FLAVONOIDS IN STROPHOSTYLES SPECIES AND THE RELATED GENUS DOLICHOPSIS (PHASEOLINAE, FABACEAE): DISTRIBUTION AND PHYLOGENETIC SIGNIFICANCE

10 Pages·1998·4 MB·English

by  Juan Pablo Pelotto

FLAVONOIDSINSTROPHOSTYLESSPECIES
ANDTHE RELATEDGENUSDOLICHOPSIS
(PHASEOLINAE, FABACEAE):
DISTRIBUTIONAND PHYLOGENETIC
SIGNIFICANCE
JUANPABLOPELOTTOandMARIAA.DELPEROMARTINEZ
CentrodeEstudiosFarmacoioguvsyBotdnkos
ConsejoNacionaldeInvestigacionesCkntifkasyTecnkas
Serrano663,BuenosAires, 1414.ARGENTINA
ABSTRACT
StrophostylesElliot:istheonlygenuswithinthePhaseolinaewithacenterofdistribu-
tionintheUnitedStates.Itcomprisesthreespecies,namelyS.hdvula(L.)Elliott,5.umbellata
(Willd.)Britton,and5,leiosperma(Torrey&A.Gray)Piper,anditisconsideredasalliedto
DolichopsisHassler,amonotypicgenusendemictoSouthAmerica.Thisstudyanalysesthe
wleiatfhfltahveonaoiimdocfonetxeanmtinfrionmgtthheetphhryeleoSgsernopehtoisctylreeslastpiecoinesshiapnsdaDmoolincghotpauxsa.[lWareaguiasroileantseidsH3a8ssglleyr-
cosidesbasedonkaempferol,quercetinandisorhamnetin.AllStrophostylesspecieswerecharacterized
bythepresenceofisorhamnetinglycosides.However,S.leiospermashowedadistinctive
profilewhileS.hdvulaandS.umbellataclusteredtogether.Incontrast,D,paraguariensis
lackedisorhamnetin-basedcompounds.Acladisticanalysisofflavonoidplusmorphologi-
caldatasupportedStrophostylesmonophylyandshowed5".leiospermaasthesistertaxonof
thecladeS.helvula-S.umbellata.
RESUMEN
StrophostylesElliottesellinicogenerodelasPhaseolinaeconuncentrodedistnbucion
enlosEstadosUnidos.Comprendetresespecies:S.helvula(L.)Elliott,S.umbellata(Willd.)
BrittonyS.leiosperma(Torrey&A.Gray)Piper,yseconsideraafinaDolichopsisHassler,un
generomonotfpicoendemicodeSudamerica.EsteestudioanalizaelcontenidodeOavonoides
foliatesenlastresespeciesdeStrophostylesyenDolichopsisparaguariensisHasslerconelobjeto
deexaminarlasrelacionesfilogeneticasentreestostaxa.Seaislaron38glicosidosdekaempferol,
quercetinaeisoramnetina.TodaslasespeciesdeStrophostylessecaractenzaronporlapresencia
deglicosidosdeisoramnetina.Sinembargo,S.leiospermamostrounperfildistintivomientras
queS.helvulaandS.umbellataseagruparonjuntas.Porelcontrario,D.paraguariensisno
sintetizocompuestosbasadosenlaisoramnetina.Unanalisiscladisticoconjuntodelosdatos
deflavonoidesycaracteresmorfologicosapoyolamonofiliadelgeneroymostroa^'.leiosperma
comoeltaxonhermanodelcladoS.helvula-S.umbellata.
INTRODUCTION
StrophostylesEUiorristheonlygenuswithinthePhaseolinaewithacen-
terofdistributionintheUnitedStates.Itscurrenttaxonomictreatment
followstheoriginalgenericconcept(Elliott 1822),butitwaspreviously
SiDA18(1):213-222. 1998
.
214 SiOA 1<S(1)
associatedwithunrelatedspeciesandreducedtoasectiono^Phaseolns(de
CandoUe 1825; Bentham 1837, 1865) until itwasrestoredasaseparate
genusincludingthreespecies(Britton&Brown 1897;Piper1926).Aset
ofcharactersprecludes mergingStrophostyleswithPhaseoh/s, namelyerect
style(notcoiled),lackofhookedhairs,pedicelsshorterthanthecalyx,and
nodesoftheinflorescencesomewhatswollen(Marechaletal. 1978).More-
over,Strophostylesplantscanberecognizedbytheirnearlyasymmetricflowers
arrangedinsubumbellate inflorescences, bractsand bracteolespersisting
throughseedmaturation,cylindricalseeds(oftenpubescent),andlinearpods.
StrophostyleshasalsobeenconsideredasalliedtoDolichopsisHassler(Marechal
etal. 1978;Lackey1983).Nevertheless,bothgeneraareeasilydistinguishable
bymanycharactersandtheirquitedistinctgeographicaldisrributions.The
monotypicgenusDolichopsis^ resemblesStrophostylesmainlyinfloralmor-
phology(purplishcorolla,keelwithanotcurvedtosomewhatcurvedbeak,
styleslightlythickeneddistally,andstigmaterminaloblique)andgeneral
appearance,butitisunlikethelatterinhavingsymmetricflowersclustered
inelongatepseudoracemesandtheuniquefruittraitssuchasoblong,very
flatpodswithoblongseedsimplantedthroughaverylongfunicleandwiththe
hilumperpendictilartotheplacenta.ThegeographicrangeoiDolichopsisisParaguay
andArgentina,inSouthAmerica,whWtStrophostylesoccursthroughouteastern
USA,easternCanadauptosouthofQuebecandextremenortheasternMexico.
PhytochemicaldataonStrophostylesspeciesarescantyandincludetheabsence
ofbothleuco-anthocyanins(Baudet 1978)andcanavanine(Lackey 1977),
andarecentreportofflavonoids(Williamsetal. 1995).Inthisstudy,we
expandedonthesurveyoffoliarflavonoidsbyconsideringalargernumber
ofsamplesofthethreeStrophostylesspecies,i.e.S.helviila,S.mnbellataandS.
leiospermci,andweaddedDolichopsisparaguariensisforcomparison.
MATERIALSANDMliTHODS
Weanalyzedtheconstitutiveflavonoidspresentintheleavesofherbarium
specimensbelongingtothethreeStrophostylesspeciesandDolichopsisparaguariensis
Samples(100-200mg)werepowderedandextractedunderrefluxwith80%
methanol(x3).Concentratedmethanoiicextractsweretwo-dimensionally
chromatographedonpaper(BAW/15%aceticacid).Compoundswereidentified
bystandardmethods(Mabryetal. 1970;Markham 1982).Theseincltided
completeandcontrolled(3min.)acidhydrolysis,enzymatichydrolysis(B-
glucosidase),co-chromatographywithauthenticmarkersandUV-Visspec-
^Doltchupmwasagenuswithtwospecies,D.paraguarienmandD,monticola(Lackey1983,
Lewis1991),butrecentlyDelgadoSalinas&Lewis(1997)createdthenewgenusOryx'n
wheretheyplacedD.monttadci.Tiierefore,D.paniy^/uinensishasbecamethetiniquerepre-
sentativeofthegenus.
1
PelottoandMartinez,FlavonoidsinScrophoscylesandDolichopsisspecies 215
troscopy. Glucosideswereseparated from theirgalactosidicanaloguesby
TLCintheappro—priatesystemaccordingtoBudzianowski(1991).
Plantmaterial. SpecimenswereprovidedbytheInstitutedeBotanica
DarwinionHerbarium(SI),SanIsidro,andtheCentrodeEstudiosFarmacologicos
yBotanicosHerbarium(BACP),BuenosAires.
Strophostyleshelvula(L.)Elliott
U.S.A.Arkansas.JeffersonCo.:Arkansasriverbottoms,220ft,17Sep1937,Demaree
2126323455((SSII))..IIlolwiano.isD.iMckcinDsoonnoCuog.:hNCos.h:oArergoyfkSlpierLiatkLea,ken,easraCnodl(cohledsetrerb,ea2chA)u,g5\A9u5gH,1J9o1n3e,s
Si>/ek14...(numberillegible)(SI).Mississippi.HarrisonCo.:nearthecoastonsand,6
Jan\9'5UDewaree30675(SI);ShipIsland,P.O.Biloxi,instabilizedsand,moist,longtrailing,
Demaree31059(SI).Virginia.PrinceGeorgeCo.:richalluvialthicketbackofsand-beach
ofJamesRiver,JordanPoint,SEVirginia, 16Sep 1938,Feriialil&Long9353(SI).Un-
hwuHloadity.movdoftheWesternReserve]15Atig1897,G.B.Asduroft.Berea0.s.ii.(SI).
Strophostylesleiosperma(Torrey&A.Gray)Piper
U.S.A.Oklahoma:5miNWofBreckenridge,25Jul1941,Gephardt747(SI).Woods
Co.:inwasteplace,hardsoil,nearAlva,24Sep\9U,Stevens2824(SI).Texas.SmithCo.:
Amigo,neglectedsandyfield,10-17Aug194'5,Moore.Jr.995(SI).MorrisCo.:Aug1891,
Carleton420(SI).
Strophostylesumbellata(Willd.)Bntton
U.S.A.Virginia.GreensvilleCo.:drypineandoakwoods,about1miNofSkipper's,
l4-\5]n[1938,Fernalct&Long8737iSl).
DolichopsisparaguariensisHassler
JPatneA.1RH9Ga8yE1eN,sfT:l.IEaNzsutAla-.nvciEoinlataLcroeeom,aRPiToryostn:ac,oDseo2p3dt°eoB.4i0/L'raSk,aPr5atz9,&°R3B5a1a'2gW6a,,l2udpeAsopvi3r'0o19a967O4(mS,bI)te.inPerseA,dRabAdoeGrrUdaeAcaYcsa.imriDanseotp,rtero3a.,
flviolaceas,creceenpastizal,n.v.'kekleichetas,'Arenas544(BACP).Depto.Boqueron:
MisionSantaRosa,21°45'S,61°35'W,Feb1981,enredadera,fl.violaceas,creceenpajonal,
n.v,'ceihlowey'.Aren—as1726(BACP).
Dataanalysis. Aclusteranalysiswasperformed onflavonoiddataof
15 herbariumspecimens.SimilaritymatrixwasmeasuredusingJaccard's
coefficientandadendrogramwasconstructedapplyingtheunweigthedpair-
groupmethodofarithmeticaverages(UPGMA).Allcalculationsweredone
usingNT-SYSprogram(Rohlf1993).
Acladisticanalysisoftheflavonoiddataplusasetofmorphologicalcharacters
wascarriedoutaccordingtothemaximumparsimonyprinciple.Outgroup
criterionwasusedforcharacterpolarization.Vignaadenanthawaschosenas
theexternalgroupwhichflavonoiddatawereobtainedfollowingthemethods
abovementioned(Pelotto,unpublishedmanuscript).Forflavonoids,char-
acterstatesthatoccurredintheoutgroupwerescoredas andthoseinthe
ingroup(D.paraguariensisandthethreeStrophostylesspecies)werescoredas
1(seeAppendix,TablesAandB).Morphologicaldataweregatheredfrom
theliteratureandincludedsomemultistatecharactersthatweretreatedas
non-additive(seeAppendix,TablesAandC).Cladogramswerecalculated
-^^ SiDA18(1)
usingtheimplicitenunierationroutine(i.e.*)oftheprogramHennig86
(Farris 1988)withallcharactersequallyweighted.
RESULTSANODISCUSSION
Chromatographicpropertiesoftheidentifiedflavonoidglycosidesand
Itsdistribution inStrophostylesspeciesandD.paragiiarienmareshown in
Tables Iand2,respectively.AlldetectedcompoundswereO-glycosidesof
flavonolswithsugarsattachedatpositions3and7oftheagliconeskeleton.
ThispatternofglycosilationisverycommonamongthePhaseolinae(Zallocchi
&Pomilio1994;Williamsetal. 1995,Pelottounpublishedmanuscript).
AllthreeStrophostylesspeciesproducedglycosidesbasedonthemethy-
latedflavonolisorhamnetinpluskaempferolandquercetin.Notwithstanding
S. helvnlaandS. nmhellatashowedverysimilarchromatographicpatterns,
whiletheflavonoidprofileof5.lewspmnawasquitedistinctive.Norhamnosides
weredetectedinS.kmpmnaanditonlysharedthepresenceofmonoglycosides
withtheothertwospecies.Basedonathree-sampleanalysiswithinafla-
vonoidsurveyofthePhaseolinae,Williamset.al(1995)havealsoreported
theoccurrenceofisorhamnetinglycosides inStrophostylesspeciesandno-
ticedthesameinterspecificdifferences.Incontrastwithourresults,Will-
iamsandco-workersisolatedfewercompoundsanddidnotdetectkaempferol
glycosidesfromleaves,althoughtheydidfromstemsand/orflowers.
Inturn,Dolichopsisparaguariensissampleswerecharacterizedbythepresence
ofkaempferolandquercetinglycosides,lackingisorhamnetin.Noticeably,
ParaguayansamplescontainedonlykaempferolglycosideswhileArgentinean
onehadkaempferolplusquercetinglycosides.However,inapreviouswork
(Zallocchietal. 1995)bothkaempferolandquercetinglycosideswerere-
portedfromonesampleofD.pcircigiiarmisisfromParaguay,butoftheeight
flavonolglycosidestheauthorsidentifiedonlyrutinandkaempferol-3-O-
rutinosidewerealsopresentinour.samples.Thesedifferencesmaybedue
tothefact thatZallocchi and co-workersanalyzed awholeplantextract
andthereforetheirresultsaredifficulttocomparewithours.
AftertheclusteranalysisS.helvnlaandS.umhellataareclosertoD.paraguariensis
thantoS.lewsperma(Fig. 1).ThisisbecauseS.helvulaandS.umhellatahave
moreglycosides(basedonkaempferolandquercetin)incommonwithD.
paraguariensisthanwithS.leiosperma,eventhoughD.paraguariensisdoesnot
produceisorhamnetinglycosides.
Cladisticanalysisresultedintwomostparsimonioustrees(length,L-
35,consistencyindex,CI =91,retentionindex,RI=70.Fig.2).Bothcla-
dogramssupportStrophostylesmonophyly butdiffer in thedepicted rela-
tionshipsamongStrophostylesspecies.Onetree(Fig.2.A)showsS.helvula
andS.leiospermaasbeingsiblingspecies,butthishypothesisneedsthepar-
allelgainofthecharacters19,22and31ontheS.umhellataandS.helvula
branches.Theothertree(Fig.2.B)supportsthecladeS.umhellata-S.helvula
1
PelottoandMARTfNEZ,FkvonoiclsinStrophostylesandDolichopsisspecies 217
Table1.Chromatographiccharacteristicsoftheidentifiedcompounds.
COLOUR" Rf(x100)
SPOT IDENTITY UV +NH3 BAW 15%aa
1 K-3-O-glLicoside+K-3-O-gahictoside DP Y 69 45
2 K-7-O-glucoside+K-7-O-galactoside Y Y 45 15
3 K-3-O-rutinoside+K-3-O-robinobioside DP Y 52 54
4 K-3-O-diglticoside+K-3-O-digalactosidc DP Y 33.5 61.5
5 K-3,7-0-ch^ducoside DP Y 29 70
6 K-3-0-rLitinoside-7-0-glucoside+
K-3-0-robinobioside-7-0-glucoside DP Y 24 75
7 K-3,7-0-tnglucoside DP Y 3 82
8 K-3,7-0-triglycoside(glu+rha+gal):|: DP Y 4 80
9 Q-3-O-glucoside+Q-3-O-galactoside DP Y 56 39
10 Q-7-O-glucoside+Q-7-O-galactoside Y YO 27 9
1 Q-3-O-rutinoside+Q-3-0-r()binobioside DP Y 40 51
12 Q-3-O-digliicoside+Q-3-O-digalactoside DP Y 26 52
13 Q-3,7-0-diglucoside DP Y 23 63
14 Q-3-0-rutinoside-7-0-glucoside+
Q-3-0-robinobioside-7-0-giLicoside DP Y 15 70
15 Q-3,7-0-triglucoside DP Y 3 78
16 Q-3,7-0-triglycoside(gki+rha+gal) DP Yo 5 79
17 IR-3-O-glucoside+IR-3-O-gaiactoside DP Y 56 42
18 IR-7-O-glucoside+IR-7-O-gahictoside Y Y 38 10
19 IR-3-O-riitinoside+IR-3-O-robinobioside DP Y 40 54
20 IR-3-O-diglucoside+IR-3-O-digalactoside DP Y 29 58
21 IR-3,7-0-digIucoside DP Y 25 68
22 lR-3-0-rutinoside-7-0-glucoside+
IR-3-0-robmobioside-7-0-glucoside DP Y 19 74
23 IR-3,7-0-triglucoside DP Y 3 80
'DP:deeppurple,Y:yellow,YO;yellow-orange
^glu:glucose,rha:rhamnose,gal:galactose
andrequiresthreereversions(characters1,9and17).Thisscenarioispref-
erabletothatportrayedonFig. 2.Asinceamutationlostisamoreprob-
ableeventthanthehomoplasticacquisitionofisorhamnetmglycosides.Even
more, ifwesupposereversalofcharacters 1,9and 17ontheS. mnbellata
branchasbeingaconsequenceofsamplmgerror(undersampling),thecla-
dogrambecomesshorterwithonly32steps(CI= 100,RI= 100)andthe
uniquesolutionofasimilaranalysis.Thus,weconsiderthetreedepicted
onFigure2.Bamoreplausibleingroupphylogeny.
Flavonoidevolutionshowsmethylationoftheflavonolskeletonasanadvanced
charactersharedbyallStrophostylesspeciesandtheabsenceofrhamnosides
inS. leiospermaasan(aut)apomorphicloss.
Morphologicaltraitsarecongruentwithflavonoiddata. Subumbellate
inflorescence(character24),persistentbractsandbracteoles(character26),
linear,teretepods(character27)andseedpubescence(character28)^\vp-
^otxStrophostylesmonophyly,andarecorrelatedwithisorhamnetinmonoglycoside
production(characters 17and 18).Strophostyleshelvulastronglyresembles
—
00
Table2.Glycosidedisrriburionintlieanalyzedsamples.CompoundsarenumberedaccordingtoTable1.K;kaempterolglycosides;Q:cjuercetinglycosides;IR:
isorhamnetinglycosides;-r:present;-:absent.
K Q IR
10 11 \2 \i 14 15 16 r 18 19 20 21 22 2t
S.helvitla -- --- -- --- --
Jones22335 -+++ + -- + -- + + + + + + -
Simek14... ++ + -- + -- + + + -- -- + - + + + + -- -- + -
ADsecmharroefte&30B6e7r5eas.n. -+++-++ ---- ++ _----++ ++ ++--- -- ++ -- ++ ++ ++ ++--- -- ++ -
DDeemmaarreeee3116204595 ++ + -.-_ ++ -----+ ++ + ---- ++ ^- ++ +- ++ + ---- ++ --
Fernald&Long9353 - + + + + + + - + - + + + -
S.umbellata -- --- -- --
Fernald&Long8737 -+++ -+- +--- + + +--- -+ - + +--^-+--
SMGC.aeorpollehreiaeotrioJpdnret.r4m972a94057 +++_--_-- ++ +++ -- ++ -- +++ +--------++ +++ -- ++ ---++i-+------ ++ +^-- ++
Stevens2824 + + - + - + + + - + - + + + - +
__-_____---------
D.paraguariemii __ -......-._. ------
AArreennaass5147426 — ___^+ ++ -_-. ++ -- -------
Troncoso&Bacigalupo3096 - + + + - + - + + + - + (y^
>
00
PelottoandMartinez,FlavonoidsinStrophostylesandDolichopsisspecies 219
JACCARD'SCOEFFICIENT
0.00 0.25 0.50 0.75 1.00
—
HEL-22335
_jHEL-9353
'UMB-8737
HEL-14...
_HEL-sn
HEL-30675
—HEL-31059
—HEL-16245
DOL-3096
_|DOL-1726
—'DOL-544
LEI-995
LEl-747
-HILEI-2824
IlEI-42()
Fk;. 1.DendrogramoftheStrophostyltsandDalichopsiispecimensconstructedfromasimi-
laritymatrix(Jaccard'scoefficient)usingtiieUPGMAmethod.Copheneticcorrelationco-
efficient,r=0.969
S. umbellata,exceptforits morefobed leafletsand largerpodsand seeds.
Strophostylesleiospermaisratherdifferentfromtheothertwospeciesbecause
ofthesmallerflowers(character25)arranged inmorepauciflorousinflo-
rescencesanditsseedsglabrousandshiningatmaturity(character29).
Similarly,bothS.helvnlaandS.umbellataaremorewidespreadandnortherly
distributed,withthefirstspeciesreachingCanada,whileS.leiospermahasa
morelimiteddistributionrangingfromsouthoftheUnitedStatestothe
extremenortheasternofMexico(Britton&Brown 1897, Marechaletal.
1978).Speciesdivergenceatchemicalandmorphologicallevelalsocorrelates
with theirecologicalfeatures;S. helvulaandS. umbellatamostlygrowin
moremesicsites,whileS.leiospermaisadaptedtoliveintomorexerichabitats.
Regardingtheevolutionofthegrowthform,overlappingthischaracter
ontoourpreferredtopologysuggeststhatannualgrowthwouldhaveevolved
independentlyinbothS.leiospermaand.S".helvula,whereasperennationwould
betheplesiomorphicstatesharedbyD.paraguariensisandS. umbellata.
Insummary, universaloccurrenceofisorhamnetin-basedcompounds in
Strophostylesspeciesisagoodchemicalcharacterindefininggenericmonophyly
whileindividualglycosidesareusefulcharacterstotracespeciesevolution.With-
inthePhaseolinaeisorhamnetinglycosideshavesporadicallybeenrecorded
infourPhaseolusspecies(Pelotto,unpublishedmanuscript)andsomeVigna
andMacroptiliumspecies(Zallocchi&Pomilio1994;Williamsetal. 1995).
Thisfactsuggests thatflavonolmethylationhasappearedseveraltimesin
thetribe,makingitavaluablephylogeneticmarkerattheinfragenericlevel.
Beyondthiscontribution,flavonoiddatafromtherelatedgeneraOxyrhynchus
andOryxisarewantingforacompleteviewofthislittlegroupofAmerican
speciesaroundVigna.
220 SlDA18(1)
A-
t
/3-7
-11-15
20,21,23
25(2),29
^
18,24,26,27(2},28
^
2,10,25(1)
B-
7,]8.24.26,27(2),28
13
Fic.2.Thetwomostparsimoniousrrees(L^ 35)^'enerateclusingthedatamatrix(see
Appendix,TableA)andVigthiiiJenunthaasoutgroup.CharactersaremajDpedonthetrees
asFollows:solidbar- non-homoplasiotisapomorphy,clearbar=homoplasiousapomorphy,
andcross=reversal.Numbersontherightofthecharactersymbolsstandforcharacter
numbers(andcharacterstate).
Pelottc)andMartinez,FlavonoidsinStrophostylesandDolichopsisspecies 221
ACKNOWLEDGMENTS
WethankthecuratorsoftheInstitutedeBotanicaDarwinionHerbarium
(SI)andtheCentrodeEstudiosFarmacologicosyBotanicosHerbarium(BACP)
forprovidingtheplantmaterial,andCONICETforfinancialsupport.We
arealsogratefultotwoanonymousreviewersforsuggestionsthatimproved
theearliermanuscript.
APPENDIXES
TableA.Datamatrixforthecladisticanalysisincludingbotiiflavonoid(characters1-15,17-23,
codifiedaccordingtoTableB)andmorphological(characters24-31,codifiedaccordingtoTableC)
datasetsandusingS/igruiddencinthaasoutgroup.
characternumber
Taxon 1 6 8 9 1(111 1213li15n18192021222i242526r28293031
y'lgnaadmanlki (1 >
D.para^^uarmiili 1 1 1 1) 1 1 :' 1
110 10 110 10 12 10
5S..iheilmvhieilUaata 10 1 10 1 110 10 111I12 10 11
.V.lampnma 1 1 1 1 I 1 I 1 I 1 1 1 1 1 1 1 1 I I 1 2 1 2 I 1
TableB.Flavonoids.CharacternumbersareasinTable1.Character16wasnotincludedbecauseno
hypothesisabouthomologycanbemadeonapartiallyidentifiedcompound.
Characterstates
1.0=absent,1=present 8.0=absent,1=present 15.0=absent,1=presenr
2.0=absent,1==present 9.0=absent,1=present 17.0=absent,1=present
3.0=present,1=absent 10.0=absent,1=present 18.0=absent,1=present
4.0=absent,1=present 11.0=present,1=absent 19-0=absent,1=present
5.0=absent,1=present 12.0=absent,1=present 20.0=absent,1=present
6.0=present,1=absent 13.0=absent,1=present 21.0=absent,1=present
7.0=absent,1=present 14.0=present,1=absent 22.0=absent,1=present
23.0=absent,1=present
TableC.Morphologicalcharacters,statesandpolarities.
character
24.inflorescence 0=pseudoracemose;1=subumbellate
25.flowersize 0=great(>20mm);1=medium(7-15mm);2=small(<7mm)
26.bractandbracteole 0=persistingnolongeranthesis;1=persistingthroughseedmaturation
27.pods 0=linear,compressed;1=oblong,veryflat,withfalsecellulosicsepta;2=
linear,cylindrical
28.seedcoat 0=smooth;1=pubescent
29.seedpubescence ?=inapplicable;0=persistinginmatureseeds; 1=absentfrommature
seeds
30.hilum 0=paralleltotheplacenta;1=perpendiculartotheplacenta
31.leaflets 0=entire;1=lobedtosomewhatlobedatbase
.
222 Sum18(1)
RVA-VMV.NCV.S
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