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THE BLUE-EYED-GRASSES (SISYRINCHIUM: IRIDACEAE) OF ARKANSAS PDF

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THE BLUE-EYED-GRASSES (SISYRINCHIUM: IRIDACEAE) OF ARKANSAS KATHLEENL. HORNBERGER DepartnimtofBiology, ScienceDivision Widener Unii'ersity Chester, PA 19013, U.S.A. ABSrRAC;T Dataobtainedfrommorphologicalcharactersandchromosomenumbermdicarethat fsoelvieunmsMpielcli.e,soSi.Saitslyannntcihamn/n//Bsihcokunl.d,b.Ve.rceacnoilgwntirzeedBifcokrnA.r,kaSn.salsa:n^SU.mail!biGcrleumenReaf.{S,.S.praunigmmstntm- Bickn.],S.nsulatnmBickn.\S.exileBickn.],andS.sagttuferiimBickn.Theseseventaxa aredifferentfromtheeightpreviouslyrecognizedbySmithinthat.V,uipttijerumBickn. Sw.asexinloetBliicsktned.,anwderSe.llainsgtleodisaisifGorueredniestainncdtSs.pperciiiesi.uouDiimstBriicbkunt.io,nplmuaspSs.wroesruelaptruomdBuiccekdn.usainndg herbariumvoucherspecimens. intr()i:>ik:tk)n The genus Sisyrinchiim consists of both herbaceous perennials and annuals with simple or branched stems that may or may not be con- spicuously winged. The flowers are epigynous, subtended by a spathe (made oftwo overlapping bracts), and have undifferentiated actinomor- phic perianths. Themucronulatc toanstulate tcpals range incolorfrom whiteoryellowwithpurplestripesandapurpleeye-ringtomoretypically blue, purple, orwhitewith ayelloweye-ring. No major revision had been done on this genus in the southeastern United States since Small (1933), which did not include the state of Arkansas. Smith(1978)recognizedeightspeciesforthestate:S. alhidum Raf., S. angustifoltumMill., S. atlanticiim L^ickn., .V, campestreBickn., S. exile Bickn., S. langloisu Greene, S. pruinosum Bickn., and S. rosulatuni Bickn. Therefore, aspartofataxonomicrevisiononthisgenusintheSE U.S. fortheSoutheasternFloraProject(Massey&Radford 1981), which includesArkansas,specialattentionwaspaidtothisstate'sspeciesinorder to update Smith'sAtlas(1978). ThisstudywaspartofadissertationcompletedattheUniversityofArkansasinFayetteviUein January 1987. SiDA 14(4);597-6()'1. 1991. 598 systi-maik:study A) Morphology Several hundred herbarium specimens ofBlue-eyed-grass forthestare wereexaminedfor24differentcharacters(Hornberger 1987a). Discrimi- nantanalysisindicatedthatofthese24characters, thefollowingprovided thebestseparationforArkansasspecies;pairedvs.singlespathes,lengthof outertoinnersparhebracts(equaltosubequalvs. unequal), connationof outerspathebract,stemwidth(includeswingoneithersideofraisedcen- ter portion), stem wing width (average valueofboth wings), and dried capsule color and shape. Flower color and shape arc also important characters, butdonotpreservewell, andare, therefore, notreadilyavail- able from herbariumspecimens. However, theyareincluded, alongwith thecharactersmentionedabove, inthetaxonomickeywhichfollowsthis section. Essentially therearetwobasicmorphologicalgroups, onewithsimple stems(.V. alhich/iuRaf.,.V.campestreBickn.,andS.sapttiferiimBickn.)and onewith branched stems{S. angustijoliuniMill., .V. atlanttcmnBickn.,S. IctnglmsiiGreene, andS. rosidatnwBickn.). Thespecieswithsimplestems arearrangedinoneoftwoways: 1)thestemshaveeithersinglespathesona scapethathasnocaulineleaf,hence, thereisnonodeor2)thestemshave pairedsessilespathes(sometimesasingleone)subtendedbyacaulineleaf. Thislattertypeofstemisconsideredtohaveonenode.Someofthesimple- stemmedspeciesmay, however, occasionallybranch, particularly5. sagit- t/JennN Bickn. (Hornberger 1987a). Species that typically have branched stemshaveoneormorenodes, eachofwhichhasacaulineleafandoneor more pedunculatespathes. B) Chromosome Number x=C8h,rowmitohsommoestreopfortthseinsptehceielsitbeeraitnugretientdriacpaltoeidtsha(tOltihveerge&nusLeiwsibsas1e9d6o2-n Oliver 1966; Goldblatt 1982; and others). Amongthesechromosomereportswerecountsofn=16fromflowerbud materialforonlytwospeciescollectedinArkansas,S.campestnBickn.and S. Liugloisii Greene (the latter taxon reported as S. pruinosnm, Oliver & Lewis 1962;Lewis&Oliver1961).1wasabletoconfirmthisnumberfori\ camptstnBickn. from twodifferentArkansaspopulations,oneinHotSpr- ing Co. and theother in Washington Co. (Hornberger 1987a). Ialsoobtainedthecountofn= 16fromflowerbudmaterialforS.langlo- isiiGreene,S. rosidatiimBickn., andS. sagittifernm Bickn., allcollected, however, in Louisiana(Hornberger 1987a, b). 599 Sisyrinchii/m alhidum Raf. was reported as n=l6 from Louisiana by Oliver&Lewis(1962). This numberwasoriginallyreportedbyBowden (1945)forapopulationcollectedinVirginia. Itwasfurthersupportedby Ingram(1964)forapopulationinTennessee. EventhoughIhavenothad theopportunitytocountthenumberforthisspecies,itappearstobeoneof the tetraploid blue-eyed-grasses. Sisyrinchiinn angmtifolium Mill, has been reported as n=48 from Louisiana by Oliver & Lewis (1962); they also reported this numberfor several populations inTexas. Ihaveseen thesespecimensand agreewith theiridentification. Hill(1984)recordedthissamenumberforaVirginia population. However, Ingram (1964, 1967) reported n=40, 44, 45 for populationsinNorthCarolina,Tennessee,andVirginia.Goldblatt(1982) feelsthattheseconflictingreportsforthesamespeciesareprobablymorea reflection ofmisidentification or incorrectcounts thancytological diver- sity. Sisyrinchuimatlanticum Bickn. has more reported diversity inchromo- somenumberthantheprecedingtaxon.Numbersrangefromn=8(Oliver 1966)ton=16(Ingram 1964;Oliver 1966;Hill 1984)ton=48(Oliver &Lewis 1962). IhaveseenthespecimenscollectedbyOliver(1966)andI agreewiththeidentifications. However, 1haveseen theherbariumspeci- men(Oliver253,ASTC)ofoneofthetwopopulationscollectedbyOliver &Lewis(1962)fromTexasthatwasidentifiedas5^.atlanticumBickn.;this specimenrepresentsS.biformeBickn.,aspeciesseeminglyrestrictedtothe GulfCoast andoffshore islands. C) Synonymy ThegenusSnyrtnchiumhasbeenmisunderstocxltaxonomicallyformore thanacentury. Becauseofsubtledifferencesinmorphology,disagreements among botanists on recognition of legitimate taxa, synonymy, or the properepithetforataxonhaveledtoaplethoraofspecies'descriptionsin theliterature.Nomenclaturalconsiderations,then,becameamajortaskof theSEU.S. revision. Holotypeswererequestedforalltaxa,andwhenthey were determined lost or nonexistent, neotypes and lectotypes were designated. TheonlytypespecimennotseenwastheoneforS. rosulatum Bickn. becauseitwasunavailableforthisstudy. Acompletediscussionof types and synonyms is included in Hornberger (1987a). Synonyms will only be listed in this paper ifthey are different from Smith (1978) and would causeconfusion if not included. KT.YTOTUF.SFr,<:.Ii:S IN ARKANSAS A. Outerspathcbratt norconnateatbase, oronlyslightlyso(1.0mmor less) fi 600 B. Spachespairedattopofstem;bractsofouterspatheslightlyunequal; sremwithasinglecaulincleafsubtendingthespathes .V.alhulumRaf. B. Spathe single at top of stem; bracts unequal with outer one I 1/2-")X lengthofinnerone;stemwithnocaulineleafsubtending A. OCute•^S'rip^'astsppha'eitth-behreabcrtascteqcuoanlnattoesautbebqausael(1.1 mmormore) V camOCpestrcBickn. DD..OOH.uuttSeetrressmppsaatt2hh.ee5bbmrraamccttoccroonnmnnoaartteeewaattidbbeaa,sseewifunoprgstmoo0r.2e9.0tmhmmamno2r..0m.om.r.emS.wi.sdu.ge.ii.St.liiferumBickn. angiiUifoltiiniMill. E. Stemslessthan 2.") mm wide, wingsless than0.9mmwide F F. Capsulespalebeigewithpurpleorbrownsutures,globoseto subglobose; tepalswhiteoryellowwithjxirplestripesand F. Cpuarpis:>ulleeesyeb-rrionwgn;ftolowbelrascku,rcgeloolbatoese.to..o.bSo.vartoeu;datteipaanlsBilcikgnh.t blue to purple, sometimes white, with yellow eye-ring; flowersrotate q G.Innerspathe bract distinctly mucronate; spathes often deflectedatbase;capsulesoblong-subglobosetoobovate G. Innerspathebractnotmucronate;spathesnotdeflSe.ctaeildciantticKDiBickn. base;capsulessubglobose S. langkisiiGreene C. Spathebractsnoticeablyunequal H H.Outerbractconnateatbaseupto2.0mm;outerbractupto3X lengthofinnerbract S.uigittijennnBickn. H.O1u/2te—r2brXactlecnogntnhatoefiatnnbearsebrmacotrethan2.0mm;outerbractonly 1 J. Stems2.0mmormorewide,wings0.9mmormorewide;tepalsJ lightbluetowhitewithyelloweye-ring;flowersrotate;capsules J. Sdatrekmsbrleoswsnthan2.0mmwide, wingslessthan0.9m9mcwni^dmtei;foluimMill. tepalswhiteoryellowwithpurplestripesandpurpleeye-ring; flowersurceolate;capsulespalebeigewithpurpleorbrownsu- '^'-"'^^ S. roiiilatiimBickn. DISCUSSION Sevenspecieswererecognizedinthisstudyandwillbebrieflydiscussed in alphabetical order. SisyrinchiumalbidumRaf.isasimple-stemmedperennialwithpaired sessilespathesatthenode,wherealargecaulincleafisfound. Flowersare usuallywhitewithyelloweye-ringsandtheglobosecapsulesdrypalebeige toastrawcolor. Populationsbloom lateMarch toAprilandarefoundin prairies, woods, and roadsides. SisyrinchiumangustifoliumMill, isthemostcommonand mostro- bust species ofBlue-eyed-grass in the state, with fairly wide leaves and 601 stemswithconspicuouswings.Thisperennialproducesstemsthattypical- lyhave 1nodewithacaulineleafwheretwopedunclesemerge.Flowersare light blue in color with yellow eye-ringsand theglobose tosubglobose capsulesdrydarkbrown.Plantsarefoundinfields,woods,oralongroadsi- desinAprilandMay. Thistaxonhasaveryconfusingnomenclaturalhis- tory which is presented elsewhere (Hornberger 1987a), but several manualscurrentlyinusehavenamesconsideredsynonyms:S.bermudtanum L. emend. Fern, and S. graminoides Bickn. (Gleason &Cronquist 1963; Steyermark 1963)- Sisryinchium atlanticum Bickn. is a branched species found in swciatthter1e-d2praniordieesa,reaansdinartehenasrtratoew.lyStweimnsgearde(te1retmemtowsildieg)h.tlySpfaltahtetseneadr,c small, often deflected at the base; bracts are equal to subequal, and the inner bract is distinctly mucronate. The oblong-subglobose to obovate capsulesdry darkbrown to black. Flowersaregenerally light bluewith yelloweye-rings, butthetepalsaresometimesdarkbluetopurple. This perennial taxon blooms from March to April. Sisyrinchium campestre Bickn. is commonly found in prairies, meadows, andgrassyareasalongroadsidesinAprilandMay. Thisperen- nial has asimple stem with asingle spatheat the topofthescape. The bractsofthespatheareverynoticeablyunequal,withtheouteroneatleast 1-1/2 to 2 timesormore the lengthofthe inner, gibbousone. Sisyrinchium langloisii Greene is a branched perennial found along grassy roadsides, prairies, and disturbed areas in March and April. Its spathebractstypicallyhaveapurplecolorationwhichisoftenrestrictedto thebaseofthespathe.ThistaxonissimilarmmorphologytoS.pnnnomm pBriecskenn.t,ianltthhoeulgahttetrh.eBpoutrhplreaxcaolhoarvaetiotnheofsatmheescpharthoemobrsaocmtesinsugmebneerra,ll2ynno=t 32(Lewis&Oliver 1961;Oliver&Lewis 1962),andhavebeenreportedto hybridize in areas of overlapping range (Correll & Johnston 1970). ComparisonofflavonoidspotprofilesbetweenapopulationfromTexasand another from Louisiana showed similar patterns (Hornberger 1987a). Morphology, chromosome number, and flavonoid chemistrysuggestthat these various populations mayactually be variationsofonelargespecies complex; therefore, I have synonymized S. pniinosum Bickn. under the oldername, S. langUnmGreene. Sisyrinchium rosulatum Bickn. is the only annual Blue-eyed-grass foundinArkansas,seeminglyrestrictedtoseveralsoutherncounties,plus PolkCounty. Flowersare yellow towhitewith purpleto maroonstripes and purple to maroon eye-rings. These flowers are urceolate, instead of rotateasdisplayedbytheflowersoftheothersixtaxa. Spathesareslender 602 a1n-d1f/o2liatciemoeLiss,lownigtehrtthheanouttheerinbrnaecrtbrsahcgth.tlGylofbaolcsaetetoatsutbheglaopbeoxseancdapsouflteesn drypalebeigewithpurpletobrownstripesalongthesutures. Populations can be found in disturbed areas ofroadsides and lawns, prairies, river bottoms,andpinewoods.Synonymsincluded.ex/^Bickn.(Smith 1978). Sisyrinchium sagittiferum Bickn. is represented only from Miller County,withpossiblehybrids{sagittiferum X Uinglonii)collectedinUnion County. Thistaxon is usuallyrepresented bysimple, leaflessstemsoften withfibrousbases.Spathesaresingle(orsometimespaired)atthetopofthe scapes, being conspicuously broader than the stems. Spathe bracts are equal or the outer one can be three times the length ofthe inner one. Flowersarebluetopurplewithyelloweye-rings,bloominMarchtoApril, andproducedarkbrown,globosetosubglobosecapsules,oftenwithsub- marginal veins. ThislattertaxonismostsimilarinmorphologytoS. campestreBickn., butdiffersinseveralimportantrespects: I)^.sagittiferumhasouterspathe bractsthatareconnateforseveralmm;S. campestrehasnon-connateouter bracts(less than 1.0 mm); 2)spathesof^. sagittiferumareconspicuously wider than the stems and dry brownish, often mixed with purple; S. campestrehasbractsthatdrygreenincolorandarenotconspicuouslywider thanthestems;and3)S.sagittiferumoftenhasfibrousleafbasesattachedto the stems, S. campestreseldom does. Sisyrimhiumsagittiferum Bickn. had MbeyenstruedcyordienddicfaorteAsrkthaantsaistsbhyoDuledmabreeeco(n1s9i4d3e)r,edbuptarntotofbAyrkSamnistahs'(1f9lo7r8a).. SUMMARY Basedondatacollected in thisstudy fromobservation, investigation, and literature review, sevenspecieso{Sisyruichiumare recognizedforthe state ofArkansas: S. alhidum Raf., S. angustifolium Mill., S. atlanticum Bickn.,^. campestreBickn.,S. langloisiiGreene,S. rosulatumBickn., and S.sagittiferumBickn. ThisinformationispresentedinSmith(1988). Dis- tributionmapswerepreparedforeachtaxonbasedonherbariumvoucher specimens. Adotindicatesthatatleastonespecimenexistsforaparticular taxoninaparticularcounty(Fig. 1).(Note:thedotinUnionCountyfor.V. sagittiferumBickn. representspossiblehybridsbetween itandS. langloisii Greene). a(:knowli;dc;i:mi;nts IthankE. B. Smithforservingasmydissertationadvisor.Manythanks toJerryShellandhiswonderfuldog,Mittens,forhelpingmecollectflower buds. I would also like to thank the Arkansas Native PlantSociety for ^ 603 g. albidumRaf — S. campestre Bickn. p. lanaloisil Greene S. rosulatuinBickn. g. flaaitt-iferumBickn. FIG 1.DocumenteddistributionofArkansasSnyrinihiiim. awardingmetheAileenMcWiUiamsScholarshipinBotanyin 1986which provided encouragementand financial support to finish mydissertation. Specialthanksarealsoextendedtothecuratorsofthefollowingherbariafor lending mespecimens that madethis studypossible: ASTC, BH, BLH, BM, FLAS, FSU, GA, GH, LAP, MA, MIN, MO, NCSC, NCU, ND, NDG, NLU, NY, PH, SMU, TENN, UAM, UARK, US, andWIS. 604 ri:fi:renc[^.s CHOORWtRoDElMLy'rLNt,,acDeW.a.e.SM..Aamne1dr94.M^..J.CA.BolJtisO.tHo3Nf2(cS2hT)r;Oo8NmIo.-so91m29.e70n.umMbaenrusalmofhitghheevrasplcaunltasr.p1l.anAtcsanofthTaecxeasa.e DEMTeAxRasEER,eseaDr.ch1F9o4u3n.datAionc,ataRleongnueer.of188th1eppv.asc(uSliasyrrmpclhainimtis,opfp.Ar4k2a5n-sa4s2.8)T.axod.um 1(1):!-88. {S!syyimhni»i, p. 21). GLEASON,H.A.andA.CRONQUIST 196^.Manualofvascularplantsofnortheastern UnitedStatesandadjacentC;anada. D.VanNosrrandCo.,NY.810pp {Si^rinchn/m pp. 219-220). GOLnDeoBtLroApTicTa,lIRnda1c9e8a2e..CShysrto.moBosto.me7(2c)y:t1o8lo6g-yi1n98r.elationtosupragenericsystematicsof HlLlV.i,rgiLn.iaM..Cas1t9a8n4.caA49(f2l)or:i6s2tic-6a8n.d chromosomal study oiiSisyrimhii/m (Iridaceae) in HORNBERGER, K. L. 1987a. SystematicsofthegenusSLsyrimhimu(Iridaceae)mthe southeasternUnitedStates. PhDDiss.,Univ. ofArkansas. 328pp. INGDiRsAs.M,,UnRi.v.19o61f49D8.u7Tbrh.heaItmna.xAo.1n0oL5omvypep,a.nCdhcryotmoloosgoymoeftnhuemgbeenrusrSeipsoyrrtisncXhCiVim.(ITraixdaocneae3)6.:4P9h7.D 1967.OntheidentityoftheIrishpopulationsofSisyrimivi/m Watsonia 6(5): 283-289. LEWIS,W.H.andR.L.OLIVER. 1961.MeioticchromosomesinsixTexasandMexican Ni'Dui.UylisantiSnyrimhuim(Iridaceae). Southw. Naturalist6(l)-45-46 MASSEY,J. R.andL.S.RADFORD. 1981.Revisedconrtibutors'guideforthevascular OLIGfVlaoErrdRa.,ot5R3t.:he1L0.s.ovu1t9h6e6a.stIenrnchUrnoimtoedsoSmteatensu.mbUenirvs.ooffpNhoarnetrhogCaarmosl.inaI..A2n1nppM.issouriBot ^ and W. H. LEWIS. 1962. Chromosome numbers of Snynmhn,YW (Iridaceae)ineasternNorthAmerica. Sida l(l):43-48. SMACLhLap,ej!.HKi.ll.193135.54Mappn.ua(lSiosfyrtihnechi./s/omu,thpeaps.te3rn27f-lo3ra3.0T)h.eUmv.ofNorthCarolinaPress, SMITH,E.B. 1978.AnatlasandannotatedlistofthevascularplantsofArkansas.Student yUrnniicohniuBmo,okpspt.or4e8,1—Un4i8v2.).ofArkansas, Fayetteville. 592pp. [OUTOFPRINT].{Sis- 1988.AnatlasandannotatedlistofthevascularplantsofArkansas.2nd STEe31dY87.2E18KR-i3Mpn8pAk2.oR)s{.K,Si,s6yrJ5i.3inAhWi.ieisi1nt9,6D3pi.pc.kF,l4soo6rn6a,—oFfa4yM6ei7t.)ts.e.vsiolulne.,TAhRe,I7o2w7a0S1t.at4e8U9nipvpe.rs{iSt,ysPyrnemshs,u/Amme'nsp^.'

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