RANUNCULUS FICARIA (RANUNCULACEAE), NEW TO NORTH CAROLINA AND AN UPDATED KEY TO CAROLINA CONGENERS

2005·3.3 MB·English

by  Alexander Krings

RANUNCULUSFlCARIA(RANUNCULACEAE),NEWTO
NORTHCAROLINAANDANUPDATEDKEY
TOCAROLINACONGENERS
AlexanderKrings AlanS.Weakley
Herbarium,DepartmentofBotany Herbarium
NorthCarolinaStateUniversity UniversityofNorthCarolina
Raleigh,NorthCarolina27695-7612,USA. NorthCarolinaBotanicalGarden
[email protected] ChapeiHill,NorthCarolina27599-3280
[email protected]
JosephCNeal EdwardCSwab
DepartmentofHorticulturalScience Herbarium,DepartmentofBotany
NorthCarolinaStateUniversity NorthCarolinaStateUniversity
Raleigh,NorthCarolina27695-7609,U.S.A. Raleigh,NorthCarolina27695-7612,U.S.A.
Joe [email protected]
ABSTRACT
PopulationsoftheexpandingexoticRanunculusficariaL.(Ranunculaceae)arereportedescapedin
NorthCarolina.VoucherspecimensarecitedandanupdatedkeytoRanunculusL.intheCarohnas
ispresented.
RESUMEN
PoblacionesdelaexoticaenexpansionRanuncu!u5/iMnaL.(Ranunculaceae),secitanescapadasen
CarolinadelNorte.SecitanlosespecimenesysepresentaunaclaveactualizadadeRanunculusL.en
lasCarolinas.
PopulationsoftheexoticRanunculusficariaL.(lessercelandine)areknown
outsideofcultivationfromthenortheasternUnitedStates,Oregon,andWash-
ington(Gleason&Cronquist1991;USDA,NRCS2005),butthespecieshasbeen
previouslycollectedintheSoutheastonlyinKentuckyTennessee,Virginia,and
Maryland(Whittemore1997).RanunculusficariaisnativetomuchofEurope
(Tutm1976;Taylor&Markham1978).ThespecieswasreportedinNewYork
Stateasearlyas1890(HoUick&Britton1891).Bythemid1940s,itwasknown
fromseveralnortheasterncoastalstates,aswellasVirginia(Sargents.n.^NCSC),
buthadnotyetreachedWestVirginia(Bell1945).Thetaxonwasnotprevi-
ouslyreportedforNorthCarolinabyRadfordetal.(1968),Pittilloetal.(1969),
Pittilloetal.(1972),Krai(1981),PittilloandBrown(1988),orWhittemore(1997).
However,apopulationofthespecieswasrecentlyencounteredinRaleigh(Wake
Co.,NorthCarolina).Twenty-nineclumpswereobserved,ofwhichseventeen
wereeitherinflowerorfruitatthetimeofcollection.Individualswereobserved
SIDA21(4):2429-2437.2005
2430 BRIT.ORG/5IDA21(4)
onlyinareasreceivingpartialshadelorasignificantportionoftheday.They
wereabsentfromportionsofthelawnreceivinglullsunlight.
VmKlu-icollcctionll.S.A.NorthCarolina.WakeC.o.:Raleigh,back)'ardlavvnolresidentialhomeon
Vanl")yke,\\eniie,inpartialshade,absentIroinportionsoflawnreceivinglullsun,11Apr2005,
Kiin,i;,s127/(AUA,\\n.AS,GA,LSI;,MISS,NCSt;,NCU,TF.X,UNA,LIS,USF,USCl1,VOB).
Thespecieshasalsc^beenobservedonaprivateproperiyinChathamCo.,where
ithadapparentlybeenplantedinthepast,butisnowescapuigshortdistances
intonearbynaturalareas.IDuetosummersenescence,vouchersofthispopula-
tionhavenotbeentakenthisyear.
Severalattemptshavebeenmadetomorphologicallydistinguishdiploid
frompolyploidplantsofR.fiairia(Marsden-jones1935;Marsden-Jones&Turrill
1952; Lawalree1955).Unfortunately,mostofthese haveproven unrefiable
(Heywood&Walker1961;Jones1966;Taylor&Markham1978).Arguingthat
toogreatareliancehadbeenphicedonchroniosomecounts,Sell(1994)recog-
nizedIivesubspecies(Table1),withthecautionthatthesetaxacouldberecog-
nizedwitheaseonlyilcultix'atedorexaminedatinterx'alsthroughtheirflow-
eringand Iruiting periods. Ifonlyseenonce in theHeld orfrom asingle
specimen,certainidentilicationwouldremaindifficult.Thefollowingkeywas
providedbySell(1994)tofacilitateidentification:
1. 5L.e0afXbl3a.5demsmto8x9cm;petiolesto28cm;flowersto60mmdiam;achenesto
2. Stemsrathertobust,butstraggling;bulbilspresentinleafaxilsafterflowering
_— subsp.ficariiformis
2. Stemsrobustanderect;withoulbulbilsinleafaxilsafterflowering subsp.
chrysocephalus
1. 3L.e5afXbl2a.2demsmto4x4cm;petiolesto15cm;flowersto40mmdiam;achenesto
3. Leavescrowdedatbasewithfewonshortsterns subsp.calthifolius
3. Leaveslesscrowdedatbaseandmorenumerousontheelongatestems
4. Bulbilsnotpresentinleafaxilsafterflowering;acheneswell-developed
subsp.ficaria
4. RulbilspiestMitinleafaxilsafterflowering;achenespoorlydeveloped subsp.
bulbilifer
LISnA,NRCS(2005)reportedonlyR.ficdriavar.huUnlcniMarsden-Joneslor
theUnitedStates.Sell(1994)pointedoutthatthisnameisillegitimate,beinga
laterhomonymo(RjuaruivarbulhifcrAlbert,whichSelltreatedinsynonymy
undersubsp.bulbHijcrl.ambinon.BasedonWhittemoi'es(1997)description
olthespecies,theIkwersizesol indi\'idualsofsubsp.huihiUjcrintheUnited
Stateswouldbeontheupperendcompai'cdwithEuropeanmdix'idualsasrec-
ognizedbySell(1994)CRtble1).
l'ollowingSell(1994),theindividualsoftheWakeCo.,NorthCarolinapopu-
lationaretentativelyreferabletosubsp.fuiniifonnisiV.W.Schwartz)Rouy&
Fouc,previouslynotreportedfortheUnitedStates.Thoughtherearesomemore
KRINGSETAL.,RANUNCULUSFKARIANEWTONORTHCAROLINA 2431
Table l.ThefivesubspeciesofRanunculusficariaL.recognizedbySell(1994).Fls=flowers;
ptio=petioles;ptls=petals.
subsp.bulbilifer subsp.calthifolim subsp.chrysocephalus subsp.fkariiformis subsp.f/for/o
Lambinon (Reichenb.)Arcangeli P.D.Sell (F.W.Schultz)
Rouy&Fouc.
Tetraploid Diploid Tetraploid Tetraploid Diploid
Ptioss15cm Ptio=s7cmlong Ptio«s21cmlong Ptioss28cm Ptios;15cm
long long long
Axillarybulbils Axillarybulbils Axillarybulbils Axillarybulbils Axillarybulbils
globular absent absent ovoidor absent
globular
FIss=25mm FIs=s30mm FIs=s60mm FlsssSOraim FIs20-40mm
diamPtls6-11 diamPtis10-15 diamPtIs18-25 diamPtIs17-26 diamPtIs10-20
X2-5mm,not X2.5-6mm, X9-15(-18)mm, X4-12mm, X4-9mm,often
contiguous notcontiguous contiguousor contiguousor contiguous
overlapping overlapping
Occursthrough- Restrictedtoeast- Occursinthe Occursinthe Restrictedto
outmostrange centralandsouth- eastern centraland westernEurope
ofthespecies, easternEurope Mediterranean western
althoughrarein region Mediterranean
Mediterranean Region
region
diminutiveplantsinthepopulation,anumberofindividualsbearleaves>4
cmdiamandpetalss^17x6miu,thatarecontiguoustooverlapping(seeTable
1foracomparisonofcharactersamongsubspecies).Onsoineplants,ellipsoid
axillarybulbilsareevident.However,subspecifictaxaofR.ficanaarenotuni-
formlyaccepted.Citingextensiveintergradationinform,Whittemore(1997)
didnotrecognizeanysubspecifictaxainhistreatmentofRanunculusL.for
theFloraofNorthAmerica.Tohelpaddressthecontinuingdisparitymtaxo-
nomictreatments,detailedstudiesareneededtoexaminethedistributionof
haplotypesandpotentialmorphologicalcorrelations.Suchanapproachhas
beenusefulforotherexpandingexoticplants(e.g.,Saltonstall2002,2003a-c;
Saltonstallctal.2004).Furtherinvestigationisalsoneededtodeterminemore
preciselythetimingandmodeofintroduction,aswellastherateofspreadof
thespecies.Surveyofherbariumcollectionsmayshedmorelightonthismat-
ter.Pertinentstudiesregardingthelifehistory,pollinationbiologyandecology
havebeenpublishedbyMarsden-Jones(1935,1936)andTaylorandMarkham
(1978).
Toimprovecollections,Sell(1994)suggestedthatspecunensshould be
takenlatewhenfruitandbulbilsaredeveloped,fiowever,atthisstageanyflow-
ersremainingopenaretypicallylateones,whicharegenerallysmallerthan
2432 BRIT.0R6/SIDA21(4)
hQ.^.RanunculusficariaL.A.Habit;B.Flowers;C.Tuberousroots.BasedonKrings1271.
KRINGSETAL,RANUNCULUSFKARIANEWTONORTHCAROLINA 2433
thosewhentheplantfirstcameintoflower.Toimproveourunderstanmgof
thedistributionandecologyofsubspecies,botanistsshouldnoteflowersizes
onaninitialvisittopopulationsandthencheckthebulbilsandfruitsatalater
date(Sell1994).
RanunculusJicaria(Fig.1)canbedistmguishedfromitsCarolinaconge-
nersbythecombinationofunlobed,reniformtosuborbicularleaves,tuberous
roots,typicallythreesepals,yellowpetalss*10mmlong(sometimespartially
fadingtowhitewithage),andpubescent,beaklessachenes.Anupdatedkeyto
Carolinacongeners,largelyadaptedfromWhittemore(1997),isprovidedbe-
low.Followingarguments presentedbyNesom(1993),wedivergefrom
Whittemore(1997)intreatingR.carolinianusDC.asaspecies,ratherthanasa
varietyofR.hispidusMichx.Distribution,habitat,andfrequencyinformation
followsWeakley(2005)(Mt=Mountains;Pd=Piedmont;Cp=CoastalPlain).
Unlessotherwiseindicated,provincialdistributionsandcommentsapply
equallytoNorthCarolina(NC)andSouthCarolina(SC).Asterisksindicateex-
oticspecies.
KEYTORANUNCULUSINTHECAROLINAS
1.Allleavesunlobed
2.Leafbladesreniformtosuborbicularororbicular,basesshallowlytodeeply
cordate.
3. Rootstuberous;petals5= 10mmlong;achenespubescent,beaksabsent
*R.ficariaL[Pd(NC);disturbedrich
forestsandbottomlands,mesicsuburbanforests,lawns,naturalized
locallyfromhorticulturalplantings;rare]
3. Rootsfiliform;petals=£3.5mmlong;achene5glabrous,beakssubulate,curved
R.abortivusL.[Mt,Pd,Cp;low
fields,disturbedareas,bottomlands,lawns,roadsides;uncommon]
2.(LesaofmebtliamdeesscoovradtaetetoinlaR/nocxe/oclfaltu/eZ,s,btahseensptertuanlcsat2e-,6rmoumndleodn-g)o.btusetocuneate
4. Petals1-3,1.5-2mmlong R.pusillusPoir.fMt(NC),Pd,Cp;
marshes,ditches,otherwethabitats;common(uncommoninMt)]
4.5P.etaPlrsox4i-m6a,l5c-a8ulminmelleoanfgblades5.9-12.2cmlong;petals5;achenesto1.8mm
long ^ R.ambigensS.Wats.
[Pd(NC),Cp(NC);marshes;rare]
5.Proximalcaulineleafbladesto5.7cmlong;petals4-6;achenes0.8-1mm
long R.laxicaulis(Torr.&A.Gray)
Darby[Cp;marshes;rare]
1.Allorsomeleaveslobedorcompound.
6. Leafystemscreepingandrootingatthenodes,orfloatinginwater(then
rootless).
7.Leaves3-foliolate.
8.Achenemargins0.4-1.2mmwide R.carolinianusDC.[Mt,Pd,Cp;swamp
forests,wetwoodlands,openmarshywetlands;uncotmmon]
.
2434 BRIT.ORG/SIDA21(4)
8.AchenemarginsOJ-0,2mmwide "R.repensL.[Mt(NC),Pd(NC),Cp;
bwmeadows,disturbedareas;uncommon]
7. Leavessimple,lobed,parted,ordissected.
9.L1e.6avmesm=l£on1gcmlong;floralreceptaclesglabrous;petalswhite;achRe.nheesde=sraceusL.[Cp;
coastalbrackishmarshes,othercircumiieulialsoils;rare]
9. Leaves3=1.2cmlong;floralreceptaclessparselyhispid;petalsyellow;
achenes5=1.8mmlong R.flabellarisRaf.
[Cp(NC);poolsinfloodplainsofsmallstreamswamps,
otherstagnantorslowmovingwaters;rare]
6.Leafystemserectorifdecumbentrootingonlyatthebase(rarelyrootingatthe
nodesinRsce/erafus),neverfloating.
10.Styleabsent;achenemarginsthickandcorky;emergentaguaticoronwet
soil R.sceleratusL,[Pd(NC),Cp;
marshes,ditches,andstreammargins;uncommon]
10, Stylepresent;achenemarginsnotcorky;varioushabitats,butnotaquatic.
11, Basalleavesvariouslyunlobedtodeeplydivided;achenesthick-lenticu-
larorasymmetricallythick-lenticulartocompressed-globose,1.2-2times
aswideasthick.
12. Stemsvillous R.micranthusNull,[Pd(NC);richforests;rare]
12. Stemsglabrous.
13, Sepalsglabrous;achenebeaks0,1-0,2mmlong R.abortivusL,
[Mt,Pd,Cp;k:)Wfields,disturbedareas,
bottomlands,lawns,roadsides;uncommon]
13.Sepalshispid;achenebeaks0,6-1mmlong R.alleghaniensisBrilton
[Mt(NC,SC?);coveforests,
richforestedslopes;uncommon]
1I, Basalleavesalwaysdeeplylobedorcompound;achenesstronglycom-
pressed,atleast3-15timesaswideasthick,
14. A15c,hePneteaslssp1i-n2osmemorlot)nagp;ilrleocseept(ascolmeestgilmaebsrosums;ooatchheinneRs.sfdiniedloyusp)a.pil-
late,eachwithahookedbristle.
16. Flowerspedicellate;sep[aMlls,P5d,Cp;disturbedareas;com"mRo.npar(vrairfelionruMst)I]
16. Flowerssessile;sepals3 "R.platensisSpreng.[Pd(NC);
15.Petals4 10mmlong;receptaclespiloseorhispid;laawcnhseannedsditches;rare]
coarselypapillate(butnotterminatinginhookedbristles),spi-
nose,ortuberculato.
17.Sepalsspreading;achenes5-9,borneinasinglewhorl,long
spinose "R.arvensisl,|Mt(NC),Pd;
fields,disturbedareas;lare]
17.Sepalsreflexed;achenes13-60,borneinovoidorglobose
heads,papillosetospinose,
18.Basalleavessimple;achenebeaks2-2.5mmlong
"R.muricatusL
[Pd(SO,Cp(SC);ditchesandmarshes;lare]
18, BasalleavescorTipound;achenebeaksto0,7mmlong.
1
KRINGSETAL.,RANUNCULUSFICARIANEWTONORTHCAROLINA 2435
19. Petals7-10mmlong;achenessparselypapillateor
sometimessmooth *R.sardousCrantz[Pd,Cp;
bwfields,disturbedareas;uncommon]
19. Petals4-5mmlong;achenesdenselytuberculate
"R.trilobusDesf.[Cp(SC);
fields,roadsides,ditches;rare]
14.A20c.heP^neetaslssmo3o-t5h,mgmlabrloounsg;orapcuhbeensecenbte.aks markedlyrecurved
R.recurvatusPoir.[Mt,Pd.Cp;bottomland
forests,coveforests,swamps,mesicslope
forests;common]
20. Petals9=7mmlong;achenebeaksmoreorlessstraight,not
markedlyrecurved.
21.Sepalsreflexedalongadefinedfold1-3mmabovebase.
22.Stembasesbulbou5,corm4ike;petals9-13mmX8-1
mm *R.bulbosusL.[Mt,Pd,Cp;fields,
roadsides,disturbedareas;common
(rareinSouthCarolina)]
22.Stem basesnotbulbous;petals7-10 x 4-8mm
"R.sardousCrantz[Pd,Cp;
lowfields,di5turbedareas;uncommon]
21.Sepalsspreading(sometimesreflexedfrombasewithage).
23. Basalleafblades3-5-parted,pentagonalinoutline
*R.acrisL.[Mt(NC),Pd(NC),Cp;pastures,
fields,roadsides,disturbedareas;common
(uncommoninPd,rareinCp)]
23. Basalleafblades3-5-foliolate,ovatetodeltateinoutline.
24.Tuberousrootsabsent R.hispidusMichx.
[Mt,Pd;richmoistforests,creekbanks,
mesictodrywoodlandsandforests,
bottomlands;common]
24.Tuberousrootspresent R.fascicularisMuhl.
exBigelow[Mt(NC),Pd;
wetflatswithprairieaffinities,rocky
barrensandgladesover
maficrocks,ultramaficoutcropbarrens,
limestonebarrens;rare]
ACKNOWLEDGMENTS
WearegratefultoCharlesW.Averre(Prof.Emeritus,Dept.PlantPathology,
NCSU)forbringingtheplanttoourattentionandthethoughtfulreviewsofa
previousversionofthemanuscriptbyMichaelDenslow,CarlKeener,andan
anonymousreviewer
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