Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 much ecological,geographic, andmorphologicalvariation aspossiblefor states ofBahiaand MinasGerais.Samplingwasdesigned tocapture as variation primarily focusedon DOI 10.1600/036364411X553144 © Copyright2011 bythe American SocietyofPlantTaxonomists Systematic Botany Pennington etal.2010 ). ogy inshapingphylogenyis underscored (e.g. Lavin 2006 ; onomic oversightsare rectified andtheimportanceofecol- 1993 ; Pennington etal.2000 ; lands thatare richinsucculenttaxa(sensu Schrire etal.2005 Prado and Gibbs ). Thesetax- lineages withapredilection toseasonallydrytropical wood- dict relatedness aswellmorphologyespeciallyamong ter species.We argue thatecologyandgeographyoftenpre- Coursetiarostrata demonstrates notonlytheprevious misclassificationof pedicel. postpollination resupination viaa180degree twistingofthe tion 1988 ). southern edgeofthedistribution (Nees &Mart.)Benth.( Lavin 1988 ). caatinga endemics, diversification events. gests thatthecaatingageographicallyconfinesevolutionary than twospecies/taxaendemictothecaatinga,whichsug- ( Queiroz 2006 , 2009 ). Ofthe77 legumegenera,36havemore total taxareported, orabout athird ofthespeciesinflora sity ofLeguminosaewith77genera,293species,and322 woodlands generallyharboringarborescent cactiandadiver- ing species, pollination resupination. The otherknowncaatinga-inhabit- by paripinnateleavesandflowersthatdonotundergo post- Coursetia cm long).More importantly thisspecieswasclassifiedwithin nosed inpartbyitslarge resupinate flowers(e.g.petals3–4 Here, wepresent amolecularphylogeneticanalysisthat Coursetia The caatingaineasternBrazilcomprises seasonallydry Coursetia ao Sampling— Taxon Craccoides substitution ofabout2–3 of thespeciesstemcladesandabout17MaforRostratastem.Theseageestimateswere biasedyoungandare associatedwith and geographically confinedcladeissuggestedbyitsphylogeneticisolationwithin coalesce withrespect tonuclearribosomal5.8Sandinternaltranscribedspacer(ITS)sequencesfor woodlands. vegetation. are richinsucculenttaxa.Thisphylogeneticsignature ismore likelytobefoundinlineagesharbored by thisthanothertype with relatively oldminimumageestimatesare suggestiveoftheevolutionarystabilitylocalpatchesseasonallydrytropi with respect tochloroplast Coursetia vicioides Abstract— Keywords— C. vicioides Sequence AnalysisrevealtheMonophylyofThreeCaatinga-inhabitingSpecies section DC.wasthoughttoincludetwodistantlyrelated 1 (2011), 36(1):pp.69–79 Herbario, UniversidadeEstadualdeFeiraSantana,Km03BR116, 44031-460,FeiradeSantana, Coursetia vicioides (DC.)Lavinbecauseofitsflowersthatundergo (Leguminosae)FromEasternBrazil:NuclearRibosomalandChloroplastDNA Three woodyspeciesof butalsotheexistenceofanoverlookedsis- . All comefrom theSouthernSertanejaDepression ofthecaatinga,andfirsttheseisherein described.Theantiquityo olsec ipra lmtto egahc hlgntc tutr hlgntc ih cnevts esnly dry seasonally conservatism, niche phylogenetic structure , phylogenetic geographic limitation, dispersal coalescence, aeil ad Methods and Materials Coursetia Sampling formorphologicalandDNA sequence

2 Coursetia rostrata Plant SciencesandPathology, MontanaStateUniversity, Bozeman59717U.S. A. wasclassifiedwithin × Coursetia , agroup characterizedprimarily 10 trnD-T −9 , was poorlyknownfrom the substitutionspersiteyear, anexpectedrateforwoodyplantlineages.MultipleDNA sequenceaccessions sequencesfor speciesfrom eastern Brazilinthe Coursetia rostrata uin Pgnci e Queiroz de Paganucci Luciano Coursetia Coursetia rostrata Benth.and

3 Author forcorrespondence ([email protected]) from easternBrazilare here classifiedintotheRostrataclade, omnctn Eio: ak . Simmons P. Mark Editor: Communicating Coursetiacaatingicola Coursetia C. vicioides a diag- was (Lavin sec-

69 . CoalescenceofconspecificnuclearDNA sequencesamplescombined with onlyonetree retained per replicate. Gapswere treated asmissing each analyzedusing theheuristicsearch parameters mentionedabovebut implemented inPAUP* 4.0b10,where 10,000bootstrapreplicates were mated withnonparametricbootstrap resampling ( Felsenstein 1985 ) as well supported( Maxtrees wassetto10,000,theconsensusofwhichcaptured allbranches most parsimonioustrees. All characterswere unweightedandunordered. tree-bisection-reconnection (TBR)branchswapping,andretention ofall These strategiesincludedSIMPLEand CLOSESTadditionsequenceswith search strategiesimplementedinPAUP* 4.0b10 Altivec ( Swofford 2002 ). of Simmons (2004) . Parsimonyandbootstrapanalysesusedtheheuristic using thedefaultparameters( Larkin etal.2007 ) orthesimilaritycriterion Codes, Ann Arbor, Michigan)and alignedwithClustalXversion2.0.10 and thoseforthe reaction conditionsforthe ITSregion were describedin Lavin etal.(2003) the analysisofITSregion. Amplification andsequencingprimers region ( Shaw etal.2005 ) were alsogeneratedtovalidatefindingsfrom Chloroplast DNA sequencesfrom thephylogeneticallyinformative variation inlegumes(e.g. Lavin etal.2003 ; Coursetia Riley-Hulting etal.2004 ). al. 1995 ) wasexhaustivelysampledwithrespect totheeasternBrazilian California). ThenuclearribosomalDNA ITS/5.8S (ITS)region ( Baldwin et lated from leaftissueusing theQiagenDNeasyplantminikit(Valencia, leaves andasouthernSouth American geographicaldistribution. combining postpollinationresupination oftheflowerswithparipinnate Coursetiahassleri ern Brazil( Lavin 1988 ). Lavin (1988) hypothesizedasisterrelationship of ern Argentina, centralBolivia,westernParaguay, andextreme southwest- Chodat. Thisspeciesoccursinseasonallydrytropical woodlandsofnorth- Coursetia was doneinaneffort todetectthepotentialsistergroup oftheBrazilian included asoutgroups were theearlybranchingspecies of as amixofMesoamericanandSouth American Coursetiaaxillaris the closestrelatives of Sphinctospermum the genera the studyof Lavin etal.(2003) and Duno deStefanoetal.(2010) where samples were mappedusingDIVA-GIS ( Hijmans 2009 ). rostrata lections ofageographicallyandphenotypicallydistinctform Estadual deFeiraSantana,).Thesespecimensincludedrecent col- were studiedfrom theHUEFSHerbarium(Herbarium attheUniversidade C.rostrata each speciesofeasternBrazilian Reverse andforward reads were assembledwithSequencher(Gene DNA SequenceDataandAnalysis— Outgroups were selectedfrom amongtribeRobinieaefollowing 1 n at Lavin Matt and Coursetia reported in Lavin (1988 ; p.92).Geographicaldistributions ofall becausethislocusshowsmuchintraandinterspecificgeographic clade.Ofnotehere istheinclusionofsamples and Olneya C. vicioides andminimumageestimatesofabout9Maforeach and Rose,and J.M.Coulter&Roseand ≥ trnD-T A. Gray, 90%)bybootstrapanalysis.Clade supportwasesti- C. vicioides Coursetia region were describedin Shaw etal.(2005) . Coursetiacaatingicola reported in Lavin (1988) , additionalspecimens Genistidium Robinia 2 , 3

. Also identifiedin Lavin etal.(2003) and becausetheyare uniqueinthegenus Coursetia Coursetiacaatingicola L., I.M.Johnstonwere identifiedas Poissonia . Inadditiontothespecimensof Total genomicDNA wasiso- C. insomniifolia and Baillon, cal vegetationthat Coursetia s ofNeotropical C. rostrata ITSratesof , , Coursetia hassleri Peteria C. rostrata Lavin)aswell samples. This tropical Coursetia , and , f this A Gray, A. Coursetia trnD-T , (e.g.

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 ing all provides strong supportformonophylyofacladecompris- rate smoothing. narrow rangeofoptimal smoothing valuesduringpenalizedlikelihood nomically sampledexhaustivelyandbroadly, anddidnotconverge ona sequences includedlimitednucleotidesequencevariation,were nottaxo- analyses were performed ononlytheITSsequencesalone.The parametric ratesmoothing(NPRS)methods( Sanderson 1997 ). Therates the rateconstant(LF; Langley andFitch 1974 ) andtheratevariablenon- ( Sanderson 2002 ). Thesewere compared totheestimatesderivedfrom age estimateswere derivedviapenalizedlikelihood(PL)ratesmoothing other ageestimatestoward youngerages(cf., Lavin etal.2004 ). Minimum of 41Ma).Thisminimumroot ageconstraintwasusedbecauseitbiasesall Lavin etal.2005 , whichhadameanageestimateof 38Maandmaximum was theminimumageestimateforRobinieaecrown clade(node74in straining theageofroot oftheRobinieaecrown cladeto31Ma,which Lavin etal.(2003 , 2005 ). Minimumratesandageswere obtainedbycon- nucleotide substitutionratesandagesofspecifiedclades,asdescribedin trees atstationaritywere retained from bothruns. [Volume 36 SYSTEMATIC BOTANY terminal taxaby743alignednucleotidesitesandthe data setscontainedlessthan0.1%missingentries.TheITSsetof90 Wilcoxon signed-rank( Templeton 1983 ) tests,asimplementedinPAUP*. ous onesusingtheKishino-Hasegawa( Kishino andHasegawa1989 ) and Coursetia enforced thenonmonophylyofconspecificITSaccessionsspecies clade andother Brazilian data becauseindelvariationwaslackingamongthespeciesofeastern 70 pled every5 Posada 2008 ). Markovchainswere run for5 ( Akaike 1974 ) implementedinjModelTest ( was selectedfortheentire ITSregion viathe Akaike informationcriterion Guindon andGascuel2003 ; at defaulttemperatures. ThenucleotidesubstitutionmodelGTR+IG of parameterswere eachinitiatedwitharandomtree andfourchainsset rate runs ofaMetropolis-coupled MarkovchainMonteCarlopermutation were estimatedseparatelyfortheITS1,5.8S,andITS2regions. Two sepa- tree andnucleotidesubstitutionparameterspace.Substitutionparameters and Ronquist2001 ; Ronquist andHuelsenbeck 2003 ) wasusedtosearch jected toanevolutionaryratesanalysis.MrBayesversion3.1( Huelsenbeck 1997 ) wasperformedtogenerateasetofphylogenetictrees thatwere sub- site substitutionratesamongtheITS1,5.8S,andITS2regions. netic analysis,whichfoundhomogeneityofbasefrequencies andamong- of divergent paralogswasalsosuggestedbyaBayesianMCMCphyloge- ITS sequenceswere detectedbysamplingconspecificaccessions. A lack ual variation(i.e.from putativeheterozygotes). Nodivergent paralogous aligned readily andrevealed nodivergent 5.8Ssequencesorintraindivid- 50°C andwassequenceddirectly. Forward andreverse reads contigedand The ITSregion wasPCR-amplifiedusingalowannealingtemperature of founded bythepresence ofparalogouscopiesITS(e.g. Bailey etal.2003 ). Treatment) for datageneratedduringthisstudy. as outgroups andinRepresentative SpecimensExamined(Taxonomic et al.(2003) and Duno deStefanoetal.(2010) forRobinieaespeciesused (acronyms following Thiers 2010 ). Voucher detailsare reported in Lavin housed atCICY, F, HUEFS,HUH,MEXU,MICH,MO,MONT, NY, andUS 2003 ; Duno deStefanoetal.2010 ). Voucher specimensforallsamplesare (S10438). Individualsequenceswere depositedinGenBank( Lavin etal. 25 terminalsby1,662alignedsitesare bothaccessionedwithTreeBASE Coursetia clade ishence somewhat earlybranchingwithin thelarger North andCentral American species( Fig. 1 ). TheRostrata are nestedwithinasubcladeof Coursetiahassleri to theRostratacladeasare otherSouth American clades. resolved. Mesoamerican Rostrata clade).Nodefinitive sisteroftheRostratacladeis ula IS Data— ITS Nuclear The program r8sversion1.71( Sanderson 2004 ) wasusedtoestimate The ITSdatasetcontainednomissingentrieswhereas thatofthe Eouinr Rts Analysis— Rates Evolutionary Phylogenetic analysisofnuclearribosomalsequencesmaybecon- from thecaatinga,were evaluatedagainstthemostparsimoni- Coursetia Coursetia clade. × 10 Coursetia 4 ornotphylogeneticallyinformativeamongtheRostrata generationsuchthat100nonautocorrelated Bayesian comprisestwosubcladesbutregardless both accessionsfrom easternBrazil( Fig. 1 ; the . Trees derivedfrom constraintsanalyses,which Parsimony analysisoftheITSdataset Coursetia Results A Bayesiananalysis( Yang andRannala Coursetia cladesare ascloselyrelated × 10 thatincludesmostly 6 generationsandsam- trnD-T datasetof trnD-T trnD-T

Coursetia eastern Brazilian Taxonomic Implications— and flowersatanthesisdonot. ing beentwistedwhereas thepedicelsbearingflowerbuds Coursetiarostrata However, uponcareful inspection,thefloralpedicelsof ers obscure thetwistedpedicelin long pendantinflorescence rachisesandresupinate flow- 3.4 clades ( Table 1 ). Thefastestsubstitutionrate,approximately one Maorlessageestimatesfortheindividualspeciescrown Ma ageestimatefortheRostratacrown contrastwiththeca. age estimatefortheRostratastemcladeandroughly nine youngest ageestimates( Table 1 uniform estimatesofsubstitutionratesassociatedwiththe ). Theapproximately 17Ma that formthenewlydetectedsistercladeof trait wasobviousin of ashortanderect toascendinginflorescence rachis,this mature legumewasborninaresupinate position.Because all showedevidenceofatwistingthepedicelsuchthat were scrutinized forpostpollinationfloralresupination, and two extremes ( Table 1 ). mate fortheRostratacrown isintermediatebetweenthese individual speciescrown clades.Thesubstitutionrateesti- 1.8 Rostrata stemclade,whereas theslowest,approximately with thesingleaccessionof tribution of that geographicallyliesonthesouthwesternedgeofdis- detected lineage(the“southwesternaccessions”in Fig. 1 ) rostrata two. All accessionsofthemore westerlydistributed ern Bahiaastheweaklysupportedsistertoremaining ing parameters.Oneofthesevalues,10 a smallrangeofoptimalpenalizedlikelihoodratesmooth- of the One ofthesewassupportedbya66bpsinsertionatthe3 western accessions”were notdetectedwiththeITSanalysis. tinct sequences couldnotbeamplifiedforthislocus).Thetwodis- clade of“southwesternaccessions”( Fig. 3 ; cally distinctsubcladeswere resolved, oneofwhichwasthe the Rostrataclade,three well-supportedandgeographi- was tothesouthernSouth American to variousNorthandSouth American clade waswellsupportedanditrevealed tobeasclose in theITSanalysis( Fig. 3 ). ThemonophylyoftheRostrata resolved here were generallyconsistentwiththoseobtained Hasagawa test; constraint trees are significantlylonger( est tree obtainedbytheunconstrainedanalysis(788).These most parsimonioustrees sixstepslonger(794)thantheshort- nonmonophyly ofthe“southwesternaccessions”results in ships are wellsupported. A constraintsanalysisenforcing the Phylogenetic RelationshipsoftheRostrataCladeand Morphology— Within theRostrataclade,three mainlineagesare resolved Eouinr Rts Analysis— Rates Evolutionary Data— trnD-T Chloroplast × × 10 10 Coursetia rostrata trnD-T are strongly supportedasacladesistertonewly −9 −9 and substitutionpersiteyear, isestimatedforthe substitutionpersiteyear, isestimatedforthe Coursetia rostrata region ( Fig. 3 ). C. vicioides p bearingmature fruits show evidenceofhav- =0.0143,Templeton test). pcmn o of Specimens Coursetia Coursetia vicioides cladesresolved assisterstothe“south- insect. Discussion placed Lvns 18) asfcto o the of classification (1988) Lavin’s Coursetia vicioides i. ) Al f hs relation- these of All 2). (Fig. Craccoides Coursetia hlgntc relationships Phylogenetic C. rostrata rs vldto yielded validation Cross Coursetia rostrata Coursetia hassleri andinthespecimens 1.0 Coursetia from easternBrazil . Suchaclassification p , produced themost =0.0141,Kashino- Coursetia vicioides in rm southeast- from C. rostrata Coursetia speciesasit Coursetia i. 4). (Fig. . Within . . The . sect. ′ end

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 Examined). those beginningwith“GQ”in Duno deStefanoetal.(2010) , andthosebeginning with“HQ”intheTaxonomic Treatment (seeRep bers followtheterminaltaxonlabels. Theassociatedvoucherinformationforaccessionsbeginningwith “AF”are reported in L the ChapadaDiamantina.Outgroups comefrom North America unlessotherwiseindicated(i.e.SA =from South America). GenBankac into aregion surrounding thenorthernChapadaDiamantina(N;see Fig. 2 ) andonesurrounding thesoutheastern(SE)andsouth 01 QERZADLVN RZLA OREI 71 QUEIROZ AND LAVIN: BRAZILIANCOURSETIA 2011] Fig. 1 . . One ofthemostparsimonioustrees derivedfrom theanalysisofITSsequences.Theaccessions Coursetia rostrata are roughly classified resentative Specimens avin etal.(2003) , for central (SC)edgeof cession num- Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 habit (mostly large shrubs tomultistemmedtreelets) com- tive withinsect. largest viaaccrescent growth. TheRostrataclade isdistinc- with thedistal-most(including theterminal)leafletbeing Craccoides render squarishseedchamberscharacterize nounced verticalimpressions onthewallsofvalves that parallel dorsal(placental)and ventralmargins and pro- clade shouldbeclassifiedwithin iting pollinatorsoffruit set.Thus,allmembersoftheRostrata lination twistingofthepedicelpossiblyasanindicationtovis- 1988 ), theconstituentsofRostratacladeundergo apostpol- 3 ). Likeothermembers of well supportedmonophyleticgroup, theRostrataclade( Figs. 1 , suggest thatthecaatinga-inhabiting data, aswellbydiagnosticmorphologies,whichtogether is clearlyrefuted byouranalysisofITSand [Volume 36 SYSTEMATIC BOTANY 72 -46.00. Thesolidbarequals250km. Rostrata group. Thenortheasterncross represents latitude-5.00andlongitude-35.00whereas thesouthwesternrepresents latit Chapada Diamantina(CD),whichisrepresented by thecentralarea lackingcross-hatching andexceptatitsperimeterisdevoid and Pernambuco(collectionsof hatched area represents theSouthernSertanejaDepression, whichiscentered primarilyinthestateofBahiaandadjacent Besides post-pollinationtwisting ofthepedicel,podswith Fig. 2 . . The distributionof . Theleaves alsoare mostlypinnate inthissection Craccoides byhavingalarge-statured woody Coursetia rostrata Coursetia vicioides Coursetia Coursetia Coursetia sect. bles, (bullets), sect. come from outlyingenclavesofcaatinga).TheSouthernSertanejaDepression surrounds theupland Craccoides trnD-T speciesforma Coursetia Craccoides C. vicioides sequence (Lavin sect. . (squares), and sister lineage, as wellcoalescenceofthe ITS sequencevariation withineachof are here distinguished atthespeciesrank.Thecoalescenceof Coursetiarostrata study demonstratesthatthe collections initiallyassignedto common tomostspeciesofsect. ous roots, andmature podswithparallelmargins, allfeatures but hasasuffrutescent habit,fascicledandfusiformtuber- Argentina andParaguayismostsimilartotheRostratagroup tion. phies ofsect. also bythisgroup possessingonlyoneofthesynapomor- ITS sequencedata(andtosomedegree the the Rostratacladewithin leaflets. Inessence,therelatively earlybranchingpositionof innate leaveswithnopronounced distalenlargement ofthe bined withpodstendingtohavesinuousmargins andparip- In additiontothemisclassification of Coursetia hassleri C. caatingicola Craccoides actuallycomprisedtwodistinctlineages that (triangles)ineasternBrazil.Thehorizontallycross- from seasonallydryforests ofnorthern , thepostpollinationfloral resupina- Coursetia Craccoides isrevealed notonlyby Coursetia rostrata Coursetia rostrata ude -17.00andlongitude s ofMinasGerais,Piauí, . trnD-T trnD-T of collectionsthe data),but sequence n its and this , Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 Treatment (seeRepresentative Specimens Examined). Chapada Diamantina.GenBankaccessionnumbersfollowtheterminal taxonlabelsandassociatedvoucherinformationisprovided i ter states(e.g. Nixon andWheeler1990 ). genes anditsdiagnosability usingqualitative orfixedcharac- as evincedbythemonophyly (coalescence)ofitsconstituent a separatelyevolvingmetapopulation (cf., de Queiroz 2007 ) is thusrankedasadistinctspecies inthatitlikelyrepresents lution foreachoftheselineages. Thenewlyresolved lineage variation inthelatter, evincesalonghistoryofisolatedevo- resolved asparaphyletic with respect tothatof 73 QUEIROZ AND LAVIN: BRAZILIANCOURSETIA 2011] Fig. 3 . . One ofthemostparsimonioustrees derivedfrom theanalysisofchloroplast Coursetia caatingicola are roughly delimitedbyadivisionnorth(N)andsouthcentral(SC)oftheupland rich insucculent taxa,andbelongtoagenus thatismostly is ecologically confined toseasonallydrytropical woodlands the three speciesare derived from asinglediversification that absent from theuplandChapadaDiamantina( Fig. 2 ). That in theSouthernSertanejaDepression andare apparently Rostrata cladeare confinedtothecaatingamostlybelow800m Boegah o te Caatinga— the of Biogeography trnD-T sequences.Thetwo Coursetia rostrata The three speciesofthe hools lineages chloroplast n theTaxonomic Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 not observedfortheITSaccessionsofthissamespecies( Fig. 1 ) The The Rostratacrown clade(= deviations (std)are alsoreported. reported assubstitutionspersitemillionyears(s/s/Ma).Standard age estimates. Ages are reported inmillionsofyears(Ma)andratesare constant andanonparametricratesmoothingapproach yieldedolder 4 YTMTCBTN [Volume 36 roplast et al.2009 ). Thegeographicalstructure observedforthechlo- limitation (sensu Hubbell 2001 ) inthisbiome(e.g. Pennington dry Neotropical woodlandsis evidenceofstrong dispersal and interspecificcladesof Donoghue 2008 ). Thestrong geographicalstructuring ofintra netic nicheconservatism(e.g.sensu Harvey andPagel1991 ; confined tothesamevegetation,isevidenceofphyloge- SYSTEMATIC BOTANY The The Rostratastemclade ing valuesthatfellwithintheoptimumrange(10 estimates whilerendering themostuniformrateestimates.Othersmooth- from amonganarrow rangeofoptimalvaluesthatminimizedtheage clade derivedfrom ITSsequences. A smoothingvalue(10 74 close-up ofupper leafletsurface(upper, 1mm).J.Pod (1cm). of cm). B.Resupinate floweratanthesis(1cm).C.Leaf (left, 1cm)andclose-upofupperleaflet surface(right,1mm).D.Pod Coursetia vicioides Fig. Table and C. caatingicola C. rostrata 4 C. rostrata . . trnD-T 1 . .

Coursetia rostrata Penalized likelihoodageandrateestimatesfortheRostrata crown stemclades) accessionsof Clade con0000001 0.0005 0.0018 0.0 0.0 crown bornfrom leafynode(1cm).F. Flower preanthesis (1mm). G.Wing petal (1mm).H.Standard petal (1mm).I.Leaf(lower, 1 C. caatingicola (left, A-D) and Coursetia Coursetia rostrata

mean age specieswithinseasonally 16. 7 (Ma) C. vicioides . . .050.0003 0.0025 2.2 9.3 1.1 0.0 -10 (age) 0.7 2.2 std i. ta was that 3) (Fig. 1.0 (right,E-H). A. Longpendantracemeof ), aswellarate mean rate (s/s/Ma) 0.0018 0.0034 1.0 ) waschosen std (rate) 0.0007 0.0004 × size ofthehaploidgenome(e.g. Zink andBarraclough2008 ). is probably afunctionalsoofthesmallereffective population Sertaneja Depression harbors mostofthediversity. Four prises sevencentersofendemismwhichtheSouthern cerrado andlowland Amazonian forests. from otherSouth American vegetationtypessuchasthe (cf., Queiroz 2006 ), atheaterofplantevolutionthatisdistinct one ofmanylegumediversificationsconfinedtothecaatinga likely tobeanartifactofundersampling.TheRostratacladeis Rostrata cladehavebeensampled,thisoldageestimateisnot val). Giventhatallthepotentialextantsisterlineagesof least 12Ma(attheyoungerendof95%confidenceinter- clade intheregion oftheSouthernSertanejaDepression ofat rates strengthens theresident ageestimateofthe Rostrata rates estimatedhere withotherstudiesevenbiasingforfaster region forwoodyperennials. Thisconcordance ofsubstitution 0.4–7.9 (mean=2.2)substitutionspersiteyearfortheITS range estimatedby Kay etal.(2006) forwoodyperennials, i.e. Furthermore, therateestimatesfallintomiddleof were madewithdifferent ageconstraintsandtaxonsampling. et al.(2003 ; pp.396–397,nodeD1).Thesesimilarestimates 17 Ma,are essentiallythesameaswasthoseestimatedin Lavin 10 The rateandageestimatesfortheRostratastemclade,3.4 According to Queiroz (2006) , thecaatingadomaincom- −9 substitutionspersiteyearandameanageofabout Coursetia rostrata bornfrom leaflessnode(scalebar=1 (1 cm).E. Ascending raceme cm)and Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 rumque quam10cmminore, typicequamfoliosubtendenti cum petalisprincipaliter lavendulis,rachi racemorum ple- morphologia florali,e.g.floribus inanthesinonresupinatis cum tum partiumvegegetativarum e.g.foliolos plures perfolium Coursetiacaatingicola 1. Racemeascending,bornfrom theaxilsofwelldevelopedleavesalongnewlygrowing stems;thelongerrachises 1. Racemependant,mostlybornonshortshootsfrom leaflessnodesontwigsproduced duringprevious growing seasons; cent cerradobiome( Queiroz 2006 ). tion thatrelatively fewcaatinga legumes(9%)entertheadja- short andlongterm.Thisideaissupportedbytheobserva- tism inseasonallydrytropical woodlandsplaysoutoverthe vegetation. Thissuggeststhatphylogeneticnicheconserva- South American areas withseasonallydry, succulent-rich 75 have disjunctdistributionsbetweenthecaatingaandother 53% are endemic.Mostof therest (29speciesorabout10%) Sertaneja Depression ( Barneby 1998 ; Queiroz 2006 ). Calliandra for example,includes40endemicspeciesof harbors endemicdiversifications.TheChapadaDiamantina, mism inthecaatinga(e.g.SouthernSertanejaDepression) Furthermore, eachofthesevenconstituentcentersende- dry tropical woodlands,andinthecaatingaspecifically. niche conservationischaracteristicofplantsinseasonally et al.(2005) and Pennington etal.(2006 QUEIROZ AND LAVIN: BRAZILIANCOURSETIA , cal woodybiomeormetacommunityrecognized by 2009 Schrire). Phylogenetic the vegetationisclassifiedwithinseasonallydrytropi- herbs, andsucculentsmainlyoftheCactaceae.Moreover, canopy, oftenspinyplantswithcaducousfoliage,ephemeral Depression corresponds totypicalcaatingawithlow totall 144 endemiccaatingalegumespecies.TheSouthernSertaneja and J.W. Grimes,and Mysanthus Tabaroa genera are endemictotheSouthernSertanejaDepression, 2011] Coursetiacaatingicola Of the274legumespeciesoccurringincaatinga,144or 2. Leaflets14–22perleaf,distinctlymore hairyonthelowersurface;inflorescence rachis5–26cmlong,stipitate 2. Leaflets32–42perleaf,equallyhairyonbothsurfaces;inflorescence rachis5.5–7.5cmlong, distinctlylessstipitat ( to bedistributed),HQ158097(ITSsequence),HQ158041 Queiroz 12815 para Jaborandi.13°35 Bahia: .ca.27kmSdeS.FélixdoCoribenaestrada 5–10 (-12);southernBahiaandadjacentnorthMinasGerais . sparsely glandular;flowersnotresupinate atanthesis;ovarysparselyornotallstipitate-glandular; seeds calyx distinctlymore denselystipitateglandularthantheadjacentinflorescence rachisorallthesestructures are 10–15 mmlong(in 5.5–7.5 cm(in glandular; seeds11–17 (-20);throughout BahiaandadjacentnorthernMinasGerais( Figs. 2 , 4 ) . calyx, andracemerachisequallydenselystipitateglandular;flowersresupinate atanthesis;ovary denselystipitate the longerrachises15–55cmlong;standard petalatanthesispredominantly whitishand30–35mmlong;pedicels, trnD-T Coursetia rostrata L.P. Queiroz, G.P. Lewis&M.F. Wojciechowski, glabrous; southeasternBahia ( Figs. 2 , 4 ) 10–11 . . mmlong,predominantly whitishatanthesis;ovaryglabrous; legumeabout4mmwide by5.0–7.5cmlong, glandular astheadjacentpedicelsandcalyces,withupto40nodes; 3–4mmlong;corolla banner sericeous; southwesternBahiaandadjacentnorthcentralMinasGerais ( Figs. 2 , 5 25–30 mmlong,predominantly lavenderatanthesis;ovarydenselysericeous;legume6–7mmwide by5.0–9.0cmlong, – 6 . ) glandular thantheadjacentpedicelsandcalyces,withupto12nodes;10–20mmlong;corolla banner , noneofwhichenterthesurrounding Southern G.P. Lewis& A. Delgado, sequence). C. caatingicola (holotype:HUEFS!;isotype:MONT!,and C. vicioides L.P. Queiroz, sp.nov.—TYPE: BRAZIL. precipue congruens, sedpraesertim L.P. Queiroz, sp.nov. quoadadspec- Goniorrhachis ′ 10 ) or5–26cm(in ) orpredominantly lavenderand30–35mmlong(in ″ S, 44°19 Blanchetiodendron Taub., asare 57(40%)of Key totheSpeciesof ′ 12 C. vicioides ″ W, 545m, Calliandra ); standard petalatanthesispredominantly whitish and Barneby Barneby sect. L. P. C OURSETIA C.vicioides the Rostratacladeofgenus to manyendemicradiations.Oneofthesmallertheseis older stemclades( Table 1 ; cf., Barraclough 2010 ). coalescence andshortcrown cladescontrastingwithmuch revealed bygeneticsamplingwithsignatures ofintraspecific graphically localizedpopulationswithsmalleffective sizesis wise render thisvegetationfire-prone. Thepersistenceofgeo- moisture regime thatminimizesgrasscoverwould other- the Pleistoceneprobably becausetheytoleratetheerratic grants tendtopersistforevolutionaryperiodsthattranscend predilection tothisvegetationtype.Onceestablished,immi- are likelytobecolonizedbyimmigrantsthatalready havea gests thatpatchesofseasonallydryNeotropical woodlands and Pennington etal.(2000) . Theevidenceprovided here sug- ing thePleistocene,asadvocatedby Prado andGibbs(1993) was drivenbythefragmentationofthisvegetationtypedur- ciation inSouth American seasonallydrytropical woodlands Pennington etal.(2004) toreject thehypothesis thatmostspe- Neotropical woodlandscorroborates evidence used by Sertaneja Depression andelsewhere inseasonally dry dular, 9–17mmlong,articulated withthecalyx,persisting early caducous, pedicelssericeous,strongly stipitate-glan- flower bracts2–3mmlong,broad atbaseandlineardistally, 12 nodes,eachwith1–2flowers atanthesisanyonetime, densely sericeous,sparselystipitate glandular, withupto long. Inflorescence withanascendingrachis5.5–7.5cm long, densely sericeousbelowthan above,stipuleslinear, 5–10mm leaf, mostlyelliptical,8–20mm longby4–7mmwide,more rachis sericeous,canaliculate,exstipellate,leaflets32–42per et leguminematuro cumusque12seminibusdiffert. minus glandularibus,ovariopilisglanduliferissparsevestito, breviore etquampedicelliscalycibusqueadjacentibusvalde (1988 , p.48)keytosectionsandspeciesof lowing keywouldreplace thefirsthalfofcouplet33in Lavin’s The SouthernSertanejaDepression ofthecaatingaishome The diversificationof Scandent shrubs 1.5–3.0 mtall;mature leaves7–16 cmlong, from theCaatinga C. caatingicola , , C. rostrata . aooi Treatment Taxonomic ); pediceland ...... 2 . , andaspeciesherein described.Thefol- Coursetia . . Coursetia e withintheSouthern , whichcomprises, Coursetia Coursetia caatingicola Coursetia vicioides Coursetia rostrata .

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 6 YTMTCBTN [Volume 36 SYSTEMATIC BOTANY 76 H. Stipitate glandularhairscoveringtheouter calyxsurface(1mm).I.Gynoecium (1cm).J.Close-upofsurface ovary. K. E. Pod(1cm).F. Flowerwithpetalsremoved toexposeandroecium andgynoecium(1cm). G.Basifixed(lower)anddorsifixed (up Fig. 5 . . Line drawingsof Coursetia caatingicola . A. Leafy floweringbranch(scalebar=1cm).B.Standard petal.C.Wing petal. D.Keelpetal(B-D,1cm). Close-up of stigma (J-K, 1 mm). Close-up ofstigma (J-K,1mm). per) anther (1 mm). per) anther(1mm). Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 n Fg. 6B–6D. Figs. in ing relatively fewseeds(lessthan10)compared to the mainrachisin C. Erect inflorescence showingnakedpedicelsfrom whichtheabortedflowershavedisarticulated.Thepedicelitselfalsoevent specimens inMay andSeptember. from March, April, May, andSeptember, whereas fruiting gins, seeds5–10. Figures 5 – 6 . glandular, 5–9 congested towards thedistalend;legumesericeous,stipitate 8–12 ovules,style18–22mmlong, withalatrorse pollen brush 1.8 tion offilaments7–11 mmlong,antherslinear-oblong, 1.5– frontal perspective);staminalsheath8–12mmlong,free por- keel rostrate, 22–25 ish nectar-guide, wingsfree from thekeel,25–30 dominantly lavender, 25–30 mm long,sericeous,stipitate-glandular;standard petalpre- 5–6 mmlong,sericeous,stipitate-glandular, calyx lobes5–6 flower aborts.Flowersnotresupinate atanthesis,calyxtube shortly aftertheflowerabscisesbutultimatelydeciduous if older flowertotherightwithpetalsshedisbeginningtwist,whichpostpollinationresupination thatcharacteri 77 QUEIROZ AND LAVIN: BRAZILIANCOURSETIA 2011] Phenology— 6. Fig. × 0.3–0.4mm;ovarytomentose, glandularstipitate,with Poorps f of Photographs × Flowering specimenshavebeencollected C. caatingicola 0.6–0.7cm,withsinuousdorsal-ventral mar- × 13–14mm,curvingtotheright(from a Coursetia caatingicola . D.Podinresupinate position(e.g.thestylewouldultimatelybendupwards ifthefruit wasregularly positioned)show- × 28–37mm,withamostlygreen- . A. Habit ofamultistemmedtreelet orshrub. B.Flowershowingtheasymmetrickeel.Thepedicelof Coursetia rostrata × 10–12mm, , whichtypicallyhas10–20 seeds perlegume.Scalebar=0.5min Fig. 6A and 2 cm forests. with limestone outcrops andinmedium tolowcaatinga in areas notassociatedwithcalciumrichsoilsoratleast not in thetallcaatingaforest whereas Coursetiacaatingicola could alsoreflect edaphicspeciationsinceallcollections of sister speciesbackintogeographical proximity. Peripatry ical isolationwithsubsequent dispersalbringingthesetwo ingicola collection site( 800 melevationbutextendsto1,150atthenorthernmost on calciumrichsoils.Thisspeciesrangesbetween500and Bahia andadjacentnorthcentralMinasGerais,Brazil( Fig. 2 ), ern SertanejaDepression ( Queiroz 2006 ) insouthcentral lands (caatinga)from thesouthwesternportionofSouth- The peripatricgeographicaldistribution of Habitat— and Distribution n and C. rostrata Souza 305 are from areas ofcalcareous outcrops couldreflect divergence bygeograph- , HUEFS). , Seasonally drytropical wood- C. rostrata zes Coursetia ually disarticulatesfrom occursmostly Coursetia caat- sect. Craccoides . Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 cence andfloralmorphologyof 305 as indicatedinthekey. pods of in axilsofinconspicuousornotfullyexpandedleaves.The Coursetiarostrata inflorescence rachis.Thependantinflorescence rachises of mature leavesthatare aslongorlongerthanthe adjacent and are bornalongnewshootsintheaxilsoffullyexpanded are markedlylessglandularthantheadjacentfloralpedicels The ascendinginflorescence rachisesof the muchlongerpendantinflorescence rachisesof and are ascendingtolaxinorientation,whichcontrastswith reminiscent of dense stipitateglandsalongthepedicel,calyx,andlegume In

estrada paraDelfino,660m,-10.80,-41.22, -41.86, W. R.Anderson36955 BRAZIL. Bahia:13kmSofCocos,3RioItaguari,520m,-14.34,-44.45, 1173 m, -12.96,-42.21, (HUEFS); ,12kmdeRio doPires paraNovoHorizonte, 600 MONT), 223/83 G.C.P. Pinto188/83 França 4718 -15.01, -42.88, 1 1 Sento Sé,MinasdoMimoso,840m,-10.31, -41.39, 540 m,-10.10,-40.23, 1 (HUEFS), 3.9 kmdeJussaraparaCentral,680m,-11.08, -42.00, 1 de SobradinhonaBA210,500m,-9.54,-40.92, (HUEFS, MONT), BRAZIL. Bahia:Morro do Chapéu,645m,-11.01, -41.41, 9156 Januária onroad toSerradas Araras, 625m,-15.52,-44.48, 1 1 Brejinho das Ametesitas, 770m,-14.26,-42.57, (leaf tissue).Caetité,Entre Brejinho das Ametistas eMorrinhosa8kmde m, -13.69,-44.25, are indicatedforcollectionsthatwere amplified for [Volume 36 ers amplifiedfrom from thisspecimen(HQ158101; Fig. 1 ) isidenticaltotheoth- the axilsoffullymature leaves. TheITSsequenceamplified short andascending,are born alongolderstemsandnotfrom SYSTEMATIC BOTANY flowers of predominantly lavenderpetals,incontrasttotheresupinate duces flowersthatare notresupinate atanthesisand thatbear mainly inreproductive morphology. narrowly ellipticleaflets.Theseperipatricsisterspeciesdiffer Both ofthesespeciesproduce paripinnateleaveswithmany bearing arostrate keelthatinturnhousesalongcurvedstyle. to 78 MONT), S SãoFélixdoCoribe,545m,-13.59,-44.32, Abaíra estrada Abaíra-Piatã, 800m, -13.27,-41.67, 520 m,-11.01, -41.43, 500 m,-9.60,-39.60, this newlyrecognized species. specimen couldrepresent anorthernphenotypicvariantof HQ158082; Abaíra, estradaCatolés-Abaíra,730 m,-13.27,-41.67, HQ158093; GentiodoOuro, 1,150m,-11.54, -42.54, HQ158092; 3kdeIacu,273 m,-12.76,-40.27, HQ158088; Morro doChapéu,lagedoBordado, 710m,-11.26, -41.08, HQ158101. MinasGerais.Espinosa,10km SemdiraMonte Azul, 600m, HQ158099. GentiodoOuro, 1,150m,-11.54,-42.54, The northernmostcollectionof Diagnosis— Coursetia rostrata C. caatingicola Other CaatingaCoursetiaSpecimensExamined— Examined— Specimens Additional (HUEFS),isdistinctiveincombiningthetypicalinflores- (HUEFS), (MEXU,MO,NY, VEN). (HUEFS), L. P. Queiroz 3995 1 Coursetia caatingicola 1 H186, HQ158067, H189, HQ158095, 1 HQ158090; Curaçá,FazentaSantaRita,490m,-9.67,-39.68, (HUEFS), C. rostrata V. C. Souza 5458 1 HQ158083; estrada entre JussiapeeBrejo deCima,620m, 1 C. rostrata F. Franca3855 D. S.Carneiro-Torres 118 Coursetiacaatingicola DNA 2121, theracemerachisesare lessthan10cmlong , incontrast,are bornalongolderstemsand (HUEFS);Curaçá,490m,-9.67,-39.68, 1 (F, MO,NY, US), H187, HQ158073, J. Costa407 2 E. B.Souza1525 E. deMelo3107 2 HQ158030; Jaugarari,40kmNSenhordoBomfim, 1 HQ158039, (petaltissue); especiallywithrespect tothelarge flower HQ158089; Morro doChapéu,Gruta dosBrejoes, Coursetia caatingicola thatbearpredominantly whitishpetals. (HUEFS), . Inaddition,theinflorescences, albeit 2 (HUEFS), HQ158034; 9 kmWdeIrecê, 770m,-11.30, (HUEFS), (HUEFS,MONT), 2 are shorterandwithfewerseeds, HQ158033; Uauá,sentidoBendengó, 1 HQ158091; 90kmWdeJacobinana (HUEFS), 1 (HUEFS), AF398858. 5kmdeDescoberto,652 Coursetia caatingicola 1 (Gnak ceso numbers accession (GenBank Coursetia caatingicola HQ158100. 13kmbyroad Wof (HUEFS), issimilarinmanyregards 1 H189, HQ158094, Coursetia caatingicola L. P. Queiroz 5117 L. P. Queiroz 12671 L. P. Queiroz 9171 V. C. Souza 5382 , whichsuggeststhis F. França2908 Coursetia caatingicola E. R.Souza127 1 1 E. R.Souza305 1 H187, HQ158070, H187, HQ158071, W. Ganev401 H189, HQ158096, 1 1 HQ158068; 9kmW HQ158081; 2km de 2 1 HQ158038. 27km ITS and Coursetiarostrata E. R.Souza305 T. S.Nunes949 D. Cardoso1865 W. R.Anderson G. C.P. Pinto C. rostrata 2 2 2 HQ158032; HQ158031; (HUEFS), HQ158040 (HUEFS), (HUEFS), (HUEFS), (HUEFS), (HUEFS), (HUEFS, (HUEFS, W. Ganev 2 trnD-T , with Souza pro- F. F. ). . .

MOL), et al.177 trecho /Itororó. Faz.BarradaNegra,330m,-15.21,-40.1, MONT), Aak, 94. A e lo a te ttsia mdl dniiain. identification . model statistical the at look new A 1974. H. Akaike, specimens ofspeciesrepresenting criticalSouth American outgroups. the manuscript.Toby Pennington andTiina Särkinenprovided herbarium anonymous reviewer provided manyhelpfulsuggestionsthatimproved Institute atMontanaStateUniversity. BenTorke, MarkSimmons,andan Funding fortheDNA sequencedatacame from theThermalBiology clade. DomingosCardoso provided photosof nosis toLatin.CarladeLimaillustratedthethree speciesoftheRostrata (FHO, MONT), -13.68, -40.77, et al.1845 Vitorino aVitoria daConquista,390m,-14.19,-40.27, Pé deSerra,440m,-13.53,-40.63, m, -14.18,-40.28, Carvalho 1845 Vitorino aVitoria daConquistanokm20,330m,-14.16,-40.25, 860m, -13.39,-42.31, (HUEFS), Rio deContas.BoaSentença,460m,-13.85,-41.53, (FHO, MONT), M.Elorsa-C.4626 MEXICO. Veracruz. (HUEFS), Abaíra. 12kmERiodeContas,610m,-13.41,-41.61, (HUEFS), estrada LivramentoparaBrumado, 490m,-13.79,-41.83, 1 1 -41.42, C.B.de 2 Cajamarca. caribaea Wied-Neuwied s.n. 2209 a Feirinha,220m,-15.25,-39.77, Harley 20004 E doRiodeContasonroad toMarcolino Moura,570 m,-13.59,-41.7, Contendas doSincorá,590m,-13.76,-40.27, -13.54, -40.62, 470m, -13.80,-41.78, -13.37, -41.60, (F), -45.46, MEXU, NY);5kmNTabocas, 10kmNWSerraDourada,700m,-14.61, by , 575m,-13.34,-44.64, (TEX), (FHO, MONT), (E), Pindai. Guirape,600m,-14.49,-42.69, et al.840 (CEPEC); Milagres. ItaberabaporIacu,520m,-12.87,-39.85, 10 kmWContendadoSincorá,730m,-13.7,-41.2, Feijão,740 m,-11.68, -41.46, , T. Jost487 (US). MinasGerais:Morro datorre 2kmdoEspinosa, 710m,-14.90,-42.82, HQ158080; , 270m,-10.06,-38.34, HQ158077; ManoelVitorino, 6kmdaFazendasReunidasdasContas,660 HQ158024. Acknowledgments. Poissoniaheterantha Poissoniahypoleuca Poissoniaeriantha Coursetiahassleri Coursetiamaraniona Coursetiacaribaea Examined— Species Coursetia Other Poissoniaweberbaueri Poissoniaorbicularis Coursetiagrandiflora 2 2 (FHO,MONT), HQ158042. HQ158063. Transactions onAutomaticControl 2 2 R. M.Harleyetal.21986 . PERU.Cajamarca. HQ158061. HQ158058. (CEPEC,TEX), (CEPEC);Prope Joazeiro, 250m,-12.32,-40.12, (CEPEC);LivramentodoBrumado, Fazenda Alagadico, 320m, (HUEFS), 1 1 A386, AF398860, 1 Santos 1589 1 HQ158078; , estradaBrumado-Ourives, 400m, -14.11, HQ158085; Abaira, 650m,-13.25,-41.67,B. H187, HQ158072, T. Särkinen2183 (CEPEC,NY);ItajúdoColonia,12kmdaestradaemdirecão (HUEFS), R. M.Harley53520 Oliveira Filho&Lima103 A. M.Giulietti1301 2 2 2 HQ158046. HQ158025. A. Bohrer 14 (HUEFS), M. daSilva-Castro 546 HQ158023; Cajamarca. 1817(BR). Chodat. ARGENTINA. Tucuman. 1 Hauman.PERU.Cuzco. (Jacq.)Lavinvar. 2 H188. HQ158086. T. R.S.Silva71 HQ158022. R. C.Duran5395 G. P. Lewis1863 (CEPEC);Brumado. BaizadesFlores, 2 (Speg.)Lillo. ARGENTINA. Salta. Hauman.PERU. Apurimac. (Grisb.)Lavin. Argentina. Jujuy. HQ158036; Brazil,Bahia,20kmSLivramento, Benth. exOerst.PERU.Cajamarca. (Harms)Lavin.PERU. Arequipa. 2 Lavin.PERU.Cajamarca. 1 HQ158035; .FazendaCurralVelho, 1 AF398849. InviaFelisbertiaadBarradaVareda, HQ158075; PédeSerra,1kmdacidade,500m, Cássio vandenBerg kindlytranslatedthediag- (FHO,MONT), ieaue Cited Literature 2 T. Särkinen2174 HQ158029. (HUEFS), G. Hatschbach39552 Santos 420 (HUEFS), (CEPEC,MEXU);,Rod.do Coursetia vicioides (HUEFS), M. daSilva-Castro 643 9 716–723. 19: (HUEFS), (MONT), R. M.Harleyetal.21661 Silva 220 (CEPEC);BanksofRioCorrente Coursetiacaribaea 1 (HUEFS), T. Särkinen2189 HQ158074; R116 trecho Manoel ochroleuca (HUEFS), (CEPEC);km6doRod.BR415, 1 2 HQ158076; 20kmdeJequiéa (FHO,MONT), HQ158026. L. P. Queiroz 2159 Gonçalves 236 1 (CEPEC);Livramentodo HQ158084; Brazil,Bahia, R. T. Pennington1801 1 HQ158087; 6kmWda 1 (CEPEC,MICH,NY); . BRAZIL.Bahia.13km A386, AF398861, (Jacq.)Lavin.PERU. Coursetia caatingicola R. T. Pennington1202 1 2 H187; Maracás, HQ158079; H182; Oaxaca. HQ158028; G. Hatschbach47874 A. M.Giulietti2433 T. Särkinen2182 L. P. Queiroz 7808 A. MdeCarvalho L. P. Queiroz 7068 Martius s.n. (Jacq.)Lavinvar. T. Särkinen2002 (FHO,MONT), T. Särkinen2035 Klitgaard 78 Hutchinson 7259 M. Lavin5787 (NY);Manoel Sobrinho 256 2 2 HQ158027. HQ158037; T. Särkinen G. P. Lewis (HUEFS), (HUEFS), (CEPEC, A. M.de R. M. IEEE (M); Silva (K, (E .

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Tue, 28 Sep 2021 12:21:26 Lvn M, . . crr , . ei , . . enntn A Delgado-Salinas , A. Pennington, T. R. Lewis, G. Schrire , D. B. M., Lavin, Lvn M, . . ocehwk , . asn C H Hge , n E Wheeler . E. and Hughes , H. C. Gasson, P. Wojciechowski , F. M. M., Lavin, sea- discontinuous of stability geographic and Floristic 2006. M. Lavin, of Systematics 1988. M. Lavin, Lri, . . G Bakhed , . . rw , . hna P A McGettigan, A. P. Chenna, R. Brown , P. N. Blackshields, G. A., M. Larkin, Lnly C H ad . ic 94. A etmto o te osac o the of constancy the of estimation An 1974. Fitch. W. and H. C. Langley, likeli- maximum the of Evaluation 1989. Hasegawa. M. and nuclear H. of Kishino, survey A 2006. Hodges. A. S. and Whittall, B. J. M., of K. inference Kay, Bayesian MrBayes: 2001. Ronquist. F. and P. J. Huelsenbeck, 2001. P . S. Hubbell, for system information geographic A 7.0. DIVA-GIS 2009. J. R. Hijmans, 1991. Pagel. D. M. and H. P. Harvey, algorithm accurate and fast, simple, A 2003. Gascuel. O. and S. Guindon, Flesen J. 18 ofdne iis n hlgne: n prah using approach an phylogenies: on limits Confidence 1985. J. Felsenstein, Dn d Seao R, . . enne-oca L L CnIz , and Can-Itza, L. L. Fernández-Concha, C. G. R., Stefano, de Duno of distribution the on perspective phylogenetic A 2008. J. M. Donoghue, Lvn M, . eede , n M F Wjicosi . 20 Evolutionary 2005. Wojciechowski . F. M. and Herendeen , P. M., Lavin, d Qerz, K d 20 pce cnet ad pce dlmtto delimitation. species and concepts Species 2007. de K. Queiroz , de for framework unified a towards entities: Evolving 2010. G. T. Barraclough, generic a – earring monkey’s Guanacaste, Silktree, 1998. C. R. Barneby, 01 QERZADLVN RZLA OREI 79 QUEIROZ AND LAVIN: BRAZILIANCOURSETIA S. C. Wojciechowski , F. M. Porter , M. J. Sanderson, J. M. G., B. Baldwin, Characteriza- 2003. Hughes. E. C. and Harris, A. S. Carr , G. T. D., C. Bailey, 2011] legumes. Wojciechowski . F. history betterexplaingeographicallystructured M. phylogeniesin and Beyra-Matos, A. Hughes, E. 2004 C. Thulin, . M. Metacommunity processes ratherthancontinentaltectonic Caribbean endemics . and 2003 . Phylogeny ofrobinioid legumes(Fabaceae)revisited: Press . CRC Florida : tion sonally dryforests: plantbiodiversity, biogeographic patternsandconserva- endemism, Chapter19 . Pp. 433 – sonally drytropical forests explainspatternsofplantphylogenyand 447 in SystematicBotanyMonographs Topo , . . isn ad . . ign 07. Cutl and W Clustal 2007. D. Higgins. J. G. Lopez, D. R. and Wilm , Gibson, A. Clustal Xversion2.0 Wallace. , J. M. T. I. Thompson, Valentin , F. McWilliam , H. rate ofmolecularevolution . Evolution data, andthebranchingorder inHominoidea . hood estimateoftheevolutionarytree topologiesfrom DNA sequence BMCEvolutionaryBiology sperms: anapproximate molecularclockwithlifehistoryeffects . ribosomal internaltranscribedspacersubstitutionratesacross angio- phylogenetic trees . Press. University Princeton Princeton: diva-gis.org . the analysisofspeciesdistributiondata . Available at http://www. biology Biology to estimatelarge phylogeniesbymaximumlikelihood . the bootstrap . ain of cations . distinctions phylogenetic and morphological Coursetiagreenmanii The 2010. Lavin. M. 11549 –11555 . 105: plant diversity . lineages duringtheTertiary . rates analysisofLeguminosaeimplicates arapiddiversificationof 1509–1522. 359: tematic Biology Transactions oftheRoyalSociety, B understanding diversityatthespeciesandhigherlevels . 1–223. Calliandra system forthesynandrous Mimosaceaeofthe Americas: partIII, Cmbl , n M J Dngu 95. Te T rgo o ncer ribo- nuclear eny . of region ITS The 1995. somal DNA:avaluablesource ofevidenceonangiospermphylog- Donoghue. J. M. and Campbell, genes . tion ofangiospermnrDNA polymorphism,paralogyandpseudo- , d. R T Pnigo , . . atr ad . . ei oa Raton, Boca Lewis . P. G. and Ratter , A. J. Pennington, T. R. eds. , Gliricidia Annals oftheMissouriBotanicalGarden . Ofr xod nvriy Press. University Oxford Oxford : . Molecular PhylogeneticsandEvolution 2 696–704. 52: Systematic Botany 9 170–179. 29: Philosophical Transactions oftheRoyalSociety . Memoirs oftheNewYork BotanicalGarden recircumscribed, andabiogeographicalappraisal ofthe 6 879–886. 56: The unifiedneutraltheoryofbiodiversityandbiogeography Evolution Proceedings oftheNationalAcademySciencesUSA Bioinformatics (Leguminosae):taxonomicandecologicalimpli- Systematic Botany Bioinformatics 8 783–791. 38: : 36. 6: 5 289–295. 35: Coursetia Journal ofMolecularEvolution Systematic Biology 1 1–167. 21: The comparativemethodinevolutionary , 35 1801–1813. 365: , 7 754–755. 17: 3 2947–2948. 23: (Leguminosae-Papilionoideae). 8 387–409. 28: Neotropical savannasandsea- 2 247–277. 82: 9 435–455. 29: 4 530–549. 54: Journal ofMolecular . Biological Series 4 pr III): 74(part Philosophical : 161–177. 3: Systematic Coursetia Sys- .

Smos M P. 20 needne f lgmn ad re erh. search . tree and alignment of Independence 2004. P . M. Simmons, C. Siripun, C. K. Miller , J. Liu, W. Farmer , S. Beck, J. Lickey , E. J., Shaw, patterns distribution Global 2005 . Lewis. P. G. and Lavin, M. D., B. Schrire, (December Manual User’s (beta), 1.71 version r8s, 2004. J. M. Sanderson, Tmltn A R. 18 ovret vlto ad o-aaerc infer- non-parametric and evolution parsimony Convergent using 1983 . R. analysis A. Phylogenetic Templeton, PAUP*. 2002. L. D. Swofford, evolution molecular of rates absolute Estimating 2002. J. M. diver- Sanderson, estimating to approach nonparametric A 1997. J. M. Sanderson, Psd, 08. joeTs: hlgntc oe aeaig. averaging. model phylogenetic jModelTest: 2008. D. Posada, Rnus, . n J P Hesnek . 20 rae 3 Bysa phyloge- Bayesian 3: MrBayes 2003. Huelsenbeck. P. J. and F. Ronquist, 2009. de P . L. Queiroz, patterns phytogeographical caatinga: Brazilian The 2006. de P. L. Queiroz, the in distributions species of Patterns 1993. Gibbs. E. P. and E. D. Prado, Pnigo, . . J E Rcado , n M Lvn . 20 nihs no the into Insights 2006. Lavin. M. and Richardson , E. J. T., R. Pennington, RlyHlig E T, . egd-aia , n M Lvn . 20 Phylogenetic 2004. Lavin. M. and Delgado-Salinas, A. T., E. Riley-Hulting, Pnigo, . . D E Pao ad . . edy . 20 etoia sea- Neotropical 2000. Pendry . A. C. and Prado, E. D. T., R. Pennington, Pnigo, . . M Lvn T Srie , . . ei , . . ltar , and Klitgaard , B. B. Lewis, P. G. Särkinen, T. Lavin, M. T., R. Pennington, Pnigo, . . M Lvn D E Pao C A Pnr , . el and Pell, S. Pendry , A. C. Prado, E. D. Lavin, M. T., R. Pennington, plant Woody 2009. Oliveira-Filho. A. and Lavin, M. T., R. Pennington, Zn, . . n G F Braluh . 20 iohnra DA ne siege under DNA Mitochondrial 2008. Barraclough. F. G. and M. R. Zink, Nxn K C ad . . hee 90. A apiiain f h phyloge- the of amplification An 1990. Wheeler . D. Q. and C. K. Nixon, Yn, . n B Rnaa . 19 aein hlgntc neec using inference phylogenetic Bayesian 1997. Rannala. B. and Z. Yang, herbaria public of directory global A Herbariorum: Index 2010. B. Thiers, Wne , . . ciln , n R L Sal . 20 h tros ad the and tortoise The 2005. Small. for phylogeneticanalysis . L. R. and hare II:relative utilityof21non-codingchloroplast DNA sequences Schilling, E. E. Winder , and globaldimensions.BiologiskeSkrifter (eds.), Balslev H. of theLeguminosae:insightsfrom recent phylogenies . In I. California. of University Friis Davis: & 2004). Distributedbytheauthor( http://ginger.ucdavis.edu/r8s/) . Mre Dekker . Marcel in 179 ences from restriction fragmentandDNA sequencedata . Pp. Associates. 151 Sinauer – Sunderland: 4. Version Methods). Other (*and Phylogenetics andEvolution Biology andEvolution and divergence times:apenalizedlikelihoodapproach . Evolution gence timesintheabsenceofrateconstancy . netic inference undermixedmodels . 627–653. Press . CRC Florida: Raton, Boca Ratter . raphy, andconservation Neotropical savannasandseasonallydryforests: plant diversity, biogeog- inferred from distributiondataoftheLeguminosae . Pp. 121 – 157 in Garden dry seasonalforests ofSouth America . tutr structure . logenies, neutralecologicaltheoryandphylogeneticcommunity historical construction ofspecies-richbiomesfrom datedplantphy- ate genuswithinaneotropical diversification . systematics of Universidade EstadualdeFeiraSantana. Biology andEvolution Biogeography sonally dryforests andQuaternaryvegetationchanges . of SciencesUSA within histories diversification the plant Andean biodiversityhotspot Contrasting . 2010. Hughes. E. C. Transactions show oftheRoyalSociety plants forest dry patterns ofbothTertiary seasonally andQuaternarydiversification. Neotropical 2004. Butterworth. C. Systematics sonally drytropical forests . diversity, evolutionandecologyinthetropics: perspectivesfrom sea- in avianphylogeography . netic speciesconcept . Biology andEvolution DNA sequences: aMarkovChainMonteCarlomethod . http://sweetgum.nybg.org/ih/ . and associatedstaff. New York BotanicalGarden’s Virtual Herbarium . Statistical analysisofDNA sequencedata 0 902–927. 80: 4 1218–1231. 14: The NewPhytologist 0 437–457. 40: 7 261–273. 27: 0: 13783–13787. 107: Strophostyles Plant diversityandcomplexitypatterns:local,regional 9 101–109. 19: 5 1253–1256. 25: 4 717–724. 14: Cladistics , d. R T Pnigo , . . ei , n J A. J. and Lewis, P. G. Pennington, T. R. eds. , Leguminosas dacaatinga Molecular Ecology 1 874–879. 31: American JournalofBotany (Fabaceae):aNorth American temper- Annual ReviewofEcologyEvolutionand 7: 605–616. 172: . : 211 –223. 6: Biological Sciences Proceedings oftheNationalAcademy Bioinformatics 5 375–422. 55: Annals oftheMissouriBotanical : 2107–2121. 9: , d . er . Nw York : New Weir . B. ed. , Molecular Biologyand . Feira deSantana : Systematic Botany 5: 515–538. 359: 9 1572–1574. 19: 2 142–166. 92: Philosophical Journal of Molecular Molecular Molecular Molecular 29: