A NEW SPECIES OF PTILIMNIUM (APIACEAE) FROM THE ATLANTIC COAST

2004·3.8 MB·English

by  Alan S Weakley

ANEWSPECIESOFPTILIMNIUM(APIACEAE)
FROMTHEATLANTICCOAST
AlanS.Weakley GuyL.Nesom
UniversityofNorthCarolinaIHerbariunn(NCU) BotanicalResearchInstituteofTexas(BRIT)
NorthCarolinaBotanicalGarden 509PecanStreet
CokerHall,CB3280 FortWorth,Texas76102-4060,U.S.A.
UniversityofNorthCarolina
ChapelHill,NorthCarolina27599-3280,U.S.A.
ABSTRACT
PtilimniumahlesiiWeakley&Nesom,sp.nov.,isdescribed[romlocalitiesineightcountiesofNorth
Carolina,SouthCarolina,andGeorgiaintidalfreshwatermarsheswithin20milesoftheAtlantic
coast.ThespecieshasbeenpreviouslyidentifiedasPcapillaceumbutdiffersfromthelatterinits
conspicuouslylargeovariesandfruits,whichatmaturityare2.7-4.2mmlongand1,8-2.2mmwide
(vs1.4-2.5mmlongand0.8-1.2mmwideinP.capillaceum),initsless-dividedleaves,andinitsin-
florescenceswithfewerumbelletsperumbelandfewerflowersperumbellet.Thenewspeciesalso
flowersandfruitsearlierthanPcapillaceum(May-earlyJune,lateMay-July,vs.midJune-earlyAu-
gust,July-September,respectively).Ptilinmiumahlesiiisrareandrestrictedtofreshwaterorslightly
brackishmarshes,whilePcapillaceumhasawidergeographicrangeandecologicaltolerance,but
thetwospeciesgrowincloseproximityintheirareaofsympatry.
RESUMEN
SedescribePtilimniumahlesiiWeakley&Nesom,sp.nov,devariaslocalidadesenochocondadosde
Carolinadelnorte,CarolinadelSuryGeorgiaenhordesdecharcasdeaguadulceenunabandade20
dmiiflilearseedenelsatcaopsotralaotslaonvtairciao.sLyafersupteocsicelhaarasmiednoteprmeavsiagmreanntdeesi,dqeunteiefniclaadmaacdoumroezP.tciaepnienll2a.c7e-4u,m2pmermo
delongitudy1.8-2.2mmdeanchura(contra1,4-2.5mmdelongitudy0.8-1.2mmdeanchuraenP.
capillaceum),porsushojasmenosdivididas,yporsusinflorescenciasconmenosumbelulaspor
umbelaymenosfloresporumbelula.LanuevaespecietambienfforeceantesquePcapillaceum
(principiosdemayo-Junio,finalesdemayo-julio,vs.mediadosdejumo-principiosdeagosto,julio-
septiembre,respectivamente).Plilimniumahlesiiesrarayrestringidaacharcasdeaguadulceo
ligeramentesalobres,mientrasquePcapillaceumtieneunrangogeograficoytoleranciaecologica
masamplia,perolasdosespeciescrecenmuyproximasensuareadesimpatria.
Anexceptionallylarge-fruitedformofPtilimniumhasbeenobservedandcol-
lectedincoastalcountiesofNorthCarolina,SouthCarolina,andGeorgia(Figs.
1,2,and3).ThisplantapparentlydrewtheattentionofthelateHarryE.Ahles,
thenworkingattheUniversityofNorthCarolinaHerbarium(NCU),andspeci-
mensatNCUincludeannotationsinhishandindicatinghisopinionthatthey
representedanundescribedspeciestobenamed"Pmacrospermum.'"IntheCaro-
linas"Guide"(Radfordetal.1964),theseplantswereincludedinthekeyand
speciessummariesas"PtilimniummacrospermumAhles."Thefollowingyear,
itwasmappedintheCarolinas"Atlas"inBrunswick,NewHanover,andPender
SIDA21(2):743-752.2004
744 BRIT.ORG/SIDA21(2)
counties,NorthCarolina,andGeorgetownandJaspercounties,SouthCarohna
(Radfordetal.1965).Threeyearslater,inthefirstprintingoftheCarolinas
"Manual,"thisentitywasplacedasasynonymofP.capillaceum(Michx.)Raf.
withthenotationthatthenamewas"anomennudum"attributedtoAhles,but
wasconfusinglyincludedinthekeyas"P.macrospermum"(Bell1968,inRad-
fordetal.);inlaterprintingsitwasalsoremovedfromthekey,presumablycon-
firmingtheintentofBellnottoincludethetaxon.Bell(pers.comm.,June2004)
statesthathewasuncertainofthetaxonomicdistinctivenessofthetaxon,be-
causeofthepaucityoiherbariummaterialavailable,andasithadnotbeen
validlypublishedhedecidednottoincludeitinthe"Manual"(seeSorrie1997
forexamplesolotherunpublishednamesbyAhles).
Thenameinsimilarformhasresurtacedinawidelyuseddatabase(Kartesz
1999),as"PtilimniummacrospermumKartesz,sp.novined."Additionally,the
taxonhasbeenconsideredtobeofconservationconcernbythenetworkof
NaturalHeritageProgramsandhasbeentrackedasararespeciesunderthe
unofficialname"Ptilimniumsp.FinNorthCarolinaandGeorgia(Franklin
2004,GeorgiaNaturalHeritageProgram2004).Thepotentialconservation
importanceoftheputativetaxondemandsthatthistaxonomicandnomencla-
turalconfusionberesolved,andthespecieshasbeenthesubjectoffieldand
herbariumstudybytheauthorssince1990.Withtheseadditionalobservations,
weconcludethatthisplantisadistinctiveandundescribedspecies.Weare
pleasedtoprovideitwithavalidname,allowingittoproceedinpolitecom-
pany,ratherthanunderatwice-nakednameCPtilimniummacrospermum"),a
non-name("Ptilimniumsp.1"),orinappropriatelyignoredunderPtilimnium
capillaceum(Michx.)Raf.Thenewepithetvalidatedherehonorstheoriginal
insightofHarryE.Ahlesregardmgthis,aswellashisgeneralcontributionsto
theunderstandingofthefloraoftheCarolinasthroughhisvoraciouscollecting.
PtilimniumahlesiiWeakley&Nesom,sp.nov.(Figs.1-3).J\ni:NORTHCAROLINA,
BrunswickCo.:tidalIreshwatermarsholtheBrunswickRiver,justV.oftheBrunswickRiver
andjustNoltheUS74-76causeway,ca.2miVv'ofWihuington,growingwithTypha
angustiJolia,Carexhyalinolepis,Fonledenciiordata,PcnUindravirginica,SagiUarialanciJolia,
RumexverliciUatus.andraxodiumdistichum,34'14'04.7"N,77"5Q'05.7"(NAD83),ManS.
Wcakky7317andRichard].I.eBlond,lOJun2004(holotypk:NCU,iS0TYi>iiS:BRIT,CLEMS,
Gl\,MO,NY,TEX,UGA,US,USCH.
Ptilimniocapillaceo(Mich.\,)Ral.siniihsbracteisinvolucraiibusdivisis,ramisbrevibusstyli,ctcostis
l5-a8t(e-r]a3h)bupserIruumcbteulslvaaml,defelvoorliubtuiss,5~s8edpedritulemrbtelloliiuilsadim,viIsriuocntiibbuuss2m-a3jo(vreirbsuuss(\2).7p-e4r.n2omdmumi,oungmibselvleurlsiuss
1.4-2.5mm),geographicaachabitationemultomagisrestrictis,ettemporepraecociilorendiac
fructilicandi.
Annual,glabrous,caulescentherbsfromashortsystemoffibrousroots,the
stemserect,4-13dmtall,unbranchedorinvigorousindividualsbranching
abovemid-stem.Lowerstemleaves(oftenwitheredbyanthesis)phyllodial(re-
WEAKLEYANDNESOM,ANEWSPECIESOFPTILIMNIUM 745
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Fig.1.HolotypeofPtilimniumablesii.Notethenearlyphyllodiallowerstemleaves,thebroadenedsegmentsofthemid-
caulineleaves,andthelargefruits(notfullymature).
746 BRIT.ORG/SIDA21(2)
1cm 2 3 4 5 6
I
Fig.2.InflorescenceofPtilmniumahlesii(fromanisotype).Notethelargefruits,thedividedinvolucralbracts,andthe
fewumbellets/umbelandfewflowers/umbellet(ascomparedtoP.capillaceum)
ducedtothehollowrachisandlackinglateralsegments,thuscloselyresem-
pbihnnngatteh,etqhueilsle-glmeeavnetsso(fwPh.efnluvpiraetsiekn,t)P.Fnlaotdtoesnuedn,%taon2dmP.mviwviidpea;rummi)d-toasnndnpulpy-
per-stemleavespinnatelydecompound,3-21cmlong,thesegmentsliliiormto
distinctlytlattened,with2-3divisionsatprimarynodesontherachis;petioles
3-25mmlong,broadlywinged,ca5-ribbed.Inflorescenceumbellate,distinctly
andirregularlyflat-topped(theraysvariableinlength),once-compound,the
umbelsterminalandaxillaryontheupperhalfofthestem,peduncles5-14cm
WEAKLEYANDNESOM,ANEWSPECIESOFPTILIMNIUM 747
1cm
Fig.3.Fruits(nearlybutnotfullymature)oiPtilimniumahlesii,iromanisotype.
748 BRIT.ORG/SIDA21(2)
long;primaryrays5-13,5-35mmlong,pedicels(2-)4-10,(3-)5-10mmlong;
involucreoflinearbracts,entireor3-parted;involuceloflinearbracts.Calyx
teethdeltoid,acutetosubacuminate,persistent;petalswhite,obovate,0.6-0.8
mmlong;stylopodiumconic,thestylesmostlyslightlyshorterthanthesty-
lopodium,ascendingtoerect-ascendingorwidelyspreading;antherslightpur-
plish.Carpophorebifidattheapex.Fruitelliptic-ovoid,slightlycompressed
laterally,2.7-3.5mmlong,1.8-2.2mmwide,themericarpslooselyconnateand
usuallyseparatedatmaturity;dorsalribs3,narrowandsharp-edged,thelat-
eralribssimilarbutwithabroad,rounded,corky-thickenedextension,theex-
tensionsoftheadjacentmericarpsconnateandformingabroad,conspicuous
bandaroundthefruit;oiltubesdarkrusty-brown.Intidalfreshwatermarshes;
floweringMaytoearlyJune,fruitinglateMaytoJuly.
Collectionsexamined.UNITEDSTATES.GEORGIA,ChathamCo.:SavannahNationalWildliicRef-
uge,alongHwy17,fresh-wateririarsh,14Jul1966.Bozcman6100(NCU,distributedas"PtiUmnium
macwspermumAhles").NORTHCAROLINA.BrunswickCo.:Wilmington,Jun18^^)8,A.sfu'S.n.(NCU).
NewHanoverCo.:1miNofUSUonWnghtsboroRoad,marsh,2Jun1949,Radford4,3.5.3(NCU);
brackishmarshontheCapeFearRiveronUS17,29Jun1963,Ahks58397(NCU)PenderCo.:swamp
SonOUNoTrHtheCaAsRtOCLaIpNeA.FeBacrauRfiovreirnCeoa.r:R0.o5cmkyiPWomotf-CHoa.mRposatdea11d1RoonadC,o.fJRuonad195330,,cRaatd-ftaoirlds5w14a2m(pNCnUe)a.r
Trichinhami'laniation,27Jun1956,Bell.1767(NCU).BerkeleyCo.:CooperRiveratthemouthof
DurhamCreek,lowsalinity,7Jun1990,McAninch2.3(NCU).Colle«onCo.:1.4miNEofAshepooon
USHwy17,swamp,26Jun1956,Bell3703(NCU).GeorgetownCo.:tidalmarshofBaruchPlantation,
20Jun1967,Barry103(USCH);BaruchPlantation,fresh-watermarshaffectedbytidalaction,17Jun
(1WmU9i6S9oCW,fFlGB)oea;fotoIrscnogctneartas.osci.wono.anns(,atUlafSlirCneWHss)ahu;t-nwenBaryatwersarupycomhta1sr6PsilMnhaa,nwyte1at31t9Jis9ouw0nn,a,1mNs9epl5l7oys,uowgRnohao9ddi0fns2o,4rfdWr(e2Us5Sshi1Cwd3Hae9)to,e(frNSSCamoUmau)prt.ishthI,Rsil2av7nedrJuRnnoeaa1rd96U(9SS,-11B87a)At.asbo9o,n5utsm.in4.
Relationshipswithinthegenus
PtiUmniumisarelativelysmallgenuscenteredinthesoutheasternandsouth-
centralUnitedStates.Thefivetosevenspeciesareseparableintosubgroupson
thebasisofdifferencesinmorphologyandchromosomenumber,althoughap-
parentdysploiddifferencesinchromosomenumbersuggestthatthesituation
maybemorecomplexthanthecurrentassessmentallows.Plantswithleaves
reducedtoawingedpetioleandextendedrachis(withlateralsegmentssup-
pressed,thestructuressometimesreferredtoas"phyllodia")arediploidsbased
onx=6(2n=12;Easterly1957;Bell&Constance1960).Theseplantshavebeen
regardedascomprisingfromonetothreespecies,P.nodosum(Rose)Mathias,R
viviparum(Rose)Mathias,andP/luviatile(Rose)Mathias)(Krai1981;Kressetal.
1994;Bartgis1997);theappropriatetaxonomicdispositionremainsunclearand
isunderadditionalstudyPti/imniumcostatum(Ell.)Raf.hasdistinctlypetiolate
leaveswithshort,crowded,verticillatesegments,longstyles,andatetraploid
chromosomecomplementbasedonx=8(In=32;Easterly1957).Onepopulation
ofP.costatumfromIllinoishasbeencountedas2n=22(Bell&Constance1960),
thisperhapsatetraploid(x=6,2n=24)havinglostonepairofchromosomes.
WEAKLEYANDNESOM,ANEWSPECIESOEPTILIMNIUM 749
Ptilimniumahlesii,P.capillaceum,andP.nuttalliiaresimilarintheirrela-
tivelyshortstyles(shorterthanorequallingthestylopodia)andprimarilydip-
loidchromosomenumberbasedonx=7(2n=14;Easterly1957,BellandCon-
stance1960).Tetraploidplants(2n=28)ofP.capillaceumwerereportedby
Easterly(1957)fromcentralGeorgia(UnadillaCounty).Thesewerefoundap-
parentlyintermixedwithdiploidplantsofthesamespeciesandwithplantsof
P.nodosum"nearbymthesameditch."Easterlydidnotcommentonanymor-
phologicaldifferencebetween theconspecificdiploidsandtetraploids.One
populationofP.capillaceumfromFloridahasbeencountedas2n=16(Con-
stanceetal.1976).ItisnotablethatthebasalandlowerstemleavesofP.ahlesii,
producedinthespringandtypicallywitheringpriortofloweringandfruiting,
havefewtonoleaflets,andthereforecloselyresemblethereduced"quillleaves"
ofthe"Harperella"groupofPtilimnium(P.Jluviatile,P.nodosum,andP
viviparum).ThissupportsthebroadercircmnscriptionofPtilimniumasin-
cludingHarperellaRosethathasbeengenerallyfollowedinrecentdecades.
PtilimniumtexenseCoulter&Rosewasoriginallydescribedasaspecies
(Coulter&Rose1909)butlater(Mathias&Constance1944-45;Eastedy1957)
interpretedtobeofhybridoriginandofintermediatemorphologybetweenP
capillaceumandPnuttallii.Thisinterpretationhasbeenacceptedandrepeated
bysubsequentworkers(e.g.,Correll&Johnston1970;Correll&Correll1972),
althoughMathias&Constance(1961)placeditinsynonymy,withoutcomment,
underPcostatum.StudyofPtilimniumcollectionsatLL,TEXshowsonlya
singleentityinTexas(interpretedhereasPcostatum)otherthanPcapillaceum
andPnuttallii.
InthefreshwatertidalmarsheswestofWilmington,NorthCarolina,three
speciesofPtilimniumgrowincloseproximitytooneanother;Ptilimnium
ahlesii,P.capillaceum,andPcostatum.Theyaremorphologicallydistinctive;
nointermediateshavebeenseen.Observationsoverthepastdecadeturther
suggestthatthefloweringtimesofthethreespeciesarelargelyorcompletely
non-overlapping,withPahlesiifloweringfirst(Maytoearlyjune),followedby
Pcapillaceum(mid-JunetoeadyAugust),andthenPcostatum(lateJulyto
August).Thephenologicseparationprovidesadditionalevidencecorroborat-
ingthestatusofPahlesiiasadistincttaxonandnotmerelyalarge-fruited
formofPcapillaceum.
Ofitsclosestrelatives,Ptilimniumahle.siiismostsimilartoPcapiHaceum,
particularlyinitsdividedinvolucralbracts,minutecalyxteeth,dorsalfruit
ribsnarrowerthantheintervals,andlateralribswithexpanded,corkymar-
ginsencirclingthefruitlikeaband(Pcostatumalsosharesthedistinctivefruit
morphology).Thetwospeciesareseparatedbythefollowingcontrasts(andsee
Figs.1and2).
1. Maturefruitelliptic-ovate,2.7-4.2mmlong,1.8-2.2mmwide;umbelsflat-topped,
irregular,theraysvariableinlengtii;umbellets5-8(-l3)perumbel,therays5-35
750 BRIT.ORG/SIDA21(2)
runswicki^
Fig.4.Mapofdocumentedcountydistribution(in,fromnorthtosouth,NorthCarolina,SouthCarolina,andGeorgia)of
Ptilimniumahlesii.
mmlong;flowers5-8perumbellet;petals0.6-0.8mmlong;leafsegmentsofmid-
stemleaves15-30(-40),capillarytolinear,0.5-2.0mmwide;floweringMay-early
June,fruitinglateMay-July;tidalfreshwatermarshes,rareinNorthCarolina,South
Carolina,andGeorgia Ptilimniumahlesii
1. Maturefruitovatetoorbicular,1.4-2.5mmlong,0.8-1.2mmwide;umbelsgener-
p(ale5lt-ya)lr9s-o2uO.n1.dp3ee-0rd.6utmombehmle,miltsohnpegh;rearlyiescaf5,-st2he1egmmreamnystlsgoenongfe;rfmalilodlwy-ersnsteeaumrsulaylleelayqvue1aslo5ri0nmolorernemgotprhe;er,uumcmabbpeeillllllaeerttys;,
usuallylessthan0.5mmwide(exceptinsubmersedleaves);floweringJune-Au-
gust,fruitingJuly-September;ditches,marshes,otherwetplaces;commonandwide-
spreadineasternNorthAmerica,rangingfromMassachussettsandNewYorkto
MissouriandKansas,southtoFloridaandTexas Ptilimniumcapillaceum
Descriptionsbyvariousauthors(e.g.,Fernald1950;MathiasandConstance1944-
45;Easterly1957)ofP.capillaceumwithfruitlongerthan3mmalmostcertainly
iofncNlCudUedcoolblseecrtviaotnisoonfsPo.fahPl.eashiilesaisi.PAcnanpoitlaltaicoenusmbaypbpoetahrCtooncsotnafnicremathnids.EAassstoecril-y
atedwiththesmallernumberofumbelletsandflowersbornebyPahlesii,as
notedinthekey,plantsproduceabout45-70flowers/fruitsperinflorescence,
comparedtothoseoiPcapillaceum,whichusuallyproduceover100.Depauper-
ateplantsofPcapillaceum,however,canhavereducedinflorescences.
.
WEAKLEYANDNESOM,ANEWSPECIESOFPTILIMNIUM 751
Itappearsthatthisspeciesisnarrowlydistributedecologicallyandgeo-
graphically,totidalmarshesassociatedwiththetidalreachesofriversinsouth-
easternNorthCarolina,SouthCarolina,andeasternGeorgia(Fig.4).
ACKNOWLEDGMF.NTS
WethankthestaffsofLL-TEX.NCU,andUSforhelpwithherbariumstudies
attheirinstitutionsandstaffofUSCUforaloanofspecimens.RichardLcBlond
(NorthCarolinaNaturalHeritageProgram)assistedwithfieldstudies.Ronald
L.Hartmanandananonymousreviewerprovidedhelplulsuggestions.
REFHRHNCES
B,ARTGis,R.L.1997.ThedistributionoftheendangeredplantPtilimniumnodosum(Rose)
Mathias(Apiaceae)inthePotomacRiverdrainage,Castanea62:55-59,
Bill,C.R.1968.Apiaceae.In:A.E.Radford,H,E.Ahles,andC.R.Bell.1968,Manualofthevas-
cularfloraoftheCarolinas.Univ.NorthCarolinaPress,ChapelHill,Pp.761-789.
Bell,C.R.andLConstance.1960,ChromosonnenumbersinUmbelliferae.II.Amer.J.Bot.
47:24-32.
Constance,L.,T.d.CHUANG,andC.R.Bell1976.ChromosomenumbersinUmbelliferae.V.Amer.
J.Bot.63:608-625.
Correll,D.S.andM.C.Johnston.1970.ManualofthevascularplantsofTexas.TexasRe-
searchFoundation,Renner,Texas.
CoRRFi1,D.S.andH.B.Cokrlll.1972.AquaticandwetlandplantsofthesouthwesternUnited
States.EnvironmentalProtectionAgency,Washington,DC.
Coulter,J.M.andJ.N.Rose.1909.SupplementtothemonographoftheNorthAmerican
Umbelliferae.Contr.U.S.Natl.Herb.12:441-451
Easteriy,N.W.I957.AmorphologicalstudyofPf/7/mn/um.Brittonia9:136-145.
Fernald,M.L.1950.Gray'smanualofbotany(ed.8).AmericanBookCo.,NewYork.
Franklin,M.A.(ed.).2004.NaturalHeritageProgramlistofrareplantspeciesofNorthCaro-
lina:2004.N.C.NaturalHeritageProgram.
GeorgiaNaturalHLRirAGEProgram.2004.SpecialconcernplantspeciesinGeorgia,[http://
georgiawildlife.dnr.state.ga.us/content/specialconcernplants.asp]
Kartesz,J.T.1999.Asynonymizcdchecklistandatlaswithbiologicalattributesforthevas-
cularfloraoftheUnitedStates,Canada,andGreenland.Firstedition.in:Kartesz,J.T.,and
C.A.Meacham.l999.SynthesisoftheNorthAmericanflora,Version1.0.NorthCarolina
BotanicalGarden,ChapelHill,NorthCarolina.
Kral,R.1981.NotesonsomequilFteavedumbellifers.Sida9:124-134.
Kress,W.J.,G.D.Ma[3dox,andC.S.Rofsil.1994.Geneticvariationandprotectionprioritiesin
Ptilimniumnodosum(Apiaceae),anendangeredplantoftheeasternUnitedStates.
Conserv.Biol.8:271-276.
Mathias,M.E.andL.Constance.1944-45.Umbelliferae.N.Amer.Fl.286:43-295.
Mathias,M.E.andLConsiance.1961.Umbelliferae.In:C.LLundell,ed.FloraofTexas,Vol.3.
TexasResearchFoundation,Renner,Texas.Pp.263-329.
752 BRIT.ORG/S1DA21(2)
Radford,A.E.,H.E,Ahles,andC.R.Beu.1964.GuidetothevascularfloraoftheCarolinas.The
BookExchange,Univ.ofNorthCarolina,ChapelHill.
Radford,A.E.,H.E.Ahles,andC.R.Bell.1965.AtlasofthevascularfloraoftheCarolinas.North
CarolinaAgric.Exp.Stat.Tech.Bull.No.165.
SoRRiE,B.A.1997.NotesonLycopuscofcer;(Lamiaceae).Castanea62:119-126.