA);CantónNangaritza, ′ 56”W, 15Oct2002(fl), ′ 59”W, 6Oct2006(fl,fr), (MO,QCNE,UNA); RíoWawaime watershed, ′ D. Reyes&C.Morales1191 21”S 78°28 J. L.Clarketal.10825 , andseveralofthecollectionswere ′ 27”S 78°26 Shuariaecuadorica ′ T. B.Croat etal.96736 17”W, 12 Apr 2006(fl,fr), H. vanderWerff etal.21704 W. Quizhpeetal.413 (MO,QCNE);cuencadel ′ 40”W, 1May2008(fl,fr) 2 ) withmostmature C. Morales&D.Reyes (AAU,F, K,MO,NY, Shuaria ecuadorica (MO,QCNE,UNA); (MO,QCNE,UNA, (O QCNE, (MO, s known is J. Jaramillo T.B. Croat (LOJA, ′ ′ 07”S 51”S

Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 7 SSEAI OAY [Volume 35 . 1. Trees, more than3mtall SYSTEMATIC BOTANY migrated from more populatedareas ofEcuador. Ingeneral, cattle pastures andsubsistencecrops bycolonists whohave Deforestation intheregion continuesfortheestablishment of have beenconducted, appropriate habitatwhere intensivebotanicalinventories and inlarge areas oftheCordillera delCóndorregion with ductive adults,butthesepopulationsare widelyscattered, It occursindenselocalpopulationsof30ormore repro- has notbeenfoundinanyformallyprotected area in Ecuador. September, andOctober. and October. Collectedinfruit inFebruary, March, April, mountain range. Ecuador-Peru border thatfollowsthecrest ofthatsub-Andean 672 1. Herbstosubshrubs, lessthan3mtall . ing herobservationson cence terminology, ElianeM.Norman(DLF;StetsonUniversity)forshar- National MuseumofNaturalHistory)forhelpfulcommentsoninflores- Garden. Theauthorsthank JohnBoggan(US;TheSmithsonianInstitution’s 00, andbytheTayor FundforEcologicalResearch atMissouriBotanical by aNationalGeographicSocietyResearch andExplorationgrant6794- National ScienceFoundation(DEB0841958toJLCand0346679DAN), Bafr-upy H T 18 h se srae opooy f the of morphology surface seed The 1983. Wojciechowski T. , F. H. M. Beaufort-Murphy, Porter , M. J. Sanderson, J. M. G., B. Baldwin, inspiring discovery-basedscience. Freeman Dyson(Institute for Advanced Study, Princeton,NewJersey)for helpful commentsonthemanuscript.Thefirstauthorwouldliketothank for facilitatingtheBayesiananalysis,andtwoanonymousreviewers for ful suggestionsontheanalyses,View-Hune Teoh (Universityof Alabama) the illustration,JohnR.Clark(SEL;MarieSelbyBotanicalGarden) forhelp- diagnosis, Alba Arbelaez (MO;MissouriBotanicalGarden) for preparing Gereau (MO;MissouriBotanical Garden) forassistancewiththeLatin Conservation andIUCNRedListcategory— Phenology— Acknowledgments 2. Bractswithintheinflorescence partlypresent. Corolla atanthesiserect relative tothecalyx;corolla nearlyactinom 2. Bractswithintheinflorescence absent.Corolla atanthesis perpendicularrelative tothecalyx;corolla zygomorphic system fordiagnosing seedmorphology . Gesneriaceae utilizing thescanningelectron microscope andanew nuclear of region logeny ITS . The 1995. Donoghue . J. ribosomal DNA:avaluablesource ofevidenceonangiosperm phy- M. and Campbell, S. C. 3. Stamens5,exserted;anthersbasifixed,2-celled,afterpollenshedwallsrecurving, theanthers thusbecomingseemingl 3. Stamens4,included;anthersbasifixed,2-celled,afterpollenshedretaining theirbasifixedand 2-celledappearance; 4. Leavesopposite,occasionallyclustered inabasalrosette . . 4. Leavesalternate,notinabasalrosette Annals oftheMissouriBotanicalGarden seed surfacereticulate withnarrow honey-comb-like wings . . not winged . . 5. Fruit acapsule . 5. Fruit aberry Collected inflowerMarch, May, September, Key totheGeneraTraditionally RecognizedintheTribes NapeantheaeandBeslerieae, 6. Leavescongestedinabasalrosette . . 6. Leavesevenlyspacedonanelongatestem .

7. Petiole5cmorlonger . . 7. Petioleslessthan2cmorbladesessile . . 9. Conspicuousstomatalclustersabsentonthelowerleafsurfaces . . 9. Conspicuousstomatalclusterspresent onthelowerleafsurfaces Shuaria Support forthisproject wasprovided bythe Shuaria ecuadorica ieaue Cited Literature 8. Peduncleselongatewithwell-developedpedicels;inflorescence bractsreduced orabsent . 8. Peduncleselongatewithsessileorreduced pedicels,inflorescence bractspresent andwelldeveloped . . 10. Inflorescence bractsabsent . 10. Inflorescence bractspresent and Neotropical GeneranotCurrently Recognizedin Assigned Tribes relative to 11. Prostrate herb;seedswingedandflattened . 11. Erect herb;seedsnotwingedandnarrowly elliptic . Peltanthera Selbyana hasnotbeenfound. 2 247–277. 82: : 220–422. 6: and hs species This Sanango , Roy , Dcio, . . 94. A eeaiain f of re-examination A 1994. C. W. Dickison, chloroplast of Congruence 1995. Knapp. S. and Dickson , E. E. A., Bruneau, listed inthecategoryNT(NearlyThreatened). conservation inmostofitsrange ( IUCN 2001 ), andconsideringtheuncertainfuture ofhabitat for estimatedrange,area ofoccupancyandpopulationsize tered populations. According totheIUCNRedListcriteria such asfuture climatechangecouldthreaten thesesmall, scat- similar habitatneartheminingsites.More long-termimpacts including set-asidesandsupportforconservationofareas of are developingplanstomitigatetheirenvironmental impacts, mining companiesthatare beginningtooperateintheregion eliminated byfuture miningoperations,butthetransnational Some ofthepopulations,includingtypelocality, maybe that maintainmostoftheirterritoryasundisturbedforest. cal distributionownedbyindigenousShuarcommunities this speciesisbetterprotected inlandswithinitsgeographi- Gaa, . . J R Kh , . . otn J E Eknadr ad . . H. C. S. and Eckenwalder , E. J. Tucumán, Morton, R. author, B. the Kohn , R. by J. W., Published S. Graham, 2. ver. NONA 1994. P. Goloboff, using approach an phylogenies: on limits Confidence 1985. J. Felsenstein, Dngu, . . R G Omta , . . mt , n J D Ple 1992. Palmer . D. J. and Smith, F. J. Olmstead, G. R. J., M. Donoghue, Fri, . . M Klesö A G Kue ad . ut . 19 etn signifi- Testing 1994. Bult. C. and Kluge , G. A. Källersjö, M. S., J. Farris, Bee, . K Bee , . edr , . rxn R G Olmstead, G. R. Erixon , P. Heidari, N. Bremer , K. B., Bremer, Bee, . 98. Te iis f mn ai sqec dt i angiosperm in data sequence acid amino of limits The 1988. K. Bremer, ...... 2 . 1151 –1167 . data in DNA restriction sitecharacterswithmorphologicalandisozyme SystematicBiology one among discordance and morphological and three moleculardatasets from Pontederiaceae congruence . Phylogenetic 1998. Barrett . Argentina . the bootstrap . Phylogenetic relationships ofDipscalesbasedon and floralanatomy. AnnalsoftheMissouriBotanicalGarden cance ofincongruence . . . nebr , . älrj , n E Brhrain . 2002. Barkhordarian . E. and taxonomic levels . Källersjö, roplast DNA markersandtheutilityofnon-codingDNA M. athigher Anderberg , Phylogenetics ofasteridsbasedon3codingandnon-codingchlo- A. A. phylogenetic reconstruction ...... 4 ...... Solanum 6 . Evolution ...... sect. 7 545–567. 47: Molecular PhylogeneticsandEvolution Taxon ...... 7 ...... 5 . Cladistics 9 783–791. 39: Lasiocarpa 3 601–618. 43: ...... 8 ...... 11 ...... 9 . . . Evolution 0 315–319. 10: . Shuaria ecuadorica Sanango racemosum Canadian JournalofBotany 9 333–345. 79: rhc ...... 3 . orphic 2 795–803. 42: . 10 ...... seed surfacesmoothand y peltateand4-celled; Cremospermopsis rbcL 4 274–301. 24: . . . 2.Vegetative Cremosperma sol be should Gasteranthus sequences. Napeanthus Anetanthus Peltanthera Tylopsacas ... . Sanango Shuaria Besleria Reldia Resia 73:

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Wbr A. 17 opee nlrseztutrn e Gesneriaceen. bei Infloreszenzstrukturen Komplexe 1975. A. Weber, 2002. L. D. Swofford, evo- Molecular 1993. Zimmer . A. E. and Reeve, E. H. Thien, B. L. Y., Suh, phyloge- A 2004. Baum. A. D. and Hileman, C. L. Draper , B. S. F., J. Smith, Ula lo, . n D A Nil . 20 position Phylogenetic 2006. 2004. Cronk . B. Neill. C. Q. and Möller A. , M. N., C. D. Wang, and C. Ulloa, Ulloa for primers Universal 1991. Bouvet. J. and Pautou, G. Gielly , L. P., Taberlet, Familial 1997. Denton. S. D. and Carroll , L. C. Brown , D. K. F., J. Smith, Gasteranthusleopardus Morton) Wiehler, GQ119606, GQ166806, Ecuador, GQ166805, Ecuador, Clark 7502 (US); Cultivated (Colombia,CostaRica, Nicaragua, Panama), Panama, Clark 7143 Cremosperma nobile Wie T J, . rn , . e , n J Tyo 90. Apiiain and Amplifications 1990. Taylor . J. and Lee, S. Bruns , T. J., T. White, Systematic 1996. Smith. A. B. and del-Arco-Aguilar , J. M. L., G. Webster, in 63–158 of Pp. cyme Gesneriaceae. pair-flowered 2004. the A. Weber, of aspects Developmental 1995. A. Weber, of systematics and morphology the to Contributions 1988. A. Weber, Welr H 18 snpi o te etoia Gseica Gesneriaceae. neotropical the of synopsis A 1983 . of H. classification Wiehler, the on report A 1976. H. Wiehler, Welr H 19 r-xmnto o of re-examination A 1994. H. Wiehler, num GQ119600, GQ166800,Ecuador, GQ166799, Bolivia, GQ166798, Peru, Ecuador, 10003 L.E.Skog7777 Clarke, AY047040, AY047099, Cultivated(China,Myanmar, Thailand), (Central America), mation withherbariumacronym inparentheses. sequence notobtained),collectionlocality, andvoucherspecimeninfor- Idnurm. A. and Boggan, K. J. Skog, E. L. Roalson, H. E. A., E. Zimmer, J.L.Clark6790 Igop Ingroup. Appendix C.V. Morton,GQ119601, GQ166801,Ecuador, Société BotaniqueSuisse Bericht derSchweizerischenBotanischenGesellschaft . Botany sequences ofribosomalDNA inWinteraceae . lution andphylogeneticimplicationsofinternaltranscribedspacer Gesneriaceae) . netic analysiswithintribesGloxinieaeandGesnerieae(: Ecuador, disyuntadeungénero guayanés . (Melastomataceae), unaespecienuevadelaCordillera delCóndor, gene regions . of Molecular Biology amplification ofthree non-codingregions ofchloroplast DNA . (*and othermethods).Version 4 . Sunderland : Sinauer Associates . lematic generaoftheLamiales. placement of p 1 2 n in 315–322 Pp. direct sequencingoffungalribosomalRNA genesforphylogenetics . BotanicalJournaloftheLinneanSociety distribution offoliartrichometypesin Berlin: Kadereit . W. J. Springer-Verlag and . Kubitzki K. eds. Avicenniaceae), including vascular Gesneriaceae . der Pflanzen and phylogenyoftheinflorescence of Klugieae -Loxonieae(Gesneriaceae),X.Development,interpretation, M A Ins D H efn, . Sisy ad . . ht Academic White. J. T. and Sninsky , J J. Gelfand, Diego. H San Press : D. Innis, A. M. Gloxinia, Goyazia,Anetanthus 1–219. based onnrDNA ITSandcpDNA 2002 . Phylogenetic relationships inthe Gesnerioideae(Gesneriaceae) tematic positionintheGesneriaceae. eune sequences . (US); Gasteranthus carinatus Titanotrichum oldhamii J. L.Clark8618 J. L.Clark4575 (US); (US); 0 1042–1055. 80: Besleria aggregata 1 1. Taxa, GenBankaccessionnumbers, (ITS, Achimenes misera (US); (US); American JournalofBotany 3 431–451. 63: Cremospserma veraguanum Gasteranthus glaber J. L.Clark8202 Systematic Botany Cyrtandromoea, Titanotrichum C.V. Morton,GQ119602, GQ166802,Ecuador, Gesneriana o. Dctldn, ails ecp Acanthaceae (except Lamiales Dicotyledons, 7, vol. L. E.Skog7903 Systematic Botany J. L.Clark6852 PCR protocols: aguidetomethodsandapplications 7 1105 –1110 . 17: M.Freiberg, GQ119607, GQ166807,Ecuador, J. L.Clark6130 Anetanthus Besleria angustiflora (US); PAUP*. (US); Wiehler, GQ119604, GQ166804,Ecuador, 5 164. 85: (Mart.) Hanst.,GQ119596, GQ166796,Bolivia, (Gesneriaceae)inferred from fourdifferent : 18–28. 1: Lindl., AY047067, AY047126, Cultivated Episcia lilacina Besleria gracilenta J. L.Clark5629 (US); Phylogenetic analysisusingparsimony sp.,GQ119595, —,Ecuador, (enraee. (Gesneriaceae). (US); (US); L.E.Skog&P. Kvist,GQ119605, 9 407–418. 29: Taxon 9 947–958. 29: (US); Sanango racemosum. Besleria ovalifolia trnL-F 2: 41–57. 121: Fritsch,GQ119597, GQ166797, 9 296–311 . 89: Wiehler, GQ119603, GQ166803, 6 65–74. 46: Taxon Besleria variabilis Aeschynanthus andersonii Hanst., AY047091, AY047150, Epithema Gasteranthus lateralis and (US); and The familiesandgeneraof 3 625–632. 43: C.V. Morton,GQ119598, Novon Croton Selbyana Phainantha shuariorum J. L.Clark7619 Achimenes, Eucodonia, trnE-T J. L.Clark7104 . Sanango Cremosperma castroa- American Journalof Beitrage zurBiologie 6 281–285. 16: Rusby, GQ119599, (Euphorbiaceae). : 374–404. 1: L. E.Skog8132 4.Itsnewsys- C.V. Morton, spacerregion Bulletin dela , three prob- trnL-F Selbyana J. L. Clark J. L.Clark (C.V. ; —= C.B. (US); (US); (US); (US); , eds. , Plant J. L. J. L. J. L. J. L. J. L. J. L. 6:

Delivered by Publishing Technology to: Smithsonian Institution Libraries IP: 160.111.254.17 on: Tue, 19 Oct 2010 21:07:38 Copyright (c) American Society for Plant Taxonomists. All rights reserved. L. P. Kvist&L.E.Skog,GQ119611, GQ166811, Panama, Ecuador, (L. E.Skog)L.P. Kvist&L.E.Skogvar. sp., GQ119609, GQ166809,Ecuador, robustus ex Kuntze,GQ119591, GQ166795,Venezuela, H.D.Clarke10351 Napeanthusrupicola Philipson, GQ119588, GQ166792,Ecuador, GQ119587, GQ166791,Panama, GQ166808, Ecuador, 7 SSEAI OAY [Volume 35 SYSTEMATIC BOTANY Clark 6331 674 Fritsch,GQ119589, GQ166793,Ecuador, J. L.Clark8017 (US); Gasteranthus wendlandianus (US); Feuillet&L.E.Skog,GQ119590, GQ166794,Guyana, J. L.Clark6421 (US); Napeanthus subacaulis Reldia minutiflora J. L.Clark8565 (US); J. L.Clark8835 minutiflora, Napeanthus apodemus J. L.Clark9157 (Hanst.)Wiehler, GQ119608, (Griseb.)Benth&Hook.f. J. L.Clark6898 var. (US); (US); GQ119610, GQ166810, J. L.Clark5651 veraguensis Napeanthus riparius (US); Reldia minutiflora J. L.Clark8619 (US); Donn.Sm., Napeanthus Napeanthus (Wiehler) (US); (US); Reldia

Rica, 13180 ica GQ119593, GQ166812,Ecuador, Guyana, (US); lius GQ119586, GQ166790,Cultivated(Ecuador), AY047141, Cultivated(Brazil), (Cuba), (US); D. A. Neill&J.L.Clark,GQ119594, GQ166813,Ecuador, Otrus Outgroups. Gand., AY047039, AY047098, Cultivated(South Africa), B. Hammel24426 (MO). (MO). Rhytidophyllum Tylopsacas cuneatum L. E.Skog8050 K. M.Redden1460 Peltanthera floribunda exsertum (MO); (US); (Gleason)Leeuwenb.,GQ119592, GQ166814, (US); Sanango racemosum Sinningia cooperi Griseb., AY047055, AY047114, Cultivated L. E.Skog7808 Shuaria ecuadorica D. A.Neill15912 Benth.,GQ119585, GQ166789,Costa (US); J. L.Clark8863 (Paxt.)Wiehler, AY047082, (Ruiz&Pav.) Barringer, D. A. Neill&J.L.Clark, (MO); Streptocarpus primulifo- Shuaria ecuador- L. E.Skog7868 (US). D. A. Neill D. A.Neill