Revision of the Section Hololeuce Bunge of the Genus
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TurkJBot 28(2004)307-347 ©TÜB‹TAK ResearchArticle RevisionoftheSectionHololeuce BungeoftheGenusAstragalus L. (Leguminosae)inTurkey MuratEK‹C‹ GaziUniversity,FacultyofArtsandScience,DepartmentofBiology,06500,Teknikokullar,Ankara-TURKEY TunaEK‹M ‹stanbulUniversity,FacultyofScience,DepartmentofBotany,34460Süleymaniye,‹stanbul-TURKEY Received:16.01.2003 Accepted:05.09.2003 Abstract: WerevisedthesectionHololeuce BungeofthegenusAstragalus L.(Leguminosae)inTurkey.Thesectionusedtocontain 20taxainTurkey,butinthisstudyithasbeenreducedto15.Anewspecies(Astragalusdumanii Ekici&Aytaç)hasbeendescribed. Inaddition,weconcludedthatA.vuraliiH.Duman&Aytaç,A.cylindraceus DC.,A.alindanus Boiss.,A.paecilanthusBoiss.&Heldr. andA.squalidus Boiss.&NoëaresynonymsofAandrasovszkyi Bornm.,A. globosusVahl,A.hirsutusVahl,A.sibthorpianus Boiss. and A.amoenus Fenzlrespectively.Thepollengrainsaretricolporate,prolate,subprolateorprolatesphaeroidalandthe ornamentationisreticulate. KeyWords: Turkey,Revision,Astragalus,sect.Hololeuce Türkiye’ninAstragalus L.(Leguminosae)CinsininHololeuce BungeSeksiyonununRevizyonu Özet: Türkiye’ninAstragalus L.cinsineaitHololeuce Bungeseksiyonununrevizyonuyapılmıfltır.Seksiyonülkemizde20taksonasahip ikenyapılanbuçalıflmaylasayı15’eindirilmifltir.Bilimdünyasıiçinyenibirtür( Astragalusdumanii Ekici&Aytaç)betimlenmifltir. AyrıcaA.vuralii H.Duman&Aytaç A.andrasovszkyii Bornm.’nin;A.cylindraceus DC.A.globosus Vahl’un;A.alindanus Boiss.A. hirsutus Vahl’un; A. paecilanthus Boiss.&Heldr. A.sibthorpianus Boiss.’un; A.squalidus Boiss.&Noë A.amoenus Fenzl’un sinonimleriolduklarıtespitedilmifltir.Polenlertrikolporat,prolat,subprolatveyaprolatsferoyidveornemantasyonuretikul attır. AnahtarSözcükler: Türkiye,Revizyon,Astragalus Sek.Hololeuce Introduction 1975),Caraganella Bunge(Podlech,1975),Theiochrus DuringthepreparationoftheFloraofTurkey,many Bunge (Podlech&Kozik,1983), Herpocaulos Bunge taxonomicalproblemsinsomegeneraandsectionswere (Podlech,1984), Caprini DC.(Podlech,1988), onlymentionedbutnotsolvedbecauseofthelimited Hypoglottidei DC.(Maassoumi,1988), Campylanthus timeandmaterial.Theeditorssuggestedthatfurther Bunge, Microphysa Bungeand Poterion Bunge(Tietz, revisionshouldbecarriedouttosolvetheseproblems 1988), Platyglottis Bunge(Podlech,1990), (Chamberlain&Matthews,1970;Davis&Hedge, Chlorostachys Bunge, Phyllobium Bungeand 1975;Davis,1975). Astragalus L.,therichestgenus Skythropus Simpson(Wenninger,1991), Megalocystis thatbelongstothefamily Leguminosae, isanexample Bunge (Tietz&Zarre,1994), Hymenostegis Bunge withmanyunresolvedtaxonomicalproblems.Thus, (Zarre&Podlech,1996), Dasyphyllium Bunge(Aytaç, certainsectionsofthegenus Astragalus havebeen 1997), Adiaspastus Bunge, Macrophyllium Bungeand revisedsincethepublicationofvolume3oftheFloraof Pterophorus Bunge(personalcommunicationwith Turkeysuchas Acanthophace Bungeand Aegacantha Zarre),and Alopecias Bunge(personalcommunication Bunge(Deml,1971), Stipetella G.Grig.(Podlech, withAkan). 307 RevisionoftheSectionHololeuce BungeoftheGenusAstragalus L.(Leguminosae)inTurkey ThefirststudyonthegenuswascarriedoutbyPallas Podlech(1988;1999;2001);Davisetal.(1988); (1800),followedbythoseofDeCandolle(1802)and Greuteretal.(1989);Maassoumi,(1989;1994;1998); Bunge(1868,1869).Theoriginsofthegenusextendto Yakovlevetal.(1996);Aytaç,(2000). the1700sandashorttaxonomichistoryofthegenusis Asitisimpossibletostudyallthespeciesatthesame givenontheinFigure1. time,itismorereasonabletostudythegenus sectionby Tournefourtcollectedsomethornyspeciesof section. Astragalus fromtheMiddleEastthathenamed ThemembersoftheHololeuce sectionaredistributed Tragacantha.Later,asseeninFigure1,severalscientists acrossthecentralandeasternpartsofAnatolia,the havestudiedthegenus.In1983,theGermanbotanist MediterraneanandBlackSearegionsinTurkey, Podlechdecidedtoseparatesomespeciesofthisgenusas Afghanistan,Armenia,Azerbaijan,Georgia, Iraq,Iran, anewgenus Astracantha, delimitedbyhaving Lebanon,TajikistanandTurkmenistan.Theworld “paripinnateleaveswith spiny rachisandflowersbornein distributionofthesectionHololeuce isshowninFigure2. axisofleaves ”(Podlech,1983).Neverthless,afterthis separation,itwasrealisedthattherearemany TheFloraofTurkey,Volume3(Chamberlain& transitionalspeciesbetweenthese2genera.Thatthey Matthews,1970),recognised20specieswithinthe boundariesofTurkey,16ofwhichwereconsidered werereunitedagainasthegenericdelimitationofthese2 endemic.Someofthespeciesareonlyknownfrom generaisnotjustified(Zarre&Podlech,1997). ancientgatheringssuchas A.sibthorpianus (1850), A. Astragalus isrepresentedbyc.10subgenera,130 andrasovszkyi (1911)and A.caudiculosus (1916).In sectionsand2500taxaworldwide.Acomparisonofthe somespecies,someimportantmorphologicalcharacters numberofspeciesinTurkeyandothercountriesisgiven aremissing.Forexample,in Astragaluscaudiculosus, the inTable1Shishkin(1965);Chater(1968);Chamberlain fruitcharacterwasnotknown.Inaddition,thereisa &Matthews(1970);Tackhom,(1974);Townsend& similaritybetweenthesections Hololeuce and Guest(1974);Hedge&Podlech(1987);Zohary(1987); Onobrychium.Someofthespeciesof Onobrychium are Tournefort (1700): Astragalus Tragacantha Astragaloides Linnaeus (1753): Astragalus Phaca DC (1802): Oxytropis Astragalus Phaca Gray (1864): Astragalus Von Bunge (1868) Astragalus Phaca Tragacantha etc. Podlech (1983): Oxytropis Astragalus Astracantha Zarre&Podlech(1997): Oxytropis Astragalus Figure1.Theshorttaxonomichistoryof Astragalus. 308 M.EK‹C‹,T.EK‹M Table1.ComparisonofthenumberofAstragalusspeciesbetweenTurkeyandotheroldworldcountries. Region Country TotalSpecies End.Species End.rate Iran 678 381 56.19% SaudiArabia 21 1 4.76% W.ASIA Iraq 145 9 6.20% Afghanistan 293 144 49.14% FormerUSSR Russia 1005 562 55.92% Europe 142 50 35.21% Europe Turkey 446 210 47.08% Syria-Lebanon 59 15 25.42% M.EAST Jordan 35 2 5.71% Palestine 52 4 7.69% India 54 4 7.40% S.ASIA Pakistan 135 26 19.25% E.ASIA China 278 22 7.91% Egypt 34 2 5.88% Algeria 39 2 5.12% AFRICA Morocco 45 9 20.00% Libya 89 24 26.96% N 0500km Figure2.WorlddistributionofthesectionHololeuce. 309 RevisionoftheSectionHololeuce BungeoftheGenusAstragalus L.(Leguminosae)inTurkey transferredto Hololeuce, whereassomespeciesof andCltweremeasuredandtheratioofClg/Clt Hololeuce aretransferredto Onobrychium.Therefore, it calculated. wasdecidedtocarryouttherevisionofsectionHololeuce - Structure:Intheopticcross-section,thecharacters presentedhere;therevisionofthesection Onobrychium ofalayerofpollen. isalsoplanned. - Sculpture: Intheopticcross-section,thesurfaceof exineanditsornamentation. MaterialsandMethods - Thelengthofporesonthepolarside(Plg)and Researchmaterialsofthesection Hololeuce were equatoriallengthweredetermined. collectedfromdifferentpartsofTurkeyinApril-August, Inaddition,thedetailsofthepollenandmorphology bothinthefloweringandfruitingperiod,between1996 wereexaminedusingascanningelectronmicroscope and1999.Inordertostudythevariationpatterns, (SEM). Thesizesofthepollenofallspeciesaregivenin specimenswerecollectedfromasmanydifferentpartsof Table2. theexistingdistributionareaofthesectionaswas possible.Inaddition,thecollectionsofAEF,ANK,ATA,B, SEMMethod CUM,E,EGE,GAZI,HUB,ISTF,K,KNYA,M,MSBandW Pollengrainsweretransferredtostubsandcoated herbariawereexamined.Theauthornameofevery withgold.Afterstudyingtheresultingphotographs,we specieswaswrittenaccordingtotheAuthorsofPlant couldnotfinddiagnosticdifferences. Names(Brummitt&Powell,2001). Taxonomictreatmentofthesection: MorphologicalStudies Astragalus sect. Hololeuce BungeinMém.Acad. Arangeofcharacteristicsthatwereconsideredtobe Imp.Sci.SaintPétersbourg11(16):104.1868. taxonomicallyimportantinthesectionwasinvestigated. Lectotype:A.hololeucus Boiss.&Buhse(Podlech, Theseincludethefollowing:stipulestructureandlength, 1990). leaflength,leafletshapeandlength,theinflorescence, bractshapeandlength,whetherthecalyxisinflatedor =A.sect.Chlorosphaerus BungeinMém.Acad.Imp. not,thecorollastandard,wingandkeelandthe Sci.SaintPétersbourg11(16):106.1868. structure,andsizeofthefruitbeak.Measurementswere Lectotype: A.chlorosphaerus Boiss.(Podlech, obtainedfromatleast10replicatesoneachof10 1990). specimensandtheaveragevalueofeachcharacterwas Scaposeorshortlycaulescent,perennialherbs. evaluated.Themaximumandminimumvaluesofthe Stipulesfreeoradnatetothepetioles.Leaves measurementsofthesepartsaregiveninthedescriptions imparipinnate;leaflets4-20-paired,withbifurcateor ofthetaxa. sub-bifurcatehairs;narrowlyovateornarrowlyelliptic, Palynologicalanalysis oblongorobovate.Inflorescenceadense,pedunculate many-floweredspike;globose,ovateorcylindrical. Thepollenmorphologyofthespecieswasexamined Pedunclegenerallywhite-hairyandalmostaslongasor usingaKyowaMicro-Lux-11lightmicroscope(10x100) longerthanleaves.Bractslinear,triangular-ovate,with andpollenpreparationsweremadebytheWoodhouse whiteorblackhairs.Bracteolesabsentor1-2.Pedicels methodandstainedwithbasicfuchsin(Woodhouse, generallyabsentorupto1mm.Calyxmembranous, 1959).Theirdescriptionshavebeendrawnupusingthe generallytubular,hairy;calyxteethlinear,triangular; morphologicalcharactersgivenbelow(Pehlivan,1995): generallywithblackhairsoutside,glabrousorwithonly - Pollentype:Theapertureshapeandarrangement whitehairsinside.Standardglabrous.Clawofwingsand onthepollen. keelonlyadnateatthebasetostaminaltube.Stamens - Pollenshape:Thedivisionofthepolaraxis(P)tothe diadelphus,filamentsglabrous.Staminaltubestraightly equatorialaxis(E). cut.Ovaryovateorelliptic,withwhitehairs.Fruitsovoid, oblong;bilocular,almostaslongascalyx,beakstraightor - Theexineandintinethickness. curved;2seedsineachlocule,Seedsrectangular- - Thepolarandequatorialdiameterofthecolpus:Clg