NEW COMBINATIONS IN FESTUCA CALIFORNICA (POACEAE)

2005·3.8 MB·English

by  Stephen J Darbyshire

NEWCOMBINATIONSINFESTUCACALIFORNICA(POACEAE)
StephenJ.Darbyshire
AgricultureandAgri-FoodCanada
CentralExperimentalFarm,Building#49
Ottawa,Ontario,KIA0C6,CANADA
ABSTRACT
TheFcstucacalifornicaVaseycomplexiscxaniined.Considerablemorphologicalvariationexists
withinandbetweenpopulationswhichrangefromwesternOregontosouthernCalifornia.Thechar-
actersofsheathpubescence,sclerenchymadistributioninthebladesolvegetativeshootsandligule
lengthareofusemdistinguishingthreetaxamthecomplex.Analysisofcharactervariationand
geographicdistributionsuggeststhatthesetaxaarcbesttreatedattherankofsubspecies;thenew
combinationsh'cstucacalifornicasubsp.hUcluockianacmdF.calijorr^icasubsp.pariihiiareprovided.
RFSUMHN
Seexaminaelcomplejob'eiliicacalifornicaVaseyExistcunavariacionmorfologicaconsiderableeny
entrepoblacionesquevandesdeelOestedeOregonhastaelSurdeCalifornia.Loscaracterespubescencia
delavaina,distribuciondeesclerenquimaenlaslaminasdeloslallosvegetativosylongituddelaligula
seusanparadifcrcnciarlostrestaxadelcomplejo.Elanalisisdelavariaciondelcaracterydistribucion
geograficasugierenqueestostaxadebentratarseenelrangodesubcspecies;Sehacenlasnuevas
combinacionesFcstucacalijornicasubsp,hitchcochianayFcalilornicasubsp.parishii.
Californiafescue,FestucacalifornicaVasey,isthelargestNorthAmericanspe-
ciesofthegenusandshowsconsiderablevariationinmanymorphologicalchar-
acters.ItISusuallyreadilydistinguishedfromotherNorthAmericanspecies
ofFestucabyitslargesizeandthepresenceofstiffhairsontheleafcollarre-
gion.Thesehairsmaybeabsentoverthebackofthesheathapexandrestricted
tothemargins(auricularregion).Occasionallycollarhairsareabsentaltogether
onsomesheaths,butexaminationofanumberofleavesonthesameplantwill
usuallyrevealhairsonthecollarmarginsofatleastsomeleaves.
Morphologicalvariation wasexaminedon113specimensofFestuca
californicasensulato,includingthreetaxathataresometimesrecognizedas
separatespecies(e.g.,Alexeev1982;Aikenetal.1997).Variationincharacters
usedtodistinguishtaxadidnotconsistentlycorrelatetoallowtheunambigu-
ousrecognitionofseparatespecies,althoughthreeinfraspecifictaxamaybe
recognizedwithsomeconfidence.Inadditiontotheirmorphologicaldifferences,
theyhavelargelyseparatedistributions,hencearebesttreatedassubspecies.
Unlessotherwisestated,thedescriptionsofleavesrefertothoseofthesterile
vegetativeshootsorinnovationswhichformalargeproportionofthedensely
caespitoseclumpsofthesebunch-grasses.Thewidthofconvolute,involuteor
conduplicateleafbladesisgivenasthewidestdiameter
SIDA21(3):14SS-1463.2005
1456 BRIT.ORG/5IDA21(3)
FestucacalifornicaVasey,Contr.U.S.Natl.Herb.1(8):277.1893.Typi::U.S.A.Caliicir-
NlA.Claklandhills,18C)2,H.N.HoltindiTISO.'UilOi.OTYPH:b'S-55(-)2l2!).
Ihvmn^kiilmiivariinstiihitus'lorr.PacilicRailnRep4l,T):l'57.1857.}-i-\tULcianstuJtito(Torr.)
Shearr.vI'ipL-r.t;ontr.U.S.Natl.Herb.I0tl);i2,1406.I'csluuunhamisubsp.cuailaimvan
anstulata(Torn)St.-Yves,Candollea2:273.U)25,Tvi'i;:L'S.A.CAi.lFORMA.MarkWest'sCreek,
30Aprl834,j.M,Bi^^clows.i].Cl.liciOTVPi.designatedbyPiper1Q0(t33:US-55621H:LsoiirioiYPi-.:
Gl-1).
Plantsdenselycaespitosc,withoutrhizomes.Culms(30-)60-L50(-275)cmtall;
internodesglabroustopubescent(oltenpubescentforupto20mmbelowthe
nodes).Prophylls2-9cm long,scabroustopubescent(atleastapically)and
usuallyscabrousorciliatcontheveins.Sheathsopentothebase,marginsover-
lapping,glabrousorscabroustopilose(sometimesonlyattheapexoronupper
margins),persistent;collarsusuallydenselypubescent,sometimesinconspicu-
ouslypubescent,withalewhairsatthemargins,orglabrous;ligules(0.2-)0.8-
l.5(-5)mm(ligulesofthecaulineleavestendtobesomewhatlongerthanthose
olthevegetativeshoots),usuallyciliateatapex,aba.xialsurfaceglabroustopu-
bescent;blades(0.5-)0.8-2(-2,5)mmwide(3-6.5nunwidewhenflat),convo-
lute,involute,conduplicateorflat,abaxialsurfaceglabrous,scabrousorbasally
pubescentorpilose,adaxialsurtacepubcrulenttodenselypubescent-piloseor
moreorlesstomentose,9-17(-19)veins,(3-)5-17(-19)adaxialribs;sclerenchyma
inamoreorlesscontinuousabaxialbandorring,usuallywithpillarsorgird-
erspresentatmostveins,sometimesreducedtosmallstrandsatabaxialand
adaxialsurlaces(Fig.1).Inflorescences(10-)15-2.5(-30)cmlong,open;branches
tereteorangular,glabroustoscabrous-pubescent(especiallyonangles)and
sometimespubescentatthebaseintheaxilsoilowerbranches,spreadingand
lax,(l-)2(-3)pernode,lowerbrancheswithadistinctpulvmus.Spikelets(8-
)10-l8(-20)mmlong,bornetowardtheendsofthebranches,usuallygreenor
sometimesreddish-purple,with(3-)4-6(-8)florets.Glumeslanceolate,glabrous
orsparselyscabrousatapex.Lowerglumes(3.5-)4.5-b.7(-8)mmlong,fvein;
upperglumes(4.5-)5-10mmlong,3veins;lemmacallusindurate,widerthan
long,scabrouslaterally;lemmas(5^)7.5-1l(-f1.5)mmlong,lancetilate,scabrous,
orpuberulent(rarelypubescent),sometimesminutelybidentate,acute,usu-
allywithanawn(0.5-)l-3(-4)mmlong;paleasslightlyshorterthantoslightly
longerthanlemmabody,emarginateorbidentate,glabrousorpubescenton
themargins,scabrousorpubescentbetweentheveinsapicallyorthroughout,
theveinsscabroustoaboutthemiddleorscometnnesrighttothebase;rachillas
scabroustopubescent(sometimessparsely)atleastonabaxialside(awayfrom
palea),usuallyglabrousontheadaxialside;anthers3,(3-)4-7.5(-8.5)mm;ovary
apexpubescent.
Distributionandhci/);ld/.—L^ryopenslopesandmoiststrcambanksinchap-
arral,thickets,openlorestsandforestopenings(coniferous,oakormixedfor-
ests).Ottenonultramaficsubstrates.Sealeveltoabout1500melevation.From
DARBYSHIRE,NEWCOMBINATIONSINFESTUCACALIFORNICA 1457
Fig.1.CrosssectionsofleafbladesfromvegetativeshootsofFestucacalifornica(A,Howell&True42564,RSA300178;B,
Crampton4882,AHUC2500V,C,Hoover8864,RSA201564;D,Roos92,RSA-POM261420;E,Banks&Boyd0509,DAO796417;
F,Banks&Boyd0429,DAO796414).A-B.Subsp.californica.CSubsp.hitchcockiana.D-F.Subsp.parishii.Sclerenchyma
tissuedistributionindicatedinblack.
westernOregontothePalomarMountainsinsouthernCahfornia(Fig.2).his
reportedasfarnorthasWashingtonCounty,Oregonintheonlinespecimen
databaseof05U(oregonstate.edu/dept/botany/herbarium/db.php).
KEYTOTHF.INFRASPECIFICTA.XAOFh'ESTUCACALIFORNICA
]. Culms30-80(-l00)cmtall;internodesusuallypubescentfor5-20mmbelowthe
nodes;lowersheathsusuallydenselyretrorsepubescent,sometimessparselypu-
bescentorglabrous;blades0.5-F2(-1.5)mmwide,conduplicate(sometimes
loosely),with(3-)5-9shallowadaxialribs(toabouthalfasdeepasbladethick-
ness);sclerenchymainsmallabaxialstrandstoacontinuousbandandadaxial
strandspresentorabsent,pillarsrarelyformed,girdersabsent;spikeletswith3-4
florets;plantsoftheSanGabrielandSanBernardinoMountainsandsouthward
subsp.parishii
i
1458 BRIT.ORG/SIDA21(3)
+=subsp.californica
1^=subsp.hitchcockiana
0=
subsp.parish!
40'
36*
32'
124' 120' 116'
Fig.2.DistributionofspecimensofFestucacalifornicaexaminedinthisstudy.+=subsp.californka.JV=subsp.
hitchcockiana. ^=subsp.pons/)//.
2
DARBYSHIRE,NEWCOMBINATIONSINFESTUCACALIFORNICA 1459
1, Culms60-180(-'275)cmta!l;inlernodesglabrousorpubescentforlessthan5mm
belowthenodes;lowersheathsusuallyglabrousorsometimespubescent(the
hairsspreadingorsometimesretrorse);blades0.8-2.5mmwide(3-6.5mmwide
whenflat),convolute,involute,conduplicateoroftenflat,with7-15(-19)deep
adaxialribs(usuallymorethanhalfasdeepasbladethickness);sclerenchymaina
continuousabaxialbandandwithadaxialstrands,formingpillarsorgirdersatmost
oftheveins;spikeletswith(3-)4-6(-8)florets;plant5morenortherlyindistribution.
22.. LmLiimgguullleeossnaga;sllhooenrgmtemrcaimlsiea(tm5e-b)mr7e.am5n-be1r(1aunmseumal0l.ly2o-n11g-(6-1m.2m)mlomng)l,oncgil;isatpeikoerlentost;(s8p-i)k1e2l-e1ts8(s8-u-2b10s)p.californica
(-17)mmlong;lemmas7-8(-8.5)mmlong subsp.hitchcockiana
Festucacalifornicasubsp.californica
Culms60-180(-275)cmtall;internodesglabrousoroftenpubescentuptoctbout
5minbelowthenodes.Sheathsglabroustopubescent;collarsusuallydensely
pubescent(atleastatthemargins),sometimesinconspicuouslypubescentor
glabrous;ligules0.2-l(-1.2)mmlong,aciliatemembrane;blades0.8-2mmwide
(3-5mmwidewhenflat),convolute,involute,conduplicateorflat,with7-f5
(-19)deepribs(usuallymorethanhalfasdeepasbladethickness);sclerenchyma
acontinuousabaxialband,withadaxialstrands,usuallyforminggirdersatmost
veins(Fig.la,b).Inflorescences15-2.5(-30)cmlong.Spikelets(8-)12-18(-20)
mm,with(3-)4-6(-8)florets.Lemmas(5-)7.5-llmmlong,usuallyentire,or
sometiiuesminutelybidentate.Chromosomenumber;2n=56(Stebbins&Love
1941).
Thisisthemostwidespreadsubspecies,occurringfromnorthwesternOr-
egonsouthtosouthwesternCalifornia.Theleafsheathsofthevegetativeshoots
aretypicallyglabrousorscabrous,althoughplantswithafewpubescentsheaths
arenotuncommon.Thehairsare,however,usuallynotstronglyretrorseand
areoftenrestrictedtowardthetopofthesheaths.Plantswithpubescentleaf
sheathsdifferfromsubsp.parishiiinpossessingsclerenchymagirdersatthe
majorveinsintheleafblades.Ligulesareaciliatemembrane,usuallyabout0.5
mmlongorless.
Selectedspecimens:U.S.A.CALIFORNIA:AlamedaCo.:WestBerkeley,1916.P.B.Kennedy,(AHUC
N57i1l5e)s;CBaenrkyeolne,y,3S.5hamsitaWRooafd,SuNnolf,acriunbgbglryassshyalselospleo,peIs,Jucnoa1s0t3^a-l),sGa.gLeaSstseobcb,in4sjJru.n27149358(,ABHCUrCam6p26t5o)n;
4882(AHUC25001).ButteCo.:NWsideofFeatherRiverCanyonnearPulgaBridge,serpentine
canyonside,elev.1400ft,11JunIQ67,/.IHowell&G.H.True42564(RSA300178);about2miSWof
ForbestownDiversionDam,NEofForbestown.yellowpineforest,ondampredsoilonsteeproadcut
nearsmallstream,lOJun1984.LAbart&P.Ahart4701(RSA-POM365944),ContraCostaCo.:Mount
Diablo,nearjunctionolsouthandwestroads,drybankandroadside,25Apr1931,ML.Bowcrman
757(RSA-POM300669).DelNorteCo.:alongSmithRiverat18-mileCreek,3miEofGasquet,in
chapparral,alt.500ft,24May1933,j.RTracy12292(RSA10111);T17NR3ESec.32,atjunctionof
roadstoBearBasinButteandBigFlat.Douglaslirwoodland,elev3500ft,27Jun1976,/^Smith8848
(RSA-POM365942).ElDoradoCo.:1miEofGeorgetown,mixedevergreenforestwithshrubunder-
story,serpentinesoil,20Aug1959,B.Crampton.5445(AHUC26540).GlennCo.:MendocinoNatl.
1450 BRIT.ORG/SIDA21(3)
For,CampHllcnJalcarea,ca,3,5miFAlderSpringsRangerStaiion,elev,ea,iOOOIt,11Jun10h3,B.
Cyaiuplon(AiluWlUCI'-mi).HumboldtCo.:Hureka,alt.0-200It,23Jun1900,/./'Trtuy1175(DAO
a6l4t9.9290)0;0HuItr,e3k0a,Maalyt.109-2p5,.0/0,F.It.Ir2tu3vju7n51ISQO{IR.S/A,F1T0r1a0c6)y;1T1275S(RR5FS:AS-ePc.OM12,2W061o8l0)A;ldWeirlSpKrwinCg:sreaelkoCnagn2ySo0n7,,
growingonserpenluie,JellreyPinewoodland.16|un1976,T.J\ldson&j.Nchon2740CRS.A273307).
LakeCo.:nearC.learLake,westernshoreollake,mK/iirs-Funis-Quenirsassoc,.27Api-1927.A.A.
DBeoeu(gllcais7-lii6r(-AmIapilUeC,5s9o4m8e;OchAaVpa4r0r4a9l),;113/j4umni1S95W5h,iBs.pCerraimnpglPomncs2,8SMlat(eAl1IlwUyC.2291.0r8o0a)d.cMuta,riynellCoo.w:pnneiaer-
6)lema,Atigl898,./,ii.Dnvy,s.(i.(RSAd'OM205834);Laguniias,7May1916,1,S.Smithsm.(A1ICC26^)4);
Mt,TaiTialpais,Bootjack,onserpentu-ie..19Jun.1938,/.7:Howell;,!92l)(RSA221132);PointReyes,scat-
teredclumpsindenselyvegetatedFlatsonNsidebehinddunes,12Jun1945,AA.Beetle3998(AHUC
10940);AngelIsland,borderbetweengrasslandandbrush.16Mayb)46.j.7://owe/I2(cS88(DAO65002).
Mendocinoc:o.:Ckiah,24May1869.U'.CHlasdalc&j.liDcivy.500(RSA-POM206181);Sherwtxid,
openwoodsondryslope,15Jun1915,AS.Hitchcocks.n.,AmerCrNat.Herb.no.487,(OAO65000);
T20NRlOWSec.33,MendocinoNatl.For,alongforestserviceroad1N02justnortholMonkeyRock
mi.xedevergreenlorest.elev58001).14Jul1977,/.ICs»n(tli,./,(CSmwc'c'vIWNe/.son9412IRSA275370),
MontereyCai.:eoasl:dhillsbetweenDelMonteandSalinas,2-3mionrd.toJohannesburgoilrd.
betweenDelMonteandSalinas,chaparralbelt,5Jun1946,A.A.Beetle428.](AHUC12343);Sargent
CCoy.p:reMosnstiFocreelslto,,ANladperaCRroeaedk,,4seMrapeynt1i6n2e7,.2M00W0icfst,e7ndMaan^yi^Ic9r607,0C0.B(.DHAaVni4h0c59i)m;5466mi6(FRSofAR1u8t0h2e1r3f)o.rNd,ap2a5
SApWro1f95K0n,o/..wS(ilreleef&s.uca[.A]1/I2CCmi13W901of);DKneov.iwlihlclaedQRuoaad,d,aTtljIuNncRti5oWnoNfVtVhle/cdrSeeeck.a25n,dCaeddairrtCrroeaedk,,g3r.a5smsiy
meadowandstrcamside,alt.f800ft,6May1986,(.,l.aPrcs.n.(RSA371706).San1-ranci.scoCo.:Daly
City,ColmaCanyon,Apr|6|p,I'JiKennedys.n.(.AHUC2691);Mi.Davidson,summit,largeclumps
amongbrnsli.22Apr1956,P.H.Raven900-f(D.AV56413).SanLuisObispoCo.:SternerCreek("Ser-
ranoCanyon"),onbrushyslopesmareaofserpentine,14May1947,R.t.Hoover7/68(DAD636020);
SantaLuciaMts..betweenRockyButteandPineMr,21Jun1950,R./:Hoover7999(DAO635967).San
M3afilar(.si5tAc6rHo4i5UdC(goC_e.A;\2W5Cl8.Ur6o)l2C')sS3.iaa2lSn2i9SsA3p)knr.iidynrSgoeasuan,tsCa1o1L.Ca:MrkanueyezaornC1o9H.0r:2oa,apIdLpmeliyoRPNCoiaylWmaArpohc,fritaDFonassiveseuL2dan-okpiteobs,rlutey.l(^eaRTvISw6OAo5S-o0PRdfOs3t,WMo1,5n871Ms56al9Moy5pa)e;1y9ao51b0n9,o3ev5Ra,e.sNHit.ieh/i\rec.niKCbfalalrhaaiimnepkaistooehnfl
Rixer31Ma\'1942.AA.Beetle&C,.L.Stcbhinylr.H4/(.AllUC7646);T40NR9\V.Sec.21,2.0miSof
SugarCreekonI'.urotiMillroad,roadsideinmixedconiferforestonmeiamor]ihicrock,4400ft,4
Jull972,/,/CS/ni()i./.().Sa\v\r/c'7M,0'/Vfe(;r((.5735(RSA273423).SolanoCo.:VacaMts.iacuigSacra-
memoValley,GatesCanyon,slopeswoodedwithIneoak,bayandchaparral,12May1958,RCrampfon
4784(AllUC24872).SonomaCo.:DuncanMillsClnadrangle,T6NRllW,nearBodegaPort,elev.25,
21Apr1634.H.S.Yale'sv/i (RSA123473.123478).andH.S.Yates3806(AHUC25861);NHslopesof
FilchMt.near1lealdsburg,lOJun16^2,l>Ruhtzolj/236(DAO291944).TehamaCo.:MendocinoNatl.
For.PaskentaDisr,CraneMillsloggingroad.Whiskey.Saddle,underconifers,elev4720ft,16jtm
N165s4i,de/IoClr(nTirii/n'(iit)yPiR/iv%e5r(aAnldlfW:Cs2i0d6e61ofJ.NTreiwnitRyivCoe.r:rNoeckwoRuitvcerrotprpaiilnIgrsoimnGCraalyifsoIr'naliaIsmCiaxmepdgrevoeurngdreoenn
withDouglasfiranddiggerline,elev1000b,7Apr1973.j./'Smif/i.5962(RSA244973);T31NRlOW
SSemci.th1.3,jCilouSnatwyyeFrin&eTRWo.adNcahloonng9B8r7o9w(n'RsSACr2e9e6k7,65G)a;rrByonoaankzawoKoitlnlganQdu,aderlaenvg2le2,00T3ft9,N7MRa7yW1S9e7c8,,3j4.,R
Shasta-'IrinityNatl,For,BearCreekatitsjunctionwithTrinityRi\ermi.xcdconilcrforest,elev3000
fi1tn,/41D5ooJluugNnlaW1s9d79i1/,r4/.t,oPMrSacsmctDiotohnn&aslloTdpWeS.,tamNtoeelsstFolonrye1ss0iu1n0N3noy(fRsCSpooArt\,2'a91l35l1iJ8su2,n)t.r1aO6irl6e4tg,ooBnH.:uLtBtWeerinfth1oo\'nnMCe&o.a:d¥.oT(lwAsIiS,mReadc5ghWeoSo7ef0c1.o4p1e(8nDNiAnWOg
786419).JosephineCo.:GaliceC"reekroad3 miItornjunctionwithRogueRiverroadalGalice,steep
DARBYSHIRE,NEWCOMBINATIONSINFESTUCACALIFORNICA 1461
serpentinehillsideonNsideofcreek,commoninpartialshadeatedgeoloak-pinewoodsb\-the
slope,elev.1250ft.10N'lay1Q74,K.L.Cbumhers,^^)}5lDA0540761).
Festucacalifornicasubsp.hitchcockiana(E.B.Alexeev)S.J.Darbyshire,comb,
etStat.nov.BASIONYM:Fc^tucahitchcotkianaE.B.Alexeev,Byull.Moskovsk.Obshch.Isp.
Prn-.,Otd.Biol,n.s.,87(2):111.1Q82,TYPE:U.S.A.CALIFORNIA,SantaClaraCo.:6N4ay1921,A.H.
WoUcy-Dod207(llOl.OTYPli:K).
Culms60-120cmtall;internodesglabrousorpubescentforupto5mmbelow
thenodes.Sheathsglabrous,scabrousorsometimesretrorsepubescent;collars
denselytosparselypubescentorsometimesglabrous;ligules(l-)1.5-6mmlong,
withorwithoutapicalcilia;blades0.8-2mmwide(3-5mmwidewhenllat),
convolute,involute,conduplicateor1lat,with7-15(-17)deepadaxialribs(usu-
allymorethanhalfasdeepasblade thickness);sclerenchymaasinsubsp.
californica(Fig.Ic).Inflorescences15-25cmlong.Spikelets8-12(-17)mmlong,
with (4-)5-6(-8)florets.Lemmas7-8(-8.5) mmlong,entireor minutely
bidentate.Chromosomenumberunknown.
Thissubspeciesisdistinguishedbyitslongerliguleswhichareusuallynot
ciliate,butoftenlacerate.Ligulesofleavesofthevegetativeshootsareusually
(1-)1.5~2mmlongwhilethoseoltheculmleavesmaybeevenlonger.Thecol-
larsmaybeglabrousorpubescent.Sclerenchymainthelealbladesontheveg-
etativeshootsiswelldevelopedwithgirderspresentatthemajorveins(Fig.Ic).
Thespikeletsandlemmastendtobesomewhatsmallerinthistaxonthanthe
othersubspecies.Althoughtherangeoverlapswiththatolsubsp.californica,
itoccursonlyincentralCalifornia(SantaClaraandSanLuisObispocounties)
nearthesouthernpartoftherangeol thetypicalsubspecies.Itislesscom-
monlycollectedthantheothertwosubspeciesandappearstohaveamorere-
stricteddistribution.Theformoftheliguleandsomewhatsmallerspikelets
andfloralbractssoclearlydistinguishthesepopulationsthatsubspecificrank
seemswarranted.
Specimensseen:U.S.A.CALIFORNIA:SanLuisObispoCo.:SeeCanyon.3May1948.R.KHoover7.512
(DAO635974);SeeCanyon,14May1904,R.F.hooverSS64(DAt3635612,635951:RSA201564);See
Canyon,lowerpartofPerlumoCanyon,claysoilIVomserpentine,14May1964,R.Z";Hoover8878(DAO
635956).
Festucacalifornicasubsp.parishii(Piper)S.J.Darbyshire,comb,novBasionym:
Fesluaiaristulatasubsp,pari.sliiiPiper,C^.ontrU.S.Natl.Herb.10(U:33,1906,¥esturaparishii
(Piper)Ilitchc.inJcpson,Fl,Calif.1:169,1912.rcslucaecilifornicavnr.parishii(Piper)Hitche.
inAbrams.Ill,fl.PacificStatesP222.1923,TYPli:U,S,A.CALIFORNIA:SanBernardinoMoun-
tains,MillCreekFalls,alt.5500ft,20Jun1901,S.B.Parish5036(lectotypI:,designatedby
Hitchcockinjepson1912:169:US-556210!).
Culms30-80(-100)cmtall;mternodesusuallydenselypubescentupto20mm
belowthenodes.Sheathsusuallydenselyretrorsepubescent,rarelyglabrousor
sparselypubescent;collarsusuallydenselypubescent,atleastatmargins(rarely
glabrous);ligules(0.2-)0.5-l,5(-2)mm,aciliatemembrane;blades0.5-L2(-1.5)
1462 BRIT.ORG/SIDA21(3)
mmwide,condupUcate(orlooselylolded),with(3-)5-9shallowadaxialribs
(uptoabouthall asdeepasbladethickness);sclerenchymainsmallabaxial
strandstoacontinuousband,withorwithoutadaxialstrandsatthemajorveins,
rarelylormingpillars(girdersabsent)(Fig.Id-l).InllorescencesKV20cmlong.
Spikelels11-16mmlong,with3-4florets.Lemmas(8)9-11.5mmlong,usually
minutelybidentate,sometimesentire.Chromosomenumberunknown.
ThissubspeciesoccursindrychaparraloropenforestsotsouthernCali-
forniaintheSanGabriel,SanBernardinoandPalomarMountains(Fig.2).Al-
thoughthelemmastendtobesomewhatlargerthantheothertwosubspecies,
overallplantstendtobesmallerLealbladestendtobenarrowerandshorter
(10-30cmlong,versususuallymorethan30cmlong)andthesclerenchymais
lessdevelopedwiththeabaxialbandoltcndiscontinuousorreducedtosmall
fasciclesoppositetheveins,pillarsonlysometimespresentandgirdersabsent
(Fig.Id-f).Leafsheathsofthevegetativeshootsareusuallydenselyretrorse
pubescent,butaresometimessparselypubescenttoglabrous.
Selectedspecimens:U.S.A.CALII-ORNIA:LosAngelesCo.:SanDimasF.xpt.For.,SanGabiiclMts.,
WolfskillnrebrcaknearBrown'sFlat,28May1042,KHH(624)63l(RSA-POM308'-)44).SanBernar-
dinoCo.:SanBcrnardmoK'lts.,7Oaks,JulK)01,A.Wividson224.5(RSA4l'5'-)66>,SanAnionioMts.,
CucamongaCanonatCdiarcoalCamp,abundantunderoaksonanorthslope,uppei'SonoranZone,
alt.4700ft.7Jun1919,i.M.Johiison2166(RSA-POM1271,6414,6415J;SanBernardinoMts.,nearGlen
Martin,neartailsonroadbelow,1Jun1941,j.Roos92(RSA-POM261420),SanBernardinoMts,,
WatermanCanyon,shadyhillside.21Apr194],/,Roos26.].](RSA-PC^M302531):SanBernardinoMts,,
RFroeodsa&lbaL,Roopoesn50p6la1c(esRSiAnf7o8r2e2s7t).wiStahnPDiiiteiigsopCoou.d:crPiaKlao,mOauroRianiig.se,kcUAog\;u^aiuTeitbci,aalWti,l5d8e0r0nefts,s9AJriema,1w95e1s.t-/,
erncrestofAguaTibiaMt.atheadofasteepdrawinthePechangaCreekwatershed,justN\Vofthe
largeQucirusa^^^yiloUawoodlandaboutthejunctionolthePalomai^Divideandnri|ipingSprings
trails,T9SRIWSW1/4NWl/4Sec.4,near3.3''25'10"N116°5938"W,elev.ca,4500ft,extremelysteep
sk)pcswithwoodlandolPieiuiutsugtfmiicnHdipuandQuercuschrysolepisandunderstoryofRil)cs
amaruin,Poly^itichum.Monanicllamailanlbnssp,/u/lhi,Carextriquetva,locallycommononmore
mesicex|")0surcswithsomeafternoonsun,25Apr1991,5,RoydH508(\')AO7'-)6415;RSA):NWPalomar
WMts.o,fCCluetvcealaVnaldleNya,tla,loFnorg.,PAagluomaaTri-bMiaagWcielTdreariln,eTss9SArRelaW,ENfEac1e/o4fSEWalg,l/e4CSreacg.,1S4,oefleuv,pp4e6r00Arftr,obyiogSceocnoe,
sprucewoodlandolPseudotsugamacwccirpa.Quenlischrysoicpis,LinunthusIloiibu}]dusJ\ctkidhi
7t0c',fVn>,at1a0.MPahaycc1l9i95a,uDn.hLi'Bmaintak.se&tc,S,.gBroayndit0i4c2s9ub(sDtrAaOte79w6i4t1h4;deRcSoAm)j:ioNsWedPgarlanoimtaersMotisl,.,aCspleecvtelNaFn,dsNlaot|li.e
For,AguaTibiaWildernessArea,NpeakofAguaTibiaMt,onNFcornerofpeak,justSoftheRiver-
sRiIdeWCoS,Fb1o/u4ndNarWy,1i/n4aSe\c'.cry4,stre3e°p25b'olw8l"-Ns1h1a6p"e5d7'd2e4p"rWe,ssbiiogncoonnethseprNucfelawnkooodflAagnudaofTiPbsicaiiMdto„lsuT^'-^)uS
macwcai'pa,Quercuschrysolepis,Rii/msIcucodermis,RibcsamarumPolystiehumiml'ru(j/i,sss|:).
c/)u./r..tuBma,nkestc.&,gSr.aBniotyidc0s5u0b9st(raDtAeOan7d96s4a17n:dyRShAu)m:usN-WricPhalsooiml,arasMptesc.t,CNJWe,vcsllaonpdeN1a7tl0,%F[osri.c,],A1gJuuanT1i9b9i5a,
WildernessArea,NEfaceofEagleCrag,SEoftheCrosleySaddle,SofCutcaTrailalongdrainagethat
parallelstrail,EofupperArroyoSeco,T9SRIWSE1/4Sec.14,elev4520ft,bigconesprucewood-
landofPseudotsuf;,amacrocarpa.Quercuschrysolepis,Q.berberidifolia,Arctostapbylosi^landulosa,
Carextriqueira,etc.,graniticsubstratewithdecomposedgranitehutnus-richsoil,aspectS,slojie
50%,15Jun1995,D,L.Banks&S.Boyd0684(DAO79(-)416:RSA).
DARBYSHIRE,NEWCOMBINATIONSINFESTUCACALIFORNICA 1463
ACKNOWLHDGMENTS
ThecuratorsoftheherbariaAHUCDAO,DAV,RSAandUSarethankedfor
kindlyprovidingspecimens,includingtypes,forstudyjochenMiilleristhanked
forhismanyhelpfulsuggestions.JacquesCayouette,WJ.Cody,andananony-
mousreviewercommentedonearlierdrafts.
REFERENCES
AiKFN,5.G.,M.J.Dailvvitz,C.L.M(Jannft,andL.L.Consaul.1997.BiodiversityamongFestuca
Poaceae)inNorthAmerica:diagnosticevidencefromDELTAandclusteringprograms,
andanINTKEYpacl<ageforinteractive,illustratedidentificationandinformationre-
trieval.Canad.J.Bot.75:1527-1555.
ArbXEEV,E.B. 1982.Newand little-knownfescues{FestucaL.)ofNorthAmerica.Byull.
Moskovsk.Obshch.lsp.Prir.,Otd.Biol.,n.s.,87(2):l09-118.[InRussian].
SibBBiNs,G.L.,jR.andR.M.Love.1941.AcytologicalstudyofCaliforniaforagegrasses.Amer.
J.Bot.28:371-382.