A REVISION OF TRISETUM AND GRAPHEPHORUM (POACEAE: POOIDEAE: AVENINAE) IN NORTH AMERICA NORTH OF MÉXICO

2005·16.2 MB·English

by  Victor L Finot

AREVISIONOFTRISETUMANDGRAPHEPHORUM
(POACEAE:POOIDEAE:AVENINAE)INNORTHAMERICA
NORTHOFMEXICO
PaulM.Petersonand
VictorL.Finot RobertJ.Soreng
UniversidaddeConcepcion DepartmentofBotany
FacultaddeAgronomia NationalMuseumofNaturalHistory
DepanamentodeProduccionAnimal SmithsonianInstitution
Casilla537,Chillan,CHILE Wasliington,(DC)20013-7012,U.S.A.
[email protected] [email protected];[email protected]
FernandoO.Zuloaga
InstitutodeBotdnicaDarwinian
Labarden200
SanIsidro1642,ARGENTINA
[email protected]
ABSTRACT
AtaxonomictreatmentofJrisetumPers.andGraphcphovumDesvinNorthAmericanorthofMexico
isgiven.EightspeciesotTrisetumarerecognized.Twos|icciesareendemictotheUnitedStates:T.
orthochacium(Montana)andT.projeclum(CaHfornia,Montana,andNevada);Trisctumccrnuuni
subsp.ccrnuum,T.cernuurttsubsp.cancsccns./.montainim,T.sibiricum,T.spicatumvar.spicalum,
andIspicatunivar.pilosiglumearefoundinCanadaandtheUnitedStates;andT.spicatumvar.
spicalumand7.spicalumvar.pilosiglumearc(oundinGreenland.Twospecies,T"aurcumandT.
Jlavescens,areintroduced.Tri.setuni[lonhundumPilg.ischosenasalectotypeforTriselumsect.
AulacoaLouis-Mane.ThegenusGraphephorumcomprisesonlytwospecies:G.melicoidcsranging
fromCanadatotheUnitedStatesandG.wolfiiendemictotheUnitedStates.
RF.SUMHN
EnelpresentearticulosepresentauntratamientotaxonomicaparaTrisctumPers.yGraphephorum
Desv.enNorteamericanortedeMexico.SereconocenochoespeciesdeTrisctum.Dosdeestasespecies
sonendemicasdelosEstadosUnidos:T.orlhochaeium(Montana)yT.projcctum(Cafifornia,Mon-
tanayNevada).Trisetiimcernuumsubspccrnuum.1'ccrnuumsubsp.canescens.T.monianum,T.
sibiricum,T.spicatumvar.spicatumyT.spicalumvatpilosiglumeseencuentranenCanadaylos
EstadosUnidos;yT.spicatumvar.spicatumyTspicatumvar.pilosiglumeseencuentranen
Groenlandia.Dosespecies,T.aureumyT.jlavescens.hansidointroducidas.Tri.setiim/lor!bundum
Pilg.sceligecornolectotipoparaTri.scUimsecc..AulacoaLouis-Marie.ElgeneroGraphephorum
comprendesolodosespecies:G.melicoides.queseencuentradesdcCanadahastalosEstadosUnidos
yG.wcil/ii,endemicodelosEstadosUnidos,
Jrisetumincludesapproximately40speciesandseveralmfraspecifictaxain
theAmericasandisdistributedfromGreenland(70°N)tosouthernSouth
America(55°S)[Clebsch1960;Hulten1959;Nicora1978,Fmot2003a,Finotet
SIDA21(3):1419-1453.2005
1420 BRIT.ORG/SI0A21(3)
al.2004).TherearctwoendemicspeciesofTrisctiimmthe1lawaiianIslands:T.
;d,lo>ncraluin(KuntlT)Trin.exSceud.and7"inacijualcWhitney(O'Conner1990).
Werecognizeeightspeciesandtwovarietiesin NorthAmerica(excluding
Mexico),17speciesinMexicoandCentralAmerica,and18speciesandseven
varietiesinSouthAmerica(Finot2003a;Finot2004;Finotetal.2004).Tri^cluin
isalsopresentinFurope,Asia,Australia,andNewZealand(Jonsell1980;Tsvelev
1983;Clayton&Rcnvoize1986;Pohl&Davidse1994;Tucker1996;Fdgar1998;
Sorengetal.2003;Fuiotctal.2004,2005).Thegenusseemstobeabsentm
Africa(Clayton&Rcnvoize1986).Ahhoughseveralspeciesweredescribedfor
Africa(Steudel1854),theywerelatertransferredtoHcUclothchonBesserex
Schult.&Schult.f.(1lubl3ard1936;Schweickerdt1939).
TAXONOMICHISTORYOl-NORTHAMI-RICANSPFCll'S
Oneof theearliesttreatmentsof TrisctumwasmadebySteudel(1854),who
recognizedliveNorthAmericantaxa:T.ccrnuumTrin.,T.groenlandicumStQud.
{=T.spicatuin(L.).K.Richt.var.spicalumiT.lahradoricumSteud.{T.spicatum
var.spicdlum), T.moUc Kunth(I.spicatum var.spicaluin),and T.paluslvc
(Michx.)Torr\=Sphcnophoiispcnsylvonica(L.) Hitchcl Buckley(1862)de-
scribed threenewspecieslorNorth America: 7risctu;ngidhrun?Buckleyh
Dc,sclicf/)i/)s;t; diti\lhonioidcs (Trin.) Munrof, 7'. intcnuplum Buckley 1=
S/)/u'n()/)/i()/i.s!ntc;-n(/)(d(Buckley)Scribn.f,and7;ainc.sccitsBuckley,sometimes
referredtoasavarietyorsubspeciesoi'f.cernuumCBeal1896;Calder&Taylor
\965),ortoHelictothchonr(inc.sTcn,s(Buckley)Clayton.Anewspeciesdescribed
by Scribner(1884), T. hullii Scribn., is now considereda synonym ol
Sphcnopholisintcrrupta(Buckley)Scribn.(Finotetal.2004).Beal(1896)pre-
sentedatreatmentof 7ri5clu))imNorthAmerica(thosefoundonlyinMexico
and/orcountriessoutharemarkedwithanasterisk)tliatincludedthefollow-
ing15species:T.ccrnuuin,7^.cit^cuxioitics(Kunth)Kunth*,7^.c/onguhu?!(Kunth)
Kunth,T.jilijoUumScribn.exBeal*,T.halluScribn.,T.ludovicianuinVasey,T.
montanun]Vasey,T.palustrc,T.panicuhiiumF.Fourn.|=7/;,sctiu7ivir/tic(Kunth)
Kunth]*,T.pratensePers.[=Tr;.sctuni/7t/vc,s'cen.s(F.)RBeauv],T.^andbergiiBeal,
T.sesquiflorumTrin., T.spicalum, I.loluccnsc(Kunth)Kunth(= Trisctum
spicatumvai:spicaturn)r>).ndT.virlcliiF.Fourn.*f-iveolthesespeciesmcluded
inBeakstreatmentwerelatertransferredtoothergenera:Trisetumpalustre,T.
ludovicianuin,and l.haUiitoSphcnopholh(Scribner1906;Frdman1965);7^
ticycuxioit/cstoPcvni.st/i/tUl'niot2003b);andl.scscpujiorunitoCalamagwstis
(Rozhevits1952;Soreng&Greene2003).Beal(1896)alsodescribedanewspe-
cies,7"sandhcrgiifromMt.Stuart,Washington,nowconsideredasynonymof
T.cernuumTrin.(1litchcock1939:Hitchcock1950;Finot2003a).Beal(1896)rec-
ognized 7^.montanun]asa x-alidspecies.Itwaslatertreated,however,asa
synomymofIspicatumbyseveralauthors(Hitchcock1928;Weber1976).
Karl Richter(1890),an Austrian botanist recognized that 7Ti,st'(inn
FINOTETAL.,REVISIONOFTRISETUMANDGRAPHEPHORUMINNORTHAMERICA 1421
suhspicatum(L.)P.Beauv.wasanilligitimatehomonymsinceitwasbasedon
AiraspicataL.,andtherefore,madethecombination,Trisetumspicatum.Beal
(1896)wasthefirstNorthAmericanagrostologisttouseTrisetumspicatum.
ScribnerandMerril (1902)describedanewspeciesforthefloraofNorth
America,T.congdoniiScribn.&Merr.,asynonymotT.spicatuminmoderntreat-
ments.Severaldifferencesbetween I.congdoniiandT.spicatumlistedbythe
authorsincludedmorerigidleaves,narrowerglumes,morescabrouslemmas
andlongerawns.Sincetheextentof variationofT.spicatumisgreat,weleel
thesedifferencesdonotwarrantrecognitionatthespecieslevel.Forthecentral
Rockies(Colorado)threespecies:T.spicatumiasT.suhspicatum),T.nuy'usVasey
exRydb.(=T.spicatum),andT.montanumVasey,werereportedbyRydberg(1906).
Louis-Marie(1928-29)inhistaxonomicrevisionofTrisetuminAmerica,
included15specieswithseveralvarietiesinNorthAmerica.InTrisetumsubgen.
HeterolytrumLouis-Mariesect.AnaulacoaLouis-Mariesubsect.Trisetum(as
"Eutnseta"),heincluded Trisetummontanum[includingtwovarieties: 7^.
montanumvarpilosumLouis-MarieandTmontanumvarshearii (Scribn.)
Louis-Marie],T.cernuuni[including1cernuumvarluxuriansLouis-Marie,T.
cernuum var. luxurians to.puhcscensLouis-Marie,and T.cernuum var.
sandhcrgii(Beal)Louis-Marie],Tccmesce?isBuckley(includingIcanescensfo.
tonsumLouis-ManeandTcanescensio.velutinumLouis-Marie),Tprojectum
Louis-Marie, Tsesquijlorum, T. hongardiiLouis-Marie [= Calamagrostis
sesquijlora(Trm.)Tzvelev],IwilliamsiiLouis-Marie[=Danthoniaintermedia
Vasey],Tspicatum[includingTspicatumvar laxius(Lange)Louis-Marie,T.
spicatumvarmajusFarwell,T.spicatumvar.molle(Michx.)Piper,T.spicatum
vanbrittonii(Nash)Louis-Marie,T.spicatumvar.pi/osig/umeFern.,T.spicatum
varalaskanum(Nash)Makeex Louis-Marie,T.spicatumvar.vUlosissimum
Lange],andT.congdonii.InTrisetumsubsect.SphenophouieaLouis-Mariehe
includedTpennsylvanicum(L.)RBeauvexRoem.&Schult.,T.interruptum,
and Thallii,nowtransferredback toSphenopholisbyErdman (1965). In
Trisetumsubsect.Graphephorum(Desv)Louis-Mane,Loius-Marieincluded7^.
mehcoiJes(Michx.)Scribn.andT.wolfiiVaseylincludingTwolfiivarhrandegci
(Scribn.)Louis-MarieandTwolfiivarhrandegeifo.muticum(Boland.)Louis-
Marie],andinTrisetumSect.AulacoaLouis-MarieheincludedTjlorihundum
Pilg.iLectotypeheredesignated:=Dielsiochloaflorihunda(Pilg.)Pilg.],andT.
trinii(Trin.)Louis-Marie[=BromusherteroanusColla].
Hitchcock(1934)describedanewspeciesfromMontana,Torihochaetum.
Later,Hitchcock(1939)gaveadetailedaccountofthegenusfortheNorthAmeri-
canflora(includingMexico),where19specieswererecognizedunderTrisetum.
ElevenofthespecieslistedbyHitchcockarefromMexicoandCentralAmerica,
andthefollowingeightrangefromtheUnitedStates,Canada,andGreenland:
T.melicoidesand T.wolfii (heretreatedundergenusGraphephorum), T.
spicatum, T.orthochaetum, Tcanescens, T.cernuum, Tjlavescens,and T
1422 BRIT.ORG/SIDA21(3)
monttajiim.Finotetal.(2004)recognized17speciesofTrisctuminMexicoand
CentralAmerica:T.angustumSwallen,Ibarhatipaleum(HultenexVeldkamp)
Finot,7".turvisctumMorden&rValdes-Rcyna,7"durangenseFinot,P.M.Peter-
son,T.Jilifolium,T.irazuense(Kuntze)Hitchc,T.ligulatumFinot&Zuloaga,
Imanha-gonzalcziaePM.Peterson&Finot,Ipalmeri,T.pinctorumSwallen,
T.pringlei(Scribn.exBeal)Hitchc, 7" rosei Scribn.&Merr.,7^.spellenhergii
Soreng,Finot&RM.Peterson,T.spicatum,T.tonduziiFlitchc,T.viride,andT.
virletti.
Hulten(195Q)studiedtheTrisctumspicatumcomplexonaworld-wideba-
sis,andrecognizedseveralNorthAmericanintraspecifictaxa:T.spicatumsubsp.
alaskanum(Nash)Hulten,T.spicatumsubsp.pilosiglumc(Fernald)Hulten,T.
sT.psipcaitcuamtusmubssupb.spm.olclocn(gMdiocnhixi.)(S1clruilbtne.n,&I,Mseprirc.a)tHuumlstuebns,p.anmdajTu.ss(pViacsaetyu)mHuslutbesnp,.
cnngdoniivar.spicatiformeHulten.LaterauthorsconcludedthatT.spicatum.is
polymorphic(Randall&rHilu1986),andmostoftheinlraspecifictaxawere
placedinsynonymy.
GraphcphorumisasmallgenusendemictoNorthAmerica,closelyrelatedto
Trisctum.Itditlerstromthisgenusinhavingtheapexolthelemmaentireandthe
dorsalawnreducedtoasubapicalmucro.Graphcphorumcomprisesonlytwospe-
cies,G.fuelicoitfcs(Michx.)DesvandG.wolfii(Vasey)Vasey(Finot&Soreng2003),
sometimestreatedunder Trisctum.Claytonand Renvoize(1986)indicate
Graphcphorumhasthreespeciesbutonlythetypespecieswasgiven.Onespecies
fromMexico,originallydescribedasGraphcphorumaltijugumE.Fourn.iscur-
rentlyacceptedasPcyritschiakoclerioidcs(Peyr.)E.Fourn.(Finot2003b;Finotet
al.2004).GraphcphorumpringleiScrib.exBeal,IromMexico,Guatemala,Costa
Rica,andPanama,wastransferredtoTrisctumby1Iitchcock(1927).
MATERIALANDMFTIIODS
Typespecimensandgeneralcollectionsfromthefollowingherbariawerestud-
ied:BA,BAA,BAFCCR,CIIDIR,CONC,FLPMFRI^,PPR,QCA,SGO,S,SIand
US.Formicromorphologicalstudies,liguleswerecollectedIromupperandbasal
culmleaveswithasharprazorbladeandmountedinlactophenol.Forleafana-
tomicalstudies,handcrosssectionsandabaxialepidermispreparationswere
madeloUowingMetcalfe(1960).Theterminologyforliguledescriptionisbased
mainlyonChaffey(1983,1984).Transversesectionsandabaxialepidermisol
leavesweredescribedlollowingtheterminologyproposedbyEllis(1976,1979).
ObservationsweremadewithaZeissMC-80microscopeequippedwithobjec-
tives20x,40x,C')3x,andlOOx.Allgrassnamesmentionedinthismanuscript,
includingthoseintheintroductionarctreatedinAppendix2.
Inthemorphologicaldescriptionsthelengthgivenforfloretswasusually
takentromthefirstorlowestfloret.Iftherewerethreeormorefloretsperspike-
letthenthesecondfloretwassometimesusedtocalculatetherange.Therefore,
fINOTETAL,REVISIONOFTRISETUMANDGRAPHEPHORUMINNORTHAMERICA 1423
whenusingourkeystodetermineNorthAmericanspecimensofTrisetumitis
besttomeasureonlythefirstorlowestfloret.
TAXONOMICTREATMENT
KEYFORDISTINGUISHINGTHEGENERATRISETUM
ANDGRAPHEPHORUMINNORTHAMERICA
1. Lemmawithlateralnervesnotprolongedintoapicaisetae,theapexentiretoslightly
bilobate;dorsalawnabsentorreducedtoashortsubapicalmucro;paleanotgap-
ing(paleatightlyenclosedbythemarginsofthelemma);paniclelax,openorcon-
tracted,neverspiciform Graphephorum
1. Lemmawithlateralnervesprolongedinto2(4)apicalsetae,theapexbidentate;
dorsalawnwelldeveloped,borneontheupperhalforthirdofthelemma;palea
gaping(paleanottightlyenclosedbythemarginsofthelemma);paniclelaxor
denselyTlowered,contractedoropen,andovateorpyramidal,oftenspiciform
__^^__^^Trisetum
Graphephorum Desv.,Nouv. Bull.Sci.Soc.Philom.Paris2:189. 1810.Type:
Gi'dpht-phoruinmelicoides(Michx.)Desv.
Perennials,looselycaespitoseorwithshortrhizomes.Culms50-100cmtall,
glabrousorpilose.Leafsheathsglabrous,scabrous,and/orpilose;bladesflat,
glabrousorpilose;ligulemembranous.Inflorescenceinpaniclesopenorcon-
tracted,glabrousbelowthepanicle.Spikelets2-or3-flowered;rachillapilose;dis-
articulationabovetheglumesandbetweentheflorets;glumesslightlyunequal,
slightlyshorterthanthespikelet;firstglume1-nerved,lanceolatetolinear-lan-
ceolate;secondglume1-or3-nerved,lanceolatetooval-lanceolate;lemmaswith
apexentiretoslightlybilobate,muticousorwithareducedawn(mucro),borne
inmediatelybelowtheapex,themucronotreachingtheapexoithelemma;
paleanotgaping;stamens3;ovaryglabrousorwithshortapicalhairs.
KEYTOTHESPECIESOFGRAPHEPHORUMINNORTHAMERICA
1. Panicles2-4cmwide,open,lax;ovaryglabrous;callushairsabout2.5mm I.G.melicoides
1. cPaalnliucslehsai1rs-10..55cmmmwildoen,gcontracted;ovaryglabrousorwithshorthairsattheapex; 2.G.wolfii
1.Graphephorummelicoides(Michx.)Desv,NouvBull.Sci.Soc.Philom.Paris
2:189.1810.AnamelicoidesMichx.,FLBor.Amer.L62.1803.Poamelicoides(Michx.)Nutt.,
Gen.N.Amer.Pi.L68.1818.TriodiamclicoitiesCMichx.)Spreng,,Syst.Veg.1:331.1825.Trisetum
mehcouics(Michx.)Scnbn.,Bot.Gaz,9:169,1884.Typh:CANADA.A\mericaBoreal,Michaux
s.n.(llOl.OTYPH:PI;ISOTYPIZ:LE-TRIN]864,0P),
ArundoairoidesPoir,,Encycl,6:270.1804.Deyeuxiaairoiodes(Poir.)FBeauv,Ess.Agrostogr.44,
152,160.1812.Agmslisairoides(Poir.)Raspail,Ann.Sci.Nat,(Paris)5:449.1825,nom.illeg.
liom.Ctilamdgrosfisairoides(Poir.)Steud.,Nomencl.Bot.(ed.2)1:249.1840.TypeU.S.A.America
Septentrionalis,Michauxs.n.(HC^IX")TYPE:Herb.Juss,,notseen),
DuponiiacooleyiA.Gray,Manual(ed.2)556.1856,GraphephorummelicoidesvarmajusA.
GrayProc,AmerAcad,Arts5:191,1861.Graphephorummelicoidesvar.cooleyi(A.Gray)Scnbn.,
Mem.TorreyBoi.Club5(4):53,1894.Trisetummelicoidessubsp.cooleyi(A.Gray)Scribn.,
1424 BRIT.ORG/SIDA21(3)
Rlmtli^raBtaOJiSZ.1Q06,Trisclummduoidcsvai:majus[A.Gray)Hitchc.RhodorakXll2):65.
\^K)8.Giaphcphorumcoolcyi(A.Gray)Farw.,Pap.MichiganAcad.Sci.1:88.1Q21Tvri-:IJ,.S.A,
MlClllCANidUil(TiYp|-:luitlound).
Pcrenninls.Culms50-100cmtall,glabrous.Leafsheathsscabrousorpubes-
cent;Hgules1.3-2mm long,oval,glabrousdorsally,apexdentatewithshort
cilia;blades80-150x2-8mm,flat,glabrous,sometimespubescentadaxially.
Panicles8-20x 2-4cm,lax,open,nodding;branchesascending;rachissca-
brous.Spikelets6-7mmlong;rachillahairy;thehairs1-1.5mmlong;glumes
unequal,shorterthanthespikelet;keelsmoothorscabrousontheupperhalf;
1irstglume4-4.5X0.5mm,shorterandnarrowerthanthesecondglume,lan-
ceolate,1-nerved;secondglume6-7X0.8mm,oval-lanceolate,3-nerved;lem-
maswithshortandstraightsubapicalawn(mucro),mtermediatenervesnot
reachingtheapex;apexacute,entire,notsetulateoraristulate;callushairy,the
hairsabout2.5mmlong;paleashorterthanthelemma,2-ncrved;apexentu'e;
lodiculesabout0.6mmlong,wuhtheapexentiretoslightlybilobate;ovary
glabrous.Caryopses3.2-3.8mmlong,glabrous;endospermsemi-liquid.
AnatomyandMicwmorpholoii^y—Ugukapiceswithouthairsorpapillate
cells,composedollongcellswithroundedends;liguleepidermiscomposedof
rectangular longcellswithstraightside walls, without prickle hairsor
macrohairs;bladesintransversesectionflattoweakIykeeled;adaxialfacewith
roundedribs,theribsnotveryprominent;furrowswiderthantheadjacentribs;
abaxial lacewithribssimilarinsizeandshapetotheadaxialribs;vascular
bundles15,rounded,situatedinthemedianportionoftheblade;vascular
bundleswithadaxialandabaxialgirders;marginalsclerenchymapresent,small;
epidermisintransversesectionlargerthanthemesophyll,theadaxialepider-
mis with epidermal cells more or less rounded, thin; bullilorm cells
inconspicous;pricklesscarce,presentonlyintheadaxialepidermis;stomata
present in bothadaxialandabaxialsurfaces;abaxialepidermiswithcostal-
intercostalzoneswelldillercntiatcd;intercostalzoneswithlongcellsrectan-
gulartolusiform,withstraightsidewallsandverticaltoobliqueterminalwalls;
intercostalshortcellsabsent;stomataintwointercostalrows;pricklesscarce;
macrohairsabsent;costalzone2or3cellswide,withlongcellssimilartothe
intercostalepidermalcellsbutshorterandnarrower;shortcellsrectangular,
withsinuouswalls;pricklesinrowsonthecostalzones.
Dislrihutinn.—Canada(Newfoundland,Ontario,Quebec)andUnitedStates
(Maine,Michigan,NewHampshire,NewYork,Vermont,Wisconsin)|1litchcock
lQ39;Karteszl998i
Specimensexamined.CANADA.Ontario:Township,2(1mISofMoonbeam,14Aug195^,A'loMoii(144'}
(US).Newfoundland:\alieyolF.xploitsRiver.Grandi^alls,12Aug\^n].M.LJc}iutld&Wiegandi591
a.lS).U.S.A.Maine:AroostookCo.:alongSt,JohnRiver,St.Francus,5Aug18^)3,ML.Fcvnald187(P):
NorthernMaine,St.FrancisRiver,Aug1902.E^lc^tonM36tP);WooseheadLake,1Sep18(58,A.E
Smith&(:.!:.SmithynAP).
FINOTETAL.,REVISIONOFTRISETUMANDGRAPHEPHORUMINNORTHAMERICA 1425
2.Graphephorumwolfii(Vasey)VaseyexCoult.,Man.Bot.RockyMt.423.1885.
TrisetumwoljnVasey,MonthlyRep.Dept.Agric.1874:156.1874.TypeU.S.A.Colorado:Lake
Co.:TwinLakes,1873,/.Wolfe^.n.(i.soTYn-S:NY-232367!,US-81781!).
Tvi:>€iumiubspicalumvar.mutnuinBol.,Bot.California2:296.1880.Typh:U.S.A.California:onthe
upperTuolumne,7500ft,1866,H.N.Bolandcr50.19(isotypps:NY-232371!,US-867860!,US-
344201!).
Perennials.Culms50-100cmtall,withshortrhizomes.Leafsheathsglabrous
orpilose;ligules1.5-2mmlong,truncate,glabrousdorsally,apexdentate;blades
60-200 X 3-7mm,flat,soft;lowerbladespiloseabaxially,sparselypilose
adaxially;upperbladesglabroustosparselypiloseabaxially,sparselypilose
adaxially.Panicle8-15x1-1.5cm,contracted,dense,erect,tingedwithpurple;
rachisstronglyscabrous,Spikelets6-7.5mmlong,2-or3-flowered;rachilla1.5
mmlong,hairy,thehairs0.5-1.5mmlong;glumesunequal,nearlyaslongas
thespikelet;keelscabroustowardstheapex;apexacute;firstglume4.2-6 x
0.5-0.7mm,shorterandnarrowerthanthesecondglume,lanceolate,1-nerved;
secondglume5-6.5x 0.8-0.9mm,aslongorshorterthanadjacentfloret,3-
nerved;lowerfloret5-5.5mmlong,;upperfloretsshorter;lemmaglabrousto
slightlyscabroustowardstheapex;apexacutetoslightlybidentate,hyaline,
muticousorwithashortsubapicalmucrof-2.2mmlong,borneat1-1.3mm
belowtheapex;callusshorthairy,thehairsabout0.5mmlong;paleaabout4
mmlong,shorterthanthelemma,2-nerved,thenervesscabrous;lodicules0.7-
0.8mmlong;apexwithtwo2teeth,sometimeswithathirdsmallertoothbe-
tweenthem;anthers0.7-0.8mmlong;ovaryglabrous,sometimeswithhairs
neartheapex.Caryopses2.4-3mmlong,withorwithouthairsattheapex;en-
dospermsolid.
Anatomyandmicromorphology.—Liguleapiceswithstilfhairsandpapil-
latecells;liguleepidermiscomposedofrectangularlongcells;shortcells
present;pricklehairsscarce;stomataandmacrohairsabsent;bladesintrans-
versesectionV-shaped,symmetric,keeled,thekeelwithawelldeveloped
sclerenchymatictissue;adaxialribsabsent;centralvascularbundlefree,with-
outadaxialsclerenchyma;secondordervascularbundleswithI-shapedscleren-
chymagirders;thirdordervascularbundlesfree,withsmallstrandsofadaxial
orabaxialsclerenchyma;marginalsclerenchymasmall;bulliformcellscon-
spicuous,mfan-shapedgroups;macrohairsabsent;abaxialepidermiswithcos-
tal-intercostalzonationpresent;intercostalzoneswithfusiformtorectangular
longcells,withlateralwallsstraight;shortcellsandstomatapresent;prickles
andmacrohairsabsent;costalzoneswithlongcellsrectangular,narrowerthan
theintercostalcells;shortcellsrectangular,withsinuoussidewalls.
Distribution—AspeciesendemictowesternUnitedStates(California,Colo-
rado,Idaho,Montana,NewMexico,Nevada,Oregon,Utah,Washington,and
Wyoming)iHitchcock1939;Kartesz1998).
1426 BRIT.ORG/SIDA21(3)
Specimensexamined.U.S.A.CALIFORNIA:MonoCo.:RavineCreek,3200m,21AuglQ54,Kr((j7()viil;as
8102(SI):YosemiteNationalPark,%00It,1Aug1^34,Barlholomewsji.(US);J.R.Swallcn1937(P).
fCt.OL14ORA^uVgDO18:^)8S,cSqhucaacihi&CoB.c:sMsacrysIh4a«l2llPUaSs)s.,N1E08V0A0DIAt,:27EJluklo1Co8.%:,jCu/stemaebnofv,es2A0n0ge(lUSL);akBeu,ffEaalsotPlaisus,ml1x0j5l0dt0
Mountain,8000ft,24Jun1958,/'RR(jven&O.ISolbrig.040(SI);PondcrosaMountam,aboveIxiglis
LWaokoet,on70s0.n0.(fUt,S)2.6WJAulSH19I01N,GMTcOnNi:l&S'poVkVainleuwCo3.:482(1P)j.unN1E8W89.MWEXISuCkOs:doCrafsst.inlXlPa)V:alWl.eyS,uk4sdSoerpJ9f49913{,.Pl)i..d
TriselumPcrs.,Syn.Pi.1:97,1805.TnicianumPom,Hncycl.Suppl.5:365.f8f7,nom.superfl.,
RchoitisiiiiaOpiz,Lotos4:104.1854,nom.sh/hi/I.,nonPA.Karsten1869.TYPE:Trisetum
Jhivcsa-iis{]..)PBeauv,Hss,Agrostogr88,153,t.18,LL1812.
AcwspcUonBesserexSchuh.&Schult,L,Syst,Veg.Mant.3:526,1827,LliOTOTYPl-(designatedby
L,K,G.Pleiffer,Nom.1:38.8Dec.I78f.):AvcnadistuhophlhiVill.|=InsrlumdistichophyUum
(Vill,)PBeauv]
Rupc:^tnnaI'rov,,Fl,Canad.:689.1862.TYPE:RupcsDiiuipuhcstcnsProv.|=JyisclumspicatHm(L)
Richi.l.
Perennialsandannuals,caespitose,sometimesshortlyrhizomatousand/or
stolonilerous.Culms16-120cmtall,erecttogeniculateatbase,glabrousorpu-
bescent.Lealsheathsglabrousorpubescent,longerorshorterthantheintern-
odes;bladesHat,conduplicate,convoluteorinvolute,softrarelyrigid;ligule
membranous.Inllorescenceinpaniclescontractedoropen,laxordensely-flow-
ered,spiciform,ovate,orpyramidal;therachisglabrous,scabrousorpubescent.
Spikelets(l-)2-to4-flowered,shortpedicellate;rachillapubescentorglabrous,
usuallyprolongedbeyondtheupperfloret;disarticulationabovetheglumes
andbetweentheilorets;glumesheteromorphic,lanceolatetoovate-lanceolate,
equalorunequal,firstglutne1-nerved,usuallyshorterandnarrowerthanthe
second,secondglume1-or,3-nerved;lemmaslanceolate,(3-J5-(7-)nerved,usu-
allyawneddorsallyormuticous,withapexandmarguishyaline,glabrousor
pubescent,slightlykeeledandcompressed,rai^elyterete;apexwith2-to4api-
calsetaeorshortawns,bidentate,or2-toothcd;centralawn fromtheupper
third,rarelythemiddle,ofthesubapicalportionofthelemma;awnexserted,
geniculateormerelydivaricate;callusshortpilose;paleanottightlyenclosed
bythemarginsofthelemma(gaping),2-keeled,hyaline,usuallyshorterthan
thelemma;stamens3,anthers0.5-3mmlong;lodicules2,membranous,often
apicallylobed;ovaryglabrousorwithshortandshiningtrichomesnearthe
apex;endospermsolidorliquid,softorhard.Caryopscscompressed,soft;hi-
lumshort,punctiform.Basicchromosomenumberx=7.
KEYTOSPHCIHSOPTRISETUMIN NORTHAMFRICA
L Platitsdelicateannuals;knownonlyfromasingleintroductioninCamden,New
Jersey S.T.aureum
L Plantsperennial;widespread.
2.Panicleslax,open;glumesshortertlianthespikelet(Sect.Tnsetum).
3, Ovaryandcaryopsishairyneartheapex.
FINOTETAL,REVISIONOFTRISETUMANDGRAPHEPHORUMINNORTHAMERICA 1427
4. Awns3.5-4mmlong,straight,shorterthanthelemma;apexofthelemma
shortlybidentate.-firstglumes4-6mmlong,neverrudimentary 7.T.
orthochaetum
4. Awns6-16mmlong,geniculate,onetothreetimesaslongasthelemma;
apexofthelemmaendingintwosetae0,5-1.5mmlong;firstglumes0.5-
5mmlong,sometimesrudimentary
5. Paniclesfew-flowered,loose;branchescapillary,theloweronesusually
nakedonthelowerthird;leafbladesglabrous 4a.T.cernuum
subspcernuum
5. Paniclesdensely-flowered,narrow;branchescloselyappressed,thelower
oneswithspikeletstonearbase;leafbladescanescenttosparsely
pilose 4b.T.cernuumsubspcanescens
3. Ovaryandcaryopsisglabrousneartheapex
6. tSrpaickteelde;tsspi2k-e4l-eftlsow5e-r9edm;mpalnoincgl;esleumsmuaatllaylgaowlnds-y5el-l9owmismh,lsonogm;eawnthhaetrsc2o-n3-
mmlong.
7. Lemmasonlydorsallyscabrous 9.T.sibiricum
7. Lemmasuniformilyscabrous S.T.flavescens
6. Slpoioskee;lestpsik2e-leotrs3-4f.i5o-w6ermedm;plaonnigc;lelsemgmraeteanlsaowmnesti3m.e5s-4timngmedlwonigt;hapnutrhpelres,
0.8-1.2mmlong 6.T.montanum
2 Paniclescontracted,spiciformtonarrowbutalwaysdensely-flowered;glumes
onlyalittleshorterthanthespikelet(Sect.Trisetaera,\npart).
8. Panicles2.5-7(-10)cmlong,denselyspiciform lO.T.spicatum
8. Panicles8-23cmlong,open,loose,few-floweredtospiciformandinterrupted
withshortascendingbranches.
9. Lemmatalawns7-12mmlong,weaklytwisted,geniculate;ligulesabout
2.5mmlong,glabrousorscabrousdorsally;ovaryhairynearapex 4b.T.cernuum
subsp.canescens
9. cLuermvmeadt;alligualwenss0.a3b5o-1u.t55m.5mmlmongl,ondge,nnsoetlytwpiilsotseeddoorrsgaelnliyc;uolvaatrey,dgilvaebrrsoeulsy
8.T.projectum
3.Trisetumaureum(Ten.)Ten.,Fl.Napol.2:378.1820.KodcnaaurcaTen.,Tract,Pitogn.
1:58.1806.Trisetariaaurea(Ten.)Pignatti.Arch.Bot.(Forii)31:51.1955.Type:(hoi.OTYPI::NAP?),
Delicateannuals,caespitose;culms18-30cmtall,erect,spreadingorgenicu-
lateatbase,glabrous.Leafsheathsglabrousorpubescent,shorterthanthein-
ternodes;ligule0.8-1.3mmlong,hyaline,apexobtuse,marginsdecurrent;blades
2-8ciTilong,1.2-3mmwide,glabroustopubescent,thehairslessthan0.2mm
long.Panicles1-5x 0.8-3cm,pyramidaltoovoid,contracted,denselyflow-
ered,oftenlobed,shiningyellowishorlightbrownish;branches0.6-3cmlong,
closelyappressedandascending,nakedbelow;rachisglabrous.Spikelets2.6-
3.3mmlong,2-or3-flowered;pedicels0.2-1.6mmlong;rachilla0.6-1.5mm
long,coveredwithstiffhairs,thehairs0.3-1.2mmlong;glumesunequal,the
secondglumelongerthanthespikelet,scaberulousonthekeel;firstglume2.1-
2.6ininlong,linear-lanceolate,shorterandnarrowerthanthesecondglume,1-
nerved,apexacuminate;secondglume2.8-3.3mmlong,ovate-lanceolate,1-or
1428 BRIT.ORG/SI0A21(3)
3-nerved,apexacute;lemmas1.8-3mmlong,glabrousorscatteredpilose,rarely
scabcrulous,ovate,hyaline,theapexbidentatewithtwosetae0.1-0.3mmlong;
awn1.3-3.'5(-6)mmlong,slightlybent,scabcrulous,Liornejustabovethemiddle;
callusobtuse,mostlyglabrous;paleas1.5-2.8mmlong,shorterthanthelemma,
hyaline,2-nerved,theapexbidentatewithtwoteeth0.2-0.4mmlong;anthers
1.2-1.6mmlong;ovarywithalewshorthairsneartheapex.Caryopsisnotseen.
Dislrihution—TrisclnmaureumisintroducedinNorthAmericaandoc-
cursnativelyintheMediterraneanRegionolEuropeinGreece,Italyjugoslavia,
andSicilia.
Comments—V^earenotinapositiontocriticallyevaluatethegenericstatus
ottheassemblageofapproximately15speciesthataresometimessegregatedas
IrisclariiiForssk.(sensuClayton&Renvoize1986).ThistaxonistreatedasTri.sct-
umaureumintheFloraolFrancebyKcrguclen(1999)andasTrisctariaaurcaby
Dogan(1985)intheFlorao\ Turkey.
S|X'cimcnsexamined.U.S.A.NewJersey:C^anulcn.Iiallasi,1-C-MartiDihilcsji.ilJS).
4a.TrisetumcernuumTrin.subsp.cernuum,Mem.Acad.hup.Sci.St.-Petersbourg,
Ser6,Sci.Math.1:61.1830.AvcncufMUKaTrin.)i<unth.Rev.cicn.i.suppL20,1833.Type:
U.S.A.Ala.sKA:Sitka,1826,j.HG.vonEschschoitz.s./i.(iioi.(Vivim l_l'-Ti^lN1884-01!;iMriYPrs:
BAA-3366!,NY-23236W,P!,US-8t779Fragm.e,\IJ-TRINU.
Avfihii\ull;aciisis]Vrcs\.Rc]]^.liaenk.i(4-^):2'i4.1830.TnsctumiiKlletU'iLsis(J.l'resI)Scnbner&
Men-exOavy,Univ,c:alil Publ,Bot.1:03,1402,TvPi::CANADA.BKPrisilCoLUMBLA:Vaneouver
Lslancl.Nootka.7".Hucukc,s.m.UsOTYPFS:iJMRiN-l929.0t!;US-865598tragm.D.
TriictumsandhcignBeal,GrassesN.Amer.2:378,1896.Triaetumcernuumvar.sandhcr^^htBeal)
Louis-Marie.Rhodora30:214.1928.Typp,:U.S.A.WASHINGTON:Mt,Stuart,7000-8000ft,
Sdndbci\[&l.e(/)ert;,S'2.i IOIati'YPP.:MSC?;Ps0TYPP:s,NY-431712!,Gil,\VS).
Trisclitinicii\uumvar,luxiintinsLouis-Mane,Rhodora30:213.1928.Typh:LJ,S,A,OkP(,(iK:Sea-
side,11Aug1899.(.'/_Shi-dr&EL.Se/-i/)/ieriZOSCHOLOTYPH:US-867917!).
Tri.sc(11)11ce/uiuuiito.]iii/v,sie)PsLouis-Mane,Rhodora30:213.1928.Trisetumccrniiumio.puhc'^ccns
G,Jones,Univ,Wash,I'uhLBiol,5:108,1936,horn,illeg.superfl,TYPE:U,S,A.C^APIPHRNIA:
1lunibdldtGo,:Hureka.30May1920,Anonvmouss.n.(nonrwvv:Ut;-2t2883),
Perennials,withstolonsandrhizomes.Culms35-85cmtall,glabrous,some-
timesgeniculate,with2-4nodes.Fcafsheathslongerorshorterthanthemter-
nodes,glabrousorrarelypilose;ligules1,3-7mmlong,membranous,longerin
theupperleaves,oval,Iimbriatc,ciliateattheapex,glabrousorpilosedorsally;
blades100-220x3.5-12mm,flat,soft,glabrousorsparselypubescentonthe
adaxially.Panicles6-30X2-5cm,lax,open,loose,few-flowered,greenorpur-
plish;branchescapillary,theloweronesusuallynakedonlowerthird;rachts
glabrous.Spikelets4.5-12x2-4mm,2-or3-flowered,notovcriapping;pedicels
c1-o4v.e5remdmwiltohngs,ticfafpihlaliarrsy,,tIhIecxhuaoiurss,1g-l2ambrmousloonrg;scgalburomuess;srhaocrhitlelrat1h.a5n-2tmhemspliokneg-,
let,veryunequaltosubequalinlengthandveryunequalinwidth;firstglumes
0.5-5XO.f-0.5mm,fincar-lanceofate,attenuate,1-nerved,sometimesrudimen-