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NEWCOMBINATIONSINNORTHAMERICAEREMOGONE (CARYOPHYLLACEAE) RonaldLHartman Richard K.Rabeler RockyMountainHerbarium UniversityofMicfiigonIHerbarium DepartmentofBotany 3600VarsityDrive UniversityofWyoming AnnArbor,Miclnigan48108-2287,US.A. Laramie,Wyoming82071-3i65,US.A. [email protected] [email protected] ABSTRACT Thefollowing18newcombinationsinlivcmogoneareproposed:Ercmogor\ecapillarisvarameriama. E.congestavar.cephaloidea,E.congcstavarcharksloncnsis,E.congestavar.crassula,E.congeslavar. glandidijera,E.congestavar.pro/i/erci,E.congestavacsimulans,E.congestavar.suhcongesta.E.congesla varsuffrutescens,E.congestavarwheelerensis,E.eastwDodiaevar.adenophora,E.franklinii,E. frankliniivar.Ihompsonii,E.kingiivarplateauensii,,E.kingiivar.rosea,E.macradeniavararcuifolia, E.macradenhivacjerrisiae.andE.macradeniavarkuschei. RFSUMHN Seproponenlassiguientesdieciochocombniacionesnuc\'asenEremogone:Eremogonecapdlarisvar americana.E.congestavarcephaloidea,E.congestavarcharlestonensis,E.congestavarcrassula,E. congestavar.glandulifera,E.congeslavarprolilera./:.congestavar.simulans,E.congestavar. subcongesta,E.congestavar.sitfjndescens.E.congeslavacwheelerensis,E.eastwoodiaevar.adenophora, E.jranklinii.E.jrankliniivar.thompsonu.E.kmgiivarplaieauensis,E.kingiivar.rosea.E.macradenia vanarcuifolia.E.macradeniavarfernsiac.yE.macradeniavarkuschei. DuringpreparationofthetreatmentoftheCaryophyllaceaeforFloraNorth America(RabelerandHartman,editors,mprep),thedecisionwasmadetorec- ognizethegenusEremogoneFenzl(=ArenariasubgeneraEremogone(Fenzl) FenzlandEremogoneastrumF.WilliamssensuMcNeill1962). NepokroeffandWagner(pers.comm.;seealsoNepokroeffetal.2001)con- ductedaDNAsurveyoftheCaryophyllaceae,usingsequencedatafrommatK andtherpsl6intron.Over79speciesconstituting38generawereincluded,with emphasisonthesubfamilyAlsinoideae.ThephylogenyillustratedArenaria (sensuMcNeill(1962),excl.MinuartiaandMoehringia)tobepolyphyletic,with twomajor,distinctclades.WhileonecladeinciudesmembersoffourofMcNeill's subgenera,includingArenariaserpyllifoliaL.,thetypeofArenaria,and the genusMoehringia,thesecondcladeincludessevenspecies(fivefromsubgenus Eremogone,threeNorthAmericanandtwoAsian,andtwoAsianmembersof subgenusEremogoneastrum).Withinthatclade,ArenariahryophyllaFern, (subg.EremogoneastrumfromAsia)clusterscloselywithA.aculeataS.Watson andA.hingii(S.Watson)M.E.Jones(subg.EremogonefromNorthAmerica), suggestingthattherelationshipsbetweenthesubgeneramayalsoneedexami- SIDA21(1):237-241.2004 238 BRIT.ORG/SIDA21(1) nation.TheresultsindicatedthattheEremogonegroup,includingbothsub- genera,isstronglysupportedasmonophyletic. EremogonewastreatedbyMcNeill(1962)asasubgenusofArenariacon- tainingeightsections(64species,ourcompilation;ca.70species(Zhengyietal. 2001))closelyalliedwithsubgenusEremogoncastrum(ca.22species).Mem- bersofsubgenusEremogonearefoundIromwesternEuropethroughAsia,east tosouthernAlaskaandnorthwesternCanada,disjuncttosouthwesternCanada andthewesternUnitedStates.ThecompositionofEremogonesensuIkonnikov (1990)excludesMcNeill'smonotypicsectionMonogoneMaxim,(theTienShan MountainsandtheAltaiRegion),twoofthespeciesfromhissectionScariosae (northernIranandTurkishArmenia),andhissect.Pungentes(twospecies,Spam andNorthAfrica). AsoutlinedbyMcNeill(1962),ArenariasubgenusEremogoneastrumis confinedtothreecenterswithtwospecies(ArenariahookenNutt.andA. JrankliniiDougl.exHook.;athird,A.pinetorumA.Nelsonisavarietyofthe former)inwesternNorthAmerica,thetype,A.festueoidesBenth.,knownfrom India,Pakistan,andChina,andtheremainder(ca.6species)Sino-Himalayan indistribution.SeventeenspeciesarenowknownfromChina(Zhengyietal. 2001);mostareSino-Himalayan,ninewerepublishedsince1979.Ikonnikov makesnomentionoiArenariasubgenusEremogoneastrumineitherofhissyn- opsesofEremogone(1973,1990). InNorthAmerica,bothMaguirc(1947,1951;treatedinMaguire(1951)as Arenariasect.PentadcnariaWilliams)andHickman(1971)recognizeda broaderconceptol ArenariasectionEremogone,includingthetwowestern American membersof subgenus Eremogoneastrum (A. hooken and A. franklinii).Maguiretreatedthosespeciesasrelatedtobutofuncertainplace- ment,whileHickmanconsideredthemtobeintermediatebetweensections EremogoneandAlsine(=Minuartia).Eremogoneaspresentedhereiscurrently acceptedinflorasofColorado(Weber&Wittmann1992andsubsequentup- datesathttp://eumuseum.colorado.edu/)andWyoming(Dorn2001). Morphologically,Eremogoneconsistsofwoodybasedperennialswitha ccspitoseormattedhabitat,filiformtosubulateleaveswithscariousbases,stiffly erectorascendingfloweringstems,andcymesthatareopentocongestedor umbellate.ThisiscontrastedwiththeremainingNorthAmerican(northof Mexico)membersof Arenaria(ca.9species),thatareannualsorperennials witheitheracaespitosehabitorascendingtotrailingstems,broader(ovateto lanceolate,sometimesnarrowlyso)leaves,andflowersoftensolitaryorinfew- 1lowered,opencymes. Ikonnikov(1973)andHolub(1974)bothnotedthatabasechromosome numberofx=11alsodistinguishedEremogonefromArenariaix=10).Ofthe approximately30speciesofEremogone,asheredefined,whichhavebeen counted,onlythreecounts(includingtwolorE.fendlerithatappearnottohave HARTMANANDRABELER,NEWCOMBINATIONSINNORTHAMERICANEREMOGONE 239 beenvouchered;thattaxonwascountedbyHartman(1971)as2n=44)arenot basedonx=11.Contrarytotheirnotion,Arenarias.s.iscytologicallymoredi- verse;whilemanycountsarebasedonx=10,countsbasedonx=11areknown intwosubgeneraaswellasfivesectionsofsubg.Arenaria(Rabeler,pers.com- pilationfromstandardchromosomeatlases). Mostofthenomenclaturalcombinationsforthe13NorthAmericanspe- ciesoiArenariatobeincludedmEremogoneareavailable(Ikonnikov1973,1974; Weberetal.1981);bycontrast,onlyafewoftheinfraspecifictaxahavetheap- propriatecombinations(Weberetal.1981;Dorn2001). ThefollowingcombinationsareneededforthegenusEremogoneinNorth AmericainanticipationoftheFloraNorthAmericatreatment(Hartman,Rabeler andUtech,inprep). Eremogonecapillaris(Poir)Fenzlvaramericana(Maguire)R.L.Hartman61 Rabeler,comb.nov.Basionym:ArenariacapillarisPoir.subsp.americanaMagu- ire,Bull.TorreyBot.Club74:41.1947.Arenariacapillarisvar.americana(Magu- ire)R.J.Davis,Madrono11:144.1951.Eremogoneamericana(Maguire)Ikonn., NovostiSist.Vyssh.Rast.11:174.1974. Eremogone congesta(Nutt.)Ikonn.var.cephaloidea(Rydb.)R.L.Hartman& Rabeler,comb.nov.Basionym:Arenarit;cephaloideaRydb.,Bull.TorreyBot.Club 39:316.1912.ArenariacongestaNutt.varcephaloidea(Rydb.)Maguire,Bull.Torrey Bot.Club74:46.1947. Eremo&goRnaebecloenrg,ecsotmab(.Nuntotv..)BaIksoinonn.ymv:arArcehnaarrlieastcoonnegnessitsa(NuMtatg.uviarre)chRa.rLl.eHsatorntemnsains Maguire,Bull.TorreyBot.Club73:326.1946. Eremogonecongesta(Nutt.)Ikonn.var.crassula(Maguire)R.L.Hartman& Rabeler,comb.nov.Basionym:ArenariacongestaNutt.varcrassulaMaguire,Bull. TorreyBot.Club7445.1947. Eremogonecongesta(Nutt.)Ikonn.varglandulifera(Maguire)R.L.Hartman& Rabeler,comb.nov.Basionym:ArenariacongestaNutt.var.glanduliferaMagu- ire,AmerMidi.Naturalist46:501.1951. Eremogonecongesta(Nutt.)Ikonn.var.prolifera(Maguire)R.L.Hartman& Rabeler,comb.nov.Basionym:ArenariacongestaNutt.varproliferaMaguire, Bull.TorreyBot.Club74:47.1947, Eremogonecongesta(Nutt.)Ikonn.varsimulans(Maguire)R.L.Hartman& Rabeler,comb.nov.Basionym:ArenariacongestaNutt.varsimulansMaguire, Bull.TorreyBot.Club74:48.1947. Eremo&gonRaebecloenrg,esctoamb(.Nutnto.v).IBkaosnionn.yvm:arArseunbacroinagfeesntdaler(Si.AW.aGtrsaoynv)arR.sLu.bcHoangretsmtaanS. Watson,Botany[FortiethParallel!5:40.1871.ArenariacongestaNutt.var subcongesta(S.Watson)S.Watson,Bot.Calif.1:69.1885.Arenariasubcongesta (S.Watson)Rydb.,Bull.TorreyBot.Club24:244.1897.ArenariahurkeiHowell,Fl. N.WAmer1:85.1897. Eremogonecongesta(Nutt.)Ikonn.var.suffrutescens(A.Gray)R.L.Hartman& Rabeler,comb.nov.Basionym:BrewerinasuffrutescensA.Gray,Proc.AmerAcad. 240 BRIT.ORG/SIDA21(1) 8:620.1873.ArenanacongestaNutt.var.sujjrutescens(A.Gray)B.L.Rob.,Proc.Amer. Acad.Arts29:295.1894.ArenariasujjrutescensCA.Gray)A.Heller,Muhlenbergia 6:96.1910. Eremogonecongesta(Nutt.)Ikonn.var.wheelerensis(Maguire)R.L.Hartman& Rabeler,comb.nov.Basionym:ArendridcongestaNutt.varwheelerensisMagu- ire,Bull.TorreyBot.Club74:48.1947. EremogHoanretmeaasntw&ooRdaibaeele(rR,ydcbo.m)bI.konnonv..vBaar.siaodneynmo:pAhroernaa(nKaeaearsntewyoo&diPaeeebRlyedsb).Rv.aLr. adenophoraKearney&rPeebles,].Wash.Acad.Scr29:475.1939. Eremogonefranklinii(Dougl.exHook.)R.L.Hartman&Rabeler,comb.nov. Basionym:ArenanafranklinnDougl.e.xHook.,Fl.BorAmer1:101.1831. Eremogonefranklinii(Dougl.exHook.)R.L.Hartman&Rabelervanthompsonii (M.Peck)R.L.Hartman&Rabeler,comb,nov\iAS\OK\u:Arenanajrankhnn Dougl.exHook,varthompsoniiM.Peck,Torreya32:149.1932{"Thompsoni''). Eremo&gonRaebeklienrg,iic(oS.mbW.atnsovo.n)BaIskioonnynm.:vaArr.epnlaantaeahu\engn\siis(S.(WMaatgsuoinr)e)M.RE..LJ.oHneasrstumbsapn. plateauensisMaguire,Bull.TorreyBot.Club74:54.1947.Arenanakingiivar. plateauensisiM-Agmxe)Reveal,GreatBasinNaturalist35:344.1975. Eremogonekingii(S.Watson)Ikonn.varrosea(Maguire)R.L.Hartman&Rabeler, comb,etstat.nov.Ba,sionym:Arenariahingii(S.Watson)M.E.Jonessubsp.rosea Maguire,Bull.TorreyBot.Club73:326.1946. Eremomgoanne&macRrabaedleenri,ac(oSm.bW.atnsovo.n)BaIskioonnny.m:vaArreanracruiiafomlaicar(aMdaegnuiiarSe.)WRa.tLs.oHnarvta-r arcuifoliaMaguire,Bull.TorreyBot.Club74:51.1947. Eremogonemacradenia(5.Watson)Ikonn.varferrisiae(Abrams)R.L.Llartman &•Rabeler,coinb.etstat.nov.Basionym:ArcnoriamacradeniaS.Watsonsubsp. ferrisiaeAbrams,111Fl.Pacil.States2:151.1944. Eremo&goRnaebemlaecr,racdoemnbi,ano(Sv.BWAaStIsOoNnY)M:IkAornenn.arviaarkkuusscchheeiiEa(sEtaws.t,w.P)rocR..LC.alHiafr.tAtcnaadn. IV.20:140.1931.ArenariamacradeniaS.Watsonvarkuschei(Eastw.)Magu- ire,Bull.TorreyBot.Club74:5L1947. ACKNOWLEDGMENTS WewishtothankKanchiGandhi(GLl)fornomenclaturaladviceandMolly Nepokroeff(SDU)andWarrenL.Wagner(US)forsharingwithusresultsof theirmolecularworkontheCaryophyllaceae. REFERENCES DoRN,R.D.200L\/a.scularplantsofWyoming.ThiirdEd.MountainWestPublishing. Hartman,R.L.1971.ChromosomenumbersinCaryophyllaceaefromWyomingandadja- centstates.Bull.TorreyBot.Club98:276-280. Hickman,J.C.]97].Arenaria,SectionEremogone,(CaryophyllaceaeinthePacificNorth- west:akeyanddiscussion.Madrono2L201-207. HoLUB,J.1974,NewnamesinPhanerogamae3.'roliaGeobot,Phytotax.9:261-275. —— HARTMANANDRABELER,NEWCOMBINATIONSINNORTHAMERICANEREMOGONE 241 HoLUB,J.1977.NewnamesinPhanerogamae6.FoliaGeobot.Phytotax.12:417-432. 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