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CALAMAGROSTIS COAHUILENSIS AND C. DIVARICATA (POACEAE: POOIDEAE: AGROSTIDINAE), TWO NEW SPECIES FROM MÉXICO PDF

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CALAMAGR0STI5COAHUILENSISANDC.DIVARICATA (POACEAE:POOIDEAE:AGROSTIDINAE), TWONEWSPECIESEROMMEXICO PaulM.PetersonandRobertJ.Soreng JesusValdes-Reyna DepartmentofBotany DepartamentodeBotanica NationalMuseumofNaturalHistory UniverisidadAutonomaAgraria"AntonioNarro" SmitfisonianInstitution Buenavista,Saltillo Washington,DistrictofColumbia20013-7012,U.S.A. Coatiuila25315,MEXICO [email protected];[email protected] [email protected] ABSTRACT CalamagrostiscoahuUensisP.M.Peterson.Soreng&Valdes-Reyna.sp.nov.,andCalamagrostisdharicata oP,cMc.urPsetoenrsconal&caSroeroeunsg,slsop,pensov.m,ftrhoemmMoeuxnitcaoianrsedofesscoruitbleideaasntderinllCusotarhauteidl,aCaanldamsaogurotsftiiwsecsotaelrinuNiIuccns\ios Leon;andCdivaricataoccursonmoistslopesandmossycliffsofsouthwesternDurango. CalamagrosliscoahuilcnsisismorphofogicallysimilartoCpurpurasccnsbutdilIcrsbyhavingshorter anthers(1-1.2mmlong),nearlysmoothligules,scaberulousrachises,glabrousorminutelyantrorsely scabroussheathsummits,andsomewhatdensebutnotspikelike,greenishpanicles,Calamagrostis divaricataseemsmorphologicallyalliedtoCpringleibutdiffersbyhavingwiderpanicles(4-10 cm)withbranchesreilexedspreadinganddivaricate,shorterspikelets(.3.4-4.3mmlong),shorter glumes(3.4-4,3mmlong)thatarebothIveined,andshorterlemmas(3,8-4.3mmlong)withan awninsertedonupper1/4to1/3. RESUMEN SedescribenyseilustranparaMexicoCalamagrostiscoahuUensisRM,Peterson,Soreng&Valdes- Reyna,sp,nov,,yCalamagrostisdivaricataRM,Peterson&Soreng,sp,nov,Calamagrostiscoahuihnsis seencuentraenladerascalcareasenlasmontafiasdelsurestedelestadodeCoahuilaysuroestedel estadoNuevoLeon:yCdivaricataseencuentraenladerashumedasyriscosmusgososdelsuroeste delestadodeDurango,CalamagrostiscoahuUensisesmorlologicamentesiinilaraC,purpurasccns perodifiereenposeeranterasmascortas(1-1.2mmlargo),ligulascasilisas,raquisescaberuloso,apices delasvainasglabrosoescasamenteconescabrosidadantrosa,ypaniculasdensasperonoespigadas, decolorverdusco.CalamagrostisdivaricataparecemorfologicamenterelacionadoaC.pringleipero ddiifviaerriecaednas,poessepeirguiplalnaiscmulaasscomratsasa(m3p,4l-i4a,s3m(4m-1l0arcgom)),gclounmarsammiafsiccaocritoanse(s3,r4e-l4l,e3xams,melxatregon)diadmabsasy uninervadas,ylemasmascortas(3.8-4.3mmlargo)conunaaristainsertadadesde1/4hasta1/3 arribadelamitad. Calamagrostisischaracterizedashavingsingle-!loweredspikelets,oneorthree- veinedglumesaslongorexceedingthefloretinlength(rarelyslightlyshorter), non-keeledlemmasthataremembranousorcartilaginous(infrequentlyhya- line),usuallywithasingledorsallyattachedawn(rarelyawnless),acalluswith acrownofhairs,caryopseswithshorthilumsandwithoutapicalhairs,and lodiculesthatareapicallymembranous(Clayton&Renvoize1986;Watson& SIDA21(1):311-320.2004 312 BRIT.ORG/SIDA21(1) Dallwitz1992).Worldwide,Calamagrostisincludesbetween 230(Watson&r Dallwitz1992)and263(reportedbyD.Clayton&rB.Simon,pevs.comm.2004) species.InNorth,Central,andSouthAmerica131speciesarereportedinSoreng etal.(2003).InMexicothelollowing11speciesolCdlamd^i^wstisarerecognized: C.crcctijoUaHitchc,C.cnantha(Kunth)Steud..C.guatcmalcnsisllitchc,C. intenncdia(J.Presl)Steud.,C.ondzcihcic(Rupr.e.\E.Fourn.)Beal,C.piinglci Scribn.exBeal,C.purpurascensR.Br.,C.rigcsccns(j.Presl)Scribn.,C.tolucensis (Kunth)Trin.exSteud.,C.validaSohnslsyn.=C.ivcvauglici Sohns;McVaugh (1983)wasthelirsttoselectthisnameasasynonym,seeArticle11.5forequally publishedspeciesintheCodeolBotanicalNomenclature(Greuteretal.2000)1, andC.vukaniLciSwallen(Chimal1987;Hspcjo-Sernaetal.2000). Whilecollectinggrassesandspecilicallysearchingloruniquelormsol TnsctuminDurango,Mexico(Finotetal.2004)inthefallof200.3thefirstau^ thorgatheredaspecunenthatseemedtobeanundescribedspecies.Atlirst glancethisspecimenappearedtobetwoorrriorcIloweredbutuponcloserin- spectionandundermagnilicationallspikeletscontaniedasingleIloret.Alter studyingmorphologicallysimilarmaterialfromMexicocollectedbythelirst author,anadditionalundescribedspecieswasloundmCoahuila.Thetwonew speciesareclearlymembersol subfamilyPooideae,tribePoeae,subtribe Agrostidinae(Sorengetal.2003,2004).Wedescribethesetwospecimensas newspeciesofCahimagm:<tis. CalamagrostiscoahuilensisRM.Peterson,Soreng&Valdes-Reyna,sp.nov.(Figs. 1,2).T\rr-MHXICO.Cc:)AtlUlLA.SierraMadreOriental,32miSEofSaltilloand8 miSI:ofJamconroadroSierraLaViga,3240m.29Sc]:)1990,RM.Pctersoi].C.R. Aundblc&j.Vcildcs-Rcyua10051b{l\o\xy\'\PF:.US'j. ACalamii\^i'osliimipunisci'niiR3r.antlicrls1-1.2mmlongis.ligulisIcrclai-\'i,i;aiis,rliucliibus scaberulis,\'aginaapicalighibra\'clscabi'cllaaninirsc.panicuhsaHquaniuindensisscclnonspicae similibiLS,viridis.reccdit, Caespitciseperennialwithintravaginalandextravaginalshootinitiation.Culms (22-)40-100cmrall,erect,glabrous;internodesglabrous.Sheaths6-28cmlong, shorterthanthemternodes,glabrous,oltenlibrousbelowwithage;margins smooth;cataphyllsandK^wersheathsglabrous;summitglabrousorminutely antrorselyscabrous.Ligules5-8mmlong,membranousoltenlacerate,Hrmer below,nearlysmooth,apexobtusetoacute.Blades8-20cmlong,2-4mmwide. Hat.apicallyacuminate,glabrous,scabrousabo\'e.Panicles(7-)9-20cmlong, 0.5-1.2cmwide,contracted,somewhatdensebutnotspike-like,greenish;ra- chisscaberulous; inlK^'escence branchesmostly 2-10cm longbelow,the tbrwaoncpheers,noadsec.eSnpdiiknegleatnsd5t.i8g-h7t.h2'mapmprelosnsge,d,1-mtolsotwlesr'eIdl,ortiiglhetrloyusanpeparrebsasseed,otnoethoer branches,darkgreenish;pedicels0.6-4 mm long,ascending,scaberulous; rachilla2.4-3mmlong,coveredwithstillhairs,thehairs0.5-1mmlong.Glumes 5.8-7.2mmlong,lanceolate,aboutccjualorsubequalinlength,longerthanthe PETERSONETAL.JWONEWSPECIESOFCALAMAGROSTISFROMMEXICO 313 hQ.l.CalamagrostiscoahuHensis{Peterson,Annable&Valdes-ReynaJ005/6).A.Habit.B.Inflorescence.C.Sheath,ligule, andblade.D.Spikelet.E.Lowerglume,dorsalview.F.Upperglume,dorsalview.G.Floret,lateralview.H.Lemma,lateral view.I.Lemma,dorsalview.J.Paleawithrachilla,dorsalview.K.Stamens.I.Pistil.M.Lodicules.N.Caryopsis,dorsal view. 314 BR1T.ORG/SIDA21(1) -C.divaricata 200miles A-C.coahuilensis k V1 /,Zacatecas''I\SaPontoLsuiis"\^^^T.a,maulipas( Fic.2.DistributionofCalamagrostiscoahuilensisandC.divaricatainMexico. floret,membranous,1-vemed,scaberulousalongveni;marginshyaline;apex acute.Lemmas5-5.7mmlong,lanceolate,membranous,yellowish-green,con- spicuously5-veinednearbase,scaberulous,awned;apexacute,entireorbifid; awn8~10mmlong,goldenbelowandpurplishabove,bornenearthebaseon thelower1/6,insertedinagroovebelowandtwistedandonce-geniculateabove; callusobtusewithwhitehairs,thehairs0.8-1.8mmlong.Paleas5-5.6mmlong, aboutaslongasthelemma,hyaline,2-veined,theveinsscaberulous,green;apex acute,entire.Lodiculesca.1mmlong,hmceolate,lobed,glabrous.Stamens3;an- sthteirgsma1-s1.2,2fmeamthelroyn,g,whpiutripslhe..COavrayroypscae.s02.-22m.5mmlmongl,onggl,abfursoiufso;rsmt,ylleisgh2t,gsreepeanriasthe.; Phenology.—FloweringinlateAugust,September,andearlyOctober. Distribution—Ccilamagmsliscoahuilensisisknown(romthemountains ofCoahuilaandNuevoLeongrowingoncalcareousslopeswithAbiesmexicana Martinez, Pinusculmirncola Andresen &Beaman, P. hartwegii LindL, Hesperostipaspartea(Trin.)Barkworth,PoarupreehtiiPeyr,Holodiscusdiscolor (Pursh)Maxim.,Quercusgreggii(A.DC.)Trel.,andscatteredPopulustremuloides Michx. Additionalspecimensexamined(par/Viypes):MEXICO.Coahuila:municipiodeArteaga,nearsummit ofCoahuiion.S\'-of.SaiiAnioniodel.asAlazanasandSFofSaltillo.3f20m,17Oct1989.P.M.Pctcr- siui.J.W(/dcs-Rcv/Kifv/.A.V^i/lcin-((/8.5'J9{ANSM,US):29Aug1986,/.A.Mel)()iui/tilTi;XJ.NiievoLeon: municipiodeGaleana,CerroelI'otosi.3600m,3Aug1989,A.Garcia-Arevalo66(ANSM,CllDlRj; cumbredeCerroPorosi.ca.Galeana.29Aug1986,/A.M<:Donald(TEX);LaderaS,26Jul1985,J.A. McDonald(THXJ;/.Ochoa-CuiUcnunil9S.iICOCA). PETERSONETAL.,TWONEWSPECIESOFCALAMAGR0STI5FROMMEXICO 315 Calamagrostis divaricataP.M.Peterson&Soreng,sp.nov.(Figs. 2, 3).Type: MEXICO.DURANGO.SierraMadreOccidental,ca.3miSofHwy40atMexiquillo, ArroyoPasoResbaloso(23°42'46.2"N-105°39'45.1"W),2520m,14Sep2003,P.M. Peterson,M.S.Gonzdkz-Elizondo&G.Tena-Gonzdlez17774CiiOLOTYri-:US!; ISOTYPES:ANSM!,CHDIR!,MO!,RSA!,US!). sApiCcaLihluivsa3v.,4m-s4t.3impymingllocnigiSsc.rgilbun,mies,\3i.i4e-al4.[3lmanmiculloinsgi-.'si,—\u0ncivmcnliast.is,Icrmammaitsirbelulsex3i.s8,-e4l.i3usmismellodnug'iasr.icaartiisst.a iiLsertasu|iera1/4-1/3,recedil. Looselycaespitoseperennialwithshortrhizomesandextravaginalshootini- tiation.Cuhiis52-96cmtall,erecttoslightlydecumbentnearbase,glabrous; internodesglabrous.Sheaths8-20cmlong,shorterorlongerthantheintern- odes,glabrous;marginssmooth;cataphyllsandlowersheathsglabrous;sum- mitglabrous.Ligules2-3.6m.mlong,membranousoltenlacerate,decurrent below;apextruncate,ciliolate.Blades15-60cmlong,1-3mmwide,flatorinvo- lute,apicallyacumitiate,glabrous,scabrousabove.Panicles9-16cmlong,4-10 cmwide,ovate,open,lax,sparselyflowered,greenish;rachisscaberulous;in- florescencebranchesmostly2-8cmlongbelow,thebranchesdelicate,ascend- ingandspreading,nakednearbase,whorled;lowerinflorescencenodeswith 3-6 branches.Spikelets3.4-4.3mm long,1-flowered,spreading from the branches,yellowish-green;pedicels1-5mmlong,ascendingtoreflcxedand spreading,scaberulous;rachilla1.6-1.9mmlong,coveredwithstiffhairs,the hairsupto1.5mmlong.Glumes3.4-4.3mmlong,lanceolate,aboutequalin length,shorterorlongerthanthelloret,membranous,l-vemed,scaberulous alongvein;marginshyaline;apexacutetoacuminate.Lemmas3.8-4.3mmlong, lanceolate,membranous,yellowish-green,5-veined,scaberulous,awned;apex acute,entire;awn4-6mmlong,yellowish,borneontheupper1/3or1/4,straight ortwistedandonce-geniculate;callusobtusewithshortwhitehairs,thehairs 0.2-1 mm long.Paleas3.8-4.4mm long,aslongorslightlylongerthanthe lemma,hyaline,2-veined,theveinsscaberulous,green;apexacute,entire.Lodi- cules0.7-0.8mmlong,lanceolate,lobed,glabrous.Stamens3;anthers1.8-2.2 mmlong,yellow.Ovary0.2-0.4mmlong,glabrous;styles2,separate;stigmas 2,feathery,whitish.Caryopses1.8-2.5tnmlong,fusiform,lightbrownish. PhcMoIog3/.—FloweringinSeptetriber Distribution.—Calamagrostisdivaricataisknownonlyfromthetypelo- calitygrowingonmoistslopesandmossyclillswithPinuscooperiC.E.Blanco, jRdurangensisMartinez,PfljyacahuiteC.Ehrenb.exSchltdl.,Quercussideroxyla BonpL,ArbutusoccidenlalisMcVaugh&Rosatti,CupressusarizonicaGreene, andMuhJenbcriiiaalaiuosacVasey. DISCUSSION Calamaii^rostiscoahuilcnsisremainedundetectedbybiologistsloralmost19 years.ThefirstcollectionsofthisspecieswereapparentlymadebyMcDonald 316 BRIT.ORG/SiDA21(1) 0.5mm h<c3Xalamagrostisdivaricata{Peterson,Gonzdlez-Elizondo&Tena-Gonzdlez17774).LHabltZAnMiscence.C.Sh&ath, ligule,andblade.D.Spikelet.E,Lowerglume,dorsalview.F.UpperGlume,dorsalview.G.Lemma.dorsalview.H.Lemma, ventralview.I.Paleawithrachilla,dorsalview.J.Palea,dorsalview.K.Stamensandpistil.L.Gynoecium,mature.M. Lodicules. PETERSONETAL.JWONEWSPECIESOFCALAMAGROSTISFROMMEXICO 317 in1985fromtheCumbredeCerroPotosi.Allspecimensthatwehaveseenfrom CoahuilaandNuevoLeonthatwerepreviouslydeterminedasC.purpurascens orC.pringleiareC.coahidlensis.Therefore,C.purpurascensisnotfoundin Mexico. Calamagrostis coahuilcnsis is morphologically similar to C. purpurascens,butcanbeseparatedfromthelatterbyhavinganthersthatare shorter(f-1.2mmlong),ligulesthatarenearlysmooth,ascaberulousrachis, sheathsummitsthatareglabrousorminutelyantrorselyscabrous,andgreen- ishpaniclesthataresomewhatdensebutnotspikelike(TableI).Populationsof C.coahuiknsisareseparatediromtheclosestknownlocalityofC.purpurasccns intheSangredeCristoMountains,TaosCounty,NewMexico(Allred1993)by over800mi. McDonald(1990)andGarcia-ArevaloandGonzalez-Elizondo (1991)firstreportedC.purpurascensfromCoahuilaandNuevoLeon,Mexico. ApparentlyManuelGonzalez-Ledesma,JohnReeder,andCharlotteReederiden- tifiedthegrassesforMcDonald(1990)sincetheyappearintheacknowledg- mentsforhavingreviewedthatfamily.Thisisnotsurprisingsincemmostchar- acteristics the rangeof variation in C.coahuilcnsisoverlapsthatof C. purpurasccns.AnadequateillustrationiOchoa983)ofthenewspeciesappears inBeetle's(f987)treatmentofthegrassesofMexico,althoughhereferredtoit asC.pringlei. ItissurprisingthatC.divaricataisknownonlyfromasingle,recentcol- lection since it occurs near a rather heavily used tourist destination. CalamagrostisdivaricataseemsmorphologicallyalliedtoCpringleiandcan bedifferentiatedfromthelatterbyhavingwiderpanicles(4-10cm)with mm branchesreflexedspreadinganddivaricate,shorterspikelets(3.4-4.3 long), shorterglumes(3.4-4.3mmlong)thatareboth1-veined,andshorterlemmas (3.8-4.3mmlong)withanawninsertedonupper1/4to1/3(Table1.). TheinfragenericclassificationofCalamagrostishasnotbeencritically tested,anditisnotusuallydiscussedfortheNewWorld,otherthantheaccep- tanceofthesegregategenusDeyeuxuibysomeSouthAmericantaxonomists (RugolodeAgrasar 1978, 1998; Villavicencio1995).Thegenus(including Deycuxia)isdividedintoatleastloursubgenerawithmanysections,subsec- tions,andseries(Rozhevits&Shishkin1963;Tzvelcv1976;Wasiljew1960).All oftheseRussianagrostologistsplacedC.purpurascensinCalamagrostissect. Deyeuxia(ClarionexRBeauv)Rchb.,andWasiljew(whoseclassilicationwas world-wideinscope)placedC.pringleiinthesamesection.Severalinfrageneric taxawereacceptedfortheNewWorld(Sorengetal.2003).Basedoncallushair length(1/2orlessthelemmalength),lemmasmembranous,andpresenceola well-developedrachiUaextention,bothofthenewspeciesappeartobelongin Calamagrostissubgen.Deyeuxiasect.IJeyeuxia.Akeylorseparatingthetwo newspeciesfromotherspeciesolCahimagrosiisinnorthernMexico(Chihua- hua,Coahuila,Durango,NuevoLeon,Sinaloa,andTamaulipas)lollows. Table1.SalientfeaturescomparingCalamagrostiscoahuilensis,C.divaricata,Cpringlei,C.purpurascens,andCvalida. Characters C.coahuilensis C.divaricata C.pringle C.purpurascens C.valida Cataphylls&lowerleaf glabrous glabrous denselyhirtellous glabrousor glabrous sheathabaxialsurface scaberulous Leafsheathsummit glabrousorminutely glabrous glabrous glabrousorminutely Denselyhirtellous abaxialsurface antrorselyscabrous retrorse-Strigulose Leafblade&ligule glabrousornearly glabrous glabrous scabrous sparselyhirtellous adaxialsurface smooth Inflorescencewidth 0.5-1.2cm 4-lOcm 2-4cm 0.8-1,5cm (15-)4-10cm Inflorescence ascendingand ascending, ascending,loosely ascendingand ascending,appressed branches, closelyappressed; reflexedspreading appressedor closelyappressed, tobasespreading;± arrangement& ifloriferoustobase todivaricate,not spreading;not spike-like; floriferoustobase spikeletplacement floriferoustobase floriferoustobase +floriferoustobase Spikeletlength 5.8-7.2mm 3.4-4,3mm (4.5-)5-5,8mm 5-9mm 4-6.5mm Spikeletcolor greenish yellowish-green palegreentopurplish yellowish-purple yellowish-purple rraacchhiillllaalhaeinrgtlhength 02,.54--13rmTmim 01..86--11..59mmmm 11.-52-2mmmm 11.-82-3mmmm 21-.53-2mmmm Glumeslength 5.8-7.2mm 3.4-4.3mm (4.5-)5-5.8mm 5-9mm 3.4-6.5mm Upperglume, one one three one (one)three numberofveins Lemmalength 5-5.7mm 3.8-4.3mm 4.5-5mm 4-7mm 3.4-5mm Lemmasurface smooth scaberulous scaberulous scaberulous smooth Lemmaveins,aspect conspicuous inconspicuous inconspicuous inconspicuous inconsp./conspicuous Lemmatalawninsertion base,lower1/6 upper1/4to1/3 lower1/3tomiddle lower1/6to1/3 nearmiddle Lemmaapex entireorbifid entire entire bifid twoorfoursetaeor withage teeth(prolongation Callushairlength 0.8-1.8mm 0.2-1mm 0.2-1mm 0.7-2mm, o1f.3v-e2i.n8s()-3.5)mm oso occasionallyabsent o Antherlength 1-1.2mm 1.8-2,2mm 2.2-2.4mm 1.5-3.5mm 1.4-2.2mm Anthercolor purple yellow yellowish-purple purpleoryellow yelloworpurple := PETERSONETAL.JWONEWSPECIESOFCALAMAGROSTISFROMMEXICO 319 KEYTOTHESPECIESOFCALAMAGROSTISINNORTHERNMEXICO 1. oPranoinclelsow0e.r5-11/.65ctomnweiadre,lcoownetrra1c/4t;eda,ntdheenrssel1y-f1l.2owmermed;lolnegmmaawnCianlsaemrategdroatstbiassecoahuilensis 1. sPaenritceldesfro1.m5-l1o0wecrm1/3w,idmei,ddnlaer,raonwdtuopopperen1/a4;ndantlhoeorsesly1.f4-l2o.w4ermedm;lloenmgm.aawnin- 2. Lemmaapexwithtwoorfoursetaeorteeth(prolongationofveins);callu5hairs 1.3-2.8mmlong;rachillahairs2-3mmlong;leafblade,ligule,andsummitof sheathsparselyhirtellous Calamagrostisvalida 2. hLaeirmsma0.a8p-e2xmenmtireloanngd;wlietahfoubtlasdeet,aeliogrutlee,etha;ncdallsuusmhamiirst0.o2f-1shmemathlonggl;arbacrhoiulsl.a 3. Upperglume3-veined;spikelets(4.5)5-5.8mmlong;lemmas4.5-5mmlong, awninsertedonlower1/3tomiddle;panicles2-4cmwide,thebranches looselyappressedorspreadingbutnotdivaricate Calamagrostispringlei 3. Upperglume1-veined;spikelets3,4-4,3mmlong;lemmas3.8-4.3mmlong, awninsertedonupper1/3to1/4;panicles4-10cmwide,thebranchesre- flexedspreading,divaricate Calamagrostisdivaricata ACKNOWLEDGMENTS WewishCOthanktheSmithsonianInstitutions,NationalMuseumoi Natural History,BiodiversityandInventoryProgram,andResearchOpportunitiesFund forsupportingtheiieldwork.AppreciationisextendedtoAliceR.Tangerinifor providingtheillustrations;SusanJ.Penningtontorpreparingthedistribution map;AlainTouwaideandDanNicolsonforcorrectingtheLatindiagnoses;Kelly Allredand RichSpellenbergforcheckingthedistributionrecordsin New Mexico;andTeresaMejia-Saulesforfindinginconsistenciesinthemanuscript. REFERENCES Allred,K.W.1993.AfieldguidetothegrassesofNewMexico.DepartnnentofAgricultural Commur^ications,CollegeofAgricultureand HomeEcononnics,NewMexicoState University,LasCruces. 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