BROMUSHALLII (POACEAE), A NEW COMBINATION FOR CALIFORNIA, U.S.A., AND TAXONOMIC NOTES ON BROMUS ORCUTTIANUS AND BROMUS GRANDIS

2005·8.1 MB·English

by  Jeffery M Saarela

BROMUSHALUl(POACEAE),ANEWCOMBINATIONFOR
CALIFORNIA,U.S.A.,ANDTAXONOMICNOTES
ONBROMUSORCUTTlANUSANDBROMUSGRANDIS
JefferyM.Saarela Paul M.Peterson
DepartmentofBotany,and DepartmentofBotany
UBCBotanicalGardenandCentreforPlantResearch NationalMuseumofNaturalHistory
UniversityofBritishColumbia SmithsonianInstitution
Vancouver,B.C.,CANADA Washington,DC20013-7012,U.S.A.
[email protected] [email protected]
JacquesCayouette
NationalProgramonEnvironmentalHealth-Biodiversity
AgricultureandAgri-FoodCanada
Wm.SaundersBuilding(#49)
CentralExperimentalEarm
Ottawa,Ontario,CANADA,KIA0C6
[email protected]
ABSTRACT
IlerbunumstudyofBromusgrandi.s,Rim"u((itnuis,andB.ortutiionusvar.haW'uIromwesternNorth
Americahasindicatedthatthesetaxaaremorphologicallyandgeographicallydistinct,andi3est
treatedasspecies.AnewcombinationBromushallii(.Hitchc.)Saarela&RM.Petersonismade.Bromin
ohraclulititiadniufsf)e,rslofwreormshB.eaotrhcsuwltiitahnduesnsbeyshhaorvtinstgifdfehnasiresly1mpmubelsocngenotrlbelssad(evss.l(voenrgsseosftghlaaibrrso2u-s4imnmB.
longj,and1-2C-3)culmnodes(vs.2-4).BromushallhdiffersfromB.gvandisbyhavingbladesthat
are7.5-16.")cmlong|vs.(13-)18-38cmlonginB.^randis],l-2(-3)culmnodes(vs.3-7),andlower
glumesthatarcl(-3)-nerved[vs.3(l)-nerved].Weprovidethefirstreportoidifferencesmthetypeol
pubescenceonthelowersheathsbetweenB.orcuttianusandB.hallii.Weconfirmthediploidchro-
mosomenumberofIn=14forB.^randis.Weincludedescriptions,synonymies,andrepresentative
specimensexatninedforB.grandis,B.hallii,andB.orcuttianus,akeytoallsectionsolBromusin
California,andakeytoBromussect.BromopslsinCalifornia.
RESUMEN
RlestudiodeBromusgrandis,B.orcuttianusyB.orcuttianusvar.halliideNorteamericaoccidental
haindicadoqueestostaxasonmorfologicaygeograficamentediterentes.yestantratadosmejorcomo
especiesdistintas.SehaceunanuevacombinacionBromushallii(Hitchc.)SaarelayP.M.Peterson.
gBlraobmruassehnalBl.iiorsceutdtiifaenruesn)c.ivaadmeas6.ionrlceuritotrieasnucsonpolorstpeenleorsrliagmidmoasscodretonssadmee1nmtempluabregsocoenmtcenso(s.c(ocnotnr-a
tralospelossuaveslargosde2-4mm),yf62(-3)nudosenelculmen(contra2-4).Bromushalliise
diferenciadeBromusgrandisportenerlaslaminasde7.5-16.5cmdelargo(contra18-38cmenB.
grandis),1o2nudosenelculmen(contra3-6),yglumasinlerioresU-3)-nervadas[contra.3(1)-
nervadasj.Proporcionamoselprimerinformedediferenciaseneltipodepubescenciaenlasvainas
inlerioresentreB.orci(ttianu.syB./ialiii.Coniirmamoselnumerodiploidedecromosomasde2)1=14
paraB.grandis.Incluimosdescripciones,sinonimias,yespecimenesrepresentativosexaminadosde
SIDA21(4):1997-2013.2005
1998 BRIT.ORG/SIDA21(4)
li,,i;)(1/1(1IS,lihtilhiyU.orntUuiitus.unaclavedctdclaslasseccioncsdciironiusenc;aliiornia,yuna
cla\x-dcHiiHiiuss(-((.BioinopsisenCaliidrnia,
BromusL.isalargegenusofapproximately160speciesthataredistributed
worldwideintemperateregions.Thegenusisdistinguishedfromothergrass
generabythecombinationol leal' sheathsthatarcconnateformostoftheir
length,awnsthatareinsertedsubapically,andhairyappendagesontheapices
oftheovary(Clayton(SiRenvoize1986).IntheNewWodd,79nativeandintro-
ducedspeciesarecurrentlyrecognized(Pavlickctal.2003),althoughnewspe-
ciesarebeingdescribed(e.g.,Saarelaetal.inreview)andthetaxonomicbound-
ariesolseveraldillicultspeciescomplexeshavebeenreevaluated(e.g.,Peterson
ctal.2002;Ojaetal.200^,Massaetal.2004).Molecularphylogeneticinvestiga-
tionshaveclarifiedseveralaspectsoftheevolutionaryhistoryofBromus.and
haveidentilledmajorlineagesinthegenus,someofwhichcorrespondtocom-
monlyrecognizedintragenerictaxa(Pillay&Hilu1990,1995;Ainouche&Bayer
1997;Saarelaetal.inpress).Anewclassificationofthegenushasnotyetbeen
proposed,pendinglurtherstudyofmorphologicalandmolecularvariation.
BroiHii.ssect.Bronio/i.si.sDumort.,thelargestofthetraditionallyrecognized
sectionsinthegenus,isanon-monophyletictaxoncomprisedofseverallin-
eagesthatoccurinNorthAmerica,Mexico,SouthAmerica,andHurasia(Saarela
etal.inpress).Thesectionischaracterizedmorphologicallybyl(.3)-and 3(5)-
nervedtirstandsecondglumes,lemmasthataredorsallyflattened,andape-
rennial habit(Smith 1970),althoughsomeofthesecharacteristicsmaybe
symplesiomorphies.InNorthAmericanorthofMexico,thereare16nativespe-
ciesinsectionBwmopsisthathavediversifiedinavarietyofhabitats(Pavlick
1995;Pavlick&Anderton,in press).Somespeciesarewide-ranging(e.g.,B.
iiiidiusP.),whereasothersarehighlyrestrictedmdistribution|e.g.,B.icxcnsis
(Shear) llitchc.].Several taxa (B. lacvipcsShear, B.orcullianusShear, B.
oiruttidDusvar /u;///i llitchc,B.p^cudoUicvipcsWagnon,andRsuksdorfii
Vascy)arerestrictedtomountainousregionsinthesouthwesternpartofthe
continent,largelymCalifornia(Pavlick1995).Othersle.g.,B.pofk-ri(J.M,Coult.)
Nash,B.richardsoniiPink,B.vul(,'c(n5(Hook.)Shear]havebroaderdistributions
inwesternNorthAmerica.WhilestudyingBmmii.sspecimensfromCalifornia
tosupportpreparationot thetaxonomictreatmentofBromuslortheSecond
HditionolthejepsonManual(Saarela&rPeterson,inprep.),itbecameclearthat
B.orcuitianusvar/ud/ii,despiteitsvarietalrank,isareadilyidentifiabletaxon
thatcan beconsistentlyandreliablydistinguishedfrombothB.orcutluuius
var.orcuttuniusandB.t^runJi.s.Thesepreliminaryobservationsstimulateda
moredetailedinvestigationofitsmorphologicalvariationandtaxonomicstatus.
Vasey(1885)namedanddescribedB.orcuttiauusbasedoncollectionsby
C.R.Orcutt Irom the mountainsnearSan Diego,andcollectionsby W.N.
SuksdorffromMt.AdamsinWashington.InhisrevisionoftheNorthAmerican
SAARELAETAL.,BROMUSHALLIIANDRELATEDTAXAFROMWESTERNNORTHAMERICA 1999
speciesofBromus,Shear(1900)recognizedB.orcuttianus,anddescribedlarger
plantsfromLaMaite,SanDiego,withpubescentsheaths,leaves,culms,and
spikeletsasanewvariety,B.orcuttianusvar.grandisShear.Iditchcock(inJepson
1912)describedplantsfromthemountainsofsouthernCaliforniawithdensely
pubescentbladesandcataphylls,andpubescentglumesandlemmasasanew
variety,B.orcuttianusvarhalliiHitchc.Hitchcock(inJepson1912)alsoelevated
B. rcuttianusvar.grandistoaspecies,B.grandis(Shear)Hitchc,nidicatnigthat
itissimilartoB.orcuttianusvarhallii.butdiffersinitsdroopingpanicleand
distinctly 3-nervedlowerglumes.Subsequentauthorshaverecognized B.
grandi.s,B.orcuttianus,andB.orcuttianusvarhalliiintheirtreatments(e.g.,
Hitchcock&Chase1951;Wagnon1952;Munz&Keck1959;Hitchcocketal.
1969;Munz1974;Wilken&Painter1993;Pavlick1995;Pavlick&Anderton,in
press)althoughClaytonand Williamson(2002onwards)synonymizedB.
orcuttianusvar. hallii with B.orcuttianus. In hisrevisionofBromussect.
BromopsisinNorthAmerica,Wagnon(1952)hypothesizedthatB.orcuttianus
varhalliiisequallyrelatedtoB.grandisandB.orcuttianus,andindicatedthat
itrequiresfurtherstudy.AmongspeciesofBromussect.BromopsisinNorth
America,B.orcuttianusvarhalliiistheonlytaxonrecognizedcurrentlyatthe
varietallevel,eventhoughthecharacteristicsseparatingitfromB.orcuttianus
varorcuttianusandB.grandisaresimilarmdegreetothoseseparatingother
morphologicallysimilarspeciesofBromus.Herewediscussthemorphologi-
cal,cytological,geographical,andecologicalcharacteristicsolthesethreetaxa,
andproposeanewcombination,B.hallii(Hitchc.)Saarela&P.M.Peterson.We
includesynonymies,speciesdescriptions,distributions,andspecimensexam-
inedforB.grandis,B.hallii,andB.orcuttianus,andkeystothesectionsofBromus
inCalifornia,andBromussectionBromopsisinCalifornia.
MATERIALANDMETHODS
Thisstudyisbasedonexaminationolover425herbariumspecimensfromCAS,
DAO,K,OSU,UBC,UC/JEPS,US,W,andWTU,includingmosttypespecimens.
Specimensweremeasuredtogeneratemorphologicaldescriptions.Dataongeo-
graphicdistribution,elevation,andhabitatwereobtainedfromherbarium
specimenlabels.CytologicalobservationsotB.grandiswerecarriedoutbythe
thirdauthoronlivingmaterialgerminated fromseedscollectedin1994
{Cayouette&DarhyshireC7947).Detailedproceduresforthechromosomecount
areprovidedinPetersonetal.(2002).Thetaxonomickeywasgeneratedthrough
studyoftheliteratureandrepresentativeherbariumspecimensatUS.
RESULTSANDDISCUSSION
Bromusorcuttianus,B.orcuttianusvarhallii,andB.grandisaremorphologi-
callysimilartaxathathavebeenrecognizedasdistinctforalmostacentury.
Examinationofmorphologicalvariabilityamongindividualsfromacrossthe
2000 BR1T.ORG/SIDA21(4)
rangeofthesethreetaxaindicatesthattheyareeacheasilydistinguishedbya
combinationolquantitativeandquahtativecharacteristics,andeachhasadis-
tinctivegeographicdistribution.Thedegreeofdillerenceamongthethreetaxa
isapproximatelythesame,althoughonetaxonisrecognizedcurrentlyatthe
varietallevelwhiletwoarerecognizedasspecies.Itwouldbeequallylogical
(althoughnotxalidnomenclaturalh)toclassifyindividualscurrentlyincluded
inliorculliLiniiSvarhalliiasanmtraspecilictaxonol B.{]^}iindis,il emphasis
wereplacedonvegetativeratherthanreproductivecharacteristicswhenmak-
ingtaxonomicdecisions.Tominimizesuchambiguity,eitheronewide-rang-
ingpolymorphicspeciescouldberecognized,oreachmorphologicallyandgeo-
graphicallydistincttaxoncould betreatedasaspecies.Wepreferthelatter
approach,andproposeanewcombination,B.hctll'n(.llitchc.)Saarcla&P.IM.
Petersontoaccommodatethis.Thismorphological-geographicalapproachto
circumscribingspeciesfollowsexistingtreatmentsofBromussect.Broniop.sis
mNorthAmerica,inwhichtaxawithconsistentmorphologicalcharacteris-
&
ticsthroughouttheirrangesaretreatedasspecies(e.g.,Pavlick1995;Pavlick
Anderton,inpress;Saarcla&rPeterson,inprep).
BnvnushallnandHonuftidnii.saredistinguishedmorphologicallybysev-
eralqualitativecharacteristics,includingthepresenceorabsenceolpubescence
ontheblades,thetypeol pubescence,andthenumberofculmnodes(Table1).
BroniiishaUiiischaracterizedbybladesthataredenselypubescentabaxially
andadaxially,lowersheathsthataredenselypubescentwithshort,stilfhairs
upto1mmlong,andculmswithl-2(-3)nodes.Incontrast,B.orcuilianushas
bladesthatareglabrousabaxiallyandadaxially(sometimeswithpilosemar-
ginstowardsthebase),lowersheathsthatarcsparselytodenselypilose(occa-
sionallyglabrous)withlong,softhairsupto4mmlongthatareeasilyvisible
withthenal<edeye,andculmswith2-4nodes.Previousauthors(e.g.,Hitch-
cock1Q5TPax'lick1995)notedthatthesetwotaxadifferinthedistributionol
thepubescenceontheleaves,buttlieywereapparentlynotawareoltheditler-
encesnotedhereinthetypeol pubescenceonthelowersheaths.Additional
morphologicalcharacteristicsdistinguishB.hallhandB.orcuttianus,butthere
issomeoverlapinthesecharacterstates.Bnvnushalliiconsistentlyhaspubes-
centglumes,lemmas,andbladesthatrangemlengthfrom7.5-16.5cmlong,
whereasB.orcuttianushasglumesthatareusuallyglabrousbutsometimessca-
brousorpubescent,lemmasthatrangeIromglabroustopubescent,andleal
bladesthatrangeinlengthIrom7-24cmlong.Themorpliologyof theinflo-
rescenecisthesamein bothtaxa,withstilf branehesthatareerectandap-
pressedtospreadingnotmorethan90°Iromtherachises.Goodillustrationsol
thegeneralhabitandinflorescencemorphologyofB.orcutlianusandRhallii
arefoundinPavliek(1995),l-)utthesedonotadequatelyemphasizethediagnos-
ticdiflerencesolthetypeol pubescencefoundonthelowerleafsheathsorthe
differencesineulmnodenumber.
SAARELAETAL.,BROMUSHALLIIANDRELATEDTAXAFROMWESTERNNORTHAMERICA 2001
Table1.DiagnosticcharactersseparatingBromusgrandis,B.hallii,andB.orcuttianus.
Characters B.grandis B.halli 6.orcuttianus
Leafblades pubescent pubescent glabrous,occasionally
pilosenearbase
Lowerleaf denselypubescent, denselypubescent, sparselytomoderately
sheaths hmaimrsluopngto1(--3) hairsupto1mmlong lpiolnogs,e,ochcaaisrsio2n-a4llmym
glabrous
Culmnodes 3-7 l~2(-3) 2-4
Leafbladelength (13)18-38cm 7.5-16,5cm 7-24cm
Glumesurface pubescent pubescent glabrous,scabrous,or
pubescent
Lemmasurface pubescent pubescent glabrousorpubescent
Lowerglumes, 3(1) 1(3) 1(3)
numberofveins
Awnlength 3-6mm 3.5-7mm (4-)5,5-8mm
Inflorescence flexuous,usually stiff,erect,ascendingand stifferect,ascending
branches spreading>90°from appressed,tospreading andappressed,to
culmaxis not>90°frc)mculmaxis spreadingnot>90°
fromculmaxis
StebbinsandLove(1941)reportedadiploidchromosomecounto(2n=14
forB.orcuttianus,andthiswasconfirmedbycountsbyWagnon(1952).Stebbins
andLove(1941)synonymizedB.orcuttianusvar.ha/hiwithB.orcuttit/nu.s,thus
itisuncleariftheyobtainedcountsformaterialthatwearetreatinghereasB.
halli.Wagnon(1952)wasunabletoobtainlivingmaterialofB.hallii,andat-
temptstogerminateseedsofB.halliiforchromosomecountsinthisstudywere
unsuccessful.ThechromosomenumberotB.halliiremainsunknown.
B/'omus/ial/iiandB.orcu11lanusditierintheirgeographicdistributionsand
mean(\.0elevations.Bromushalliiisanarrowlydistributedspeciesthatoccurs
insouthernCaliforniainthemountainsotKern,Fresno,LosAngeles,Monterey,
SantaBarbara,SanBernardino,andTularecounties,athighelevations(\x=6658
11;n=29).Bromusorcuttianushasamuchwiderrange,occurringinthemoun-
tainsthroughoutCalifornia,Oregon,southernWashington,easternNevada,and
southeasternArizona(Piper1906;Wagnon1952;Kearneyetal.1960;Pavlick
1995),generallyatlowerelevations(|u=5306ft;n=40).
BromushalliiandB.grandisaredistinguishedmorphologicallybyacom-
binationofvegetativeandreproductivecharacteristics(Table1).Thedistribu-
tionandtypeofpubescenceontheblades,sheaths,andspikeletsissimilarinB.
halliiandB.grandi.s,althoughtwospecimensolB.grandisseen(Wol/688cSand
2002 BRIT.ORG/SIDA21(4)
SHvcns2829)havelongerhairsupto3mmlongonthebladcsandsheaths.The
taxadilterint:heirbladelengths,numberofcuhnnodes,numberofnerveson
thelowerglume,andinflorescencemorphology.Bromusluilliihasbladesthat
are7.5-16.5cmlong[vs.(13-)18-38cmlonginB.grandisil-2(-3)culmnodes
(vs.3-7),andlowerglumesthatarel(3)-nerved[vs.3(l)-nerved].B/o/mi.shallii
haspanicleswithstill branchesthatareerectorspreadingnotmorethan90°
Iromtheculma.xis,whereasB.ii^randishaspanicleswithflcxuousbranches
thatarespreadingoltenmorethan90°Iromtheculma.xis.Inimmaturespeci-
mensol B.\!^nu]disCthosccollectedinlateMayandearlyJune),thepaniclesap-
pearsimilarinmorphologytothosefoundinB.orcultiauusandB./ud/ii,thus
thischaracteristicshouldonlybeusedonfullymaturespecimens.Agoodil-
lustrationoftheinflorescencemorphologyofB.grandi:^isfoundinHitchcock
(1951).Wagnon(1952)indicatedinhistaxonomickeysthatB.haUidiffersfrom
B.grandismhavingcucullate(boat-shaped)bladetips,butthoroughexamina-
tionofthischaracterindicatesthatseveralindividualsofB.gru/k/i.ssharethis
characterstate,makingthisanunreliablecharacterStebbinsandLove(1941)
reportedadiploidchromosomenumberol2/i=14torB.grcn]dis.Wagnon(1952)
didnothaveireshmaterialofB.grandisforchromosomecounts.Ourcytologi-
calobservationsofR^t^rcnuii.sconfirmthcdiploidnumber2n=f4.LikeB.hidlii,
B.grandisisanarrowlydistributedspeciesendemictosouthernCalifornia,
knownfromthemountainsinthesoutherncountiesandinthecoastalranges
fromSantaCruztoSanDiegocounties.Bro/iiusgrandisgenerallyoccursatmuch
lowerelevations(j.i=3267ft;n=25)thanB.haUii(}.i=6658ft;n=29).
Recognitionol B.halliiasadistinctspeciesincreasesthenumberofspe-
ciesinBromussectionBromopsisinCaliforniatofl,fOofwhicharenative,and
onethatisintroduced(B.incn}]isLeyss.).Keysdistinguishingthesectionsof
Bromusandthespeciesofsect.BromopsisinCaliforniaarepresentedbelow.A
revisedkeydistinguishingall34nativeandintroducedspeciesolBromusin
CaliforniawillbepublishedintheSecondL.ditionofthejepsonN4anual(Saarela
andPeterson,inprep).
TAXONOMICTRHATMHNT
Bromusgrandis(Shear)Hitchc,Fl.Calil.1:175.1912.Ihomusonuttiamisy'dv.^uatdis
Shear,Bull,Div.AgrostoLU.S.D.A.23:41lOOOHrn/Mo/isis^i^rcduiis(Shear)Mdlub,rdluK.cobot
Phytotax.8t2):167,1073.Tvi'l':I'.S.A,CAl.iroRNlA..SanDiegoCo.:LaMaite.25KmIHHlC.R.
Orn(((472(iioi(TrYi'i::liS-8UTlV).
H}-oiuu\porlcnvar,assimiUsBuntOavy,Univ.C;alil.Publ.Bot1;55,1902.TVPH:U.S.A.CALll-ORNlA:
southside,San|aeinioMts,1001.M.M.Hall2228(HOLOTYPEUC-37692!:1SOTYP1-:US-865453
Iragm.e,\I'(').).
Looselycaespitoseperennial,CaiIhis70-180cmtall,erect,glabroustopubes-
cent;nodes3-7,darkbrown,rctroi'sclypilosebelow,oftencoveredbysheaths.
Leal sheaths3.5-12cm long,closedlormostoftheirlength;sheathsdensely
SAARELAETAL.,BROMUSHALLIIANDRELATEDTAXAFROMWESTERNNORTHAMERICA 2003
pubescent,hairsupto1mmlong;marginssmooth,occasional!yhyalineatapex;
collarsdenselypilose,withhairsupto2mmlong,auriclessometimespresent;
cataphyllsandbasalsheathspubescent,sometnnesshredding;ligules1-3mm
long,membranous,denselypubescenttopiloseadaxially,glabrousabaxially,
apexobtuse,erose;blades(13-)18-38cmlong,3-12mmwide,Hat,membra-
nous,apicallycucullate,sparselytodenselypubescentadaxiallyandabaxially,
hairsuptol(-3)mm long;marginsglabrous,sparselypapillateorscabrous.
Panicles15-26cm long,6-17.5cmwide,open;lowerbranches5-26cmlong
with1-6spikelets,flexuous,noddingandusuallyspreadingmorethan90°Irom
therachises,pubescent;lowerinflorescencenodeswith1-4branches.Spike-
lets2.5-3.5(-4.5)cmlong,4-9-flowered,teretetoconspicuouslydistichousat
maturitywithrachisvisible;glumessubequal,shorterthanlemmas,pubes-
cent,marginssometimeshyaline;lowerglumes5-8.5mmlong,3(l)-nerved,apex
acute;upperglumes7-10(-12)mmlong,3(5)merved,apexacute;lemmas11-14
mmlong,marginspubescent,backsglabroustodenselypubescent,marginal
hairssometimeslonger;backsflatteneddorsally;apexentireorminutelybifid,
theteethnotgreaterthan0.2mmlong;awnedjustbelowapex,awns3-6mm
long,straight;paleas8-11mmlong,shorterthanlemma,backsglabrousor
slightlypubescent,marginsusuallypubescent;apexacute;anthers3-5mm
long,yellowish-orange,Caryopsesabout9mmlong,linear,darkbrown.Chro-
mosomenumber2n=14.
Distribution.—EndemictosouthernCalifornia;SanGabriel ]\4ts.,Santa
LuciaMts.,SanBernardinoMts.,SanAntonioMts.,SanJacintoMts.,SanRafael
Mts.,andTopatopaMts.
Habitat.—Dryareasmopenpinewoods,hillsides,androckyslopes;eleva-
tion1200-8000It.
S\^p)e2c\iDmuenncsaenxas.mni.n(eCdA.S)U;.SK.mAg.sCaClainfoyronniar:oFarde4s.n.3omCio.W:joufncItlioornseNshoofeSBoeuntdh,f3to)r0k0offt,K6iJnugnsfR9i6v0e,r,H1owN4daly
35^^0(CAS);Kaiser,27junf935,Bullard95(UC/JEPS).LosAngelesCo.:SanDiinasCanyon,San
GabrielMts,1750ft,4Jul1Q33,Wheekr1894(CAS,US);SanGabrielMts.3500ft,4Julf933,Ewan
7857(US);SanGabrielMts,FastFork.BigSantaAnitaCanyon,2250ft.30May1931,Ewan4249(US);
LittleSantaAnitaCanyon.2July1902,Abrams2632(CAS,US);SanGabrielMts.BellCanyon,2300
ft,8Jun1936,W/iecler4134(CAS);SanGabrielMts,IndianCanyonCamp,2mioffSoledadCanyon,
3000ft,27May1936,Wo!f7845(CAS,W);SanGabrielPeak,9Jul1900,Dudleyi.n.(CAS);SanGabriel
Mts.,990m,1Sep1994,Cayouette&DarhyshircC7947(DAO);Summitol'Mt.Wilson,29Jun1902,
Abrams2600(CAS,US);SanGabrielMts,BigTujungaCreekbetweenColdwatcrandWickiupCan-
yons,fOJul1992,Ross667l(UC/JEPS);HorseFlats,SanGabrielMts,16Aug1991.Koss5873(UC/JEPS);
RoundtopMtn.SanGabrielMts,ChilaoFlat,14Aug1991,Ross5780(UC/JHPS).MaderaCo.:0.5miS
of-Southfork.17jun1933,Hormay26(UC/JEPS).MontereyCo.:PicoBlanco,11May190f.Davy7339
(UUSS));;LiStatnlteaSuLru,c1i4aMMatsy,1J9u01n,1D9a01v,yD7a3v8y5(7U6S9)1;(cUoSa)s;tTraasnsgaejaWraoHfoKtinSgprCiingtsy,RJJuunn11990217.,HSlwmacllre3n359986((CUASS),;
GavilanPeak,3000ft,1Julf86LBrewer740(US);AlderCreek,SantaLuciaMts,3000ft,23Jul1958,
Hardbam3789(CAS);ArroyoSecoRiverCanyonnearHangingValley,2Jun1957,HowcJl32566(CAS);
ArroyoSecoRiver,SantaLuciaMts,5Jun1959,Hardbam4796(CAS);CruikshankTrail,SantaLucia
2004 BRIT.0R6/SIDA21(4)
Mts,26Ma)'IOh\Howill1580(CAS);FrancesSimesllastingsNaturalHistoryReservation,Santa
IiiciaMts,22JulK)44.Jjii.stlule.s.n.(CAS);1-raneesSimesHastingsNaturalHistoryReservation,up-
pei"t'armclVallc)',about2miNPolJame^burj;.22Augl'-H2,Durhams.n.(C;ASJ;NaciinicntoSum-
inii,Santa],ueiaMts,2801^1it,liS|uni^)n^.Mini;20886(CAS);nearCarmclRiver,261un\^)0i.Dudley
s.iL(C;AS);nearNaeiniienloSiiininitCamp,SaniaLuciaMts,26Jun\^)V,Raven10987(CAS);Santa
LuciaMts,bankbyroadfromArroyoSecoCamptoEscondidoCamp,24jun1956,HowiK980(CAS);
SantaLuciaMi>,betweenMiddleandSouthForksofIlevil'sCreeknearC;anogasFalls,2000It,23
Mayl'-)83,C((f7'e/i(cr2f8(,CAS);SantalaiciaMts,Landels-1[illBigCreekReserve,CamboaPointSec-
twiiotnh,2A7nMthaoynly'-)C8r2,ee(ikc,ncciat.i5cavir/:u,ki;m/e,'W>,3N9.WUCoAfS)S;aSnanMtiaguLeulciitaoMRtas,ncShanRuMiingsu,el39C0reeml,<,5bjeulnow19c99o,nfWliuleknecne
15707(CAS);SantaLuciaMts,,'^outhForkOevMsCaimm,alongGamboaTrailnearsaddleotTwin
PeakandLeak3850,4junl'-)81Ccir/H-ndT2,57(CAS);SantaLuciaMis,lunDOl,/3(n'V76.5.3,7718.7720
(.UC/JFPS),OrangeCo.:Santiagopeak,20)uni'-)27,Swalien640(US);SilveradoCanyon.,640m,3
Sep1994,CayoueUe&DavbyshuvC79,5,i(HAC);SantaLuciaMts,MillCreek,25May1941,tioover
5260(K).PlacerCo.:NbrancholMiddleForkofA\mericanRiveratroadcrossingFoiForestHill,20
lulK)52,5(t4'l'i/isetal51.12(CAS).RiversideCo.:SanJacintoMts,khJUvild.Area,NofStrawberry
Valley,betweenBearTrapC\nandLilyCj-eek,5200-5500ft,IAug1996,White^442(CAS);Colo-
r;idoOcsert,1889,P,(/mcrs./i(US);FullerCreek,29Jun1936,rales6,38,3(UC/JEPS).SanBeniioCo.:
PinnaclesNationalMonumeni,3|unK)3^),Stebbins2715(UC/JFPS).SanBernardinoC:o.:SanIk'r-
ii.u\linoMts.29May&-28|un1888,Pti/is/i20.5.HUS);SanBernardinoMts.,.330011,15|un.ReedlOIl
(US);SanBernardinoMts,29May1890,I'ciislis.n.(US);.SanBernardinoMts,roadbelowHighland
andRunningSprings,26Jun142,Bccilc.i645(L:BC);.SanAntonioMts,8000ft,28jull917,/()/ins;()n
/4t'7(UC/JEPS,US);NofSnowc;anon,5500ft,20JunDOLI'ansb.50.i8(C;AS);SailBernardinoMis,
250011,5JunL)17,Fun.'.li/I.itH(CAS,UC/JFPS);Sanf5ernardinoMis,roadbetweenHighlandand
RuiniIngSprings,26junf942,BcciIf3645(CAS);SanBernardinoMts.24May1888,F(i?-|,s/i,s,ir(C;ASJ;
.SanBernardinoMts,3000ft,29Jun1888,Finish20.5.3(CAS);SanBernardinoMts,Dobbstrail,Mill
Creek,2|ul,C.yawjoyd.s.n.(CAS);SanBern.in.linoNtlI'oresl.road38.nearAngclus,1654m,2Sep
1994,C((\'(iiic((cif-Pt/r/'v.sliMcC7949(DAO);0,5miSofSantaAnaRn-er26Junf937,>(;le,s6688(UC/
IFPS)SanDiegoCo.:ClevelandNationalForest,30Jun^'H,HiitbeiKk13160(US),SanLuisObispo
Co.:beiweeiiRockyButteandPineMt„SantaLuciaMts,21Jun1950,Hoover7998(CAS,UC/JF.PS);
nearRockyfUitteFireLookout.12JuneF)64.Hoover9076(CAS).SaniaBarbaraCo.:SanRafaelMts,
PolreroonSFsideolCachuiiiaIVak,400t)i1.12|un1961,BidIclcv4491(CAS,US);NsideolLaCumbre
SP19ae5an1kt,,aPSoYalnnletasardMYson.une.nzi(CaMAitSns),s;.3W650m0kiiIrt1k.;o5oflDSCeaocnltd1a9S5Bp8a,rri1bn'agorUaCa,arnd1y2so.Snn.e.p(CS1Aa9Sn9)t4;a,RBCaoarmybeoaurraco.KCcIa6n&jyuoDnna,1r9bS6y5a,nstbPanoleYlna(re.Jdz9s7M.1nt.s((,DCA7AOSJ))u;l;
bankofCammoCielo,FofandnearSanMarcosPass,2000ft,fOjunF)58,Pollard.sn.(\V);ZacaLake
ForestReserve,19-30JunDOo.Pastwood719(UC/JEPS).SantaCruzCo.:nearEagleRock.2500ft,
HF.casr.Jsyc2I7nt3a1ke(CdA,Su)n;.S1a0ntmaiCWruzo,f2M7atJhuner19138J,ul.S1i9l5v1c,u.sS(c2/8)/2)9M1.(sC.A5S0)0.0T(uUCo/ljulm4\nSe).CoV.e:ntTuuroaluCom.n:eRReidveRreeaft
Canyon.TopatopaMts.2800 5500IJ,81tin1908.Al'ni/ti.s&McGre.^oi"/6/(US);aboveMurietla-Sauia
Ynezdivide,SantaYnezMts,OjaiDistrict.4500ft.27Jun1963.Po/dods.n.(CAS);MatihjaCanyon,3
|unl945,Poll(:irti,s./i,(CAS);MurrietaCanyon.l5Jun1946,Po/dod.s.n,(CAS);RedReefCanyon,Topatopa
Mts,2800-3500h,8Jun1908,Abeams&MeGre^or161(CAS);Santa^'nezMts,CammoCiclo,3500ft.
29Jul1967,Pollards.n(CAS);UpperNorthFork,MatilijaCanyon,OjaiDistrict,27Jun1962,Pollards.n.
(CAS);VenturaRiverBasin,CammoCielo,4(XX1ft.25May1^)46.Pollards.n.(CAS);SantaYnezMts,Cljai
toCuyamaValleyRoad,NForkofVenturaRiver.2.2mibelowWheelers1lotSprings.1200ft,2fMay
1935,W()I/'68,S8(CAS);0.5miNofWhiteacrePk,LSJun193'S..SmionlduIn(20(UC/JEPS).
Broinushallii(Hitchc.)Saarela&P.M.Peterson,comb.nov.Broriiicsonuitianusvdr.
haliiiIliiclic.Fl,Calif1:175.1912.Tvri-:U.S.A.Cai.II-ORNI.A;westside..SankicmtoMts,27Jun
1901,i/.Af./iti/!2,301(1KM.OTYPi;:US-412894!).
SAARELAETAL.,BROMUSHALLIIANDRELATEDTAXAFROMWESTERNNORTHAMERICA 2005
Looselycaespitoseperennial.Culms90-150cmtall,erect,puberulentorocca-
sionallyglabrous;nodesl-2(-3),darkbrown,retrorselypilosetodenselypu-
bescentbelow.Leafsheaths6-12.5cmlong,1/4-3/4aslongasinternodes,closed
formostoftheirlength,denselypubescent,hairs0.3-1 mmlong;margins
smooth,occasionallyhyalineatapex;collarsdenselypilose,withhairsupto2
mmlong,auriclesabsent;cataphyllsandbasalsheathsdenselypilose,some-
timesshredding;ligules0.5-2.5mm long,membranous,sparselytodensely
pubescentadaxiallyglabrousabaxiallyapexobtuse,erose;blades7.5-16.5cm
long,3-12mmwide,flat,membranous,apicallycucuUate,denselypubescent
abaxiallyandadaxially,hairs0.2-0.5mmlong,occasionallylongerhairsupto
1.2mmonmarginsnearbase;marginsscabrousorsmooth.Panicles5-16cm
long,2-11cmwide,opentodenselybranched;brancheserect,ascendingand
appressedtospreadingnotmorethan90°tromtherachises,pubescent;lower
inflorescencenodeswith1-2branches,lowerbranches3.5-11cmlongwith1-
2(-3)spikelets,upperspikeletsoccasionallysessileandspike-like.Spikelets2.5-
3.5(-4.5)cmlong,3-7-flowered,teretetoconspicuouslydistichousatmaturity
withrachisvisible,lowestrhachilla2.5-4mmlong;glumessubequal,shorter
thanlemmas;sparselytodenselypubescent,marginssometimeshyaline;lower
glumes5-8(-9)mmlong,l(3)-nerved,apexacute;upperglumes(7-)8-9mm
long,3-nerved,apexacute,acuminatc,ormucronate,themucrolessthan1mm
long;lemmas10-14mmlong,5-to7-nerved,sparselytodenselypubescent
acrossback,marginalhairssometimeslonger,upto0.5mmlong,backsflat-
teneddorsallytoslightlykeeled;apexentireorminutelybifid,theteethnot
greaterthan0.2mm;awnedjustbelowtheapex,awns3.5-7mmlong,straight;
paleas9-13mmlong,shorterthanthelemmas,backsglabrousorslightlypu-
bescent,marginsusuallypubescent;apexacute;anthers3-6mmlong,yellow-
ish-orange.Caryopses9-11mmlong,linear,ambertodarkbrown.Chromosome
numberunknown.
Distribution.—EndemictosouthernCalifornia;knownonlyfromFresno,
Kern,LosAngeles,Monterey,SantaBarbara,SanBernardino,andTularecounties.
Habitat—Dry,openorshadyareasonhillsides,rockyslopes,andpine
woodsinthemountains;elevation5200-8800ft.
Specimensexamined.U.S.A.California:I'rcsnoCo.:betweenVicletleMeadowandBullfrogLake.9
Aug1940,HowcU16079(,CASJ;BubbsCreekCanyonuivicniityofVidetieMeadows,9,500-10.000ft,
21935J8u,lHo19w4e8l,lH3o3w9e2l6l(2C4AS9)4;2W(oCAoSd)s;CSriemepks,o1n9JMuela1d9o10w,.CMliedmdelnesFs.onr.k(CoAfS)t.heKeKrmngsCo.R:iv1e.r5,m6i0W00offt.G2r5eeJnu-l
hornSummit,5200ft,22Jun1970,Howell&Inic46599(CAS);BreckenridgeMountainRoad,0,4mi
F-olBarrelSpring,6500ft,19Aug1958,'iwissclivann4725(CAS);GreenhornMountains,1.5miSFof
Summit,5000ft,BJul1962,Howell383I2(CAS);GreenhornPassRoad,about0.5miEofSproutSpring,
GreenhornRange,5750ft,13Aug1958,Twtssclmann4666(CAS);GreenhornRange,NVv'ofCalfCreek,
17Sep1963,Twisselmann9010(CAS);GreenhornRange,GreenhornPassRoadjustSofTigerFlat,
6600ft.13Aug1958.Iwissehnann4653(CAS);KernPlateau,roadtoBartolasCountryEofLittle
CannellMeadow.7200ft.12Jul1966.Howell&True41851ICAS);KernPlateau,trailtoLittleC:anncll
2006 BRIT.0R6/SIDA21(4)
MeadowabovePineFlat,7450It,24Jul1964,Twissclmunii9X80(CAS);OldKernvilleRoad,4.4miF.
olGreenhornSummit.GreenhornRange,5200ft,21Jun1957,Twissclmann3716(CAS).LosAngeles
Co.:Buckhorn,SanCuibnclMts.0500It,20Jul19)].Duidn.i52HCAS.OSl!,US);SanAntonioMts.S
Forkoll.ytleCreek,dOOOIt,15Jul1917,/o/m.sdxi14.54(CAS,US);SanAntonioCanyon,27Jun1927,
Swollen679(US);SanBernardinoMts,2miNolBigPineCamp,25Jun1942,Beetle3640(CAS);San
GabrielMts,BlueRidge,8000-8500ft,I8jul1947,Howell23389(CAS);SanAntonioMountains,Prairie
ForkofSanGabrielRiver,6500ft,6Jul1918,/i)liiis(()n2070(CAS);SanGabrielMis.,1820in,1Sep
31399144,(CCaA^S',ouKc,lO/eSU&,•DU(Si)r;|)vS,asn/itiareLuC7c9i4a(SM't(sO,AJOu)n.19M0o1i,uDcarvcyy7C7o0.:9lTUaSs)s,aj7a7r1a0(IOloStU)S;prNinWgs,olJuTnass1a9j0a1.raIiRlo/iaiedr
summit,CJiewsRidge,LosPadresNationalForest,15Jun1973,Giijjin362,5(UC/JFPS).SanBernar-
dinoCo.:SanBernardinoMts.LittleBearValley,Aug1907,VVi/!)i(r107/tUS);SanBernarelinoMts,22
JSBuaalnrt1J9oa0nc2i,FnlAtatols),Mrtt2s(2,m5sW127mf9,o9r2k(CSoAefSpS,1nK9o)9;w4,SCaCrneaeyBkoe,uren5ta0tr0ed0i&n1o1,nMJauiylsb.v19iB0he1ia.rrHeVua(lll7le92y5.,50368(6D0(AC0OAf)St;,,fKS9)aJ;nuSlBa1en9r0Gn0,oajrr(g)d/oiinne.ioso.Ms.tM»ist.s,(.C,DAeSSe)op;f
CSrweaelkl,en6570000(ftO,SU19,JUuSlj;1289m9,iHWaoilfIB.aMSr'toUnO;Flaatl,xi2v6eJAurnr1o9w3h7.ea)d;((Lca,kse6,69.S4a(nUCB/eJrFnPaS)r.diSnaoniMatsB,arIbJaulra1c9:2o7.,:
MissionPine,SanRafaelMts,6200ft,25Apr19^0,Hoffmau90(US);SanBernardinoMts..Sam,iAna
CanyonaboveCdarksRanch,7400h,|ul1920,Oiii|icH59(US).TulareCo.:trailIromBakeovcnto
TempleionMeadows,8800It,17Jul1950.Howell27005(LIS,CAS);KaweahRiverValley,29lul1801,
Covi11c(S'Fun.sd)ItJ346(US);KingsCanyonNaiionalPaik.belowMistballs,S.ForkKingsRiver,13Jul
1927.Swallen770(US);CopperCreektrailtol.raiiitct^a^n,11Jul1927,Swollen7.53(US);Aug1897,
Ihtdleys.n.(CAS);KernPlateau,about2miN\VofBeachMeadow;8200ft,7Aug1967,HowellfvTrue
4.5419(CAS);KernPlaieau,NofCainMeadow,ca.7200ft.24|un19oo,Iwisselmonn12421(CAS);
KernPlaieau,NorthManterCreek,BigMeadow,7700-8000ft,24Jun1970,HowelliS'TriJC46696(CAS);
MonarchLakesTrail,ca.8000It,19Jul1951,Howell27943(CAS);PortugesePass.7400ft.5Aug1957.
iwisselman3946(CAS);QuakingAspenRoad,1.5miNolleeCaTck,6600ft.8]ul1959,Iwi.s.seloiunii
5497(CAS);Se(.|uoiaNaiionalForesi,Llo\'dMeailowsBasin,1.25miWoltheP)JesBoysCampinihe
FreemanRedwoodGrove,5800ft,19Jun1973.^bevoch2770(CAS);SequoiaNationalForest,Lloyd
MeadowsBasin,alongUSPSroad22-S82,HbaseoftheNeedles,about3mifromthePylesBciysCamp,
m5a8t0e0lyft,24mJiunNW197o4f,tShleiePvyolde;s3B.o5y8s0C(Ca>m\pS).;5S7e0qtu1oliia,1N0atluilonUa)l74,FoSrleisrvt(,>c!L.l'o.y5d716M(eCaAdSo);ws.SeBqausoiina,Naaptpiroonx.ii-l
Forest,nearBelknapCreek,26-28Jul1441.lioii\^iilup>.Wiggins.&Ferri.s2655(CAS);SequoiaNil
Forest,NlimitofSequoiaNt!Parkroad.2010in.5.Sep1994,Cawuette&DarbyshiieC79.55(DAO);
LongMeadow8000''-H^OO11.14Jun1888.l\(lmcr2.5.5(K,US);Mt.Sillunan.ClovertJrek,29Jul1806,
PuilleyH.S'KCAS).
BromuFsolioar(c.uretotboita.nPuhsyioVtaasxey8,(2)B;ol6t8..G1a97z3..T1v0i':i;2:23U..S.1A.88t^5.,.'\t.iBiwoni^inoipa:simu>tnhue!mnoainmai(aVuaisseyn)eairlolSuabn.
Diego.1884,C.R.Omii((.s.n.(IKM(vrvi'i.-;tis!),
Bioinii\|i)ii(/iv/4ivl/iisMerr.Rhodora4:140.b)02.J\cr:l^.S.A.Orii.on:CrookCo.:BlackButte,
opendrypineforests.19|ulU'Ol,Cmnk2677(llOl.OTVl'i::unknown;t.SOTVl'i;:US!).
Looselycacspitoscpcfcnnial.CAilms^)0-150cmtall,erect,glabrou.stopubes-
cent;nodes2-4,darkbixiwii,retrorsch'pilosetodenselypubescentbelow.Leal
sheaths.3.5-11cmlong,usually1/3-2/.3aslongasintcrnodes,closedlormost
ol theirlength;lowersheathssparinglytodenselypilose,hairs2-4mmlong,
ogicncsassiionnoaoltlhy,ogclcaabrsoiuosn;alulpypheyralsihneeatahtsappeuxb;ecsoclelnatr,sphialiorsseuwpitthoh1amirmsulpontgo;4mmarm-
longorglabrous,auriclesabsent;cataphyllsandbasalsheathsglabrous,some-