NEW TAXA AND NEW COMBINATIONS IN NORTH AMERICAN CIRSIUM (ASTERACEAE: CARDUEAE)

2004·6 MB·English

by  David J Keil

NEWTAXAANDNEWCOMBINATIONSINNORTH
AMERICANCIRSIUM(ASTERACEAE:CARDUEAE)
DavidJ.Keil
BiologicalSciencesDepartment
CaliforniaPolytechnicStateUniversity
SanLuisObispo,California93407,U.S.A.
[email protected]
ABSTRACT
SixnewvarietiesareproposedinNoriiiAinericanCirsium(Asteraceae:Cardueae):C.arizonicum
(A.Gray)PetrakvantenuisectumDJ.Keil\^arnov,.C.eatonii(A_GrayjB.L.Rob^var.viperinumD,J.
Keil,var.nov.,C.edulcNutt.var.wenatchcnscI.^J,Keil,var.nov,C.occidentale(Nutt.)Jeps.varlucianum
D.J,Keil,varnov,C.scariosumNutt,varrobustumD.J,Keil,varnov,C.scariosumNutt,varloiyabcnse
DJ,Keil,varnovAdditionally,twentynewcombinationsarepresented:C.arizonicum(A,GrayJ
Petrakvanbipinnatum(Eastw,)DJ,Keil,eoinb.nov,C.arizonicum(A,Gray)Petrakvar.chellyense
(RJ.Moore&Frankton)DJ.Keil,comb,etstat.nov.C.arizonicum(A.Gray)Petrakvarroihrockii(A.
Gray)DJ.Keil,comb,etstat.nov,C.clavatum(M.E.JonesJPetrakvar.americanum(A.Gray)DJ,Keil,
comb,nov,C.clavatum(M.E,Jones)Petrakvarosterhoutii(Rydb,)DJ,Keil,comb,etstat.nov,C.
cymosum(Greene)J.T.Howellvarcanovircns(Rydb)DJ,Keil,comb,etstat.nov,C.ealonii(A.Gray)
B.L.Rob.varcrioccphalum(A.Nelson)DJ,Keil,comb,nov,C.eatonii(A,Gray)B.L.Rob.varhesperium
(Eastw.)DJ.Keil.comb,etstat,nov,C.eatonii(A,Gray)B,L.Rob.varpeckii(L.EHend.)DJ.Kcil,
comb,etstat,nov,C.eatonii(A,Gray)B.L.Rob.varvarclokeyi(S,EBlake)DJ,Keil,comb,etstat,nov,
C.eduleNutt.varmacounii(Greene)DJ,Keil,comb,etstat,nov,C.horridulumMichx,var
megacanthum(Nutt.)DJ.Keil,comb,etslat.nov.C.inamocnum(Greene)DJ.Keil.comb,nov,,C.
inamoenum(Greene)DJ.Keilvardavisii(CronquistJDJ.Keil,comb,etstat.nov,C.ochrocentrumA.
Grayvar.martinii(PBarlow-Irick)DJ.Keil.comb,etstat,nov,C.pukherrimum(Rydb.)K.Sebum,van
aridum(R.D.Dorn)DJ.Keil.comb,etstat,nov,C.scariosumNutt,varamericanum(A.Gray)D,J,Keil,
comb,no\'„C.scariosumNutt,vancitrinum(PetrakjDJ.Keil,comb,nov,C.scariosumNutt.\'an
coloradensc(Rydb.)DJ.Keil,comb,etstal.nov,C.scariosumNutt.varcongdonii(RJ.Moore&
Frankton)DJ.Keil,comb,etstat.novLectotypesaredesignatedlorseveralofthesetaxa.
RESUMF.N
SeproponenseisvariedadesnuevasenCirsiurn(Asteraceae:Cardueae)deNorteamerica:C.arizonicum
(A.Gray)PetrakvartenuisectumDJ.Keilvarnov,C.eatonii(A.Gray)B.L,Rob.var\iperinumDJ.Keil,
varnov,C.eduleNutt.varv/enatchettseDJ.Keil,vannov,C.occidentale(Nutt.)Jeps.varlucianumDJ,
Keil.varnov,C.scariosumNutt.varrobustumDJ.Keil,varnov,C.scariosumNutt.vantoiyahenseDJ.
Keil.varnovSepresentanademasveintecombmacionesnuevas:C.arizonicum(A.Gray)Petrakvar
bipinnatum(Eastw.)DJ.Keil,comb,nov,C.arizonicum(A.Gray)Petrakvarchellyense(RJ.Moore&
Frankton)DJ.Keil,comb,etstat.nov,C.arizonicum(A.Gray)Petrakvarrothrocfeji(A.Gray)DJ.Keil,
comb,etstat.nov,Ccla\atum(M.E.Jones)Petrakvaramericanum(A.Gray)DJ.Keil,comb,nov,C.
cla\atum(M.E.Jones)Petrakvarosterhoutii(Rydb.)DJ.Keil,comb,etstat,nov,C.cymosum(Greene)
J.THowellvancaiiovircns(Rydb.)DJ.Keil,comb,etstat.nov,C.eatonii(A.Gray)B.L.Rob.van
eriocephalum(A.Nelson)DJ.Keil,comb,nov,C.eatonii(A.Gray)B.L.Rob,vanhesperium(Eastw.)DJ.
Keil,comb,etstat.nov,C.eatonif(A.Gray)B.L.Rob.vanpeckii(L.FHend.)DJ.Keil,coinb.etstat.nov,
C.eatonii(A.Gray)B.L.Rob.varvarclokeyi(S.FBlake)DJ.Keil,comb,etstat.nov,C.eduleNutt.var
SIDA21(1):207-219,2004
208 BR1T.ORG/SIDA21(1)
macounii(Greene)DJ,Keil,comb,etstat.nov..C.horridu\uinMichx.varmegacanthum(Nutt)D.J.Keil,
comb,etstat.nov..C.inamoenum(GreeneJD.J.Keil.comb,nov.,C.inamoenum(Greene)DJ.Keilvar
davisii(Cronquist)DJ.Keil,comb,etstai.nov,C.ochrocentrumA.Grayvar.martinii(P.Barlow-lrick)
DJ.Keil,comb,etstat.nov.,C.pulcherrimum(Rydb)K.Schum.vararidutn(R.D.Dorn)DJ.Keil,comb,
etstat,nov.,C.scariosumNutt.var.ameiicanum(A.Gray)DJ.Keil,comb.nov.C.scariosumNutt.var.
citrinum(Petrak)DJ.Keil,comb,nov,,C.scariosumNutt.var.coloradense(Rydb.)DJ.Keil,comb,et
stat.nov,C.scariosumNutt.vancongdonii(RJ.Moore&I'rankton)DJ.Keil.comb,etstat.nov,Tambien
sedesignanlectotiposdcvariosdcestostaxa.
PreparationofataxonomictreatmentofCirsiumMill.(Asteraceae:Cardueae)
fortheupcomingAsteraceaevolumesoftheFloraofNorthAmericahasen-
tailedareevaluationofmanyofthepublishedtaxa.Theonlycomprehensive
treatmentofNorthAmericanCirsiumspecies(Petrak1912,1917)islongoutof
date.Revisionsorcytotaxonomicstudieshavebeenpublishedforseveralspe-
ciesgroups(Frankton&Moore1961,1963;Moore&Frankton1962,1964,1965,
1966,1969,1974;Ownbey1952;Ownbey&Hsi1963,1969;Ownbey&Olson
1969;Ownbeyetal.1975),andmanyofthespecieshavebeenincludedinstate
orregionalfloristictreatments(e.g.,Cronquist1955,1994;Dorn1992;Gardner
1974;Flowell1960;Keil&Turner1993;Munz&Keck1959;Munz1974;Weber
1996;Weber&Wittmann1987;Welsh1983;Welshetal.1993),butthesetreat-
mentshavebeenunevenandlimitedmscope.Politicalboundariesoftenmark
theapplicationofdifferingtaxonomicconstructs.InmyinvestigationsIhave
examinedNorthAmericanCiniumasawholeandacrossthebreadthoi its
geographicrange,
Cirsiumwelldeservesitsreputationasataxonomicallydiflicultgenus.One
ofthemostchallengingaspectsforataxonomiststudyingNewWorldCirsium
isthepresenceofspeciescomplexesthatareapparentlyevolutionaryworksm
progress.Someofthethistles,especiallyinthemountainouswesternpartot
NorthAmerica,arefrustratmglypolymorphicwithmosaicandoverlapping
patternsofvariationandintergradationoicharacters.Farlytaxonomists,bas-
ingtheirworkonalimitedsamplingofthemorphologicaldiversity,named
manyoftheformsasspecies,andthefiteratureisrobustwithspeciesnames.
Theinfillingthatresultsfrommorecollectorsvisitingmorelocalitieswithin
therangesofthesecomplexeshasblurredtheboundariesbetweenmanyofthe
proposedspeciesandoftenaddedformsthatdonotTit"thecharactersotnamed
species.AsIfacedthechallengesofpreparingtheFNAtreatmentIrecognized
thatmaintainingsomeofthenamedentitiesasspecieswouldforconsistency
requireafurtherproliferationofspeciesnames.
1havechosentogotheotherwayInsteadcontinuingtheproliferationof
ill-definedmicrospeciesItreattheplantsinquestionasrapidlyevolving,only
partiallydifferentiatedassemblagesofracesthathavenotreachedthelevelof
stabilitythatisusuallyassociatedwiththeconceptofspecies.Inamolecular
phylogeneticinvestigationKelchandBaldwin(2003)foundunusuallylowrates
KEIL,NEWTAXAANDNEWCOMBINATIONSINNORTHAMERICANCIRSIUM 209
ofmoleculardivergencerelativetotheecologicaldiversityof NewWorld
Cirsiumandsuggestedthatthegenushasundergonearapidecologicaldiver-
sificationonacontinentalscale.Certainlythereismuchvariationwithinthese
groupsthatdeservesaleveloftaxonomicrecognition,orthatatleastshouldbe
mentioned,butIthinkitmuchmoreprudenttorecognizetheseasinfraspe-
cifictaxa—entitiesthatmaybeexpectedtofreelyintergrade—thanspecies.I
viewspeciesasentitiesthatshouldbeconsistentlyrecognizable.Withinalarge
genussuchasCirsiumtheamountofinfraspecificvariationandthedegreeof
geographicalseparationamongracesvariesfromspeciestospeciesandfrom
geographicalracetogeographicalrace.Racesoccupyingcontiguoushabitats
oftenintergradefreelyinzonesofcontactwhereasracescurrentlyisolatedby
geographicorhabitatbarriersoftenhavemorediscretevariationpatterns.Both
situationsmayoccurinasinglespecies.
InfraspecifictaxainCirsiumhavebeenrecognizedinvarioustaxaatthe
rankofform,variety,andsubspecies.IntheFloraofNorthAmerica(Floraof
NorthAmericaEditorialCommitteei996)thetrivialrankofform,represent-
ingsporadicvariantswithoutageographicrange,isnotused.SomeCirsium
variantsrecognizedmthe pastasvarietiesaretheequivalentofforms.In
Cirsiumbothvarietyandsubspecieshavebeenusedinthecontextofmorpho-
logicallydistinguishablegeographicalraces.Therankofvarietyhasbeenused
morewidelyTheFloraofNorthAmericaGuideforContributors(FloraofNorth
AmericaEditorialCommittee1996)stipulatesthatonlyoneinfraspecificrank
(subspeciesorvariety)berecognizedwithinagenus.Asusedbymostworkers
inCirsiumtaxonomythechoiceofrankhasbeenamatterofpreference,andI
viewthemasequivalent.Ihavechosentousevarietyratherthansubspecies
becauseonlyonechangefromsubspeciestovarietyhastobemadetofollow
theFloraofNorthAmericaGuidelineswhereastheuseofsubspeciesasthe
infraspecificrankforformallyrecognizedgeographicalraceswouldrequire
manymorenomenclaturalinnovations.
TheoverallproductofmystudieswillbedetailedintheFNAtreatment
(Keil,inpress).Presentedbelowaresixnewlyproposedvarietiesandvalida-
tionottwentynewcombinationsforNorthAmericanCirsium.
Cirsiumarizonicum(A.Gray)Petrakvarbipinnatum(Eastw.)D.J.Keil,comb,
nov.BASIONYM:CnicusdrummonditTorn&rA.Grayvar.hipinnatusEastw.,Zoe
4:8.1893.Type:COLORADO:JohnstonCanonnearwhereitjoinsMancosRiver,
Sep1892,Eastwoods.n.(holotype:CAS;isotype:MIN).
Cirsiumarizonicum(A.Gray)Petrakvarchellyense(RJ.Moore&Frankton)
DJ.Keil,comb,etstat.novBasionym:Cirslumchcllye7^seR.].Moore&Frankton,
Canad.J.Bot.52:547.1974.Type:ARIZONA.ApacheCo.:CanyondeChelly,1Jul
1961,Norman680(holotype:DAO).
210 BRIT.ORG/SIDA21(1)
Cirsiumarizonicum(A.Gray)Petrakvanrothrockii(A.Gray)D.J^Keil,comb,et
Stat.nov.BasionYM:Cmcusrothrock11A.Gray,Proc.Amer.Acad.Arts17:220.1882.
Type:ARIZONA:RockyCanyon,1874,Rothrock289(lectotype,herechosenfrom
syntypes,GH).Gray(1882)describedCnicusrothrockiibasedupontwocollec-
tions{Rothrock289andLemnion2794).
MooreandFrankton(1974)selectedRothrock289asthelectotypecollectionof
Cnicusrothrockii.However,theycitedaslectotypespecimensolthisgathering
frombothGHandUS.AccordingtoArticle9.2oftheInternationalCodeof
BotanicalNomenclature(Greuteretal.2000)"alectotypeisaspecimenoril-
lustrationdesignatedfromtheoriginalmaterialasthenomenclaturaltype."
MooreandFrankton'sdesignationoftwospecimensatdifferentinstitutionsas
lectotypefailstherequirementthatasinglespecimenserveinthatrole.Iagree
withtheirdesignationofRothrock289asthelectotypecollectionandhere
choosetheGHspecimenaslectotypeforthetaxon.
Cirsiumarizonicum(A.Gray)PetrakvartenuisectumD.J.Keil,varnovType:
CALIFORNIA.SANBernarlmnoCo.:NewYorkMountains,KeystoneCanyon,2.1
roadmifromIvanpahRoad,ca.1670m.Pinusmonophylla-]uniperusosteospcrma
woodland;scatteredinrockywashchannel;associates:Fallugiaparado.xa,Prunus
fasciculata.Guticrreziasarothrae,Atriplcxcanescens,Eriodictyonangustijoliuni,
YuccahreviJolia,OpuiUiaechinocarpa,Me^rLodorascahra,Ephedranevadensis,Pen-
stemor]sp.,Machacranlhcracanesccns,Purshiamexicana,Muhlenhergiaporteri.
Ericamerialinearijolia.Salviadorrii,Lyciumcooperi,Yuccabaccata,Berhenssp.,
R/iirstrilobata,Sphaeralceasp.,Ipomopsisagrcgaia,Daturawrightii,Artemisia
dracunculus,andAchnatherumhynienoides,5Nov1999,Keil28401(holotype—
floweringstemofplantD,OBI;ISOTYPES[theletterinparenthesesrepresentsthe
individual],ASU(A),BRY(C),CAS(B),CDA(C),DAO(D),GH^D),MIN(D),MO
(C),NY(C).OBI(A,C),RSA(A),TEX(D),UC(13),US(D)),
C:aulibusetcostisabaxialibusglabrisadromenrosistricliomatibustenuibusnon-septatis,spinis
pnncipaiibusloliorum5-30mmlongis.ctcorollisrubellopurpureisdistinguatur
Stemsthinlyarachnoidtomentose,± glabrate,withoutseptatetnchomes.
Leavesdeeplydivided,oftennearlytothemidvein,abaxiallyarachnoidtomen-
toseorsometimesglabrate,withoutseptatetrichomes,adaxiallythinlyarach-
noidorglabrate;principalmarginalspines5-30mm,oftenstout;caulineleaves
narrowedatbasetotruncateor±clasping,butnotoronlyslightlydecurrent.
Involucrescylindrictocampanulate;phyllaryspines5-25mm,stout.Corollas
reddishpurple,25-35mm,thetube10-13,throat5-8mm,lobes10-13.5mm;
styletips1-2mm.
Paratvi'i:s:CALIFORNIA.SanBernardinoCo.:NewYorkMountains.CarruthersCanyon,1615-1645
m19,356,NMouvn1z99193,86Ke2i(lUC2)8.40Ne2v(aNdaM,C.ClOaBrIk.COoS.:C)S;WNselwopYeooifTNMWounStpariinnsg.MKtse.y,sutpopneerSCplrairngk,C1a5n8y5onm,.y1e3lOlcotw
pinevegetation,7000-7600It,9Jul1970,Becitley.113.55(LINLV);CharlestonMrs,.CharlestonPeak,
gravellywash;withPinusscopulorum,2270m,18Jul1938,ClokevSlSZ(PH,UC.UTC(2j,WTUC2)j;
CharlestonMts.,KyleCanyonatbridge,washwithaspens,2300m,3Aug1935,Clokcy5609(UC);
ChariesronMts,,CharlestonPark,gravellywash,yellowpmebelt,2300m,25Jul1936,Clohey5609
1
KEIL,NEWTAXAANDNEWCOMBINATIONSINNORTHAMERICANCIRSIUM 21
(UC,UTC,WTU);CharlestonMts.,LeeCanon,inlimestone,8000ft,4Aug1913,HcllcfJJ057(PH,
UCj;CharlestonMts,PmeCr,6700ft,4Sep1927.jaeger2202(UC);CharlestonMts.,nearCharleston
KpyalrekCreasnoryto,nplRedn.ti(fuNle,v8ad0a00Rtfet.,11257S)ejpus1t92W5.oJfaejguenrctsi.no.n(wUiC)t;hSNperviandgaRaRtneg.e,157T,oiPyoanbdeerNoastaiopnamleFfoorreesstt,,
6930ft,17Aug2002,Keil29112(OBI);SpringMountains,ToiyabeNationalForest,LasVegasSkiand
SnowboardareainUpperLeeCanyon,36,299°N,115.696°W,subalpinelorestwithlimberpine,bristle-
conepine,whitefir,quakingaspen,wa.\currant,andcommonjuniper,9200ft,17Aug2002,Keil
29114(OBI);RedRockCanyonRecreationLands,MouthofVelvetCanyon,withCeltisreticulata,
Quercusturhinella,andChrysothamnuspantcidatus,4400ft,6Jun1990,Learyetal.4006(UNLV);
SpringRange,NVv'ofLasVegas,upperLeeCanyonbetweenskiareaandsaddleatheadofWallace
Canyon,coniferforestwithPinusflexilis,P.longaeva,AbiesconcoloKandPopulusIremuloidcsonsteep
limestoneslopesandcanyonbottom,8700-9900ft,31Aug1986,Sandersetal.6881(OBIj.
TheCirsiumarizonicumcomplexisagroupofcloselyrelatedtaxaotthesouth-
vv^esternUnitedStatesandnorthwesternMexico.Thisspeciescomplexwasre-
visedbyMooreandFrankton(1974),whorecognizedeightspecies,threeolthem
newlydescribed.InMooreandFrankton'skeytospeciesoftheCanzonicum
complex,vartenuisectumkeystoCirsiumnidulum(M.E.Jones)Petrak,and
theseauthorscitedachromosomecountfromtheSpringMountainspopula-
tionofvar.tenuisectumasC.nidulum.MooreandFrankton'sratherimprecise
rangemapdoesnotincludetheNewYorkMountainsforC.nidulumorany
othermemberoftheC.arizonicumcomplex,andthereisnoindicationthat
MooreandFranktonwereeuvareot thethistlesfromthismountainrange.
CirsiumnidulumisthenamethathasbeenusedfortheNewYorkMountains
thistleinvariousfloras(fiowell1960;MunzandKeck1959;Munz1974;Keil
andTurner1993,2002).AsapartofmyresearchonCirsiumfortheFloraof
NorthAmericaproject,Ihaveexaminedmanyspecnnens,includmgtypes,from
throughouttherangeofthevariousmembersoftheCirsiumarizonicumcom-
plex,andIhaveconcludedthattheCanzonicumcomplexshouldbetreatedas
asinglepolymorphicspecies.IthasbecomeevidentthatthenameCirsium
nidulumhasbeenwidelymisapplied;thetypeisaveryspinyplantofvar
arizonicum.TheplantsfromtheSpringMountainsandNewYorkMountains
representapreviouslyunnamedvariety.
Cirsiumclavatum(M.E.Jones)Petrakvar.americanum(A.Gray)DJ.Keil,comb,
nov.Basionym:CnicuscarlinoidesSchrankvar.americanusA.Gray,Proc.Amer.
Acad.Arts10:48.1874.Type:COLORADO:RockyKits.,lat.39-41°,HallandHarbour
339(lectotype,lierecliosenfromsyntypes,GH).
ThevarietalepithetamericanumwasbaseduponsyntypesfrombothColo-
radoandCalifornia.ThelatterarereferabletoCirsiumremotijolium(fiook.)
DC.Gray(1874)citedtwoColoradocollections,Greenes.n.andHallandHarbour
3'^2.Cronquist(1994)statedthat"Hall&Harbour342,RockyMts.,Colo.,isthe
firstcollectioncitedandhasbeentakentobethetype;originalatGH!"1agree
withthechoiceofHallandHarbour342andamvalidatingitsdesignationas
lectotype.
212 BRIT.ORG/SIDA21(1)
Cirsiumclavatum(M.E.Jones)Petrakvar.oslerhoutii(Rydb.)D.J.Keil,comb,et
Stat.nov.BASlONYMiQirduusostfc'WioutiiRydb.,Bull.TorreyBot.Club32:131.1905.
Typh:COLORADO.FAGi.liCo.:RedCHff,1902,Ostcrhoui2706(ilOi.OTVPi;:NY;
ISOTYPE:RM).
Cirsiumcymosum(Greene)J.T.Howellvar.canovirens(Rydb.)DJ.Keil,comb,et
Stat.nov.Basionym:CarduuscanovirensRydb,,Mem,N,Y.Bot.Card.1:450.1910.
Typh:MONTANA,MadisonCo.:JackCreekCanon,7000ft,15Jul1897,Rydhcrg
andBcs.st'v5213(holotypp,:NY).
Cirsiumeatonii(A.Gray)B.L.Rob.var.eriocephalum(A.Nelson)DJ.Keil,comb,
nov.Basionym:CirsiumcriocephalumA.Gray,nom,illeg,;Carduushookerianus
Nutt.Ivan]eriocephalusA.Nelson,inJ.M.Coult.&A.Nelson,Man.Bot.RockyMts,
585.1909.Type:COLORADO:headwatersofClearCreekandthealpinendgesly-
ingEofMiddlePark,1861,Parrys.n.(lf.ototypf,,herechosenfromsyntypes,GH),
ThistaxonwasoriginallynamedCirsiumeriocephalumbyGray(1863),but
thisname wasalaterhomonym.BecauseoftheillegitimatestatusofC
eriocephalum,apriorableautonymwasnotcreatedbypublicationolC;rsium
criocephalumA.GrayvarIciocephalumD.C.Eaton(1871).However,Nelson's
recognitionofCarduushookerianusvar.erwcephalusestablishedthisepithet
atthevarietallevelwithNelsonastheauthoroftheepithet,notGray[ICBN
Art.58].Nelsondidnotciterankformfraspecifictaxabuthistrinomialcombi-
nationsaretobetreatedasvarieties(ICBNArt.35.4).
Cirsiumeatonii(A.Gray)B.L.Rob.varhesperium(Eastw.)DJ.Keil,comb,et
Stat,novBasionym:CnicushesperiusEastw.,Bull.Calil.Acad.Sci.ser3,1:122.1898.
Typp:COLORADO:LaPlataMts.,Mt.HesperusonBearCreekDivide,Aug1892,
Eastwoods.n.(holotypf:CAS).
C:irsiumeatonii(A.Gray)B.L.Rob.var.peckii(L.F.Hend.)DJ.Keil,comb.etstat.
novBasionym:CirsiumpeckiiL.FHend.,Madrono5:97.1939.Type:OREGON.
HarneyCo.:AlvordRanch,EbaseofStcensMc,6Jun1927,Irlcndcrson8521(ho-
LOTYPE:ORE).
Cirsiumeatonii(A.Gray)B.L.Rob.var.clokeyi(S.F.Blake)DJ.Keil,comb,etstat.
novBasionym:Cir.siumclokeyiS.EBkd<e,Proc.Biol.Soc.Washington51:8.1938.
TYPE:NEVADA.Cl.ARKCo.:CharlestonMts.,6Aug1937,Clokey7456(holotype:
US;ISOTYPES:BRY,DAO,GA,MO,PENN,PH,RM,UC).
Cirsiumeatonii(A.Gray)B.L.Rob.var.viperinumDJ.Keil,varnov.Type:NE-
VADA.WhitePineCo.:SnakeRange,HumboldtNationalForest,SnakeCreek
Canyon,aboveJohnsonLake,steeprockyslope,granite,common,11,000It[3353
m],10Aug1964,Ho/mgrcnandReveal}588{\iolotype:UTC;isotypes:WTU,MIN).
Ckianpgiits,uHssetirsarleoanrgiascshinmoiisdepiasppriri2c0ho-i2n5atmibmusdniosnr-isngeupatiautrls,corollaslavandulisadpurpureis,29-35mm
Plantserect,2.5-4dm.Leavesglabrousornearlysoonbothsurfaces.Capitula
1-5,subsessileorshort-peduncled,mraceme-likeorcorymbiformcapitulescence.
Involucres3-5cm,thinlyarachnoidwithnon-septatetrichomes;outerphyl-
KEIL,NEWTAXAANDNEWCOMBINATIONSINNORTHAMERICANCIRSIUM 213
larieswithnumerouslateralspines;spinetipsofphyllariesstout.Corollaslav-
endertopurple,29-35mm,thetube9-12.5mm,thethroat9-12mm,thelobes
9-11mm.Longestpappusbristles20-25mm.
Paratypi;s:NEVADA.WhitePineCo.:SnakeRange.Mt.Washington,bristleconepinelorestjustSof
thepeak.T12N,R68E,Sect.11,plantsgrowingwithPinu.sonlimestonegravelslopes,11,300ft[3444
ml.9Aug1985,Tiehm10107(CAS,COLO,MINI,OSC,UNLV);MountMoriah,NendofSnakeRange,
moistlimestonederivedsandytorockysoil,onS-facingslopewithPhlox,Aquilegia,Crepisnana.
Al/ium,andEri^(^cron,T18N,R69E,Scct10,11840ft(3500m).2Aug1999,Nilcsetal.ScS'JcS(UNLVj;12
miSWofBaker,HumboldtNationalForest,SnakeRange,NsideofMt.Washington,T12N,R68E,F.
1.2ofSect.11,withscatteredbristleconepine,11,000ft,12Sepf983,Goodrichelal.20063(BRY);13mi
SWofBaker.SnakeRange,SslopesofMt.Washington,T12N,R68E,N1/2ofSect.14,upperedgeof
bristleconepme-limberpinelorest.11,000ft,12Sep1983,S.Goodrichetal.20063(BRYj.
Cirsiumeatoniivarviperinumisapparentlyendemictoupperelevationsof
theSnakeRangeofWhitePineCounty,Nevada.Capitulaofvar.viperinumare
similarinsizetothoseofvarclokeyi.However,varviperinumisashorterplant
(2-4dmvs.4-15dm)withlongercorollatubes(9-12mmvs.3.5-7mm),and
longerpappusbristles(longestbristles20-25mmvs.16-18mm).Rangesofthe
twovarietiesareseparatedbyabout340km.PutativehybridsbetweenC.
mamoenumandC.eatoniivarviperinumareknownfromtheSnakeRange.
CirsiumeduleNutt.varmacounii(Greene)D.J.Keil,comb,etstat.novBasionym:
CarduusmacoumiGreene,OttawaNaturalist16:38.1902.Typf;BRITISHCOLUM-
BIA:ChilliwackValley,13Jun1901,Macoun26'^51(lectotypl,herechosenfrom
syntypes,NDG).
CirsiumeduleNutt.var.wenatchenseD.J.Keil,varnovTypf:WASHINGTON.
ChelanCo.:NasonCreek,banksofstreams,660m,30Jul1893,Sandbergand
Leihgerg626(holotype:ORE;isotypes:CAS,UC).
Capitulisgeneralitersolitariis,inpedunculis10-30cmportatisetin\'olucris3-4cmaltiset4-5cm
diametrodistinguatur.
Capitulamostlysolitary;peduncles10-30cm.Involucres3-4cm,4-5cmdiam.,
moderatelyarachnoid;phyllaryapiceslong-acicular,widelyspreading,spine
tips5-15mm.Corollas29-33mm,thetube9-11mm,thethroat8-12mm,the
lobes9-10mm.Styletips3-4mm.Cypselae4.5-6mm,darkbrown;longest
pappusbristles20-25mm.
Paratvpks;WASHINGTON.ChelanCo.:WenatcheeMts.,trailtoSnowLakes,rockygraniticalpine
slopes,11Augf952,4000ft,7hompson]7i35(WTU);YakimaRegion,1883,Brandegee922(UC);Nason
City2000-3000ft.,Jul1893,SandhergandLeihergs.yr(MIN);YakimaRegion,Wenatchie.Aug188.3,
6500ft.rweetJy.s.n-(YUj.KittitasCo.:Mt.Stuart,Cascades.Augf898.Elmer.12(7(MIN).
Cirsiumedulevai.wenatchenseisapparentlyendemictotheWenatcheeMoun-
tains,intheeasternCascadeRangeofcentralWashington.Littleisknownof
itsvariationorhabitat.Becausenoneofthespecimensarecompleteindividu-
als,theoverallstatureoftheplantisunknown.1amnotawareolanyrecent
collectionsofthistaxon.
214 BRIT.ORG/SIDA21(1)
Cirsium horridulumMichx.v^\r.megacanthum(Nutt.)D.J.Keil,comb,etstat.
nov.Basionym:CirsiummegacanthumNutt.,Trans.Amer.Phil.Soc.2ndsen,7:421.
1841.Typh:LOUISIANA:banksofMississippinearNewOrleans,Littles.n.(lf.cto-
TYPE,herechosenfromsyntypes,PH).
Cirsiuminamoenum(Greene)D.J.Keil,comb.nov.Basionym:Carduusinamoenus
E.L.GreeneFl.Fran.479.1897;CarduusundulatusNutt.varnevadensisGreene,
Proc.Acad.Nat.Sci.Philad.44(1892):361.1893(homotypicsynonymofC.
mamoenus).Typr:CALIFORNIA:EofTruckeeornearthatplace,Aug1883,Grccfif
s.n.(lectc)TYPH,herechosenfromsyntypes,ND-G).
Cirsiuminamoenum(Greene)DJ.Keilvar.davisii(Cronquist)DJ.Keil,comb,et
Stat,novBasionym:CirsiumdavisiiCronquist,Leafl.W.Bot.6:46,1950.Type:
IDAHO.BannockCo.:UniversityofIdahoi^arm,Pocatello,8Jun1931,Daviss.n.
(holotype:WS).
Cirsiumoccidentale(Nutt.)Jeps.var.lucianumDJ.Keil,var.nov.Type:CALIFOR-
WNIAo.lSEanboLuunidsaOrbyisspiognCof.o:rCCuueessttaaRRiiddgeg,eaBlootnagnimcaalmArreoaa,dc(a.US4.P3SmRid.W29oSf1LH3w)y1,310m1i,
scatteredalongdisturbedroadside,serpentine,2460ft,22Jun1989,Juna/?andAyers
4073(holotype:SBBG;isotype:OBI).
Habituerecto,capitulisconspicuepedunculatis,involucris2-4cmdiametro,atropurpureis.floccoso-
arachnoideis,etcorollaspurpureo-atrorubens20-24mmdistinguatur
Plantserect,3-20dm.Capitulalong-peduncled,elevatedwellabovelowerle^wes;
involucres2-4cmdiameter,darkpurple,floccose-arachnoid;middlephyllary
tipsgenerally5~8mm,f-3mmwide,straigbtorupwardlycurved,ascendmg
tospreading.Corollas20-24mtrt,darkpurplishred.
bPaerlaotwypTesV:lCoAwLeIrFsOjuRsNtIEA.olSWaenstLCuiusesOtbaiRsipdogeCoR.d:,Laot.seaPsatderrensbNoattainoincaallaFroeraesbto,uWnddaarc)i'nsgigcnl.ecaar.e3dmsiLiWpe
ofHwy101,22Jun1989,Ajyer.siiudjunak75-iA(OBI,SBBGJ;0..3miiromhwy101alongWestCuesta
RidgeRd,.chaparral,Slacingslope,roadside,ca,1540ft.4jun2000,BushakraandBu^hahra35
(rOiBdIg)e,W36o(l.UuC)p,pe37r(LMoIpNezAC,5a8n(yAoSnU,J.opCeenrrWo-Aflatcoi,n1g3sMloapye,1595J8u,nH1a9r6d4,haHmoo3v2e8r88(9S3B2BG()O:BIn):earridsguemNmiWtooff
CuestaPass,serpentinearea,disturbedroadside,17Jun1965,Hc)over9430(OBI);BlackButteResearch
NaturalArea,SEbaseolPeak3,steeprockyopeningatedgeolQuercusagnfoliawoodlandatmargin
ofunburnedarea,2May1995.HMcsaJ2258(CDA,OBIJ:CuestaRidgeBotanicalArea,Nofmainroad,
JustSofKSBYtelevisionlowers,ca,2,7miVv'ofHwy101,28Jun1989,/imal;andAyers4098{SBBGG);
CuestaEastinareaburnedbyGayMt.andLasPilitasfires,firesuccessionareainchaparral,17May
1b9e8l7o,wKesiulmmetiatl.o2f0T16V2p(eOaBkI.J;WSanofLCuuiessOtbaisGproa,de18a7nd,LHewmymo10n1.4J28(PJHu)n;a19l6o8n,gSrmoitaldit1o0c0y9p5re(sSsBBfGo)r;esSt
entrancetoCuestaBotanicalArea,serpentinesoil,7Jun1971,2500ft,Wwic97(OBI),
arsiumoccidentalevar.lucianumisrestrictedtoSanLuisObispoCounty,Cali-
forniawbereitoccupiesanarrowcorridoralongandadjacenttothemainridge
otthesouthernSantaLuciaMountains.InThejepsonManualKeilandTurner
(1993)treatedtheseplantsasanatypicalraceofC.occidentalevar.calijornicum.
Theyresemblesmall-headedplantsofvar.calijornicumbutdifferintheirdark,
reddishpurplecorollasanddarklypigmentedinvolucres.Theyapproachthe
KEIL,NEWTAXAANDNEWCOMBINATIONSINNORTHAMERICANCIRSIUM 215
rangeso(var.californicumandvar.venustumbutarenotknowntogrowwith
eitherofthesevarieties.Wells(1983)andKeilandTurner(1992)reportedthe
occurrenceofhybridswarmsbetweenthesetaxainwhichemergentpheno-
typesdifferingfromeitherparenthavebeenobserved.Cirsiumoccidentalcvar.
lucianummayrepresentastabilizedemergentformderivedbyprehistorichy-
bridizationbetweenvar.californicumandvarvenustum.
CirsiumochrocentrumA.Grayvarmartinii(P.Barlow-lrick)D.J.Keil,comb,et
Nstoavto.nno9v:3B2A0.Si1O9N99Y.MiTCyipfr:siNumEoWchrMoEtX'IeCnOtr.uCmaAt.rGornayCos.u:bsNp.MmHarwtiyni1i2,P.5BamriioEw-oIfriScakn,
FranciscoRiverbridgeatReserve,5800ft.elev,28Jul1997,Barlow97-8(holo-
TYPH:US;ISOTYPES:ASU,MO,NMC,UNM).
Hsi(1960)manunpublisheddissertationproposedrecognitionofthistaxonas
aspecies,notingitsresemblancetoCochrocentrum.Bariow-Irick(1999)for-
mallydescribeditasasubspeciesofCochrocentrum,demonstratingitsin-
tergradationwithtypicalC.ochrocentrum.AsindicatedaboveIhavechosento
recognizevarietiesratherthansubspeciesinmyFNAtreatmentofCirsium,
andImakethechangefromsubspeciestovarietymdeferencetotheFNAGuide
torContributors.
Cirsiumpulcherrimum(Rydb.)K.Schum.var.aridum(R.D.Dorn)DJ.Keil,comb,
etStat,novBasionym:CirsiumaridumR.D.Dorn,Vase.Pi.Wyoming,2nded.304.
1992.Type:WYOMING.FremontCo.:ca.6miNofSweetwaterStation,barren
chalkyhills,6700ft,26Jul1990,Dorn5105(holotype:RM).
CirsiumscariosumNutt.varamericanum(A.Gray)DJ.Keil,comb.nov.Basionym:
CirsiumacauleAUionivaramericanumA.Gray,Proc.Philad.Acad.Nat.Sci.
1863:68.1863.Type:COLORADO:RockyMts.,lat,39-41,HallandHarbour.339(ho-
L0TYPE:GH).
Cirsiumscariosum Nutt.varcitrinum(Petrak)DJ.Keil,comb,novBasionym:
Cirsiumquercetorum(A.Gray)Jeps.varcitrinumPetrak,Bieh.Bot.Centralbl.
35:363.1917.TYPE:CALIFORNIA.SanDiegoCo.:Warner'sHotSprings,10Apr1913,
Eastwood2629(holotypi::CAS).
CirsiumscariosumNutt.varcoloradense(Rydb.)DJ.Keil,comb,etStat.nov.
Basionym:CarduuscoloradensisRydb,,Bull.TorreyBot.Club32:132.1905.Type:
COLORADO:PagosaSprmgs,28Jul1899,Baker644(holotype:NY;isotypes:MO,
ND-G,POM).
CirsiumscariosumNutt.varcongdouii(RJ.Moore&Frankton)DJ,Keil,comb,
e4t5:S1t7a3t8,.n1o96v7,BTayspieo:nyCmA:LCIiFrOsiRiNmIiAe,onMgodonmoiCRoJ.,:MBoriodrgeep&orFtraVanlklteoyn,,3.C5anmaid.NJ,WBootf.
BridgeportalongU.S.Llwy395,edgesofwinteroverflowofBuckeyeCreek,6505
ft,Bacigalupietal8015(holotype:DAO).
Cirsium scariosum Nutt.varrobustumDJ.Keil,var.novType:CALIFORNIA.
SlskiyouCcy:NsideofStatelineRd.,Hwy161toTulelake,5.2kmEofHwy97,10.5
216 BRIT.ORG/SIDA21(1)
kmNEofDorns,T48N,RIE,sect..16,14Jun1977,FuHcr20216(HOLt)TYPi:;mounted
on3sheets,CDA;ISOTYPE:CDA).
Habitucaulescentecaulibusproximaleramosis,capitulisevidcnterpedunculatis,apicibus
phyllarioruminteriorumexpansisscariosiserosodeniatis.elcorollasalbidisdistinguaiur.
Plantscaulescent,2.5-7dm.Stemssolitaryfrombase,oftenverystout,proxi-
mallysimpleanddistallybranchedorbranchedthroughout,leafy,glabrous,
villouswithseptatetrichomes,orarachnoid-tomentose.Leavesoblanceolate
toelliptic,deeplypuinatelylobed,villouswithseptatetrichomesorthinly
arachnoid-tomentose,abaxiallythinlytodenselyarachnoid-tomentose,villous
withseptatetrichomesalongthemidvein.Capitula3-many,evidentlype-
duncledatstemtips(inageclusteredaxillarycapitulasometimesdeveloping),
subtendedand±overtoppedbyupperleavesorthese±reduced.Involucres
2.5-4cm;outerandmiddlephyllarieslanceolatetoovate,thespinetips1-6
mm,slenderto±broadandflat;tipsofinnerphyllarieslinear-acummatcor
morecommonlyexpandedasascarious,erose-toothedappendage,oitencon-
torted.Corollaswhite,30-36mm,thetubesf4-22mm,thethroats7-12mm,
thelobes5-10mm;styletips6-8mm.Cypselae4-6.5mm;pappi22-32mm.
VP.a-l\lRAeTyYPWEs:oCtADLoIrFnOs,RNoIecAu.rrSiitsigkiiynoluarCgoe.:paGatzcehlelse,,1m6oJiulst19m8e2,a5dlio(w/svv,;>2.)i0,(JCuDnA)1Q;05E,oHfeMlcleCrlo8i0i7d7a(lPoH)n;gBHuwtyte.
89,7-7miVVotShastaCo.line,roadsidedrainageinlodgepolcpinelorest.24Jull'-)Q2,KcilandKcil
2,3571(OBU;5miNoiGazelle,HJul1948.Prv()r.3704(CDA);4miNotGazelle,26Jul1950,Pryois.n.
(CDA);withoutlocality,22Jul1954,Huse,s.n.(CDA);Hwy.99,2miSofGazelle.T42N.R6W,roadside
meadow,9Jul1959,Fuller2738(CDA).OREGON.KlamathCo.:Keiio,bankofKlamathRiven(\]ul
1920,Peck9279(D.S);l"!miSolDiamondLake,moistground,7Jun1937,1'cck197.56(WILLU).Lake
Co.:9miSolSummerLakeP.O.,swampymeadow,19Jun1941,Peck20809(UC,WILLU);alongjohnson
Creek,SummerLake,12Jul1928,Con.sI(;nce9478(ORE);6miNEofLakeview,29Junl937,PecH9624
(WILLU).
Cirsiumscariosumvar. rohustum isknownonlyfrom northernCalifornia
(.SiskiyouCounty)andsouth-centralOregon(KlamathandLakecounties).
CirsiumscariosumNutt.varloiyabensef").J.Keil,varnovTypH:NEVADA.Lani^lr
Co.:NofToiyabeRange,SilverCreek,10.5miEofNevada8A,12airlinemiNof
Austin,T21N,R44E,Sect.22,edgeofwetmeadow,17Jul1970,7000It,Holmii^rcn
andHolmgren4476(ilOLOiVPE:MIN;lsotypf.S:ASU,WTLJ).
1labitucaulescentecorollaspurpureislobis"15-8mmlongisdistinguatur.
Plantssubacaulesccnttoerect,0.5-5.5dm.Stemsusuallysimple,leafy,glabrous
tovillousortomentoscwithseptatetrichomes,sometimesthinlyarachnoid
tomentose.Leavesoblongtooblanceolateorelliptic,pinnatelylobedorolten
unlobed,adaxiallygreen,glabrousorvillouswithseptatetrichomes,abaxially
greenand±glabroustowhitc-tomcntose.Capitula1-10+,sessileorshort-pe-
duncled,crowdedatstemtips,usuallysubtendedand±overtoppedbyupper
leaves.Involucres2-3cm;outerandmiddlephyllarieslanceolatetoovate,the
spinetips2-4mm,slender;tipsofinnerphyllariesacuminateandentireor