Meclatisi n Clematis: Yellowflowering Clematis species
Systematic studies in Clematis L.(Ranunculaceae) , inclusive of cultonomic aspects Promotor: dr.ir L.J.G. van derMaese n Hoogleraar in de Plantentaxonomie UNO8?Z\,Z% ' 1
Meclatisi n Clematis: Yellow flowering Clematis species
Systematic studies in Clematis L. (Ranunculaceae),inclusiv eo f cultonomic aspects
WillemA .Brandenbur g
Proefschrift terverkrijgin g van degraa d van doctor opgeza gva n derecto r magnificus van Wageningen Universiteit, dr.CM . Karssen, inhe topenbaa r te verdedigen op maandag 26jun i 2000 desnamiddag st e half tweei n deAula .
*JY-\ ^OjX^\ Clematis'Bravo '
Date of publication: 8Jun e 2000
Brandenburg, WillemA . Meclatisi n Clematis: Yellow flowering Clematis species -Systemati c studies inClematis L. (Ranunculaceae), inclusive of cultonomicaspect s/ Willem A. Brandenburg ISBN 90-5808-237-7 Subject Headings: Systematics, Cultonomy, Phylogenetics,Phytogeography , Morphology, Cytology, Palynology, Clematis, Meclatis,Ranunculacea e CONTENTS 0 Preface 1 1 Historical survey of classification and delimitation of the genus Clematis L. 5 1.1 Pre-Linnaean treatments of Clematis. 5 1.2 Classification of Clematis from Linnaeus onwards 7 1.3 Phylogenetic analysis of the genus Clematis 17 1.3.1 Cladistic analysis of subdivisions of Clematis and of related genera 18 1.3.2. Biogeography of Clematis 38 1.4 Interspecific crosses in Clematis 54 1.4.1 Introduction toth e experiment 54 1.4.2 Material andmethod s 54 1.4.3 Results and discussion 57 1.4.3.1 Seed set and offspring of diallel and othercrosse s 58 1.4.3.2 Pollen tubegrowt h experiments "* 66 1.4.3.3 Discussion andconclusion s 78 1.4.4 Speciation and species concepts in Clematis 79 1.5 Description of thegenu s Clematis 82 1.5.1 Growth habit 82 1.5.2 Leaves 85 1.5.3 Synflorescence and inflorescence 85 1.5.4 Flower 89 1.5.4.1 Perianth 92 1.5.4.2 Nectar leaves 94 1.5.4.3 Stamens 94 1.5.4.4 Pistils 97 1.5.5 Fruit 98 1.6 Summary description of Clematis 98 2 Clematis sectionMeclatis (Spach ) Baillon 99 2.1 Description of the sectionMeclatis (Spach ) Baill. 99 2.2 Distribution of the sectionMeclatis (Spach ) Baill. 101 2.3 Morphological analysis of the sectionMeclatis (Spach ) Baill. 106 2.3.1 Material andmethod s 106 2.3.2 Results 112 2.3.2.1 Factor analysis 112 2.3.2.2 Principal coordinate analysis 115 2.3.2.3 Cluster analysis 116 2.3.2.4 Phylogenetic analysis 125 2.3.2.5 Conclusions 128 2.4 Species descriptions 129 2.4.1 Keyt oth e species of Clematis sect.Meclatis 129 2.4.2 Clematis orientalis L. 130 2.4.2.1 Lectotypification of Clematis orientalis L. 130 2.4.2.2 Thetypotyp e of Clematis orientalis L. 134 2.4.2.3 Thetyp elocalit y of Clematis orientalis L. 136 2.4.2.4 Synonymy and description of Clematis orientalis L. 139 2.4.3 Clematisgraveolens Lindley 147 2.4.4 Clematis intricata Bunge 151 2.4.5 Clematis ispahanica Boissier 156 2.4.6 Clematis serratifolia Rehder 160 2.4.7 Clematis tibetana O. Kuntze 164 2.4.7.1 Clematis tibetana O. Kuntze subsp.tibetana 165 2.4.7.2 Clematis tibetana O. Kuntze subsp.tangutica (Maximowicz ) Brandenburg comb.nov . 169 2.4.7.3 Clematis tibetana O. Kuntze subsp.vernayi (C.E.C .Fisch. ) Grey-Wilson 175 2.5 Cytology 179 2.5.1 Material andmethod s 179 2.5.2 Results and discussion 181 2.5.3 General discussic# 186 2.6 Pollen morphology 188 2.7 Isozymepolymorphis m inMeclatis 191 2.7.1 Material andmethod s 191 2.7.2 Results anddiscussio n 192 3 Classification of cultivated plants 195 3.1 History of classification schemes concerning cultivated plants 195 3.2 Principles of open classification outlined 198 3.3 The taxon concept and cultivated plants 206 3.4 Theculto n concept 210 4 Classification ofcultivate d Clematis 215 4.1 Clematis asa garde n flower -th e first modern treatise of cultivated Clematis 215 4.2 Cultivarclassificatio n of large-flowered Clematis 216 4.3 Introduction intocultivatio n of Clematis sect.Meclatis 217 4.4 Cultivars of Clematis sect.Meclatis 220 4.4.1 Clematis'Aureolin ' 221 4.4.2 Clematis'Bil l McKenzie' 222 4.4.3 Clematis'Bravo ' 223 4.4.4 Clematis'Burford ' 225 4.4.5 Clematis'Corry ' 226 4.4.6 Clematis'Drake' s Form' 227 4.4.7 Clematis'Golde n Harvest' 227 4.4.8 Clematis'Helios ' 227 4.4.9 Clematis'Lambton' s Park' 228 4.4.10 Clematis'Orang e Peel' 228 4.4.11 Clematis'Wallsal ' 230 4.5 Conclusions 230 5 General discussion 231 5.1 Clematis species withregar d to various species concepts 231 5.2 Crossability dataan d their value for plantbreedin gresearc h 231
VI 5.3 Evolution and domestication of Clematis 238 5.3.1 Evolutionary aspects of Clematis within the Ranunculaceae 238 5.3.2 Domestication of Clematis 239 5.4 Cultivar documentation 241 5.4.1 Cultonomy, identification andcharacterizatio n 241 5.4.2 Clematis cultivars andthei r validation of cultivation and usage characteristics 242 6 Summary 245 Samenvatting 247 7 Curriculum vitaeWille mA .Brandenbur g 249 8 References 251 9 Index to Clematis species in chapters 1-5. 285 Appendix 1 (ondisk ) 289 Appendix 2 (ondisk ) 293
vn Stellingen behorende bij het proefschrift, getiteld: Meclatisi n Clematis: Yellow floweringClematis species -Systemati c studies in Clematis L. (Ranunculaceae), inclusive of cultonomic aspects
I
Het onderscheid tussen houtige en kruidachtige Clematissoorte n heeft tot dusver een goede interpretatie van declassificati e van het genusi n dewe g gestaan.
Dit proefschrift, hoofdstuk 1.
n
Hetonderschei dtusse nsynflorescenti ee ninflorescenti ei snutti gbi jhe tbeschri jve nva nplanten ;he t veronachtzamenva ndi tonderschei dbemoeilijk td einterpretati eva nbloemgestelle ni ngener aal s Clematis.
Troll, 1964,1969 . Ditproefschrift , hoofdstuk 1.
m
Clematis is een goed genus om de hypothese te toetsen dat daar waar de grootste variatie gevonden wordt het ontstaansgebied is dan weljuis t niet.
Ditproefschrift , hoofdstuk 1.
rv
Dehie rvoorgesteld eindelin gva nClematis sect .Meclatis (Spach )Baillo nmaak tduidelij k datd e aanwezigheid van variatie nog niet onmiddellijk aanleiding moet zijn een nieuwe soort of een infraspecifiek taxon tebeschri jven .
Ditproefschrift , hoofdstuk 2.
V
Depollenmorfologi e van Clematis verdient systematische aandacht.
Ditproefschrift , hoofdstuk 2. VI
Clematis is als genus van sierplanten buitengewoon interessant omdat de complete reeks van bloemkleuren eri n voorkomt. vn
Hetfei t date rverwant eniet-kruisbar e planten naastkruisbar eminde rverwant esoorte n bestaan maakthe t biologisch soortsbegrip onhoudbaar: een waarschuwing voor plantenveredelaars!
Dit proefschrift, hoofdstuk 1 en5 .
vm
Decultiva rgroe pal seenhei dva nclassificati ei steru gt evoere no p19 deeeuws etuinbouwkundig e classificaties van sierplanten, waaronder declassificati e van toen in cultuur zijndeClematis.
Dit proefschrift, hoofdstuk 3e n4 .
DC
Detraditionel ebehandelin gva nkunstmatig ehybride si nd eplantensystematie ki see nfundamentel e misvatting van demogelijkhede n en deresultate n van demodern e plante nveredeling .
Dit proefschrift, hoofdstuk 3.
X
De introductie van het begrip culton was noodzakelijk om goede, flexibele classificaties van cultuurplanten metbetrekkin g tot debetrokke n eenheden (cultivar, cultivargroep) te ontwikkelen.
Hetterscheid &Brandenburg , 1995a, 1995b Hetterscheid et al., 1996 Dit proefschrift, hoofdstuk 3.
XI
Datvoo rcultuurplante nme tbehul pva nbotanisch eclassificatiesysteme n geenclassificatie s zijnt e maken,val ta ft eleide nui the tfei tda te rvoo rgee nenkel ebelangrijk ecultuurplan tzo 'n classificati e bestaat die algemeen is aanvaard.
II Brandenburg &Schneider , 1988 Ditproefschrift , hoofdstuk 3.
xn Destudi eva nhemicyclisch ebloeme nverdien tmee raandach tva nontwikkelingsbiologe nda nto tn u toe het gevalis .
xin Genoomgrootte in termen van het aantal chromosomen per genoom kan nog steeds niet goed wordenverklaard .D eRanunculacea ezij ni ndez eee ngoe d 'proefkonijn' vanweged everschillen - de, systematisch verspreide aantallen (x=6, 7 of 8) en de daarmee verband houdende chromo- soommorfologie.
Ditproefschrift , hoofdstuk 2.
XIV
Dehuidig emethode sva nfylogenetisch e analysehebbe nd eplantensystematie kto tee nwetenschap - pelijkedisciplin egemaak tme ttoetsbar ehypotheses ;d emoleculair ebiologi ei see nzee rkrachti g hulpmiddeldaarbij .
Ditproefschrift , hoofdstuk 5.
XV
Heti szorgwekken dda td ebasi sva nd ewereldvoedselvoorzienin g steedsmee rafhankelij k wordt van steeds minder multinationale ondernemingen.
XVI
Ind eWesteuropes ecultuu ri sgee nplaat smee rvoor 'verliezers' ;'winners 'e nwins ti swa te ri nhe t centrumva nd eaandach tstaat .Da tdaarme eoo kd ekuns tva nrouwe nda nwe lverliesverwerkin g verloren dreigt tegaa n isechte r geen winst maarech tverlies .
xvn Rouwen iseerde r kunst dankunde .
Ill xvm Anton Brucknerwa sbehalv eee nbegenadig dcomponis t ookee n zeergoe dorganist : dathi j als componisthe torge lnooi tui the too gheef t verloren,blijk tui the tfei t datme tnam ei nzij n latere symfonieen (7-9) de rusten zo zijn gekozen dat de muziek langzaamwegebt : deze symfonieen komen dan ookhe t beste tot hunrech t in grotekerke n zoals dedomker kt eLiibeck .
XIX
De zesde symfonie van Gustav Mahler heeft ten onrechte de bijnaam Tragische Symfonie' gekregen; echt tragisch in deuitvoeringspraktij k van deze symfonie is alleen dat een verkeerd gekozen tempoi n deeerst e 24mate n desastreus is voor deverder e uitvoering van de symfonie.
XX
Verkeer en verkeerd schelen maaree n letter en dat kan helaas maar al tevaa k op de openbare wegworde n waargenomen.
IV PREFACE
In 1975, this Clematis study started as an experimental systematics project under supervision of Dr. R.A.H. Legro (Brandenburg, 1976, 1977a, 1977b). The aim of this study was to reveal the degree of relationship between Clematisspecie s of (potential) economicimportanc ean dt oovercom ecrossin gbarrier sbetwee nthem .Thi sstud ywa sth e continuation ofearlie r workb yBarendrech t (1972).Barendrech t alreadyindicated , that many cultivated Clematis species were assigned to speciest owhic h theyeithe r did not belong or with which they cannot be any longer solely linked because of repeated interspecifichybridization .Thi sreferre despeciall yt oth egrou po fyellow-flowere d species, initiallyindicate da sClematis ser .Orientates Prantl :e.g .C. orientalisL. ,a si twa slabelle d in cultivation, showedremarkabl e differences toth e wild plant. The above project was later taken as the starting-point for an extended experimental systematicsprojec t tob ecarrie d outi nth eframewor k ofth eresearc hprogramm eo fth e then AUDept . Taxonomy of Cultivated Plants andWeeds . In the period 1978 - 1984, the research project was still focussed on the crossabilitypotentia lo fth especie sconcerned .I nthi speriod ,i tbecam eclea rtha tClematis is not a very suitable plant to be used in hybridization studies because of complex germination behaviour, slow growth to an adult flowering plant and large variation in flowering season per species.I twa s therefore decided toredefin e theprojec t in amor e morphologic taxonomic way;th econsequenc e ofthi sbein g alarg e seto f chromosome counts that can hardly be used for the new project. Also other data sets remained incomplete (pollen morphological andbiochemica l data). In the period 1984 - 1987,th e guidance of the project was laid in the hands of Prof.Va nde rMaese n andth eprojec t waslargel yfocuse d onth e systematics ofyellow - flowered Clematis spp.,a sfo r thesespecie sa representativ e datase tcoul db eobtained . Besidesthat ,dat awer ecollecte dt ocompil ea ninternationa lregiste rfo rClematis cultivars . Thelis to flarge-flowere d Clematiscultivar s hasbee ncomplete d upt o 1991an dwil lb e published separately. Thestructur eo fthi sthesi sreflect s somewhatth edirectio no fth eevolutio no fbot h
1 theevolutio ni nthinkin gconcernin gthi sparticula rresearc hprojec tan dm yothe rpersona l developmenti nsystematic so fcultivate dplants ,whic hwa sdirecte dtoward sprinciple san d concepts in classification of cultivated plants, and towards the development of an unequivocalnomenclatur eo fcultivate dplants ,culminatin gi nth edevelopmen to fth eope n classification concept,a ple afo ra consequen tapplicatio no fth ecultiva rgrou pconcep tan d the establishment of the culton concept (Brandenburg et al., 1982;Brandenburg , 1984, 1986a, 1986b; Brandenburg & Schneider, 1988;Hetterschei d & Brandenburg, 1995a, 1995b;Hetterschei d et al., 1996). Chapter1 dealswit hth egenu sa sa whol esurveyin gan danalysin gphylogeneticall y andbiogeographicall yth eclassificatio n ofsection san dsubsections .Th ecrossabilit ydat a obtainedbetwee n 16specie swer eanalyse da swel lan dviewe di nth eligh to f systematic evidence. Chapter 2 deals with the systematics of the Clematissect .Meclatis (Spach) Baillon.Base do nmorphologica lcharacter sthi ssectio nha sbee nanalyse dusin gmultivariat e andphylogeneti c analysismethod san dgivin gris et oa ne wclassificatio n ofthi sthu s far ambiguouslyclassifie d section.Th edat awer ecomplete dwit hdistribution ,cytologica lan d pollenmorphologica ldata .Isozym edat awer esample dincompletel yt osuc ha nextent ,tha t they only will be surveyed indicating their potential value but without any conclusive contribution ast othi ssystemati canalysis .Chapte r 3 consistso fa treatis e ofconcept si n systematics of cultivated plants, needed to deal properly with cultivated Clematisi n Chapter4 .Chapte r 5i sth egenera l discussion ofth ethesi sparticularl y dealing withth e fundamental problemsbrough tu pb ythi sthesi san dth epossibilitie sfo rapplication s from this and similar research in plant systematics.
Completiono fthi sthesi sha sbee nroughl ydon ei nthre eperiods .I nth eperio d198 7 -1991,th edat awer eanalyze dan dth egenera lsetu po fth ethesi sha sbee nmade .I n199 3 and 1994th efirs t versiono fth ethesi sha sbee nwritte nan di n 1999th elas tone .Th efac t thati tha stake ns olon gi sa matte ro fpersona lconditio nan dm yow nresponsibility .Th e fact thati tha sbee nfinalize d islargel ydu et o astimulatin genvironmen t for whichI a m deeply grateful.
Manystudent s ofth esmal lDepartmen tTaxonom yo f Cultivated Plants andWeed san d
2 laterth eDepartmen to fPlan tTaxonom yhav emad eimportan tcontribution st oal lsystemati c worko n Clematistha ti fno tdirectl yrelevan tt othi sthesi swa sa tleas timportan tfo rm y thinking about it. Thelat eR.A.H .Legr omad em eenthusiasti cfo r Clematis,th elat eProf .J.H .va n derVee nstimulate dm et ostar twit hpublishin gm yidea so nsystematic so fcultivate dplants . Prof.L.J.L.D . van Griendsven, ir. C.A.A.A.Maenhout , dr.N.G .Hogenboom , dr. CM. Colijn-Hooymans and dr. A.J. van Tunen respectively kept saying that Ireall y should finalize thisthesis . Ithan km ycolleague si nsystematic so fcultivate dplant san deconomi cbotan yi nth e Netherlands,Ronal d vande nBerg ,Nynk eGroendijk-Wilders , Marietd eGeus ,Wilber t Hetterscheid, Marjan Boonefo r their numerous discussions and suggestions on various topicsbu tespeciall yfo rthei rpatienc ewit hm ewhe ndealin gwit hClematis. Ruu dva nde r Meijdenintroduce dm eint oth efiel do ffloristic san dw eha da lo to fdiscussion so nth eedg e ofbot h sides: systematics of cultivated plants and systematics of wildplants . Josva nd eVoore nwa salread yinvolve di nClemati swor klon gbefor e Istarte da s astuden to nth esubject .Jos 'ope nmind ,innovativ ean dunorthodo xapproac han dstron g support really kept me going. Besides that we spent a lot of time together which was indispensable.Anj a vande rNeu ttoo ka kee n interest in Clematis orientalis andrelate d speciesan dmad ea majo rcontributio nt osettin gu pth erevisio no fClematis sect .Meclatis. Youbot hhav ealway ssupporte dm ebot hprofessionall yan dpersonall y insuc ha wa ytha t Ia mstil lactiv ei nth efields o fsystematic so fcultivate dplant san deconomi cbotan ytha tdi d not goa tal l without sayingdurin g themos trecen t period of my life. Iwoul d like tocommemorat e all myfriend s for their interest inmysel f and this undertaking.Al li nm yfamil y whokep tm eo ntrac kt ofinaliz e thisthesi sI woul dlik et o thank. There were times that Imad ething s difficult for youb yno t wanting to talk aboutit . Hanneke,w ear ebot ho ntrac kagai nan dconfiden t forth efuture .Th efac t thatI stil lma y talkabou tClemati si ssignifican tfo ryou rinteres ti nm ean dm yundertakings .I admir eyou r initiative tog oint o fashion andclothin gdesign .Wouter , Ia mhapp yt o seetha tyo u are studying sound engineering: keep cool,b ehapp y and gofo r it! 1. HISTORICALSURVE YO FCLASSIFICATIO NAN DDELIMITATIO NO FTH E GENUS CLEMATIS L.
1.1. Pre-Linnaean treatments of Clematis.
Etymologically,th ewor dClematis ha sbee nderive dfro micX.T||i a(klema) ,whic hmean svin e (Loudon, 1869).A OUTOVKX,nuaTeio v(phyto nklemateion ) is avine ,a climbin g plant. Accordingt oWittstei n (1856),variou sclimbin gplant swer eindicate d with Clematisi n Dioscorides' herbals and mediaeval herbals based upon it:
'Kkfyiaxv; Diosc. ist aber Vinca minor und eine andere KATUJOITK; desselben Schriftstellers ist warscheinlich Polygonum convolvulus;dahingege n stimmt KXTIUOUTK; Diosc. mit Clematis cirrhosa und eine andere KXriuaraic, Diosc. ist Aristolochiabaetica (nich tA. clematitis).' Thename sClematis an dClematitis wer euse dconcurrently ,th eforme rbein gth eLatinize d formo fKXnucmc; ,th elatte rbein gth eLati ntranscriptio naccordin gt oBacke r(1936) .Thi s explanationonl yhold swhe nKArmarau ;ha sbee nuse di nGree kfor mderive dfro mth eLati n transcription. Bothname s remain concurrent until post-Linnaean classifications. ©e6(ppaoroc,(Theophrastos )mentione da wil dvin ei nhi sHistoriaNaturali s(±32 0 BC)unde rth enam e aGpaysvTi(atrag6ne) : "Theyals osa ytha ta ver ygoo dfir e sticki smad efro m thewoo dwhic hsom ecal l traveller'sjoy :thi si sa tre elik eth evin eo rthe vwil dvine' ,which ,lik ethese ,climb su p trees(Hort , 1916,Englis h translation ofth eHistori aPlantarum , BookV ,6 ,9). ' Theophrastos'classificatio n wasbase do nbot hmedicina lo ragronomi ccriteri aan dbotanica l characters.Hi swor kstrongl yinfluence dth ebotanica lan dmedica lliteratur etill th e17 *century . Herbalsu pt oth e16 thcentur yar ebase do nknowledg ehande ddow nfro mancien ttime swit h asprincipa lsourc evariou srevise dversion so fDioscorides ' 1Mcentur yherba l(se eDioscoride s etal. ,1934) ,whic hwas ,however ,inferio rt oTheophrastos 'wor kb ylac ko fhi sow nbotanica l observations,an ddu et omisinterpretation so fsom eo fth eplant sconcerned .Onl ysom elate r 16th and 17thcentur y herbals consist of descriptions and data compiled byth e authors themselves(e.g .Brunfels , 1530;Clusius , 1601;Fuchs , 1542;Lobelius , 1576).Th enam e Clematiscovere da ttha timet a numbe ro fspecie stha tar estil lassigne dt oth egenus ,suc ha s C rectaL. ,C. vitalba L .an dC. viticellaL. ,a swel la sa rang eo fothe rplants ,suc ha sVinca minorL .an dV. majorL .unde rth ename sClematis daphnoides an dClematis daphnoides majorrespectivel y (Dodonaeus, 1583). Themai nreaso nfo rincorporatin gClematis spp .i nherbal swa sthei rmedicina luse . EspeciallyC. rectawa sknow ni nthi srespect .Va nde rNeu tan dPfeiffe r (1982)surveye d severalDutc hherbal sfo rmedicina luse so fClematis: puportedl ydiureti can ddiaphoreti c applications,expellin ggal lan dphlegm ,agains tvariou sdisorder san dischia scoul db ecure d withth eplants .Moreover ,Va nde rNeu tan dPfeiffe r (1982)mentione dtha tth eyoun gsprout s ofC. rectaan dC. viticellawer eeate na sa vegetable .Fo rC. rectathi si sremarkable ,a sthi s speciesi sals omentione da smos teffectiv e init smedicina lcharacteristics .Th emedicinall y effective chemical compounds areapparentl y absenti nyoun gsprouts .I n contemporary ethnobotanicliterature ,mentio ni sstil lmad eo fyoun gsprout so fvariou sClematis spp .eate n cooked asvegetabl e (Usher, 1974).
Clusius (1611) was one of the first to praise C.viticella a s an ornamental. He especially mentioned the double-flowered form inthi s context. BothRa y(1686,1724 )an dTournefor t (1700,1703)mor eo rles sestablishe dth e genusconcep t(Stafleu , 1971;Stearn ,1957) .Man yLinnaea ngeneri cname sar edirectl yo r indirectlyderive dfro mthei rpublications .A thir dprincipa lsourc eo fgeneri cname sfo rLinnaeu s wasBauhin' sPinax (1623) .Befor e 1736alread yRa yan dTournefor tfrequentl yreferre dt o Bauhin. Alreadyi nth e17 *an di nth efirs thal fo fth e18 *century ,ther ewa ssom edisagreemen t aboutwhic hspecie sshoul db eassigne dt oth egenu s Clematis. TakingLinnaeu s'Species Plantarum,ed .I (1753 )a sa starting-point ,i ti sremarkable ,tha ti nthi swor kAtragene wa s establisheda sa separat egenus ,wherea si nClematis specie swer ebrough ttogether ,tha twer e formerlyassigne dt oanother 'germs'Flammula,e.g .b yBauhi n(1623 )andDilleniu s(1732) . The species C. cirrhosa L., C crispa L., C.integrifolia L.,C. vitalbaL. , C.viticella L. werecommonl yassigne dt oClematis o r Clematitis,wherea sC. flammula L.,C. rectaL . andC. viornaL .wer evariedl yinterprete d asbelongin gt oeithe rFlammula orClematis. Thesedifferen tearl yopinion sar eth estarting-poin tfo rth edisagreemen to ngeneri cdelimitatio n of Clematis until this veryday . 1.2. Classification of Clematisfro m Linnaeus onwards
Apart from separatingAtragene, Linnaeu s (1753) subdivided Clematis into twogroups : *Scandentes 1. C. viticella 2. C viorna 3. C. crispa 4. C orientalis 5. C vitalba 6. C. cirrhosa 7. C.flammula *Erectae 8. C recta 9. C. integrifolia Thestatu so fth etw ogroup sremain sunclear ,a sLinnaeu sdi dno tspen da wor do nthe mi nhi s workso nclassification ,no ri nth evariou sedition so fth eGenera Plantarum. A sth eforma t heuse di sprincipall ydesigne dt ob ea nidentificatio nformat ,th etw ogroup sar et ob eregarde d asinforma l indications(Stearn , 1957;Stafleu , 1971).However ,th ename sar eindicativ e enough. InAtragene, Linnaeu s distinguished: 1. A. zeylanica( =Naravelia zeylanica(L. ) DC), 2. A. alpina( =C. alpina(L. )Mill ,p.p.) , 3. A. sibirica( =C. alpinap.p. ) and 4. A. capensis (=Anemone capensis (L.)Lam.) .
Althoughsom eauthor sstil lrecogniz eAtragene b yth epetaloi dstaminodia ,th evie wo fMille r (1768) toinclud eAtragene i n Clematis, is widely accepted.
Moenchemphasize di nMethodu sPlantaru m(1794 )bot hnumbe ran dfor mo rstructur eo f flower characters.Base d onthes echaracters ,h edefine d several 'classes', amongwhic h Thalastemon(Gcdauoc; ,thalamos ,i sreceptacle ,an d0Tr) u&>v ,stemoon ,stamen )with"stamin a receptaculoinserta' .H einclude dth egener aClematitis an dViticella i nthi s"class 'a swel la s variousothe rgenera ,no wconsidere dt obelon gt oth eLeguminosae ,Liliaceae ,Rosacea ean d otherfamilies .H edistinguishe dViticella fro m Clematitis( =Clematis) b yshort-hair ystyle s and glabrous ovaries.H e distinguished the following species: Clematitis crispa Clematitisflava (= Clematis orientalis L.) Clematitis vitalba Clematitis integrifolia Viticella deltoidea( = Clematis viticellaL. ) Withhi sgeneri cname sMoench 1referre d topre-Linnaea n Clematitis(Tournefort , 1700; 1703),an dViticella (Dillenius ,1732) .Followin gth ecriteriafo r separatingViticella fro m Clematitis, iti sremarkabl etha t Clematitis crispai s not grouped under Viticella, asdi d Dillenius(173 2cite db yMoench) .Th eMoenc h systemi sinconsisten ti nit s infrageneric classification of Clematitis and Viticellaan d has not found general acceptance. His classification schemeo f "classes'di dno t survive either.
Persoon(1805 )largel ymaintaine dth eclassificatio nb yLinnaeus ,bu textende di twit hspecie s describedmeanwhile .Hi streatmen to fClematis i sth elas ton ewit ha subdivisio ni nScandente s andErectae ,obviousl ymean ta sa ninforma lsubdivision .Quit eremarkabl ei sth eplacemen to f ** Viornaunde rAtragene, referrin gt ospecies ,late rclassifie dunde rth esectio nCheiropsis byD eCandoll e(1818) .A sh eassigne dth especie sC. viornaunde rClematis * Scandentes, iti sa nobviou smistake .Persoo n transferred someearlie r described Clematis speciest o Atragene.Thes especie s(e.g .C.florida Murra ye xThunb. )occasionall yproduc epetaloi d staminodia,henc e (semi-)double flowers.
A.P.D eCandoll e (1818)i nhi sRegn iVegetabili s SystemaNatural eha da broa dvie wo f Clematis.Withi nth etrib eClematideae ,h edistinguishe donl ytw ogenera : Clematis,inclusiv e ofAtragene, an dNaravelia, whic hLinnaeu s (1753) assigned formerly toAtragene. D e Candolle subdivided Clematis into4 sections,base d oninflorescenc e and fruit characters: sect.Flammula DC. achenes withbarbate-plumos e styles; sect. Viticella (Moench) DC. achenes with short, +barbat e styles; sect.CheiropsisDC. aninvolucru munde rth eflower ,achene swit hbarbat e styles;
I Although Moench variously spelled his name asMoenc h andMonch ,th e standard author citation ascompile d byBrummit t and Powell (1992)i sfollowe d (see also entries 6165 -616 9i n Stafleu and Cowan (1981)). sect.Atragene (L. )DC . numerous petals,achene s withbarbat e styles. DeCandoll eregarde dth ecoroll ai nth eusua lClematis flowers a sabsen texcep ti nflowers o f sectionAtragene. Withinsectio nFlammula, h edistinguishe d5 informa lgroup s(i.e .withou tindicatin gnam ean d rank): § 1. many-branched, panicle-like synflorescences, pinnateleaves ; § 2. many-branched, panicle-like synflorescences, ternateleaves ; § 3. 3-flowered cymes,o r terminally solitaryflowers; § 4. solitaryflowers, pinnat eo rternat eleaves ; § 5. solitary flowers, simpleleaves . Heattribute d 85specie st o Clematis, of which 70 were assigned to sectionFlammula. DeCandoll econsidere dth egenu sNaravelia t ob emonotypic .Th eonl yspecie sN. zeylanicawa sbase don Atragene zeylanica L .Th egenu swa sdistinguishe dfro mClematis byhavin gpinnatisec tleave swit hth ebasa lpai ro fleaflet spresen tan dth eothe rleaflet sreduce d totendrils ;th eflowe rha sth epresenc eo fstaminodi ai ncommo nwit hsectio nAtragene. A s therear eals oClematis species ,suc ha sC. afoliata Buchanan ,tha tsho wreductio no fleaflet s totendrils ,th e separate status ofNaravelia DC.ma yb e doubted.
Spach(1839 )propose di nhi sHistoir eNaturell ede sVegetau xa narro wdelimitatio no fth e genusClematis an dbase dthi so nearlie rtreatment so fMoenc h(1794 )an dpartl yReichenbac h (1837),apar tfro mreference s topre-Linnaea nliterature .Withi nth etrib eClematideae ,h e distinguished many genera, which areno w generally regarded tob e sections ofClematis: AtrageneL . (=> sect.Atragene (L. ) DC); Cheiropsis (DC.) Spach (=> sect. Cheiropsis DC); Viticella Moench (=> sect. Viticella (Moench) DC); ViornaRchb . (campanulateflowers ;th egenu sha sbee n based on pre-Linnaean references, a.o. Dillenius (1732)b y Reichenbach, 1837); MeclatisSpac h (thenam ebein ga nanagra mo fClematis, thisgenu s hasbee nestablishe db ySpac hfo r yellow-flowered Clematis species andconsiste d of twospecies ,C. orientalis L. and C. glauca Willd.); Clematis L. (=> sect.Flammula DC.) Within Viorna and Clematis, hedistinguishe d sections: Viorna sect.Euviorna Spach (woody climbers); sect. Viornium Spach (perennials). Clematis sect. Vitalba Spach (woody climbers); sect.Flammula Spac h (suffruticose +climbers) ; sect.Aspidanthera Spac h (woody climbers, anthers with an appendage). Spach'sclassificatio n -an dtherefor eals oth eview so fMoenc han dReichenbac h- ha sno t beengenerall yaccepted ,althoug hhi sgeneri cname spla ya rol ei nmoder nclassificatio no f Clematis at the sectional level (Meclatis,Viticella andViorna).
Loudon(1844,1869 )kep ttw ogener awithi nth etrib eClematideae : ClematisandAtragene; thetrib eitsel fwa sdistinguishe di nth eRanunculacea eb yvalvat eo rinduplicat ebu daestivation , evergreeno rdeciduou sclimbin ghabi tan ddecussat elea fposition .Withi nClematis Loudo n recognized 4 sections: FlammulaDC ; Viticella (Moench) DC; Cheiropsis DC; AnemonifloraLoudo n Anemonifloraha sno tbee nmaintaine db ylate rauthors ,bu twa sreduce dt oth eran ko fserie s (Montanae CK.Schneid.).
Within the tribe Clematideae, Bentham (1862) recognized two genera: Clematisan d Naravelia.Clematis was subdivided intothre e sections: Viticella (Moench)DC . Cheiropsis DC. Flammula DC. According to Bentham (1862), the characters distinguishing the Clematideae from the Anemoneaear eth ebu daestivatio n(valvat evs .imbricate )an dth elea fpositio n(decussat evs . scattered).Bentha m estimated that there were 100specie s withinClematis.
Inth eperio d 1867-1869Baillo npublishe dhi sHistoir ede sPlantes .I nvolum e1 (1867)h e
10 describedman ymorphologica lfeature so fth eflowe rwithi nClematideae .Fo rClematis h e focussed onflowe r bud aestivation, dehiscence of anthers andtransien t forms between androeciuman dperianth .H elargel yenhance dth eclassificatio nb yD eCandoll e(1818) ,bu t addedhi sow nobservation san dreferences .Baillo nsubdivide dClematis int oseve nsections : Atragene(L. )DC . Naravelia(L. )DC ; the section wasregarde d as derived fromAtragene. Cheiropsis DC. Meclatis(Spach )Baill . Viorna (Rchb.) Baill.; syn.Muralta Adans .e xEndl . Viticella (Moench)DC . Flammula DC. Kuntze(1885 )wrot ea monograp ho nth egenu sClematis, i nwhic hh ecombine ddetaile d observationswit ha rathe rpeculia rclassification .Hi sinfrageneri cclassificatio n seemspartiall y tohav ebee nderive d from Linnaeus',althoug hh edi dno tstat ei texplicitly .Hi s informal classification, i.e.withou t stating anyrank , isa s follows: a. Scandentes (woodyplant s or subshrubs climbing with ranking petiolules); 1. Scandenteseperulatae (flowerin gbranche sdevelopin ga tth eyoun ggrowth) ; 2. Scandentesperulatae(mostl yreduced ,flowerin gbranche sdevelopin gfro m buds atth e last yearsgrowth) ; b.(=) 3. Escandentes(non-climbin gperennials ,subshrub so rshrubs ;b=a 3indicatin g that theseplant s areregarde d tob ederive d from the Scandentes). Hetrie dt ointerpre tal lspecie sa sderive dfro mC. vitalba,whic ha sa nexercis ei sworthwhil e doing, but he failed to pinpoint consequences, as isi n essence the intention of modern phylogenetic analysis. Kuntzedefine d speciesa sbroadl ya spossible ,s oh ecombine dman yspecie sknow n atth etime .Althoug hh emad esom erathe rpeculia rcombinations ,base do ninsufficien tmaterial , studyingsom eo fthes emerger si sworthwhile .H ewas ,b ydoin gso ,reall yth efirs tClematis authorwh odeviate dfro mth etraditiona ltypologica ltaxonomi ctreatmen tan dwh odare dt o interpretinfraspecifi c variation.H ewas ,however ,inconsequen ti ndealin gwit hvariatio nb y creatingexhaustiv einfraspecifi c classifications,whic har estrictl yhierarchica li nth esens etha t the broader the infraspecific variation he described the more extensive his hierarchical classification was.Hi streatmen twa sno taccepte db ylate rauthors .Furthermore ,hi simplici t statingo finfraspecifi c rankswa scumbersome .A sa nexample ,hi streatmen to fC . orientalis
11 ispresente d below (ranks interpreted byth epresen t author): C.orientalis L. subsp.a normalis var. l.flava (Moench) Kuntze var. 2.daurica (Pers. ) Kuntze subvar. a.persoonii Kuntze subvar. b.thomsonii Kuntze subvar. c.dyeri (Clarke ) Kuntze var. 3.albida (Klotzsch ) Kuntze subvar. a. obtusifolia Hook.f. &Thomso n subvar.b .massoniana (DC.) Kuntze subvar. c.vulgaris (Trautv. ) Kuntze subvar. d. angustifolia (Ledeb.) Kuntze subvar. e.fasciculata Kuntze subsp.p graveolens (Lindl.) Kuntze var. 1.lindleyana Kuntze var. 2.hookeriana Kuntz e var. 3.aitchisonii Kuntz e var.4 .subtripinnata Kuntze subsp.y thunbergii (Steud.) Kuntze var. 2.lutea (Jacquem. ) Kuntze var. 3.intricata (Bunge) Kuntze var.4 .glabrescens Kuntz e var. 5.pauciflora Kuntze subsp.6 brachiata (Thunb.) Kuntze var. 2.subglabra Kuntz e subsp.e wightiana (Wall.) Kuntze var. 1. typica subvar. a.glaucescens (Fresen.) Kuntze subvar. b.inciso-dentata (Rich.) Kuntze var. 2.longecaudata (Ledeb.) Kuntze var. 3.pseudobuchananiana Kuntze var.4 .hoffmanni Vatkee x Kuntze subsp.(simensis (Fresen.) Kuntze var. 2.brevifoliola Kuntz e var. 3.longifoliola Kuntze Furtherexaminatio no fth eabov eclassificato n ofC. orientalis wil llear ntha ti ti sa mixtur eo f trueC. orientalisan dallie dspecie so nth eon ehan dwit hspecie stha tar emisclassifie d for various reasons onth e other hand (see Chapter2) .
Prantl(1888 )wa son eo fth efirs tauthors ,wh otrie dt oclassif y thegenu sClematis bot hb y
12 consideringal lavailabl emorphologica lcharacters ,an db yinterpretin gth epolarit yo fcharacte r states.Thi s resulted in the following classification: Clematis L. sect.Pseudanemone Prant l ser.Spatulifoliae Prantl ser. Villosae Prantl sect. Viorna (Rchb.) Prantl ser. Crispae Prantl ser. Tubulosae Decne. ser.Atragenae Prant l ser. Cirrhosae Prantl sect. Viticella (Moench) DC.emend , ser.Euviticellae Prantl ser.Floridae Prant l sect.Flammula DC.emend . ser.Rectae Prant l ser. VitalbaePrant l subser.Euvitalbae Prantl subser. Saxicolae Prantl subser.Dioicae Prant l subser. Hexapetalae Prantl ser.Aristatae Prant l ser. Orientates Prantl sect.Naravelia (DC. ) Prantl emend. Theran ko fserie sa scurrentl yapplie dha sbee nderive dfro mPrantl' s'Gruppe' .I nreference s antedatingPrant lhi sname sfo rserie shave ,however ,variedl ybee nconsidere dt ob eeithe r subsectiono rserie snames ,se ee.g .Tamur a(1968a ,1987) .Prant lformulate dth ehypothesi s thatth eorigina lform so fClematis, i.e .thos eform smos tsimila rt oth egenu sAnemone L. , weret ob efoun di nth ePalaeotropics .Th esectio nPseudanemone shoul db emos tsimila rt o thisgrou pan di sindee drestricte dt oth ePalaeotropics ,havin git sdistributio nare ai nAfrica . SectionNaravelia wa sregarde da sa nearl ydivergin gsection .Sectio nFlammula originate d inth ePalaeotropics ,bu tha sdisperse dal love rth eworld ,excep tfo rth epola rregions .Sectio n Viornaoriginate dan dremaine drestricte dt oth eNorther nHemisphere ,wherea sser .Atragene has dispersed to subarctic and mountainous regions and ser. Cirrhosaesettle d in the Mediterranean.Sectio nViticella originate di nCentra lan dEaster nAsia .I tno wals ooccur s inth e Mediterranean area. Ast oth ehighe rclassificatio no fClematis withi nth eRanunculaceae ,Pran dstresse d
13 thatther ear en oparticula rreason swh yther eshoul db ea separat etrib eClematidea eDC . Althoughpredominantl ypresent ,th evalvat eaestivatio no fflowe rbud si sno ta uniqu echaracte r andth estructur eo fth einflorescenc ean dfrui ti ssimila rt oAnemone. Th eonl ydistinguishin g characteri sth edecussat elea fposition .Prant lregarde d Clematisa sbelongin gt oth etrib e Anemoneae. Both Prantl's classification and his phylogenetic hypothesis contain some inconsistencies, but wereth ebas eo f modern Clematisclassification .
Koehne (1893) followed Prantl's classification ofClematis:
sect.Flammula DC. ser. Vitalbae Prantl ser. Orientales Prantl
sect. Viticella (Moench)DC .
sect. Viorna (Rchb.) Prantl subsect. Euviorna Koehne subsect. Connatae Koehne subsect. Escandentes Koehne subsect.Atragene (L. ) Koehne It isremarkabl e that he subdivided the section Flammula in series, whereas he erected subsectionsi nsectio nViorna withou tan yargument .Furthermore ,Atragene wa sreduce dt o theran ko fsubsectio ncontrar yt oth eview ,stil ladopte db ysom ecurren tauthors ,tha ti tshoul d be aseparat e genus.Anyhow , Koehne clearly denoted ranks inhi s classification.
Hooker (1837) showed aplat eo f aspecime n assignedt oa the nnewl yrecognize dgenu s ClematopsisBoje re xHook. ,bu tproduce dn odescription .Hutchinso n(1920 )presente da firstsurve yo ftha tgenus ,separate dfro mClematis mainl yo nit sflowe rbu daestivation ,bein g imbricateinstea do finduplicative . Clematopsisi smor eo rles ssimila rt oPrantl' ssectio n Pseudanemone.Brummit t(1976 )an dRayna l(1978 )presente dth emos trecen tsurve yo f Clematopsis. Tamura (1987) again included Clematopsis inClematis.
Inhi sManua lo fCultivate dTree san dShrub s(197 42 nded .12 thpr. first; publishe di n1923) ,
14 Rehderclassifie d theClematis species ,importan ti ncultivation .H epartl yfollowe dPrant l (1888), Koehne(1893) ,an d Schneider's (1904) Ulustriertes Handbuch der Laubholzkunde: Clematis L. sect.Flammula DC. ser. Vitalbae Prantl ser.Rectae Prant l ser. Orientales Prantl ser.Montanae C.K.Schneid. sect.Atragene (L. )DC . sect. Viticella (Moench)DC . sect. Viorna (Rchb.) Prantl ser. Cirrhosae Prantl ser. Crispae Prantl ser. Connatae (Koehne) Rehder ser. Tubulosae Dene. Tamura (1968a) published an extensive classification of the Ranunculaceae, based on phylogeneticconsiderations ,a sfinal par to fa serie so fpublication so nth ephylogeneti cvalu e ofcharacter swithi nthi sfamily .Withi nth etrib eAnemonea eh edistinguishe dthre esubtribes , including the Clematidinae, asdefine d byLots y (1911): 'Die Clematidinae weichen von alien anderen Gruppen der Anemoneae durch gewohnlichefaltig-klappig eAestivatio nde sKelche sun dgegenstandig eBlatte rab .E s sind4 bi smehrer ekronblattartig e Kelchblatter vorhanden,nu rdi eUntergattun g Atrageneha tlinear eKorollenblatter ,de nander nfehl tdi eKorolle .Di eFruch tis tein e NuB,of t wiebe iPulsatilla mi tauswachsende m federformigen Griffel.' Withinth eClematidinae ,Tamur ainitall ymaintaine d4 genera :Archiclematis Tamura ,Clematis L.,Clematopsis Boje re xHook. ,zndNaravelia DC ,althoug hth eprincipa ldistinguishin g tribalcharacters ,flowe rbu daestivatio nan dlea fposition ,wer en oconstan tcharacte rstates . Tamura classified Clematis into 12section s anddistinguishe d 15subsections : Clematis sect. Viorna (Rchb.) Prantl subsect. Viorna (Rchb.) Tamura subsect. Connatae Koehne subsect. Crispae (Prantl) Tamura sect.Bebaeanthera Edgew. (Paratragene Tamura) sect.Atragene (L. )DC . sect.Meclatis (Spach )Baill . subsect. Orientales (Prantl) C.K.Schneid. subsect. Tanguticae C.K.Schneid.
15 sect. Clematis subsect.Pierotianae (Tamura )Tamur a subsect. VitalbaeTamur a subsect.Dioicae (Prantl ) Tamura subsect.Aristatae (Prantl) Tamura subsect. Papuasicae H.Eichler subsect. Crassifoliae (Tamura) Tamura subsect.Rectae (Prantl ) Tamura subsect.Angustifoliae Tamur a sect. Cheiropsis DC. sect.Lasiantha Tamur a sect. Viticella (Moench)DC . subsect.Floridae (Prantl ) Tamura subsect. Viticella sect.Patentes Tamur a sect.Pterocarpa Tamura sect.Fruticella Tamura sect.Naraveliopsis Hand.-Mazz. In 1987,Tamur a modified hisclassificatio n considerably: Clematis subgen. Campanella Tamura sect. Campanella Tamura sect. Tubulosae (Decne.) Kitag. sect.Bebaeanthera Edgew. sect.Atragene (L. )DC . sect.Meclatis (Spach ) Baill. subsect. Orientales (Prantl) C.K.Schneid. subsect. Tanguticae C.K.Schneid. sect.Pseudanemone Prantl subgen. Viorna (Rchb.) Tamura subgen. Clematis sect. Clematis subsect.Clematis subsect. Pierotianae(Tamura ) Tamura subsect.Dioicae (Prantl ) Tamura sect. Cheiropsis DC. sect.Lasiantha Tamur a sect.Aspidanthera Spac h sect.Naraveliopsis Hand.-Mazz. subgen.Flammula (DC.)Peterm . sect.Flammula DC. subsect.Rectae (Prantl ) Tamura ser.Rectae Prant l
16 ser. Uncinatae Tamura subsect.Angustifoliae Tamur a subsect. Crassifoliae (Tamura) Tamura sect.Pterocarpa Tamura sect. Viticella (Moench)DC . subsect. Floridae(Prantl ) Tamura subsect.Patentes Tamur a sect.Fruticella Tamura Comparedt ohi searlie rclassificatio n in1968 ,Tamur ano wintroduce dth esubgenu srank . SubgenusCampanella i snewl ydefine dan dbase do nth ene wsectio nCampanella. On eo f thesubgenera ,Viorna, i sjus ta nupgrade dsection .Others ,subgenu sClematis an dsubgenu s Flammula,ar ene wcombination so fearlie rdescribe dsections .A sthi sclassificatio n shows someinconsistencie san di smor ecumbersome , theclassificatio no f196 8ha sbee nchose na s starting-point for further examination.
1.3. Phylogenetic analysiso fth egenu sClematis
Inorde rt ojudg eth eintegrit yo fth egenus ,cladisti canalyse shav ebee ncarrie dout ,base do n morphological characterso f subdivisions of Clematisan do frelate dgener a(1.3.1.) ,an d analysiso fth egloba ldistributio nof Clematis(1.3.2.) .I ti spreferabl et ocarr you tthi styp eo f analysiswit hscore so fcharacters .B ythem ,i ti sals opossibl et ojudg eth eintegrit yo fth e subdivisionso fa genus .Th edatabase sbuilt ,however ,wer ebase do nintegra ldat afo rth e subdivisionsthemselve sa sthe ywer edefine db yPrant l(1888 )an dTamur a(1968a) .Thi s restrictsth epossibilit yt odra wdetaile dconclusion sfro mth eanalysis ,bu tgloball yprovide s insighto nvariatio npattern si nth egenus ,bot hwit hregar dt oit sdistributio n(Barton , 1989; Cox,1990;Cracraft , 1975;Humphries , 1979;Humphrie setal. , 1988;Nelso n&Platnick , 1981;Rave n& Axelrod , 1974;Rosen , 1975,1978;Schuster , 1976;Thome , 1978,1983; Wiley,1975,1977,1978,1979a ,1979b ,1979c ,1980,1981 ;Wulff , 1950;Zande e& Roos , 1987)an dt oit soveral lvariatio n(Ax ,1985;Bremer&Wanntorp , 1978,1981;Brooksetal., 1984;Brook s& Wiley ,1985;Eldredge ,1985,1989;Eldredge&Cracraft , 1980;Estabrook , 1972,1978;Fun k& Brooks , 1990;Hennig, 1950,1965,1966;Humphrie s&Funk , 1984; Loconte & Stephenson, 1990, 1991; Minelli, 1993; Nelson, 1978, 1979, Queiros &
17 Donoghue,1990a ,1990b ;Sluiman ,1985 ;Sober ,1975,1983,1984,1988 ;Stevens , 1991; Stuessy, 1990;Wiley , 1981).
1.3.1. Cladistic analysisof subdivisions ofClematis and of related genera
Ast oth edelimitatio no fth egenu sClematis, ther ear eman yunresolve dissues .A numbe ro f taxa, often described as separate genera, have been included in the genus s.l., such as Atragene(Linnaeus , 1753,vs .Miller , 1768),Naravelia (D e Candolle, 1818,vs .Prantl , 1888),Clematopsis (Prantl ,1888 ,vs .Boje re xHooker ,1837/Hutchinson , 1920),Meclatis (Spach,1839 ,vs .al lothe rauthor safte rhim) ,Viorna (Reichenbach ,1837 ,vs .D eCandolle , 1818),Viticella (Moench ,1794 ,vs .D eCandolle ,1818)andArcfa'cfemaft'.s(Tamura , 1968a, 1970vs .e.g .Rehder , 1974)ar eincluded .Whethe ro rno tClematis ca nb econsidere da sa monophyleticgenu si sth emai ncriteriu mt odecid eupo nit sdelimitation .I nth emoder nvie w onbotanica lclassification ,monophyl yi sconsidere dt ob ea stric tconditio nfo rrea lsystemati c entities. AsFarri s (1991; cf. Farris, 1974;Platnick , 1977)pu t it: 'Monophylyca nb edefine d (thoughno trecognized )withou treferenc et ocharacte r evidenceonl ybecaus emonophyleti cgroup shav erea lan dindependen thistorica l existence.Paraphyleti c andpolyphyleti cgroup shav en osuc hexistence ;the yar e nothingbu tth echaracter sb ywhic hthe yar edelimited .Withou t characters,paraphyl y andpolyphyl ymea nnothing .I ti sth erealit yo fmonophyleti cgroup stha tultimatel y distinguishes phylogenetic systematics from syncretistic taxonomies'. Prantl's (1888) andTamura' s(1968a ) classifications havebee n chosenfo r phylogenetic analysis,a sthes eclassification s arerelativel yrecen tan dwidel yaccepted .A dat ase twa s constructedbase do nthei rrespectiv epublications .A sthes epublication sdiffe r inthei rusag e ofdescriptiv echaracters ,th edat aset sals oslightl ydiffer .Table s1. 1an d1. 5surveytheuse d charactersfo rbot hanalyses .Table s1. 2an d1. 6presen tth etw odat amatrices .Th edat aset s wereanalyse db yHennig8 6releas e 1.5 (Farris, 1988),usin gmhenni gan db bfo r branch swapping.Th ecladogram sthu sobtaine dwer esubjecte dt osuccessiv eweightin gusin gth e consistencyinde xo fcharacter sa sweightin gcriteriu m(Carpenter ,1988;Farris ,1969,1989) . Astric tconsensu stree ,usin gFarris 'nelse nalgorithm ,wa sconstructed ,i fsevera lequall y parsimonoustree swer eobtaine dfro m theanalysis .Thi stre egive sa nimpressio n of the
18 informationconten to fth euse ddat ase t(th edegre eo fhomoplasy )an dvisualize suncertai n steps inth e tree (Anderberg &Tehler , 1990;Miyamoto , 1985).
In the case of Prantl (1888), thetrib e Anemoneae (Anemones.l .an d Clematiss.l. )wa s includedi nth edat amatri xt oge ta bette rsigh to nth ebordercase so fth einvolve dgenera .Th e results are presented in figures 1.1, 1.2 and 1.3. As in all produced trees the section Homalocarpusseem st ob eth esiste rgrou po fClematis, th eanalysi sha sals obee ncarrie d outwit hAnemone sectio nHomalocarpus a soutgroup ,th eresult sbein gpresente di nfigur e 1.4.
19 Table1.1 .Characte rse tfo rPrantl' sclassificatio n ofth eAnemoneae ,use di nphylogeneti c analysiswit hHennig86 .
Character character states 1 Habitus perennial 1 subshrub 2 shrub 3 2 Leaf simple 1 composite 2 3 Leaf palmately nerved 1 pinnately nerved 2 4 Leaf scattered 1 alternate 2 decussate 3 5 Leaf irregularly serrate/dentate 1 serrate2 lobed 3 entire 4 6 Leaf pilose 1 nervespilos e underneath 2 glabrous 3 7 Leaf herbaceous 1 pergamentaceous 2 coriaceous 3 8 Leaf tendrils absent 1 present 2 9 Bracts free 1 +fuse d 2 involucral 3 10 Infl. simplecyma e 1 composite cymae2 fasciculate 3 heterotactic 4 solitary fls. 5 11 Infl. flowering onth eyoun g wood 1 flowering onth eol dwoo d 2 12 Flower flat, erect 1 broadly campanulate 2campanulat e 3 tubulose, urceolate 4 13 Flower hermaphrodite 1 unisexual 2 dioecious 3 14 Tepal n=41 n=52 n>6 3 15 Tepal herbaceous 1 coriaceous 2 fleshy 3 16 Tepal ovate 1 rhomboid 2 obovate 3 17 Tepal imbricate 1 valvate 2 18 Tepal villose atmargins ,glabrou s 1ditto ,bu tpilos e inside 2 ditto,bu t pilose outside 3 19 Tepal acute 1 acuminate 2 mucronate 3 20 Staminodia absent 1 incidentally present 2 present 3 present,petaloi d 4 21 Filament filiform 1 dilatate 2 rugulose 3 22 Filament glabrous 1 ciliate 2 23 Anther shorter 1 aslon g as2 longer than filament 3 24 Anther connective not elongate 1 elongate 2 25 Ovary ovate 1 rhomboid 2 26 Ovary ovules 2-4 1 ovules 1 2 27 Fruit ovate 1 rhomboid 2 rhomboid, dorsiventrally costate3 fleshy 4 28 Fruit glabrous 1 pilose 2 villose 3 29 Fruit style not elongate 1 2x elongate 2 >2x elongate 3
20 ft en en CN CN CN CN CO - CN CN CN - - CN - CN CM CN - 8 - en co CM CN - CN CN en en - CN en en en CN CN CM CN CN - CN - 8 CN CN ~ CN CN CN CN CN CN CM CN CN - CN - S CN a CN a CN CN CN CN CN CN CN - - - - CN - CM - CN CM CN -
«N CN CN CN - CN CN CN CN CN CM CN CN CM CN -
o CN
0\ ------~ - ~ - - - - - CN - CN CN - CM - CN - - - oo ------CO cO - - CN CN - - - - ~ CN - ~ - - CN CO en
CN CN CN CM CN CM CN CN CN CN CN CN CN CM CN
CN - co CO CN CM en en
•Q S *Tt CM CN CN CO t ^f CN CN Tf Tf CN CN CN •s OS co CO CO - co co en en en oo 00 CN oo oo *- e _o « o l« en - CM CO en CO -* en en en - - - - «3 - TT co en en en CO CO en CO en en en en en en CO en CO o CM eN CN CN - CM CN CM CN CN A 1 i § 09 Q j. 8 •s a a 4) U. 9 c 1 1 •S to 53 1 * * to Si c > 1 § & s i •s e "5 a S 1 s 1 •1 ••a to S 1 11 1 2 2 S § 1 ! o •s a ss 5 s 3D s. 1 i 1 5 1 1 1 I i1 I 1I 3 1 1 W 65 O S W 03 •P ->H 01 ' ^ ~ T3 01 •H oi o •P (0 (0 -"•* oo 00 00 a _o o a o u «en feb I u b O ai oo Z 0) (0 <—» < <0 fl CU A r-.H 10 X! ai (0 fl ~ (S (0 'S 0) +J A •H 10 -H H 4J 0c (0 \-> H e N>, +>| CO ^ a) co •H •H \> <0.-^( Q) c •H > 4-1 (0 H e IC -. -. +J \<0 •H rH o o •c CO CO 10 ~ 10 e •P ~ CO o 6 3 •H •H s 10 •ri (1) (0 10 O 3 0) (U •P •P -.a •H +J H s ~ H •rl e to 10 ~<0 10 rH +J 10 u 0) (0 *-*r* •P a a* s 10 6 G o 10 6 CO Q) •«~iH B •a CO u co H c •H O (0 > a u c e ••u u S• H O co tx, •H — » > » > -H •• V) •H ••cooco-cotacocococflricocj > co co -H -H in -H a) co ID -H A co co co •• co u 0)w 4JV4d)»-l-Hi-HM3M-H..(BWffl a) M ai COCOlHCOCDCOCOCfl-'SCOCVfc-H c id (0 13 10 fl) 10 CO-'CO—W^cOfl CO CO-r l CO M CO o •P -rH (0 -H £ -H -H M O a a) (0 a IO a) —H co a> CO ^rH C ClOTJJS-H+JMCiOaJ M- -H i-t 01 c (0 fl (0 (0 (0 (0 wQ) co < 10 O 3 ft-H C (0 O U 10 •H rH 10 10 co rH 4J 0) -P H 10 . a O 4J H •P-P U ft 0 * (0 •H 0 +J giococooa^iorH •P « u (0 £ 3 •H CO •H 10-rl r( •P r co -H •c nah-HHigoBH (0 M-H h 0 >-H O X X u O 0 •H > a a h •P •H 3- 3 h •H CD 10 -H 2} H u 3 5 Cfl % o < s Ei w c o !*3 uto X r c^ c> CN m 3 r> c- • - r4 8 r> - cs c c o c> c e- r> c r r c> c^ r r c- r p- iH c- r c- c- c c- c o r r r r c- 61 8 O i (30 ^H 2? 9\ c .. '53 '•d en 90 c^ > .U H c- r- c c> c- o c> c- r c c> r- r r 3 c c^ cs T—1 1«H I"' 1-4 r r c- r> c^ <4-H 1-1 o c« • - K O r r r r i-H o > 0\ c f cn T3 « .^ 00 f~ s ^ on v« o §2?S V) c c c c c c c c T—l C •*r c c c c c c c c c c c c c c c c - 2 o o •S ' • o to c c o o c> c- c r> r c- r r r c- r r .00 oo 3 f^ c c c- r C r r o r r r r c r c r r r c r CS 2 o « V3 X ' , U i-i c c c c c c c c c c c c c c a ro CO CM O u y s o •a a) u 00 en n n c V c o o CO H o •fi I 00 3 PH M C M 3 bft C J•e3 si; u is « > «t/i o« o3 C/3 I* 1) a, aJ 3 t-i O •^ f> r- c- c- c- e- c- (^ rc-or-r-orcs .5 6. N C* EEEEEEE^3EEEEE X S"- 6 (3 ' , 1H c c c c c c c fcccc — c^n B I" TJ- •2o Asi nal lequall yparsimoniou scladogram sAnemone sectio nHomalocarpus adjoin s Clematis,th edat ase twa srestricte dt oClematis sens uPrant lan dsectio nHomalocarpus. Theresultin gcladogra m(character sweighted )i spresente di nfigur e1.4 .Considerin gsectio n Pseudanemonea sa separat egenu s{Clematopsis), onl yth evalvat ebu daestivatio nappear s tob ea synapomorph yfo rClematis an dNaravelia, th elatte rseparate db ylea ftendril san d appendagesa tth eanthers .Th etre elengt ho f16 9step san dth eman ycharacter swit heithe r aconsistenc yinde x0 o rweigh t0(2,3,9,11,12,14,18,19,20,22,23,2 4an d26 )indicat e 28 thatth einformatio nconten to fthi sse ti slo wan dtha tdetaile dconclusion sar eno tpossible .Th e remaining character with a multiple statement at one ancestral node is character 10 (inflorescence structure).Th eevolutionar yhistor yo fClematis ca nonl yb eclarifie d aftera detailed analysis of inflorescence structure andflowe r characteristics. Although the information content of the data set concerned israthe r low, some conclusions mayb e drawn: Section Clematis and its series Vitalbae appear tob epolyphyletic ; Thesam ehold sfo rth esectio n Viorna,althoug hthi scanno tb ea stron gstatemen t regardingth efac tthatAtragene i smostl yclassifie d assection .Th epositio no fth e seriesTubulosae ha st ob efurthe rascertained ,bein gals orelate di nclassification s to sections Clematis andFlammula (seeTamura , 1987); The section Viticellalook sparaphyletic ,i nthi srespec tth e specieso fth e series Crispae havet ob e analysed. Takingint oaccoun tth etreatmen tb yTamur a(1968a) ,th esubtrib eClematidina esens uTamur a (i.e.Clematidina ea sdefine db yLots y(1911) )wa sincluded ,especiall yt oconside rwhethe r the separate generaArchiclematis, Clematopsis andNaravelia ar e interposed between sectionso rsubsection so fth egenu sClematis sens uTamura .Th eresult sar epresente di n figures 1.5,1.6an d 1.7. 29 Table1.5 .Characte rse tforTamura' sclassificatio n ofth eClematidea ewit hAnemone a soutgroup , used in phylogenetic analysis withHennig86 . Character character states 1 Habit perennial 1 subshrub 2 shrub 3 2 Leaf simple 1 composite 2 3 Leaf palmately nerved 1 pinnately nerved 2 4 Leaf scattered 1 alternate 2 decussate 3 5 Leaf irregularly serrate/dentate 1 serrate2 lobed 3 entire4 6 Leaf pilose 1 nerves piloseunderneat h 2 glabrous 3 7 Leaf herbaceous 1 pergamentaceous2 coriaceous 3 8 Leaf tendril absent 1 present 2 9 Bracts free 1 fused 2 10 Infl. simplecyme s 1 composite cymes2 fasciculate 3 heterotactic 4 solitary fls. 5 11 Infl. floweringo nth eyoun gwoo d1 flowering onth eol d woodi 12 Flower flat, erect 1 broadly campanulate 2campanulat e 3 tubulose,urceolat e 4 13 Flower hermaphrodite 1 unisexual 2 dioecious 3 14 Tepal n=41 n=52 n>6 3 15 Tepal herbaceous 1 coriaceous 2 fleshy 3 16 Tepal ovate 1 rhomboid 2 obovate 3 17 Tepal imbricate 1 valvate 2 18 Tepal villose atmargins ,glabrou s 1 ditto,bu tpilos e inside 2 ditto,bu tpilos e outside 3 19 Tepal acute 1 acuminate 2 mucronate 3 20 Staminodia absent 1 occasionally present 2presen t 3 present, petaloid 4 21 Filament filiform 1 dilatate 2 rugulose 3 22 Filament glabrous 1 ciliate 2 23 Anther shorter 1 aslon g as2 longer than filament 3 24 Anther connective not elongate 1 elongate 2 25 Ovary ovate 1 rhomboid 2 26 Ovary ovules 2-4 1 ovules 1 2 27 Fruit ovate 1 rhomboid 2 rhomboid, dorsiventrally costate 3 fleshy 4 28 Fruit style notelongat e 1 2x elongate 2 >2x elongate 3 30 S - - - - CS CS CS CS en CS CN en - CS CS CS CS - - - - CS CS CS CS - - fi - - - - 8 - - - - CS - CS CS - - - CS CS CS CS CS CS CS - CS CS CS CS CS CS CS (N a CS CS CN a 3 - - CS - CS CS CS CS CS CS CS CS CS CS CS CS f5 - - CN - CS CS CS CN CS CS CS CS CS CS CS CS CS CS CS - CS CN e o\ CS CS CS CS - - - - CS CS CS CS CS CS CS - 90 - - - en CS CS - - - CS CS - CS ------CS CS ~ - - - - - t~ - CN - cs CS CN CS CS CS CS CS CS CS CS CS CS CS CN CN CS CS CS CS CS CS CS » en CS CS CS CS en CS CS *-* CS CS CS CS - - - - CO en CS CS CS - CS t^ CS CS CS - CS - - - C3 OO O cs CN CS CS - - - CO en CS r^ - - - - cs CS CS o « - - - - CS CS CS - CS - - CS - - - CS en - ~ - - - CS CS - - - 3 •fl - - - - en en en - CS ------^t •* ^f - - en en -* - CS - ^ - CS en en en CO en en on m m en en m en en en o CI en en en en en en en en u m - cs CS CS cs CS CS CS - CS CS CS CS CS CS CS CS CN CS CS CS CS CS CS CS CS CS en en CS CS CS en en m m m en en en en en en en en en en en en en •fi - - - - - a B 09 Q it •3 K 3 * 4» 1 1 I 1 4) a i j CD ft s I 1 , 11 9 •Si 1 | 2 S e I e 1 •s I | .1 1 » 1 5 i ! i S 1 I a 1I * 1 I 3 1 •4. oil 1/ —> -H 0) ^-01* •H 01 -H -H +J -H 4-> 4-> -eHn (C 4-> U] g ^^ n3 HI JJ 6 10 •H ON 10 iH •H (C a e -P <-* C -» IT u c IH a m ol >- * c 01 01 fa01 H 6 n c h S3 rH (.)C J ai •• -H H 0 » u H R > 0 -H •• •• —c •• o 0 -H > c c c o c •H •• o o •• o c w •P-* c o o •• c +> 4-1 0 +> o ai •H c o ~ o o 01 o in 4-> O -H C 01 01 w a) -P xi O-H +> O 01 01 st tn o 3 0) -P o -H,Q Si 3 Si 0) 01 If) U III •P 3 3 01 3 01 " 01 01 O 01 01 -3 Hi — 0) A 3 01 XI 0) w •w. 01 U *— 3 01 01 A 3 01 is *" o o: 01 re ^3ll-« 01 -H 43 re 01 ~C 01 -^ 01 ID .s? 01 0J •P 0 re m 01 — 4»H rj '53 u. •§ B o Ml -a > OINONH NIMHriNNN HHHHHPilM o 14 14 3 U. U LJ u OH CM n 00 vo Ml M ON L oe m i J u 00 PI 2 u a o c o CM (A u ca L in o to •3 J3 CO H M01 l C I O W i-l in O n. •• 1 -S, CM 00 CM CM CM CM co CM CM CM CM CM CM CM CM CM - CM •5 —' 5 ' CM S CM CM CM CM CM CM CM CM CM - CM CM CM CM CM CM CM CM CM - - - - '. CN c3 AC S X X X X X X X X X X X X X X X X X X X X X X X X 8 CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM - - - 3 6 a =3 2. J3 ... O CM u , <£ 00 O ... CM CM ~ CM CM CM - CM CM CM CM CM CM •a 0 0 1 CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM - - - - o •a ... .'PJ SP 1 § - CM CM ------~ - CM CM CM CM CM CM CM CM - - - - 00 CM CN ' CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM - HI o en en o en en CM CM CM CM 3 CM CM U ^H l-~ a 2 •QS3 7^ £ 1 *-H cej ^ 2 en CM o e ^f ^1- en CM CM CM -^ 1 - - - t •* - •* ~ - - - - - CM CM -tf c* » c x O T3 00 X X X X X X X X ON c3 X X X X X X X X X X X X X X X X X o 00 t~ CM 0 2 '•3 CN T) C CM CM CM CM CM CM CM » - CM CM - S3 f . C3 c3 w u^ 1 ^ tfl co - rr en 2 •S T. 0 "? CN O t co en CO en en en en en CO en en CO en CO CO CO CO CO CO CO CO en en *-t CM en en CM en en en en CO en en CO en CO CO CO CO CO CO CO CO >0 S3 - - - ^ ,1- CN S fN = S § ' 1 O CO u 13 0 CN 2 2 00 N 0\ c 8 8 O 3 <*> s 8 CO § 3 3 9 3 3 $ 9 bp , •S> (N.. cej O en CN >> 1 pa - 00 E Fromth edat amatri xo fTamura' sclassification ,thre ecladogram sar epresente d(figure s 1.5, 1.6 and 1.7). Although the first one (characters unweighted) seems to make clear, that Clematisi swel lseparate dfro mth eothe rgener aAnemone, Archiclematis, Clematopsis an d Naravelia,th esecon dcladogra mmake sclea rtha tsuc ha conclusio ni sno tjustifie db yth e data,a tleas ta sfa ras Archiclematis an dNaravelia ar econcerned .Wit ha tre elengt ho f219 , aconsistenc yinde xo f7 2an da retentio ninde xo f83 ,ther ei sa lo to fincongruenc ewithi nth e dataset .Tabl e1.7 .support sthi sshowin gth echaracte rstate so fhypothetica lancestor swit h manymultipl estatement sa tcertai ncharacter s(10,12,15,16,2 4an d27 ;1 5an d2 4exclude d fromth ecladogra mb yweigh t0) .Characte r 12(gros sflowe rmorphology )i nparticula ri sa dissolvingcharacter ,an dsimilarl ycharacte r1 6(tepa lmorphology) .Lookin ga tcharacters ,th e datase twoul db egreatl yimprove db yanalysin gth eClematis inflorescenc e structurepe r species(Tobe ,1979,1980d ;Troll ,1964,1969 ;Weberling ,1981) .Thu sfar ,onl yfragmente d informationi spresent .Th esam ehold sfo rothe rflowe rcharacters ,especiall ythos econcernin g morphology,textur ean dvenatio no ftepals ,an dnecta rleaves .A st ovegetativ echaracters ,th e growthhabi tviz .th egrowt hmode lshoul db eanalyse di nmor edetai lpe rspecies ,a sth e relationbetwee nherbaceous ,suffruticos e andwood yspecie si sno tsolved ,an dneithe ri sth e presenceo rabsenc eo fth eclimbin gcapacit y(Baillon , 1867;Bell, 1974;Decamps ,1975 , 1976,1979;Hall eetal. , 1978;Jeannoda-Robinson , 1977;Sterckx,1897;Tomlinson ,1984) . However,thi scladogra mreveal stha tonl yClematopsis ma yb emaintaine da sa separat e genus,an dtha ti tseem slogica lt oinclud eArchiclematis an dNaravelia i nClematis. Pinnatel y nervedleave san dvalvate/induplicat ebu daestivatio nar ethe nsynapomorphie sfo rClematis, inchidingArchiclematisan dNaravelia. Th edecussat elea fpositio ni sa synapomorphy ,share d byClematis an dClematopsis, andth epinnatel ynerve dlea fi sa nautapomorph yfo rbot h Clematis and Clematopsis. The decussate leaf position istherefor e the only remaining charactert odistinguis hth etrib eClematidea eaccordin gt othi sdat aset ,but ,especiall ywit h respectt oNaravelia, decision swhethe ro rno tt oinclud ethes egener ai nClematis hav et o waitunti l further databecom e available. Bearingi nmin dth emodes tinformatio n contento fth edat aset ,som econclusion s concerning classification canb e drawn: Thesectio nClematis i spolyphyletic ;th econclusio nagree swit hth eon efro mth e analysis of Prantl's classification. 36 Thesectio n Viticellai spolyphyletic ;th econclusio ndisagree swit hth eresult sb y analysingPrantl' sclassificatio nprovide dtha tPrantl' sclassificatio ni srathe rparaphyleti c thanpolyphyletic . Subdividingth esectio nMeclatis int oth esubsection sOrientates an dTanguticae i s dubious,a sth edistinguishin gcharacter s(inflorescenc estructur e[10 ]an dgros s flower morphology[ 12] )caus ea multipl estatemen ti ncharacte rstate sfo rthei rhypothetica l ancestor (seefurthe r Chapter2) . Thesectio nViorna seem st ob emonophyletic ,bu tthi sdisagree swit hth eanalysi so f Prantl's classification. MaintainingArchiclematis an dNaravelia a sseparat egener ai sdubious ;inclusio ni n Clematis seems tob emor e appropriate. Althoughgroup so fconvenienc ear eals opresen ti nTamura' sclassification ,th enumbe ro f monophyleticgroup sincrease d(figur e1.7) .Th esectio nClematis turne dou tt ob epolyphyleti c hereals oan dth esectio nViticella i st ob eregarde deithe ra sparaphyleti co ra spolyphyletic . Thecharacter snumbe ro ftepals ,flowerin g onth eyoun go rol dwoo dan dth epresenc eo f staminodiahav ebee nuse dt odetermin eth erelativ epositio no f serieswithi nth esectio n Viticella.Fro mobservation si nth ecultivate dassortmen to fClematis cultivar si ti sobviou stha t precisely these characters are very weak and not as decisive as Tamura supposed. Furthermore,a sspecie sdelimitatio nwithi nth esectio nViticella i sonl ygradua l(Brandenbur g andva nd eVooren ,1986,1988a) ,ther ear estron greason sno tt omaintai na ninfrasectional , supraspecific classification within this section. Asi nTamura' sclassificatio nth esectio nViorna ha sbee nrestricte di nit sdelimitatio n compared to Prantl's classification (Cirrhosaean d Atragene are considered separate sections), Viorna sensu Tamura is amonophyleti c group. Theonl yremainin gpolyphyleti csectio ni ssectio nClematis whic hindee dshow sman y divergenttrait s(perennial svs .shrubs ;se xpolymorphisms ;variou sflowe ran dfrui tcharacters) . Asi ti scircumscribe di nliteratur eu pt oan dincludin gTamura ,thi ssectio nha st ob eregarde d asa 'res tgroup'i nal lclassifications .Th esubsection so fClematis sect .Clematis i nTamura' s system (1968a) consist of restricted sets of affiliated species. As such they should be considereda srea lentitie sa sresul to fth eevolutionar yhistor yo fClematis an dtherefor eequall y ranked:the yhav et ob eregarde da stru esections .Th esubsection so fth esectio nMeclatis ar e clearlysiste rgroups ,a sdistinguishe db yTamura ,bu tlookin ga tth especie sleve l(fo rmor e detailsChapte r2 )th edistinguishin gcharacter sappea rt ob egraduall ychangin gove rthes e subsections.Sinc esimila rreason sappl yt oth esectio nViticella no tt omaintai nsubsections , 37 theywil lno tb emaintaine di nth esectio nMeclatis either .Fo rth etim ebeing ,th esubsection s ofViorna ar epreliminar ymaintained ,sinc ethe yhav eno tbee nsubjecte dt odetaile dstud yi n this investigation. Theresultin g sectional classification of Clematis is: Clematis L. sect.Angustifoliae (Prantl ) Brandenburg comb.nov . sect.Aristatae (Prantl) Brandenburg comb.nov . sect.Atragene (L. )DC . sect.Bebaeanthera Edgew. (Paratragene Tamura) sect. Cheiropsis DC. sect. Clematis (ser . Vitalbae Prantl) sect. Crassifoliae (Tamura) Brandenburg comb.nov . sect.Dioicae (Prantl ) Brandenburg comb.nov . sect.Fruticella (Tamura) Brandenburg comb.nov . sect.Lasiantha (Tamura ) Brandenburg comb.nov . sect.Meclatis (Spach ) Baill. sect.Papuasicae (H.Eichler) Brandenburg comb.nov . sect.Naraveliopsis (Hand.-Mazz.)Brandenbur g comb.nov . sect.Pierotianae (Tamura ) Brandenburg comb.nov . sect.Pterocarpa (Tamura ) Brandenburg comb.nov . sect.Rectae (Prantl ) Brandenburg comb.nov . sect. Viorna (Rchb.) Prantl subsect.Viorna subsect. Connatae Koehne subsect. Crispae (Prantl) Tamura sect. Viticella (Moench)DC . Thisclassificatio n hasbee nuse dfo r theanalysi si nsectio n 1.3.2o nth ebiogeograph yo f Clematis. 1.3.2. Biogeography ofClematis Thelarg ediversit ywithi nClematis ha sbee ninterprete dvariously :Th enumbe ro fspecie st o beaccepte di slargel ydependen to nth einterpretatio no fdistributio npattern swithi nth egenus . Accordingt ocurren tview so nth eevolutio no fflowerin gplants ,Ranunculea nplant sar e consideredbya.o.Stebbins(1950,1974)andTakhtajan(1969,1980,1991)t oconstitut eth e moreprimitiv eplan tgroup samon gth eAngiospermae .Thei rdistributio ni scosmopolita nexcep t forpola rregions ;the yoccu ri nal lclimati cregions ,althoug hmos tabundantl yi nth etemperat e 38 zoneo fth eNorther nHemisphere ;the yoccu rbot hi nmontan eregion san dplains ,i nwoodland s andope nvegetation .I nsom echaracters ,suc ha swoo danatomy ,lea fvenatio nan dflowe r structure,the yexpres scharacte rstat ecombinations ,tha treflec tthei rorigi na tth edivergenc e betweenMonocotyledon san dDicotyledon s(Dahlgre nan dBremer ,1985) .The yals osho w alarg edifferentatio no fflowe rtypes ,althoug hth ebasi cphenomeno no fhemicycl yi stypica l throughoutth efamil y(Brouland , 1935;Eyde ,1975;Leppik , 1964;Meicenheimer, 1978; Tobe,1976a ,1976b ,1980a ,1980b ,1980c ,1980d) .Ther ei ssom epalaeobotanica levidenc e forth eabove .Krassilo ve tal .(1983 )foun d inSiberi ai nAlbia ndeposit sfossi lremain so f bisexualflower swhic hreflec tth egenera lflowe rstructur eo fRanunculacea eo rPaeoniacea e (flattenedreceptacle ,wit h3- 5follicle san dremain so fbot hperiant han dandroeciu m(Batten , 1984). Clematisshow sman yprimitiv efeature si ncompariso nt oothe rgener ao fth efamily : thebasi ctyp eo fflowe rstructur e(withou tspecialize dnecta rleaves ;Prantl ,1888) ,th ebasi c typeo fmetacentri can dsubmetacentri cchromosome s(Gregory ,1941) .I tals oshow ssom e advancedcharacters ,expresse di nth emajorit yo fit sspecies :decussat elea fpositio n(Haccius , 1942)an dth ewood yperformanc eo fstem san dbranche so fth emajorit yo fit srepresentatives , this state is considered to be derived by comparison with the stem anatomy of related herbaceous plants (Smith, 1928; Sterckx, 1897;Tepfer , 1960). Duet oit srelativel ylon gevolutionar yhistory ,Clematis show sparticula rdistributio n patternsa st osections ,whic hhav et ob eunderstoo di nterm so fth eorigi nan dmovemen to f continents,an di nterm so fdispersa lmechanisms .Geographi cdistributio no frepresentative s of the sectionsAtragene, Meclatis, Viorna and Viticella arepresente d in figure 1.8. Thesectio nAtragene i slargel yconfine d toth emontan eregion so fNorther nan d CentralEurope ,Norther nan dCentra lAsi aan dNort hAmerica .Specie so fth esectio noccur s inman ydisjunc tareas .Representative sfro mNort hAmeric aresembl eth eEuropea none st o such an extent that uniting them pairwise into one species has to be considered, the distinguishingcharacter sno tbein gconstan ti nexpression .Thi si si nlin ewit hth emovement so f continentsi nth epas t(Eurasi aan dNort hAmeric ahavin gonc ebee nfused) ,a sca nb esee ni n figure 1.9. 39 Legend Atragene Viticella Vioma Meclatis Figure1.8 .Geographi cdistributio ntrend si nClematis classicatio nfo r4 sections .Furthe r explanationi ntext . Thesectio nMeclatis i sdisperse dfro mTurke yt oKore aalon gmountai nchain s and inth emontan eregio no fth eHimalaya san dTibe t(fo rdetail sse efigur e2.1) .Representative s ofthi ssectio nar epresen ti nsom eadjacen t areas(Himalaya ,Kashmir ,Tibet) ,bu tals oi n disjunctarea s(Korea) .Thi sfac tan dthei rmutua lgenetica laffinit ycombine dwit hdifference s betweenpopulations ,tha tar eonl yminor ,mak eplausibl etha tther ei sa restricte dnumbe ro f variable speciesan d not 10o rmor e(cf . 1.4 and Chapter2) . Thesectio nViorna i slargel yconfine dt oth eUS Aan dMexic oapar tfro monl ya fe w taxai nEurop ean dth eFa rEast .Th eoccurrenc eo frepresentative so fthi ssectio ni nth eeaster n andwester nextreme so fEurasi amus thav eforme don eare awit hth eViorna distributio ni n NorthAmerica ,becaus eth eEurasia ncounterpart shav ethei rparalle lspecie si nNort hAmeric a (e.g. C.integrifolia L.vs . C.ochroleuca Aiton;C. fusca Turcz. vs.C. pitcheriTorr .& A.Graya.o.) . 40 The section Viticella has itsrepresentative s intw o separateareas : Mediterranean area (C.viticella L. subsp.viticella), inclusiv e of thePortugues e Atlanticregio n(C .viticella subsp.campaniflora (Brot .Fon tQue re xO .Bolo san d Vigo); Eastern China / Japan (C.florida Murray ex Thunb.; C.patens C.Morren and Decne.). Althoughinterspecifi c hybridizationbetwee nthes especie sha sgive nris et oman ylarge - flowered cultivars,i tdoe sno tsee mt ooccu r spontaneouslyan di shampere db yvariou s isolatingmechanism s(se e1 A) .I nbot hregion sspecie so fthi ssectio nhav eoverlappin gareas , butbecaus eo f differingflowerin gtime san da differentia l ecologicalpreferenc ethe yappea r tob e wellisolated . This sectioni srestricte d toEurasi a andha sn ocounterpar t inNort h America,o rw eshoul dfin dsuc ha counterpar tamon grepresentative so fth esectio nViorna. Iti sworthwhil elookin ga tth edistributio n ofth evariou ssubdivision so fth egenu sove r continentso rpart so fcontinents .Th eresult so fthi sar epresente di ntabl e1.9 ,an dfigure s 1.10, 1.11an d1.12 .Fo rthi ssurve yth eclassificatio no fTamur a(1968a )wa smodified :th esectio n Patentes,an dth esubsection sFloridae an dViticella wer ecombine dint osectio nViticella, andth esubsection sOrientates an dTanguticae wer ecombine dint oth esectio nMeclatis. Th e rationale of these decisions hasbee n discussed in 1.3.1. 41 Eurasia India South America! North America Africa Australia Antarctica Laurasia Gondwanaland Pangaea Initial division ofsupercontinen t intotw o subcontinents Further division intocurren t continents Fusionbetwee ncurren t continents Figure 1.9. Schematicalpresentatio n of continental shifts in thepast . 42 •a 2 1 •o 8 1^ s § 8 z " 'S 3 o § < z Si a 5 -a § •a 1 •a 1? "Si.11. 1 •s o -o w c s « « a w t ^ >, « « s » « rl 5 I 1 I S 3 5 C i< C afa 6 fl. J O W S U SB 1 -Si •a I £ I G g e ISJasI s i •8 0) c _cg "SIi x I* o 0) T3 C o '5b c3 X> O 5 d 3 60 vio con cri beb atr ori pie vit dio an pap era rec angch e las vii pte fru nar arc cle naa Clematis sections and subsections and related genera. Figure 1.11.Distributio n of taxa over regions. Forexplanatio n of taxon acronyms, seetabl e 1.8. 20 ° 10 eur me ca him ind chi afr sib jap ich ina anz nca sa Global regions. Figure 1.12. Distribution of taxaove rregion s2 . For acronymso f regions, seetabl e 1.8. 45 Abundancei nvariatio ni sindicativ efo rth eorigi no ftaxa ;tw oattitude sconflictin gi npar thav e been summarized byWile y (1981): Theregio ni nwhic hth emos tabundan tvariatio ni spresen ti sth elikel ycentr eo forigi n ofth etaxo nconcerned ;th emos tprimitiv erepresentative sar et ob efoun di nothe r marginalarea s(Croizat , 1962,1964;Croizatetal,1974 ;Nelso n&Platnick ,1981) ; Theregio ni nwhic hth emos tabundan tvariatio ni spresen ti sth ecentr eo forigi nan d primitiveform so fth etaxo nconcerne dhav ebee nmaintaine dthere ,wherea smor e derivedform shav edisperse dt oothe rareas ;thi si sth eessenc eo fwha ti sformulate d byHenni g (1966) asth eprogressio n rule(Cracraft , 1975). Tose ewhethe rcurren tdistributio no fClematis divulge smor einformatio n onorigins ,th e distributiondat ao ftabl e1. 8wer esubjecte dt ofurthe r analysisusin gHennig86 .Th eanalysi s wasperforme dtwice :on ewit hth etax aan dthei rdistributio nove rgloba lregions ;on ewit h global regions and the abundance of occurring taxa. The first approach facilitates the comparisonwit hcladogram so fsectio n 1.3.1.,especiall ythos eo ffigur e 1.5,1.6an d 1.7. Therear efou requall yparsimoniou stree safte rsuccessiv eweightin g•withArchiclematis o rwit h Clematopsisa soutgroups ,respectively .I nbot hcase sth estric tconsensu stree sshowin gth e questionmark so fth eresult sb ymultifurcat efork sar epresente di nfigur e 1.13.Th eoutgroup s were chosen as inspired by Tamura (1970), whopostulate dArchiclematis alternata as ancestorfo rClematis an da sgleane dfro mHutchinso n(1920) ,wh oconsidere dClematopsis aprimitiv egenu so fth etrib eClematidea ebecaus ei tshare scharacter swit hboth Anemone (budaestivation )an dClematis (lea fposition) .Consequently ,th eanalysi so fdistributio ndat a per global region has been carried out with Laurasia and Gondwanaland as outgroups respectively;moreove rther ei son eadde dwit hPangae aa soutgrou pwit ha postulate dzer o distribution.Pangae awit ha postulate don edistributio nha sals obee ntested .Accordin gt o Nelsonan dPlatnic k(1981) ,th eon edistributio nstart sfro mth eassumptio ntha tClematis i s alreadypresen ti nit sancestral ,undifferentiate d form.Bein gundifferentiate d andal lcurren t subdivisionso fth egenu sar ederive dfro mth eancestra lform ,i tha sbee nscore don eove ral l currentsubdivision so fth egenu sa st ob epresent .Th ezer odistributio ndoe sno tconside r derivationfro mth eancestra lfor man dstate sth ecurren tsubdivision so fClematis a sbein g absent in Pangaea. The one distribution results in a less parsimonious cladogram after successive weighting and isno t presented. Aftersuccessiv eweighting ,i nal lthre ecase sa singl emos tparsimoniou scladogra mresulte d from theanalysis . 46 01 W 0) •H Minl CM 0> tM 6 o in e o s <4-H n O rt CS -a a o a x> •Rc/3 T3 A) CO Ul J3 •H 10 •H 0) •P M to CO 10 a) an 10 (0 0) (0 -H Figure1.14 .Are acladogra mo farea san dtax aliste di ntabl e1.10 .afte r successiveweightin gwit h Gondwanaland as outgroup;weight s and states of distributional nodes listed in table 1.9. 48 to a I §« O 0 O 0 O - O 0 0 0 0 0 0 c 53 41 O 0 O 0 O O O 0 0 0 0 0 0 - £ « - - O 0 O O O 0 0 0 0 0 0 c o> o 3 i - 0 O 0 O - O 0 0 0 0 0 0 c 1 - - O 0 O O O 0 0 0 0 0 0 c «.• - 0 O 0 O O O 0 0 0 0 0 0 c •P- - 0 O 0 O O O 0 0 0 0 0 0 c j£ 09 O 0 O 0 O O O 0 0 0 0 0 0 c 6 I O O 0 0 0 0 0 0 - at I'*' : O 0 O - - O - 0 0 0 0 0 0 c O 2:.:. ! - - - - - O - 0 - 0 - 0 - - •s s « ~ 0 O 0 O ~ O - 0 0 0 0 0 c T3 O 0 O 0 O O O 0 0 0 0 0 0 c 1) M O 0 O 0 O O O 0 0 0 0 0 0 c T3 8- a '•5 o O 0 O 0 O O O 0 0 0 0 0 0 c V ------O - 0 - 0 - - - - •8 o - 0 O 0 O O O 0 0 0 0 0 0 c c i a _o s~ O - 0 0 0 0 0 0 c 3 - 0 - O 0 - 0 0 0 0 c x> « 3 - - •fi i - - - - O O O 0 0 0 0 0 0 c O 0 O 0 O O O 0 0 0 0 0 0 c 3 S a - ~ - - O O O 0 0 0 0 0 0 c "J O - - ~ - - O - 0 - 0 - 0 0 c I U J- .C SO OS •8 V-4 t-4 8 a 3 3 3 s s 8 ? =5 chi f=29= =14l a =3 him =1 me ^C =2 ca L26 ' =0 eur =25 =7 sib C =12nc a W =8 jap 1L2 2 =15g o =6 afr =21> =4 ind £19 1 13s a -C 18J 11an z t 17=! ij=9ic h l==16===10in a Figure 1.15.Are acladogra mo farea san dtax aliste di ntabl e 1.8.afte r successiveweightin gwit h Laurasia asoutgroup ;weight s and states of distributional nodes listed in table 1.10. =13 sa H=301 =16 pg =11 anz 29 =9 ich • 0 =10 ina =4 ind 1==2& =15g o 25 =6 afr r =8 jap =12nc a C =7 sib L22J =0 eur UJ -2 ca lUoJ =1 me iLiJ =3 him Q chi UvC la Figure1.16 .Are acladogra mo farea san dtax aliste di ntable .1.8 .afte r successiveweightin gwit h Pangaea asoutgroup ;weight s and states of distributional nodes listed in table 1.11. 50 I bo ON C s CO 60 §« - o © o © © o o © o © © © c > o o o © - © © CT© © © © © c r- u o o © © o © o o © © © © 6' (J n«• - - u • 3 1* - o © o o © © © © © © o o - vo i o o © o o © © © © © © © 1 - - in I o o © © © © o o © o o o © - •> "" o o © o © o © © © o © o © - © P2 «e o o © © © © © © © © © © © c CO o o © © - o © © o Sb 1 i O § en o o © © © © © o - - - o © c •a -* co "o t o o © o •S g a - - © © © © © © © © o © © - a o o © © o © o o © o © © o c iS en •a 2. a-! 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Thebes tpresentatio ni stherefor edependen to nfurthe rdetails .Fro mfigur e 1.11,i tappear s thatsom esection sar ewidespread ,wherea sother sar eendemi can dconfine dt oa restricte d area.Th eregio no fmos tabundan tvariatio ni sn odoub tChina ,bu tthi svariatio ndoe sno t includeman yendemics ,whil ebot hprimitiv ean dadvance dpart so fth egenu s(an drelate d genera)ar erepresente dher e(Tamura ,1970) .O nth eothe rhand ,man yendemi csubdivisions , sharingremarkabl echaracters ,suc ha sdioecy ,occu ri nAfrica ,Sout hAmerica ,Indochin aan d Australia.Wit hClematopsis distribute di nAfric aan dth eviewpoin to fPrant l(1888 )tha t Clematisi ssuppose dt ohav eoriginate di nth epalaeotropics ,thi si si nfavou ro fa developmen t froma norigi ni nGondwanaland .Th eare acladogra mo ffigur e1.1 4i si nagreemen twit hthi s hypothesis,a si sbu tt oa lesse rexten ttha to ffigur e 1.16,wherea sth eare acladogra mo ffigur e 1.15.i sno tfull yi nagreemen twit hth ehypothesi so fTamur a(1970) ,Archiclematis bein gth e ancestoro f Clematis withit s distribution inChin aan dIndochina . Asth eanalysi sha sbee ncarrie dou ta tth eleve lo f subdivisionso f Clematiswit h considerationo frelate dgenera ,bu twithou tconsideratio no fth enumbe ro fspecie sinvolve d persubdivision ,n ofir mconclusio nca nb edrawn ,excep ttha tcombinatio no fcontinenta l movements(se efigur e 1.9.;Krutzsch ,1989 )an dcurren tdistributio npattern so fClematis subdivisionsan drelate dgener apoint si nth edirectio no fa Gondwanalan dorigin .Thi sagree s withClematopsis a sprogenitor ,it sAfrica ndistributio nan dth eproportio no fendemi centitie s stilli nGondwani ccontinents .Th eMalaysia nconnectio ni sstil la lin kt oth eothe rhypothesis . Onth eothe rhan dwhe nth eide ao fcentre so forigi ni sno tadhere dto ,i ti spossibl et oindicat e areaso fendemism :Africa ,Australia ,an dChin aan dNort hAmerica ,whic hi smor eo rles si n agreementwit hth epalaeobotani cfinding s of Krassilove t al.(1983) . Furtheranalysi sbase do nbot hspecie sdescription san ddistributio ndat ai sneede dt o reacha conclusio nan dt otes tth etw ohypothese ssummarize db yWile y(1981) .Chin ai sth e areao fmos tabundan tvariatio nlookin ga tth enumbe ro fpresen tsubdivision so fClematis. FollowingCroiza t(1962,1964) ,Croiza te tal .(1974 )an dNelso n& Platnic k(1981) ,thi sstil l doesno texclud eClematopsis fro mbein gth epostulate dancesto ro fClematis. I ncas eon e adherest oth eprogressio nrul e(Crisc i& Stuessy ,1980;Hennig ,1966 ;Cracraft , 1975),Chin a shouldb eth ecentr eo forigin ,whil ether ei sa proble mi nfindin ga mos tprimitiv eform ,a s Archiclematis occurs outsidethi sarea . 53 1.4. Interspecific crossesi n Clematis Fromhorticultura lliteratur ei ti sgenerall yknow ntha tmos to fth ecurren tClematis assortmen t originatedfro mman yinterspecifi ccrosse so fwhic hth emos timportan tone sha dalread ybee n madei nth e19 thcentur y(Moor e&Jackman , 1872).Sinc ethes ecrosse swer ecarrie dou tb y chance,hardl yan yinsigh twa sgaine di nth egeneti cbreedin gpotentia lo fClematis species .Th e hybridizationexperiments ,reporte dhere ,wer ecarrie dou tt orevea lpotentia l interspecific relationships and toevaluat e hybridization data inth eligh t of systematic research. 1.4.1.Introduction tothe experiment Tocharacteriz erelationship sbetwee nClematis species ,a dialle lhybridizatio nschem ewa s carriedout .Specie swer eselecte deithe rb ythei ravailabilit yi nth eWageninge nClematis collectiono rb ythei rimportanc efo rClematis cultivation .See dse tand ,i nsom ecases ,polle n tubegrowt hwer escored .I na restricte dnumbe ro fcrosse sals oth eprogen ypopulation swer e observed.Th ecomple xgerminatio nbehaviou ro fmos tClematis specie spreclude d further extensiono fthes eexperiments .Th egerminatio nbehaviou ra spresen ti nmos tClematis specie s hasbee ndescribe di nliteratur e(Barton ,1967;Blair ,1959;Kinzel ,1913;Niethammer , 1928). Adoubl esee ddormanc ymechanism ,wit htemperatur ean dligh ta ssuggeste d important factors, prevents seed germination for aperio d of one ortw oyears : themorphologicall ymatur esee dcontain sa nimmatur eembry oan di ssurrounde db y animpermeabl e fruit wall; afterth efirs twinte rseason ,th efrui twal lha sbee nweathere dt osuc ha nexten ttha ti t becomespermeabl ean dth eembry ostart st odevelo pi nth efollowin gseason ,bu ti s notcapabl et ogerminate ; aftera secon dwinter ,seed sgerminate ,bu tsometime stw osuc hcycle sar erequired . 1.4.2. Materialand methods Bydesignin gth edialle linterspecifi chybridizatio nexperiment ,Rehder' s(1974 )classificatio n wastake n asa starting-point . Thisreferenc e isa goo drepresentatio n ofth ehorticultura l taxonomictreatmen to ftax aimportan ta sornamenta lan dmainl yfollow sth eclassificatio no f 54 Prantl (1888): Section SerieSeriess Species used Viorna Rchb. Crispae Prantl C.integrifolia L. TubulosaeDecne . C.heracleifolia DC. Connatae Koehne Cirrhosae Prantl C.cirrhosa L. Atragene (L.) DC. C.alpina (L. )Mill . C. macropetala Ledeb. Viticella (Moench)DC . C.campaniflora Brot. C.viticella L. C. patens C.Morrene t Decne. FlammulaDC . MontanaeC.K.Schneid . C.montana Buch.-Ham . exDC . Rectae Prantl C.recta L. Vitalbae Prantl C.vitalba L. Orientales Prantl C.orientalis L . C.glauca Willd . C.serratifolia Rehder C.tangutica (Maxim. ) Korsh. Weuse dpopulation sfro mnatura lprovenance so rS rpopulations,excep tfo rC. macropetala and C.patens. During the experiment, populations from natural provenances of C. macropetalahav eno tbee navailabl et ous ,an do fC. patens w eha da nofte nreproduced ,bu t originallynatura lpopulatio n at ourdisposal . Crosseswer emad edurin gth eperio d197 8-198 3i nth egreenhous et oavoi dcontaminatio n by adverse weather conditions and to prevent uncontrolled cross-pollination. From 1 Septembert o1 Ma yther ewa sadditiona llightin gwit hincandescen tligh tt oensur ea neffectiv e daylengtho f1 4hrs .Unde rthes econdition splant scontinue dt ogro wan dflowere ddurin gth e periodMarc h/ April ,enablin gcrosse sbetwee nearl ywinter -an dspring-flowerin gspecie san d summer-flowering species.I norde rt overif ypolle nqualit yunde rthes econditions ,polle n sampleswer echecke dfo rvitalit yb yi nvitr ogerminatin gi nva nTieghe mcells ,i na hangin gdro p of 12.5%saccharose ,10 %Brewbake rmediu man d0.2 %aga rsolution .Polle nsample swit h 55 agerminatio npercentag eles stha n50 %wer eexclude dfrom th ehybridizatio nprogramme . Pollensample smaintaine dmor etha n50 %germinatio npercentag eb ystorag ea t5 ° Ci na vacuumexsiccato rfo ru pt o10 5days ,whic happeare dt ob esufficien t toovercom evariou s gapsi nflowerin gperiod .Crosse si nth eperio dfro m 1981-1983wer emad efo rthi sdialle l hybridizationscheme ;fro m 1978unti l198 1onl yincidenta lcrosse swer emad et ooptimis eth e techniqueo fartificia lhybridizatio ni nClematis, an dt overif ycertai nhybridization smad eo r supposed by Clematis nurseries inth epas tan dno t dealt with here further. Flowerswer eemasculate da tth ematur ebu dstage ,jus tbefor eth etepal sstretc han dsprea d out,o ri nth ecas eo ftubula ro rurceolat eflowers ,whe nth ecoroll aopen sa tth eapex .Th e stagefo rremova lo fth eanther sappeare dt ob ecrucial ,a searl yremova ldamage sth estigma s andrender sthe mnon-functional .Whe nth etepal sar ealread yspreading ,th eoute rstamen s alreadydehisc ethei rpollen .Durin gth espreadin gtepa lstag eemasculatio ni sconvenient ,bu t useo fth eflower si sno tadvisabl efo rcrossin gdespit eprotandry .Self-compatibilit yt ovariou s degreesan dth ehemicycli cnatur emake sselfin glikel ya ttha tstage .Pollinatio n followed immediately throughout theday . Inprofusel yflowerin gaccessions ,crosse swer ecarrie dou to nsevera lflower so fth e sameplant .See dse twa saverage dpe rflower .Som eflower swer euse dt oanalys eth epolle n tubegrowth . In selected cases,seed s were sown to testviabilit y and to observeprogenies . Pollentub egrowt hwa sobserve db yepifluorescenc emicroscopy .Th epreparation swer emad e accordingt oth emetho db yKh oan dBae r (1968)wit h some modifications: Collectflower s3- 7day safte rpollination ,dependin go nweathe rcondition s(u pt o1 0 days after pollination good slides canb eprepared) . Removalo fpistil sfro mth ereceptacle ;pistil swer esoake di na Her rsolutio nt oclea r tissues.I ncas eo fClematis pistil spellucidit yi sabsolutel ynecessar ybecaus eo fth e pubescence of styles and ovaries.Compositio n of theHer r (1971) solution: lactic acid 85% 2part s chloric hydrate 2part s phenole 2part s xylol 1 part clove oil 2part s Rinse 2x inethano l orxylo l and lx in demineralized water. 56 Macerate pistils in INNaO H at60° C during lV2-3hrsa t 60°C. Rinsewit hdemineralize dwater ,transfe rt oanilin-blu estai nsolutio nan dincubat edurin g 20-24hrs .Compositio n of the stain solution: 7gK3P04.3H20 I gAnilin-blu e (Merck, Anilinblau W.S.C .1 ,nr . 42755) II demineralized water Squashpistil sslightl yi nglycero lan dmoun tslide swit hCanada-balsam ;stor eslide s dark andcool . Observationswer emad ewit ha Zeis sepifluorescenc emicroscop ewit hth eexcitatio n/ barrie r filtercombinatio n02 .Photograph swer emad ewit ha M6 3Camer asystem .Polle ntub egrowt h is shownb yyello w fluorescence of callose along pollen tubes. The number of seeds obtained after hybridization was counted. Of each interspecific combinationa selectio no fcrosse swa sthe npu tt ogerminate .A ssee ddormanc yi nClematis maylas ttw oyears ,th eseed swer esow ni nsee dtrays .I fther ewa sn ogerminatio na tonce , thesetray swer elef t alonefo rhal fa yea ri na coo l(5-10 cC)an ddar kplace ,the nth etray s wereplace da troo mtemperatur e2 0° C fo rtw omonths .Whe nther ewa sstil ln ogerminatio n some of the seeds weretake n out to examine whether there was an embryo and if sot o determineth edevelopmenta lstag eo fth eembryo .I fth echeck swer epositive ,th etray swit h theremainin gseed swer elef tcoo lagai nfo rhal fa year .Afte rth esecon dperiod ,th eseed s should germinate,whic h somedi dt ovaryin gdegrees .F,' s wereobserve dt ojudg e their performance. 1.4.3. Resultsand discussion Thissectio ndeal swit hth eresult so fth ehybridizatio nexperiment si nthre eparagraphs .Th efirs t paragraphi sdealin gwit hsee dse tan deventua loffsprin go fth ehybridizatio nexperiments ;th e secondparagrap hwit hth epollen-pisti lrelationship si nth einterspecifi chybridizations ;an dth e third paragraph consists of adiscussio n of both earlier sections. 57 1.4.3.1. Seed set and offspring of diallcl and other crosses Withinth eframewor ko fth edialle lhybridizatio nscheme ,interspecifi ccrosse swer eexamine d for seed setcharacteristics ,result s are showni ntabl e 1.12 and figures 1.17,1.18an d 1.19. 58 co CN CN >n f«N CN in CN ~W "7^ CO V m o o m o o O CN o 00 O o CO CO CN CO in CO in CN CN in O O o o o CO O r^ CN CN CN CN CN CN CN CO o O O o O ON o o ' o - CN c 00 co CN co •n CN m CN -* CN CN >n co CN CN CO m CN •*t CN CN CO CO O o cn o o o CN O © o o c CN CN in o o o 00 o in CN o o o CO © in 00 (N ON cn co CO CO CN in 00 CO CO CO CN CN O O CO OO o «n CO CN CO CN 1^ O <* ^ CN 00 ON m CN en CN CN CO 00 o o O cN o O o CO in O O o ' o o CN O <* CN yo CO in CO CN m en O t*- o o 00 CN m o o «n o CN O CO CN in CN 'o (U OH 13 CO •* CN CN CN CO v» O "* o o o O O O o • OS o s CN T3 B CO in CN T JO 00 o o o o CN o in o o m c in 00 >n CN CN m CN CO CN CN CN CO fO CO o o m O O O o O in OO o CN T3 i "5 1 3 1 "S 1 a I f 11 Of a. I E i I 0) i i s i u u o V) 00 iH 1-4 Fromtabl e1.12 asurve yo fsee dse tb ypollinatio npe rspecie swa scalculate d(figur e1.17) , and seed set on mother plants (figure 1.18),an dcorrecte d for thenumbe r of crossespe r combination infigur e 1.20 and table 1.13. Table 1.13. Interspecific crosses of Clematis species.Femal e andmal econtributio n per species in seeds/cross. Column number • 1 2 3 4 ' Clematis species 1. C. tanzutica 25.4 19.3 13.6 4.5 2. C. orientatis 10.9 10.1 26.2 30.4 12.9 69 16.3 9.3 4. C. serratifolia 27.7 16.0 24.7 8.1 5. C. vitalba 4.0 4.2 5.3 5.5 6. C. montana 3.5 0.4 3.4 0.4 7.C. recta 2.0 1.6 4.1 4.0 8. C. camwniflora 2.0 1.8 2.4 2.2 9. C. viticella 1.5 1.4 4.5 5.1 10. C. patens 7.4 4.0 6.1 3.9 11. C. alpina 0.1 0.1 0.7 0.7 12. C. pitcheri 3.5 2.3 2.8 1.7 13. C. heracleifolia 13.3 11.9 8.3 7.7 14. C. integrifoUa 3.7 3.7 5.4 5.6 1 15 C rirrhma 3 5 4.0 50 56 Legend of column numbers: 1 Female contribution within andbetwee n species 2 Female contribution between species 3 Male contribution within andbetwee n species 4 Male contribution between species Thefirs ttw ocolumn spe rspecie sar et ob ecompare dwit hfigur e 1.17;th elas ttw owit hthos e infigur e 1.18.Therelativel ylarg enumbe ro fseed spe rcros sfor C tangutica,C. orientalis, C.vernayi an d C. serratifoliaimplie s more genetic affinity between these species than betweenon eo fthe man dan yo fth eother so fth ediallel .O nth eothe rhan dth enearl yzer o successi ncrosse swit hC. alpina doe sno timpl ytha tthi sspecie si scompletel yisolated ,a sfo r bothth efemal ean dmal econtributio nther ei sn odifferenc ebetween "withi nan dbetween "an d 60 "between"species .On ema yexpec ttha t"withi nan dbetween "a tleas tequal s"between "specie s forbot hth efemal ean dmal econtribution .Thi seffec t wasgenera lexcep tfo ri nC. orientalis. This speciesha da bette rmal eperformanc e for "between" thanfo r "withinan dbetween " species.Thi sma yb ea nindicatio nfo ra nincompatibilit ymechanis mi nthi sspecies ,s oi n198 6 extrapollination swer ecarrie dout .O nth eothe rhand ,C. montana ha salread yshow nt ohav e aself-incompatibilit ymechanis m(Brandenburg ,1984a) ,bu tapparentl yals oa relativel ypoo r capacity for cross-fertilization in comparison with other species involved. Arelativ econtributio nfemale/mal epe rspecie swa scalculate dt ojudg eth ebarrie rvs . penetrationcapacit yo feac hspecie s(figur e 1.19;Hogenboom , 1973).I nabalance dsituatio n (i.e. barrier capacity = penetration capacity) the quotient equals 1. The quotient of C. tangutkai sremarkabl yhig han dattracte dfurthe rinvestigation .I ti scause db yrelativel ylarg e amountso fseed sproduce dfro mcrosse swit hC. vitalba,C. patens, C. heracleifolia an dC. cirrhosaa spolle n parent. A sampleo f the obtained Fj-seeds were grown and seedlings appearedt ob eexac tcopie so fth einitia lmothe rparent ,wherea sth eF 2-progeniesbehave dlik e aS jo fth emotherplan t (thisS ,wa ssow nalongsid eth eF 2o fth einterspecifi c cross),thu s indicatingtha tn ohybridizatio nha dtake nplace .Thes eresults ,excep tfo rC. vitalba,wer e checkedan dconfirme d byepifluorescenc e ofpolle ntube so nC. tanguticastigmas .Thi s phenomenon, adventive embryony,i sno t observed in otherinterspecifi c combinations. Interspecific combinationswit hC. heracleifolia,C. integrifoliaan dC. cirrhosaa smothe r ledt osee dset ,bu tseed swer enon-viable .Seeds ,take nfro m seedpopulation stha tdi dno t germinatea tall ,wer edissecte dan di tturne dou ttha tth eembry owa slacking .Whethe ro rno t therewa sa naborte dembry ocoul dno tb everifie danymore .Therefore ,i tcanno tb econclude d fromthes eexperiment stha tfrui tdevelopmen tcoul db eparthenocarpous .B ypollinatin glarge - flowered Clematiscultivar swit hnearl ynon-viabl epolle ni nisolatio nbags ,an dth eresultin g emptyachenes ,i ti sknow ntha tparthenocarpou sfrui tdevelopmen tdoe soccu ri nClematis. However,interspecifi chybrid so fC. heracleifoliaan dC. integrifoliaar ereporte dunde rth e nameC xjouinianaC.K.Schneid .Hybri dprogen yfro mC. heracleifolia an dC. vitalba ha s beenreported ,o fwhic hth ecultiva rC .'Mrs .Rober tBrydon 'i sstil li ncultivatio nan dindee d showsintermediat echaracteristics .Mainl ybase do ncrosse sbetwee nC. integrifoliaan dC. viticella, severalcultivar shav ebee nraise d from the 19thcentur yonward ;fo rinstanc eC. 61 was alreadyreported in 1852an d is still in cultivation. In order to establish nomenclatural stabilityaroun dthi sgrou po fcultivars ,th eClematis Diversifolia Groupwa scircumscribe da s cultivargrou p(Brandenbur g andva nd eVooren ,1988b) . Fromthes e experiments, a cross polygon has been constructed (figure 1.21).Thi s showstha tmos tspecie sinvolve di nth ehybridizatio nprogramm ear egeneticall yisolated . The roughsee dse tdat ahenc eca neasil yb emisleadin gb yth ephenomeno no ffruit developmen tdu e toparthenocarp y oradventiv eembryon y(a si sobserve d forC. tangutica). Seedse t 1,400 - 1,,„ ) Withinan dbetwee nspecie s 1,200 ^1 Betweenspecie s 1,000 (-] 800 600 200 • ill ^llM..!. 1. .Ill tan ori ver ser vita mart rec cam viti pat alp pit her int cir Clematis Figure 1.17. Interspecific crosseso fClematis species:see dse tb ypollinatio npe rspecies ;fo r abbreviations,se efigure 1.21 . 62 Seedse t 1,600 I I Within and between species ^1 Between species _ flu VM Hi l .rkrifin ser vita man rec cam viti pat alp pit her int cir Clematis Figure 1.18. Interspecific crosses of Clematis species: seed set on mother plants; for abbreviations,se efigure 1.21 . Log female / male contribution 10 tan oh ver ser vita mon rec cam viti pat alp pit her int cir Clematis Figure 1.19. Interspecific crosseso f Clematisspecies :relativ econtributio n female/male per species;fo r abbreviations,se efigure 1.21 . 63 Seeds/cross tan ori ver ser vita mon rec cam viti pat alp pit her int cir Clematis Femalecontributio n seeds/cross Femalecontributio n seeds/cross • Within andbetwee n species Between species Malecontributio n seeds/cross Malecontributio nseeds/cros s • Within andbetwee n species Between species Figure 1.20. Interspecific crosseso f Clematis species:femal e andmal econtributio npe r speciesi n seedspe rcross ;fo r abbreviations, seefigure 1.21 . 64 1. c.tengutfca 1. ten 2. c. orientalls 2. ori a c. Vernayl ReciprocalcrosscrahavsmsusiK toi hybri d »WK1 a ver 4. c. serratifolia 4. «0f 5. a vitalba 5. vita 6. c. mOfrtSLfWl UMatMtcras sItaspraAictdtMcf , 6. mon 7. a reCta. butn ogans*A»o n occurs. 7. rec 8. a campantftom —~- 8. cam a ytuCmm pj prated sesdsafte radvsnisv sembryony . 9. viti a a 10. pat 10. a 11. alp 11. a pitched UnllsBBraojc&sha spfcstjc»<4vi40i 8seed . 12. pit 12, a ia h&r ia a i/rfeg/ffofa 14. int a c/frr»osa 15. dr 14. 15. Figure 1.21.Interspecifi c crossesi nClematis: cross polygon ofinvolve d species. 65 1.4.3.2. Pollentub e growthexperiment s Hogenboom(1973 )introduce dth eter mincongruit yi norde rt oapproac hmechanism so f interpopulationaldivergence .Althoug hinterpopulationa lrelation sar esubjec tt obiosystematic s research,an dhybri dinviabilit yi so finteres tt oevolutionar yresearc h(Coyne ,1974) ,u pt ono w theter mincongruit yha shardl ybee nuse di nbiosystematics .Accordin gt oHogenboo m(1975) , at leasttw omechanism sfo r non-functioning inpistil-polle nrelationship soccur : incompatibility:preventin go rdisturbin gth efunctionin go fth erelationships ,althoug hth e potentialfo r functioning of bothpolle n andpisti li s complete; incongruity:non-functionin gdu et oincompletenes so fth erelationship ;geni esystem so f bothpartner sd ono t fully fit together. Ina nincongruen tcombinatio nth epenetratio ncapacit yo fth epolle ni sto orestricte dcompare d withbarrie rcapacit yo fth epistil ,fo rwhateve rreason .Incongruit yi scoheren twit hevolutionar y divergencebetwee npopulations ,wherea sincompatibilit yi sa resul to fa positiv eselectio n pressurewithi npopulations ,favourin ggene stha tpreven tself-fertilization .Nowadays ,breedin g progammeshav ebee ndevelope dwhic hmak eus eo fincongruit yi norde rt oobtai nF jhybrid s (Hogenboom, 1979b). Althoughth econcep to fincongruit yha sprove dt ob euseful ,ther ear estil lcontroversie s withrespec tt oth enee dfo rdistinctio nbetwee nincompatibilit yan dincongruit y(Hermse nan d Sawicka, 1979;Hogenboom , 1979a;Pandey , 1979).Workin gwit hNicotiana L. ,Pande y (1979)state dtha tS-gen epolymorphis mha sbee ndevelope db yevolutionar yprocesses ,thu s leadingi nth efirs tplac et ointerspecifi cincompatibility ,an di nth esecon dplac et oinfraspecifi c incompatibility.Comparin gHogenboom' sconcep to fincongruit ywit hPandey' shypothesis , Hermsenan dSawick a(1979 )cam et oth econclusio ntha tinterspecifi crelationship sar emor e simplyan dmor ewidel yexplaine db ymean so fincongruity ,tha nb ymean so fcomple xS-gen e polymorphism.Althoug hsom eeffect slik epollen-tub egrowt hinhibitio nca nb ecause db ybot h incompatibilitya swel la sincongruity ,man yphenomen a(pisti llengt hvs .maximu mpolle ntub e growth,o rpolle nunabl et ostic ko nalie nstigm alobes )ma yonl yb eexplaine db yth econcep t ofincongruit y(Hogenboom , 1983b).Th e sameconcep ti sapplicabl et orelate dfield s of intimaterelationship s (Hogenboom, 1983a). 66 Withinth eframewor ko fth edialle lhybridizatio nscheme ,interspecifi ccrosse swer eexamine d byepifluorescenc e microscopy,th eresult so fwhic har et ob efoun di ntabl e 1.14.Onl yfo r somecombination spolle ntub egrowt hwa sobserve db yU Vmicroscopy .Thi swa sdu ebot h toth erestricte damoun to favailabl eflower san dt olabou rintensit yo fthi swork .Fo rth esectio n Meclatis, table 1.15show s theresult s and observations onresultin g F!plants . 67 VI w* y Q , ^ , , , , , , , u , , S EC O Q , , , , , W PL, , , , % , , fi *H y o- , , , , , , , , , « Q ^ a B M * S3 y C Q u y , , , , , , , Q Q Q § 2 fH l-N , , Q Q , o , , , , , , , , , o 1 § , < , o o , w c- , , , , w o o 'o ©\ 0) o5 o o X 00 e w SB w <3 S r- * * o o Q 8 w o PQ u. a O # « y o , , Q , , CO , , , , , 8 , B fO o tu X o feb 1 « "o (^_ t_ *m * y * § CO Q o § 1 Q pa A u •b 3 1 2 3 4 1.C. tangutica B D E D a a a a 2. C. oriental™ D D C/D G ab b be b 3. C. vernayi C/G* D H F/G c be c cd 4. C. serratifolia A/G* C/D G E b (d) d Legend A Pollentub egrowt hinhibitio no nstigm apapilla esurface ;n ogerminatio no f pollen; B Pollengerminate sincompletely ;i fgerminatin gpolle ntub egrowt hi sinhibite d in stigmao r style; C Pollen tubegrowt h isinhibite d in stigma; D Pollen tubegrowt h isinhibite d inth euppe r half of the style; E Pollen tubegrowt h isinhibite d in thebasa l half of the style; F Pollen tube growth isinhibite d inth eape x of the ovary; G Pollentub egrowt hi sno treachin gth eeg gb yconfuse d growthi nth eovar y ending up in akno t of pollen tubes; G* Variable reactions among whichreactio n G; H Pollen tubegrowt h isnorma l andleadin g to fertilization. ? Results ambiguous. a Plants are similar to C. tangutica; b Plants are similar to C. orientalis; c Plants are similar to C. vernayi; d Plants are similar toC. serratifolia; (d) - Juston eplan t similar toC. serratifolia; ab Plantsar eintermediat ebetwee nC. tanguticaan dC. orientalis;etc .fo rb e anded. Table1.1 5show stha tdespit eo fth epresenc eo fvariou sform so fpolle ntub einhibition ,ther e wasstil lconsiderabl esee dset ,i nal lcase sproducin gseedlin gpopulations .Therefore , in additiont oth edialle lhybridizatio nprogramme ,i n198 6extr acrosse sbetwee nspecie so fth e 69 sectionMecte wwer emad et oobtai na detaile dcharacterizatio no finfrasectiona lhybridizatio n behaviour,an dt ofind ou twhethe rther ear edifference s betweenprovenance swithi nspecies . Results arepresente d intabl e 1.16. Table1.16 .Epifluorescenc emicroscop yo fpolle ntub egrowt hi ninfrasectiona l crosses 1986 inth e sectionMeclatis (Spach ) Baill. and characterization ofresultin g seedling populations. 1 2 3 4 1.C. tangutica B/H G/H E/G/H G a a/ab c 2. C. orientalis G/H A/G/H G/H G/H ab b be 3. C. vernayi G/H G/H H G/H c be c 4. C. serratifolia B/C G/H B/C H Legend: seetabl e 1.15. Different codes indicate different accessions. Comparison of tables 1.15 and 1.16show s that with regard to pollen tube growth the accessionsuse di n198 6wer egenerall ymor eproductiv ei ncombinations .Furthermore ,a s fertilization wassuccesfu li nthos ecombination swit hC. tanguticaa sfemal eparent ,an dth e offspring isquit esimila rt oC. tangutica,C. tanguticapossesse smor edominan tallele si n relationt oth eothe rspecie so fth esectio nMeclatis, althoug hadventiv eembryon ycanno tb e excludedcompletel y(F 2-populationshav eno tbee nscreene di nal lcases) .C. serratifolia appearedt ob emos tisolate dfro mth eothe rMeclatis species .Wit hbot hC. tanguticaan dC. vernayia spolle nparent sfertilizatio n didno ttak eplace ,wherea sbetwee nth ethre eothe r speciesfertilizatio n wasoccasiona l andreciproca l (figure 1.23).S odifference s between accessions areimportan t in selecting parentsfo r plantbreedin g programmes. Apartfro mth edia lle ischeme ,som eothe rinterspecifi c crosseswer emad ebetwee n putativerelate dspecies ,suc ha sC. chrysocomaFranch .wit hC. montana an dC. ispahanica 70 Boiss.wit hC. orientalis.Result so fpolle ntub egrowt hi ncombination sbetwee nC. montana and C. chrysocoma arepresente d in table 1.17. Table1.17 .Epifluorescenc emicroscop yo fpolle ntub egrowt hi ncrosse swithi nan dbetwee n the speciesC. montanaan dC. chrysocoma. cf • 1 2 1.C. montana B/C/G*/H G/H 2.C. chrysocoma C/D/G* B/C/G*/H Legend: seetabl e 1.15. Different codes indicate different accessions. Theresult swit hC. montana an dC. chrysocoma sho wtha tself-incompatibilit yan dincongruit y reactions maycoincid e inth e samecomple x ofrelate d species.Bot hC. montana andC . chrysocoma showvariabl epolle ntub egrowt hbehaviour .I nselling sth epolle ntub ewa s inhibitedi nth estigm a(figur e1.22) ;i ninfraspecifi c crossesth ephenomen avar yfro minhibitio n inth estigm at ocomplet efertilization .Reciproca lcrosse sbetwee nbot hspecie sar ereciprocall y different: C.montana a smothe rplan tshow sfertilization ,wherea sth ereciproca lcombinatio n shows allpattern s of pollentub e inhibition. The results of crosses between C. ispahanicaan d species of the section Meclatis are presented intabl e 1.18. 71 Table1.18 .Epifluorescenc emicroscop yo fpolle ntub egrowt hwit hC. ispahaniea a sreceptiv e parent and various species of the sectionMeclatis aspolle n parent. cr" • 1 2 3 4 l.C B/H D/G* G/H D/G* ispahaniea Legend pollen parent: 1. C. ispahaniea 2. C. tangutica 3. C. orientalis 4. C. vernayi Legend: seetabl e 1.15. Different codesindicat e different accessions. C. ispahanieai sa poorl yknow nspecies ,whic hi sregularl yconfuse dwit hC. orientalis,bein g distinguished byit s suffruticose habit. In two combinations, but not with C. orientalis, fertilizationwa sobserved ;resultin gseed sfaile dt ogerminate .Th eembryos ,presen ti nth eseed s when sown, apparently aborted. 72 Figure1.22 .Polle ntub egrowt hinhibitio na sself-incompatibilit yreaction .C. montana self - pollinated. Thepolle n tubes areinhibite d in the stigma. 73 74 75 76 Figure 1.23. Variouspattern s ofpolle n tubegrowt h inrelatio n toincongruit y assee nb y epifluorescence microscopy. Interspecific crosses: a. C.montana X C. chrysocoma:vigorou spolle ntub egrowth ,bu tpolle ntub ebecome s uncoordinated in growth when approaching the ovary; b. C.montana X C. chrysocoma: same as a.; c. C. montanaX C. chrysocoma: pollentub egrowt hinhibite dwhe napproachin gth e ovary; d. C.chrysocoma X C. montana: pollen tubegrowt h normal,leadin gt o fertilization; e. C. viticella X C.integrifolia: pollen tube growth normal,leadin g to fertilization; f. C.integrifolia X C.viticella: pollen tube growth inhibited in the style; g. C. viticella X C.campaniflora: pollen tube growth normal, leading to fertilization; h. C.campaniflora XC. viticella: pollen tube growth normal, leading to fertilization; i. C. orientalisX C. tibetana subsp.vernayi: pollentub egrowt hnormal ,leadin gt o fertilization; j. C. tibetana subsp.vernayi XC. orientalis: pollentub egrowt hnormal ,leadin gt o fertilization; k. C.tibetana subsp . vernayiX C. tibetana subsp. tangutica: pollen tube growth normal, leading to fertilization; 1. C. tibetanasubsp . tanguticaX C. tibetana subsp. vernayi:polle n tube growth normal, leading tofertilizatio n . 77 1.4.3.3. Discussion and conclusions BetweenClematis specie sa nincongruit ysyste mi soperatin ga swel la sa nincompatibilit y system within species, as already shown earlier (Brandenburg, 1984a). According to Brewbaker(1957) ,polle ntub egrowt hinhibitio na tth esurfac eo fo rjus tinsid eth estigm ai sa n indicationfo rth egametophyti csyste mo ftrinucleat especies .Eas t(1940 )classifie d several Ranunculeanspecie si nthi scategory .Ver ycomple xself-incompatibilit y systemsar ereporte d inRanunculus acris L .b yLundquis te tal .(1973) .Fro mth eno tver ypronounce dsyste mo f self-incompatibility inRanunculacea ewit htrend spresen tbot hi nMonocotyledone s and Dicotyledones, De Nettancourt (1977) concluded that there must have been acommo n ancestralsyste mo fself-incompatibilit yfo rth eAngiosperma ebefor eth ediversificatio nint o Monocotyledones andDicotyledones . Withrespec tt ointerspecifi chybridizatio nvariou sphenomen ao fimpede dpenetratio n havebee n observed: pollenno tgerminatin go nth estigm asurfac ean di fsom egrain sgerminat eth etube sar e inhibited in stigma orstyle ; inhibition ofpolle n tubesjus t below the stigma; inhibition of pollen tubes atvariou splace s in the style; pollentube sformin ga kin do fhaustoriu mi nth eneighbourhoo do fth eeg gbu tno ti nit . Thesephenomen aar eno tfull ysimila ri nreciproca lcombinations ,a sca nb elearne dfro mtabl e 1.14.I ntabl e1.1 5progen ydat aan depifluorescenc e datao ftabl e1.1 4wer ecombine d for Meclatis.The yd ono tagre ewit heac hother .Th eadditiona lcrosse si n198 6sho wtha tther e isvariatio nbetwee naccession so fdifferen t provenances:th edisagreemen tbetwee ndat ao f table1.1 5find sit sexplanatio ni nth evariatio nbetwee npopulation so fdifferen t provenances ast othei rcrossability .Simila rresult sar eknow nfro mothe rplan tgroups ,suc ha sbetwee n Brassicaoleracea L.an d related species (Snogerup, 1980),Cucumis meloL. , Cucumis sativusL .an drelate d species (Khoe tal. , 1980; Kroone tal. , 1980; Ramachandran etal. , 1983),an d Lycopersiconesculentum Mill ,an drelate dspecie s(Rick , 1950,1995;Ric ke t al.,1979) .Thi scrossabilit ybehaviou ri ncombinatio nwit hdistributio narea smake snecessar y further thought on species concepts ast o Clematis. 78 1.4.4. Speciation andspecies conceptsin Clematis Inorde rt ostud ybiosystemati crelationship sbetwee n species,tw oaspect so fresearc har e important: to gather genetic information oninterspecifi c barriers; to obtain ecological information on the adaptive abilities of the taxa concerned. Thecombinatio no fthes etw otype so finformation ,leadin gt oa genecologica lapproac ho f species,ha sprove nt ob ea goo dstarting-poin ti nbotanica lclassificatio n (Anderson,1949 ; Briggs& Walters ,1984 ;Camp , 1951;Cam p& Gilly ,1943 ;Clausen , 1951;Clause ne tal. , 1940, 1947, 1948; De Wet, 1981;Grant , 1971;Hogenboom , 1973;Pickersgill , 1981; Stebbins, 1950;Valentine , 1975). Species definitions are subject tomuc h discussion intaxonomy . Wagner (1984) presented arathe r cumbersome butpractica l definition: 'Speciesi sa convenien ttaxonomi ccategor ytha tdefine sa uni to forganismi cdiversit yi na give n timefram ean dcompose do findividua lorganism stha tresembl eon eanothe ri nal lo rmos to f theirstructura lan dfunctiona lcharacters ,tha treproduc etru eb yan ymeans ,sexua lo rasexual , andconstitut e adistinc t phylogenetic linetha tdiffer s consistently andpersistentl y from populationso fothe rspecie si ngap si ncharacte rstat ecombination sincludin ggeographical , ecological,morphological ,anatomical ,cytological ,chemical ,an dgenetic ,th echaracte rstate s ofnumbe ran dkin dordinaril yuse dfo rspecie sdiscriminatio ni nth esam ean drelate dgenera , andi fpartiall yo rwholl ysympatri can dcoexisten twit hrelate dspecie si nth esam ehabitats , unablet o cross or, if able to cross, ablet o maintain the special distinctions.' Thespecie si nthi ssens ei sa multidimensiona lentity ,tha tca nb einterprete di nthre emai nways : a genetical unit, which forms an adaptive complex; anecologica lunit ,i nwhic hgeneti cinformatio nca nfreel yb einterchange dwithi nth e framework of acertai n response to an environment; anevolutionar yuni t(an ds oo fa certai ndescen tt ob eregarde da sa nindependen t lineage) in time and space, which forms an adaptive complex in which genetic information can freely be interchanged. Atfirs t sight,th ethre eapproache s seemt ob esimilar ,bu tthe yquit ediffe r inthei rwa yo f analysis. The genetical approach, which islargel ybase d onth e interpretation of genomic differences (behaviour,structur ean dnumbe ro fchromosomes )make sal lothe rdiagnosti c charactersuse di nplan tsystematic sdependen to nsuc hdifference s andlead st ocomplicate d 79 classification schemes,a sca nb esee nfro mtreatment si ngras ssystematic s(Dewey ,1984 ; Estes&Tyrl, 1982;Love ,1982 ;Raven ,1975;Sakamura , 1918;Tateoka, 1960;Wenzel & Hemleben, 1982). Ecologicalstudie shav ereveale dsystemati cvariatio nrelate dwit habioti can dbioti c factorsi nth eecosystem .Th eresult so fthes ear emeaningful ,i fcombine dwit hdat aconcernin g thereproductiv e strategy of theplant s concerned and their genetic background. Inth eevolutionar yapproach ,th edirectio no fevolutio no rth edevelopmen to fth e variationo fa certai nplan tgrou pha st ob ederive dfro mdat aset so fcurren tplan tcollections . Usingmorphologica ldata ,extracte dfro mliterature ,thi swa sdon efo rClematis an ddeal twit h in 1.3.1. In this approach the response of the genetic variation in a population to the environmenti simplicitl ydeal twit hb ytake nint oaccoun tth edistributio ndat ao fth etax a concerned.I nthi srespec thybridizatio nha scause dman yproblem si nphylogeneti cdat asets , asth eOTU' swer esuppose dt ob eindependen tlineages .Fun k(1985 )an dWagne r(1980 , 1983) adressedth eproblem , but did not solveit . Ina nattemp tt osolv eth elac ko fconsistenc yi nspecies 'definitions , Kornet(1993 ) establishedth econcep to fcomposit especie sstartin gfro mthre emai napproache s ofspecies : themorphologica l species -dependen t onversio n of approach -,base do neithe r morphologicalsimilaritie so rshare duniqu echaracte rstate s(Adanson , 1763;Nixo n andWheeler , 1990;Wagner , 1984); thebiologica lspecies ,base do ninterbreedin gabilit yo findividual san d- dependen to n versiono fapproach-eithe rpotentia lo rrea l(Dobzhansky , 1935;Mayr, 1940,1976, 1978, 1982); theinternoda lspecies ,base do ncommo nmembershi po fth egenealogica lnetwor k betweentw opermanen tsplittin gevent so ra splittin geven tan da nextinctio neven t (Hennig, 1966;Kornet , 1993;Nixo n andWheeler , 1990). The composite species concept byKorne t (1993) is defined as follows: NA composit especie si sth ese to fal lorganism sbelongin gt oa noriginato rinternodon ,an dal l organisms belonging to anyo f its descendant internodons, excluding further originator internodonsan dthei rdescendan tinternodons' ;internodon sbein gth eequivalen to finternoda l speciesi nth esens eo fNixo nan dWheele r(1990 )an dth eoriginato rinternodo nbein gv an internodondistinguishe db yhavin gsom equalit yQ 'an dqualit yQ being vth epropert yo fa n internodon within the life span of which acharacte r statebecome s fixed'. Thecomposit especie sconcep ti sno tteste dfo rit sgenera lapplicabilit ythu sfar .Especially ,it s consequences for hybridization, when occurring, arecompellin g for further study.Th e 80 compositespecie sconcep tha sa sconsequenc etha thybridizatio nbetwee nspecie scanno t occurspontaneously ,o rels especie sthough tt ob eseparat ehav et ob ecombined .Th eresult s under1.4 .ar eobtaine db yartificia lhybridization ,starrin gfro mthought saroun dth ebiologica l speciesconcep tan devolutionar yconsideration sassigne dt otha t(Dobzhansky ,1935 ;Lotsy , 1916;Solbrich ,1968 )an dtherefor esa ysomethin gabou tgeneti crelatednes srathe rtha nabou t species'border-linecase sbearin gth ecomposit especie sconcep ti nmind .The ymay ,however , behelpfu li nformulatin ghypothese so nho wspecie shav eevolve dfro meac hothe ra son eo f theconsequence so fth ecomposit especie sconcep ti stha tne wspecie sbranche sof f from existingspecie san dtha treticulat evariatio npattern sma yexis t(Wanntorp ,1983 )rathe rtha n thata n ancestralcomple x splitsint otw o new species (Kornet, 1993). 81 1.5. Description ofth egenu s Clematis 1.5.1. Growth habit Clematis is rather polymorphic in growth habit. The predominant habit within the Ranunculaceaei sth esingl estemme dperennial ,o fwhic hth eseasona lgrowt hi sende db yon e terminalflower o ra ninflorescence ,wherea sth enex tyear' sgrowt hdevelop sfro mgeophyti c sideaxe so rrhizomes .Thi sgrowt htyp ei sprevailin gi nmos tAnemone specie san di nsevera l Clematisspecies ,suc ha sC. integrifolia.Th ebranchin gpatter no fth erhizom ei ssimila rfo r allClematis species .Th edegre eo fwoodiness ,o rpersistenc et oth enex tgrowt hseason ,an d thedegre eo fbranchin go fabovegroun dpart si nsubsequen tyear sar eth evaryin gfactor so n theabov etheme .Som especie sdi ebac kt oth egroun dan dar eperennia lherbs .Other sdi e backt oth ebasal ,woody ,overgroun dparts ,wherea sth edevelopmen to fth erhizom ewarrant s sufficient numbero fne waxes ;the yfor msubshrubs ,suc ha sC. texensisBuckley .Th ewood y typeo fthi shabi tshow sn orea ldifferenc ebetwee nvegetativ ean dgenerativ esid eaxes ;the y formvigorous ,wood yclimbers ,suc ha sC. vitalba. Othe rwood yplant swithi nClematis sho w twotype so f aboveground stems: vegetative stems developing from the rhizome orbasa l aboveground nodes; generativestem sdevelopin gfro mhighe rnode san dendin gu peithe ri nsynflorescence s consistingo fsevera laxillar yan d(composite )cymes ,o ri n(composite )cymes ,o ron e solitary flower; these stems diebac k after thegrowt h season. Thistyp ei srepresente db yC. viticella.A nextrem eo fthi smode li stha tthes egenerativ estem s arever ymuc hreduced ,an dflowering occur si nth enex tgrowt hseaso ni nth eaxil so fth eon e yearsgrowth ,a si nC. alpinaan dC. montana. Hallee tal .(1978) ,Tomlinso n (1984)an d Jeannoda-Robinson (1977)hav emad edescription so fgrowt hmodel so fwood yplant san d perennials.Clematis largel yfit sth emode lo fTomlinson .Thi smode lstart sfro ma rhythmi c annualgrowt ho fa rhizom ewit hroot sa tth ebasa lnodes .Th erhizom eend si na nuprigh tste m withfolia rnode san dfinally node scombinin gfoliag ean dinflorescences .Cremer s(1973,1974 , 1975)mad eclea rtha tther ema ywel lb eseparat egrowt hmodel sfo rclimbin gplants ,a sthe y havesometime sa growt hhabi tintermediat ebetwee nthos eo fwood yan dherbaceou splants . 82 Moreover,rhizom eorganizatio nha st ob econsidere di nrelatio nwit hth evegetativ esprea do f theplant s (Bell, 1974). Thegrowt hform sar eexemplifie d withphotograph so fC. integrifolia,C. heracleifolia an d a schematic presentation of Tomlinson'sgrowt h model in figure 1.24. Especiallyi nwood yclimbers ,th edistinctio nbetwee ntypica lgrowt hform si nClematis is only gradual, as can be seen from species such as C. patens. Prantl (1888) already described thecoherenc e of apparently different growth forms. t " 1 I I I a Modelo fTorrttirBo n • Nodeswit hfolage wxt Mflorascww M Nodeswit hfoliag e 83 Figure 1.24. Basicgrowt hhabi t ofClematis a^Tomlinson'sgrowt h model;b. ,c , d. C.integrifolia; e. C. heracleifolia. (Photographs:Jo sva nd eVooren ) 84 1.5.2. Leaves Leafmorpholog yi nClematis i sver yvariable ,rangin gfro msimpl et oternatel yo rpinnatel y composed leaves.Simpl eleave s occur onlyi n someperennials , such asC. integrifolia, whereasconsequen tternate ,o rbiternat eleave soccu ri nsom ewood yclimbers ,suc ha sC. columbiana Torr. &A.Gra y andC. alpinarespectively . Inmos tClematis spp. ,th eon ebasi clea fmorpholog yvarie sconsiderabl ybetwee n individualplants ,an dshow smoreove reffect so fpositio nwithi nindividuals ,suc ha sreductio n oflowe rleave s(thes ema yeve nb esimple) ,an dgradua lchang et obracts ,bracteole san d bracteolules. Kuntze (1885) remarked that it is sometimes impossible to describe composite Clematisleave swit hth eusua l descriptiveterminology .Man y Clematis spp.,suc ha sC. flammula,bea rleave so fwhic hth ebasa lleaflet so fth efirs torde rar eagai nsubdivide don e tosevera ltimes ,wherea stowar dth eto pthe yar ejus tsimple ,wit hintermediat evariousl ylobe d forms inbetween . Such leaves heterme d flammuliform. Inperennia lClematis spp. ,th eone-yea rol drhizomes ,an dlate rthei rgeophyti csid e axes, bear much reduced scaly and simple leaves. The lateral branches, bearing the inflorescences, develop from therhizom e axils. 1.5.3. Synflorescence and inflorescence Thebasi cstructur eo fth eClematis inflorescenc e isth ecym e(figur e 1.25a),mostl yborn e axillaryalon ga youn gshoo tan dbearin gbracteole san dbracteolules ,whic hloo klik ereduce d leaves.I nClematis spp .wit hmultipl ecompoun dleaves ,bract sar esimila rt oleaflet so fth efirs t order,bracteole st oleaflet so fth esecon dorde ran dbracteolule st oleaflet so fth ethir dorder . Insom especies ,however ,bracteolule sar ereduce dt omino rscales ,whic hdisappea rsoo n after flower buds open. Insom eClematis specie swit hsolitary ,axillar yflowers ,th ebasi ccymos estructur eo f theinflorescenc eca nb erecognize dfro mflowe rstalks ,bearin gbracteole sa tth eplac ewher e thepeduncl eend san dth epedice lcommence s(figur e 1.25b).I nthes ecases ,th einflorescenc e 85 maysometime sbea rtw oo rthre eflowers ,thu sshowin gth eorigina lpatter no fth einflorescence . Inothe r Clematis species,suc h asC. orientalis, theinflorescenc e isa composit e cyme,i.e .a cym eo fwhic hth esid ebranche san dth eto pbranc hfor magai npartia lo rentir e cymes (figure 1.25c).Thi s canb erepeate d several times,leadin g tocluster s of flowers. Figure 1.25. Schematic presentation of Clematisinflorescences . a. dichasium ground plan. b. reduced dichasium: solitary flower c. multiple dichasium b'. bracteole b". bracteolule Troll (1964) used the term synflorescence for flowering branches, that are characterizedb ya gradua ltransitio nfro mvegetativ egrowt ht oinflorescenc e structures.I n variousplants ,thi sappear st ob ea pragmati c approach.I nCompositae ,fo r example,th e inflorescence isalway sa flowe r head(capitulum) ,surrounde db ya ninvolucrum ,bu tth e synflorescence may vary greatly in size and shape. As a rule, in all cases of derived, inflorescencestructure sdescriptio no fth esynflorescenc ei sworthwhil econsidering .I nplant s with gradual transition from vegetative to flowering branches, the description of the synflorescenceadd st oth eunderstandin go fth eplan tmorphology .I nCruciferae ,fo rinstance , theinflorescenc ei salway sa raceme ,bu ti fw eloo ka tth etota lo fraceme san dthei rposition , synflorescences areeithe rabsen to rgreatl yvar yals oi nsiz ean dstructure .Th esam ehold sfo r Ranunculaceae. In Clematis, some species bear thyreoid synflorescences, of which the inflorescence properlyremain sdeterminate ,base do nth ecymos estructures ,wherea sth e synflorescence has an undeterminate character, acting more or less as a raceme. The characterizationo fth esynflorescenc estructure ,bein gthyreoi dinstea do fbein ga rea lthyrsus , ist ob ejustifie db yth efac ttha tth eto pstructur ei si nfac teithe rdeterminate ,bein ga termina l cymetha ti smuc hreduce dan dbear softe n abortedflower buds ,o rundeterminate ,bein ga terminal cymewhic h partially orwholl ychange s overt ovegetativ e growth endingb ya vegetative bud asth e start of next year's growth. Theconfusio n aboutinflorescenc e structureha smuc htrouble dClematis literature , rangingfro mth emisapplicatio no fterms ,suc ha sbracts ,bracteole sandbracteolules ,t oth e misinterpretation offlowering branche sb yvariou sauthor s(Baillon , 1867;Prantl , 1888; Tamura,1968a) .Th eimportanc eo fa consisten tterminolog yi nthi srespect ,i sth emor eeviden t asa Clematis cultiva rgrou pclassification ,principall ybase do nhabit ,synflorescenc e and inflorescence morphology exists since more than a century (Moore &Jackman , 1872; Brandenburg &va nd eVooren , 1988a;Brandenburg , 1989a). 87 either flower or vegetative bud for next season's growth •flower 1 bract 2 bracteole 3 bracteolule Figure 1.26. Schematic presentation of Clematissynflorescences . 88 1.5.4. Flower Atfirs tsight ,Clematis flowers see mt ohav ea simpl estructur ewithou tspecialize dorgans . Normallyth eflower i s hemicyclic, in having an undifferentiated, +cycli c perianth, an androeciumconsistin go fnumerou sspirall yarrange dstamens ,an da gynoeciu mo fnumerou s spirallyarrange dapocarpou spistils ,eac hpisti lformin ga nachen e(Schoffel , 1932).Clematis hasthi sflowe rmorpholog yi ncommo nwit hth eothe rgener ao fth etrib eAnemoneae .I tca n beconsidere da sth efirs tvarian to fth emos tprimitiv estructur eo fth eflowe r(Dilche r& Crane , 1984;Kosuge&Tamura,1989;Ra6ner ,1931;Trapl ,1912) .Th edistinctiv echaracte ri stha t theClematis flowe rha stw owhorl si nit speriant hi nlie uo fth edecussat elea fposition .A serie s oftransec tslide so fth eflowe rdevelopmen to fC campaniflorab yAckerman s(1983 ;figur e 1.27)demonstrate sthis ,usin gth eapproac hb yMeicenheime r(1978,1979 )an dth eanatomica l dissectiontechniqu eb yTob e(1976,1980a , 1980b). Thepresenc eo ftw owhorl scanno t alwaysclearl yb erecognize di nth earrangemen to fnecta rleave s- i fpresent ,a si si nsectio n Atragene. Inth eandroeciu man dth egynoecium ,th epresenc eo ftw owhorl sca nonl yb e deducedi npar tfro mth eabov ementione dserie so fslide sb ycomparin grelativ eposition san d dimensionso fstamen san dpistil srespectively .Compariso no fth eresult sb yAckermans , Meicenheimeran dTob ereveal sth eagreemen to fAckerman san dTob etha ttw owhorl si nth e flowerarrangemen to fClematis coul db eshow na sopposit ei nRanunculus L. ,analyse db y Meicenheimer.Apar tfro mhi sanatomica lwork ,Ackerman sseparate dflowe rpart spe rflowe r ofsevera lClematis specie san dmounte dthe mo npape ri nth eexistin gorde rshowin gth e relationbetwee npositio nan ddimensio no fflowe rparts .Comparin gflower part si nthi swa y provedth epresenc eo ftw owhorl si nth eflowe rarrangemen to fClematis, irrespectiv eo fth e speciesconcerned . VanHee l (1981,1983) studied theformatio n of free carpels,typica lfo r Clematis. 89 90 $& Figure 1.27.Exceip to fa serie so ftranssection so fflowers o fC. campaniflora.Legend : a youngbud ,tepal sdevelope d(tepal s+ floral base ) b maturebud ,transectio na tbas e(tepal s+ androeciu m+ floral base ) c immaturebu d(tepal s+ androeciu m+ gynoeciu m+ ovule sdeveloping ) d same bud as b, but transection in the middle of the bud (tepals + androecium + gynoecium+ ovule smature ) ft) -floral base ;t -tepals ;a - androecium ; g-c/d- gynoeciu m / carpels developing;g-c/ m- gynoecium/ carpel smature ;o - ovule. (Photographs:Gu yAckermans ) 91 Prantl(1888 )analyse dth efloral morpholog yo fth eRanunculacea efro m- a sh epu t it -a phylogeneti c point of view: 'Esdiirft ezunachs tzugegebe nwerden ,das sbe ieine rAnzah lvo nGattunge n(wenigsten si n ihrengroBere nArtenzahl )uberhaup tnu rei neinfache s Perigonvorhande n ist,da skein e Sonderung in Kelch und Krone erfahrt, das auch weder als Kelch noch als Krone zu bezeichnenist ,wei lebe nkei nGrun dz ude rAnnahm evorliegt ,das sde rander ediese rbeide n Perigonteile wiederum verschwunden sei' (p.15) . ForRanunculaceae , Prantl settled for the following basic flower structures outside the androecium: 1a . Simplepetaloi dperianth ,necta rleave s(Honigblatter )absent :a.o .th emajorit yo f Anemonean d Clematis species; 1b . Pedantwit hdifferentiate d calyxan dcorolla ,necta rleave sabsent :a .o .Anemone § Knowltonia; 2a. Simpleperianth , either tending towards corollar, ortoward s calycious perfomance, a nectar leaves present: a.o. Clematis zeylanica; P staminodia present: a.o. Clematis§ Atragene; 2b. Perianth with differentiated calyx and corolla, nectar leaves present. Prantl made adistinctio n between nectary andth eter m nectarleaf , introduced byhim : Onlynecta rsecretin gtissu ei sindicate dwit hnectarium ;suc htissu ema yb elocalize d onan yflowe r parts, such as stamens in Clematis§ Viorna; whereas nectarleave sar eleaf yorgan swit ha smai nfunctio nnecta rsecretion ;necta rleave sar e supposed tob e derived from stamens after losing theirreproductio n function. Withinth eRanunculaceae ,nectari aoccu ro nnorma lstamen s(Clematis § Viorna);o nnecta r leaves(mos tRanunculaceae ;Werth ,1941) ;an do novarie s(Caltha an dmos tTrollius spp. ; Prantl,1888) .Thei rlocatio nshow su pb ytreatin gfilament swit hFehling sreagens ,a sClematis nectarlargel yconsist so fdisaccharide s(Dauman nan dSlavikova ,1968) .Clemati sflower sar e notver y specialized inpollinator s (Knuth, 1898,1904). 1.5.4.1Periant h Tepalsusuall y4 ,bu ti nsom especie s5-1 0o rmore ,petaloid ,aestivatio nvalvate ,induplicat e orwit hag esometime simbricat e(Godley ,1977) .Tepa lpositio ni sdistichous ,eac hpai rha s similarlyshape dtepals .Wit hregar dt ovenation ,thre elargel yparalle lmai nvein sar econnecte d 92 byreticulate mino rveins .Shap ean dsiz eo ftepa lmargi noutsid eth emai nvein svar ydependen t onbu daestivation .I nsom esections ,suc ha sViticella, ther ei sa nextende dmargi n (figure 1.28b),wherea si nother sth emargi ni sjus treduce dt oa villos emargi noutsid eth elatera lvein s (figure 1.28c). 11m 12 Figure 1.28. Polymorphism in Clematis tepal morphology. a. basal shape of tepal. b. tepal with showymargin sbesid e theveins . c. tepal with hardly anymargi n (mostly lanate) beside thelatera l veins (11 and12) . m. midrib. 11. left lateral vein. 12. right lateralvein . 93 Theape xo ftepal si smostl yacuminat et oacut ean dsometime smucronate .Th ehairines so fth e outeran dinne rsid eo ftepal sma yvar yindividuall ywithi nspecies ,althoug hi nth ebu dstag eth e margin isa tleas t pilose,mostl yvillose . 1.5.4.2. Nectar leaves Nectarleave sar eno tcharacteristi cfo rClematis a sa whole ,bu tfo rcertai nsection so fth e genus,suc ha sAtragene. Thei rshap ean dsiz e(figur e 1.29)ma yvar yfro malmos tpetaloi d organst onon-functiona l stamenso fwhic hth efilament sar estrongl ydilatate ,a se.g .i nsectio n Naraveliaan di n someAtragene species .I nothe r sections,som e species showa strong tendencytoward sth edevelopmen to fstaminodia ,whic har eessentiall yno tdifferen tfro mnecta r leaves,thu sleadin gt odouble-flowere d forms.I nsom especies ,thi stendenc yi sexpresse ds o frequently,tha tafte rthei rintroductio nt oEurope ,suc hform swer eoriginall ysuppose dt ob e the wild typeform . This wasth e case with Clematisflorida Thunb .e x Murray. Apartfro ma stron gmidrib ,th evenatio no fnecta rleave si slargel yreticulate .Th e nectaries arelocate d on and alongth e midrib of those leaves (Heyting et al., 1980). 1.5.4.3. Stamens Stamensar enumerous ,positione di ntw owhorl sa tth eflowe rbase ,th ewhorl sgraduall y developingfro meithe rth eperiant ho rth ewhorl so fnecta rleaves .However ,du et oth espiral likestructur eo fth eflower ,ther ei sn ofixe dnumbe ro fstamen spe rflower ,i nthi srespec tther e areconsiderabl edifference s between Clematisspecies :C. viticellaha sa naverag eo f 60 stamenspe rflower ,wherea sC. alpina an dC. montana hav ea naverag eo f13 0stamen spe r flower. 94 • t Figure1.29 .Polymorphis mi nnecta rleave so fClematis flowers .Blac kdot smar kth eventra l location of nectaria (Heyting et al., 1980). a. Staminodial nectarleaf ;stamina lderivatio n is still easily recognized. b. Spatulate nectarleaf ;filamen t part is still recognizable. c. Petaloidnecta rleaf ;th estamina lderivatio ni sonl yt ob ededuce db yform stransitiona l tofunctiona l stamens 95 Figure 1.30. Clematisorientalis stamen . Dark spotted areaa tth eto po f thefilamen t marksth eplac e wherenectari a are localized. 96 Filamentsfiliform, o rdilatate ,glabrous ,ciliat eo rpilose .Full-grow noute rstamen s sometimeshav emor eelongate dfilament stha nth efilament so finne rstamens .Filamen tshap e issometime sa characte ri nsectiona lclassification ,bu ti ntha tcas ei ti sinconsistentl yapplied . Anthershav etw otheca ean da connectiv eeithe ra slon ga sth eanther ,o relongate d (sectionCheiropsis), o rwit ha nappendi x(sectio nNaravelia). Althoug hthes econnectiv e charactersar econsisten ti nthes etw osections ,the yma yals ooccu ri nothe rsections .I nsom e sections, sucha s section Viticella, connectivesar eneve rpresent . Arenecta rleave st ob econsidere dtransforme dstamina ,thu sbein gnon-functiona l staminao rstaminodia ,a tfirs t sighti nman yClematis spp .ther ear enorma llookin goute r stamina,whic har ehoweve rsteril ean dhenc estaminodial .Thes estaminodi afunctio nnormall y inever yrespec tbu tfo rpolle nproduction :nectari aar efoun di nth euppe rpar to fth e filament (figure 1.30.). 1.5.4.4. Pistils Pistilsar eapocarpous ,numerou san dpositione di ntw owhorl sa tth eflowe rbase .Du et oth e spiral-likestructur eo fth eflower ,ther ei sn ofixed numbe ro fpistil spe rflower ,a si sth ecas e inman yothe rRanunculacea ean dals oi nRosacea e(Bessey ,1898) .I nthi srespec tquit elarg e differencesbetwee nClematis spp .exist ,th eaverag enumbe rpe rflowe rrangin gfro m3 0t o 130.Gradua ltransition sbetwee nstamin aan dpistil ssometime soccur ,du et oth ewhor l structureo fth eflower .Steril eorgan soccu rthe nwit ha rudimentar yovary ,a style/filamen tan d a sterileo r areduce dfertil e anther atth etop . Thestigm ai sglabrous ,no tmarkedl ytwo-lobed ,wit ha stick ysurface ,ben ta tth etop . Apartfro mcolou ran dsize ,ther ei shardl yan ystructura lvariatio ni nstigm amorpholog y throughout thegenu sClematis. Stylesar eplumose ,wit hbranche do rsingl ehairs .Apar tfro mcolou ran dsize ,styl e morphologythroughou tth e genus Clematis isquit e uniform. Ovariesar eellipsoid ,rhomboi do rdeltoid ,contai ntw ointegument san dl-6(-8 )ovules , ofwhic honl yth elowes ton ei sfunctional .Th eothe rovule sabor ti na nearl ystag eo rreduce , remaining non-functional. 97 1.5.5. Fruit Afterfertilization ,one-seede dachene sdevelo pfro mth eovaries .Ther eare ,however , indicationstha tfruit s alsodevelo pafte rpolle ntub egrowt hint oth estyle ,du et oadventiv e embryony orparthenocarp y (see 1.4.).Achene s areellipsoid , rhomboid ordeltoid ,an d sometimesstrongl ydorsiventrall yribbed .Th efrui twal lca nb erathe rwoody ,leathery ,smoot h orintermediat ei nstructure .A visuall ymatur efrui tcontain srelativel ymuc hendosperm ,an da smallprematur eembryo ,tha tdevelop sfurthe rdurin gth elon gdormanc yperio d(Schaepp i& Frank, 1962;Tamur a& Mizumoto , 1972;cf . 1.4.). Aremarkabl efeatur eo fsevera lClematis spp .i sth estrongl yelongated ,plumos estyl e atth eto po fth eachene .I ti sa conspicuou scharacte ro fsevera lClematis sections ,suc ha s Atragenean dMeclatis, whic hcontribute sconsiderabl yt oth eornamenta lvalu eo fit splant s in cultivation and inth ewild . 1.6. Summary description ofClematis Woodyclimbers ,subshrub so rperennials .I fclimbing ,plant sar edoin gs owit hwindin g petiolules. Leaves opposite, rarely or occasionally alternate, simple or compound. When compound, leaves areon et oman y times regularly orirregularl y ternately orpinnatel y subdivided. Inflorescencesbasicall yone -t omany-flowere d cymesa tth eto po fyoun gstem so r axillaryo nyoun go rol dgrowth , organized inraceme-lik e synflorescences. Flowerswit h4-8(-many )tepal si na nundifferentiate d perianth,numerou sstamen san d mostlyman y apocarpous pistils forming achenes sometimes with plumose,elongate d styles. Agenu so fabou t15 0species ,disperse dthroughou tth eworl dexcep tfo rpola rregion s withit smai ndistributio nare ai nth enorther ntemperat ezone .Th egreates tdiversit yoccur si n theFa rEast . Themajorit y ofth elarge-flowere d cultivarsan da considerabl eamoun to fsmall - flowered onesar eo fhybri d origin and cannot be assigned to aparticula r species.Thei r classification willb edeal t withi nChapte r4 . 98 2. CLEMATIS SECTIONMECLATIS (SPACH) BAILLON 2.1. Description of thesectio nMeclatis (Spach)Baill . Spach (1839)describe dMeclatis asa separate genus withtw o species: Meclatisorientalis (L. )Spac h (basionym: Clematis orientalis L.) Meclatissibirica Spac h (synonym: Clematis glaucaWilld .p.p. ; Clematis intricata Bunge) Thecharacter s todistinguis hMeclatis from other Clematis spp.ar e according toSpac h (1839): 'Sepales4 ,petaloides ,pendan tl'epanouissemen tetale so urevolutes ,divergents ,e n prefloraisonimbrique spa rle sbords .Petale snuls .Etamine spaucisenees ,conniventes ; filetslanceote s(d umoin sle sinterieurs) ,comprimes ;anthere slineaires-oblongues , inappendiculees.Style slongs ,filiformes , obtus.Nucule scoriaces , subfusiformes, tetragones,u npe ucomprimees ,legeremen tmarginees :bord stranchants .Gynophor e subglobuleux.' Afurthe r extensive description bySpac h presents thefollowin g characters: 'Ramulesflorifere s axillairese tterminaux ,dichotome s(!) ,brachies .feuille s aux bifurcations, nusinferieurement , 3-15 flores' (!); 'Feuilles petiolees, glauques (!),molles , nonpersistantes'; 'Fleurs legerement odorantes (!),asse zgrandes ,nutantes ,disposee s (..)e ncym e subfastigiees' (!); 'Sepalesjaunes (!) ,plane s(!) ,5-nerve s(!) ;le s2nervure smarginale se tlamedian e saillantes,carenees ;le s2 intermediaire stre sfines) ,acumines ,cotonneu xau xbords , plus longqu e les etamines'; 'Filets violets (!), cilies'; 'Nucules petites,lisses ,pubescente s dememe squ el egynophor e (!).' Thecharacte rstates ,indicate db yexclamatio nmark sabov e(presen tauthor) ,d ono talway s hold or are absent: dichotomydoe sno toccur ,bu ti sth eresul to fa someho wartificiall yaborte dtermina l twig; solitaryflower s also occur; glaucousleave sar epresen ti nman yMeclatis specimen sbu tno ti nall ,wherea sthe y also occur inothe r sections, such as Viorna; insom especie shav earemarkable ,i fno todd ,fragrance ,bu tthi scharacte rvarie si n intensity; subfastigiate inflorescences wereno t observed byth epresen t author; tepalcolou ri spredominantl yyellow ,bu tvariatio noccur srangin gfro m tingedo r 99 spotted with violet to full violet asa n exception (Spachterm s tepals sepals); tepalsma yb eflattene d(plane )i nsom eMeclatis specimens ,bu tcertainl yno ti nall , whereas thepresenc e of 5nerve s isa misinterpretatio n of thetepa l venation; filaments(an dtepal sa swell! )indee dofte nar etinge dwit hviole to rar eeve nmarkedl y violet,bu t manyMeclatis specimen sjus t haveyellowish-gree n filaments; Iti ssuppose dtha tSpac hmean tb ygynophor ea tleas tth eandrogynophore ,a sth e transition zonebetwee n androphore and gynophore isofte n not markedly present. Thesecharacter sar eno tunequivocal :non ei suniquel ydistinctiv efo rMeclatis. Therefore , Meclatiscanno tb emaintaine da sa genus .Apar tfro m thecombinatio no fsom echaracte r states (bright yellow flowers, pinnatisect leaves andrathe r small,pubescen t achenes),i t possesses,however ,a physiologica ltrai twhic hmake sthi sgrou pa systemati centit ywithi nth e framework ofClematis: Meclatis spp .lac kth estric tan dlon gsee ddormancy ,whic hi ss o characteristicfo rmos tClematisspp. (Barton,1967;Blair , 1959;Kinzel,1913;Niethammer , 1928)an dothe rrepresentative so fth etrib eAnemoneae .Afte rsowin gwithi nthre eweek sth e seedsgerminat ereadil yan dseedling sar egrowin gwithi non eyea rint oflowerin gplants .Thi s lacko f seed dormancy isclearl y of interest for Clematis cultivation andbreeding . Clematissectio nMeclatis (Spach )Baillo n(1867 )- Histoir ede splantes ,vol .1:52-62,87 . Paris;type : C.orientalis L. (Lectotype Dillenius 2868, OXF-DBLL). Basionvm: MeclatisSpac h(1839 )- Histoir enaturell ede svegetaux ,vol .7 Le sClematidees :257-284 . Librairie encyclopedique deRoret , Paris;type :Meclatis orientalis (L. ) Spach. Homotypic synonyms: Clematisserie sOrientates Prant l(1888 )- Beitrag ezu rMorphologi eun dSystemati kde r Ranunculaceen. Botanische Jahrbiicher9 :225-273 ; Clematis section Clematis subsection Orientates(Prantl )Tamur a(1967 )- Morphology , ecologyan dphylogen yo fth eRanunculacea eVI .Annua lreport so fth eColleg eo f General Education OsakaUniversit y 1967:13-35 . Description:Wood yclimbers ,flowerin gi nsummer ;(June)-July-Septembe ro nth eyoun g growtho ftha tseason .Leave sver yvariabl ei nshap ean dsize ,pinnatel yo rpartl yo rwholl y bipinnate,i ncertai ncase stripinnatel ysubdivided .Inflorescenc eaxillar yand/o rtermina lrangin g fromsolitar yflower st o3-many-flowere dcymes ;peduncl erangin gfro mver yshor t(almos t absent)t opredominand ypresent ,thu sgivin gth eimpressio no fver ydifferen t inflorescential structuresbetwee nspecies ;pedice lcurve da tth eapex .Flower swit h4 ,occasionall y5 tepals , 100 yellowo rgreenis hyellow ,sometime svariousl yligh tpurplish-brow ninsid eo rspotte do rringe d withred-viole toutside ,lanat ea tth eincurve dmargin .Stamen snumerous ;filament sdilatate , towards the base pilose, or ciliate; anthers without an elongated connective or other appendages.Pistil snumerous ;ovarie sellipsoi do rrhomboid ;style slon gpubescent .Achene s rhomboid,variousl yribbe da tth emargin ,pubescent ,dar kbrow nwit ha mor eo rles s fibrous appearance; styles variously elongate,persistent , covered with long erecthairs . 2.2. Distribution ando f thesectio nMeclatis (Spach) Baill. Thenatura ldistributio no fth esectio ni sextensive ;specie sar efoun di nTurkey ,Syria ,Iraq , Iran,Afghanistan ,Georgia ,Armenia ,Azerbaidzhan ,Kazakhstan ,Turkmenistan ,Uzbekistan , Kirgiziya,Pakistan ,th eHimalaya nregio no fIndia ,Nepal ,Bhutan ,P.R .o fChin a(Tibet , XinjiangUygurZiziqu ,Kansu ,Shansi ,Inne rMongolia ,Sichuan ,Yunnan) ,Mongolia ,Russia n Federation (regions adjacent to abovementione d countries), andNort h and South Korea. Apartfro mthei rnatura loccurrenc ei nth eabov eregions ,som especie shav eescape d fromcultivation .Ther ei sevidenc efro mherbariu mspecimen stha trepresentative sof Clematis sect.Meclatis hav ebee nnaturalised .I nCanad a(Alberta) ,C. tibetanasubsp .tangutica ha s been escaped (Turner 2644 (S)), whereas in the neighbourhood of Georgetown (USA, Colorado), C.intricata must have been naturalised (Weber &Salamu n 12937 (C,S)) . Outsidesect .Meclatis, ther ear emor eexample so fnaturalizin gClematis populations ,suc h asC. viticellaalon gth erive rMaa si nBelgiu man dth eNetherlands .A sfa ra sI know ,thi sha s neverle dt olastin glarg echange so nth evegetation ,bu tremain srestricte dt osmal lisolate d populations. Basedo ndistributio ndat ao fspecimen scite dfo rth etax adescribe di nth efollowin g sections,track s(t ob eunderstoo da sth egraphi cpresentatio no fth ecircumscriptio no fth etota l range of monophyletic taxa; Camp, 1947;Wiley , 1981)ar epresente d in figure2.1 . 101 C. orientate ' a C. ispahanha • b C.intricat e C. graveolens C. serratlfolia V^r~%^ •* 1 C.tibetana subsp .tibetana "2 C. tibetanasubsp .tangutica •• 3 C.tibetana subsp .vernayl Figure 2.1. Tracks of taxa within Clematis sect.Meclatis (Spach ) Baill. 102 Count Midpoint 2 166 0 429 2 692 7 955 13 1218 11 1481 16 1744 11 2007 8 2270 10 2533 9 2796 4 3059 4 3322 0 3585 3 3848 0 4111 2 4374 0 4 8 12 16 20 Histogram frequency Clematis orientalis L 102case s Mean 2009.931 Std.Dev . 847.239 Count Midpoint 3 1908 0 2132 0 2356 1 2580 2 2804 4 3028 8 3252 6 3476 4 3700 10 3924 4 4148 3 4372 2 4596 1 4820 1 5044 0 5268 1 5492 0 2 4 6 8 10 Histogram frequency Clematis tibetana subsp. vernayi (C.E.C.Fischer )Grey-Wilso n 50case s Mean 3639.800 Std.Dev . 737.429 Figure2.2 . Characteristicdifferenc e inrang eo faltitude sbetwee ntw o taxaof Clematis sect.Meclatis. 103 Therear emainl ytw otype so fhabitats ,t owhic htax ao fth esectio nMeclatis ar eadapte d (figure 2.2): low-altitude,dr yhill san dlowlands ,alon gstream san dgorge so nrock yslope su pt o +3.000m abov e sealevel ; montaneregions ,alon gstream san dgorge so nrock yslopes ,altitud erangin gfro m +2.500m t o+5.00 0m . Speciation patterns coincide withborderline s between the above types of habitats.Th e mountainchain so fElbur z(Iran) ,Hind uKush ,Chitral ,Panjshir ,Kashmir ,Tibeta nplateau ,an d Altaifor meffectiv egeographica lbarrier sbetwee ntax ao fth esectio nMeclatis (se eals oAh , 1978;Endler, 1977;Good, 1964;Gould, 1984;Goul d&Eldredge ,1977;Ledebour ,1830 , 1841;Raven, 1979;Rave n&Axelrod , 1974;Roche, 1974;Rosen, 1975,1978;Schuster , 1976;Tarling ,1982) .Wher edistributio narea sma yoverlap ,tax aar eecologicall yisolate d principally by altitude. C. orientalis and C. serratifolia occur at lower altitudes; C. graveolensan dC. intricata occura tlowe raltitudes ,bu tinhabi tmontan eregion sa swel l(se e further speciesdescriptions) ;C. tibetanai sdisperse di nmontan eregions :it shabita trange s from 2.500-5.000m abov e sealevel . Oneo fth emai nproblem so finterpretin gClematis sect .Meclatis specie si sthei rrespons et o particularhabitats .If ,fo rexample ,a specie si sadapte dt o montaneregions ,plant sgrowin g athighe raltitude ssho wa dwarf ,non-climbin ghabitus ,wherea splant sa tlowe rrange ssho w vigorousgrowt htrailin gove rothe rvegetatio no rrock yslopes .Thes edifferen tpopulation swer e oftenconsidere dt ob edistinc tspecies ,bu tcompariso ni na singl eexperimenta lgarden ,a swa s donea tth eA UDept .o fPlan tTaxonomy' snursery ,similaritie sar es ogrea ttha tconspecificit y is obvious.Thi s feature is not unique for thisparticula r section of Clematis, but canb e observedi nothe rpart so fth egenu sa swell .Th eabov ephenomeno nha smisguide dGrey - Wilson (1986,1989)i nhi streatmen t ofMeclatis. Hedescribed ,beside s C. tangutica, a separatespecie sC. pamiralaica, wit ha sonl ydistinguishin gcharacte rit snon-climbin ghabit . Accordingt oGrey-Wilson ,C. pamiralaica occur sonl ya thighe raltitudes .Plant sfro msimila r provenancesi nth eWageninge ntria lfield sprove d- unlik eth esingl epopulatio ngrow na tKe w - to climb asothe rpopulations . 104 Phenotypicplasticit yi sver ydifficul t tointerpre ti nterm so fth edelimitatio nbetwee n species and infraspecific taxa, sincether ei sadaptiv e andnon-adaptiv e components are involved(Morisse t& Boudin ,1984) ,on ecanno tavoi dstudyin git .Therefore ,ecologica leffect s onth eplant shav et ob ecarefull y examined,befor e concluding on species delimitation. Ranunculeanspecie sappea rspeciall ypron et oenvironmentall yinduce dvariation ;se eth eplan t habit of e.g. Aconitum,Anemone, Aquilegia, Delphinium, andRanunculus. In all these genera,classificatio n tendedt oignor esuc hecologica lresponses ,resultin gi na nexces so f described species. 105 2.3. Morphological analysis of thesectio nMeclatis (Spach) Baill. Sincether ei sa lo to fconfusio n onth edelimitatio no fspecie swithi nth esectio nMeclatis, a numericalanalysi so fth esectio nha sbee nundertaken ,base do nmorphologica lcharacter si n ordert o determine thediscontinuitie s between species. 2.3.1. Materials andmethods Forth emorphologica l analysis of the sectionMeclatis, 20 6herbariu m specimens were selected. They were apriori assigned to species names (see table 2.1. and Appendix 1 available ondisk) . Table2.1 .A prior iassignmen to fClematis sect .Meclatis specie suse dfo r morphological analysist oherbariu mspecimen suse di nth emorphologica lanalysi s(fo rspecime ndetails ,se e Appendix 1 available on disk). - Nos. 1- 5, 126-127 C.hilariae Kovalevsk . -Nos . 6- 11,132-13 4 C.serratifolia Rehder -Nos . 12-2 4 C.akebioides Veitch -Nos . 25-51 , 128-131 C.glauca Willd. -Nos . 52-6 6 C.tangutica (Maxim.) Korsh. -Nos . 67- 81 C.tibetana Kuntz e - Nos. 82-9 9 C.graveolens Lindl. - Nos. 100-120 C.vernayi C.E.C.Fisch. - Nos. 121-125 C. vernayi/tangutica - Nos. 135-148 C.ispahanica Boiss. - Nos. 149-206 C.orientalis L. Thelis to fherbariu m specimensuse di sprovide di nAppendi x 1 (availableo ndisk) .Al l specimensha dbee ncollecte da tnatura lsites ,thu sreflectin gthei rnatura lphenotype .Followin g astandardise dforma tbot hqualitativ ean dquantitativ echaracter swer escored .Qualitativ e charactersar epresente di ntabl e2.2. ;quantitativ echaracter si ntabl e2.3 .togethe rwit hthei r frequencystatistic s(minimu man dmaximu mvalue ,median ,mea nstandar ddeviation ,variance , kurtosisan dskewnes so fth ecurve) . Therati o1/2 ,a smeasur et ocharacteris eth epositio na t whichth egreates twidt hi sreached ,i sexplaine dfo rtepal si nfigur e2.3 .Th erati oi sals ouse d 106 for leaflets andachenes . Toanalys eth edat aset ,th esoftwar e packagesSPSS-PC 4an dGENSTAT 5wer e used.I nanalysi sprocedures ,bot hqualitativ ean dquantitativ echaracter swer eprocesse dt o createa similarit ymatrix .Th econtributio no fa variat et oth esimilarit yi scalculate da sEuclidea n distancesbetwee nquantitativ echaracters ,a sJaccard' scoefficien t forbinomia lqualitativ e charactersan da sManhatta ndistance sfo rpolynomia lqualitativ echaracters .Th esimilarit y matrixha sbee nuse dt operfor m principalcoordinat eanalysi san dhierarchica lclusterin g (averagelinkage) .Wit hth equantitativ echaracter sonl y(excludin gth efrui tcharacter sa sto o manyvalue sar emissing )principa lcomponen tanalysi san dconsequen thierarchica lclusterin g (average linkage based onth e principal component scores) havebee n performed. Only quantitative characters were used as the analysis is based on the correlation matrix of characters.Correlation sbetwee nqualitativ ean dquantitativ echaracter sar emeaningless . Whethero rno tqualitativ echaracter ssho wgenera lcorrelatio nha sbee nanalyse db yx 2-test andcalculate d significance scores(P) . Afterhavin gestablishe dOTLT so nmorphologica lsimilarity ,th egroup sscore sfo rth equalitativ e characterswer estartin gpoin tfo rphylogeneti canalysi susin gHennig86 ,releas e 1.5.Th e restrictiont oth equalitativ echaracter swa sbase do nth eobservatio ntha tmos to fth ediagnosti c characters werepresen t inth equalitativ e dataset .Th eOTU' swer eanalyse db yimplici t enumeration, followed by 'bb' and successive weighting according toFarri s (1988). 107 Ratio 1/2: Parto f greatest lengthfro mape xunti l intersectionwit hgreates twidt h Parto f greatest lengthfro mbas eunti l intersectionwit hgreates twidt h epalform s usedfro mfig . 1.28,bu tals oapplicabl et oothe rplan tshape s Figure 2.3. Ratio 1/2 asa measur e to characterise thepositio n of the greatest width. 108 Table 2.2.Qualitativ e characters usedfo r themorphometri c data matrix - Vegetativeparts. Character Character state Code Character Character state Code 1 shape of young wood round/angular 1 8 leaflet incision entire 0 costate 2 + serrate 1 + serrate/dentate 2 2 indumentum of glabrous 0 + serrate/dentate/lobed 3 young wood pillose 1 + dentate/lobed 4 + serrate/lobed 5 3 shape of old wood round/angular 1 serrate 6 costate 2 serrate/dentate 7 serrate/dentate/lobed 8 4 stem habit not climbing 0 dentate/lobed 9 climbing 1 serrate/lobed 10 irregularly lobed 11 5 leaf blade structure simple 0 pinnate 1 9 leaflet apex acute 1 pinnate and bipinnate 2 acuminate 2 bipinnate 3 mucronate 3 ternate 4 bitemate 5 10leafle t base cordate 1 triternate 6 cuneate 2 angustate 3 6 petiole glabrous 0 pillose 1 11 indumentum of glabrous 0 upper side of leaflets 7 leaflet shape elliptic 1 ovate 2 12indumentu m of glabrous 0 ovate/lanceolate 3 ower surface of leaflets at veins 1 lanceolate 4 overall 2 lanceolate/linear 5 linear 6 13indumentu m of glabrous 0 petiolule at base 1 overall 2 14 petiolule not twining 0 twining 1 109 Table2.2 .Qualitativ e characters used for themorphometri cdat a matrix -flower andfruit Character Character state Code Character Character state Code IS flower position (1) terminal 1 27 tepal margin flat 1 terminal and axillary 2 exduplicate 2 axillary 3 induplicate 3 16 flowers per infl. many 1 28 tepal apex blunt 1 one 2 acute 2 acuminate 3 17 flower stalk straight 1 mucronate 4 nodding at the top 2 regularly bent 3 29 tepal base cuneate 1 angustate 2 18 bracts absent 0 present 1 30 indumentum of glabrous 0 tepals abaxial pilose 19brac t shape simple/entire 1 simple/lobed 2 31 indumentum of glabrous 0 composite 3 tepals adaxial pilose 1 20 flower position (2) pending 1 32 indumentum of glabrous 0 upright 2 tepal margin lanate 1 21 flower shape open/flat 1 33 gradual transition absent 0 trumpet-shaped 2 from tep. to stamens present 1 campanulate 3 tubulate 4 34 indumentum of absent 0 filaments at base 1 22 flowering abundant in a short time 1 overall 2 profuse (longer period) 2 few flowers 3 35 filaments not dilatate 0 dilatate 1 23 tepals free 0 + overlapping, imbricate 1 36 indumentum of absent 0 margins touching, valvate 2 ovary present 1 24 tepal curvature not recurved 0 37 indumentum of absent 0 recurved 1 style present 1 25 tepal shape elliptic 1 38 shape of achene rhomboid 1 ovate 2 ovate 2 ovate/lanceolate 3 obovate 3 lanceolate 4 lanceolate/lineariform 5 39 achene lustre not shiny 0 lineariform 6 shiny 1 26 tepal texture herbaceous 0 40 achene surface not ribbed 0 fleshy 1 ribbed 1 41 pericarp woody 1 fibrous 2 110 o o C3 u •fi o1 4= B- O E 53 2 o > _ a a CI t 1 I 2.3.2.Results Asth e list of scored characters,presente d intable s 2.2.an d 2.3. isth egenera l listuse dfo r all Clematisobservations ,som echaracter sdi dno tsho wan yvariatio nwithi nsect .Meclatis. Thes ewer e omittedfro mfurthe r analysis:indumentu mo ftepa lmargi n(alway slanate) ,indumentu mo fovar y (alwayspresent) ,indumentu mo fstyl ean dachene sshin y/ dul l(the yar eal ldull) .Th esam ehold sfo r numbero ftepal spe rflower :withi nsectio nMeclatis thi snumbe ri s4 ,onl yoccasionall y5 .Th ex 2-test ongenera lcorrelatio nbetwee nqualitativ echaracter sshowe dtha tmos tcharacter sar egenerall y correlated exceptfo rth eshap eo fth eol dwoo dan dth ebrac tcharacter ;th elatte rcharacte rwa s omitted asth e absence of bracts in herbarium specimens wasi n some cases artificial. 2.3.2.1. Factor analysis Factoranalysi swa scarrie dou twit hth equantitativ echaracter si ntabl e2.3 .apar tfro m the fruit characters.Facto rloadings ,an dscore sar epresente di nfigur e2.4 ;th efacto rscore sals oi ntabl e2.4 . 112 Table 2.4.Characte r scores perfacto r byfacto r analysis.Characte r numbers, seetabl e 2.3 «^^^ Factor^ Factor 1 Factor 2 Factor 3 'Character " ^_^ 1 -0.132 0.054 0.061 2 -0.004 -0.023 0.840 3 -0.205 0.758 0.011 4 0.257 0.375 0.069 5 -0.049 0.618 -0.445 6 0.027 0.523 -0.366 7 -0.010 -0.081 -0.035 8 0.037 0.852 -0.310 9 0.260 0.356 0.188 10 -0.514 0.099 -0.381 11 -0.363 -0.058 -0.690 12 0.111 -0.116 0.317 13 0.113 0.134 0.198 14 0.055 -0.120 -0.051 15 0.815 0.000 0.029 16 0.448 0.312 0.191 17 0.119 0.022 -0.068 18 0.107 -0.112 0.674 19 0.783 -0.073 -0.064 20 -0.104 0.083 -0.050 21 0.580 -0.015 0.462 22 0.119 -0.173 -0.064 23 0.638 0.038 -0.009 24 0.072 0.223 0.108 25 0.669 -0.061 0.046 113 Loadings 1.5 123456789 10111213141516171819202122232425 Characters • Factor 1 Factor 2 Factor 3 Figure2.4 .Facto rdiagra man dloading sfo rth efirs tthre efactor s(loading spe rcharacte rpe rfacto r next to each other andcumulative)i n separate with the quantitative characters 1-25 of table2.3 . 114 2.3.2.2.Principa l coordinate analysis Startingfro ma nOT Ub yOT Umatri xo fdistances ,principa lcoordinat eanalysi sha sbee ncarrie d outt orevea ldistance sbetwee nspecimen swit hth eai do fa se to forthogona lvariate s(principa l coordinates).Th efirs t five principal coordinates explain 48.9%o f thevariation , seetabl e2.5 . Table 2.5.Characteristic s of thefirs t 5principa l coordinates. P.C. Latent root Expl.variationMai n contribution of characters 1 7.2059 15.01% indumentum characters of vegetative parts flower stalk/flower position tepal morphology 2 6.6469 13.85% apex of leaflet/leaflet width shape of flowers/blooming/number of flowers per inflorescence tepalsrecurved/indumentu mo ftepal sdownside/tepa l length number ofpistil s 3 3.5402 7.37% tepal ratio 1 / 2 filament dilatate 4 3.2275 6.72% petiolulelengt h achene length 5 2.8584 5.95% baseo f leaflet number of bracteoles perbract / bract stalk length stylelengt h Thescore so fspecimen salon gth efirs t5 principa lcoordinate sresul ti n1 0plots ,o fwhic hth e fourwit hprincipa lcoordinat e1 alon gth ex-axi sar edisplaye di nfigur e2.5 .Fro mthi sdisplays , 115 itca nb e concluded that: Alongth efirs tprincipa lcoordinat especimen s 136-14 8ar eseparat efrom th eothe r specimens; Alongth e secondprincipa l coordinate specimens aregraduall y spread overth eplot ; Along thethir dprincipa l coordinate specimens 82- 9 9an d 177ar emor eo rles s separate from theres t of specimens in a scattered group; Alongth efourt hprincipa lcoordinat especimen s61,11 6an d14 0ar eisolate dfro mth e rest; Alongth efift hprincipa lcoordinat en oclea rgroup sbu tindividua loutlier sar eseparated . Specimensseparate dalon gth efirs tprincipa lcoordinat emus thav ea.o .distinc tcharacteristic s withrespec tt oth eindumentu mo fvegetativ eparts ,flowe rstal kan dflowe rposition ,whic hare , however, all characters that show high phenotypic plasticity. Along this axis separated specimens 136-14 8hav ebee npreviousl ylabelle dClematis ispahanica Boiss .Principa l coordinate2 sho wmuc hcontributio no fflowe ran dflowerin gcharacteristics .The yar erelativel y scatteredove rth erelevan tplots ,henc eexplainin gwh ythes echaracter shav eno tcontribute d mucht oth eclassificatio n ofClematis sect .Meclatis. Principa lcoordinat e3 ha son eo fit s contributionsb ytepa lrati o1/ 2(figur e2.3) ,whic haffect sth eflowe rmorpholog yi nit sgenera l appearance.A sther ear en ogroup so fspecimen sdistinguishe db ythi scoordinate ,specia l flowermorpholog ydoe sno tcontribut et oth edistinctio nbetwee ngroups .Th ethir dprincipa l axisha sfurthe rreceive dit smai ncontributio nfro mfilamen tdilatation .Alon gthi saxi sspecimen s 82-99appea rt ob eseparate di na scattere dgroup .Thes especimen shav ebee npreviousl y labelledClematis graveolens Lindl . Thefourt han dfift h principalcoordinat ed ono tprovid e anyclea r separations between observed specimens.Mos t groups inth eplot s are highly overlapping. 2.3.2.3. Cluster analysis Insearc ho fsubdivision so fth e20 6specimen sthre efurthe r approacheshav ebee napplied . Firstly,a si ti smathematicall yth emos tpur eagglomerativ edevice ,th esingl elinkag eapproac h hasbee napplie dt oth edat at oproduc ea nonrooted ,minimu mspannin gtre e(Gowe r& Ross , 1969).Th eminimu mspannin gtre ei sactuall yno ta tre ebu ta networ k connectingal lindividual s bya se to fstraigh tline sjoinin gpair so fpoints ,whos elength sar eequa lt oth edissimilaritie s 116 betweenindividuals ,an dwhos esu mi sminima l(Chatfiel d& Collins ,1980) .Th eminimu m spanningtre ei shelpfu li nrevealing group sbetwee nindividuals ,wherea sothe rclusterin gdevice s tendt oimpos ea phylogeneti cstructur eo nth eindividua lconcerne drathe rtha nrevealin gthei r mutualrelationship. Th estructur eo fth eminimu mspannin gtre efo rth e20 6Meclatis specimen s ispresente di nfigure 2.6 .Thi sfigure confirm sth econclusion sfro mth eprincipa lcoordinat e analysistha tC. graveolensan dC. ispahanicaappea rt ob eseparat egroups .Furthermore , C. orientalisresult sa sa rathe rcohesiv egroup ,an dres tgroup sappea rcombinin gpart so fth e apriori groups C.akebioides, C. glauca, C.hilariae, C. tangutica, C. tibetana an d C. vernayi.T ovisualis eth edat ai na cluste rdendrogram ,th eaverag elinkag eroutin ewa sapplie d toth edat at ostres sbetwee ngrou pdifferences .Th edendrogra mi spresente di nfigure 2.7 .Thi s dendrogrami si nagreemen twit hth eminimu mspannin gtre ewhere ,again ,C. ispahanicaan d toa lesse rexten tC. graveolens ar eseparated .C. serratifolia i spartiall yseparated ,wherea s C. orientalisi slargel yseparated .Th eres tgroups ,consistin go fth eothe ra priori assigne d species,appea rscattere dthroughou tth edendrogram .Th eoveral lsimilarit y( +60% )withi nth e sectionexplain sth eincomplet eseparatio no fgroup sb yth eaverag elinkag eclusterin gdevice . Based on the correlation matrix of quantitative characters without the fruit characteristics(thes echaracter sha dto oman ymissin gvalues) ,principa lcomponent s( aderive d seto forthogona lvariates )hav ebee ncalculated .Wit hth eai do fthi sse to fvariates ,agai na clusteranalysi s(averag elinkage )ha sbee ncarrie dou tagain ,se efigure 2.8 .Thi sanalysi sshow s a further mixing upbetwee n groups duet o therathe r high overall similarity. Figure 2.5.Principa l coordinate scatter plots for 206Meclatis specimens (Appendix 1 available ondisk) . a pc 1 vs.p c2 b pc 1 vs.p c 3 c pc 1 vs.p c4 d pc 1 vs.p c5 117 Clematisispahanica Boiss. 136-139,14 2 -148 | |Remainin g species Clematis graveolens Lindl. 82-99 Clematis ispahanica Boiss. 136 -139, 141 -148 I I Remaining species -1.0 -0 118 Clematisgraveolens Lindl. I 82-99 \Clematis ispahanica Boiss. 1136-139,141-148 Remainingspecie s -1.0 -0.8 -0.6 -0.4 -0.2 0 0.2 pcl Clematisgraveolens Lindl. I 82-99 IClematis ispahanica Boiss. '136-139,141-148 I IRemainin g species -1.0 -0.8 -0.6 -0.4 -0.2 0.2 pel 119 140 b •92 Clematisgraveolens Lindl . Clematis intricata Bunge Clematis ispahanica Boiss. Clematis orientalis L. Clematisserratifolia Rehder Clematistibetana Kuntze * Specimens (numbers,se eApp . 1 ondis kavailable ) Specimenswit hdiscutabl esystemati cassignmen t Figure2.6 .Minimu mspannin gtre eo f20 6Meclatis specimens ;distance si nth efigur ear e presented asunifor m (seefo r similarities Appendix 2 available on disk). 120 isp. or. i. g. ti.g . i/or. s. ti/i/or. Figure2.7 .Cluste rdendrogra mo f20 6Clematis sect .Meclatis specimen sbase do nqualitativ e and quantitative characters andusin g theaverag elinkag ecriterion . isp.=C. ispahanica;or. =C. orientalis;i. =C. intricata; g.=C. graveolens;ti.=C .tibetana ; i/or.=C. intricatawit hsom eC. orientalisspecimens ;s.= Cserratifolia; ti/i/or =C. tibetana with some specimens of C. intricata andC. orientalis. 121 g+ Figure2.8 .Cluste rdendrogra m(averag elinkage )usin gprincipa lcomponent sa sse to fvariate s derivedfro mth ecorrelatio nmatri xo fquantitativ echaracter sth efrui tcharacter sexcluded . g+=C.graveolens+ someC. tibetana specimens. 122 Table2.6 .Legen dt ofigur e2.7 . U sendt o figure2. 7 o s o S O S O S O S O S O S O S 1 139 27 181 53 149 79 92 105 43 131 190 157 106 183 21 2 136 28 172 54 193 80 93 106 41 132 191 158 109 184 23 3 138 29 174 55 166 81 82 107 42 133 197 159 52 185 20 4 141 30 151 56 179 82 89 108 202 134 25 160 58 186 16 5 137 31 163 57 182 83 90 109 37 135 26 161 59 187 31 6 142 32 161 58 28 84 95 110 203 136 32 162 74 188 108 7 143 33 158 59 29 85 88 111 204 137 33 163 101 189 107 8 144 34 152 60 30 86 86 112 160 138 22 164 103 190 115 9 146 35 159 61 12 87 87 113 167 139 18 165 24 191 117 10 140 36 186 62 96 88 13 114 192 140 14 166 121 192 17 11 148 37 154 63 198 89 83 115 201 141 15 167 124 193 114 12 145 38 180 64 84 90 45 116 38 142 1 168 120 194 64 13 147 39 169 65 85 91 19 117 11 143 76 169 78 195 60 14 135 40 188 66 94 92 105 118 8 144 113 170 112 196 62 15 184 41 200 67 97 93 2 119 9 145 77 171 67 197 54 16 27 42 153 68 98 94 127 120 7 146 79 172 44 198 102 17 177 43 156 69 91 95 3 121 10 147 119 173 47 199 104 18 178 44 164 70 99 96 4 122 205 148 75 174 40 200 110 19 68 45 171 71 133 97 5 123 206 149 100 175 46 201 122 20 185 46 195 72 134 98 34 124 48 150 118 176 53 202 123 21 175 47 196 73 71 99 39 125 51 151 111 177 72 203 125 22 157 48 150 74 73 100 126 126 36 152 116 178 49 204 132 23 176 49 165 75 69 101 128 127 130 153 63 179 66 205 129 24 183 50 162 76 70 102 6 128 194 154 57 180 55 206 131 25 155 51 189 77 80 103 50 129 170 155 56 181 65 26 173 52 187 78 81 104 35 130 199 156 61 182 168 0= 0 rder; S= S r >ecimen 123 Table2.6 .Legen d tofigur e2.8 . Legendt o figure2. 8 O s O S o S O S O S O S O S O S 1 4 27 149 53 110 79 173 105 178 131 38 157 13 183 89 2 3 28 155 54 127 80 137 106 176 132 105 158 19 184 83 3 12 29 154 55 117 81 190 107 180 133 11 159 6 185 92 4 124 30 156 56 141 82 203 108 189 134 57 160 34 186 82 5 65 31 164 57 9 83 166 109 199 135 62 161 49 187 84 6 14 32 75 58 136 84 193 110 194 136 23 162 39 188 94 7 68 33 183 59 147 85 145 111 198 137 28 163 40 189 119 8 56 34 191 60 153 86 162 112 201 138 29 164 48 190 66 9 63 35 177 61 179 87 205 113 139 139 133 165 51 191 116 10 118 36 181 62 171 88 170 114 200 140 8 166 46 192 59 11 55 37 140 63 186 89 187 115 30 141 7 167 47 193 64 12 109 38 196 64 197 90 204 116 37 142 10 168 41 194 90 13 52 39 185 65 172 91 182 117 35 143 80 169 42 195 111 14 58 40 195 66 192 92 61 118 43 144 81 170 44 196 93 15 107 41 18 67 158 93 206 119 126 145 67 171 25 197 87 16 102 42 72 68 165 94 160 120 128 146 120 172 101 198 88 17 54 43 22 69 138 95 157 121 125 147 45 173 1 199 91 18 60 44 70 70 146 96 161 122 121 148 69 174 71 200 85 19 115 45 32 71 143 97 150 123 122 149 106 175 103 201 96 20 151 46 76 72 142 98 132 124 123 150 113 176 17 202 95 21 2 47 74 73 144 99 188 125 131 151 114 177 73 203 97 22 100 48 20 74 184 100 202 126 5 152 31 178 78 204 98 23 175 49 15 75 148 101 174 127 26 153 108 179 112 205 86 24 152 50 21 76 135 102 130 128 129 154 50 180 77 206 99 25 167 51 53 77 168 103 159 129 134 155 24 181 79 26 169 52 27 78 36 104 163 130 104 156 16 182 33 0 = 0 Irder; S= S i )ecim 124 2.3.2.4.Phylogeneti c analysis From the morphometric analysis, it is clear that not all apriori groups are sufficiently distinguishable,bu tbot hC. ispahanicaan dC. graveolens appea rt ob eseparat egroups .T o a lesser extent, the same holds for C.serratifolia an d C. orientalis.Th e rest group is invariablymutuall yintermixe dan doccasionall yintermixe dwit hth eabov especies .Thi sleave s us with rest groups of parts of C. akebioides,C. glauca, C.hilariae, C.tangutica, C. tibetanaan dC. vernayi,o fwhic hth erelatednes st oC. orientalis ha sbee nvariedl ystresse d or neglected. Phylogeneticanalysi sha sbee ncarrie dou tt oexamin eth erelatio nbetwee nOTU' si n viewo fpolarit yo fcharacte rscores .C. ispahanicaha sbee npostulate da soutgrou pb yit s characterstate s"perennia lo rsubshrab "an d"multi-flowere dbot haxillar yan dtermina lcymes , formingthyreoi dsynflorescences" .Implici tenumeratio nresulte di n1 0equall yparsimoniou s cladogramswhic hwer efurthe rreduce dt oon ecladogra mafte rsuccesiv eweightin gaccordin g toFarri s(1988) .Th edat amatri xi spresente di ntabl e2.7. ,th eresultin gcladogra mi nfigur e 2.9. and thecorrespondin g ancestral nodes with their character states intabl e2.7 . =0 C. ispahanica \=13j 3 C. orientalis ir=- =4 C. intricata I 12! i, 1 C. graveolens tilHil 1, =2 C. serratifolia 11=1(r i i. 6C . tibetana subsp. tibetana IL=9| 1=5 c. tibetana subsp. tangutica IL=8=!i=7 c. tibetana subsp. vernayi Figure2.9 .Cladogra mo fClematis sect .Meclatis; th eonl yon eafte rsuccessiv eweightin g according toFarri s (1988).Fo rcharacte r states atancestra l nodes, seetabl e2.7 . 125 Table2.7 .Dat amatri xo fClematis sect .Meclatis, a suse dfo rphylogeneti canalysis ,se efigur e 2.9. Taxa 0 1 2 3 4 5 6 7 •Character i i 7 •> 7 7 7 7 7 •> n i n 1 1 1 1 1 4 n i 1 1 1 1 < i T. s 1 1 T 1 (. n , n 1 , 1 , , 1 s 1 i 1 4 1 S 7 * a 4 s A i. 1 7 a 0 7 1 7 ID i i 7 1 1 1 i \r n n n n n n w n n 1 n i , id n , I 1 i , , , ti ? i i 7 i i i •» i« , i I 1 i 7 I 7 17 , 7 7 7 7 7 7 7 i« , 1 i 1 7 1 1 1 M 7 1 i 1 1 1 11 1 11 i 1 1 1 1 1 1 w i 1 7 7 •i 7 7 « n 7 7 7 7 7 7 7 7J n n n 1 n n n n « s i 7 4 i i 7 7* n n n (1 n n n n 1 t i 1 i i T i •)» 7 i l 1 l i 1 VI 1 i i 1 n n i ii n n n 1 n n n n u n 7 7 7 7 7 7 7 « 1 n 1 1 1 « i i 1 , 7 1 jn i i n , 1 n n 0 41 7. 1 l '• 2 ?• '• '• Legend:Characte rnumber san dstat ecoding ,se etabl e2.1 .Characte r3 ha sbee nlef tou tth e analysisbecaus eo fto oman ymissin gvalues ;character s 11,18,32,33,36,37 and3 9ar e excluded for showingn ovariation . 126 Table 2.8. Character states atancestra l nodes ofth ecladogra m offigure 2.9 . Ancestors• 8 9 10 11 12 13 *Character s 1 2 2 2 2 2 2 2 1 1 1 1 1 1 4 1 1 1 1 1 1 5 3 3 3 3 3 3 6 1 1 1 1 1 1 7 3 3 3 3 34 34 8 34 34 4 4 4 4 9 1 1 1 1 1 2 10 3 3 3 3 3 3 12 0 0 0 0 0 0 13 1 1 01 01 01 01 14 1 1 1 1 1 1 15 3 3 3 3 3 2 16 2 1 1 1 1 1 17 2 2 2 2 2 2 19 3 12 12 12 12 1 20 1 1 1 1 1 1 21 3 3 1 1 1 1 22 2 2 2 2 2 1 23 2 2 2 2 2 2 24 0 0 0 0 0 0 25 2 2 2 2 3 4 26 0 0 0 0 0 0 27 3 3 3 3 3 3 29 1 1 1 1 1 1 30 1 1 1 1 1 1 31 0 0 0 0 0 0 34 2 2 2 2 2 2 35 1 1 1 1 1 1 38 1 1 1 1 1 1 40 0 0 0 1 1 1 41 2 2 2 2 2 2 Legend: Characternumber s andstat ecoding , seetabl e 2.1.; characterweightin g (char. • weight): 1-10 ; 2- 0;3 -10 ; 4- 3 ;5 - 0 ;6 - 0 ;7 - 0 ; 8-10 ; 9- 0 ; 10 -10; 11 -0 ; 12 -0 13 -10; 14 -10; 15 -10; 16 -10; 17 -10; 18 -1; 19 -10; 20-10 ; 21 - 3; 22-10 ; 23 -10 24- 4 ; 25 -10; 26-10 ; 27-10 ; 28-10 ; 29- 0 ; 30 -10; 31 -10; 32-10 ; 33-10 ; 34 -10 35 -10; 36-10 ; 37 -10; 38-10 ; 39-10 ; 40 -0 . Character3 ha s beenlef tou tth e analysisbecaus eo fto o manymissin gvalues ;character s 11, 18, 32,33 ,36 ,3 7an d 39ar eexclude dfo r showing no variation. 127 2.3.2.5. Conclusions Thenumerica lanalysi salon ema ysuppor tth evie wt olum pth eres tgrou ptogethe rint oon e aggregatespecies .Inconsisten tassignmen tt oth e6 a priori group si sfurthe revidence .Th e phylogeneticanalysis ,however ,show stha tth eresul twoul dthe nb ea polyphyleti cgrou po rth e lumpingo fC. graveolensan dC. serratifoliaint oth eres tgroup .Non eo fthes eoption si s acceptable,a sbot hsolution s willresul ti nth eorigina lsituation :on eo rtw over yvariabl e species with implicit discontinuities. Ifw eals oconside rth egeographi cdistributio npattern so fspecimen s(se efigur e2.1) , theclassificatio nint osi xspecie si ssupporte db yth ecombinatio no fth eminimu mspannin gtree , thedistributio ndat aan dth ecladogram .Furthermore ,th eabov ecombinatio nsupport sth e mergero fC. tangutica,C. tibetanaan dC. vernayia son especies .A sthe yhav ei ncommo n theirpreferenc et ohighe raltitude si nmontan eregion san dsho wa tth esam etim ethei row n geographicdistribution ,the yar emaintaine da tth esubspecifi cran kfollowin gcommo npractic e ast osubspecie s(Meikle ,1957;Fuchs ,1958 ;cf .Hamilto n&Reichards ,1992) .Th eresultin g classification for the sectionMeclatis is: Clematis orientalis L. Clematis graveolens Lindl. Clematis intricataBung e Clematis ispahanica Boiss. Clematis serratifolia Rehder Clematis tibetanaKuntz e subsp. tibetana subsp.tangutica (Maxim. ) Brandenburg subsp.vernayi (C.E.C.Fisch. )Grey-Wilso n The apriori assumed species C.akebioides an d C. hilariae were scattered throughout dendrogramsan dth eminimu mspannin gtree .Thei rdiscriminatin gcharacter sappeare dt ob e highlyvariable , theirdescriptio nbein gbase do na smal lse to f specimensfro monl ya fe w locations.The yar etherefor eno tmaintaine da tan yrank .C. akebioides i slargel yclassifie d underC. tibetanasubsp .tibetana an dsubsp .tangutica; C. hilariaeunde rC. intricata.Th e nomenclatural consequences willb edeal t with in section 2.4. 128 2.4. Species descriptions 2.4.1. Key toth especie so f Clematis sect.Meclatis la Plant habit shrubby orherbaceou s C.ispahanica 2.4.5. lb Plant habit woody climbers 2 2a Filaments ciliate C. graveolens2.4.3 . 2b Filaments dilatate 3 3a Leaves commonlybiternate , leaflets regularly serrate/dentate C.serratifolia 2.4.6 . 3b Leaflets variablyi n shape,no tregularl y incised 4 4a Inflorescences (l-)3-7 flowers, tepals laterrecurve d C.orientalis 2.4.2. 4b Inflorescences 1-3 flowers, tepals later spreading 5 5a Inflorescences usually with 3open ,fla t flowers, tepals herbaceous C.intricata 2.4.4 . 5b Inflorescences with 1-3 broadlyt onarrowl ycampanulat eflower s 6 6a Inflorescences 3flowers , narrowly campanulate, tepals notflesh y C.tibetana subsp .tibetana 2.4.7 . 6b Inflorescences l(-3) flowers, narrowly campanulate, tepals thin,flesh y C.tibetana subsp .tangutica 2.4.7 . 6c Inflorescences l(-3) flowers, broadly campanulate,tepal s thick, fleshy C.tibetana subsp .vernayi 2.4.7 . 129 2.4.2. Clematis orientalisL. 1 2.4.2.1. Lectotvpification of Clematis orientalis L. The name C.orientalis date s back to the first edition of Linnaeus'Species Plantarum (Linnaeus,1753:543) .Th eprotologu econsist so fLinnaeus'diagnosti cphrase-name ,tw o earlierpolynomial scite di nsynonym yan da nindicatio no fth etyp elocality .Th eLinnaea n diagnosis"Clematis foliis compositis: foliolis incisis angulatis lobatis cuneiformibus" i sno t taken from anyo f hisearlie r works. Inth eLinnaea nHerbariu m(LINN) ,on especime nha sbee nassigne db yLinnaeu st o C. orientalis(LIN N712.7) .Bot hannotation so nthi sshee t"orientalis "an d- o nth evers o- "Clematisorientali sapi ifoli ofl .reflex oT.C." ,hav ebee nmad eb yLinnaeus .Whe nusin g specimensfo rdescription si nSpecies Plantarum ed .I ,Linnaeu salmos tinvariabl yindicate d onth esheet sth enumbe runde rwhic hh edescribe dth especies .Suc ha referenc ei sabsen ti n theannotation so nLIN N712.7 .I twa stherefor ealmos tcertainl yadde dt ohi sherbariu mafte r 1753.Consequently ,i tcanno tb eregarde da sa syntype ,no ra sa putativ echoic eo flectotype . Inth eLinnaea nHerbariu ma tStockholm ,ther ei son especime nunde rth enam eC. orientalis (IDCfich e no.22 45) .O nth evers oo fthi sshee tther ei sa nannotatio n "Dahla Linn eP "i n Dahl'shandwriting .Thi sspecime nwa sapparentl yreceive db yAnder sDah lfro mLinnaeus . However,th eshee ti sunannotate db yLinnaeu san dther ei sn oevidenc etha th eregarde d ita s belongingt othi staxo no rtha th eha dstudie di tbefor e publishinghi sSpecies Plantarum account.It ,therefore ,i sno ta syntyp ean dmus tb eexclude dfro mconsideratio na sa possibl e lectotype.Remarkably , two specimens of C. orientalis arepresen t in the Hortus Siccus Cliffortianus (BM),althoug hLinnaeu sdi dno tdescrib eth especie si nHortus Cliffortianus (1738).Thes especimen smus teithe rhav ebee nadde dt oClifford' s Herbariumafte rLinnaeu s studiedit ,o rh ema yhav eoverlooke dthes especimen swhe nwritin ghi sHortus Cliffortianus account.W ehav eno tbee nabl et olocat ean yothe rrelevan tspecimen si nan yo fth eothe r Theparagraph s 2.4.2.1., 2.4.2.2.an d 2.4.2.3.hav e been largelybase d onth eearlie r publication byBrandenbur g et al.(1987) . 130 general Linnaeanherbari a (i.e.BM ,H ,MW ,SBT ,UPS) .Th efirst synony mcite di nth e protologue of C. orientalisi s "Clematis orientalis, apiifolio, flore e viridi-flavescente posteriusreflexo", a sa slightl yaltere dphrase-nam efro mth eCorollarium Institutionem b y Tournefort (1703):"Clematitis Orientalis,Apii folio, flore eviridiflavescente, posterius reflexo".I n17 38 ,Linnaeu svisite dParis .H ema ypossibl yhav esee nth efou rspecimen so fC . orientalisi nTournefort' sHerbariu m(P-TO) .Ther eis ,however ,n opositiv eevidenc etha th e thoroughly studied Tournefort's Herbarium (Stearn, 1957)an d sow eexclud ethes e four specimens from consideration for lectotypification. Thesecon dsynony mcite dunde rC. orientalisi s "Flammulascandens, apiifolio glauco"fro mth eHortusElthamensis o fDilleniu s(1732) .Thi sunaltere dquotatio nrefer st o aphrase-name,a descriptio nan da nillustration .Linnaeus'ow ncop yo fthi sbook ,whic hh e usedi npreparin gSpecies Plantarum, i sno wa tJen a(JE )havin gbee nsol db ySmit h(Schmidt , 1965).Tw obeautifull ycoloure dcopies ,execute db yDilleniu shimself ,ar ea tOxfor d(OXF ) anda tLondo n(BM) .Sinc ether ei sn opositiv eevidenc etha tLinnaeu srelie do nspecimen sstil l inexistenc efo rhi sdescriptio n ofC. orientalis, theplat e"Flammula scandens, apiifolio glauco", Dill.Elth . 144,1.119,f . 145(1732 )i shereb ydesignate d asth electotyp e ofC. orientalis (figure 2.10). 131 P.144- T. CAR. ^2amnui/a ^capidenj/feJora^iuq/auca. Figure 2.10. Dillenius Hortus Elthamensis 144,t . 119, f. 145 (1732). Lectotype of C. orientalisL . 132 Figure2.11 .Dillenia nspecime n286 8(OXF) .Typotyp eo fC. orientalisL .(photograp hb y courtesyo fOXF) . 133 2.4.2.2. TheTypotyp e of Clematis orientalis L. Ifth echoic eo fa nillustratio na slectotyp ei sinevitable ,i tca nsometime sb efurthe r supported byth eexistenc e of atypotype , i.e.a specime n onwhic h theillustratio n has been based. Dilleniusdescribe di nhi sHortus Elthamensis plant swhic hwer egrow ni nth egarde no fJame s Sherarda tEltham .A tth esam etime ,h ecollecte dspecimen sfro mthi sgarden .Thi scollectio n formedth enucleu so fth eDillenia nHerbariu m(OXF )whic hwa soriginall yscattere dthroughou t theSherardia n Herbarium,itsel fbuil tu pi nth eperio d 1680-1790.Druc eseparate d both collections (Druce &Vines , 1907). Therear etw ospecimen so fC. orientalis i nth eDillenia nHerbariu m(286 8an d2868 2), thefirs to fwhic hcorrespond sremarkabl ywel lwit hDillenius'illustration ,bein gmirror-images , apartfro m oneo fth ebasa lnodes .W esurmise dtha tth eanomalou snod emigh tb edu et oa twistingo fth este mi nth eregio no fth enod ean dw ear ever ygratefu l toDr .D .Mabberle y (pers.comm . 1986)fo rkindl yconfirmin g thatthi si sth ecase .Specime n286 8show sth e following annotations (figure 2.11): '119. 145.144'(topright), 'Clematis orientalis L." (bottomright ) inth ehandwritin g of G.C.Druce , 'Europe' (bottomright, stamped) , 'Herb. Sherard' (bottomright) i nth e handwriting of John Sibthorp, '2868' (bottomright) i n thehandwritin g of William Baxter. Apinne d label atth ebotto mlef t shows usth e following: '2868' (redink) , 'Clematideorient .Api ifolio ,fl .e virid iflavescente ,posteriu sreflexo ,Coroll .20 ' (brownink) , 'Flammulascandens .Api ifoi oglauc oHort .Elth. 'wit ha ninitia l'D 'o rv O'i nth esam e handwriting (blackink) . 'fig 145orientalis' (pencil). Thelabe lshow su sfou rdifferen thandwriting so fwhic hth eon ei npenci li srecent .Th enumbe r 2868i sagai ni nWillia mBaxter' shandwriting .Th eTournefor tphrase-nam ei nbrow nin ki s neitherb yJame sno rWillia mSherard ,no rb yDillenius .I ti sver ysimila rt oth ehandwritin go n oneo fth efou rspecimen so fC. orientalis i nth eD eJussie uHerbariu m(P-JU10493) .I ti sno t byAntoin eo rBernar dd eJussieu ,bu tresemble sth ehandwritin go fSebastie nVaillan t(1669 - 1722),wh olive di nPari s(Heine ,pers .comm. ,1986) .W ehav eno tbee nabl et oidentif yth e 134 handwritingi nblac kink .Becaus eo fth estrikin gresemblanc ebetwee nVaillant' shandwritin g andth ebrow nin khandwriting ,connection sbetwee nth eSherard san dParisia nbotanist sshoul d beconsidered .Willia m Sherard(1658-1728 )aske dDilleniu si n 1721t ocom eove r from Germanyan dt owor kfo rhi mi nOxfor darrangin gth eSherardia nHerbariu man dpreparin ga n improvedversio no fBauhin' sPinax (whic hwa sneve rfinished) .Afte rWilliam' sdeath ,hi s brotherJame saske dDilleniu st owrit ea boo ko nth eplant si nhi sgarden ,whic hwa spublishe d in173 2a sHortus Elthamensis. Willia mSherar dwa sEnglis hconsu la tSmyrn a(Izmir )fro m 1703t o1717 .H etravelle dextensivel yi nEurope ,an dwa so ngoo dterm swit hmos tcontem porarybotanists .Hi scorrespondenc ewit hthe mha sbee npreserve di nth elibrar yo fth eRoya l Societyi nLondon .I tclearl yproves ,tha tther ewa sa ttha ttime quit ea nexchang eo fbook san d plants.Correspondenc ewit hVaillan t (MSS.Sh .528-545 )doe sno tshe dan yligh to nth e Tournefortphrase-nam ei n- possibl y- Vaillant' shandwritin ga si toccur so nth elabe lo fshee t 2868. Accordingt oPast i(1950) ,Tournefor t hadpromise dSherar dspecimen sfro m the Levant for his Pinax.Afte r learning that Sherard would got o Smyrna, Tournefort also promisedt ogiv ehi mal lne wplant sfro mth ePari sBotanica lGarden .However ,Tournefor t died,befor eh ewa sabl et ofulfil lhi spromises .Returnin gfro mSmyrna ,Sherar dstaye di nParis , whereh ehope dt oreceiv eth epromise dplants ,bu th ewa sno tallowe dt otak ethe mwit hhi m toOxford . Hewa sagai n deniedpossessio n ofthos eplant so nanothe roccasio n in 1722. Sherardthe n spent sixweek st ostud yTournefort' sLevan t specimens.Howeve rth e four specimensi nP-T Od ono tsho wSherard' shandwritin gan ds ow ehav eno tbee nabl et osho w directconnectio nbetwee nth eVaillan tlabe li nOxfor dan dVaillant' sspecimen si nP .Accordin g toCloki e(1964) ,ther ear eTournefor t specimensi nth eSherardia nHerbarium ,s oanyho w someexchange ,an dperhap sincludin gspecimen sfro mVaillant ,occurre dbetwee nPari san d Oxford. Fromcorrespondenc ebetwee n Sherardan dD eJussie u (MSS.Sh .276-292) ,w e knowtha tD eJussie udesire dplant sfro mEnglan di nexchang efo rSpanis hplant san dbook s oncoins .I tis ,therefore ,possibl etha tTournefor tan dVaillan tmaterial ,th elatte rformin gth e nucleus of Paris collections,coul dhav ebee n sent to Oxford. Afterhavin gbee nseparate dfro mth eSherardia nspecimens ,th eDillenia nsheet shav e beenremounte d(Druc e& Vines , 1907).I ti simportan tt obea rthi si nmin dwhe nstudyin g 135 these early specimens.Th efollowin g four categories of specimens should be distinguished: specimenslabelle db yDilleniu sand/o rHumphre ySibthorp ,whic hfor mth ebasi sfo r theillustration s inHortus Elthamensis; rathershrivelle dspecimens ,labelle db yDilleniu s(bearin gphrase-name sfro mth e HortusElthamensis commonly with theannotatio n "duplicate"); specimens from James Sherard's garden added after publication of the Hortus Elthamensis in 1732; specimens not originating from thegarde n ofJame s Sherard atEltham . Consideringth eresemblanc ebetwee nillustratio nan dspecimen ,shee t286 8apparentl yha st o belongt oth efirs t categorydespit eo fth eabsenc eo fa Dillenia n label.Th eoccurrenc eo f Vaillant'shandwritin go nshee t286 8coul db eexplaine dthroug hinterchang eo flabel safte r remounting. Theillustratio ni nHortus Elthamensis differ sfro mshee t286 8i ntha tfruitin gbranchlet s weredepicte dseparatel yfro mth eplant .Thes ecoul dconceivabl yhav ebee ndepicte dfro m sheet2868 2,bu tther ei sn odirec tevidenc efo rthi sa sthi sshee tonl yshow srecen thandwriting . Becauseo fth estrikin gresemblanc ebetwee nth edepicte dplan tan dshee t2868 ,a ninterchang e oflabel s wasassume dan dth eDillenia nspecime n286 8wa sconsequentl ydesignate da sth e typotypeo f "Flammulascandens, apiifolio glauco",Dill .Elth . 144,1.119,f . 145 (figure 2.11; originalreference : Brandenburg et al., 1987). 2.4.2.3. Thetyp e locality of Clematis orientalis L. TheICB N(1988 )doe sno tstat eanythin gabou ttyp elocalities .Suc hdetai li sno trequire db y therules ,bu ti squit einterestin gi nit sow nright. I na narro w senseth etyp elocalit yo fC . orientalisshoul d beJame s Sherard's garden atEltham .I ncas eo f acultige n this isver y plausible.However ,C. orientalis i sa specie soccurrin gi nth ewild .I tis ,therefore ,worthwhil e tracingth enatura lprovenanc eo fth etypotyp emateria lan di ti smeaningfu l toindicat ethi s provenance astyp elocality . From 1700unti l 1702Tournefor t travelled inth eLevan tb yorde r of theKin go f France.Accordin gt oBecke r(1957) ,h ecollecte dplant si nth esurrounding so fTrebizond e (Trabzon),th esurrounding so fErzurum ,th esource so fth eEuphrates ,Moun tOlympu snea r Brousse(Bursa )an dMoun tSypilu snea rSmyrn a(Izmir) .Supplementar yt oth eInstitutiones 136 Rei Herbariae (1700) he published in 1703 his Corollarium in which he mentioned: "ClematitisOrientalis, Apii folio, flore eviridiflavescente, posterius reflexo". Thi si sth e earliestrecor do fC. orientalis,a sfa ra sno wknown .Tournefor t describedan dsometime s depictedplant scollecte ddurin ghi sjourne yi nhi sboo kRelation d 'un Voyagedu Levant, publishedposthumousl yi n1717 .H eneithe rdescribe dno rdepicte dC. orientalis,bu tstate d thath esen tsom eseed sunfortunatel yunname dt ofriend si nth eNetherland s(Wijnands ,1983) . Miller(1768 )mentione dtha tmos tbotanica lgarden swoul dhav ebee nprovide dwit hseed s fromth eHortu si nParis .I nvie wo fSherard' sgoo dconnection swit hTournefor t andothe r botanistsi nEurope ,i ti slikel ytha tth eC. orientalisplant si nJame sSherard' sgarde na tEltha m stemmeddirectl yo rindirectl yfro mplan tmateria lintroduce db yTournefort .I nP-T Other ear e fourspecimen sassigne dt oC. orientalis, al lo fwhic hwel lresembl eth etw ospecimen si nth e DillenianHerbarium .On eo fthes especimen s(P-T O2520) ,i nth egenera lpar to fTournefort' s Herbarium, is labelled: "ClematisArmenia, Apii folio". The remaining three are in the supplement under nr. 30 pag. 20,whic h refers to the thirtiest species onpag e 20i n the Corollarium.On eo fthe mshow sn oannotation .Th efirs to fth eothe rtw obears "Clematitis oriental. Apiifolio glaucoflore fl. viridipetali reflexo Cor.Inst." (beneat h in another handwriting "C orientalisL.") ,an dth e secondbear s "Clematis orientalis apiifol.fl. ex viridiflavescente posterius reflexo T.Cor". The handwriting on the specimens in the supplement,apar tfro mth enote "C. orientalis L. "i sTournefort's ,a si stha to nP-T O252 0(Dr . H.Hein epers .comm .1986) .Th elatte rspecime nhoweve rcontain sa nindicatio nwher ei twa s found:Armenia .I nth eabsenc eo fan yothe rindication swher eTournefor tfoun dC. orientalis, weconside r'Armenia 'o ftha ttime i nrelatio nt oTournefort' srout eo ftrave li.e .th esource so f theEuphrate san dth esurrounding so fErzuru ma sth etyp elocalit yo fC. orientalis(figur e 2.12). 137 VA 45,., ~A^J ^fVl ~$£\ / \*-^C ^"^ Vr \ 1 \\ \M ^ v\0' \ 40, \ s***'**"'—\ \ \ ^••'si^Z- *jv'- T ^\—~~xp^ , \ Trebizonae/ ~*l—\lr**'"\ o 1 ^T^ TO Smy sy~^?5> YJ/7'\ ( ^- Ij-N^ \TP SCALE 200 300 400 500 0 5OO MILES X*^/j /3» •«> I! *0 20O «oo eoo eoo KIL OMETERS .*.^> CONIC PROJECTION 85 4\5 ^®f% —^Z.—" \ Figure 2.12.Ma po f type locality of C. orientalisL . Route takenb y Tournefort. 138 2.4.2.4. Synonymy anddescriptio n of Clematis orientalis L. Cf. Meyer (1831);Miille r(1857) ; Synonymy: Homotvpic svnonvm: Meclatis orientalis (L.) Spach Heterotypic synonyms: Clematislongecaudatahedeb.,inFlomrossica,\ol. 1 (1842)Clematideae,l-5(Lectotyp e Claus, s.n., Desert Casp., 1834B ;isotyp ea tLE) . Clematisalbida Klotsch ,i nDi ebotanische nErgebniss ede rReis eseine rKonigl .Hohei tde s PrinzenWaldema rvo nPreusse nau fCeylon ,de nHimalay aun da nde nGrenze nvo n Tibet gesammelte Pflanzen (1862), 1311.4(holotyp eHoffmeiste r s.n., B). Clematis orientalisL . subsp.orientalis var.flava (Moench) O.Kuntz e Clematis flava Moench, in Methodus plantas horti botanici et agri marburgensis a staminum situ describendi (1794;typ eno tseen) . Clematis orientalis L. subsp. wightiana(Wall. ) O. Kuntze var.longecaudata (Ledeb.) O.Kuntze . Clematisorientalis L .subsp .orientalis var .normalis an dvar .albida (Klotsch )O . Kuntze 'vulgaris', 'angustifolia'an d 'fasciculata' ClematisorientalisL. var .hindukushensis Grey-Wilson ,i nKe wBulleti n4 4(1989) :33-6 0 (Type Grey-Wilson and Hewer 1224,Afghanistan , Djam (Ghowr);holotyp e K, isotypeW) . ClematisorientalisL. var.robusta Grey-Wilson ,i nKe wBulleti n44(1989) :33-6 0(Typ e Grey-Wilsonan dHewe r1688 ,Afghanistan ,betwee nUrgu nan dQazide h(Badakshan - Wakhan);holotyp e K,isotyp eW) . Clematisorientalis L .var .baluchistanica Grey-Wilson ,i nKe wBulleti n4 4(1989) :33-6 0 (TypeCrookshan k373 ,Pakistan ,Uran g(Bahawalgawar) ;holotyp ean disotype sK) . Clematisorientalis L .var .tenuifolia (Royle )Grey-Wilson ,i nKe wBulleti n4 4(1989) : 33- 60. Clematistenuifolia Royle ,i nIllustration so fth eBotan yan dothe rbranche so fth e NaturalHistor yo fth eHimalaya nMountain san dth eFlor ao fCashmere ,vol . 1 (1839) Ranunculaceae, 43-51 (lectotype Royle s.n., LJV,se e Lauener, 1978). Clematis orientalis L.var . latifolia Hook.f. &Thomso n (lectotype Royle s.n., LIV) Clematisglobosa Royle . Clematis orientalis L. var.globosa (Royle ) Mukerjee. Clematis orientalis L. var.daurica (Pers. ) O.Kuntze . Clematisdaurica Pers. ,i nSynopsi splantaru mse uenchiridiu mbotanicum ,vol . 1 1358Clematis, 98-100.Pari s (Holotype Patrin 10502,P-JU) . Clematis dauricaLedeb .nom .illeg . Clematisglauca Willd., in Berlinische Baumzucht 65 (1796): 89-94, t.4, fig 1 (holotypeWilldeno w 10474,plan tcultivate di nth eBotanica lGarde nBerlin , B-WILLD.) Clematis orientalis L. var. obtusifolia sensu Trautv., nonHook.f . &Thomson . 139 Clematisorientalis L .subsp .orientalis var .daurica (Pers. )O .Kuntz e 'persoonii' Clematis orientalis Krylov,no nL . Clematis orientalis L.var . obtusifolia Hook.f. &Thomso n f. oblongifolia Regel. Illustration, seefigur e 2.13. Description: Woodyclimber , up to 8mtall , flowering in summer; (June-)July-August(- September).Branchlet sangula rt ocostate ,sparsel ypilose .Leave sver yvariabl ei nshap ean d size,imparipinnat ewit h(3-)5-7(-9 )leaflets ;petiole s2-1 0cmlong ;leaflet soblong ,lanceolate , elliptico rovat et obroadl yellipti co rbroadl yovate ,herbaceous ,greyis hgreen ,glabrou so r sparselypilose ,3-6 5x 5-5 0mm ,1 -1 0x a slon ga swide ,entire ,lobed ,clef talmos tt oth ebas e in1 mainpar tan d1 or2 smalle rlatera lparts ,o rcompose do f3 leaflet so fsecon dorder ,th e partsmostl ywit ha fe wteeth ,acut eo racuminat ea tth eapex ,mor eo rles sabruptl ynarrowe d andcuneat ea tth ebase ;petiolule spilose ,5-3 0m mlong .Inflorescence stermina lan daxillary , being3-many-flowere dcyme sonl yo nbranchlets ,apica lflowers o fcyme ssometime saborte d orabsent ;peduncl e5-8 0m mlong ;pedicel spilose ,5-5 0m mlong ,curve da tth eapex ,bu t elongate and straight whenfruiting . Flower buds often darkred ,ovoi dwit h acuteapex ; aestivationinduplicate .Flower sactinomorphic ,hemicyclic ;tepal s4 ,yellow ,pal eo rgreenis h yellow,sometime s lightpurplis hbrow ninsid eo rtinge dwit hred-viole toutside ,oblon go r elliptic,11-15(-20 )x 4-7. 5mm ,2- 4x a slon ga swide ,acut ea tth eape xwit hbroadl ycuneat e basestouchin geac hother ,lanat ea tth eincurve dmargin ,les slanat ewithi ntha na tth emargin , sparselypilos e to villose outside, spreading, recurved later. Stamens numerous, 20-40; filamentsdilatate ,yellow ,mostl ydar kred-purple ,toward sth ebas epilose ,u pt o1. 5c mlong ; anthers yellow,u pt o0. 5 cmlong .Pistil s numerous,5-1 1m mlong ;ovarie sellipsoi d or rhomboid,pubescent ,+ 1 m mlong ;style slong-pubescent ,4-1 0m mlong ;stigma sstraigh to r slightlyhooked ,glabrous .Achene srhomboid ,slightl yribbed a tth emargin ,pubescent ,dar k brown,2-4x1- 2mm ;elongat epersisten tstyles ,25-5 5m mlong ,covere dwit hlon gerec thairs . 140 141 142 Q'abM Figure2.13. a.C . on'e«ta/wL.(drawingbyMarietdeGeus;1 .Par to fflowerin gbranch ,0.7x ;2: flower, 1.3x; 3.tepal , 1.5x; 4. stamen,4.7x ; 5.pistil ,4.7x ; 6.frui t head,0.7x ; 7.achene ,2x ;8 . longitudinalsectio no fachene ,5.3 x (1,3,K.P .an dE .Buttle r2028 4M ;2,4,5 ,K.P .an dE . Buttler 20053M ;6 , 7, 8,K.M . Guichard T/127/60BM) . b. C.orientalis L. (drawing byMarie t deGeus ;pop .78018 ;Br a 185,186WAG ) c. C.glauca Willd . (Willdenow, 1796),synony m of C. orientalis L. 143 Distribution: Russian Federation (Khabarovskiy Kray, Buryetskaya ASSR, Dagestan AutonomousRegion) ,Kazakhstan ,Turkmenistan ,Uzbekistan ,Chin a(Xinjian gUygu rZiziqu) , India (Himachal Pradesh), India/Pakistan (Jammu and Kashmir), Armenia, Georgia, Azerbaidjian,Pakistan ,Afghanistan ,Iran ,Turkey ,N-Syria ,Easter nAegea nIsle s(se efigur e 2.14). Specimens examined: USSR: KhabarovskiyKray: Khrebet Ket-Kap, E. Serova & E. Ryschowa2 3(NY) ;Buryetskaya ASSR: Dka ,Schren ks.n .(U) ;Kazakhstan: Alakol ,Schren k s.n.(K ,M ,U) ;Lepsa ,Kareli n& Kirilof f 1130(G ,K ,M ,NY) ;Hi ,A .Rege ls.n. , 1876(M) ; ZailitskiyAlatau ,A.K .Skortso vs.n. , 15-9-1963(M) ;Khrebe tKaratau ,N.V .Pavlo v114 8 (B);Khrebe tChatkalskiy ,A.K .Skortsov ,s.n. ,3-10-196 3(M) ;Dzhambul ,J .Raikov as.n. , 1917(C ,G ,K ,MO ,NY ,S) ;Uzbekistan: Shimgan,Barano v& Raikov as.n. ,23-8-192 4(B , G, K,MO ,NY , S), Fedchenko s.n., 19-8-1897 (G),L.P .Velikano v s.n., 22-9-1962(C) ; Chircik,M .Capu s2 an d3 (P) ; Syr-DarvinskayaOblast ,A .Michelso n29 5(S) ;Karankul , O.N. Korowina & N.M. Oernomorskaya s.n., 9-8-1976 (G, LD, M); Tadjikistan: Kafirnagan,Newissk y466 0(M);Khrebe tGissarskiy ,P.N .Ovczinniko v208 8(NY) ,V .Vasa k & A. Zlatnik s.n., 23-5-1974 (M), D.H. Wilken, R. Hebb & T. Crovello 29 (NY); Turkmenistan: Farab-Pristan, N.Androso v s.n., 2/23-9-1902(C ,G ,K ,M ,MO ,NY ,S) ; Ashkabad,D .Litwino w 23(G ,M) ,E .Rege l81 2(NY) ,P .Sinteni s 848(B ,BM ,G ,K ,L , MO),110 8(G) ;Kazandhik ,P .Sinteni s 1287(WAG) ;Krasnovodsk ,D .Kareli ns.n. , 1834 (G);DagestanskayaASSR: Akhty ,Th .Alexeenk os.n. ,5/17-8-189 8 (B,G);Azerbaidzhan :Kirovabad ,Fedoseeffs.n. ,Aug .1899(B),R.F.Hohenackers.n. ,1834(G) ,Aug./Sept . 1838 (G,L ,M ,P ,WAG) ;Georgia: Tbilisi,K .Browic zs.n. , 27-7-1968(K) ;Armenia: Khrebet Gegamskiy,V.Vasa k s.n., 12-7-1975(M) . CHINA:Xinjiang UygurZizhiqu: Kumishi,Fedchenk o s.n., 2-8-1897(G) ;Kuerchu ,A . Regels.n. ,25-8-187 7(K) ;Erti xHe ,G.N .Potani ns.n. , 1876(K) ;Yining ,A .Rege ls.n. ,5-7 - 1877 (BR, K, M); Shuiding, A. Regel s.n., 8-7-1877 (K); Suoche, C. Persson 228 (S); Bositan, C.Persso n 522(S) . INDIA:Himachal Pradesh: Spiti ,V .Jacquemon t64 4(K) ;Nahan ,V .Jacquemon t110 7(K) ; Simla,J.H . Lace 530(OXF) . INDIA/PAKISTAN:Jammu and Kashmir: Shigar,W .Koel z965 5(NY) ;Basha ,R . Scott 144 Russell 1785(BM) ;Hunza ,F .Lobbichle r46 9(M) ,O .Poluni n641 4(BM ,G) ;Chalt ,O . Polunin645 3(BM ,G) ,J.E .Winterbotto m93 2(K) ;Gilgit ,G.M .Gile sAl l(G) ;She rQila ,Dr . Giles 519 (K);Imit ,F .Schmi d201 5 (BM,G) . PAKISTAN:Bahawalganar. Urang ,H .Crookshan k37 3(K) ;Chitral: Chitral ,A .A .Barret t 14873(K) ;KillaDrosh ,A.A .Barret t1483 5(K) ,Hamilto n1786 1 (K),S.M.Toppi n59 6(K) ; Kurram: Darband, J.E.T. Aitchison 415 (K,P) . AFGHANISTAN:Badakshan: WakhanCorridor ,H .Roeme r27 4(M) ;Urgun ,C .Grey - Wilson& T.F .Hewe r 1668(K) ;Kunar: Bashga lriver ,D .Podlec h 16638(G ,M) ; Takhar: Farkhar,P .Furs e816 3(K) ;Paktia: Orgun ,O.H .Vol k7 l/754 b(M) ;Baghlan: Ashra friver , H.F.Neubaue r437 4(B) ;Bamian: Shiba rPass ,R .Gibbon s074 7(K ,MO) ;Doabi-Mekh-i - Zarin,P .Furs e828 0(K) ,Griffit h s.n. (CGE);Bamian ,Carte r61 9(K) ; Band-i-Amir ,A . Dieterle75 6(G ,M) ;Surkhoy ,D .Podlec h1887 2(M) ;Ghowr: Djam ,C .Grey-Wilso n& T.F . Hewer 1224 (K); Faryab:Sangilak , M. Capus s.n. (K); Helmand: Lashkari Bazar, G. Frumkin 54(G) . IRAN:Khorassan: Dogharun ,Herb .Bung es.n. ,Aug .185 8(G) ;Mirabad ,Assad i& Masoum i 21266(K) ;Kuh-e-Mish ,K.H .Rechinge rfil. 142 4(BM ,K ,S) ;Sharifabad ,Herb .Bung es.n. , June/July185 8(L) ;Fars: Siva nriver ,J .Bornmulle r198 0(K) ;Mazandaran: Elburz ,Gaub a 19(B) ;Amol ,P .Furs e906 3(K) ;Esfahan ,Aucher-Elo y402 5 (B,P) ,402 9(G) ,A.C .Trot t 783(K) ;Lorestan: Bisheh ,M .K0i e73 5(C) ;Kermanshah: Kermanshah,C .Haussknech t s.n., Sept. 1867(BM) . TURKEY:Kars: Igdir ,P .Furs e912 3(K) ;Hakkari: Dize ,P.H .Davi s& O .Poluni n2402 2 (BM);Artvin: Artvin ,W .Andronak i s.n., 11/24-7-1907(K ,LD) , 18-7-1907(LD) , 20-7- 1907(G) ,Jul y 1907(WAG) ;Erzurum: Erzurum, H.H. Calvert &J . Zohrab 753(CGE , OXF);Soylemez ,Frase rJenkin s 2408 (BM);Tortum ,Stainto n &Henderso n 6155(K) ; Gumusane:Gumusane ,E.K .Ball s& W.B .Gourla y198 4(BM ,K) ,P .Sinteni s711 7(LD) ; Erzincan:Tanyeri ,K.P .Buttle r& R .vo nBottme r2265 5(M) ;Erzincan ,P .Sinteni s299 8(BR , G,K ,P) ;Eldzig: Elazig,K.P .& E .Buttle r2028 4(M) ;Tokat: Tokat,P .Furs e918 3(K) ; Samsun:Bafra ,P.H .Davi s& O .Poluni n2496 0(BM ,K) ;Amasya: Amasya ,Manissadjia n 639(B ,K ,LD) ,J .Bornmulle r 1318(BM ,BR ,K ,LD ,M ,S) ; Kayseri: Urgiip,W .Kotte s s.n.,28-7-193 3(M) ;Nevsehir: Nar,G .Rope r8 6(BM) ;Avcilar ,K.P .& E .Buttle r2005 3 145 (M);Kastamonu: Kosen,P .Sinteni s488 1(B ,LD ,M) ;Ankara: Kalecik,K.M .Guichar d T/127/60(BM) . SYRIA:Halab: Barsa Dagh, C.Haussknech t s.n., 1865(K) . GREECE: Kos,W .Barbe y 602(G) . Figure 2.14. Distribution of C. orientalis L. 146 2.4.3. Clematis graveolens Lindley Clematisgraveolens Lindley ,i nJourna lo fth eHorticultura lSociet y(London )(1846) :307 - 308. Holotype:Munro , s.n. Chinese Tartary (CGE-LindleyHerb.) . Synonymy: Homotypic synonym: Clematisorientalis L. subsp.graveolens (Lindl. )O .Kuntz ei nMonographi cde rGattun g Clematis. In:I .Urba ne tal .(eds. )- Verhandlunge n desBotanische n Vereinsde r Provinz Brandenburg (1885), 83-202.Berli n Heterotypic svnonvms: Clematisorientalis L .subsp .graveolens (Lindl. )O .Kuntz evar .aitchisonii O .Kuntz ei n Monographicde rGattun gClematis. In : I.Urba ne tal .(eds. )- Verhandlunge nde s BotanischenVerein sde rProvin zBrandenbur g(1885) ,83-202 .Berli n(typ eAitchiso n 614, 718,Pakistan , Darband (Kurram);holotyp e B,isotype s BM,H-W , K,P) . Clematisorientalis L . subsp. thunbergii(Steud. ) O. Kuntze var. intricata(Bunge ) O. Kuntze p.p. in Monographic der Gattung Clematis.In : I. Urban et al. (eds.) - Verhandlungende sBotanische nVerein sde rProvin zBrandenbur g(1885) ,83-202 . Berlinpro parte. Clematisparvifolia Edgew.i nTrans .Linn .Soc .2 0(1851) :2 5(Type :Edgewort h 1051, India, Kundau andBea sValleys ;holotyp e K, isotypeOXF) . Description:Wood yclimber ,u pt o4 mtall ,flowerin g insummer ; [(June-)July-August(- September).Branchlet s mostlycostate ,initiall y pilose, laterglabrous .Leave s variously bipinnate,sometime spinnat eo rtripinnat ewit h(15-)25-30(-35 )leaflets ;th epetiole spilos ea t the base, 1.3-6.5 cm long; the leaflets very variable in shape and size, ovate to lanceolate/lineariform,herbaceous ,green ,glabrou so rslighti ypilos ea tth eveins (12- )18-23( - 50)x 6-2 0mm ,1 - 8x a slon ga swide ,irregularl ylobe do rdentate ,acut eo racuminat ea tth e apex, cordate or angustate at the base; the petiolules pilose, (4)-10-13(-30) mm. Inflorescences 1-3(-7)-flowere daxillar ycymes ,i fmor etha n3 flower spe rinflorescenc eth e secondaryflower sno tfull ydeveloped ;peduncl e27-60(-100 )m mlong ;pedicel spilose ,25-8 5 mmlong .Flower sopen ,fla tt obroadl ycampanulate ;tepal s4 ,brigh tyello wo rlim eyellow , ovate-obovate,1 1-17(-20 )m mx 6-1 0mm ,+ twic ea slon ga swide ,blun tt oacuminat ea tth e apex,cuneat et oangustat ebase stouchin geac hother ,tomentos ea tmargins ,slightl ypilos ebot h insidean doutside ,spreading .Stamen snumerous ,(30-)40-60(-70) ;filament sciliate ,greenis h yellow,sometime stinge dwit hviolet ,pilose ,u pt o1 cmlong ;anther syellow ,2- 3m mlong . 147 Pistilsnumerous ,80-100 ;ovarie srhomboid ,pubescent,+ 1 mmlong ;style slon gpubescent , 3-9m mlong ;stigma sstraigh to rslightl yhooked ,glabrous .Achene srhomboid ,markedl yribbe d atth emargin ,slightl ypilos eo rglabrous ,dar kbrown ,2-4x1- 3mm ;elongate ,persisten tstyles , 20-35(-50) mm long covered with longerec thairs . Illustration: See figure 2.15. Distribution:Nepal ,Jamm uan dKashmir ,Indi a(Punjab) ,Pakista n(Peshawa ran dRawalpindi) , Afghanistan (Kandahar). See figure 2.16. Specimensexamined :NEPA LChongnea rTibrikot ,Polunin ,Syke s& William s333 9(BM , E,UPS) ;Dunaih i along Behririver , Shrestna 5266(G) ; INDIA Kundan and Bean Valleys, Sangla, Edgeworth 1051, (K, OXF); Chamba to Darmtawer,Falcone r 6 (K, M);Saraha n (Punjab), Ram Baksh432 3(K) ; JAMMU and KASHMIR, Kishtwar, Clarke 31450B (BM);Garki , Rich 1314(K) ; AFGHANISTAN, KurrumManda h (Kandahar), Aitchison 614,718(BM ,FI-W ,K) ; PAKISTAN(Peshawar) ,Mansehr aHazara ,Duthi e742 3(BM ,K) ,Kalapan iHazara ,Stewar t 27748(G) ,Swa tValley ,Weatherhea d 100(BM) ;(Rawalpindi) ,Kulu ,Larji ,Parke r336 3(K) , Murree,Stewar t403 4(K) ,Naran iDag ,Drummon d432 6(BM) ,Kabis ,Drammon d14474 , (BM). 148 Figure 2.15.Illustration s of C. graveolens. a. Illustration from Lindley(1846 ) b. 1 Polunin, Sykes &William s 3339(E) . 2Detai l of 1. 3 and4 Flower s of specimen 5396o f theFores t Research Institute,Dehr aDu n(E) . 149 ^^"fc^-'t'-w^w Figure 2.16.Distributio n of C. graveolens. 150 2.4.4. Clematis intricata Bunge Clematisintricata Bung e(1833) -Enumeratio nPlantaru mChin aBorealis ,Mem .Sav .Etr . Acad.SCi .St.-Petersb .2:7 5(1833) .Holotype :Bung e1/ 3(1831) ,Chin aBoreali s(LE) ,cf . Limpricht, 1922. Synonymy: Homotvpic synonyms: Clematis orientalis L.var . intricata (Bunge) Maxim. Clematisorientalis L . subsp. thunbergii(Steud. ) O. Kuntze var. intricata(Bunge ) O. Kuntze p.p. in Monographic der Gattung Clematis.In : I. Urban et al. (eds.) - Verhandlungende sBotanische nVerein sde rProvin zBrandenbur g(1885) ,83-202 , pro parte. Heterotypic synonyms: Meclatissibirica Spach . Clematis orientalis L.var .glauca Maxim . Clematis glaucasens u Sargent, nonWilld. ; sensu Rehder; sensu Ling. Clematishilariae S .Kovalevskay ai nNot .Syst .Herb .Inst .Bot .Acad .Sci .Uzbekista n1 8 (1967): 34 (Type: Kovalevskaya, s.n., 17-9-1966,plan t cultivated inth e Uzbek BotanicGarde n(Tashkent )fro m seedscollecte di nth ePami rvalley ;holotyp eLE , isotypes BM,E , G,K) . Clematischrysantha var .monantha Tamur ai nKitamur aAdd .Corr .Fl .Afghan .9 2(1966) : 92 (Holotype Yosii 801, Afghanistan, Ishkashim, KYO [notseen]) . Clematis chrysantha var.paucidentata Tamur a in Kitamura Add. Corr. Fl. Afghan. 92 (1966):9 2 (Holotype Yosii 488,Afghanistan , Qazideh, KYO [notseen]) . Clematissarezica Dconn .i nNovost i Sist.Vyssh .Rast . 14(1977) :23 1(Type : Dconnikov 5830, Uzbekistan, SaryTuga i (Pamir);holotyp e LE,isotyp eK) . Illustration: Ling (1980) inFlor a Reip.Pop .Sin .28 : 142,f . 15;se eals o figure 2.17. Description:Wood yclimbers ,u pt o4 mtall ,sometime swit ha compact ,bush yhabitu sdu et o environmentalconditions .Branchlet sangula rt ocostate ,th eyoun gtwig spilose ,late ralmos t glabrous.Leave svariousl ycomposite ,pinnate ,bipinnate ,sometime sbiternat ewit h(5-)7-15( - 21)leaflets ;th epetiole s12-5 7m mlong ;th eleaflet srangin gfro movate/lanceolat et onarrowl y lanceolate, green orgreyis h green, 16-56 x 2-47 mm, 1-10 x aslon g aswide ,entire ,o r irregularlyserrate ,dentat eo rlobe dwit ha fe wincision sa tth ebas eo fth eleaflet ,sometime s cleftalmos tt oth ebas ei n 1 mainpar tan d 1 or2 smalle rlatera lparts ,o rcompose do fthre e leafletso fsecon dorder ,th epart ssometime swit ha fe wteeth ,acute ,acuminat eo rmucronat e atth eapex ,mor eo rles sabruptl ynarrowe do rcuneat ea tth ebase ;th epetiolule sglabrou so r 151 sparselypilose ,5-3 8m mlong .Inflorescence smostl yaxillar ybu tsometime stermina l 1-3- floweredcyme s- occasionall ymor etha n3 flower spe rcym ei nvigorou sgrowin gspecimen s -;peduncl e(3-)8-23(-52 )m mlong ;pedicel sglabrou so rsparsel ypilose ,20-6 7m mlong , straighto rnoddin ga tth etop .Flower s+ pending ,open ,fla to rbroadl ycampanulate ,profusiv e flowering;tepal s4(-5) ,brigh tyello wsometime sveine do rwit ha flus ho fviolet ,herbaceous , ovatet o lanceolate, (12-)14-22(-24) x 4-10 mm, +twic e as long as wide,acuminat e or mucronatea tth eape xwit hbroadl ycuneat ebase stouchin geac hother ,lanat ea tth eincurve d margin,pilos e inside,glabrou s outside, spreading. Distribution: China (Hopei, Shansi), Russian F. (Gorn Altai), Kirgiziya, Uzbekistan, Tadzhikistan (see figure 2.18). Specimensexamined :CHIN AHopei ,Hsiao-Li-Chen ,Li u 12684(NY ,S) ;Hsiao-wu-tai - shan/Che-tao-kou,Smit h29 4(BM ,LD ,S ,UPS) ;Peking ,Davi d290 4(P) ;Pei'ping ,Bushell , s.n., 7-1869,(K) ,Carles ,s.n. , 1882,(BM) ;CHIN AShansi ,Taiyman ,Licen t216 3 (BM); Limpricht680 ,Taiyuan gfu /T'ai-yian-che n(S) ;Pingyang ,Serr eA67 3(UPS) ;Yan gCh'en , Serre,A76 4 (G);Chiao-ch'eng ,Smit h 7214 (BM,LD ,S ,UPS) ; RUSSIAN F., Gorn Altaisk, Elias,Weber , Tomb &Krasnaboro v 4411(NY) ; KIRGIZIYA, Issyk-kul Ozero,Brochere l 13(G) ; TADZHIKISTAN, Tianshan, Appleton 674(K) ; UZBEKISTAN, SaryTugai ,Dconniko v583 0(K ) 152 153 Acuteti p atape x oftepa l t •• • <•( .1 4 K'V. ifc-fi-'--^ Figure 2.17.Illustration s of C. intricata Bunge a.Drawin gb yMarie td eGeu s b. Photograph of flower (Bra58WAG ) c.Photograp h of holotype (LE) 154 Figure 2.18.Distributio n of C. intricata Bunge. 155 2.4.5. Clematis ispahanica Boissier Clematis ispahanica,i n Boissier, Diagnoses orientalum novarum (1845), 1 (6):3-4; Ranunculaceae,Flor aOrientali s(1867) , 1: 1-98. Lectotype(designate dhere) :Auche rElo y4026 ,Iran ,Isfaha n(Ispahan )(G) ,isotype sFI-W , G, K,LE ,P .Paratypes : Kotschy326 6(B ,G , K, M),Kotsch y 374,638(FI-W ) Synonymy: Heterotypicsynonym : ClematispseudoorientalisO .Kuntze ,i nMonographi cde rGattun gClematis. In :I .Urba n etal .(eds. )- Verhandlunge nde sBotanische nVerein sde rProvin zBrandenbur g(1885) , 83- 202.Berli n(Syntyp eAucher-Elo y4025,4026 ,P ;sinc eAuche rElo y402 6i salread ychose n aslectotyp eo f C. ispahanica, C. pseudoorientalis is anome nillegitimum) . Illustration: seefigur e 2.19. Description:Slende rerec tperennia lo rsubshrub ,u pt o2 mtall ,summe rflowering ,July-August . Twigsno ttwining ,round/angular ,sometime st ocostate ,initiall yslightl ypilose ,late rglabrous . Leavespinnate ,sometime sbipinnat eo rbiternate ,wit h7-9(-27 )leaflets ;th epetiole svariabl e inlength ,23-8 5mm ;th eleaflet slinearifor mt onarrowl ylanceolate ,herbaceous ,pal egree no r greyishgreen ,glabrous ,variabl ei nsize ,28-6 7x 4-2 1mm ,3-1 0x a slon ga swide ,entire ,o r partlyserrate ,dentate ,lobe da tth ebase ,o rirregularl ylobed ,acut eo racuminat ea tth eapex , angustatea tth ebase ;th epetiolule spilos ea tth ebase ,erec to rslightl ytwining ,5-2 1m mlong . Inflorescences,termina lan daxillar ycyme swit h7-1 7flower spe rcyme ,th ecyme sbein gpar t ofa large ,loos ethyreoi dsynflorescenc ea tth eto po fth eyoun gtwigs ;peduncl e7-6 2m mlong ; pedicelsglabrous ,22-7 0m mlong ,+ straight ,o ra tth eto psomewha tnodding ,bu twhe nfruitin g elongatean dstraight .Flowe rbud spal egreen ,ovoid ,acut eo racuminat ea tth eapex .Rower s open,flat ,+ upright ,abundan tflowerin gi na rathe rshor tperiod ;tepal s4 ,cream ywhit et opal e yellow,lanceolat et olineariform ,9-2 5x 2- 8mm ,3- 4x a slon ga swide ,blun to racut ea tth e apex,angustat ea tbases ,hardl yno ttouchin geac hother ,lanat ea tth eincurve dmargin ,shightl y piloset oglabrou sinside ,glabrou soutside ,spreading ,sometime srecurvin glater .Stamen s relativelyfew ,20-30 ;filament sdilatate ,greenis hwhite ,sometime stinge dwit hviolet ,o rdar k violet,glabrous ,u pt o1 cmlong ;anther sgreenis hwhite ,pal eyello wo rviolet ,2- 3m mlong . Pistilsnumerous ,+ 40 ,rangin gfro m30-80 ;ovarie sellipsoid ,pilose ,1 m mlong ;style slong - 156 pubescent,4- 6m mlong ;stigma sstraigh to rslightl yhooked .Achene srhomboid ,slightl yribbe d atth emargin , glabrous, 2-3 x 2mm ; elongate persistent styles, 16-33m mlong ,pubescent . Distribution: IRAN (Esfahan), seefigur e 2.20. Specimensexamined :IRA N(Isfahan) ,Bornmuelle r1 8(B) ,198 1 (G),Forough i95 8(K) ,793 7 (K),794 9(K) ,795 0(K) ,Furs e315 8(K) ,906 2(K) ,Hewe r149 2(G ,K) ,Kotsch y326 6(B , G,K , M), Kotschy 374,638 (FI-W),Pravit z 225(S) ,Rechinge r 56185 (B, M). 157 ^*." Figure2.19 .Illustration s of C. ispahanica Boiss. aFlowe r of C. ispahanica (drawing byMarie t deGeus ) b 1/2 Isotypes of C. ispahanica (Aucher-Eloy402 6FI-W ,LE ) 3Kotsch y 641,3266 (FI-W) 4 Kotschy 374,638 (FI-W) 158 Figure 2.20.Distributio n of C. ispahanica. 159 2.4.6. Clematis serratifolia Rehder Clematisserratifolia Rehde ri nMitteilunge nde rDeutsche nDendrologisch eGesellschaf t19 : 248(1910) ,an di nFedde ,Repert .Spec .Nov .Reg .Veg . 13:36 2(1914) ;Holotype :J.G . Jack, s.n., 1905,Kore aPin gYan g(A) ,isotyp e(LE ,MO) . Synonymy: Heterotypicsvnonvms : Clematisorientalist, var .se/rat aMaximowic zi nBull .Acad .Sci .St .Petersb .9:21 1 (1879) p.p. (holotypeGoldenstad t 255(10-8-1872 ;LE ) Clematisintricata Bung evar .serrata (Maximowicz )Komaro vi nAct .Hort .Petrop . 22: 289 (1904)p.p . Clematisserrata (Maximowicz )Komaro vi nAlis .Key ,PI .Fa rEas tReg .USS R1 : 549(1931 )p.p . Clematis orientalis L. var.wilfordi Maximowiczi n Bull.Acad . Sci.St .Petersb .9 :21 1 (1879)p.p .(typ eWilso n1150 ,Plastu n(Russia nFiManchuria) ;holotyp eLE ,isotype s G, K,M , P). Clematiswilfordi (Maximowicz )Komaro vi nAlis .Key ,PI .Far .Eas tReg .USS R1 1.168.(1931 )p.p . Illustration:figur e2.21 . Description:Wood yclimber ,u pt o5 mtall ,producin gvigorou sstolons flowering, in summe r (July-)August-September.Branchlet scostate ,sparsel ypilos eo rglabrous .Leave sbiternate , sometimesth efirs torde rdivisio npinnate ,no tternate ,with+ 9leaflets ;th epetiole s35-6 5m m long;th eleaflet soblong-lanceolat et oovate-lanceolate ,herbaceous ,dar kgreen ,glabrous ,28 - 70x 12-3 0mm ,2- 4a slon ga swide ,regularl yserrate ,sometime swit ha pai ro fbasa llobes , acuteo racuminat ea tth eapex ,angustat eo rcordat ea tth ebase ;th epetiolule sglabrou so rwit h afe w scatteredhairs ,strongl ytwining , 11-35m mlong .Inflorescenc eaxillary ,(l-)3(-5) - floweredcymes ,sometime sth elatera lflower so fcyme saborted ;peduncl e3-2 0m mlong ; pedicelssparsel ypilose ,25-7 0m mlong ,curve da tth eapex ,bu twhe nfruitin gelongat ean d straight.Flowe rbud softe nreddish ,o rgree nyellow ,ovoi dwit ha nacuminat eapex .Flower s +nodding ,+ open ,flat ,profus eflowering ;tepal s4 ,pal eyello wtinge do rveine dwit hviolet , ovate-lanceolatesometime snarrowl ylanceolate , 16-25x 6- 8mm ,2- 4x a slon ga swide , acuminatea tth eapex ,wit hbroadl ycuneat ebase stouchin geac hother ,lanat ea tth eincurve d margin,glabrou soutside ,sparsel ypilos einside , spreading,no trecurvin glater .Stamen s 160 numerous,20-35 ;filaments dilatate ,greenis hyellow ,tinge dwit hviole to rdar kviolet ,pilose ; anthersgreenis hyellow ,tinge dwit hviole to rviolet ,u pt o3 m mlong ;incidenta loccurrenc eo f staminodiaa stransitiona lfor mbetwee ntepal san dstamens .Pistil snumerous ,80-100 ;ovarie s ellipsoido rrhomboid ,pubescent ,+ 1 m mlong ;stigm aslightl yhooked ,glabrous .Achene s rhomboidt o(ob)ovate ,no tmarkedl yribbe da tth emargin ,pubescent ,brow nwit hfibrou s performance,2-3x1- 2mm ;elongat epersisten tstyle su pt o4 c mlong ,covere dwit hlon gerec t hairs. Remarks:Th enam eClematis koreana Komaro vi sfrequentl yuse di ncultivatio nfo rplant s belongingt oC. serratifolia. I tdiffer sfro mC. serratifolia i nhavin gmostl yternat eleave swit h broadlyovat eleaflet swit hcordat ebase san dsolitar yyello wt oviole tflowers .I ti sdoubtfu l whetherthi sspecie sitsel fi si ncultivation .C. koreana is ,lik eC. serratifolia endemi ct oKore a and Manchuria. Distribution: Endemic to Koreaan dManchuri a (seefigur e 2.22) Specimens examined: Jack 18-9-1905 (A, MO);Komaro v 710 (P);Pareja s H968 (G); Skortsov, 13-9-1967 (C,FI ,LD , M, MO);Trotte r 33(K) ;Uchiyama , 12-8-1902(LD) ; Wilford 1150(G ,K ,LE ,M ,P , S);Wilso n 8927 (BM,MO) ;Wilso n 10711(BM ,MO) . 161 X? Figure 2.21. Illustrations ofC. serratifolia Rehder a. Holotype,J.G .Jack , s.n., 1905,Kore aPin g Yang (A) b. Illustration from Riekstina (1990). c.Photograp h from Riekstina (1990). 162 Figure2.22 .Distributio n of C. serratifolia. 163 2.4.7. Clematis tibetana O.Kuntz e Clematistibetana O .Kuntze ,i nMonographi cde rGattun gClematis. In :I .Urba ne tal .(eds. ) - Verhandlungende sBotanische nVerein sde rProvin z Brandenburg,83-202 . Holotype:Strache y& Winterbotto m3 ,India ,Mila mKumao n(Utta rPradesh) ,K ;isotypes : BM,BR,P).Cf.Hara(1978). Description:Wood yclimber ,u pt o5 mtall ,flowerin gi nsummer ;(June-)July-September . Branchletscostate ,pilose .Leave scommonl ypinnat eo rpartl yo rwholl ybipinnat ewit h7-2 1 leaflets;th epetiole s14-7 2m mlong ;th eleaflet svariabl ei nshap ean dsize ,rangin gfro movate , lanceolatet olinear ,herbaceous ,glaucou so rgreen ,+ glabrous ,sometime spilos edownsid ea t veins,15-7 5x (3-)8-22(-44 )mm ,1.5-1 0x a slon ga swide ,leafle tincisio nver yvariabl ebu t mostofte nirregularl yserrate ,dentate ,sometime swit h1 o r2 basa llobes ,acut ea tth eapex , cordate,cuneat ean dmos tofte nangustat ea tth ebase ;th epetiolule spilose ,(2-)5-29(-45 )m m long.Inflorescence saxillar yo rtermina lcyme swit hcommonl y1 or3 flower s- i ncas eo fa solitaryflowe rth elatera lflower so fth ecym ear eaborted ;peduncl e0-6 5m mlong ,thu sleadin g torathe rdifferen t lookinginflorescence sa tfirs tsight ;pedicel spilose ,noddin ga tth etop ,25 - 120(-320)m mlong .Flowe rbud sgreenis hyellow ,yello wsometime swit ha flus ho fviole to r spottedwit hviolet ,occasionall yviolet ,ovoid .Fl ower svariabl yshaped ,rangin gfro malmos t open,flat ,broadl ycampanulat et onarrowl ycampanulate ;tepal s4(-5) ,basa lcolou rbrigh t yellowo rgolde nyellow ,eithe ro rno ttinge dwit ho rspotte dwit hviolet ,raril ycompletel ydar k violet,ovat et olanceolate ,16-3 5x 4-2 1 mm,1.5- 4x a slon ga swide ,acute ,acuminat eo r mucronatea tth eape xwit hcuneat ebase stouchin geac hother ,lanat ea tmargins ,glabrou so r piloseinside ,mostl yglabrou so rles scommonl yslightl ypilos eoutside ,i nan ycas espreadin g later.Stamen snumerous ,30-60 ;filament sdilatate ,variousl ycoloure dfro mgreenis hyello wt o darkviolet ,pilose ,2-1 0m mlong ;anthers ,greenis hyellow ,u pt o5 m mlong .Pistil snumerous , + 100;ovarie sellipsoi do rrhomboid ,pubescent ,+ 1 mmlong ;style spubescent ,4-1 1 mm long;stigm aslightl yhooked ,glabrous .Achene srhomboid ,raril y(ob)ovate ,no tmarkedl y ribbed,wood yo rfibrou sappearance ,2-6x1- 2mm ;elongat epersisten tstyle su pt o4 c mlon g (in cultivationu pt o7 c m long!),covere dwit h longerec thairs . 164 2.4.7.1. Clematis tibetanaKuntz e subsp. tibetana Synonymy: Heterotypic svnonvmv: Clematisakebioides (Maxim. )Veitc hi nNe whard yplant si nWester nChina ,9 .p.p .(Chin a Kansu,holotyp e Potanin s.n., 22-7-1885LE ,isotyp eK) . Clematisladakhiana Grey-Wilso ni nClematis orientalis (Ranunculaceae )an dit sallies .Ke w Bulletin 44:33-6 0(holotype :Thomso n s.n. Kashmir, Nubra, K) Clematisorientalis L .var .acutifolia Hook.f .an dThomso ni nFlor aIndica ,vol . 1. (1855) Clematideae, 3-12 Syntype:Thomson , s.n. Ladakh,K) . Clematisorientalis L .subsp .orientalis var .daurica (Pers. )O . Kuntze subvar. thomsonii O. Kuntze in Monographic der Gattung Clematis. In I. Urban et al. (eds.) - Verhandlungende sBotanische nVerein sderProvin zBrandenbur g(1885) ,83-20 2 (Type:Thomson , s.n.; holotype B,isotyp eK) . Clematisorientalis L .subsp .orientalis var .daurica (Pers. )O .Kuntz esubvar .dyeri Clark e ex O. Kuntze in Monographic der Gattung Clematis.I n I. Urban et al. (eds.) - Verhandlungende sBotanische nVerein sde rProvin zBrandenbur g(1885) ,83-20 2 (Holotype: Clarke 30329B,Kashmir , Askole,K) . Illustration: seefigur e2.23 . Description:Wood yclimber ,u pt o4 mtall flowering, i nsummer ;(June-)July-September . Branchletscostate ,pilose .Leave spinnat et obipinnat ewit h9-2 1leaflets ;th epetiole s33-7 2 mmlong ;th eleaflet slanceolat et olineariform ,herbaceous ,glaucous ,+ glabrous ,30-7 5x 3-3 4 mm,2-1 0x a slon ga swide ,acut ea tth eapex ,angustat ea tth ebase ;th epetiolule spilos ea t thebase ,10-4 5m mlong .Inflorescenc e (1-)3(-7)-flowere d axillaryo rsometime stermina l cymes;peduncl e5-5 5m mlong ;pedicel spilose ,40-9 0m mlon gcurve da tth eapex .Flowe r buds greenish yellow to yellow or greenish violet to violet, ovoid, acuminate. Flowers campanulate;tepal s4 ,yellow ,spotte do rtinge dwit hviole tt odar kviolet ,ovat et obroadl y lanceolate,16-2 7x 4-2 1 mm,1.5- 4x a slon ga swide ,acute ,acuminat eo rmucronat ea tth e apexwit hcuneat ebase stouchin geac hother ,lanat ea tmargins ,pilos einside ,glabrou soutside , spreadinglater . Remaining characters, seespecie s description. Remarks:It smor eo rles sglaucou sfoliag ean dit stepal swit hviole tspot smak ethi ssubspecie s different and easilyrecognizabl e from C. tibetanasubsp . tangutica. Distribution:Chin a(Tibet) ,Indi a(Utta rPradesh) ,Jamm uan dKashmir ,Pakista n(Baltistan) . See figure 2.26. 165 Specimens examined: CHINATibet ,La-ma-ts' o Shank'ou,Meebol d 3366(G) ; INDIA (Uttar Pradesh),Milao n Kumaon,Strache y &Winterbotto m 3(BM ,BR , K,P) ; JAMMUan dKASHMIR ,Askole ,Clark e30329 A(K) ,Chenure ,Koel z253 7(K ,L ,NY) , HusheSaltar ajunction ,Ludlo w 369(BM) ,Panamik ,Ludlo w540,54 2(BM) ,Shushah , Ludlow83 5(BM) ,Leh ,Ludlo w& Sherif f 8476(BM ,UPS) ,Sto kNullah ,Ludlo w& Sherif f 8552(UPS) ,Nubr avalley ,Meinertzhage ns.n. ,July ,(BM) ,Schomber g27 ,(BM) ,Dras/Karg U road, Stainton 7981(K) ; PAKISTAN (Baltistan), Sopor, Koelz 9593(NY) . 166 167 /fitG"? • Figure 2.23.Dlustration so f C.tibetana Kuntze subsp. tibetana a.Drawin gb yAnj a vande rNeu t (Bra285WAG ) b. Aquarelle byMarie t de Geus (Bra255WAG ) 168 2.4.7.2. Clematis tibetanasubsp .tangutica (Maximowicz) Brandenburg comb.nov . Basionym: Clematisorientalis L. var. tanguticaMaxim . Type: Przewalski 105,Chin a Kansu,Terr aTangutorum ;holotyp eLE ,isotype sE ,K ,P ;syntype sPrzewalsk i 171,Chin a Kansu,Terr a Tangutorum, LE;Przewalsk i 185, Tibet,Zai'dam , K,LE ,P) . Synonymy: Homotvpicsvnonvms : Clematis tangutica (Maxim.) Korsh. in Fragmenta florae Turkestaniae: 1. Clematis tangutica; 2. Clematis orientalis var. roschanica. Bulletin de l'Academie ImpeYialede s Sciences deSt.-Petersbour g Veser. ,vol .9 (1898):399-400 . Clematis tangutica(Maxim. ) Korsh. subsp.tangutica. Heterotypic svnonvms: Clematis akebioides (Maxim.) Veitch in New hardy plants in Western China, 9. p.p. (China Kansu Holotype Potanin s.n., 22-7-1885 LE,isotyp eK) . Clematis eriopodaKoehne ,i n Clematis. Deutsche Dendrologie (1893), 152-160;567 . Clematis chrysantha Ulbr.i nFedd eRepert .Beih . 12(1922) :37 4(Lectotyp e Limpricht 2086, China Kansu, Tsokadu B;paratyp e Limpricht 2146,B) . Clematis alpinasens u Kapoor, nonMill . Clematistangutica (Maxim. )Korsh .var .obtusiuscula Rehde r& Wilso nin :C.S .Sargen t (ed.) - Plantae Wilsonianae an enumeration of the woody plants collected in Western Chinafo rth eArnol dArboretu mo fHarvar dUniversit ydurin gth eyear s 1907, 1908 and 1910, vol. 1 Clematis (1913), 319-343 (Wilson 2487, China Sichuan, Ta-p'aoshan; holotype BM,isotype sE ,K) . Clematis tangutica (Maxim.) Korsh. subsp. obtusiuscula (Rehder & Wilson) Grey-Wilson. Clematis tangutica (Maxim.) Korsh. subsp. mongolica Grey-Wilson in Clematis orientalis (Ranunculaceae) and its allies. Kew Bulletin 44: 33-60 (holotype: Jeffrey 1436,Mongolia , Tula River N.E.Ula nBator) . ClematispamiralaicaGrey-Wilso ni nClematis orientalis (Ranunculaceae )an dit sallies . Kew Bulletin 44: 33-60 (Type: Tolmatcheva 4367, Tadzhikistan, Pamir, Murghab;holotyp e K, isotypes BM,C ,E , G,LD ,LE ,MO ,S) . Clematis orientalisvar .akebioides Maxim. InAct a Horti Petrop. 11:6 (1890). Clematis glauca var. akebioides (Maxim.) Rehder and Wilson in Sargent Plantae Wilsonianae 1:342 (1913). Illustration: Botanical Magazine t.771 0(1900) ;Revu eHorticol e 1902,p .528 .se eals o figure 2.24. Description: Woody climber, up to 5m tall, sometimes with a compact, bushy performance due to environmental conditions (altitude, drought). Branchlets costate, pilose.Leave spinnat e orpartl yo rwholl ybipinnat ewit h7-15(-20 )leaflets ; thepetiole s 18-70 mm long; the leaflets ovate to lanceolate, irregularly serrate-dentate or slightly lobed (1-2lobe s atth ebase) ,herbaceous , green, glabrous above,pilos e atvein sbelow , 15-56x 6-19mm ,2- 5 xa slon ga swide ,acut ea tth eapex ,cordate ,cuneat eo r angustate atth ebase ;th epetiolule spilos ea tth ebase ,5-2 0m mlong .Inflorescenc e mostlysolitar y 169 atth eapex ,cordate ,cuneat eo rangustat ea tth ebase ;th epetiolule spilos ea tth ebase ,5-2 0 mmlong .Inflorescenc emostl ysolitar yaxillar yan dtermina lflower s- occasionall y2- 3flower s togetheri na cym e- ;peduncl e0-2 5m mlong ;pedicel spilose ,a tth eto pnodding ,7-3 2c m long,profus eflowering .Flowe rbud sgreenis hyello wchangin gove rint obrigh tyello wwhe n maturing, ovoid, acuminate. Flowers narrowly campanulate; tepals 4, bright yellow, herbaceous,ovate-lanceolate ,18-35x8-1 2mm ,2- 3x a slon ga swide ,strongl yribbed ,acut e oracuminat ea tth eape xwit hbroadl ycuneat ebase stouchin geac hother ,tomentos ea tmargins , glabrousinside ,glabrou so rwit ha fe wscattere dhair soutside ,spreadin glater .Achene sovate , pilosewit h persistent plumose stylesu pt o 7c mlon g in cultivation. Forth eremainin g characters seespecie s description. Distribution:Mongolia ,Chin a(Shansi ,Kansu ,Xinjian gUygu rZiziqu ,Tsinghai ,Sichuan , Yunnan), Kazakhstan, Tadzhikistan, Jammuan d Kashmir (seefigur e 2.26.). Specimensexamined :MONGOLIA ,TulaRiverN.E .Ula nBator ,Jeffre y 1436(K) ;CHINA Shansi, Lu Yak Shan, Smith 8122, (LD, S, UPS); Kansu, Przewalski 105 (K, LE,P) , Przewalski 171 (LE),Zaidam ,Przewalsk i 185(K ,LE ,P) ;Xinjian g UygurZiziqu ,Chaka , Tsinghai,Deas y4 5(BM ,C) ,Deas y77 ,Ma nKhol avalle y(BM) ,Aksu ,Merzbache r8 9(B) , Imyltshvalley ,Raicheng ,Merzbache r 1236(B ,M) ,Jarkand /Serek-kol ,Norsted t4 (S ,U) ; Yunnan,Mekoj g Sakvin,Forres t 13381 (K); Sichuan,Song'pan ,Smit h291 3 (LD,UPS) ; Tsinghai, Kokonor, Rock 13277 (C,UPS) ; KAZAKHSTAN, Dzungarskiy Alatau,Mynas ,Merzbache r 1022(M) ; TADZHIKISTAN, Woshikiniozero ,Rantsjoem , Konnov 3076(TAD) ; JAMMU and KASHMIR,betwee n Daan dHanla ,Rupshu , Koelz 2288b (K,NY) . 170 A A w* AA 6 6 ^ ^r 1 { A A A A AA i A V/: \J A v A v V A A A / A :"\ \ A A 0 V v O0UG0Q A A 0 / < \ /\ ^ \y v/ -' \j ^ ^5^ A A 0 0 0 U 0 0 0 A A A A A A AI \ A V A W *0 171 172 SA"> W ') 173 Figure2.24 .Dlustration s of C. tibetanasubsp .tangutica (Maxim. ) Brandenburg a. Variationi nflower siz ean dshap eo ftepal s(Population s76045,76054,77017,7702 0 and 77022) b. Leaf morphology (population 76045) c. Fruitinghea d and acheneo f plant 75-4 d. Achenes:shape ,dorsa l view and longitudinal section (plant 77017-4) e. 1 holotype of C. tibetana subsp. tangutica (Przewalski 105 LE); 2/3 isotype (Przewalski 105LE) ;4 paratyp e(Przewalsk i 171LE) ;5 paratyp e(Przewalsk i18 5 LE). 174 2.4.7.3. Clematistibetana subsp .vernayi (C.E.C.Fisch. )Grey-Wilsoni nClematis orientalis (Ranunculaceae) and itsallies .Ke w Bulletin 44: 33-60 Basionym:Clematis vernayi C.E.C .Fischer ,i nBull .Misc .Inform . Kew 1937(1937) : 95 (Holotype: Cutting &Vema y57 ,Chin aXizang ,Gyantse ,K) . Synonymy: Homotvpicsvnonvm : Clematis tibetana O.Kuntz e subsp.vernayi (C.E.C.Fisch. ) Grey-Wilsonvar .vernayi. Heterotypic synonyms: Clematis chrysanthaUlbr .var .brevipes Tamur ai nAct aPytotax .Geobot .2 3(1968) :3 0 (Holotype Namikawa 124,Nepal ,Pijehl , KYO [notseen]) . Clematistibetana sens uHar a& William si nA nenumeratio no fth eflowerin gplant so fNepa l H (1979):16 . Clematis orientalis auct.no n L.i nBull .Dept .Med . PI.Nep .7 (1979):33 . Clematistenuifolia sens uLing ,no nRoyl ei nFlor eRe iPopulari sSinica e2 8(1980) :140 ,f . 41. Clematistibetana O . Kuntzesubsp .vernayi (C.E.C.Fisch. )Grey-Wilso nvar . laciniifolia Grey-Wilsoni nClematis orientalis (Ranunculaceae )an dit sallies .Ke wBulleti n44 : 33-60 (Holotype: Stainton, Sykes &William s 2130,Nepal , Kali Gandaki,BM) . Clematistibetana O .Kuntz esubsp .vernayi (C.E.C.Fisch. )Grey-Wilso nvar .dentata Grey - Wilsoni nClematis orientalis (Ranunculaceae )an dit sallies .Ke wBulleti n44:33-6 0 (TypeRoc k14124 ,Chin aXizang ,Radj aRive rGorge ;holotyp eK ,isotype sC ,E ,P , S). Clematis tangutica (Maxim.) Korsh. var.pubescens M.C .Chan g etP.P .Ling . Illustration: Botanical Magazine t.4495 (1850);se eals ofigur e 2.25. Description:Wood yclimber ,u pt o4 mtall ,flowerin gi nsummer ;(June-)July-September . Branchletsangula rt ocostate ,pilose .Leave spinnat eo rbipinnat ewit h7-2 1leaflets ;th epetiole s 14-64m mlong ;th eleaflet s ovatet onarrowl ylanceolate ,herbaceous ,green ,+ glabrou s upside,pilos ea tvein sdownside ,18-5 0x 6-4 4mm ,1- 8x a slon ga swide ,incidentall yentire , butcommonl yirregularl yserrate ,dentat eo rlobed ,clef talmos tt oth ebas ei n1 mai npar tan d 1o r2 smalle rlatera lparts ,o rcompose do f3 leaflet so fsecon dorder ,th epart ssometime swit h afe wteeth ,acute ,acuminat eormucronat ea tth eapex ,cordate ,cuneat eo rangustat ea tth e base;th epetiolule spilose ,2-2 6m mlong .Inflorescenc emostl ysolitar yaxillar yan dtermina l flowers- occasionall y2- 3flower stogethe ri na cyme ;peduncl e0-6 5m mlong ;pedicel spilose , 25-120m mlong .Flowe rbud sgreenis hyellow ,late rturnin gt obrigh tyellow ,broa dovoid , acuminateo rblunt .Flower sbroadl ycampanulat et oalmos topen ,flat ;tepal s4 ,brigh to rgolde n yellow,sometime swit ha flus ho fviole t(sometime sth ecolou rchange st oa brigh torang eyello w 175 whenflower swilt) fleshy,, sometime sver ythick ,broa dovat et olanceolate ,17-3 0x 7-1 6mm , 2-3x a slon ga swide ,acut ea tth eape xwit hbroadl ycuneat ebase stouchin geac hother ,lanat e atmargin ,pilos einside ,glabrou soutside ,spreading .Stamen ssometime swit hdar kviole t filaments.Achene swit hshow ylon gplumos estyle su pt o6 c mi ncultivation .Fo rth eremainin g characters, seespecie s description. Remarks:Differin gfrom C. tibetanasubsp .tangutica b yit sflesh yspreadin gtepal si nopen , flatt obroadl y campanulateflowers. This form wasver y longi n cultivation under themisapplie d nameC. orientalis. Distribution: China (Sichuan, Tibet),Nepal ,Bhuta n (seefigur e 2.26.). Specimensexamined :CHIN ASichuan ,Tung-Ch'e nSan-k'ou ,Smit h366 3(LD ,S ,UPS) , Drachogi,Smit h447 8(UPS) ,Batang-Paan ,Souli e3016bis ,(P) ,Sungp'an ,Wilso n313 1 (BM, K),313l a(BM) ,3132 a(K) ,Ta-p'as-chan ,Wilso n248 7(E ,K) ;Kansu ,Tsinghai ,Farre r& Purdom52 2(E ,M) ,Xhungyasha nShuTca ,Trippne r19 6(M) ;Xinjian gUygurZiziqu ,Su-kai - t'i,Ludlo w60 7(BM) ;Tibet ,nea rGyantse ,Cuttin g& Verna y38,5 7(K) ,Ludlo w7 5(BM) , Karlung,Humphrey s501 6(BM) ,Tongkyuk ,Kingdo nWar d608 1 (BM),Lhasa ,Ludlo w& Sheriff867 8(BM) ,Tsogo :Pasu mTso ,Ludlow ,Sherif f& Elliot t1392 8(BM) ,Tsangp ovalley , Simbiteng,Ludlow ,Sherif f& Taylo r447 2(BM) ,Tsangp ovalley ,Timpa ,Ludlow ,Sherif f& Taylor515 1(BM ,E) ,Ch uKyabden ,Ludlow ,Sherif f &Taylo r619 9(G ,UPS) ,Tongolo , Soulie92 1(P) ,Tzeku ,Souli e92 3(P) ,Samada ,Spence r Chapman 911(K) ; NEPAL,Tukule ,Grey-Wilso n& Phillip s36 6(K) ,Namdo ,Grey-Wilso n& Phillip s69 3(K) , Marsiandivalley ,Lownde sL107 8(BM ,UPS) ,Barbun gKhola ,Kakkotgaa ,Polunin ,Syke s &William s108 6(BM) ,Dunaih iBehr iriver ,Polunin ,Syke s& William s231 4(BM ,UPS) ,Kal i Gandaki, Yara, Stainton, Sykes &William s 2130 (BM),Muktinath , Stainton, Sykes & Williams 5645(BM) . 176 Figure2.25 .Dlustratio no fC. tibetana subsp .vernayi (C.E.C.Fisch. )Grey-Wilso n(drawin g Anja van derNeut) . 177 Figure 2.26.Distributio n of thesubspecie s of C. tibetana. # Clematis tibetanasubsp . tibetana • Clematis tibetanasubsp . tangutica • Clematis tibetanasubsp . vernayi 178 2.5. Cytology 2.5.1.Material and methods Fourdifferen t methodst omak esquas hpreparation so froo ttip st ostud ysomati cmetaphas e chromosome morphology, have been used to characterise Clematis sect. Meclatis chromosomes.Th eprocedure s aredescribe d andth erational e ofthei rus ei sgiven . Methodfo rFeulge nstainin go fmitoti cClematis chromosome s(Darlingto n& L aCour ,1967) : Germinate; Pre-treatth eroot swit h0.002 M8-hydroxyquinolin efo r4 hour sa troo mtemperature ; Eventually, keepth eroot si n 1:3 aceticethanol ; Macerate 8minute s in IN HC1 of 58°C; Stain with Feulgen for 4hours ; Rinseth eroo ttip si nrunnin g tapwater ; Squash themi n45 % acetic acid; Makepreparatio npermanen twit hEuparal ,o rdehydrat ei nthre esteps ,rins ei nxylo l andmoun t inCanad aBalsem . Becauseo fth especifi cbiochemica lreactio nbetwee nstai nnucleotides ,th eFeulge nstainin g methodi sver ysuitabl et odescrib eth emetaphas ekaryotype .Chromosome swer emeasure d inmetaphas e plates inpreparation s according toth e above schedule. Method for adsorptive staining of mitotic Clematis chromosomes: Germinate; Pre-treat the roottips with 0.002M 8-hydroxyquinoline for 4 hours at room temperature; Eventually, keep theroot s in 1:3 aceticethanol ; Macerate 8minute si n IN HC1 of5 8°C ; Stainingprocedur ea troo mtemperatur e(2 0°C )5 - 2 0minute s(1 )an d(2) ;3 0minute s (3). (1) l%carminin 45% acetic acid,o r 45% propionic acid; (2) 1%orcei ni n 45% acetic acid,o r 45% propionic acid; 179 (3) haematoxylin (Henderson &Lu , 1968); Squash in 45% acetic acid; Makepreparatio npermanen twit hEuparal ,o rdehydrat ei nthre esteps ,rins ewit hxylo l and mount inCanad a Balsem. Sometimes,preparation swer emad epermanen taccordin gt oth equickfreez e method after Conger and Fairchild (1953). Theacetocarmi nstainin gwa suse dfo rth epurpos eo fchromosom ecounting .I fth e stainingwa sweak ,subsequen tpreparation so fth esam eserie swer estaine dwit hcarmi ni n propionic acid. Theorcei nstainin gwa sals ouse dfo rcountin gpurposes .Detail so fth echromosom e morphology were,however , more clearlyexpresse d than with carmin staining. Haematoxylin is an unusual stain for chromosome studies. It proved to be an appropriatemetho dfo rstainin gsmal lchromosome s( 1-2u )i ncytologica lwor ki nBegonia an d Kalanchoe.Applyin gthi sstainin gprocedur et oClematis chromosome sappeare dt ob eusefu l as quick method to screen metaphases for details such as satellites. Methodfo rGiems aC-bandin go fmitoti cClematis chromosomes .Th euse dmetho di sslightl y modified from Greilhuber (1973) and Schwarzacher et al.(1980) : Germinate; Pre-treatth eroot swit h0.002 M8-hydroxyquinolin efo r4 hour sa troo mtemperatur e (20°C); Eventually,kee pth eroot si n 1:3 acetic ethanol; Keepth eroot s in aquadest for 20minute s onth e dayo f squashing; Transfer roots into INHC 1 for 5minute s atroo m temperature (20°C); Macerateth eroot si na nenzym esolutio ncontainin g1 % cellulas ean d0.5 %pectinas e in 0.01M citrate buffer (pH4.0 ) at 38%C for 25-30minutes ; Soften the roots in45 % acetic acid for 25-30minute s atroo m temperature (20°C); Squah theroo t meristems ina dro po f 45% acetic acid; Removeth ec oversli pafte rfreez edryin gi nliqui dnitroge nan dai rdr yth eslide sfo rtw o days; Immerse the slides in45 % acetic acid at60 CCfo r 25-30minutes ; Denatureth epreparation si n6 %BaO Hsolutio nfo r6- 7minute sa troo mtemperatur e (20°C) and wash them inrunin g tap water for 1 hour; Incubate the slides in45 % acetic acid at 60°C for 20minutes ; Rinseth eslide si n0.04 Mphosphat ebuffe r(p H6.8 )an dstai nwit h4% Giems asolutio n (in0.04 Mphosphat e buffer atpH6.8 ) for 10-15minutes ; 180 Rinse the slides again inphosphat e buffer and airdry ; Mount the slides inEuparal . ClematisC-bandin gpattern swer estudie di npreparations ,mad eaccordin gt oth e aboveschedule .I nthi spreparations ,th echromosome swer emeasure dan dth erelativ epositio n of bands towards thecentromer e weredraw n with the aid of acamer a lucida. Astandar dkaryogram ,base do nmeasurement swit hth emicroscop emicromete ri s presentedbelo wtogethe rwit hth emean so fdata .Th ecentromer epositio ni sexpresse di nth e shortar m/lon gar mratio ,a swa sdon eb yKalkma n(1984 )an dd ePutte ran dva nd eVoore n (1988),bu t seeals o Denver Study Group (1960),Leva n et al.(1964) . Microscopesuse dwer evariou svariant so fth eZeis sStandar dline ;measurement swer e partlycarrie dou ta ta magnitud eo f1250x ,usin ga 1 OO x(plan)apochromati cobjectiv ePhas e contrast (PH3).Photograph s were made with Zeiss MC63Photomicroscop e equipment. 2.5.2.Results and discussion Thebasi cchromosom enumbe ri nClematis i sx = 8 ,th emajorit y ofspecie sbein g diploid:2 n= 2 x= 16 .Th eoveral lchromosom emorpholog yi na Clematis genom ei sno t subjectt olarg evariatio nan dt ocharacteris ea sfollow s(nomenclatur eafte rth eDenve rStud y Group, 1960;se eals ofigur e 2.27.an d2.28.) : threelarg e (±8u) metacentric chromosomes; threerelativel y short (+6-6.5u) submetacentric chromosomes; tworelativel y short (+5.5u)telocentri c chromosomes. Oneo rtw otelocentri cchromosome sma ybea rsatellite s(secondar yconstriction so n theshor tarm .N orelatio nbetwee nClematis classificatio nan dth eoccurrenc eo fsatellite sca n beindicated . Polyploidyoccur sonl yoccasionall yi nth egenus .2 n= 4 x= 3 2ha sbee nreporte da s 181 presenti nsevera lspecies ,bu ti ti sneve ra characteristi ctrait ,occurrin gconsistently ,fo ran y Clematisspecies .I nth eWageninge ncollection ,tetraploid shav ebee ndetecte di npopulation s of C. tibetana subsp.tangutica and subsp. vernayi. Focusingo nClematis orientalis an dit sallies ,n oconsisten tdifferenc ebetwee nC. orientalisan dC. tibetana subsp .ha sbee nobserve di nth eoveral lchromosom emorphology . Intabl e2.9. ,th eresult so fmeasurement so nsomati cmetaphas echromosome sar epresented . Ingeneral ,th echromosom emorpholog ybetwee nthes especie si ssimilar ,whic hca nb esee n fromth eidiogram so fbot hspecie si nfigur e2.28 .Th esatellite stha toccasionall yoccu rar eno t drawn,a sthe yar einconsisten ti nappearance .Th eidiogram so fC. orientalis an dC. tibetana subsp.vernayi wer edraw nfro mmetaphas eplate sstaine daccordin gt oth eGiems aprocedure ; heterochromaticband swer eonl ydrawn ,i fthe ywer epresen ti nal lplates .Th eidiogra mo fC. tibetana subsp.tangutica wer edraw nfro mmetaphas eplate sstaine daccordin gt oth eFeulge n procedure.Measurement sbetwee nbot hmethod sca nb ecompare da sbot hpreparatio nan d stainingprocedure sar ebase do na simila rreactio nt oDN A(Sharm a& Sharma ,1972) .Th e adsorptivestainin gprocedur ewa suse dfo rchromosom ecounting so fpopulation st omak esur e thatn opolyploi dvariatio ncoul dinfluenc e theresult so f the interspecific hybridization programme.Thes eresult sar eno tfurthe rdetaile da sth emajorit yo fpopulation si sregularl y diploid.Onl yi nC. tibetanasubsp .tangutica tetraploi dplant san deve nsom etriploi dplant s weredetecte di ncertai npopulations .Photograph so frepresentativ emetaphas eplate sar e presentedi nfigur e2.27 .Th epopulation suse dar evouchere db yherbariu mspecimen sa s mentioned inth e legendo f this figure. 182 m ON o CN m CM cn ON NO cn Q CM o 8 o d O ^ d O d d d d NO NO CN CM m m ON CM s NO NO p m 00 in m ON in d in d in d d in d d CM en NO cn oo m 00 oo C~ in CM ON a CM ON s CM ON o o d d d d f« d d d d d in m •* in in 00 NO NO m p § d d in d r^ d in d •* in d NO ON CM ON in in 00 o ON a in NO 3 p CM d d d d d d SO CM 00 •* OO cn ON CM O CM ON NO 00 s CM en O CM 2 NO d NO p NO in in d in d o 00 CM NO ON 00 NO m in m Q NO CM en d NO d d en d d d CM NO NO CM 00 o ON m - 00 en s m CM m 00 •* NO CM CM NO CM d d NO d o NO NO NO in ON in cn NO Q ON CM cn cn en ON cn o t-; O ON o CM •* d d d d CM d d NO 00 cn in o in CM in m in CM NO oo en s r-' CM rr' cn d d cn d 0, cn o CM O in CM CM 00 in in s 3 s ON ON d oo in d in o d d d d d d r> o d d d CM 00 00 cn ON 00 m cn NO ON 3 ON ON m s rn d •* d en d cn NO ON o r- in oo NO in in in ON a r- p o f- ON o 00 00 o d d d d o « d d d d o NO CM CM o m ON NO o »n in S ON cn 00 ON 5 OO in 00 en cn •* p r-' en d d 00 d o m NO in in in in 00 ON o ON o ON 3 sCM d d o d § d CM a, CM d c CM ON cn in cn ON CM 00 o NO O in ON in 3 in in 00 2 00 ON 00 o 1 00 d •* •* d o > < < -J! < < < i / a OS £ £ £ £ t 2 / u•a a. SB 1 1 3 OS J 1 .5 ! u I i 9 E •I i 0 J, i j 0 t V **• ,*. ^ mitt «" S \ Mi'4 * • s / ^ - Figure2.27 .Somati cmetaphas eplate sstaine dwit hGiems ao fC. orientalisL . (82374-9; specimen vande rNeu t (WAG);photograph s Josva n deVooren) . 184 a Idiogram of Clematis orientalis L. diogram of Clematis tibetana subsp. vernayi (C.E.C. Fischer) Grey-Wilson diogram of Clematis tibetana subsp. tangutica (Maxim.) Brandenburg Figure2.28 .Idiogram sconstructe dafte rsomati cmetaphas eplates ,staine dwit hGiems a(a ,b ) andFeulge n (c). a C.orientalis L.(BR A 185,186WAG ) b C.tibetana subsp .vernayi (C.E.C.Fisch.)Grey-Wilso n (BRA 67,7 3WAG ) c C.tibetana subsp .tangutica (Maxim.) Brandenburg (BRA 66, 119WAG ) 185 2.5.3. Generaldiscussion The cytotaxonomic study of Langlet (1932) has contributed considerably to the understanding of the Ranunculaceae. Numbers of chromosomes per genome showed correlationswit hrelationship stoward sth eoveral lshap eo fchromosomes .Moreover ,bot h aspects could be related to the higher classification of the Ranunculaceae based on morphologicalcharacters .Withi nRanunculaceae ,th emos tfrequentl yoccurrin gbasi cnumbe r ofchromosome spe rgenom ei sx = 8 ,whic hi sth enumbe ro fa.o .mos tAnemone spp .an d Clematis;beside stha tx= 7i sals ocommonl ypresen t(e.g .Aquilegia an dHepatica). Gregor y (1940)an dOkad aan dTamur a(1979 )compare dth eclassificatio n ofRanunculacea ebase d onmorphologica lcharacter sb yPrant lwit hanothe ron etakin gint oaccoun tchromosom efor m andnumber ,an dshowe da certai ndegre eo fagreemen tbetwee nbot happroaches .Rothfel s etal .(1966 )studie dchromosom esiz ean dDN Aconten to fAnemone an drelate dgener aan d found apositiv e correlation. Clematis spp.ar enormall y diploid (2n=2x=16) andonl yoccasionall y tetraploid (Meurman&Thermann , 1939;Gregory , 1940;Kurita , 1956,1957,1958a, 1958b,1960 , 1962, 1964; Fedorov, 1969). Gregory (1940) reported C. mandshurica Rupr. and C. paniculataThunb .bein gtetraploid ,bu tcounting s atWageninge n(unpubl. )indicat etha tthes e arepopulationa ldeviation sa tth eutmost .Meurma nan dTherman n(1939 )alread yshowe dtha t withinClematis ther ei sno tmuc hvariatio na st ochromosom efor man dsize ,a conclusio n whichha sbee nconfirme drepeatedly .Th eoccurrenc eo fsatellite so nth eshor tar mo non eo r bothtelocentri cchromosome sha sappeare dt ob ea ninconsistenc ythroughou tClematis spp . andneve ra consistent ,characteristi celemen to nan ygenome .T olocalis eheterochromati c zonesi nchromosome swit hGiems astainin gha sprove nt ob ea usefu l tooli ncharacterizin g genomestha totherwis eloo kver ysimilar ,a sca nb elearne dfro mcytotaxonomi cstudie swit h Allium(Kalkman , 1984; dePutte r& va nd eVooren , 1988)an dLactuca (Koopma n etal , 1993).Comparin gth eidiogram so fC. orientalisan dC. tibetanasubsp .vernayi lead st oth e conclusiontha tGiems aband soccu ri nC. orientalisa tth etermina lzone so fth echromosome s andtha tC. orientalisshare smos to fthes eband swit hC. tibetanasubsp .vernayi. However , thelatte rshow sdoubl eband si nth elon garm so fchromosom e3, 4an d8 .A si nbot hspecies , 186 plants are analyzed of a single population it is not possible to conclude that there are interspecificdifferences .Furthe ranalyse shav et orevea lwhethe rthi spolymorphis mwithi nth e section Meclatis occurs atth e populational level, as it does in other studies,e.g .Scilla (Greilhuber, 1973),o rtha ti tconcern sinterspecifi cdifferences .Withi nth esectio nMeclatis, onceth eoccurrenc e ofB chromosome s hasbee nreporte d (Shambulingappa, 1965),bu t althoughi nth epresen tinvestigatio nw ehav ecounte dchromosome sfo ral lpopulation suse di n theinterspecifi chybridizatio ndiallele ,B chromosome shav eneve rbee nobserve dwithi nth e sectionMeclatis no r inothe r Clematis spp. 187 2.6. Pollen morphology Withinth eframework o fthi sClematis study ,ther ewa sconsiderabl einteres ti neithe rfresh o r storedpolle n(se e1.3.2.) ,an dadditiona lobservation so fth epolle nmorpholog ywer emad e withth eai do fScannin gElectro nMicroscopy .Moreover ,polle nmorpholog yha sprove nt ob e useful for plant systematic studies (Blackmore, 1984;Tobe , 1974). Pollen slideswer emad ea tTFDL-DL O(Gol dplating ;andincludin go rexcludin g pretreatmentb yrinsin gwit hchlorofor mt oremov eth eextexin elayer) .Observation swer emad e with aJEO LS.E.M . Kumazawa(1936 )an dWodehous e(1936 )studie dalread yth epolle nmorpholog yo f Ranunculaceae.Fro mthei rstudies ,i ti sobviou stha tth ebasi ctyp eo fpolle ni nRanunculacea e isth etricolpat eellipsoida ltyp ewit hdee pcolpe san dn odistinc tger mpore s(Erdtman , 1952). Polleno fClematis sect .Meclatis mee tthi sbasi ctyp eo fpollen .Not eth escabrat esurfac eo f pollena tth epola rside san di nth eequatoria lvie wbetwee nth ecolpes .Withi nsectio nMeclatis, pollen has been observed of C. ispahanica, C. tibetana subsp. tangutica,an d subsp. vernayi,C. orientalisan dC. serratifolia. Photograph so fpolle ni nbot hpola ran dequatoria l position ofC. tibetanasubsp .vernayi ar epresente d infigure s 2.29aan db . Kumazawa(1936 )note dapar tfrom th eabov ebasi ctyp eothe rpolle ntype si nRanunculacea e with no clear colpes but being polyporate. Within the Anemoneae, he depicted this phenomenonfo rAnemone coronaria L .an dfo rC. texensisBuckl. ,C. viornaL. ,C. stans Sieb.e tZucc , andC. patensMorr . etDecne. ,wherea sh edepicte d thebasi c typefo r C. apiifoliaDC . and C.fusca Turcz. var. mandshurica Takeda. Our S.E.M.-observations confirm Kuwazaga's drawings ast oC. patens (figure 2.29c). Asn o light microscopical observations were made and no measurements were carried out on Clematis pollen morphology,i ti sno tpossibl et omak emor edetaile dcharacterisation so fpolle npe rspecies . Itis ,however ,obviou stha tth edifferenc e betweenbot htype si ss olarge ,tha tfurthe r pollen morphological studyi s expected tob e significant for Clematissystematics . 188 189 Figure2.29 .S.E.M.-photograph s of Clematis pollen. Approximate magnitude atphotographs :4200 x a. C.tibetana subsp.vernay i -equatoria l view; b. C.tibetana subsp .vernay i -pola r view; c.C. patens. 190 2.7. Isozyme polymorphism inMeclatis Since morphological characters are more or less affected by environmental conditions (Crawford, 1985),an dkaryotypi canalysi sdoe sno tsho wmuc hvariation ,electrophoreti c analysiso ftota lprotein swa scarrie dou ti norde rt oge tsom einsigh ti nvariatio npattern swithi n Clematis sect. Meclatis. This experiment was carried out in 1984. Since then, the developmentsi nelectrophoreti cresearc hhav egreatl yprogressed .Th eexperimen ti sonl y reportedher ebecaus eo fth eresult sshowin gstartin gpoint st ofurthe relectrophoreti cresearc h withinClematis. 2.7.1.Material and methods Plantmateria li suse do f C orientalis (1 voucheredb yBR A18 5 and 186;2 vouchere db y BrandenburgBR A166) ,C. serratifolia ( 3vouchere db yherbariu mspecime nBrandenbur g BRA170),C. 'Bravo'( 4vouchere db yherbariu mspecimen sVa nde rNeu t35,3 6an d37 )an d C. tibetanasubsp .vernayi ( 5vouchere db yherbariu mspecime nBrandenbur g 188).Th etw o C.orientalis individuals were seedlings from different populations. Youngfres hleave s wereyielded ,groun dan dfroze nwit hliqui dnitrogen .0.5 gdowe xbuffe r (4gdowe xi n5m lphosphat ebuffe r [2.68gNa 2HP04.7H20an d3 .l gKH 2.P04i n100m ldem i water])wa sadde dt o1 g lea fsampl et obin dphenoles .Th esample s werelef tfo r4 5minute s at4°C .Th elea fsample swer esqueezed .12.5u l0.25 Mtris-borat ebuffe r (pH=7.9)wa sadde d to50u llea fextrac tan dcentrifuge d (15000tpm)fo r2 0minutes .Th esample swer ethe ndialyse d for 16hr st oremov eundesire dsalt ycompounds .A tthi sstage ,th eextract sca neithe rb euse d immediately for further analysis or stored in a refrigerator. ThePAG Egel swer eo f 12.5m l0.25 Mtris-borat e buffer (pH=7.9),5 0m l stock solution(75 gacrylamid ean d2 gmethylene-bis-acrylamid ei n50 0m ldem iwater) ,37.5m ldem i water,1m lsodiu msulfit esolutio n(43m gsodiu msulfit ei n1m ldem iwater) ,1m lammoniu m persulfatesolutio n(30m gammoniu mpersulfat ei nm ldem iwater )an d0.0 3m lTEMED .Afte r 30minute s of polymerisation the gel wasread yfo r use. Theelectrod e buffer is atrisborat e buffer (pH=7.9). 191 Verticalrunnin gwa scarrie dou ta t1 0° Cafte rprerunnin gdurin g4 5minutes .Befor e soakingth ewick swit hlea fextrac t(30ul) ,25u lDMS Ostai nwa sadde dt o10 0JLX Ilea fextract . Running timei s approximately 2hrs.an d ends when thevoltag e of 500V hadbee n reached. Gelswer estaine dwit hCoomassi eblu eo resteras eovernigh tunde rcontinuin gturning . 2.7.2.Results and discussion Froma tleas tthre erun sslot so fth esam eindividua lwer ecompared .Du et oth echaracte ro f anaspecifi cprotei nstain ,ther ewer elot so fhardl ydistinc t(overlapping )bands .Th eband s conspicuousi nal lreplication swer escreene dan dar epresente di nfigur e2.30c .Thi sfigur e showssom edifference s withinan dbetwee nspecie si nthes ebands .Bot hprovenance so fC. orientalissho wpolymorphis mi nband san dd ono thav esuc hpronounce dband sa si nwit hC. serratifolia.C. tibetana subsp.vernayi a swel la sC. 'Bravo'(whic h ascultiva rha sbee n selectedfro m hybridizationwit hC. tibetana) sharea tleas ton epronounce dban dwit hC. orientalis. Comparisonwit hothe rincidenta lrun smak eclea rtha tfurthe r analysiswit hisozym e specificstain sma ycontribut ei ndeterminin ggeneti cdistance sbetwee nspecie so fClematis sect.Meclatis. 192 Aspecific protein staining 1 C. orientalis 2 C. orientalis 3 C. serratifolia 4 C. 'Bravo' 5 C. tibetana subsp. vernayi H Figure2.30 .a .Isozym epolymorphis maspecifi cprotei nstainin go fClematis sect .Meclatis. b.Isozym epolymorphis mesteras eprotei nstainin go fClematis sect .Meclatis. c .Schemati c presentation of pronounced protein bands by PAGE of 5 specimens of Clematissect . Meclatis. 193 3. CLASSIFICATION OF CULTIVATEDPLANTS 1 3.1. History of classification schemesconcernin g cultivated plants Atth edaw n of taxonomic consciousness, cultivated plantsrathe r than wildplant swer e subjecto fclassification .I nthos eday seconomicall yan dculturall yimportan tplant s- usefu l orpoisonou s andcultivate d plants -wer e almostth eonl y species treated, asi s obvious from herbals sinceTheophrastos 'time . With the publication of Linnaeus'Specie s Plantarum, 1st May 1753,attitude s towards classification ofcultivate dplant sbecam ecleare rthroug hth econsequen tintroductio no f binomial nomenclature (Jarvis, 1986; Wijnands, 1986a). Some cultivated plants were regardeda sspecie sseparat efro mthei rwil do rweed yrelative san dputativ eancestors ,e.g . Ribes uva-crispavs . Ribesgrossularia, wherea s others were regarded as unnamed or namedvarietie swithi nspecies ,e.g .Brassica oleracea (Oos te tal. , 1989),Daucus carota (Wijnheijmere tal. ,1988 )andDianthus caryophyllus (d eLange ne tal. ,1984) .sometime s combinations ofthes eapproache s areused ,e.g. ,Brassica rapa vs. Brassica campestris (Ooste tal. , 1987)an dLactuca sativa vs . Lactucaserriola (d eVrie s &Jarvis , 1987). In other cases Linnaeus hesitated which of both approaches should be chosen, e.g. Beta vulgarisinclusiv eo fth eBeta maritima, vs .Beta maritima a sseparat especie s(Letschert , 1993;Linnaeus , 1753,1762). From Linnaeus until the second half of the 19thcentury , therewa s adecreasin g interesti nth etaxonom yo fcultivate dplant s(Heiser , 1986).Botanist sbecam eawar eo fth e hugevariatio ni nth ePlan tKingdom .The ymainl ydescribe dan dname dplant sfro mal lpart s ofth eworld ,focusin gthei rattentio no nth ehithert ounknow nplant srathe rtha nt oth ewell - known cultivated plants of those days. Cultivated plants were classified at various infraspecific ranks.Thes erank swer elargel ydetermine db ytraditio nan dno tb ynumbe r 1 Thischapte rha sbee nlargel ybase do nth eearlie rpublicatio nb yBrandenbur g& Schneider (1988; 3.1. and 3.2.), and the publication by Hetterscheid & Brandenburg (1995a; 3.3.an d 3.4.). 195 andmod eo fdistinguishin gcharacters .Vegetabl ean dsilvicultura lcro pplant swer eusuall y classifieda ssubspecie so rvarieties ,whils tornamentals ,includin gwood yornamentals ,wit h thesam ekin do fdistinguishin g characteristics wereterme dvarietie so rforms !Th eonl y rationalebehin di tma ywel lb etha tvegetable s andsilvicultura l cropsar epredominantl y reproduced generatively, whereasmos tornamental s arepropagate d vegetatively. There wasa remarkabl econnectio nbetwee nattitude stoward sclassificatio n andvariou ssector s of plant cultivation. In the 19thcentury , especially after Alphonse de Candolle's Origine des Plantes Cultivees (1883), cultivated plants attracted much attention again. In 1866, Alefeld publishedhi sLandwirtschaftlich e Flora,i nwhic hh eintroduce dth eter m'variet ygroup' , whichbecam elate rth esystemati ccategor yco nvariety .Wit hth eco nvariet ya specia lran k was created between subspecies and variety to designate crops, or, sometimes, crop groups.Th econvariet y soonbecam ea commonl yuse d systematic category, applied for several cultivated plants (Grebenscikov, 1949,1950; Mansfeld, 1950). As reviewed by Stearn (1986), the first edition of the International Code of Nomenclature for Cultivated Plants (ICNCP) was published by Stearn in 1953 after decadeso fdiscussio nabou tth edesirabilit yt omak ea distinctio nbetwee ncultivate dan d botanicalvarieties .Th econvariet ywa sadopte di nth e1953,19 58 ,an d196 1edition so fth e ICNCP as a means of cultivar classification. This attitude was almost parallel with statements aboutclassificatio n ofcultivate dplant s (Mansfeld, 1953,1954;Helm , 1954, 1963;Danert , 1962).Furthe r developments increatin g special categories for cultivated plants weremad e in an attempt to solve complex classification problems like thosei n Brassica. Jirasek (1966), among others, created an extensive hierarchy of special categories, summarized below: specoid (spd.) species subspecioid (subspd.) subspecies cultiplex (cpl.) subcultiplex (subcpl.) convarietas (convar.) subconvarietas (subconvar.) provarietas (provar.) subprovarietas (subprovar.) 196 conculta(cone. ) subconculta (subconc.) cultivar (cv.) subcultivar (subev.) In later work, Jirasek deleted the subcultivar, as it is in contradiction with the statement inICNC P(Brickel l et al., 1980),Art . 10secon d paragraph: "Thecultiva r isth elowes tcategor y underwhic hname s arerecognize d inthi sCode. ' Jeffrey (1968)trie dt oreduc eth enumbe ro fcategorie san dt odevelo pa synthesi s of differing opinions: species subspecioid or subspecies convanetas (subconvarietas) provarietas (subprovarietas) cultivar In the 1969 edition of the ICNCP (Gilmour et al., 1969), a new approach to cultivar classificationwa sadopte db yreplacin gstatement sabou tth econvariet ywit hth econcep to f cultivar-group.Fro mtha ttim eonward si twa spossibl et osubdivid elarg eassortment so f crop plants into cultivar-groups, defined by characters of agricultural importance. By discardingth etraditiona lbotanica lclassificatio n ofth econvariety ,an dreplacin gi twit hth e cultivar-groupconcept ,i ti spossibl et ocreat eflexibl eclassification s whichca nb eeasil y replacedafte rbein gsupersede db yfurthe r developmentsi nplan tbreeding .Thes eso-calle d openclassifications , startingfro m thecultiva rhav eprove nt ob epractica lan dusefu l for geneban kdocumentatio n(Brandenbur ge tal. ,1982 ;Brandenburg ,1983 )an dfo rregisters , statutory and nonstatutory, of cultivated plants (Brandenburg & Schneider, 1985; Duyvendak et al., 1981). The problem left is the linkage between botanical classification and cultivar classification indevelopin gon eunequivoca lclassificatio n systemfo rcultivate dplants .A t several occasions it was proposed to do so at the subspecies level (Harlan & de Wet, 1971;d eWet ,1981 ;Pickersgill ,1986 )wit hth eapparen tdrawbac ktha tsubspecie swoul d 197 bea nambiguou ster mt ob euse dfo rdifferen t entities(Brandenburg , 1983,1984a),distin guishingbot hgeographicall yan dmorphologicall ydistinc tpopulation san dwild ,weed yan d cultivatedpopulations .Proposal st olin kbotanica lan dcultiva rclassificatio n atform aleve l have similar drawbacks (Brickell, 1973;Wijnands , 1986b). 3.2. Principles ofope nclassificatio n outlined Strictlyfittin gcultivar sint oth ehierarch yo fbotanica lclassificatio nwoul dlea dt osuggestive , pseudoexact non-information, which has already given systematics a bad name. The suggestiontha tmos tcultivar sca nb eassigne dt oa specie sca nb ebelie db yjus t following thedevelopment si nplan tbreeding ,especiall yi nwid ehybridizatio nprogramme san di nth e field of incorporating desired new traits bymolecula rbiologica l methods. Theintroductio no fa specia lnomenclatur efo rhybri dcultivate dplant si sth edenia l ofthes edevelopments ,sinc efo rbotanica lclassificatio nther ei shardl yan ynee dfo rspecia l provisions in nomenclature and classification. The International Code of Botanical Nomenclature(ICBN ;Greute re tal. , 1994)admitte di ntha trespec ttha tth eprefi xnotho - to allrank s isno t obligatory. Whether ornamentals,vegetables ,frui t crops,agricultura l orsilvicultura lcrops , many cultivars are of (complex) hybrid origin. With the advanced techniques of plant breeding,suc ha sth eapplicatio no fmolecula rbiology ,somati chybridizatio nan dvariou s methodst opreven tdyin go fyoun gembryo sproduce db yhybridizatio no fdistantl yrelate d plants(Feldmann ,1983) ,cultiva rclassificatio ncanno tb elinke dwit hbotanica lclassificatio n atan ya priori rank .Althoug hthi slinkag ewil lofte n happen atth especifi c levelo ra tth e genericlevel ,i ti scertainl yno talway sth ecase .Apar tfro malread ymentione dshortcoming s ofth eICNCP ,i ti sdifficul t -i fno timpossibl e- t oclassif yunequivocall yal ldifferen t kinds ofhybrids ,a sbot hICB Nan dICNC Pcontai na se to frule sprovidin gdefinition s ofterm s and guidelines for hybrid nomenclature (Brandenburg, 1984a, 1986a, 1986b). Although the ICNCP thus far is meant to be additional and, consequently, subordinate,t oth eICBN ,th eHybri dAppendi x(Appendi x 1 ofth eICBN )i sunsuitabl efo r general usage with respect tohybrid s raised incultivation . This is due to the following 198 statements: 'Ahybri dbetwee nname dtax ama yb eindicate db yplacin gth emultiplicatio nsig n between the names of taxa' (Art.H.2) . 'When all the parent taxa can be postulated or are known, a nothotaxon is circumscribed soa st o include all individuals (as far asthe y canb e recognized) derived from thecrossin g ofth estate d seto fparen t taxa (i.e.no tonl yth eF [bu t subsequent filial generations and also backcrosses and combinations of these). There can thus be only one correct name corresponding to a particular hybrid formula; thisi sth eearlies tlegitimat enam e(se eArt .6.3 )i nth eappropriat eran k (Art.H.5) ,an dothe rname st owhic hth esam ehybri dformul aapplie sar esynonym s ofit'(Art.H.4.1). The first statement is not meant to indicate hybrid cultivars which have to be synthesizedeac hgeneratio nb ycrossin go fparen tline san dimplie stha ta hybri dcultiva r maybea ra nothotaxo ndesignation .I ti sno tobligator yt ocombin eth enothotaxo nnam e witha cultiva repithet .Th eresul ti stha tplan tpopulation swhic hmee trequirement so fth e cultivardefinitio nma yb eofte ncovere db ya nothotaxo nname ,thu sbein ghidde ncultivars . C. xjackmani iTh .Moor eha sbee ndescribe da sinterspecifi c hybridbetwee nC. viticella 'Hendersonii'an dC. lanuginosaLindl .e tPaxt. ,th elatte rno wbein gregarde d acultiva r (Brandenburg &va n de Vooren, 1986; 1988a). From the original publication (Moore, 1864)i ti sclea r thatth e author had aparticula r clonei nmind .Thi s cloneha sals obee n describedi nMoor e& Jackma n(1872 )unde rth enam eC. xjackmanii. Sinc eothe rclone s ofth esam einterspecifi c crossreceive dcultiva repithet si nmoder nlanguage ,i ti slogica lt o consider C. xjackmanii acultivar : C. 'Jackmanii'. Thesecon dstatemen tdoe sno tagre ewit hagricultura lan dhorticultura lpractic efo r plantdenominations .Sinc ei ti spossibl et obree ddifferen t cropsfro m reciprocalcrosse s (i.e.xBrassicoraphanus vs.xRaphanobrassica; Oost, 1984),i twa sconfusin g thatthe y shouldbea rth esam ename !Repeate dbackcrosse sar euse dt otransfe rdesire dcharacter s tocultivate d plants.I fry echaracter s are introduced intowheat , andi f thery e influence remainsrecognizabl eafte rbackcrossing ,th eresultin ghybri dcultivar shav et ob eassigne d xTriticosecale. They are usually regarded in common practice to be wheat cultivars. Registration authorities are left in confusion, asunde r Art. H.4.1a distinctio n between wheatan dtritical ecanno tb emad e(Baum , 1971a, 1971b;UPOV , 1984; Gupta&Baum , 1986),a sth edogmati capproac hdoe sno tprovid eu swit hth epossibilit yt onam e different 199 cropforms .Whe ndealin gwit hintergeneri ccultivate dhybrids ,i ti sworthwhil econsiderin g to define cultivar-groups. For the reciprocal intergeneric hybrids, the oldest available intergeneric name can wellb e combined with cultivar-groupepithet s making clear the varioustype so fcultivate dplant sconcerned .Th ebackcros ssituatio nma ywel lb esolve dt o definecultivar-group si nth erecurren tparen tspecies .Tritical eca nb eregarded ,doin gso , asa cultivar-grou p in Triticum aestivum L. TheICNC P(Brickel l etal. , 1980) alsocontain s rules concerning thenamin go f hybridcultivate dplants ,mainl ybase do nth emos trecen tversio no fth eHybri dAppendi xa t thedat eo fth eICNC Ppublication ,bu twit haddin gth epossibilit yo fusin gth eter mgrex . TheICNC P (Brickell et al., 1980) states in Art. 18(mark s inbol db yth epresen t author): 'Acollectiv eepithe tma yals ob ea wor do ra phras eo fno tmor etha nthre eword s in a modern language. For the purpose of this article, an arbitrary sequence of letters,a nabbreviation ,o ra numera li scounte da sa word .Al lderivative sfro mth e combination of the sametw o ormor eparenta l specieshav eth e same collective epithet in a modern language except where established custom or special circumstances demand otherwise, asfo r example,i norchids. ' Thegroupin go fseedlings ,belongin gt oon eprogeny ,i ngrege s(sing ,grex ) undercollectiv ename si sentirel ydifferen t fromcultivar-grouping .Recommendatio n18 A reads, 'Aphras euse da sa collectiv eepithe tma ycontai na wor dsuc ha sHybrid ,Hybrids , Cross, Crosses, grex (abbreviated g., Latin for swarm or flock), etc., indicating the collectivenatur eo fth eunit' .Thi simplie stha ta gre xi sbasicall ya populatio no findividual s ofth esam eparentage .Ther ei sn onee dt odescrib ecultivar swithi na grex ,but ,o fcourse ,i t ispossible . So,a gre x can be an independent entity,par t of which does not necessarily meet therequirement s of the cultivar definition. Looking atclassificatio n and registration of cultivated orchids according to the ICNCPan dth especia lHandboo ko norchi dnomenclatur ean dregistration ,on emus tcom e to the conclusion that with the purpose to create clarity and an unequivocal, stable nomenclatureth eresul ti sconfusing ,leavin gth euse rwit hname dseedlin gpopulation stha t aresometime sonl ysegregatin gpopulations ,bu tma yals oconcu rwit hth edefinitio no fth e cultivar.Th econclusio ni stha tforthcomin gedition so fth eICNC Pshoul dneithe rcontai n statements concerning grex nor references to the Hybrid Appendix, except for the 200 pragmaticon et oth enothogenu st owarran ti nal lcase sth econnectio nbetwee nbotanica l andcultiva rnomenclature .I nth eforthcomin g editiono fth eICNCP ,article sconcernin g grexwil ltherefor eb edelete dwit hth eon ereferenc et oth efac ttha ti ti spossibl et omak ea n informal classification bygreges . Theter m cultivar-groupi s defined in the ICNCP (Brickell et al., 1980),Art .26 : 'Whena species ,interspecifi chybri do rintergeneri chybri dinclude sman ycultivars , anassemblag eo fsimila rcultivar sma yb edesignate da sa group .Thi scategor yi s intermediate between species and cultivar. It is not an essential part of the full cultivarname .I fuse dbetwee nth especie snam eo rcollectiv enam ean dth ecultiva r name,th enam e of the groupi splace dbetwee n parentheses (roundbrackets). ' This definition contains some contradictory elements: "Interspecific or intergeneric hybrids may also include assemblages of similar cultivars," so cultivar-groups, making it impossible to rank cultivar-groups exclusively intermediate between species and cultivar! thewor d"essential "ha sbee nmisplace di nArt .26 ,becaus eo fwha ti sstate di nart . 27,secon d sentence,th ecultivar-grou p namei sno ta par to fth ecultiva rnam ea t all,le talon ea nessentia lpart .Althoug hi ti sno ta nessentia lpar to fth eful l cultivar name,ther ei sa nindicatio n wheret oplac ea cultivar-grou p designation in a full cultivar name, but there is nowhere else in the Code a guidance how to form cultivar-group names,excep t implicitly inth eexample s given for Art.26 . Therefore, itwoul db edesirabl et oinclud erule sprovidin ga statemen tho wt oestablis ha cultivar-group,base do non eo rmor estandard ss otha teverybody ,n omatte rwher ei nth e world,ca nmos tlikel ydetermin eo ngenera lappearanc ewhethe ra cultiva rca nb eassigne d to a certain cultivar-group under local circumstances. This is especially important for obligate cross-fertilized crop plants like many vegetables and fruit crops.Moreover , a statementshoul db emad ea st owhethe ra cultivar-grou pi so floca lsignificanc e ori smean t tob e valid worldwide. Anot e added to Art. 26states : 'In complex crops, for example, in apples and some cereals, a hierarchy of categories has been applied, the use of which isno t governed bythi s Code.' Thisnot ei sconfusing , sinceth econcep to fcultivar-groupin gdoe sno tagre ewit ha concep t ofhierarchica lrelationship samon gcultivar-groups .A hierarch yunnecessaril yhamper sth e connectionwit hbotanica lclassificatio n and,consequently ,nomenclatur ea sgoverne db yth e ICBN.O nth econtrary ,thi snot eshoul dexplicitl ystat etha tcultiva rclassificatio ni ncultivar - 201 groupsreplace smor ehierarchica lcultiva rclassifications ,eve ni ncomple xcro pplants .Th e currentconcep ti ssimpl ean di ti stherefor efeasibl ean dconceivabl et ojudg ewhe nan di fa cultivar-groupclassificatio n shouldb ereplace db ya mor eu pt odat eone .I tthu sdescribe s current variation inlarg e assortments which undergorapi d changeb yplan t breeders' efforts. Themos trecen teditio no fth eICNC P(Trehan ee tal. , 1995)contain s statements onth enomenclatura l consequences of mergingo r splittingcultivar-group s toadap tth e classification toshiftin g cultivarassortment s (Art.4.6) .I nbot hcase sol d cultivar-group namesshoul db ereplace dt oavoi dtha tth esam enam ereflect s variouscircumscriptions . Althougha tfirs tsigh tthi swil lno tserv ea stabl enomenclature ,i tdoe sjus ttha tb yit ssimpl e connection toon ecircumscription , asoppose d toth eforme r convarietal classifications, whichstrictl yfitte d thebotanica lhierarchy .Circumscriptio n impliestha t distinguishing characters are presented with the included character states, whereas description omits ranges of variation. Circumscription thus provides arepresentativ e text concerningth e identityo fth eparticula runi tconcerned .Descriptio nonl yprovide sa diagnosti ctext ,whic h isno tnecessaril yrepresentativ e inal lit saspects .Thi sdifferenc e necessitates aseparat e basicconcep ti nclassificatio n ofcultivate dplant san dform sth emai ndifferenc e between cultivar-group classification and former convarietal classification because of its consequences (seefigur e 3.1; sections 3.3.an d 3.4.). 202 Subspecies GEML J bot. Species var. 203 Figure3.1 . a- Botanica l(closed )classification , ahierarchica lsyste mo fmutua lexclusiv etax a(genus , species, subspecies,botanica l variety). b- Ope nclassificatio no fcircumscribe dcultivar s(the yma ypartiall yo rwholl yoverlap) ,lai d overth ebotanica l classification of 3.1a. c -Anothe r open classification ofcircumscribe d cultivars (genus,species ,cultivar) . d -Ope n classification of cultivars of lc, but partially classified in two cultivar-groups (dottedlines) . 204 The cultivar-group concept has already proven its usefulness, as can be learned from several examples: Lactuca sativa (Rodenburg, 1960), Dahlia (Anon., 1969), Lilium (Leslie, 1982),Brassica rapa (Oos t &Toxopeus , 1986),Clematis (Brandenburg &va n deVooren , 1982,1986,1988a),Narcissus (Donald , 1986),Dracaena (Bose tal. ,1992) , Aster (Hetterscheid & van den Berg, 1996), Philadelphus(Hoffman , 1996) andBeta vulgaris (Lange et al., 1999). From these examples, it is clear that cultivar-group classification should agree withcommo npractice .Sinc ei t mustb eusefu l for breeders, growers,registrars ,merchant san dal lothe rconceivabl euser s(no tt oforge tth e"public") ,i t should rely on clear characters, directly connected with growth conditions and/or consumer's usage.Biosystemati crelationshi p(degre eo frelationshi p anddescendance) , therefore,i sno ta soun dbas efo rcultivar-grou pclassification .I tma yagre ei nsom ecases , buti tusuall ywil llea dt oconfusio n asca nb eexemplifie d withRosa. Althoug hbreeder slik e torefe rt odescendance ,Polyanth arose san dFloribund arose scanno tb edistinguishe di na n experimentalfield .I ncommo npractice ,thes egroup shav en osignificanc e atall .A bette r procedurewoul db et odefin eRosa group so ncharacteristic ssuc ha sflowe rsiz ean dflowe r colour, occasionally combined with other economically important traits (Buck, 1967). Another reason not to use biosystematic evidence is that documentation about old, sometimesextinct ,cultivar si softe nver ypoo ran dstatement sabou tthei rdescendanc ear e not rarity either false, putative orjus t lacking. Requiring biosystematic criteria would precludecultivar-grou pclassificatio n inman yassortments .Thi swoul db eth ecas ei nman y ornamentals,a si nClematis (Brandenbur g& va nd eVooren ,1986) .Furthermore ,cultivar - groupswoul dthe nsugges tbiosystemati crelationship sbetwee ninvolve dcultivars ,whic h canb eraril yproven .Cultivar-grou pclassificatio nbase do nbiosystemati ccriteri aseem sa service to plant breeders. This will often not beth e case, sincebreeder s would then be lookingfo rne wbreedin gmateria lunde rsupposedl yallie dcultivar swhic hma yb ei nfac t genetically distant (Baum, 1981). Various registration authorities, both statutory and nonstatutory, have already included cultivar classifications in their registers, despite the absence of any article or recommendation inICNCP . If cultivar-groups are meant to be applied worldwide, for reasons of stability, it is 205 recommendedt ous eth eoldes tor ,i ndoubtfu l cases,th ecurren tnames ,a si sdon ewit hth e cultivarclassificatio n of Clematis (Brandenburg andva n deVooren , 1986).Sometime s common crop names, if not confusing and if not in conflict with Art. 27,ca n be used (Brandenburg, 1984b;Schneider , 1984).I ti seve npossibl et ohav etranslation sfo rthe m through theMultilingua l Glossary of Common CropName s (Koster and Schneider, 1982). In case of cultivar-groups of local significance, growth conditions or easy identification characteristics areofte n indicated in the names.A sperformanc e in other environmentalcondition sca nb equit edifferent , thecontex to fth eclassificatio n shouldb e stated clearly. In conclusion, cultivar-groups are useful in cultivar classification and serve to standardize common practice without implyingbiosystemati c relationships within thegroups . 3.3. Thetaxo nconcep t andcultivate d plants Inorde rt ounderstan dth ereasonin gbehin d theintroductio n ofa ne wbasi cter mfo r the taxonomyo fcultivate d plants,i ti sworthwhil e considering thebackground s ofth eter m taxon, as it is used in both botanical and zoological taxonomy. In 1926,Meyer-Abic h introducedth eter mtaxo na sa philosophica lconcep toppose dto "phylon" .It sapplicatio n wasneve rfollowed .I nJun e 1948,a conferenc ewa sconvene di nUtrech t(th eNetherlands ) todiscus san dprepar eproposal st oamen dth eICB Na tth eStockhol mBotanica lCongres s tobeheldin 1950.La m(Proceedings ,Chron.Bot . 12)proposedto'..indicat ea taxonomi c groupo fan yran kwit hth eter mtaxon... ' (p.12) .Th eexac twordin go fth eproposa lt ob e submittedwas,'.... ;taxonomi cgroup so fan yran kwill ,i nth eRules ,generall yb ereferre dt o astax a (singular: taxon);...'. This proposal was accepted and incorporated in the 1952 ICBN but the word "rank"ha dbee nchange d into "category",whic hwa schange dbac ki nsubsequen tcodes . Thewordin go fthi sproposa lwa slate rgive nth estatu so fa separat earticl e(art.l) ,whic h has not been changed since. Theintroductio no fth ewor d"taxon "i nth ebotanica lsociet ywa sthu sdefinitel ytie d toranking .Tax a are assigned tocategories ,whic h arepar t of anaxiomati c hierarchical 206 ranking system (thetaxonomi c hierarchy). Zoologists have adopted the word taxon as used by botanists. Simpson (1961) devotes anextensiv e discussion onth e subject: 'Ataxo ni sa grou po frea lorganism srecognize da sa forma l unita tan yleve lo fa hierarchic classification'; 'Ataxonomi ccategor yo rsimpl ya categor yi sa clas sth emember so fwhic har eal l thetax aplace da ta give nleve li na hierarchi cclassification .Th eran ko fa categor y iseithe r its absolute position in agive n hierarchic sequence of categories or its positionrelativ et oothe rcategories .Th eran ko fa taxo ni stha to fth ecategor yo f which it isa member. ' Taxa,thu stie dt oth etaxonomi chierarchy ,sho wimportan tmutua lrelations ,viz .exclusio n andinclusion .Tax ao fon eran ki non eclassificatio n alwaysexclud eeac hother .Tax ai non e classification aton eran kar eal linclude di non eo rmor etax aa tth enext-highe rrank .Thi s leadst oth etota lexclusio no foverla pbetwee ntax ai non eclassificatio n (seefigur e 3.1a). Overlapbetwee n taxa with the samenam ebu t figuring in different classifications, isusual . Since taxa areth e main subjects in evolutionary discussions,th e taxon concept shouldb erestricte dt orepresen tgroup so f organisms thatar ebase d on theevolutionar y assumptions(thei rontology) .An ds othi sleave sn oroo mfo rgroup so fcultivate dplant st o betreate d asprope r taxa, asthi s would createnon-evolutionar y "noise". Iti sthu s apparent thatth etaxo n concept nowadays has twonotions : aclassificator y devicetie d toth etaxonomi c hierarchy; anevolutionar yconnotation ,whic hi sbes texpresse db yth ewidely-hel dview ,that , if taxahav e tohav e anyrelevancy , they should bemonophyletic . Consequently,asid efro m agenera lclassificatio n -reflectin g theresult so f evolutionary discussions -i n the sense of McKelvey (1982), special classifications may serve other purposes. Simpson (1961) states: 'We must thus accept the possibility and in fact the need not only of many classifications butals oo fman ykind so fclassifications , thatis ,o f classifications based ondifferen t sorts of relationships and serving different purposes.' 'Teleologicalclassification sdefin eset s(again ,no ttaxa )accordin gt othei rusefulnes s or lack of it, usually with respect to man. Such sets might be, for example: domesticatedanimals, [ ].The yd ono thav emuc hgenera lscientifi cinterest ,an d againi nmoder nusag ethe yrequir eth eprio rclassificatio n oforganism so nsom e other system.' Thisdefinitio no fteleologica lclassification s isdirectl yapplicabl et oth ephilosoph ybehin d 207 classifying cultivated plants. Schwanitz (1967) defines cultivated plants as follows: 'Die Kulturpflanzen sind das Ergebnis von Evolutionsvorgangen, die sich in vorgeschichtlicherun di ngeschichtliche rZei tbi si nunser eTag ehinein ,teil sunte r demunmittelbaren ,teil sunte rde mmittelbare neinflu Bde sMensche n vollzogen habenun d heutenoc h vollziehen.' Schwanitz'definitio n showsambiguities .O nth eon ehand ,classificatio n ofcultivate dplant s startsfro mevolutionar yprocesses ;o nth eothe rhan di tinclude shuma ninfluence ,albei ti n part indirect. It stresses the very beginning of the domestication process, whereas the nowadays general process of domestication (formulating the demands of a new plant, breedingan dreproducin git )i sunderemphasized .A mor emoder ndefinitio n ofa cultivate d planti saccordin gt oHetterschei d &Brandenbur g (1995a),adapte d from Trehane(pers . comm.): 'A cultivated plant is one, whose origin or selection is due to the activities of mankind. Such plants may arise either by deliberate or chance (garden!) hybridizationo rb yfurthe r selectionfro mexistin gcultivate dstoc ko rthe yma yb e selected from a wild population and maintained as an entity by continuous cultivation.' Therelevan tpoin ther ei semphasize db ydeliberat ean db yman .Plant sar ecultivate db y man, deliberately to improve his standard of living, either by improving his diet (agriculture/horticulture),surroundin ghimsel f withornamenta lplant s (horticulture),o r improving/ maintainin ghi senvironmen to na large rscal e(silviculture/forestry) .I norde rt o do so, man has to manipulate plants. From the moment a plant is taken from nature (selected)b yma nan dpropagate do rmaintaine dunde rhi sow ncontrolle dcircumstances ,i t isn olonge rexclusivel ysubjec tt oth eforce so fevolution .Fro mirrespectivel yan ysurve yo f plant breeding research and results, it can be concluded that design, production and reproduction of new cultivars are labour-intensive human activities. Sinceth ecultiva ri sth ebasa luni to fcultivate dplan tclassification , itsnatur ei sall - importantt oth equestio nwhethe ri tca nb ea taxo no rnot .Th eman yan dofte n complicated waysi nwhic hcultivar sar eproduce dan dreproduce d(ICNCP ,Brickel le tal. ,198 0art . 11; Trehanee tal. ,1995 ;art .2.7-2.17) ,illustrate sthei rstatu sa sproduct sof , often large-scale, commercial industry. Forinstance ,present-da ytechnologica ldevelopment slea dt oth eintroductio nof "synthetic " 208 cultivarsi nwhic hgeneti cinformatio n ofwidel ydifferen t taxa(o rcultivar sderive d from them) isbrough t to expression. These cultivars will defy any attempt of ranking inth e taxonomichierarch y(se eGrosse r& Gmitter , 1990,fo ra nexampl eo fsyntheti ccultivar si n Citrus). Grosser andGmitter' spape r showsth eradicall y different natureo f "processes" (and order in which they are allowed to operate), leading to cultivars, compared to processes in natureleadin g toe.g . species. Another important, non-taxon, character of cultivars is homogeneity and its retention duringreproduction . Thistypicall y industrialdeman d serves togai n andkee p confidenceo fconsumer san dgrowers .On emus tb eabl et osafel ybu ya larg estoc ko fe.g . immatureplant s(o rplant-parts )o fa cultiva rtha tmeet scertai ndemand san dthat ,afte r a while,indee dshow sth erelevan tcharacter sinstea do fa narra yo funwante dvariation .Tax a on the other hand, are conceptualized to show variation as a result of evolution and, consequently,ar edescribe db ytaxonomist sallowin gan dincludin gthi svariation .Contrar y to this, present-day newly developed cultivars that show much variation are usually regardeda sinferior ,a totall ydifferen t (teleological)approach .O nth eothe rhand ,primitiv e cultivarso rlandrace ssho wthi sunwante dvariation ,bu thav eth eusefu l plasticitya sthei r purpose, and stilld oharbou r genestha tca nb e selected. Cultivarscanno tg oextinc t ase. gspecie sd oi nnature .Materia lo fa cultiva r may disappear from the face of the earth for a certain time but the same cultivar may be reconstituteda tan ytime ,i fparenta lmateria li sbein gconserved .Thi si sespeciall yclea ri n hybrid cultivars that arebein g created again every season bymakin g aparticula r cross betweenmaintaine dinbre dlines .Ne wtechnique smak ei tpossibl et oprepar ea 'recipe 'fo r obtaining(synthesizing )a certai ncultivar .Furthermore ,i ti sstate di nth eICNC P(Brickel le t al., 1980art . 10,not e 1:Trehan e et al., 1995 art.2.18 , art. 3.1), that cultivars are being recognizedirrespectiv eo fthei rwa yo forigin .Whe nplant shav ebee nproduce dtha texactl y matchth edescriptio no fa nexistin gcultiva r(o ron eo fwhic hn omateria li sextan t atth e time),the yar eregarde dt ob etha tcultivar !Th ecultiva rtherefor ei sphilosophicall ya clas s concept. The "class" (the cultivar) is being circumscribed and all plants meeting the circumscriptionar egroupe di nth eclass .Th eclas sma yb eempt yfo ra whil ewhe nn oplan t existstha tfit sth ecircumscription ,bu tsuc hplant sma y"appear "a tan ytim ean drefil lth e 209 class(th ecultiva rcome sint oexistenc eagain) .Thi si stypicall yth e"behaviour "o fa class like entity. Logically any grouping of cultivars is a class as well, whereas taxa are singularitiesi nnatur e(the ycom eint oexistenc ean ddisappea rbu tneve rreappear )an dar e regarded individuals. Cultivars thus result from amixtur e of natural and man-enforced processes,th e influenceo fbot hprocess-group sbein gver ydifferen t fromcultiva rt ocultivar .Thes edays , thedevelopment ,maintaintenanc ean dmultiplicatio no fne wcultivar shas ,i nman ycountries , developedint oa genuine ,large-scal eindustry .Man yne wtechnique shav ebee ndevelope d andth eapplicatio no fhybridization ,somati cfusion ,mutation-breeding ,geneti cengineering , etc. enhances the awareness that the final result - the cultivar - is a typical "industrial product" andmuc hles sa membe ro fMothe rNature .Thi sincrease d shift inontolog yha s not been recognized by most taxonomists/systematists. A classification of such man- influenced and/or man-made entities,canno t beterme d "natural". In view of the above-stated, a classification of cultivars can only be a special- purposeclassification ,thi spurpos ebein gman' schoic eo fan yse to fcharacters ,relevan tfo r thecultivatio nan dus eo fcertai ncultivars .Suc hclassification s areteleological ,an ddefin e classes andno ttaxa !I nconclusio n systematic categories of cultivatedplant snee dt ob e recognized asa ne w systematic concept, different from the taxon concept. 3.4 Theculto n concept Thepresent-da y useo f thetaxo n concept obviously doesno tproperl y coverth e essencean didentit yo fcultivate dplants .Th econceptua lshif ti nlookin gupo ncultivar sa s industrialproduct sbein gmanufacture d insteado fevolving ,call sfo ra ne wconcep twit ha teleologicalinclination .I ti spropose db yHetterschei d& Brandenbur g(1995b )t ointroduc e the term "culton (plural: culta)" into the systematics (including classification and nomenclature) of cultivatedplants : Aculto ni sa grou po fcultivate dplants ,base do non eo rmor euser-criteria . Aculto nmus thav ea nam eaccordin gt oth erule so fth eInternationa l Code of Nomenclature for CultivatedPlants . 210 Theculto na sa classification-categor y isessentiall ydifferen t frombotanica lcategorie sa s referredt oi nICBN .Cult aprincipall ydef yextensiv ehierarchica lrankin ga ssee ni nprope r taxaan dar eno tnecessaril ycomplementar yt oeac hothe ri na classificatio n (seebelow) .A cultonan da taxo nma yb eentirel yo rpartl yo rtemporaril yco-extensive .I norde rt oavoi d any further conceptual and practical confusion between taxa and culta, a number of nomenclaturaldevice srelevan tt oth enamin go fcult amus tb ereformulate d oromitte di n bothICNC Pan dICBN .Invokin gth eculto nconcep tan dit snomenclatura l ramifications justlystresse sth eonl yrelevan tessenc eo fcultivate dplants ,tha ti simprovin gman' sstandar d of living. The term "culton" is not entirely new. During discussions at past horticultural congresses(Hamburg ,1982 ;Davis ,1986) ,th eide atha tgroup so fcultivate dplant sd ono t behavea sprope rtaxa ,wa srecognized .A tth eDavi scongress ,F .Schneide r(Netherlands ) and C. Brickell (U.K.),jointl y proposed to introduce the term "culton", with a similar inclination asuse d here,viz . parallel to "taxon". The discussion stopped there and the introductiono fth ene wconcep twa sno tpursue dan yfurther .Afte rman yyear so fobscurity , theter mwa ssuddenl yintroduce di nth e"glossar yo fplan ttaxonomy "i npar t1 o fth eNe w RoyalHorticultura lSociet yDictionar yo fGardenin g(1992) .It sdefinitio n theredoe sno t nearlydescrib e itsorigina l intention accurately. TheDictionar y states: 'ataxonomi cuni tdescribin gdistinc tplant soriginatin gi no rmaintaine di ncultivation . This term has been proposed to reflect the fact that not all such entities can be consideredo rtreate da scultivars .I tstand si nsomewha tartificia lcontras tt otaxon , whichi twa shope dwoul dcom et omea na uni to fnamin gfo rwil dplant sregulate d byth eBotanica lCode. ' Forreason sstate dabove ,a culto nencompasse s(not describes )group so fplant san dnot justplants .Th esecon dsentenc espeak sonl yo fcultivars ,wherea sculto ni sno trestricte dt o thatcategory .Althoug hi ti sstate dwha tth ewor dtaxo nwa shope dt ocom et omean ,i ti s apparentfro mit spresen tda yusag etha tthi si sexactl ywha tit smeanin gi stoday ,namel ya unit of naming wildplant s and solegitimisin g theintroductio n of theculto n concept. Thecriterio no fusefulnes s isbasi ct oclassification s ofculta .Th echaracter supo n whichsuc h classifications mayb ebase dar eentirel ydependen to nman' ssubjectiv echoice . 211 Therefore,culto nclassification san dtaxonomi cclassification s arever ydistinct .Fo rinstance : toa grou po fpeople ,i tma yb erelevan tt oa grou po fpeopl et odivid ea numbe ro fcultivar s intoanumbe ro fcultivar-group sbase do nflower-colour . Foranothe rgrou po fuser si tma y bemor eimportan tt ogrou pth esam ese to fcultivar s onth ebasi so fpest-resistance .Th e resulti stw odifferen t classifications forth esam ese to fcultivars .The yca nonl yb ejudge d onthei rusefulnes s andthe yma yb erejecte d forlac ko fi tbu tbot hma yals ob eretained .I n such acase ,on ecultiva r mayb efoun d in twodifferen t cultivar-groups without anyone bothering. Contraryt othis ,whe na give nnumbe ro ftax ai sclassifie d inmor etha non eway ,a taxonomistwil linevitabl ychoos eon eclassificatio ntha th ethink sbes treflect sevolutionar y relations amongth etaxa .Th emer efac t thattaxonom y workswit h theaxio mtha tlif e is monophyletican da ssuc hsupport sonl yon etru ephylogeny ,lead st oth estatemen ttha to f twoo rmor ealternativ eclassification s ofth esam ese to ftaxa ,a tmos ton ema yb eth e"true " one! Choicetherefor ei sinevitabl ean dinheren tt oth esystem ,contrar yt oth eabov estate d for culton classifications. Theextensiv ehierarchica lnatur eo ftaxonomi cclassifications ,force sth etaxonomis t todefin e (often morphologically) everytaxo n aboveth especie sleve lt owhic ha certai n speciesi sassigned .Startin gfro m thecultiva r(basa lcategory) ,suc ha mechanica lhierarch y (see e.g. Jirasek, 1966) would lead to anumbe r of subordinate cultivar-groups (orne w categories), which have to be defined at every "level". The required large number of characterst odefin ethes egroups ,woul db eforce dupo nth eclassification ,whic hwoul db e inseriou sconflic twit hman' sfre echoic efo ron eo ra fe w "useful" (teleological)character s topropos ea satisfactor y "one-level"classification .Anothe rdrawbac kwoul db ea genera l inflationo frank san dcategories ,wherea sw efee ltha tth esingl ecategor yo fcultivar-group , could encompass all classificatory needs presently known above the cultivar "level" (Hetterscheid &Brandenburg , 1995a, 1995b;Hetterschei d et al., 1996). Complementarityi sanothe raspec tabsen tfro mculto nclassification .Whe na larg e numbero fcultivar so fa cro pi si nnee do fcultivar-grou pclassification ,th echoic efo r useful characterswil ldefin eth enumbe ro frelevan tcultivar-groups ,bu tthes ema yno tnecessaril y coveral lcultivars .Inevitabl ya numbe rma yremai ntha td ono thav ean yo fth echaracter s 212 defining thecultivar-group s (seeBo se t al., 1992fo r an example inDracaena fragrans KerGawl.) .I nthi s situation therei sn onee d for amechanis m that would automatically assignth eremainin gcultivar st oa cultivar-group .Thi sgrou pwoul dhav et ob edefine db y thenon-existenc e of characters,whic hi scontrar yt oth epositiv echoic eo f characters to define cultivar-groups. Iti s also contrary toclassificatio n philosophy ingeneral . Classificationsi nwhic hentitie sa ton eleve ld ono tnecessaril yfil ltha tleve lentirel y toproduc eth enext-highe rleve lentity ,ar edesignate dope nclassification s asoppose dt o closed classifications, like taxonomic ones (Brandenburg et al., 1982; Brandenburg, 1986a).Classification so fcultivate dplant sare ,b ythei rnature ,ope nclassification sb yvirtu e of the absence of obligatory hierarchy and complementarity. 213 4. CLASSIFICATION OFCULTIVATE D CLEMATIS 4.1. Clematisa s a gardenflower - th efirst moder ntreatmen t ofcultivate dClematis In 1872, Moore & Jackman published their book on cultivated Clematis, entitled 'Clematisa sa garde n flower'. Alreadyafte r publication itbecam eth ever yhandboo ko n thesubjec tan dremaine ds oa tleas tunti lth enineteen-thirties .Havin gproduce da standar d for Clematis, Moore and Jackman also realised that both growers, amateur and professional, need quite different classifications than botanists do.Th e subtitle of their secondchapte rcontain sth emeaningfu l statement:'Classificatio n unimportantfo rGarde n purposes -Cultura l Classification preferable'. Twomor ecitations : 'Thesectiona lgroups ,then ,whic hw esuggest ,ar eintende dt ob estrictl ycultura l and seasonal, and aret ob es oregarde d -i nfact , asbein g framed entirely for the guidancean dconvenienc eo fth ecultivator ,an dno ta shavin gan yspecia lrelatio nt o thebotanica l affinities of thevariou s plants'; 'Thismod eo fclassificatio n willb efoun d tobrin gtogethe ral lthos e speciesan d varietieswhic h aresimila ri nhabi tan dcharacter ,an dwill ,moreover ,assis tu si n arranging,i nsom eintelligibl eorder ,th einstruction sw eshal lhav et ooffe rregardin g thecultivatio n of the different types ofClematis.' Theseed so fth ecurren tdefinitio n forth ecultiva rgrou par ealread ypresen ti nth eabov e quotations.Th e"sectiona lgroups "b yMoor ean dJackma nconsis to fspecies ,hybrid san d cultivars. By lack of a separate term distinguishing botanical varieties and cultivated varieties,i t was logical nott obothe r toomuc h about it, although their nomenclatureo f (cultivated) varieties isalmos talway si nmoder nlanguage ,a si ti sb yothe rproducer so f cultivars attha t time,an d asi t was inothe r ornamentals, such asRosa cultivars . The Moore &Jackma n system of "sectional groups", referred to as e.g. Patens type,appeare dt ob ever ystabl ea sfa ra slarge-flowere d Clematiscultivar sar econcerned . Thecultiva rgrou pclassificatio n oflarge-flowere d Clematis, asi ti spresente di nth enex t sectioni slargel ybase do nth e"sectiona lgroups "o f1872 .Fo rth esmall-flowere dClematis groupingsar eles sstable .Moor ean dJackma n(1872 )define dth eGraveolen styp earoun d the species C. campaniflora, C fusca, C. grahami, C. grata, C graveolens, C. 215 orientalis, C. viorna,C. virginiana andC. vitalba. Thisi sa rathe rpeculia r combination of species,showin ggrea tdifference s ingrowt h performance, pruningrequirement s and vegetativepropagation ,a swel la sornamenta lvalue .I ncurren tcultiva rgroups ,onl ycultivar s figure.Therefore ,thi san dother "sectiona lgroups "i nsmall-flowere d Clematishav eneve r caughto n ascultiva rgroups . 4.2. Cultivar classification of large-flowered Clematis Thedistinctio nmad ebetwee nsmall-flowere dan dlarge-flowere d Clematisha sbee nan di s rather arbitrary, andi sbase do ntraditiona lusag ean dcurren t practice andno t strictlyo n flower size. Large-flowered Clematiscultivar s are considered to belong to C. florida Thunb. ex Murray, C. lanuginosa Lindl. and Paxt, C.patens Morr. and Decne., C. viticella L., andC. texensis Buckl.,o r areinterspecifi c hybrids between twoo rmor eo f thesespecie swit hvariou sdegree so fintrogressio nfro m oneo rmor eo fth eothe rspecies . Small-flowered Clematiscultivar sar econsidere dt obelon gt oth eremainin gspecies .Thi s distinction was alreadyformulate d bySpingar n (1935) andha sbee n applied sincethen . To classify large-flowered Clematis cultivars, it is important to know the characteristics of theparenta l species (Brandenburg, 1981,1984c, 1985,1989a, 1989b; Brandenburg &Va nd eVooren , 1982,1984,1986,1988a, 1988b). Asfa r asrelevan t for cultivation,thes echaracteristic sha sbee nuse dfo rth ecircumscriptio no fth ecultiva rgroups . Thecultiva rgroup sar eo fmondia lsignificanc eand ,excep tfo rth eTexensi sgroup ,stabl efo r moretha n onecentury . Asurve yo fthes ecultiva r groupsi spresente d intabl e4.1 . More details will be presented in the forthcoming checklist of the large-flowered Clematis cultivarso fth eInternationa lClematis Registe r(Brandenbur g& Va nd eVooren ,i nprep.) . Jouin (1907) and Spingarn (1935) already produced authorative surveys of Clematis assortments after Moore andJackma n (1872). 216 Table 4.1. Cultivar groups of large-flowered Clematis. Florida group Plants flowering on the old or ripened wood, mostly with semi- doubleo rdoubl eflowers ; flowering timespring-summer ; woody climbers. Jackmaniigrou p Plantsprofusel yflowerin go nth eyoun ggrowt hdurin ga lon gperiod ; flowering time summer-autumn; woodyclimbers . The group isoriginall y based on C. 'Jackmanii' (C. xjackmanii Th. Moore) Lanuginosagrou p Plantsflowerin go nshor tsid eaxe so nth eyoun ggrowth ;ver ylarg e flowers spread over the whole plant; flowering time summer- autumn;wood yclimbers . Patensgrou p Plantspredominantl yflowerin g onth eol do rripene dwood ;mostl y with singleflower s havingpointe dtepals ;flowerin g timespring - summer; woodyclimbers . Texensisgrou p Plantsprofusel y floweringo nth eyoun gwoo ddurin ga lon gperiod ; +bell-shape d flowers; flowering time summer; subshrubs. Viticellagrou p Plantsprofusel y flowering duringa rathe rshor tperiod ; flowering timesummer-autumn ; woodyclimbers . 4.3. Introduction intocultivatio n of Clematis sect.Meclatis Asearl ya si n1700 ,Clematis orientalis wa sintroduce dint ocultivatio nb yTournefor t (for details,se e2.4.2) .I ti sremarkabl eho wfas tth especie sha ssprea dove rEuropea nbotani c gardensan dprivat egarden so fwealth ybanquer san dmerchants ,a sca nb esee nfro mthei r correspondence(Va nde rNeut ,1983) .A tth esam etime ,Dilleniu sdescribe dth especie si n Hortus Elthamensis (1732),i tha dbecom e awell-know n species inth eNetherlands ;i n Leidenan dAmsterdam .Althoug hth especie swa sgrow nal love rEurope ,n ocultivar swer e selectedfro m it.Fro mth ebeginnin go fth e 19thcentur yonwards ,plant swer eintroduce d from other areastha n the original provenance (Turkey). 217 Willdenow (1796) described the species C. glauca in 'Berlinische Baumzucht, oder Beschreibungde ri nde nGarte nu mBerlin ,i mFreie nausdauernde nBaum eun dStraucher , fiirGartenliefhabe run dFreund ede rBotanik' .Th edescribe dplan twa salread ygrow ni nth e BerlinBotani cGarde nsinc e 1752unde rth enam e C. orientalis.A ne wsee dlot ,receive d by Willdenow a few years before 1796, induced him to compare his plants with the Dillenianplate .Base do nleafle tmorpholog yan dth eindumentu mo fth etepals ,h edecide d to describe both accessions as a new species: C.glauca. His extensive description in Germanan dth ebeautifu laccompanyin gplat ear eth efirs to fman ystatement si nhorticultura l literatureho wt odistinguis hC. glauca fro mC. orientalis,thu sresultin gi ndivers eattitude s concerningth evariabilit yo fbot hspecies .Th eorigina lherbariu mspecimen so fC. glauca prove to belong to C. orientalis. C. glauca is therefore a heterotypic synonym of C. orientalis. Inhorticultura l literature,th enam eha sobtaine d aseparat emeaning .Thi sha s complicated Clematis sect.Meclatis nomenclatur ea grea tdeal .Apar tfro m glaucabein g aninfraspecifi c epithetoni nvariou scombinations ,ther ewa sstil la par to fth evariatio ntha t mightb econsidere da separat especie san dwhic hanyho wperforme dquit edifferentl y from C.orientalis. Theseplant s originate from Siberia, Mongolia andth eNorther n plainso f China.I twa sBunge ,wh orecognize dthi si n 1833 andh edescribe da separat e speciesC. intricata(cf . Bunge, 1854). The species itself had been distributed over many botanic gardens,althoug hi twa susuall ylabelle dC. glauca.C. glauca referre d alsot opopulation s ofC. orientalis. So,th erang e of variation inC. intricata had never been clear. Inth emiddl eo fth e 19thcentury ,ther ewer evariou sbotani ctraveller s collecting specimensi nCentra lAsia ,an dth efa reas to fRussia ,Mongoli aan dChina .The ydi dthi s eithero nbehal f ofthei rmissio nb yGovernmen t (Bunge,Przewalski ,Potani nfo r Russia; Clarke,Edgeworth ,Griffith , Stracheyan dWinterbotto m forEngland )o ra smissionarie s (David from France) (cf. Bretschneider, 1898).Thei r collections gavea ne w impulsei n Clematisgrowing .Man ylarge-flowere d Clematisresulte dfro m theseactiviities ,an dth e interesti nsmall-flowere d specieswa sdefinitel yaroused .Systematist sdescribe dth evariou s forms ofMeclatis either asseparat e specieso ra sinfraspecifi c combinations.Th eplant s wereintroduce dint ocultivatio na sseedlin gpopulation san ddi dno treceiv efurthe rattentio n ast o selection. 218 Complicating aspects of the introduction into cultivation were the discrepant interpretations bybotanist s andhorticulturists .C. graveolens asdescribe db yLindle yi n 1846,wa sknow nb yit sheav ybu tunpleasan t scent.Th eplan tdepicte d anddescribe db y Hooker under the name C. graveolensi n Botanical Magazine t. 4495 was all but C. graveolensLindley .Afte rHooker' spublication ,th enam ewa sambiguou si nit smeanin gi n horticulture. The later published C. vernayiC.E.C.Fisch . -her e reduced to C. tibetana subsp.vernayi - fit sth eplat eb yHooke rperfectly .Th esam ehold sfo rth eintroductio no f C.tangutica (Maxim. ) Korsh. intocultivation . The publication by Andre (1902;More l actually wrote the paper!) contributes much to recognize its value for the Clematis assortment.Horticulturist safte rhim ,however ,gav earestricte dinterpretatio no fth emateria l involved, ending up with two views about C. tangutica: s.s. sensu Andre (according to horticultural literature) and the original view, s.L, by Maximowicz (1889), with his description of C.orientalis var.tangutica. Inth e20 thcentury ,C. tangutica wasdistribute dove rth eworl dan dbecam esoo n verypopula ramon gClematis enthusiasts .Whe ni nth elat ethirtie sC. vernayials obecam e available,th einteres t for yellow-flowered Clematisincrease d again:i nth eNetherlands , therewer ebreedin gan dselectio nprogramme sa tBoskoo pan dinitiall yals oa tWageningen , ofwhic hsom ecultivar sar edescribe di nth enex tsection .Th ever yreaso nbehin di twa san d -ma yb estil li s- th ewis hfo ra yellow ,large-flowere d Clematiscultivar ,whic hwoul dse t off another trend in Clematiscultivation . In thebeginnin g of the 20thcentury , C. serratifoliawa srecognize d as adistinc t speciesb yRehde r(1910) .Materia lwa sdistribute dfro mth eArnol dArboretum .Especiall y plants with the combination of the pale yellow tepals and markedly dark violet stained filamentsmad eman yhorticulturist sbeliev etha tthi swa sth ever ydistinctio no fth especies , whereasi ti sonl ypar to fit svariation .It sdelimitatio nwit hC. intricatawa sa furthe r subject todiscussion ,bu tnowaday si ti swel lagree dtha tit srelativel ylat eflowerin gi nth esummer , itstypica lpal eyello wflowe rcolou ran dit sbiternat eleave swit hregularl yserrat eleaflet sar e thedistinguishin g characterso fth especies .Nevertheless ,th eplant swit hth edar kviole t filaments arever y showy and iti shope d thatthe ywil lgai n in popularity. Flowervariant swit hviole tspots ,jus ta flus h ofviole to reve ndar kviole tcolour s 219 havealway sconfuse d taxonomistsa st o Clematis sect.Meclatis, because horticulturists wereofte n impelledt ogiv ethes evariant sseparat enames .Rehde r(1920 )describe dsuc h plants under the name C. glauca var. akebioides f. phaeantha Rehd., which is synonymous to C. intricata var. purpurea described by Y.Z. Zhao (1979). The occurrenceo fviole tcolouratio n oftepal si svariabl ean dtherefor e inm yopinio ni ti sno t worthwhilemaintainin gnames ,base do nthi sphenomenon ,a tan yrank .I agre ewit hGrey - Wilson(1989 )tha ti fan yo fthes eplant sar estil li ncultivation ,worthwhil egenotype s should be given cultivar names. Under the present International Code of Nomenclature for CultivatedPlant s(1995 ) 'Phaeantha' asepithe ti s allowed,bu tm ypreferenc e isa fanc y namei nmoder n language to avoidan yconfusio n concerning theidentit y of thecultiva r indicated. Atth ebeginnin go fth e20t hcentury ,ther ewer estil lauthor s(Fine t& Gagnepain , 1903;Krasheninnikov , 1937)wh olik eKuntz e(1885 )reduce dal lspecie st oon especies : C. orientalis.Surveyin gth ehorticultura lliteratur e(Bailey ,1917,1976 ;Bean ,1970 ;Boom , 1980; Rehder, 1974; Fisk, 1975; Johnson, 1998; Kriissmann, 1976; Lamarck, 1786; Lavallee, 1884;)o nClematis sect .Meclatis, iti sremarkabl eho wvagu eth edescription s ofth eassigne dspecie sare ,an dho weveryon ejus trepeat serror si npreviou sliterature .Thi s is especially striking where it concerns the maintenance of a broad description of C. orientalis,an d at the same time maintains an pseudo-exact narrow difference with C. glauca(th e indumentum of thetepal si s aver y variablecharacter) . 4.4. Cultivars of Clematis sect.Meclatis Aselectio no fyellow-flowerin g Clematiscultivar si ssurveye dhere .Recentl yth eassortmen t hasbee nextended .A concer nabou tthi sassortmen tespeciall yan dt oClematis i ngenera li s thatan yne wintroductio nvarie slittl ebeyon dth ealread yexisting .Ther ei sreall ya nee dfo r cultivarswit himprove dornamenta lvalue ,mor eresistanc eagains tpest san ddisease san d better growth performance. 220 4.4.1. Clematis'Aureolin' . Winner: Experimental Station for Arboriculture, Boskoop. Introduction into cultivation: 1979. Description: Grootendorst (1979) in Dendroflora 15/16:62 . Selectedfro mvariou sinterspecifi ccrosse swithi nClematis sect .Meclatis wit ha srecurren t parent C. tibetanasubsp .tangutica. Woodyclimbe r with thehabitus ,foliag e andinflorescenc e similart oC. tibetana subsp. tangutica, profuse flowering in the summer, (June-)July-September. Flowers nodding, broadly campanulate with tepals 4, lanceolate, bright yellow (HCC3,aureolin) , +4 0x 20mm.Achene s with showy longplumos e styles. Remark:Materia lo fthi scultiva rwa sconsidere dmeritoriou sb yth eTria lCommitte eo fth e "Koninklijke Vereniging voor Boskoopse Culturen" in 1979. Conserved standard (designated here):Brandenbur g 201(WAG) . Illustration: Seefig . 4.1. Figure 4.1. Clematis 'Aureolin'. 221 4.4.2. Clematis'BillMcKenzie' . Winner: unknown. Introduction intocultivation : 1969b yMrs .Finni s (UK). Description:Plan tsimila rt oC. tibetanasubsp .tangutica ,bu tmor evigorousl ygrowin gan d large yellow,campanulate , noddingflowers, u pt o 2cmlong . Remark: Material of thisplan t received anR.H.S .Awar d of Merit in 1976. Conserved standard: Wilders 390(WAG) . Illustration: seefig . 4.2. Figure4.2 . Clematis'Bil lMcKenzie ' (photograph byA .va nde rNeut) . 222 4.4.3. Clematis'Bravo' . Winner: Agricultural University, dept. Plant Taxonomy (former dept. Taxonomy of Cultivated Plants and Weeds;Brandenburg/Va n deVooren) . Introduction intocultivation : 1981. Description: Woody climber with the habit, foliage and inflorescence similar to C.tibetana subsp . vernayi,u p to3. 5 mtal l (therelativ e compactness of theplant s mayb e duet o thegrea t masso flarg eflower sproduced ;a tth etim eo fflowerin gvegetativ egrowt hceases) ,profus e floweringi nth esummer ,(June-)July-September .Flower snodding ,broadl ycampanulat e withtepal s4 ,broadl ylanceolate ,brigh tyello wgoin gove ri ngolde nyellow ,+ 4 5x 2 5mm . Achenes with showy longplumos e styles upt o 7c mlong . Conservedstandar d(designate dhere) :Va nderNeu t35,36,3 7(WAG) ,includin galcoho l collection (Van derNeu t 35). Illustration: seefrontispiece , fig.4.3 and fig.4.4. Figure 4.3. Clematis'Bravo ' (Photograph byR.A.H .Legro) . 223 Figure4.4 . Clematis'Bravo ' (drawingb yMarie t deGeus) . 224 4.4.4. Clematis 'Burford'. Winner: Treasures of Tenbury, England. Introduction intocultivation : 1975 Description: Plantsimila rt oC. tibetanasubsp .tangutica, excep tfo rit salmos tglobula rshape dflowers . The tepals arebrigh t yellow andrelativel y fleshy. Remark:Plant so fthi scultiva rwer eoriginall yintroduce dint ocultivatio nunde rth enam e 'Burford Variety'; such names are illegitimate under the International Code of Nomenclature, 1980,Art .31c : 'Ono rafte r 1Januar y 1959,ne wcultiva rname si nth efollowin gfor mar einvalidl y published: (c) Names including the word variety (or var.) or the word form. However, whenvar .denote svariegated ,th enam ei sno trejecte d butth ewor di s written in full.' Invalidlypublishe dname sar ea tth esam etim eillegitimate .Fo rreason so fstability ,th enam e ishereb yreplace d by alegitimat e equivalent. Conserved standard (designated here): Brandenburg 202 (WAG), Van de Laar 5335 (Experimental Station for Arboriculture, Boskoop). 225 4.4.5. Clematis'Cony' . Winner: Zwijnenburg, Boskoop. Introduction intocultivation : 1975. Description: Selectedfro m thecros sC. tibetanasubsp .tangutica x C.'Orang ePeel' .Wood yclimbe r upt o3.5 mtall .Fre eflowering i nth esumme r(June-)July-September .Leave ssimila rt oC. tibetanasubsp .vernayi. Flower sbroadl ycampanulat egolde nyellow ,+ nodding .Tepal s 4,rathe r fleshy. Remark:Materia lo fthi scultiva rwa sconsidere dmeritoriou sb yth eTria lCommitte eo fth e "Koninklijke Vereniging voor Boskoopse Culturen" in 1975. Conserved standard (designated here): Van de Laar 4994 (Experimental Station for Arboriculture, Boskoop). Illustration: seefig .4.5 . Figure 4.5. Clematis'Corry ' (photograph byA .va n derNeut) . 226 4.4.6. Clematis'Drake' s Form'. Winner: unknown. Introduction intocultivation : unknown. Description: R.H.S.Dictionar y of Gardening (1993). Plantsimila rt oC. tibetanasubsp .tangutica, bu tmor evigorousl ygrowing .Flower sbrigh t yellow,large ;tepal s+ 4 0x 20mm ,narrowl ycampanulate .Achene swit hshow yplumos e styles.Furthe r details thus far unknown. 4.4.7. Clematis'Golde n Harvest' Under this name, a selection is cultivated in the Netherlands, rather similar to C. serratifolia. Theplan t has paleyello w flowers with darkviole t stamens. 4.4.8. Clematis'Helios' . Winner: Experimental Station for Arboriculture, Boskoop. Introduction intocultivation : 1988. Description: Van deLaa r (1988) inDendroflor a 25:72 . Woodyclimbe ru pt o1.75 mtall ,cease sgrowin gwhe ni tcome sint oflower .Flowerin gtim e May-October.Leave ssimila rt oC. tibetanasubsp .tangutica. Profus e flowering.Flower s mostlysolitary ,sometime s3-flowere daxillar ycymes ,nodding ,open ,flat ,tepal s4 ,brigh t yellow (RHS 14B),lanceolate , 35-45 x 15-20mm, 2-2.5 x aslon g aswide ,acut e atth e apex,whic hi sslightl yrecurve d whenful l flowering.Achene swit hlon gplumos eshow y styles. Remark:Th eTria lCommitte eo fthe "Koninklijk eVerenigin gvoo rBoskoops eCulturen " issued apositiv ejudgemen t onmateria l of this cultivar in 1988. Conserved standard (designated here): Preferably a specimen from the Van de Laar collection, stillt ob e designated. 227 4.4.9. Clematis'Lambton' s Park'. Winner: unknown. Introduced into cultivation: unknown. Description: R.H.S.Dictionar y of Gardening(1993) . Plant similar to C.tibetana subsp . tangutica. Flowers nodding, large, up to 2cm long, bright yellow.Furthe r details thus far unknown. 4.4.10 Clematis'Orang e Peel'. Ludlow,Sherrif f andEllio thav emad eextensiv ecollectio ntrip sthroughou tTibe tan dNepal . 19Octobe r 1947the ycam e across afruitin g specimen of C. tibetanasubsp . vernayi at Kongbo near Kyimdong-Dzong: Chin-Tung (Tibet, 29'N 93'25"E, alt. 3600m). They recorded: 'Branch of seed taken'. The seeds of this specimen were introduced into cultivation in England under the name C. 'Orange Peel'. It is therefore a generatively reproducedcultivar ,wit ha sdistinguishin gcharacter sth ebroadl ycampanulat eflower swit h thickflesh ytepals ,tha ttur nfro mgolde nyello wt odee porang edurin gflowering .Foliag ei s veryvariabl ei nshap ean dsize .Thi sha scause dsom econfusio n whethero rno ton eha sth e "correct" 'OrangePeel 'o rnot .Fro mth especimen savailabl efo rthi sstudy ,I hav etherefor e designated a composite conserved standard to warrant the identity of this "collective" cultivar. Conserved standard (designated here): Caldwell s.n., 7-10-1967, Exeter (BM);Va n de Laar 2451, 2452 (Experimental Station for Arboriculture, Boskoop);Brandenbur g 282. Illustration: See fig.4.6 . 228 Figure4.6 . Clematis'Orang e Peel' (photographs R.A.H.Legro) . 229 4.4.11 Clematis'Wallsal' . Winner: Guernsey Clematis Nursery Ltd. Introduced into cultivation: 1986. Description:Plan t similart oC. tibetana subsp.tangutica, flowers brigh tyellow .Leave s trifoliolate topinnate .Achene s with longplumos e styles,u pt o 7cmlong . Conserved standard (designated here):Wilder s 391(WAG) . 4.5. Conclusions SurveyingClematis sect .Meclatis i ncultivation ,i ti sremarkabl etha tmos ti fno tal lcultivar s aresimila rt oa subspecie so fC. tibetana.Th eintroductio no fthes eplant sa tth een do fth e 19th andth e middle of the 20th centuryha s received the attention bybreeders ,bu t so far they have only made combinations within the section. True C. orientalis and C. serratifolia, haveno thithert obee n used verymuc h inthes ehybridizatio n programmes, whereas especially C. serratifolia has interesting characters such as propagation by producing lots of rooted stolons. The assortment of Clematissect . Meclatiscultivar s is too small to necessitate classification intocultiva rgroups .A ssoo na sC . orientalis,C. intricataan dC. serratifolia play a more prominent role in Clematis breeding, it is worthwhile considering to circumscribe acultiva rgrou paroun d cultivars similart oC. tibetana andon earoun d C. orientalis,C. intricata an d C serratifoliabecaus e of their different reaction to growth conditions, pruning requirements and propagation, and their predicted difference in ornamental value. 230 GENERAL DISCUSSION Thestructur eo fthi sinvestigatio npermit sth ediscussio no fthre ebiologica ltopic srelatin gth e fieldso f plant systematic research, plantbreedin g research andevolutionar y biological research.I ti sworthwhil econsiderin gthes erelationship si ntime si nwhic hplan tsystematic s asa disciplin e hasbee n questioned as aseparat e discipline tob emaintained . Iti sa tth e sametim eimportan tt oadvocat eth esignificanc eo fplan tsystematic si na realisti cscenari o and not in anuncritica l approach to use systematic conclusions to predict relationships betweentaxa ,thei rimportanc efo rplan tbreedin gor ,mutually ,t opredic tfro mcrossabilit y data in an absolute way the relations between taxa (Barber, 1970). Both have been frequently done. It will be shown in the next paragraphs that systematics are indispensable for modernbiologica lresearch .A mode lwil lb epresente dho wplan tsystemati cresearc hca n beinvolve d in allkind s of botanical research facing diverse types of scientific questions. 5.1. Clematisspecie s with regard tovariou sspecie s concepts ManyRanunculea ngener aar echaracterize db yshowin gmorphologicall yalmos tcontinuou s variationpattern sb ywhic hi ti sdifficul t todefin especies .Thi sphenomeno ni scommo nb y plantfamilie swhic har eabundan ti ndynami co rdisturbe dhabitats ,suc ha sAsteracea ean d Poaceae.Ranunculacea ear ea relativel yol dfamil yo f Dicotyledonousplants .Althoug hth e Ranunculaceae areles sabundan ti nabov ementione d habitats- an dles s successful than Asteraceaean dPoacea ei fw ecoun tnumber so fspecie s- the ysho wuniqu ecombination s of old and derived characters, as can be demonstrated by traits of flower morphology throughout the family: hemicyclic flowers and at the same time various forms of zygomorphy. A confusing situation in Clematissystematic s has always been the mixture of species conceptsa suse di nmonograph san dconsequentl yi ngarde ntextbooks .Whateve rconcep t 231 used, it isimportan t tob e used consistently throughout amonograph . Kuntze's system (1885)ha sno tbee nfollowed , insteadi twa scriticise dver ymuc hb yhi scontemporaries , buth ewa sabou tth efirs t torais eth eissu eo fusin ga specie sconcep tan dthe n following thisconsistently .Whe nspecie so fa genu sbecom epopula rgarde nplant san dman ycultivar s havebee nraised ,i ti sextremel yimportan tt omak ea stric tdistinctio nbetwee nth egenus ' botanicalclassificatio n andth edesir eb yma nt oclassif y thegarde nplant sb ythei rusage , ornamentalvalu eo rhorticultura lcharacteristics .A tth emoment ,a monographi ctreatmen to f a genus mixes upboth , soth ebotanica l species aredistinguishe d bycharacter s that are relevant to horticulture and gardening, and that have little to do with modern plant systematics(Render ,1920,1974) .A tth esam etime ,a cultiva rclassificatio n willsuffe r from theopposite ,i fi ti spartl ybase do ncharacter srelevan tt odescribin gnatura lpopulation san d tophytogenie s(Baum ,1981;DeWet ,1981;Pickersgill , 1986).Insystematic si tmean stha t elementsar euse dfro m allspecie sconcept san dcombination sbetwee nthem ,althoug hth e application of theLinnaea n andnatura l species concepts are most abundant. The Linnaean species concept isbase d on his sexual classification system.Th e generaar eclassifie d bynumber so fpart san dothe rcharacter so fflower s (figure5.1) ,an d for species Linnaeus used vegetative characters: in case of flowering plants his first characters tob euse d arethos eo f leaf morphology. Hetende d tous e asfe w aspossibl e characters whichinevitabl ylead st oa nartificia l classification. Hestate dtha tspecie sar e createdb yGod ,bu th eals oaccepte dmanmad evarietie soriginate dwithi nspecies .Wa shi s statement onth ecreatio no fspecie sinitiall yver yfirm , laterh eaccepte d somespecie sa s being derived.Especiall y in Clematis, there areindication s for this.Linnaeu s basically adoptedhi sspecie sconcep tfro mRa yan dafte rhi mther ewer eonl ya fe wcolleague swh o adoptedsimila rspecie sconcepts .Withou tLinnaeu sthi sartificia lspecie sconcep twoul dno t havebee nver yinfluential :i ti sjus ttha tSpecie sPlantaru mi ssuc ha landmar ki nsystemati c history,compilin gove r40 %o fth eworl deconomicall yimportan t(flowering )plan tspecies . Throughagricultura lbotany ,hi sspecie sconcep tcontinue dt ob einfluentia lu pt ono wa sh e simply included agronomic characters to distinguish e.g. Brassicarapa fro mBrassica campestris, Ribesgrossularia from Ribesuva-crispa, Triticum aestivum fromTriticum hibernum etc. 232 Cbrirf; LINNiGI.M.D. "METHODUS ptantarum JEXUALIS inSLTTEMATE NATURAE deftripte Polyandri GD.EHRET. ftUfcheidetb: Lugd.bat:/T3(T fecit W ediAt Fig.5.1 .Linnaeus 'classificatio n system.Polyandri anea rth elette rN indicatin gth eclas st o which Clematis belongs (Polyandria Polygynia, shown herei sPolyandri a Monogynia). Adanson (1763)introduce da natura lspecie sconcept ,whic hwa sabov eal lmad e popularb ylate rsystematists .Th eprincipa ldifferenc e withth eLinnaea nspecie sconcep t 233 wastha ti ttoo kint oaccoun tno tonl ya fe wbu tal lobserve dmorphologica lcharacter san d consequentlywa sinterpretin gvariatio ni nterm so foveral l(dis)similarity .I nmor eelaborat e treatises of economic plants it was especially this species concept that was implicitly applied.Th erecentl ypublishe dmonograp ho fClematis b yMagnu sJohnso n(1998 ) suffers from such a combined treatment, and has therefore restricted value despite the overwhelming amount of collated data. Dobzhanski (1935) introduced the biological species concept thus integrating genetic and systematic knowledge. Basically, the biological species concept states the speciesa sa nentit ywithi nwhic hindividual sca nmat ean dtherefor egenerativel yreproduce . Consequently alo t of experimental hybridization work was carried out to characterize speciesrelationships .Muc hinterspecifi chybridizatio nwor kha dalread ybee nbuil ti nplan t breeding schemesi norde rt obroade n the(narrow )bas eo fman ycro pplan tspecies .Th e biologicalspecie sconcep tbecam ever ypopula ramon gplan tbreeder sa sthe ysa wi ta sa scientific base behind their hybridization work. With respect to Clematis - and other ornamentals- interspecifi chybridizatio nha dalread ybegu ni nth eearl y19 thcentur y(Moor e & Jackman, 1872) and it continues until now. These hybridizations produced food for thoughto nclassificatio no fespeciall ycultivate dplants ,an dwer eth efactua lbeginnin go fth e definition of thecultiva rgrou pan dth ebasi c concept behind it:culton . Remarkably the speciesconcep t remained untouched for long,thu s leavingbotanica l systematics witha completerang eo finterpretation so fwha ta specie sshoul dbe ,despit eal linstabilit ytha ti t causest onomenclatur e (Brandenburg, 1991;Brandenbur g &Schneider , 1983,1985) . 5.2. Crossability data andthei r value for plant breeding research Whereasplan tsystematic saim sa tordenin gth ePlan tKingdom ,classifyin gorganism sb yal l characters at our disposal, plant breeding by making use of plant systematic evidence causes entropy in the systematic framework: as soon as species relationships has been revealedb yplan tsystemati cresearch ,plan tbreedin gdisturbe sth eintegrit yo fth especifi c entitiesconcerned .Understandin g therelationshi pbetwee n both disciplines willhel pi n 234 determining thevalu e of systematic datafo r plant breeding. Beforeth eer ao fmolecula rbreeding ,th elimit so fplan tbreedin gwer edetermine d bypossibilitie st omak ecrosse sbetwee nindividua lplant seithe rbelongin gt oon especie s (intraspecifichybridization )o rt odifferen t species(interspecifi chybridization) .A tth een do f the 19th century and the first three decades of the 20th century, many European and Americanhorticulturist swer econfronte db ya lo to fne wintroduction so fornamenta lplants . Inthei rextensiv ecollection sthe yoccasionall yfoun dchanc ehybri dseedlings ,an dlate rthe y deliberatelymad ecrossing sthemselve seithe rt oconfir mth eputativ eparentag eo fchanc e seedlingso rjus tt ocreat ene wcultivars .Th esprea do fgeneti cknowledg ei nth eearl y20 th century induced many interspecific hybridization experiments in genera such as Anthirrhinum, Galeopsis and Nicotiana to study biosystematic, genetic and even ecologicalquestions .I nal lthes estudies ,i twa sassume dtha tinterspecifi c crossabilityi s directlycorrelate dt oth edegre eo frelationshi pbetwee n species.Consequently ,th eter m specieswa sbiase db yth evie wtha tal lindividual stha tcoul db emutuall ycrosse d should belongt oth esam e(biological )species .I nth esecon dhal fo fth e20t hcentury ,afte r Camp and Gilly (1943) and Camp (1951) had introduced experimental research in plant systematicsan dcalle dthi spar to fth edisciplin ebiosystemat yo rbiosystematics ,mor ean d moreevidenc ewa sfoun dtha tth ebiologica lspecie sconcep twa sa noversimplificatio n of reality and therefore not applicable. With the Aquilegia study, Grant (1952, 1971) delivered oneo f theclassica l examples in this respect. Hedemonstrate d that sympatric Aquilegiaspecie swer eperfectl yisolate di nnatur ea tcertai nlocalitie s(n ohybrid sfound) , whereasa tth esam etim ethes especie sbrough ttogethe ri non eexperimenta lgarde ngav e unlimitedlyris et ointerspecifi c hybridsb ymanua lcrossing .Althoug h these species are genetically closely related, they are also isolated by their specialization to different pollinators(hawkmoths ,bumblebee san deve nbirds) .B yrigorousl yapplyin gDobzhansky 's biological species concept, however, the concerned Aquilegia species had to be consideredt ob eon especies ,a swoul db evali dfo rth especie so fClematis sect .Meclatis too (this thesis).B y considering their coevolution with pollinators and the consequent adaptations- trend si nvariatio n- th erecognitio no fsevera lspecie si nAquilegia b yGran t (1952,1971)an di nMeclatis (thi sthesis )ha sbee njustified . Anderson(1949 )surveye da 235 loto fothe rstudie spointin gi nthi sdirection .B ytakin gint oaccoun tadaptiv etrend si nplan t variationth eevolutionar yspecie sconcep twa sdefined .Gran t(1971 )distinguishe dsevera l speciation mechanisms: 1. Primary speciation a. Geographical speciation (Giliaspecies ) b. Quantum speciation (especially in birds) c. Quantum speciation with chromosomal repatterning (Clarkia species) d. Sympatric speciation (postulated ontheoretica l grounds) 2. Hybrid speciation with sexual reproduction a. Hybridspeciatio nwit hexterna lbarrier sa spollinator s(Carex, Delphinium and Ophrysspecies ) b. Recombinational speciation (e.g. Crepisan dNicotiana species ) c. Amphiploidy (e.g.Brassica an dNicotiana species ) 3. Hybrid speciation with asexual or subsexual reproduction a. Apomictic speciation (e.g.Festuca an dPoa species) b. Permanent ornumerica l hybridity (e.g.Oenothera biennisan dth eRosa caninacomple x by special meiotic mechanisms) 4. Wallace effect (Lycopersicon, Solanumspecie s by incongruity) Figure5. 2Hybridizatio n polygon (generalized figure offigur e 1.21). 236 Thenatur eo fspeciatio nmechanism sha sbee nreveale db yth ecombinatio no fth eanalysi s ofnatura ldistributio npatterns ,cytologica linvestigation san dexperimenta lhybridizatio n polygons such as the generalized one from fig. 5.2. In these schemes the results of interspecifichybridizatio nexperiment sar eschematicall ydepicte dcharacterizin gth ecrossin g behaviourbetwee nspecie sunde rartificia lcondition s(cf .Cai n& Harrison ,1958 ;Carson , 1985;Coyne , 1974; Davis, 1978;Davi s& Heywood , 1965;Dobzhansky , 1935;Ehrlich , 1961;Funk , 1985;Hutchinson , 1923; Huxley, 1940, 1958). The evolutionary species concept has the advantage overth e biological species concept that it takes into account the aspects of reproduction biology, distribution mechanisms and adaptive trends,bu t it doesno t deal with trends such asdivergen t and convergent developments, the systematic development of characters and geographical regularities inditributio npatterns .B ytakin gthi sint oaccount , thephylogeneti c species conceptwa sintroduced ,base do ncladisti canalysi smethod srathe rtha npheneti cmethods . Theanalysi so fchapte r2 clearl yreveal sth epowe ro fth ecladisti capproac ha soppose dt o purely phenetic approaches.Especiall y where similarity israthe r large as isth e casei n Clematissect . Meclatis,bu t also in other Clematis sections, it has been clearly shown againtha tpheneti capproache sgiv einsigh ti nth eoveral lstructur eo fth evariatio nwithou t revealingth edirectio no fth edevelopmen ti nvariatio ntrends ,wherea sth ecladisti canalysi s provides a higher resolution by taking into account the direction of variation trends. Moreover,b ystudyin gthes etrend splan tsystematic sha sbecom efo r thefirst tim ea rea l science with hypotheses that can be either proven or rejected. One problem with the phylogeneticspecie sconcep ti sit sfocu so nmonophyleti cgenealogies ,thu sno tdealin gwit h evidence from interspecific hybridization that has also been described from natural complexes.Korne t (1993) triedt ocombin ebot hint oon etheore mi norde r tob eabl et o analyzereticulat ecomplexes ,i nwhic hinterspecifi c hybridizationwa son eo fth edrivin g forces. Iassum etha ti twil lb eonl ypossibl et orealiz ehe ranalysi sb yaddin g moredat a sets,containin gmolecual rdata .Molecula rbiolog ymake si tpossibl et odelive ra nobjectiv e set of data in which the relatedness of genomes - and therefore of populations or even species- ca nb echaracterized . Combiningthi swit hmorphologica ltrends ,tha tstan d for certain adaptations to the habitat, and hybridization data, that stand for the nature of 237 isolationbetwee npopulation so reve nspecie sb ysexua lmeans ,w ema ydevelo pa specie s concepttha twil lb euniversall yapplicabl ean dtha ti sreflectin gbot hit scurren tbiologica l meaningan dit sphylogeneti cderivation .Th eapplicatio no fsuc ha specie sconcep tha sals o theadvantag e thathypothese s canb eclearl yformulated , tested andconsequentl y either acceptedo rrejecte d ina biologicall ymeaningfu l way(cf . Estabrook, 1972;Farris,1971 , 1974, 1980, 1985; Fink, 1982;Ghiselin , 1981,1984,1987 , 1988;Gilmour , 1961;Hull , 1976;Humphrie s& Funk , 1984;Meglitsch , 1954;Rieppel , 1991)despit eo fth eadvers e opinion of some authors (e.g.Cronquist , 1987). Application of aphylogeneti c species concept in themoder n sense (cf. Kornet, 1993)provide sa magnificen t startingpoin tt oestablis ha fir mbas efo rinfraspecifi c botanical -thu sclose d- classification .Man ything shav ebee nsai dabou tinfraspecifi c taxaan dthei r application (Baum, 1981; Burtt, 1970; DuRietz, 1930; Fuchs, 1957; Hamilton & Reichards, 1992; Meikle, 1958;Pickersgill , 1986; Stace, 1986, 1989; De Wet, 1981; Wijnands, 1986a,1986b )rangin gfro mrathe rartificia l systemsa spragmati ccombination s ofcurren tuse so finfraspecifi c categoriest omor efundamenta l considerations,bu tnon eo f themtakin gint oaccoun tth especie sconcep tt ostar tfrom . Notdoin gso ,implie stha ti ti s notpossibl et odefin e infraspecific categories,thu sprovidin ga nobjectiv e wayt oappl y them. 5.3. Evolution and domestication ofClematis 5.3.1. Evolutionaryaspects of Clemati s withinthe Ranunculaceae As aRanunculea n genus, Clematisha sman ycomple x features, andye ti ti s considered primitive within the family. Its flower morphology is basic, being hemicyclic, actinomorphous. However, the leaves are decussate and the stems woody. The woody stemshav eth e anatomy of perennial plants (Smith, 1928),an dth ephyllotaxi s is not a criticaldiscriminatin gcharacte r(Tepfer , 1960)a sw ese esevera lgradua ltransition sthroug h analternat elea fpositio n (e.g.i nClematis alternata) within Clematisan dth eoccurrenc e 238 of alternate phyllotaxis in other Ranunculean genera as well. Looking at cytological characters,th ebasi cnumbe ro fchromosome si nClematis i sx = 8 ,th enumbe rtha toccur s mostfrequentl y inRanunculacea ethoughou tth egener atha tar econsidere dmos tprimitiv e (Gregory, 1940).I ngeneral ,I agre etha tClematis ha st ob eregarde da sprimitiv eamon g Ranunculean genera, sharing manycharacter s with another primitive genus,Anemone. From thedat apresente d in this thesis,i t cannot concluded with certainty where Clematis originated onearth .Fo rtw ohypothese s there is some support: - Fora Laurasea norigi nth esuppor ti st ob efoun d inpalaeobotanica levidenc ean d the fact that almost every variation trend present in Clematisi s represented in especiallyth eChines eare a(Croizat , 1962,1964;Croiza te tal. , 1974;Krassilov , 1983;Nelso n &Platnick , 1981); - For a Gondwanean origin the presence of the variation trend from Anemone throughClematopsis t oClematis i sver ysupportiv e(Brummitt ,1976 ;Hutchinson , 1920;Rayna l 1978;Tob e 1980c). Inm yopinion ,i ti sonl ypossibl et omak ea bette rdiscriminatio nbetwee nbot hhypothese s by a molecular biological approach, especially by looking at conserved regions in chloroplastan dmitochondria lDN A(cf .Hoo te tal. ,1994) .T om yknowledge ,suc ha stud y islackin gfo r Clematisan drelate dgenera .Molecula rgenetic smak ei tpossibl et otes tth e abovekin do fhypotheses ,t otes tth ecladisti canalysi so fth egenu sa sbee ncarrie dou tfo r this thesis and to shed a better light on the subdivision between Monocotyledons and Dicotyledons,whic hi sessentia lfo ra bette runderstandin go fsimilaritie san ddissimilaritie s betweenbot hgroups ,thu scontributin gtoward sa bette ran dsustainabl eexploitatio no fthei r variation, and to test thephylogenetica l assumptions byTamur a and coworkers (1958, 1962, 1963,1964,1965,1967,1968a, 1968b, 1970, 1987;Tamur a &Vogel , 1993). 5.3.2. Domestication o/Clematis Domestication of Clematisdate s back in Europe towards the late Middle Ages. The Europeanwil dspecies ,especiall yClematis recta, wer eofte nplante di nmedicina lgardens . Although there are no exact data to find there are some indications that in China domestication of Clematis dates back at least to the 8thcentur y BC. (Usher, 1974).I ti s 239 therefore likelytha tth efirs t Clematisspecie stha twer eintroduce dfro mChin aan dJapa n werealread ycultivate dplant si fno tcultivars .Rober tFortun epointe dou ttha th efoun da n interesting Clematisi na garde nalon ghi srout et oNanking .Late ri twa sname dClematis lanuginosa (Brettschneider, 1898). Remarkably, this species was never found in the Chinesewil dflor a(Ling ,1980) ,thu sgivin gevidenc etha ti twa sindee da cultivar : Clematis 'Lanuginosa'. The assumed period of domestication that maydat e back to the western earlyMiddl eAge s(!) ,suggest stha tth econcerne d speciescomple xma yb eregarde da sa compilospecies(Harla n& D eWet , 1963).Th elon gperio do fdomesticatio nfo rcultivate d plantsi nChin asuggest ssimila rmigratio npatterns ,e.g .th esil kroute ,a sdescribe dfo rothe r cultivated plants bySaue r (1957, 1967a, 1967b, 1988). Domestication of Clematisha s always been focused on flower colour, size and other morphologicalcharacter ssuc ha sdoubl eflowers ,anther sbein gvariousl ytransfere dint oa kindo fpetaloi dorgan so rt ojus tanothe rwhor lo ftepals .Thi sfocu s wass ostron gtha ti t hasle dt ophysiologica lproblem si nlarge-flowere d Clematiscultivars .Cultivars ,suc ha s 'Prins Hendrik', once popular as a cutflower in the Netherlands, could only be grown graftedo nClematis vitalba. Man yo ftodays 'large-flowere dcultivar sseriousl ysuffe r from theClematis wil tdisease .Thi si sno ta rea ldisease ,bu ta physiologica lproblem :b ylac ko f waterth estem sge taerate dan da sa consequenc eo fth ecombinatio no fai ri na naqueou s environment (the stem), fungi, such asAscochyta clematidina or the common Botrytis cinereafor m colonies in xylem vessels thus blocking the water supply of the plant. Clematiswil toccur smos tprominentl ywhe nth eplant sar eabou tt oburs tint oflower .Th e flowerbud sar ebi gan drequir ea lo to fwate rt oexpand .Whe nplant sthe nar egrowin gi na sandy soilunde r sunnycondition s andwaterin gi sno t adequate,the ywil ljus t collapse. Clematis has arathe r superficial rooting system. Consequently, the one-sided focus on profuse flowering withlarg eflower s withoutselectio ntoward sa stronge rrootin gsyste m causesthes eproblems .Althoug hth esmall-flowere d Clematiscultivar sd ono tgenerall y suffer fromth eClematis wil tdiseas e- a sthei rflowerin g isi nbalanc ewit hth ecapacit yo f theroo tsyste m- ,thei rabilit yt ocompet eunde rgarde ncondition si susuall yrathe rwea k and should be improved by breeding and selection. The samehold s true for resistance against pests anddiseases . 240 5.4. Cultivar documentation Therear emor ecultivar s offlowering plant si nth eworl dtha nwil d species.Takin g into account allth eobsolet ecultivar sth edifferenc e willb eeve nmore .Despit ethi sfact , the systemo fcultiva rdocumentatio ni sno tye tver yadvanced .Differen t topicso fconcer nrang e from aspectso fcultonomy ,identificatio n andcharacterizatio ntoward scultivatio nan dusag e characteristics. 5.4.1. Cultonomy, identification andcharacterization If we recognize that cultivated plants result from domestication rather than from spontaneous evolutionaryprocesses ,i t isimportan tt oaccep ttha t for classification and nomenclaturalpurpose s(Chapte r3) .B ydoin gso ,i ti spossibl et oestablis ha open ,flexibl e classificationo fcultivars ,adaptabl etoward sne wbreedin gdevelopment swithou tstickin gt o aneverlastin gnomenclatur efo rth eentitie sbetwee ncultiva ran dbotanica ltaxa .Th eculto n conceptclearl yleave sspac efo rsuc hflexibilit ythu screatin gcultiva rgrou pclassification s thatca n easilyb eupdate d oraltered . Theculto n concept also leaves roomfo r re-useo f cultivarepithet sunde rcertai nconditions .Merel yth efac ttha tmor ecultivar sar eobsolet e block theprovision s for goodnames .Re-us e mechanisms havet ob e developed for the International Code of Nomenclature for Cultivated Plants. By not putting re-use mechanismsi nplace ,th edevelopmen twil linevitabl yg otoward sth ereplacemen to fcultiva r epithetsb ytrad ename so reve ntrad emarks ,a developmen ttha ti salread yrathe rpopula r in ornamentals. Another aspecti stha to fIntellectua lPropert yRights .I norde rt o stimulateplan t breeding,ther eshoul db ea reasonabl emechanis mt oge tth ebreede rpai dfo rhi s efforts. Sucha mechanis mha sbee nlai ddow ni nth ePlan tBreeders 'Right slegislatio nunde rth e internationalUPO VConventio na srevise di n1993 .Th eproble mwit hthi slegislatio ni stha t iti sonl yimplemente di na restricte dnumbe ro fcountrie swhil ei nsom eothe rcountrie sPB R legislation isonl yuse dd efacto . Inorde rt oensur econtro love rthei rproducts ,breedin g 241 companies, especially as for horticultural crop plants, have often chosen to label their cultivarswit htrad emark san donl yhav ea cultiva repithe tfo rregistratio npurposes .I nthi s way,th e significance of the cultivar name isreduce d in communication. People dono t realize,however ,tha tth eonl yon et oon erelationshi pbetwee ncultiva ran dnam ei stha to f thecultiva rnam eincludin gth ecultiva repithet .A trad emar kma ychang ebetwee ncultivars , thusgivin gth econsume rn owarran twhatsoeve rabou tth etru eidentit yo fplan tmateria ljus t bought.I ti son eo fth ebigges tchallenge si nsystematic so fcultivate dplant st oaddres sthi s pointproperl yan dt osolv eit .Th efac ttha tInternationa lRegistratio nAuthoritie sjus tden y thisfac ti sa mos tworryin gattitude ,tha tca nplac ecultonom y- an dwit hi ttaxonom y- i na backyard position. Strengthening the relationships between statutory and nonstatutory registrationauthoritie si sa priority .Th eattitud eb ynonstatutor yregistratio nauthorities ,a s frequently demonstrated bye.g . theRoya lHorticultura l Society, ason eo f the principal nonstatutoryregistratio nauthorities ,jus tt oprescrib estatutor yregistratio nauthoritie sho wt o acti nmatte ro fdisagreement ,ignorin gth efac ttha tstatutor yregistratio nauthoritie shav et o fulfill legislativerequirements ,i sfa rfro mconstructive .I nthi ser ao finformatio ntechnolog yi t mustb epossibl et ocreat emechanism st oovercom ethi skin do fproblem san dt owor ko n practicalsolution si nth espiri to fwha tha sbee nachieve db yPlantScop e(Aalsmeer )i nth e Netherlands. 5.4.2. Clematis cultivars andtheir validation ofcultivation andusage characteristics Validation of cultivation andusag e (vcu)characteristic s hasbee nlargel ycarrie d outb y nonstatutoryorganisations ,suc ha sth eKoninklijk eVerenigin gvoo rBoskoops eCultur ei n theNetherland so rth eRoya lHorticultura lSociet yi nth eUK .Althoug hi ti so fgrea tvalu e thatther ear esuc hvc utrial sanyhow ,th emethodolog ybein gapplie di nthes enonstatutor y trials- o rsometime sonl yo nsit ejudgement s- i smerel ywatchin gth eornamenta lvalu ean d theperformanc e ofth ecultivar s aton e sitewithou t replications andofte n a singletime . Consequently,on eca nhardl ybas eseriou srecommendation sfo rpractica lhorticultur ean d gardeningo nthem .Ther ear ehoweve rgoo dreason st ose tu pa goo dschem eo fvc utrial s for themai n arboricultural cropplants : 242 • to improve theinformatio n about themos tpopula r assortments; • torestric t the assortments toth ebes t performing cultivars; • toformulat e goodbreedin gobjective s andt oestablis hgoo dbreedin gresearc ht o meetthes e objectives. The country thatfirs t organizes such vcu trials isth eon etha t will secureit sexpor t position! 243 SUMMARY Thegenera lclassificatio n ofth egenu sClematis (Ranunculaceae )wa ssubjec to fstud yi n chapter 1.Base do nspecie scharacte rscores ,th einfrageneri c classification wasanalyze db y applyingHennig8 6a sphylogenetica lanalysi spackage .A sresul to fthi sanalysi sClematis wa s subdivided into 18sections ,on e of them subdivided in 3 subsections. Theworl ddistributio no fClematis wa sals ostudie dwit hHennig86 .I twa sno tpossibl e topostulat e the areao f origin of the genus Clematis with theavailabl e data set. Ainterspecifi c crosspolygo nwa smad ean danalyze db ysee dse tan dpolle ntub e growth.It ssystemati csignificanc e withregar dt o Clematisan di ngenera lwa sdiscussed . Dependento nth eadopte dspecie sconcept ,thes ecrosse sar ecrucia lo rjus tacademic .Th e adoptiono fth ephylogeneti cspecie sconcep tmad etha tthi schoic efo rClematis i sacademic . Nevertheless, iti s useful information for plantbreeders . Agenera ldescriptio no fth egenu sClematis wa spresente dwit hsom ebackgroun d informationo ncertai ncharacters ,suc ha soveral lhabitus , nectarleave san dth epositio no f nectaries. Chapter 2wa sdevote dt o Clematis sect.Meclatis. Thisparticula r sectionconsist so fth e yellow-flowering Clematisspp. ,tha tar egainin gpopularit yi ngardening .Man yeffort s were directedt orevea lth especie sdelimitation .I tappeare dtha tth epheneti cmethodolog yi so f restrictedvalu ei nsuc ha comple xo fquit esimila rspecies .Usin ga combinatio no fmethods ,th e phylogeneticanalysi sb yHennig8 6finall yreveale dth especie sdelimitation :Clematis orientalis, C.graveolens, C. intricata,C. ispahanica, andC. tibetana.C. tibetana was subdivided intothre esubspecies :subsp .tibetana, subsp.tangutica an dsubsp .vernayi. Well-know n 'horticulturalspecies 'suc ha sC. tanguticaan dC. vernayiwer ereduce di nran kan dother s sucha sC. glauca an dC. akebioides wer ereduce dt osynonymy .A summar yo fchromosome , pollen andisozym e data was presented. Chapter3 wa sfocusin go nmor efundamenta l aspectso fsystematic so fcultivate dplants .I tha s beenshow ntha tth ecultiva rgrou pi so fcrucia limportanc ei nclassifyin g cultivars,tha tth e 245 classificationprincipl efo rcultivate dplant si sope ninstea do fclose dan dconsequentl ytha tth e basalter mi nsystematic so fcultivate dplant sfo ra nentit ycanno tb etaxon ,bu tshoul db ea ne w term culton (plur.culta ;cultonom y for cultavs .taxonom y for taxa). Clematisi son eo fth efirst gener afo rwhic ha cultiva rgrou pclassificatio n waspresente di na systematicway ,a swa soutline di nchapte r4 .A shor tsurve ywa sgive no fth eintroductio nint o cultivation ofyellow-flowerin g Clematis spp.,an da majo r parto fth e yellow-flowering Clematiscultiva rassortmen tha sbee ndescribed .S ofa rn ocultiva rgroup sar eneede dfo r these cultivars. 246 SAMENVATTING Dealgemen eclassificati eva nhe tgenu sClematis (Ranunculaceae )wa sonderwer pva nstudi e inhoofdstu k 1.Me tgebruikmakin gva nkenmerkscore spe rsoor twer dd e infragenerische classificatiegeanalyseer dme tbehul pva nHennig8 6al sfylogenetisc h analysepakket .He t resultaatva ndez eanalys ewa sda tClematis wer dingedeel di n1 8secties ,waarva nee nwer d onderverdeeld in 3subsecties . Dewereldwijd everspreidin gva nClematis wer deveneen sbestudeer dme tbehul pva n Hennig86. Met de gebruikte set van gegevens was het niet mogelijk om het gebied te postuleren waar Clematis moetzij n ontstaan. Eeninterspecifiek ekruisingspolygoo nwer dgemaak te ngeanalyseer daa nd ehan dva n zaadzettinge npollenbuisdoorgroei .He tsystematisc hbelan ghierva nme tbetrekkin gto t Clematise ni nhe talgemee nwer dbesproken .Afhankelij k vanhe tsoortsconcep twaarme e wordtgewerk tzij ndi tsoor tkruisinge nwezenlij kda nwe lacademisch einformatie .Uitgaand e vanhe tfylogenetisc h soortsconceptzij nd ekruisinge nacademisch eaanvullend einformatie . Niettemin blijft hetbruikbar e informatie voor plantenveredelaars. Eenalgemen ebeschri jvin gva nhe tgenu sClematis wer dgecompleteer dme tenig e achtergrondinformatie overbepaald ekenmerken ,zoal sd ehabitus ,d ehoningbladere ne nd e positieva n nectarien. Hoofdstuk 2 is gewijd aan Clematis sect. Meclatis. Deze sectie bestaat uit geelbloeiendeClematis soorten ,di etoeneme ni npopularitei ti ntuinen .Vee linspannin gi s verrichto mto td esoortafbakenin g tekomen .He tblee kda tfenetisch emethode snie ttoereken d zijn inee ndergelij k soortscomplex waarin desoorte nee nhog emat eva n overeenkomst kennen.Me tgebruikmakin gva nee ncombinati eva nanalysemethodes ,leidd ed efylogenetisch e analyseme tHennig8 6to tee ngoed esoortsafbakening : Clematisorientalis, C. graveolens, C. intricata, C. ispahanica en C. tibetana. C. tibetana werd onderverdeeld in drie ondersoorten:subsp .tibetana, subsp .tangutica e nsubsp .vernayi. Bekend esoorte nui td e tuinbouwzoal sC. tanguticae nC. vernayi werde ni nran gverlaag de nander ezoal sC glauca enC. akebioides werden tot synoniemen gereduceerd. 247 Daarnaast werd een samenvatting van gegevens over chromosomen (x=8), pollenkorrels en isozymen. Hoofdstuk3 i sgerich to pmee rfundamentel easpecte nva nd esystematie kva ncultuurplanten . Erword taangetoon dda td ecultivargroe pva nwezenlij kbelan gi sbi jd eclassificati eva n cultivars,da the tbasisbegri pvoo ree neenhei di nd ecultuurplantensystematie knie ttaxo nka n zijn,maa ree nnieu wbegrip ,culto n(meervou dculta ;cultonomi evoo rcult avs .taxonomi evoo r taxa)moe tworde n ingevoerd. Clematisi see nva nd eeerst egener awaarvoo ro pee nsystematisch ewijz eee ncultivar-groe p classificatiewer dgemaakt .Di tword tduidelij kgemaak ti nhoofdstuk4 .D eintroducti eva nd e geelbloeiendeClematis soorte ni nd ecultuu rwer dbeknop tsamengeva te neveneen sva nee n grootgedeelt eva nhe tsortimen tgeelbloeiend eClematis cultivar si sbeschreven .Momentee l ishe tno gnie t noodzakelijk omvoo r dezecultivar s cultivar-groepen te creeren. 248 CURRICULUM VITAEWILLE M A. BRANDENBURG Born:Marc h 9,1953, Warffum, theNetherlands . Married, one son. Education: Primary School (Groningen) 1965 Gymnasium P 1971 Wageningen Agricultural University Study Plant Breeding (N13) 1978 Plant Breeding Taxonomy of Cultivated Plants Genetics Career: Wageningen Agricultural University 1978- 1987 Respectively atth e departments Taxonomy of cultivated plants and weeds,an d Plant Taxonomy RIVRO,hea d of methodology research 1987 -1990 CRZ,CPRO ,hea d of dept.Cultiva r Strategy 1990- 1994 CPRO, section leader Economic Botany 1994 -1999 Plant Research International, senior scientist 2000- Economic Botany Membership Committees: International Commission for theNomenclatur e of Cultivated Plants (IUBS) International Commision on Horticultural Nomenclature and Registration (ISHS) International Nomenclature Committee (ISTA) International Nomenclature Committee (IOPI) Commission on Genetic Modification (COGEM) Freetime: Music byMahle r and Bruckner, walking andcycling . Board member of alarg eDutc h funeral society; stimulating amongother s the organization of activities onmourning . 249 REFERENCES 1 Ackermans, G., 1983. Clematispatens C. Morr. et Decne. nader bekeken. LUvg . Taxonomie van cultuurplanten en-begeleiders ,Wageningen , 37pp + bijlagen. 2Adanson , 1763. Families desplantes .Ranunculaceae . Vincent,Paris . 3Alefeld , F.G.C., 1866.Landwirtschaftlich e flora. Wiegandt andHempel ,Berlin , 364pp. 4 Ali,S.I. , 1978.Th eflora o fPakistan :som egenera lan danalytica lremarks .Note sfro m the RoyalBotani c Garden Edinburgh 36:427-439 . 5 Anderberg, A., &A .Tehler , 1990.Consensu s trees, anecessit y in taxonomiepractice . Cladistics 6: 399-402. 6 Anderson, E., 1949.Introgressiv e hybridization. Wiley,Ne w York, 109pp. 7 Andre,E.F. , 1902.Clematis tangutica. RevueHorticol e528 . 8 Anonymous, 1969.Tentativ echecklis tan dinternationa lregiste ro fDahlia names . Royal Horticultural Society, London. 9 Ax, P., 1985.Ste m species andth e stem lineageconcept . Cladistics 1:279-287 . 10Backer , C.A., 1936.Verklaren d woorden boek der wetenschappelijke namenva n dei n Nederlande nNederlands-Indi ei nhe twil dgroeiend ee ni ntuine ne nparke ngekweekt evaren s enhooger e planten.Noordhoff , Groningen, 664pp. 11Baillon , H., 1867.Histoir e desplantes , vol.1, 52-62, 87.Paris . 12Bailey ,L.H. , 1917.Th estandar d Cyclopediao fHorticulture ,vol .II ,Clematis, 787-798. MacMillan, London. 13Bailey ,L.H. , 1976.Hortu s Third. Clematis, 281-285.MacMilla n Publ.,Ne w York. 14Barber , H.N., 1970.Hybridizatio n and theevolutio n of plants.Taxo n 19: 154-160. 15Barendrecht , C.J., 1972. Het Clematis sortiment op het Laboratorium voor Tuinbouwplantenteelt, Landbouwhogeschool, Wageningen. 16Barton ,L.V. , 1967.Bibliograph y of seeds.Columbi a University Press,Ne w York, 858pp. 17Barton,N.H. , 1989.Foundereffectspeciation.In:D.OtteandJ.A.Endler(eds.) Speciation 251 andit sConsequences . Sinauer Associates, Sunderland MA,229-256 . 18Batten ,D.J. , 1984.Palynology , climate and the development of Late Cretaceousfloral provincesi nth eNorther nHemisphere : areview .In :P .Benchle y(ed. ) Fossilsan dclimate , Geological Journal, Special IssueNo . 11, 127-164. 19Bauhin , C, 1623. Pinax theatri botanici. Basel,546pp . 20Baum ,B .R. ,197 1a .Th etaxonomi can dcytogeneti cimplication so fth eproble mo f naming amphidiploids of Triticum and Secale. Euphytica 20: 302-306. 21Baum.B.R. ,1971b .Nominagenericaconservandaproposita,proposal322 .Taxon20 :644 - 645. 22Baum,B.R. , 1981.Taxonom yo fth einfraspecifi cvariabilit yo fcultivate dplants . Kulturpflanze 29:209-239 . 23Bean,W.J. , 1970. Treesan dshrub shard yi nth eBritis hIsles ,vol . 1,A-C .Clematis, 633- 665. John Murray, London. 24Becker , G., 1957.Tournefort . Lesgrande s naturalistes francais, vol.2 .Paris . 25Bell,A.D. ,1974 .Rhizom eorganizatio ni nrelatio nt ovegetativ esprea di nMedeola virginiana. Journal of theArnol d Arboretum 55:458-468 . 26Bentham ,G. ,1862 .Ranunculaceae .In :G .Bentha m& J.D .Hooke r(eds. ) Generaplantaru m adexemplari a imprimis inherbarii s kewensibus servata definita 1: 1-10. London. 27Bessey.E.A. ,1898 .Th ecomparativ emorpholog yo fpistil so fth eRanunculaceae ,Alismacea e andRosaceae .Botanica l Gazette 26: 297-312. 28Blackmore ,S. , 1984. Pollenfeature s andplan tsystematics .In : V.H.Heywoo dan dD.M . Moore (eds.) Current concepts inplan t taxonomy, 135-154.Academi c Press,London . 29Blair,F.M. ,1959.Raisin glarge-flowere dClematis fro mseed .GardenJourna l9:11,14-15 , 29. 30Boissier ,E. ,1845 .Diagnosi sPlantaru mOrientalu mNovarum ,No .6 ,Clematis ispahanica 1-2. Leipzig. 31Boissier , E., 1867.Ranunculaceae . Flora Orientalis, 1: 1-98. H. Georg, Basel,Geneve . 32Boom , B.K., 1980. Flora der cultuurgewassen van Nederland, deel 1 Nederlandse Dendrologie, lOedr . Clematis,174-179 .Veenman , Wageningen. 252 33Bos ,J.J. ,P .Graven ,W.L.A .Hetterschei d &J.J . vand eWege , 1992.Wil dan d cultivated Dracaenafragrans. Edinburgh Journal of Botany 49: 311-331. 34Brandenburg ,W. , 1976.Ee ntaxonomisch everkennin gva nd egraa dva nverwantscha ptusse n soortenbinne nhe tgeslach t ClematisL .Dee l 1: Overzichtva nhe tgenu sClematis L .L H vakgr. Taxonomie van Cultuurgewassen en -begeleiders, Wageningen. 35Brandenburg ,W. , 1977a.Ee ntaxonomisch everkennin gva nd egraa dva nverwantscha ptusse n soortenbinne nhe tgeslach tClematis L .Dee l2 :Kruisinge nbinne nhe tgeslach tClematis L . LHvakgr . Taxonomie van Cultuurgewassen en -begeleiders, Wageningen. 36Brandenburg ,W. , 1977b.Ee ntaxonomisch everkennin gva nd egraa dva n verwantschap tussensoorte nbinne nhe tgeslach tClematis L .Dee l3 :Experimente dtaxonomisch ebenaderin g vanhe tgenu sClematis L .e nd ecytologisch emogelijkhede ndaarvan .L Hvakgr .Taxonomi e van Cultuurgewassen en-begeleiders , Wageningen. 37Brandenburg ,W.A. , 1981.Historicalbackgroun dan dtaxonom yo fcultivated , large-flowered Clematis inEurope . Kulturpflanze 29:321-323 . 38Brandenburg ,W.A. , 1983.Taxonom yo fcultivate dplant swit hregar dt oth ebreedin g value of the accessions. Genetika 15:325-335 . 39Brandenburg ,W .A. ,1984a .Biosystematic san dhybridizatio no fhorticultura lplants .In : W.F. Grant (ed.) Plantbiosystematics , 617-632.Academi c Press,Do nMills ,Canada . 40 Brandenburg,W.A. , 1984b.Th eimplication so fth eus eo fcommo nplan tnames .UPO V Publication 341:37-40 . 41 Brandenburg,W.A. , 1984c.Internationa lregistratio no fcultivate dClematis.ICS Newsletter 1:54-55 . 42Brandenburg , W.A., 1985.Th e international Clematis register. ICSNewslette r 4:47-51 . 43Brandenburg ,W.A. , 1986a.Objective si nclassificatio no fcultivate dplants .In:B.T . Styles (ed.)Infraspecifi c classificationo fwil dan dcultivate dplants ,87-98 .Clarendo nPress ,Oxford . 44Brandenburg ,W .A. ,1986b .Classificatio no fcultivate dplants .Act aHorticultura e182 :109 - 115. 45Brandenburg ,W .A. ,1989a .Clematis. In:S.M .Walter se tal .(eds. )Th eEuropea n Garden Flora HI, 357-364.Cambridg e University Press,Cambridge . 46Brandenburg ,W.A. ,1989b . Clematis- ee nkeu rva nkleinbloemigen .Groe i& Bloe i 1989 (11): 38-41. 253 47Brandenburg ,W .A. ,1991 .Th enee dfo rstabilize dplan tname si nagricultur ean dhorticulture . In:D.L .Hawkswort h(ed. )Improvin gth estabilit yo fnames : needsan doptions . Regnum vegetabile 123: 23-31. 48 Brandenburg,W.A. ,A .va nderNeut,&C.E .Jarvis ,1987 .Lectotypificatio nan d description of Clematis orientalis L. (Ranunculaceae). Taxon 36: 119-126. 49Brandenburg ,W.A. ,& F .Schneider , 1983.Plantentaxonomi ei nonderzoe ke n dagelijks gebruik. Bedrijfsontwikkeling 14:69-74 . 50Brandenburg ,W.A.. &F.Schneider , 1985.Th eimplication sofplan ttaxonom yfo ragricultura l research. Chronica Horticulturae 25: 1-3. 51Brandenburg ,W.A., &F.Schneider , 1988.Cultivargroupin ginrelatio nt oth eInternationa l Codeo fNomenclatur e for Cultivated Plants.Taxo n 37: 141-147. 52Brandenburg ,W.A. ,E.H .Oost ,& J.G .va n deVooren , 1982.Taxonomi caspect s of the germplasmconservatio no fcross-pollinate dcultivate dplants .In :E .Porcedd u& G .Jenkin s (eds.) Seed regeneration of cross-pollinated species, 33-41. Balkema, Rotterdam. 53Brandenburg ,W.A. ,& J.G .va nd eVooren , 1982.Taxonom yan dhistor yo f large-flowered cultivated Clematis inEurope . 21st Int.Hort . Congr., vol.n , 1976.Abstr . 54Brandenburg ,W.A. ,& J.G .va nd eVooren ,1984 .Preparatio no fa tentativ echecklis to flarge - flowered Clematis cultivars.IC SNewslette r 2:25-26 . 55Brandenburg , W.A., &J.G . van de Vooren, 1986.Geschiedeni s van degrootbloemig e Clematis. Dendroflora 22(1985 ,publishe d in 1986):29-32 . 56Brandenburg , W.A., &J.G . van de Vooren, 1988a. Large-flowered Clematis; species, hybrids andcultivars .Clemati s 88: 69-75.(Letland , InRussian) . 57Brandenburg ,W.A., &J.G .va nd eVooren ,1988b .Th eCfemari sDiversifoli agroup . Clematis International 1988:7 . 58Brandenburg ,W.A .& J.G .va nd eVooren ,i nprep .Checklis to flarge-flowere d Clematis cultivars. 59Bremer ,K. ,& H.-E .Wanntorp ,1978 .Phylogeneti csystematic si nbotany .Taxo n27 : 317- 329. 60Bremer,K.,&H.-E .Wanntorp ,1981.Thecladisticapproachtoplantclassification . In:V.A . Funk& D.R .Brook s(eds. )Advance si ncladistics ,87-94 .Ne wYor kBotanica lGarden ,Ne w York. 254 61Bretschneider ,E. , 1898.Histor yo fEuropea nBotanica lDiscoverie si nChina .Facsimile , Zentral-Antiquariat derDDR ,Leipzig , 1167pp. 62Brewbaker ,J.L. , 1957.Polle ncytolog yan dincompatibilit y systemsi nplants .Journa lo f Heridity 48:217-277 . 63Brickell ,CD. , 1973.Problem si nhorticultura lnomenclature .In :P .Gree n(ed. )Plant swil dan d cultivated, 102-113.Hampton . 64Brickell , CD., 1980.Internationa l Codeo f Nomenclature for Cultivated Plants. 65Brooks ,D.R. ,J.N .Caira ,T.R .Piatt ,& M.R .Pritchard , 1984.Principle san dmethod so f phylogeneticsystematics :a Cladistic sworkbook .Universit yo fKansa sMuseu mo fNatura l History, Lawrence,Kansas . 66Brooks , D.R., & E.O. Wiley, 1985. Theories and methods in different approaches to phylogenetic systematics.Cladistic s 1: 1-11. 67BroulancLM. , 1935.RecherchessurrAnatomiefloraledesRanunculacees.LeBotaniste 27(1- 6): 1-278. 68Brummitt ,R.K. ,1976 .A reconsideratio no fClematopsis (Ranunculaceae )i nAfric a with special reference toMalawi .Ke w Bulletin 31: 156-162. 69Brummitt ,R.K.,&CF .Powell ,1992 .Author so fplantnames .Roya lBotani cGardens ,Kew , 732pp. 70Brunfels , O., 1530.Herbaru m vivae eicones. Strassbourg,266pp . 71Buck , G.S., 1967.Garde nros eclassification . American RoseMagazin e 19:6-7,24-25 . 72Bunge ,A.A .von ,1833.Fjiumerationplantarum,quasmChinaborealicollegitann o1831 .St . Petersburg,73pp . 73Bunge ,A.A .von , 1854.Memoire spresente sa l'Academi eImperial ede sScience sd e St. Petersbourg(pa rdiver ssavants )torn .VH ,5 .St .Petersbourg .(Beitra gzu rKenntni sde rflor a Russlands). 74Burtt ,B.L. ,1970 .liifraspecifi ccategorie si nflowerin gplants .Biologica lJourna lo fth eLinnaea n Society 2:233-238 . 75Cain ,A.J., &G.A.Harrison ,1958 .A nanalysi so fth etaxonomist' sjudgemen to faffinity . Proc. Zool. Soc.Londo n 131:85-98 . 255 76Camp ,W.H. , 1947.Distributiona lpattern si nmoder nplant san dth eproblem so f ancient dispersals.Ecol . Monogr. 17:159-183 . 77 Camp,W.H. , 1951. Biosystematy.Brittoni a 7: 113-127. 78 Camp,W.H. , &C.L . Gilly, 1943. The structure and origin of species. Brittonia 4:323-385 . 79 Candolle,A.L.P.P .de , 1883.0riginedesplantescultivees.GermerBailliereetCie, Paris, 377pp. 80Candolle ,A.P .de ,1818.Regnivegetabilissystemanaturalisvol.I :Clematis, 130-169. Paris. 81Carpenter ,J.M. ,1988 .Choosin gamon gmultipl eequall yparsimoniou scladograms .Cladistic s 4: 291-296. 82Carson ,H.L. ,1985 .Unificatio no fspeciatio ntheor yi nplant san danimals .Systemati cBotan y 10:380-390 . 83Chatfield ,C ,& A.J .Collins ,1980 .Introductio nt omultivariat eanalysis .Chapma nan dHall , London, 246pp. 84Clausen ,J. , 1951.Stage si nth eevolutio no fplan tspecies .Cornel lUniversit yPress ,Ne w York,206pp . 85Clause nJ. ,D.D .Kec k& W.M .Hiesey ,1940 .Experimenta lstudie so nth enatur eo f species I.Effec to fvarie denvironment so nNort hAmerica nplants .Carnegi eInstitutio no fWashingto n Publication 520:1-452 . 86Clause nJ. ,D.D .Keck &W.M .Hiesey ,1947 .Heredit yo fgeographicall yan d ecologically isolated species.America n Naturalist 81: 114-133. 87Clause nJ. ,D.D .Kec k& W.M .Hiesey ,1948 .Experimenta lstudie so nth enatur eo f species in.Experimenta lresponse so fclimati crace sof Achillea. Carnegi eInstitutio no fWashingto n Publication 581: 1-129 88Clifford , H.T., &W . Stephenson, 1975.A n introduction to Numerical Classification. Academic Press,Ne w York, 229pp. 89Clokie.H.N. ,1964 .A naccoun to fth eherbariaofth edepartmen tofbotan yi nth euniversit y of Oxford. Oxford, 280pp. 90Clusius ,C , 1601.Histori a rariorum plantarum. Plantijn, Antwerpen. 91Clusius ,C , 1605-1611.Exoticoru m libri decern.Plantijn , Antwerpen. 256 92Conger , A.D., &L.M .Fairchild , 1953. Aquickfreez e method for making smear slides permanent. StainTechnolog y 28:281-283 . 93Cox ,C.B. , 1990.Ne wgeologica ltheorie san dol dbiogeographica l problems.Journa lo f Biogeography 17: 117-130. 94Coyne ,J.A. , 1974.Th eevolutionar y origin of hybrid inviability. Evolution 28:505-506 . 95Cracraft , J., 1975.Historica l biogeography and earth history: perspectives for a future synthesis.Annal s of the Missouri Botanical Garden 62:227-250 . 96Crawford ,D.J. ,1985 .Electrophoreti cdat aan dplan tspeciation .Systemati cBotan y10 : 405- 416. 97Crisci , J., &T . Stuessy, 1980.Determinin g primitive character states for phylogenetic reconstruction. Systematic Botany6 : 112-135. 98Cremers.G. ,1973 .ArcWtecturedequelqueslianesd'AfriquetropicaleI.Candollea28 : 249- 280. 99Cremers,G. , 1974.Architectur edequelqueslianesd'Afriquetropicalell.Candoile a29 :57 - 110. 100Cremers , G., 1975.Su r la presence de dix modeles d"architecture vegetative chezle s Euphorbesmalgaches.ComptesRenduesdel'AcademiedesScience sPari s281: 1575-1578. 101Croizat ,L. , 1962.Spac etim eform : thebiologica l synthesis.Croizat , Deventer, 880pp. 102Croizat ,L. ,1964 .Thought so nhig hsystematics ,phytogen yan dflora lmorphology ,wit ha not e onth e origin of theAngiospermae . Candollea 19: 17-96. 103Croizat , L., G. Nelson & D.E. Rosen, 1974. Centers of origin and related concepts. Systematic Zoology 23: 265-287. 104Cronquist , A., 1987.A botanical critique of cladism. Botanical Review 53: 1-52. 105Dahlgren,R.,&K.Bremer ,1985.MajorcladesoftheAngiosperms.Cladisticsl : 349-368. 106Danert,S. ,1962.UberGliederungsproblemebeiKulturpflanzen.KulturpflanzelO : 350-358. 107Darlington ,CD. , &L.F .L aCour , 1976.Th ehandlin go fchromosomes .Georg eAlle n& Unwin Ltd.,London , 201pp. 108Daumann ,E. ,& Z .Slavikova , 1968.Zu rBlutenmorphologi e der tschechoslowakischen 257 Clematis-Arten. Preslia 40:225-244 . 109Davis,P.H. ,1978 .Th emovin gstaircase :a discussio no ntaxonomi cran kan daffinity . Notes form the Royal Botanic Garden Edinburgh 36: 325-340. 110Davis ,P.H. ,& V.H .Heywood ,1965 .Principle so fangiosper mtaxonomy .Olive r& Boyd , Edinburgh, 556pp. 111Decamps,O. ,1975.StructurenodalecotyledonaireetfohairedesC7«nari. s (Ranunculaceae). Bulletin de Societe Botanique deFranc e 122:125-138 . 112Decamps , O., 1976. Ontogenese des Renonculacees - essai d'utilisation demethodes quantitatives.Universit e Paul Sabatier,Toulouse , 311pp. 113Decamps ,O. ,1979 .Ontogeni enodal eche zle sRenonculacees .Bulleti nd eSociet eBotaniqu e deFranc e 126,lettre sbotanique s 1979 (4):461-471 . 114Denve rStud yGroup ,1960 .A propose dstandar dsyste mo fnomenclatur eo fhuma n mitotic chromosomes. Acta genetica 10:322-328 . 115Dewey , D.R., 1984. The genomic system of classification as a guide to intergeneric hybridizationwit hth eperennia lTriticeae .In :J.P .Gustaffso n (ed.)Gen emanipulatio ni nplan t improvement, 209-279.Plenu m Press,Ne w York. 116Dice , L.R., & H.J. Leraas, 1936. A graphic method for comparing several setso f measurements. Contrib.Lab .Vert . Genet. Univ.Michiga n No.3 : 1-3. 117Dilcher ,D.L. ,1974 .Approache st oth eidentificatio n ofangiosper mlea fremains .Botanica l Review 40: 1-157. 118Dilcher , D.L., &P.R . Crane, 1984.I npursui t of the first flower. Natural History 93: 56-61. 119Dillenius ,J.J. , 1732.Hortu s Elthamensis seu plantarum. London, 437pp. 120Dioscorides , P., J. Goodyer & R.I. Gunther, 1934. The Greek Herbal of Dioscorides. University Press,Oxford , 701pp. 121Dobzhanski ,Th. ,1935 .A critiqu eo fth especie sconcep ti nbiology .Philosoph yo fScienc e2 : 344-355. 122Dodonaeus ,R. ,1583 .Stirpiumhistoriaepemptade sse xsivelibr i30 .Plantijn , Antwerpen, 859pp. 123Donald ,K. , 1986.Th eRoya lHorticultura lSociety :It srol ea sth einternationa l registration 258 authority for Narcissus. ActaHorticultura e 182:381-386 . 124Druce , C.G., &S.H . Vines, 1907.Th eDillenia n herbaria. Oxford. 125Duncan ,T. ,R.B .Philips ,& W.H .Wagner ,Jr. ,1980 .A compariso no fbranchin g diagrams derived byvariou spheneti c and cladistic methods. Systematic Botany 5:264-293 . 126DuRietz ,G.E. ,1930 .Th efundamenta lunit so fbiologica ltaxonomy .S vens kBotanis k Tidskrift 24:333-428 . 127Duyvendak ,R. ,B .Luesin kan dH .Vos ,1981 .Delimitatio no ftax aan dcultivar so fre d fescue (Festuca rubraL . sensu lato).Rasen-Turf-Gazo n 3: 53-62. 128East ,E.M. , 1940.Th edistributio no fself-sterilit y inflowerin gplants .Proceeding so f the American Philosophical Society 82:449-518 . 129Edgeworth , P., 1851.Clematis parvifolia. Transaction s of theLinnea n Society 20:25 . 130Ehrlich ,P. , 1961.Ha sth ebiologica l speciesconcep toutlive dit susefulness ? Systematic Zoology 10: 167-176. 131Eldredge,N. , 1985.Unfinishe d synthesis:biologica lhierarchie san dmoder n evolutionary thought. Oxford University Press,Ne w York, 237pp. 132Eldredge ,N. , 1989.Macroevolutionar y dynamics, species,niche s and adaptive peaks. McGraw-Hill, New York, 226pp. 133Eldredge ,N. ,& Cracraft , 1980.Phylogeneti cpattern san dth eevolutionar yprocess :metho d andtheor yi n comparative biology. 349pp. 134Endler ,J.A. ,1977 .Geographi cvariations ,speciation ,an dclines .Princeto nUniversit yPress , Princeton, 246pp. 135Erdtman ,G. ,1943 .A nintroductio nt opolle nanalysis .Chronic aBotanic aCompany ,Waltham , Mass., USA, 239pp. 136Estabrook ,G.F. ,1972 .Cladisti cmethodology :a discussio no fth etheoretica lbasi sfo r the induction of evolutionary history. Annual Review of Ecology and Systematics 3:427-456 . 137Estabrook ,G.F. , 1978.Som econcept sfo rth eestimatio no fevolutionar yrelationship s in systematic botany. Systematic Botany 3: 146-158. 138Estes ,J.R. ,& R.J .Tyrl , 1982.Th egeneri cconcep tan dth egeneri ccircumscriptio ni nth e Triticeae: an end paper. In: J.R. Estes, R.J. Tyrl, & J.N. Brunken (eds.) Grasses and 259 grasslands, systematicsan d ecology, 145-164.Universit y of Oklahoma Press, Norman. 139Eyde ,R.H. ,1975 .Th ebase so fangiospermphylogeny :flora lanatomy .Annal so fth eMissour i Botanical Garden 62:521-537 . 140Ezelarab ,G.E. ,& K.J .Dormer , 1963.Th eorganizatio no fth eprimar yvascula rsyste mi n Ranunculaceae.Annal s of Botany27 :23-38 . 141Farris ,J.S. ,1969 .A successiv eapproximation sapproac ht ocharacte rweighting . Systematic Zoology 18:374-385 . 142Farris,J.S. ,1971 .Th ehypothesi so fnonspecifit yan dtaxonomi ccongruence .Ann .Rev .Syst . Ecol.2 :277-302 . 143Farris.J.S. ,1973a .A probabilit ymode lfo rinferrin gevolutionar ytrees .Syst .Evol .22 :250 - 256. 144Farris ,J.S. , 1973b.O ncomparin gth eshape so fevolutionar ytrees .Systemati cZoolog y22 : 50-54. 145Farris ,J.S. ,1974.Forma ldefinition so fparaphyl yan dpolyphyly .Systemati cZoolog y23 : 548- 554. 146Farris ,J.S. ,1980 .Th einformatio nconten to fth ephylogeneti csystem .Systemati cZoolog y28 : 483-519. 147Farris ,J.S. , 1985.Th epatter n of cladistics.Cladistic s 1: 190-201. 148Farris ,J.S. , 1988.Henni g -Hennig8 6 reference. 149Farris,J.S. ,1989 .Th eretentio ninde xan dth erescale dconsistenc yindex .Cladistic s5 :417 - 419. 150Farris ,J.S. , 1990.Phenetic s in camouflage. Cladistics 6:91-100 . 151Farris ,J.S. , 1991.Henni g defined paraphyly. Cladistics 7:297-304 . 152Fedorov , A.A., 1969.Chromosom e numbers of flowering plants.Koenigstein , 926pp. 153Feldmann,M. ,1983 .Gen etransfe rfro mwil dspecie sibt ocultivate dplants .Genetik a15 :145 - 161. 154Finet ,A. ,& F .Gagnepain , 1903.Contribution sa l aflor ed e1'Asi eoriental e- Ranunculacea e 1.Clematis L.Bulleti n del a Soci6teBotaniqu e deFranc e 50:518-557 . 260 155Fink ,W. ,1982 .Th econceptua lrelatio nbetwee nontogen yan dphylogeny .Paleobiolog y 8: 254-264. 156Fischer ,C.E.C. , 1937. Clematisvernayi. Bulleti no fMiscellaneou sInformatio nKe w 1937: 95. 157Fish , R.K., 1970.Megagametogenesi s in Clematis and its taxonomican d phylogenetic implications.Phytomorpholog y 20: 317-327. 158Fisk ,J. , 1975.Th e Queen of Climbers.Westleton , 88pp. 159Fletcher ,H.R. ,J.S.L .Gilmour ,G.H.M .Lawrence ,E.L .Littl eJr. ,G .Nilsson-Leissner ,& R . DeVilmorin ,1958 .Internationa lcod eo fNomenclatur efo rCultivate dPlants ,ed .2 .Regnu m Vegetabile 10,28pp . 160Fuchs ,L. , 1542.D ehistori a stirpium commentarii insignes.Basel , 896pp. 161Fuchs ,H.P. , 1958.Historisch e Bemerkungen zumBegrif f der Subspezies.Taxo n 7:44-52 . 162Funk,V.A. ,1985 .Phylogeneti cpattern san dhybridization .Annal so fth eMissour iBotanica l Garden 72:681-715 . 163Funk ,V.A. ,& D.R .Brooks , 1990.Phylogeneti c systematicsa sth ebasi so f comparative biology. Smithsonian Institute,45pp . 164Ghiselin , M.T., 1981.Categories , life and thinking.Behavio r Brain Science4 :269-313 . 165Ghiselin ,M.T. ,1984 .'Definition' , 'Character',an dothe requivoca lterms .Systemati cZoolog y 33:104-110. 166Ghiselin ,M.T. ,1987 .Specie sconcepts ,individuality ,an dobjectivity .Biologica lPilosoph y2 : 127-143. 167Ghiselin , M.T., 1988. Theindividualit y thesis,essences , andlaw s of nature. Biological Philosophy 3: 467-471. 168Gilmour ,J.S.L. , 1961.Taxonomy .In :MacleodandCoble y(eds. )Contemperar y botanical thought. Quadrangle Books,Chicago . 169Gilmou r et al., 1969.Internationa l Codeo f Nomenclature for Cultivated Plants. 170Godley,E.J. , 1977.Imbricat esepal si nC/ema/w .Ne wZealan dJourna lo fBotan y15 :775 - 776. 261 171Good ,R. , 1964.Th egeograph yo fth eflowering plants .Longmans ,Gree nan dCo. , Ltd., London, 518pp. 172Gould,S.J. , 1984.Towar dth evindicatio nofpunctuationa lchange .In :W.A.Berggre n and J.A .Va nCouverin g(eds. )Catastrophe san deart hhistory :th ene wuniformitarianism , 9-34. Princeton University Press,Princeton , New Jersey. 173Gould ,S.J. ,& N .Eldredge , 1977.Punctuate dequilibria :th etemp oan dmod eo f evolution reconsidered. Paleobiology 3: 115-151. 174Gower,J.C.,&G.J.S.Ross , 1969.Minimu mspannin gtree san dsingl elinkag ecluste ranalysis . Applied Statistics 18: 54-64. 175Grant , V., 1971.Plan t speciation. Columbia University Press,Ne w York,435pp . 176Grebenscikov ,I. ,1949 .Zu rmorphologisc hSystem .Einteilun gvo nZea mays L .unte rbes . Beriicksichtigung der sudbalkanischen Formen.Zuchte r 19:302-311 . 177Grebenscikov ,I. ,1950 .Zu rKenntni sde rKurbisar tCucHrMapepoL.nebsteinige nAngabe n iiber Olkurbis.Zuchte r 20: 194-207. 178Gregory ,W.C. ,194 1 (1940).Phylogeneti can dcytologica lstudie si nth eRanunculacea eJuss . Transactions ofth eAmerica n Philosophical Society 31:443-497 . 179Greilhuber ,J. ,1973.Differentia l stainingofplan tchromosome safte rHC 1treatment s (Hy- bands). Osterr. Bot.Z . 122:333-351 . 180Grey-Wilson ,C , 1986.Clematis orientalis - a muc hconfuse d species.ThePlantsma n7 : 193-204. 181Grey-Wilson ,C , 1989(1991) .Clematis orientalis (Ranunculaceae )an dallies .Ke wBulleti n 44:33-60 . 182Grootendorst , F.J., 1979.Clemati s 'Aureolin'. Dendroflora 15/16:62 . 183Grosser ,J.W .& F.G .Gmitter ,Jr. ,1990 .Somati chybridisatio no fCitrus wit hwil drelative s for germplasmenhancemen t and cultivar development. HortScience 25(2): 147-151. 184Gupta,P.K.,andB.R.Baum . 1986.Nomenclatur ean drelate dtaxonomi cissue si nwheats , triticales and someo f their wildrelatives .Taxo n 35: 144-149. 185Haccius,B. ,1942 .Untersuchunge niibe rdi eBlattstellun gde rGattung Clematis. Botanisches Archive 43: 470-486. 262 186Halle\ F., R.A.A. Oldeman & P.B. Tomlinson, 1978. Tropical trees and forests: an architectural analysis,Springe rVerlag ,Berlin . 187Hamilton ,C.W.,&S.H.Reichards , 1992.Curren tpractic ei nth eus eo fsubspecies ,variet y and forma in theclassificatio n of wild plants.Taxo n 41:485-498 . 188Hara.H. ,1978 .Ne wo rnoteworth yflowerin gplant sfro measter nHimalay a(21) - Clematis tibetanaO .Kuntze .Journa l of Japanese Botany 53: 135. 189Hara ,H. ,& S.L.J .Williams ,1979 .A nenumeratio no fth eflowerin gplant so fNepa lJJ :16 . London. 190Harlan ,J.R. , &J.M.J , deWet , 1963. Thecompilospecie sconcept . Evolution 17:497-501 . 191Harlan ,J.R., &J.M.J ,d eWet ,1971 .Toward sarationa ltaxonom yo fcultivate dplants . Taxon 20:509-517 . 192Hartwig ,J. , 1892.Illustrierte s Geholzbuch, VHI(2nd .ed.) ,Pau lParey ,Berlin ,656pp . 193Hauser, D.L., & W. Presch, 1991.Th e effect of ordered characters on phylogenetic reconstruction. Cladistics 7:243-265 . 194Hedge,I.C.,&P .Wendelbo ,1978.Pattern so fdistributio nan dendemismi nIran .Note s from theRoya l Botanic Garden Edinburgh 36:441-464 . 195Henderso n S.A., &B.C .Lu , 1968.Stai n Technology 43:233 . 196Heel ,W .A .van ,1981 .A S.E.M.-investigatio no nth edevelopmen to ffre ecarpels .Blume a27 : 499-522. 197Heel , W.A. van, 1983. The ascidiform early development of free carpels, aS.E.M. - investigation. Blumea 28: 231-270. 198Helm ,J. ,1954 .Lactuca sativa i nmorphologisch-systematische rSicht .Kulturpflanz e 2:72 - 129. 199Helm ,J. ,1963 .Morphologisch-taxonomisch eGliederun gde rKultursippe nvo nBrassica oleracea L. Kulturpflanze 11: 92-210. 200Hennig ,W. ,1950 .GrundzugeeinerTheori e derphylogenetischenSystematik. Deutscher Zentralverlag,Berlin ,370pp . 201Hennig ,W. , 1965.Phylogeneti c systematics.Ann .Rev .Entomol . 10:97-116 . 263 202Hennig ,W. , 1966.Phylogeneti c systematics.Universit y of IllinoisPress ,Urbana , 263pp. 203Herr ,J.M. , 1971.A ne wclearing-squas htechniqu efo rth estud yo fovul edevelopmen t in Angiosperms.America n Journal of Botany 58:785-790 . 204 Heiser, C.B., 1986.Economi c botany: Past and future. Economic Botany 40:261-266 . 205Hermsen ,J.G.Th. ,& E .Sawicka , 1979.Incompatibilit y andinconguit yi ntube r bearing Solariumspecies .In :J.G .Hawkes ,R.N .Leste ran dA.D .Skeldin g(eds. )Th ebiolog yan d taxonomy of the Solanaceae, Academic Press,London ,445-453 . 206Hetterscheid ,W.L .A. ,& W.A .Brandenburg ,1995a .Culto nversu staxon :Conceptua lissue s incultivate d plant systematics.Taxo n 44: 161-175. 207Hetterscheid ,W.L.A. ,& W.A .Brandenburg ,1995b .Th eculto nconcept :settin gth estag efo r anunambiguou s taxonomy of cultivated plants.Act aHorticultura e 413:29-34 . 208Hetterscheid , W.L.A., &R.G .va n denBerg , 1996.Cultonom yo fAster. Act a Botanica Neerlandica 45(2) : 173-181. 209Hetterscheid ,W.L.A. ,R.G .va nde nBerg ,& W.A .Brandenburg ,1996 .A nannotate dhistor y ofth eprinciple so fcultivate dplan tclassification .Act aBotanic aNeerlandic a4 5(2) : 123-134. 210Heyting , J., W.A. Brandenburg, A.H. van Keulen & J.G. van de Vooren, 1980. Nectarafscheidingi nClematis bloemen .Projec t004.761/80. 5Taxonomie ,L HWageningen . 211Hickey ,L.J. , 1977.Earl ycretaceou sfossi levidenc efo rangiosper mevolution .Botanica l Review 43:2-104 . 212Hoffman,M.H.A. , 1996.CultivarclassificationofPMde/p/ii«L.(Hydrangeaceae). Acta Botanica Neerlandica4 5(2) : 199-210. 213Hogenboom,N.G. , 1973.A mc>de lfo rmcongruit yi nmtirnat epartne rrelationships .Euphytic a 22:219-233 . 214Hogenboom,N.G. , 1975.Incompatibilit yan dincongruity :tw odifferen tmechanism sfo rth e non-functioning ofintimat epartne rrelationships .Proceeding so fth eRoya lSociet yLondo n SeriesB 188:361-375 . 215Hogenboom,N.G. ,1979a .mcompatibilit yan dincongruit yi nLycopew/con .In :J.G .Hawkes . R.N.Leste r& A.D .Skeldin g(eds. )Th ebiolog yan dtaxonom yo fth eSolanaceae ,435-444 . Academic Press,London . 216Hogenboom ,N.G. ,1979b .Exploitatio no fincongruity ,a ne wtoo lfo rhybri dsee dproduction . 264 In: A.C.Zeven ,& A.M .va nHarte n(eds. )Broadenin gth egeneti cbas eo fcrops ,299-309 . Pudoc, Wageningen. 217 Hogenboom,N.G. ,1983a .Bridgin ga ga pbetwee nrelate dfields o fresearch : Pistil-pollen relationshipsan dth edistinctio nbetwee nincompatabilit yan dincongruit yi nnonfunctioni nhost - parasite relationships. Phytopathology 73:381-383 . 218Hogenboom ,N.G. ,1983b .Incongruity :Nonfunctionin go fintercellula ran dintracellula rpartne r relationshipsthroug hnonmatchin ginformation .In :J .Heslop-Harriso n& H.F .Linsken s(eds. ) Cellularinteractions ,Encyclopedi ao fPlan tPhysiology ,ne wseries, .Springe rVerlag ,Berlin . 219Hooker ,J.D. ,1900 .Clematis orientalis var .tangutica. Curtis'Botanica lMagazin e12 6tab . 7710. 220Hooker ,J.D.,&Th.Thomson , 1855.FloraIndica,vol .l.Clematideae,3-12 .W .Pamplin , London. 221Hooker ,J.D. ,& Th .Thomson , 1872.Ranunculaceae .In :J.D .Hooke r(ed. )Flor ao f British India, vol. l(l),l-30. L.Reev ean dCo. ,London . 222Hooker ,W.J. , 1837. 223Hooker ,W.J., &J .Smith , 1850. Clematisgraveolens. Curtis' sBotanica lMagazin e7 6tab . 4495. 224Hoot ,S.B. ,A.A .Reznice k& J.D .Palmer , 1994.Phylogeneti crelationship sin Anemone (Ranunculaceae)base do nmorpholog yan dchloroplas tDNA .Systemati cBotan y19: 169- 200. 225Hort ,A. ,1916 .Theophrastu senquir yint oplant san dmino rwork so nodour san dwate rsigns . LoebClassica l Library,London , 2vol . 226Hull ,D.L. , 1976.Ar e species really individuals? Systematic zoology 25: 174-191. 227Humphries ,C.J. ,& V.A .Funk ,1984 .Cladisti cmethodology .In :V.H .Heywoo dan dD.M . Moore (eds.) Current concepts inplan t taxonomy, 323-362.Academi c Press,London . 228Humphries ,C.J. ,P.Y .Ladiges ,M .Roos ,& M .Zandee , 1988.Cladisti cbiogeography .In : A.A.Myer s& P.S .Giller s(eds. )Analytica lbiogeograph y- a nintegrate dapproac ht oth e study of animal andplan t distributions, 371-404. Chapman &Hall ,London . 229 Hutchinson, J., 1920.Clematopsis, aprimitiv e genus of Clematideae. Kew Bulletino f Miscellaneous Information 1920:12-22 . 265 230Huxley ,J.S. , 1940.Th e new systematics. Oxford University Press,Oxford , 583pp. 231Huxley ,J.S. , 1958.Evolutionar yproces san dtaxonom ywit hspecia lreferenc et ogrades . UppsalaUniv .Arsskr .6 :21-39 . 232Ikonnikov , 1977.Clematis sarezica. Novosti Sist.Vyssh .Rast . 14: 231. 233Jarvis ,C.E. ,1986 .Th eLinnaea nplan tnam etypificatio nproject .Act aHorticultura e182 : 79- 88. 234Jeannoda-Robinson ,V. , 1977.Contributio na 1 'etud ed el'architectur ede sherbes .These . Academied eMontpellier . Universite des Sciences ettechnique s duLanguedoc .76pp . 235Jeffrey , C, 1968.Systemati c categories for cultivated plants.Taxo n 17: 109-144. 236Jeffrey , C, 1982.Kingdoms ,code s and classification. Kew Bulletin 37:403-416 . 237Jensen ,U. , 1968.Serologisch eBeitrag ezu rSystemati k derRanunculaceae . Botanische Jahrbucher 88:204-268 . 238Jensen ,U. , 1971.Zu rSystematischeStellun gde rHelleborina e(Ranunculaceae) .Taxo n20 : 747-758. 239Jirasek,V. ,1966 .Th esystematic so fcultivate dplant san dthei rtaxonomi ccategories .Presli a 38: 267-284. 240Johnson ,M. , 1998.Slakte t Clematis. Sodertalje. 241Jouin,E. , 1907.DieinDeutschlandkultivierten,winterhartenC/emah\y.Mitteilungende r Deutschen Dendrologischen Gesellschaft 16:228-238 . 242 Kalkman, E.R., 1984.Analysi s of the C-bandedkaryotyp e ofAllium cepaL . Standard system of nomenclature and polymorphism. Genetica 65: 141-148. 243 Kho,Y.O.,&J.Baer , 1968.Observin gpollentube sb ymean so ffluorescence .Euphytic a17 : 298-302. 244Kho ,Y.O .A.P.M .denNij s& J .Franken ,1980 .Interspecifi chybridizatio ni nCucumis L. R. Thecrossabilit y of species, aninvestigatio n of in vivopolle ntub egrowt h and seed set. Euphytica 29:661-672 . 245Kinzel,W. , 1913.FrostundLichtalsbeeinfluBendeKraftebe i derSamenkeimung.Ulmer , Stuttgart, 170pp. 266 246Klotzsch ,J.F. ,1862 .Di erxrtamschenErgebniss ede rReis eseine rKonigl.Hoheitde sPrinze n Waldemarvo nPreusse nau fCeylon ,de nHimalay aun da nde nGrenze nvo nTibe tgesammelt e Pflanzen. Berlin 164pp. 247 Knuth,P. , 1898.Handbuc h derBlutenbiologi e n, 1,1-55.W .Engelmann .Leipzig . 248 Knuth, P., 1904.Handbuc h derBlutenbiologi e 111,1,192-301.W .Engelmann , Leipzig. 249Koehne,E. , 1893.Clematis. Deutsch eDendrologie ,152-160 ;567 .Ferdinan dEnk eVerlag , Stuttgart. 250Komarov ,V.L. , 1904.Clematis intricata var.serrata. Act aHort iPetropolitan i 22:289 . 251 Komarov,V.L. , 1931.Clematis serrata. Alls. Keyfo rth e Plantso fth e FarEas t Region USSR 1:549 . 252Koopman,W.P.M.,DeJong ,J.H.&I.M.DeVries , 1993.Chromosom ebandin gpattern si n lettuce species (Lactucasubsect .Lactuca, Compositae).Plan t Syst.Evol . 185:249-257 . 253 Kornet,D.J., 1993.Reconstructingspecies-Demarcation si ngenealogica lnetworks .Instituu t voorTheoretisch e Biologie- Rijksherbarium/Hortu s Botanicus, Leiden, 120pp. 254 Korshinsky,S. , 1898.Fragment aflora e Turkestaniae: 1. Clematistangutica; 2.Clematis orientalisvar .roschanica. Bulleti nd el'Acad6mi eImperial ede sScience sd eSt.-Petersbour g Ve ser.,vol .9 :399-400 . 255 Koster,H.,&F.Schneider . 1982.A multilingua lglossar yo fcommo nplant-name sI .Fiel d crops, grasses andvegetables .2n d ed.ISTA ,Zurich . 256 Kosuge,K.,&M.Tamura,1989 .Ontogeneti cstudie so npetal so fth eRanunculaceae .Journa l of Japanese Botany 64:65-67 . 257 Kotschy,C.G.T. , 1843.Abbildunge nun dBeschreibunge nneuerun dseltenerThier eun d Pflanzen in Syrien (und imwestliche n Taurus gesammeltvo nTh . Kotschy),vol. 1Botanie . 258 Krasheninnikov,I.M. , 1937(1970) .Clematis. In:V.L .Komaro ve tal .(eds. )Flor ao f the USSR, vol.VJJ ,240-250 .Moscow-Leningrad . English translation Jerusalem. 259 Krassilov, V.A., P.V. Shilin & V.A. Vakhrameev, 1983. Cretaceous flowers from Kazakhstan. Review of Paleobotany andPalynolog y 40: 91-113. 260 Kroon, G.H., J.B.M. Custers, Y.O. Kho, A.P.M. den Nijs & H.Q. Varekamp, 1980. Intersepcifichybridizatio ni nCucumis L .I .Nee dfo rgeneti cvariation ,biosystemati crelation s andpossibilitie s to overcome crossablity barriers.Euphytic a 28:723-728 . 267 261 Kovalevskaya,S. ,1967 .Clematis hilariae. Notula eSystematica eHerbari iInstitut i Botanici Academiae Scientiae Uzbekistan 18:34 . 262Kriissmann , G., 1976.Handbuch e derLaubgeholze , Bd. 1.Pau lParey ,Berlin . 263 Krutzsch,W. , 1989. Paleographyan dhistorica lphytogeograph y (paleochorology)i nth e Neophyticum.Plan t Systematicsan dEvolutio n 162: 5-61. 264 Kumazawa, M., 1936. Pollen morphology in Ranunculaceae, Lardizabalaceae and Berberidaceae. Japanese Journal of Botany 8:13 . 265 Kuntze, O., 1885. Monographic der Gattung Clematis. In: I. Urban et al. (eds.) Verhandlungen desBotanische n Vereins derProvin z Brandenburg, 83-202.Berlin . 266Kurita,M. ,1956 .Cytologica lstudie si nRanunculaceae .Botanica lMagazin e(Tokyo )69 : 239- 242. 267 Kurita, M., 1957.Chromosom e studiesi nRanunculacea e IV. Reportso fth e Biological Institute of theEhim eUniversit y 2. 268 Kurita,M. ,1958a .Chromosom estudie si nRanunculacea eVHJ .Report so fth e Biological Institute of theEhim eUniversit y 5 269 Kurita, M., 1958b. Chromosome studies in Ranunculaceae XI.Memoir s of the Ehime University, UB 3(1): 13-22. 270 Kurita, M., 1960.Chromosom e studies in Ranunculaceae XVI.Memoir s of theEhim e University, UB4(1) :53-58 . 271 Kurita, M., 1962.Chromosom e studies in Ranunculaceae XX. Memoirs of theEhim e University,JJ B4(3) :31-37 . 272Kurita , M., 1964.Chromosom e studies inRanunculacea e XXII.Memoir s of theEhim e University, UB5(1) :31-36 . 273Laar ,H.J . van de, 1980.Clematis 'Helios' .Dendroflor a 25:72 . 274Lam , 1950.Minute s of the Utrecht Conference. Chronica Botanica 12: 12. 275Lamarck ,J.B.A.P.M .de ,1786.EncyclopedieMethodiqueBotanique,vol.2,Clematite , 41- 45. Panckouke,Paris . 276Lange ,W. ,W .A .Brandenbur g& T.S.M .d eBock ,1999 .Taxonom yan dcultonom yo f beet (Beta vulgaris L.).Botanica l Journal of the Linnean Society (London) 130(1) :81-96 . 268 277Langen , F.R. de, Oost, E.H, and C.E. Jarvis. 1984. Lectotypification ofDianthus caryophyllus L.an dD. chinensis L. (Caryophyllaceae). Taxon33 :716-724 . 278Langlet , O., 1932.Ube rChromosomenverhaltniss e undSystemati kde r Ranunculaceae. Svensk Botanisk Tidskrift 26: 381-400. 279Lauener ,D.A. ,1978 .Th etypificatio no fRoyle' sRanunculaceae .Note so fth eRoya lBotani c Garden Edinburgh 36: 127-138. 280Lavallee ,P.A.M. ,1884 .Le sClematite sagrande sfleurs ,t.2 1 (C.orientalis). Baillieres , Paris, 83pp. 281Ledebou r C.F.von , 1830.Flor a altaica, H:373 .Berlin . 282Ledebour , C.F.von , 1841-1853. Flora rossica, vol. 1 Clematideae, 1-5. 283Leppik ,E.E. ,1964 .Flora levolutio ni nth eRanunculaceae .IOW AStat eJourna lo f Science 39: 1-101. 284Leslie ,A.C. , 1982.Th einternationa l LilyRegister . 3rded .Roya lHorticultura l Society, London, 388pp. 285Letschert ,J.P.W. ,1993 .Beta sect .Beta: biogeographica lpattern so fvariation ,an dtaxonomy . Wageningen Agricultural University Papers 93-1.155pp . 286Levan ,A. ,K .Fredg a& AA .Sandberg , 1964.Nomenclatur e forcentromeri cpositio n on chromosomes. Hereditas 52:201-220 . 287Lewontin ,R.C. ,1974 .Th egeneti cbasi so fevolutionar ychange .Columbi aUniversit yPress , New York, 348pp. 288Limpricht,W. ,1922 .AufzahlungdervonDr .W.LimprichtinOstasiengesammeltenPflanzen . Feddes Repertorium Beihefte XII,II-teil :373-376 . 289Lindley , 1846.Journa l of theHorticultura l Society (London) 1846:307-308 . 290Ling , 1980.Flor aReipublica e Populi Sinicae 28: 142,fig. 1 5 291Linnaeus ,C , 1738.Hortu s cliffortianus - Clematis: 225-226.Amsterdam . 292Linnaeus , C, 1753.Specie s plantarum (ed.I ) Clematis: 543-545.Stockholm . 293Linnaeus , C, 1762.Specie splantaru m (ed.II) .Stockholm . 269 294Lobelius ,M. , 1576.Plantara m seu stirpium historia. Plantijn, Antwerpen, 671pp. 295Loconte ,H. ,& D.W .Stevenson , 1990.Cladistic so fth espermatophyta .Brittonia42 : 197- 211. 296Loconte ,H. ,& D.W .Stevenson , 1991.Cladistic so fth eMagnoliidae .Cladistic s7 :267-296 . 297Lotsy,J.P. ,1911.Ranunculaceae .V6rtrageuberbotanischeStammesgeschichte,Bd .3 ,Tl . 1:566-587 .Fischer , Jena. 298Lotsy ,J.P. , 1916.Evolutio n bymean s of hybridization. M.Nijhoff , Den Haag, 166pp. 299Love ,A. ,1982 .Generi cevolutio nofthewheatgrasses.BiologischesCentralblat t 101:191 - 212. 300Loudon ,J.C. ,1844 .Arboretu me tFructicetu mBrittanicum ,vol .1,2n ded. ,Clematis, 232- 246.London . 301Loudon,J.C , 1869.A nencyclopaediao ftree sandshrubs.Clematideae ,2-17 ;1112 . Warne and Co.,London . 302Lundqvist ,A. ,U .?)sterbye ,K .Larse n& I .Linde-Larsen , 1973.Comple x self-incompatibility systems inRanunculus acrisL . andBeta vulgaris L. Hereditas 74: 161-168. 303Mansfeld , R., 1950.Da s morphologische Systemde r Saatgerste.Ziichte r 20: 8-24. 304Mansfeld ,R. ,1953 .Zurallgemeine nSystematikderKulturpflanzen .I .Kulturpflanz e1 :138 - 155. 305Mansfeld ,R. ,1954 .Zurallgemeine nSystemati cde rKulturpflanzen .H .Kulturpflanz e2 : 130- 142. 306Markham,E. , 1951.Th elarg ean dsmal lflowere dClematis ,3rd .ed .Countr yLife , London, 126pp. 307Mayr ,E. , 1940.Speciatio n phenomena inbirds .America n Naturalists 74:249-278 . 308Mayr ,E. ,1976 .Evolutio nan dth ediversit yo flife .Belkna pPres so fHarvar dUniversit yPress , Cambridge, Mass761pp . 309Mayr ,E. , 1978.Origi nan dhistor yo fsom eterm si nsystemati can devolutionar y biology. Systematic Zoology 27:83-88 . 310Mayr ,E. ,1982 .Th egrowt ho fbiologica lthought .Belkna pPres so fHarvar dUniversit yPress , 270 Cambridge,Mass. ,974pp . 311Maximowicz , C.J., 1877(1879) .Specie splantaru m novarumJaponia ee tMandshuriae . Bulletin del'Academi eImperia l des Sciences de St.Petersbur g 22: 210-211. 312Maximowicz , C.J., 1889.Flor a tangutica, 3.St .Petersburg . 313McKelvey , B., 1982.Organizationa l systematics -taxonomy , evolution, classification. University of California Press,Berkeley , 511pp. 314Meeuse ,A.D.J. ,1973 .Co-evolutio nofplanthost san dthei rparasite sa sa taxonomi ctool .In : V.H.Heywoo d (ed.)Taxonom y and Ecology, 289-316.Academi c Press,Ne w York. 315Meglitsch , P.A., 1954.O n thenatur e of the species.Systemati c Zoology 3:49-65 . 316Meicenheimer,R.D. , 1978.Comparativ estud yo nflora lmorphogenesi so fRanunculus. MS Thesis,Washingto n State University, Pullman, 139pp. 317Meicenheimer.R.D. ,1979 .Relationship sbetwee nshoo tgrowt han dchangin gphyllotaxi so f Ranunculus. American Journal of Botany 66:557-569 . 318Meikle ,R.D. , 1957.Wha t isth e subspecies? Taxon 6: 102-105. 319Meurman ,O. ,& E .Therman ,1939 .Studie so nth eChromosom eMorpholog yan dStructura l Hybridity inth egenu s Clematis. Cytologia 10: 1-14. 320Meyer , C.A., 1831. Verzeichniss der Pflanzen, welche wahrend der 1929-1830 unternommeneReis ei mCaucasus.. . gefunden worden sind. St.Petersburg , 241pp. 321Meyer-Abich , A., 1926.Logi k der Morphologie imRahme neine r Logik der gesamten Biologic Springer, Berlin. 286pp. 322Miller ,P. ,1768 .Gardener' sdictonar y(ed .8) .Joh nan dFranci sRivington ,London , 1380pp. 323Minelli , A., 1993. Biological Systematics. Chapman andHall ,London , 387pp. 324Miyamoto ,M.M. ,1985 .Consensu scladogram san dgenera lclassifications .Cladistic s1 :186 - 189. 325Moench ,C , 1794. Methodusplanta short ibotanic ie tagr imarburgensi sa staminu m situ describendi. Marburg, 780pp. 326Moore ,Th. , &G .Jackman , 1872.Clematis asa garde n flower. London, 160pp. 271 327Morisset ,P. ,& C .Boudin , 1984.Th ebiosystemati cimportanc eo fphenotyp cplasticity .In : W.F. Grant (ed.)Plan tbiosystematics , 293-306.Academi c Press,Do n Mills,Canada . 328MS S Sherard: 276-292,528-545 .Roya l Society. London. 329Mttller ,C , 1857.Walpe r Annales Botanices SystematicaeIV :3-9 . 330Nelson ,GjJ. , 1978.Ontogeny ,phylogen yan dth ebiogeneti claw .Systemati cZoolog y27 : 324-345. 331Nelson ,G.J. ,1979 .Cladisti canalysi san dsynthesis :principle san ddefinition swit ha historica l noteo nAdanson' s Families desPlante s (1763-1764). Systematic Zoology 28: 1-21. 332Nelson ,G. ,&N .Platnick , 1981.Systematic sandbiogeography :cladistic san dvicariance . 567pp. 333Nettancourt ,D . de, 1977.Incompatibilit y inAngiosperms . Springer, Berlin,230pp . 334Neut ,A .va nder , 1983.Typificati e vanClematis orientalis L .Landbouwhogeschoo l vg. Taxonomieva n cultuurgewassen en -begeleiders,Wageningen , 49pp. 335Neut ,A .va nder ,& E .Pfeiffer , 1982.Introducti ee ngeneeskrachtig ewerkin gva nEuropes e soortenva nClematis L .i nbotanisch eliteratuu rto tomstreek s 1753.Landbouwhogeschoo l vg.Taxonomi e van Cultuurgewassen en -begeleiders,Wageningen , 31pp. 336Nicholson ,G. , 1888.Th eIllustrate dDictionar yo fGardening ,div.2 ,Clematis,337-340. L. Upcott Gill,London . 337Niethammer , A., 1928. Fortlaufende Untersuchungen iiber den Chemismus der Angiospermensamenun ddi eauBere nnaturliche nwi ekunstliche nKeimungsfaktore n I.de r EinfluB desFroste s auf die Keimungsfahigkeit. Biochemische Zeitung 197:241-244 . 338Nixon ,K.C. ,& Q.D .Wheeler , 1990.A namplificatio n ofth ephylogeneti cspecie s concept. Cladistics 6: 211-223. 339Okada ,H. ,& M .Tamura ,1979 .Karyomorpholog yan drelationshi po fth eRanunculaceae . Journal of Japanese Botany 54:65-77 . 340 Oost, E.H., 1984. xBrassicoraphanus Sageret or xRaphanobrassica Karpechenko? Cruciferae Newsletter 9: 11-12. 341Oost,E.H. ,W.A.Brandenburg ,&C.E .Jarvis ,1989 .Typificatio no fBrassica oleracea L. (Cruciferae)an dit sLinnaea nvarieties .Botanica lJourna lo fth eLinnaea nSociet y(London ) 101:329-345 . 272 342Oost ,E.H. ,W.A .Brandenburg ,G.T.M .Reulin g& C.E .Jarvis , 1987.Lectotypificatio n of Brassicarapa L. ,B. campestris L .an dneotypificatio no fB. chinensis L .(Cruciferae) .Taxo n 36:625-634 . 343Oost .E.H. ,an dH .Toxopeus . 1986.Scop ean dproblem s ofcultiva r groupformatio n as exemplified inBrassica rapa L .Act aHorticultura e 182: 117-123. 344Pandey,K.K. , 1979.Th egenu sNicotiana: evolutio nofincompatibilit yi nflowerin gplants .In : J.G. Hawkes, R.N. Lester and A.D. Skelding (eds.) The biology and taxonomy of the Solanaceae,421-434 .Academi c Press,London . 345Past iJr. ,G. , 1950. ConsulSherard :Amateu rbotanis tan dpatro no flearning , 1659-1728. Type-written thesis.Universit y of Illinois,253pp . 346Persoon , C.H., 1805.Synopsi s plantarum seu enchiridium botanicum, vol. 11358pp. , Clematis, 98-100.Paris . 347Pickersgill , B., 1981.Biosystematic s of crop-weed complexes.Kulturpflanz e 29:377-388 . 348Pickersgill ,B. ,1986 .Evolutio nofhierarchica lvariatio npattern sunde rdomesticatio nan dthei r taxonomictreatment .In :B.T .Style s(ed. )Mraspecifi cclassificatio no fWil dan dCultivate d Plants, 191-209.Clarendo n Press, Oxford. 349Platnick ,N.I. ,1977.Paraphyleticandpolyphyleti cgroups .Systemati cZoolog y26 : 195-200. 350Podlech ,D. ,& A .Dieterle ,1969 .Chromosomenstudie na nafghanische nPflanzen . Candollea 24: 185-243. 351Prantl,K. , 1888.BeitragezurMorphologieundSystematikderRanunculaceen. Botanische Jahrbucher9 :225-273 . 352Putter ,M. ,d e& J.G .va nd eVooren , 1986.Identificatio n ofAllium cepa L.cultivar sb y means of statistical analysis of C-bandedchromosomes .Euphytic a 39: 153-160. 353Queiro s K.de ,andM.J .Donoghue ,1990a .Phylogeneti csystematic so rNelson' sversio no f cladistics? Cladistics 6:61-75 . 354Queiro s K.de ,andM.J. Donoghue ,1990b .Phylogeneti csystematic san dspecie srevisited . Cladistics 6:83-90 . 355Ramachandran ,C ,W.A .Brandenbur g& A.P.M .de nNijs , 1985.Infraspecifi c variation in C-bandedkaryotpy e andchiasm afrequenc y in Cucumis sativus(Cucurbitaceae). Plant Systematics and Evolution 151: 31-41. 273 356RaBner ,E. ,1931 .Primitiv eun dabgeleitet eMerkmal ei mBliitenba ueinige rRanunculaceae . Planta 15: 192-243. 357 Raub,D.M., 1993.Extinctio n-ba dgene so rba dluck ?Oxfor d UniversityPress , Oxford, 210pp. 358Raven ,P.H. ,1975 .Th ebase so fAngiosper mphylogeny :cytology .Annal so fth eMissour i Botanical Garden 62:724-764 . 359Raven,P.H. ,1979 .Plat etectonic san dsouther nhemispher ebiogeography .In :K.Larse n and L.B.Holm-Nielse n (eds.) Tropical Botany, 3-24.Academi c Press,London . 360Raven , P.H., & D.I. Axelrod, 1974. Angiosperm biogeography and past continental movements.Annal s of theMissour i Botanical Garden 61:539-673 . 361Ray ,J. , 1686.Histori a Plantarum, vol. 1.London . 362Ray ,J. ,1724 .Synopsi smethodic astirpiu mBrittannicarum ,thir deditio nedite db yDillenius . London, 512pp. 363Raynal ,J. , 1978.Clematopsis, genr eafricano-malgache : typesbiologique se ttaxonomie . Adansonia, ser.2,18 : 3-18. 364Rehder, A., 1910. Clematis serratifolia spec. nov. Mitteilungen derDeutschen Dendrologischen Gesellschaft 1910: 248. 365Rehder.A. , 1915.Cfe/n«fi5serrar(/bZ!a.Repeitoriumspecierumnovarumregni vegetabilis 13:362 . 366Rehder ,A. ,1920 . Clematisglauca Willd .var .akebioides f.phaeantha, f.nov .Journa lo f theArnol d Arboretum 1: 195. 367Rehder ,A. , 1974.Manua lo fCultivate dTree san dShrubs ,2nd .ed. , 12thprin tClematis, 206-220.MacMilla n Company, New York. 368Rehder ,A. ,& E.H .Wilson ,1913 .In :C.S .Sargen t(ed. )Planta eWilsoniana ea nenumeratio n of the woody plants collected in Western China for the Arnold Arboretum of Harvard University during theyear s 1907, 1908an d 1910,vol . 1Clematis, 319-343. 369Reichenbach , H.G.L., 1837.Handbuc hde snatiirliche nPflanzensystems , 227.Dresden , 346pp. 370Rick ,CM. , 1950.Pollinatio nrelation so fLycopersicon esculentum i nnativ ean d foreign regions.Evolutio n 4: 110-122. 274 371Rick ,CM. , 1995.Tomato .In :J .Smartt&N.W .Simmond s(eds. )Evolutio no fcro pplants , 2nd ed.,452-457 . 372Rick , CM., J.F. Fobes & M. Holle, 1979. Genetic variation inLycopersicon pimpinellifolium:Evidenc eo fevolutionar ychang ei nmatin gsystems .Plan tSystematic s& Evolution 127: 139-170. 373Ridley.M. ,1989 .Thecladisti csolutio nt othespecie sproblem .Biologica lPhilosoph y4 : 1-16. 374Rieppel , O., 1991.Things ,tax aan d relationships. Cladistics 7:93-100 . 375Roche ,E. ,1974 .Paleobotanique ,Paleoclimatologi ee tderiv ede scontinents .Universit eLoui s Pasteur de Strasbourg, Sciences Geologiques Bulletin 27:9-24 . 376Rodenburg,CM. , 1960.Varietie soflettuce ,a ninternationa lmonograph .LV.T. ,Wageningen , 228pp. 377Rosen ,D.E. ,1975 .A vicarianc emode lo fCaribbea nbiogeography .Systemati cZoolog y24 : 431-464. 378Rosen,D.E. ,1978 .Vicarian tpattern san dhistorica lexplanatio ni nbiogeography . Systematic Zoology 27: 159-188. 379Rothfels ,K. ,E .Sexsmith ,M .Heimburger ,& M.O .Krause , 1966.Chromosom esiz ean d DNAconten to dspecie so fAnemone L .an drelate dgener a(Ranunculaceae) .Chromosom a 20:54-74 . 380Royle ,J.F. ,1839 .Dlustration so fth eBotan yan dothe rbranche so fth eNatura lHistor yo fth e HimalayanMountain san dth eFlor ao fCashmere ,vol .1 Ranunculaceae,43-51 .W.H .Alle n and Co.,London . 381Sakamura , T., 1918. Kurze Mitteilung iiber die Chromosomzahlen und die Verwantschapsverhaltnisse der Triticum-Arien. Botanical Magazine (Tokyo) 32: 151-154. 382Sauer ,J.D. ,1957.Recen tmigratio nan devolutio nofth edioeciou samaranths.Evolutio n11 : 11-31. 383 Sauer, J.D., 1967a. The grain amaranths and their relatives: a revised taxonomican d geographic survey.Annal s ofthe Missouri Botanical Garden 54: 103-137. 384 Sauer, J.D., 1967b. Plants and Man on the Seychelles Coast: A study in historical biogeography, University of Wisconsin Press,Madison , 132pp. 385 Sauer,J.D. , 1988.Plan tmigratio n -Th edynamic so fgeographi cpatternin gi nsee dplan t 275 species.Universit y of California Press,Berkeley , 282pp. 386Schaeppi ,H. ,& K .Frank , 1962.Vergleichend-morphologischeUntersuchungenube r die Karpelgestaltung,insbesonder edi ePlazentatio nbe iAnemoneen .Botanisch eJahrbuche r Systematik 81: 337-357. 387 Schmidt,H. , 1965.De r"Hortu sElthamensis "au sde rBibliothe kCar lvo nLinne .Fedde s Repertorium Specierum Novarum 70:69-108 . 388 Schneider, C.K., 1904.Illustrierte s Handbuch derLaubholzkunde , 1: 273-294.Gusta v Fischer Verlag,Jena . 389 Schneider, F., 1984.UPO V and nomenclature. UPOV Publication 341: 9-10. 390Schoffel ,K. ,1932 .Untersuchunge niibe rde nBliitenba uderRanunculaceae .Plant a(Berlin ) 17:315-371. 391Schuster ,R.M. ,1976 .Plat etectonic san dit sbearin go nth egeographica lorigi nan ddispersa l ofangiosperms .In :C.B .Bec k(ed. )Origi nan devolutio no fAngiosperms ,48-138 .Columbi a University Press,Ne w York. 392Schwanitz ,F. ,1967 .Di eEvolutio nde rKulturpflanzen . Bayerischer Landwirtschaftsverlag, Miinchen,463pp . 393 Schwarzacher,,T. ,P .Ambros ,& D .Schweizer , 1980.Applicatio no fGiems abandin g to Orchid karyotype analysis.Plan t Systematics and Evolution 134:293-297 . 394 Scopoli,J.A. , 1760.Flor a Carniolica.Wien ,608pp . 395 Scotese,C.R. , &W.S .McKerrow , 1990.Revise dworl dmap san dintroduction .In :W.S . McKerrow& C.R .Scotes e(eds. )Palaeozoi cPalaeogeograph yan dbiogeography .Geologica l SocietyMemoi r No. 12,243-242 . 396 Shambulingappa, K.G., 1965.Th e occurrence of Bchromosome s in Clematis. Current Science 34: 670-671. 397Sharma ,A.K .& A .Sharma ,1972.Chromosom eTechnique s- Theor yan dPractice ,2nd .ed . Butterworths,London , 575pp. 398Simpson ,G.L. ,1961 .Principle so fanima ltaxonomy .Columbi aUniversit yPress ,Ne wYork , 247pp. 399 Sluiman, H.J., 1985. A cladistic evaluation of the lower and higher green plants. (Viridiplantae). Plant Systematics and Evolution 149:217-232 . 276 400 Smith, E.P., 1928. A comparative study of the stem structure of the genusClematis. Transactions of theRoya l SocietyEdinburg h 55(3/26):644-664 . 401Snogerup ,1980 .Th ewil dform so fth eBrassica oleracea grou p(2n =18 )an dthei rrelation s toth ecultivate dones .In :S .Tsunoda ,K .Hinat a& C .Gomez-Camp o(eds. )Brassica Crop s andthei r wild Allies, 121-132.Japa n Scientific SocietyPress ,Tokyo . 402 Sober, E., 1975.Simplicity . Clarendon Press, Oxford. 403 Sober,E. ,1983 .Parsimon yi nsystematics :philosophica lissues .Annua lRevie wO fEcolog y and Systematics 14:335-358 . 404 Sober, E., 1984. Conceptual issues in evolutionary biology: an anthology. MITPress , Cambridge,Mass. ,725pp . 405 Sober, E., 1986.Parsimon y andcharacte r weighting. Cladistics 2:28-42 . 406 Sober,E. ,1988 .Reconstructin gth epast :Parsimony ,Evolutio nan dInterference .MI TPress , Cambridge, Mass.,265pp . 407 Solbrig,O.T. , 1968.Fertility , sterilityan dth e speciesproblem . In:V.H .Heywoo d (ed.) Modern methods inplan t taxonomy, 77-96.Academi c Press,London . 408 Spach,E. ,1839 .Histoir enaturell ede svegetaux ,vol .7 .Le sClematidees ,257-284 .Librairi e encyclopedique deRoret ,Paris . 409 Spingarn,J.E.,1935 .Th elarge-flowere dClemati shybrids .A tentativ echeck-list .Th eNationa l Horticultural Magazine 1935:64-91 . 410 Spome,K.R., 1977.Som eproblem sassociate dwit hcharacte rcorrelations .Plan t Systematics andEvolution , Suppl. 1: 33-51. 411 Sporne,K.R., 1980.A re-investigatio no fcharacte rcorrelation samon gdicotyledons .Ne w Phytology 85:419-449 . 412 Sprague,T.A.,1933 .Th edate so fpublicatio no fRoyle' sillustrations .Ke wBulleti n1933 : 378- 390. 413 Stace,C.A., 1986.Th epresen tan dfutur einfraspecifi c classificationofwil d plants.In :B.T . Styles(ed. )Infraspecifi c classificationo fWil dan dcultivate dPlants ,9-20 .Clarendo nPress , Oxford. 414 Stace,C.A. , 1989.Plan tTaxonom yan dBiosystematics ,secon dedition .Edwar d Arnold, London, 264pp. 277 415 Stafleu, F.A., 1971. Linnaeus and the Linnaeans. Oosthoek, Utrecht, 386pp.(Regnu m Vegetabile79) . 416 Stafleu, F.A., &R.S .Cowan , 1981.Taxonomi cliterature ,vol .3 ,secon dedition . Bohn, Scheltema& Holkema ,Utrecht ; W.Jun k Publ., The Hague.(Regnu m Vegetabile 105). 417 Stanley, S., 1986. Earth andLif e through Time.W.H .Freeman , New York,690pp . 418 Steam,W.T. ,1943 .Royle' sillustration so fth ebotan yofth eHimalaya nMountains .Journa lo f theArnol d Arboretum 24:484-487 . 419 Steam,W.T. ,1953 .Internationa lcod eo fNomenclatur efo rCultivate dPlants .RHS ,London , 29pp. 420 Steam,W.T. ,1957 .A nintroductio nt oth eSpecie splantaru man dcognat ebotanica l works ofCar lLinnaeus .In :C .Linnaeu s- Specie splantarum .A facsimil eofth efirst editio n 1753.Ra y Society, London. 421Steam ,W.T. ,1986 .Historica lsurve yo fth enamin go fcultivate dplants .Act a Horticulturae 182:19-28 . 422 Stebbins,G.L. ,1950 .Variatio nan devolutio ni nplants .Columbi aUniversit yPress ,Ne wYork , 643pp. 423 Stebbins, G.L., 1974.Flowerin g plants. BelknapPress .Cambridge ,Mass. ,399pp . 424 Sterckx,R., 1897.Contributio na l'anatomi e desRenonculacees.TribudesClematidees . Archives Institute Botanique Liege 2: 3-117. 425 Steudel,E.T. , 1821.Nomenclato rbotanicu senumeran sordin ealphabetic onomin aatqu e synonyma planta imposita. Stuttgart, 904pp. 426 Stevens,P.F.,1991.Characte rstates ,morphologica lvariation ,an dphylogenetica lanalysis :A review. Systematic Botany 16:553-583 . 427 Stuessy,T.F.,1990 .Plan ttaxonomy ,th esystemati cevaluatio no fcomparativ edata .Columbi a University Press,Ne w York, 514pp. 428Takhtajan ,A.L. ,1969 .Flowerin gplants ,origi nan ddispersal .Olive rBoyd ,Edinburgh ,310pp . 429Takhtajan , A.L., 1980.Outlin eofth e classification offlowerin g plants (Magnoliophyta). Botanical Review 46:225-359 . 430Takhtajan ,A.L. ,1991 .Evolutionar ytrend si nflowerin gplants .Columbi aUniversit yPress , 278 New York, 241pp. 431Tamura ,M. ,1958 .Ranunculacea eo fNorth-Wester nNepa lcollecte db yK .Nishiok aan dO . Namikawai n 1958.Act a Phytotaxonomicae tGeobotanic a 23: 11-13. 432Tamura ,M. ,1962 .Taxonomica lan dphylogenetica lconsideratio no fth eRanunculaceae .Act a Phytotaxonomica et Geobotanica 20: 71-81. 433Tamura ,M , 1962. Morphology,ecolog yan dphylogen yo fth eRanunculacea eI .Annua l reports of the Collegeo f General Education OsakaUniversit y 1962,115-126. 434 Tamura,M. , 1963.Morphology ,ecolog yan dphylogen yo fth eRanunculacea eII . Annual reports of the College of General Education Osaka University 1963,141-156. 435Tamura ,M. , 1964.Morphology ,ecolog yan dphylogen yo fth eRanunculacea eHI .Annua l reports of theColleg e of General Education OsakaUniversit y 1964,25-38 . 436Tamura ,M. ,1965 .Morphology ,ecolog yan dphylogen yo fth eRanunculacea eIV .Annua l reports of the College of General Education OsakaUniversit y 1965,53-71 . 437Tamura , M., 1966. Clematischrysantha var . monantha and Clematischrysantha var. paucidentata. InKitamur a(ed. )Addend a etCorrigend a FloraAfghanistanic a 92:92 . 438Tamura ,M. ,1967 .Morphology ,ecolog yan dphylogen yofth eRanunculacea eVI .Annua l reports of the Collegeo f GeneralEducatio n OsakaUniversit y 1967,13-35 . 439Tamura ,M. ,1968a .Morphology ,ecolog yan dphylogen yo fth eRanunculacea eVII .Annua l reports of theColleg e of General Education OsakaUniversit y 1968,21-43 . 440 Tamura, M., 1968b. Clematis chrysantha var. brevipes. Acta Phytotaxonomica et Geobotanica 23: 30. 441Tamura ,M. ,1969 .Morphology ,ecolog yan dphylogen yofth eRanunculacea eVIE .Annua l reports of the College of General Education OsakaUniversit y 1969,41-56. 442Tamura ,M. ,1970. Archiclematis, aprecursor ygenu so fClematis. Act a Phytotaxonomica etGeobotanic a 24: 146-152. 443Tamura ,M. ,1987 .A classificatio no fgenu sClematis. Act aPhytotaxonomic ae tGeobotanic a 38: 33-44. 444Tamura ,M. ,& Y .Mizumoto ,1972 .Stage so fembry odevelopmen ti nrip eseed so rachene s ofthe Ranunculaceae. Journal of Japanese Botany 47:225-237 . 279 445Tamura , M., & Y. Mizumoto, 1974. The cotyledon and growing point inth e Monocotyledonousembryo so fShibateranthispinnatifida an dAnemone flaccida. Journa l of Japanese Botany49 : 123-128. 446Tamura ,M. ,& S .Vogel ,1993 .Ranunculaceae .In : K.Kubitzky ,J.G .Rohwer &V .Bittric h (eds.) TheFamilie s and Genera of Vascular Plants.,563-583 .Springe r Verlag, Berlin. 447Tarling ,D.H. ,1982 .Lan dbridge san dplat etectonics .Memoi rSpecialeGeobio s6 :361-374 . 448Tateoka , T., 1960.Cytolog yi n grass systematics: acritica l review.Th eNucleu s 3:81-110 . 449Tattersall ,I. ,&N .Eldredge ,1977 .Fact ,theor yan dphantas yi nhuma npaleontology .Amer . Sci.65 :204-211 . 450Templeton ,A.R. , 1989.Th emeanin go fspecie san dspeciation :A geneti cperspective .In : D. Otte andJ.A . Endler (eds.) Speciation and its Consequences, 3-27. Sinauer Associates, Sunderland, MA. 451Tepfer ,S.S. ,1960 .Th eshoo tape xan dearl ylea fdevelopmen ti nClematis. America nJourna l of Botany 47:655-664 . 452Theophrastes , 320BC.Histori a Naturalis. 453Thorne , R.F., 1978.Plat e tectonics and angiosperm distribution. Notes from the Royal Botanical Garden, Edinburgh 36: 297-315. 454Thorne ,R.F. ,1983 .Propose dne wrealignment si nth eangiosperms .Nordi cJourna lo fBotan y 3:85-117. 455 Tobe,H. ,1974 .Morphologica lstudie so nth egenu sClematis Linn .I .Polle ngrains .Scienc e Reports of theTohok u University, fourth series,Biolog y 37: 47-53. 456Tobe ,H. ,1976a .Morphologica lstudie so nth egenu sClematis Linn .II .Note so nbrac t and floral axisi n sectionParatragene Tamura. 457Tobe ,H. ,1976b .Morphologica lstudie so nth egenu sClematis Linn .UJ .Flora lanatom yo f Clematis tosaensis Makino. 458Tobe ,H. ,1979 .Morphologica lstudie so nth egenu sClematis Lirm. IV .Vascula rAnatom y ofth eInflorescenc eAxis ,wit hsom eConsideratio no fth eEvolutio no fth eFlora lShoo twit h SimpleAxillar y Inflorescences. 459Tobe ,H. ,1980a .Morphologica lstudie so nth egenu sClematis Linn .V .Vascula r Anatomy of theCaly x Region inFour-Sepale dFlowers . 280 460Tobe ,H. ,1980b .Morphologica lstudie so nth egenu sClematis Linn .VI .Vascula rAnatom y ofth eAndroecia lan dGynoecia lRegion so fth eFlora l Recpetacle. 461Tobe ,H. ,1980c .Morphologica lstudie so nth egenu sClematis Linn .VII .Reinvestigatio no f Clematis WilliamsiiA .Gra yan dProposa lo f ItsTaxonomi cTransfe r to Clematopsis. 462Tobe , H., 1980d. Morphological studies on the genus ClematisLinn . VITI.Flora l and Inflorescence Anatomyi n Clematispatens withEight-Sepale dFlowers . 463Tomlinson ,P.B. ,1984 .Vegetativ emorpholog y- Som eenigma si nrelatio nt oplan tsystematics . In:V.H .Heywoo d andD.M .Moor e(eds. )Curren t conceptsi nPlan tTaxonomy ,49-66 . Academic Press,London . 464Tournefort , J.P.de , 1700.Institutione s reiherbariae .Paris . 465Tournefort , J.P.de , 1703.Corollariu minstitutione m reiherbariae .Paris . 466Tournefort , J.P.de , 1717.Relatio n d"unvoyag ed uLevant .Paris . 467Trapl , S., 1912. Morphologische Studien iiber den Bau und das Diagrammde r Ranunculaceeenblute. Beihefte zumBotanische m Centralblatt 28: 247-281. 468Trautvetter , E.C. von, 1841.D e novo systemate botanico: brevem notitiam. Annales (botanique) 1841(3):509-528 . 469Troll , W., 1964.Di eInfloreszenzen , Tl.I .VE B Gustav Fischer, Jena,615pp . 470Troll ,W. , 1969.Di e Infloreszenzen, T.H , 1.VE BGusta vFischer , Jena,630pp . 471Tromp , J., 1986. Boskoops Koninklijke (1861-1986). Koninklijke Vereniging voor Boskoopse Culturen, Boskoop,226pp . 472Ulbricht , 1922.Clematis chrysantha. FeddesRepertoriu m Beihefte 12:374 . 473UPOV ,19S4. Draft: Triticale distinguished from rye, soft wheat and hard wheat (French ) 1984: 11-20. 474Usher ,G. ,1974 .A dictionar yo fplant suse db yman .Clematis, 158-159.Constable ,London . 475Valen ,L.M .van , 1988.Species ,set s andth ederivativ e natureo f pilosophy.Biologica l Philosophy 3:49-66 . 476Valentine ,D.H. ,1975 .Th etaxonomi ctreatmen to fpolymorphi cvariation .Watsoni a10 : 385- 390. 281 477 Veitch,J. , 1912.Hard yplant s ofWes t China,9 . 478Vries ,I.M .de ,an dC.E .Jarvis . 1987.Typificatio n ofseve nLinnaea nname si nth egenu s Lactuca L. (Compositae:Lactuceae) . Taxon 36: 142-154. 479Wagner , G.P., 1989.Th eorigi n ofmorphologica l characters andth ebiologica lbasi so f homology. Evolution 43: 1157-1171. 480Wagner , W.H.Jr. , 1970.Biosystematic s and evolutionary noise.Taxo n 19: 146-151. 481Wagner , W.H., 1980. Origin and philosophy of the groundplan-divergence methodo f cladistics. Systematic Botany 5: 173-193. 482Wagner ,W.H. ,1983 .Reticulistics :th erecognitio no fhybrid san dthei rrol ei ncladistic san d classification.In :N.I .Platnic kan dV.A .Fun k(eds. )Advance si nCladistics ,63-79 .Columbi a University Press,Ne w York. 483Wagner ,W.H. ,1984 .A compariso no ftaxonomi cmethod si nbiosystematics .In :W.F .Gran t (ed.)Plan tbiosystematics , 643-654.Academi c Press,Do nMills ,Canada . 484Wallace ,A.R. , 1855.O nth ela wwhic hha sregulate dth eintroductio no fne wspecies .Ann . Mag.Natur . Hist. 16,2 : 184-196. 485Walper , G.G., 1848.Annale s Botanices Systematicae Vol. 1:3-4 .Lipsiae . 486Wanntorp ,H.-E. ,1983 .Reticulate dcladogram san dth eidentificatio no fhybri dtaxa .In : N.I. Platnickan dV.A .Fun k(eds. )Advance si nCladistics ,8 1-88 .Columbi aUniversit yPress ,Ne w York. 487 Weberling.F., 1981.MorpholgiederBliitenunderBlutenstande.EugenUlmer, Stuttgart, 391pp. 488Wenzel ,W. ,& V .Hemleben ,1982 .Acomparativ estud yo fgenome si nangiosperms .Plan t systematics and Evolution 139:209-227 . 489Werth,E. ,1941 .Di eBlutennektarienderRanunculaceenundihrephylogenetisch ebedeutung . Berichte derDeutsche n Botanischen Gesellschaft 59:246-256 . 490Wet , J.M.J, de, 1981. Species concepts and systematics of domesticated cereals. Kulturpflanze 29: 177-198. 491Wheeler ,Q.D. ,& K.C .Nixon ,1990 .Anothe rwa yo flooking at th especie sproblem :a repl y to deQueiro s and Donoghue. Cladistics 6: 77-81. 282 492White ,M.J.D. , 1978.Mode so fspeciation .W.H .Freema nan dCo. ,Sa nFrancisco ,455pp . 493Wijnands , D.O., 1983. The botanyo f theCommelins .Balkema ,Rotterdam , 232pp. 494Wijnands ,D.O. ,1986a .Linnaeus' sattitud etoward scultivate dplants .Act aHorticultura e182 : 67-77. 495Wijnands , D.O., 1986b.Forma , Phoenix of bedrieger?Ericultur a 16(60):21-26 . 496Wijnheijmer ,E.H.M. ,W.A .Brandenburg ,& C.E .Jarvis ,1988 .Lectotypificatio no fDaueus carota L. (Umbelliferae). Taxon 37 (1): 175-184. 497Wiley ,E.O. ,1975 .Kar lR .Popper ,systematics ,an dclassification-arepl yt oWalte rBoc k and other evolutionary taxonomists. Systematic Zoology 24:233-243 . 498Wiley ,E.O. , 1977.Ar emonotypi cgener aparaphyletic ?- a respons et oNorma n Platnick. Systematic Zoology 26:352-355 . 499Wiley ,E.O. ,1978 .Th eevolutionar yspecie sconcep treconsidered .Systemati cZoolog y27 : 17-26. 500Wiley ,E.O. , 1979a.Cladogram s andphylogeneti c trees.Systemati c Zoology 28:88-92 . 501Wiley,E.O. ,1979b .Ancestors ,specie san dcladograms.-Remark s onth esymposium . In: Cracraft andEldredge(eds. )Phylogeneti canalysi san dpaleontology ,211-225 .Columbi a University Press,Ne w York. 502Wiley ,E.O. , 1979c.A nannotate dLinnea nhierarchy ,wit hcomment so nnatura ltax aan d competing systems.Systemati c Zoology 28:308-337 . 503Wiley.E.O. ,1980 .Phylogeneti csystematic san dvicariancebiogeography .Systemati cBotan y 5: 194-220. 504Wiley ,E.O. ,1981 .Phylogenetics :th etheor yan dpractic eo fphylogeneti csystematics .Joh n Wiley and Sons,Ne w York. 505Willdenow , 1796.Clematis glauca. Berliner Baumzucht 65: 89-94,t.4 ,fi g 1. 506Wittstein ,G.C. ,1856 .Etymologisch-botanische sHandworterbuch ,2nd .ed .Palmun dEnke , Erlangen, 952pp. 507Wodehouse ,R.P. ,1936 .Polle ngrain si nth eidentificatio nan dclassificatio no fplants .VII .Th e Ranunculaceae. Bulletin of theTorre y Botanical Club 63:495-514 . 283 508Wulff ,E .V. ,1950 .A nintroductio nt ohistorica lplan tbiogeography .Chronic aBotanic a10 , 233pp (translation of the 1932Russia n text). 509Zandee ,M. , &M.C .Roos , 1987.Component-compatibilit y inhistorica l biogeography. Cladistics 3:305-332 . 510Zeylinga ,A.E. ,& G.H .Kroon , 1965.A metho dfo rmakin groot-ti psquashe s permanent without removal of thecove r slip.Euphytic a 14:36-38 . 284 INDEX TO CLEMATIS SPECIES IN CHAPTERS 1-5. Pagenumber si nbold=description ;italics=illustration ;name sbetwee nbracket sar ereduce d tosynonyms . {Archiclematis alternata) 46 Clematis afoliata 9 (Clematis akebioides)106 , 117, 125, 128, 165,16 9 (Clematis albidd) 139 Clematis alpina7 , 55,59 ,60 ,65 ,68 ,82 ,85 ,94 , 169 Clematis alternata23 8 Clematis apiifolia18 8 Clematis 'Aureolin' 221 Clematis'Bil l McKenzie' 222 Clematis'Bravo ' 191,192,193,223-224 Clematis'Burford ' 225 Clematis'Corry ' 226 Clematis campaniflora 55,59 ,60 ,65 , 77,89 ,91, 21 5 Clematis chrysantha 151,169 ,17 5 Clematis chrysocoma 70, 71, 77 Clematis cirrhosa 5,6 ,7 , 8,55 ,59 ,60 ,61 ,65 ,6 8 Clematis columbiana 85 Clematis crispa5, 6 ,7 (Clematis daurica)13 9 Clematis'Drake' s Form' 227 (Clematis eriopoda) 169 Clematis'Eriostemon ' 62 Clematisflammula 6,7 ,8 7 (Clematisflava) 8 , 139 Clematisflorida 8 ,41 ,94 ,21 6 Clematisfusca 40, 188,21 5 (Clematis glauca) 9,55 ,99,106,117,125,139, 143,151, 169,218 ,22 0 (Clematis globosa)13 9 Clematis'Golde n Harvest' 226 Clematis grahami21 5 Clematis grata21 5 Clematis graveolens 104, 106, 116, 117,118,119, 120,121,122, 125, 128, 129,147- 150, 215,21 9 Clematis'Helios ' 226 Clematis heracleifolia 55,59 ,60 ,65 ,68 ,83 , 84 (Clematis hilariae) 106, 117, 125, 128, 151 Clematis integrifolia 6,7 , 8,40 ,55 ,59 ,60 ,61 ,65 ,68 , 77,82 ,83 ,84, 8 5 Clematis intricata 99, 101,104, 120,121, 128, 129,151-155, 160,218 ,219 ,220 ,23 0 Clematis ispahanica 70,71,72,106,116,117,118,119,120,121,125,128,129,156- 159, 188 285 Clematisx jackmanii 199,21 7 Clematis'Jackmanii ' 199,21 7 Clematisx jouiniana 6 1 Clematis koreana16 1 {Clematis ladakhiana) 165 Clematis'Lambton' s Park' 228 Clematis lanuginosa 199,216 ,24 0 Clematis'Lanuginosa ' 240 {Clematis longecaudata) 139 Clematis macropetala 55 Clematis mandshurica 186 Clematis montana 55,59 ,60 ,61 ,65 ,68 ,70 ,71 , 73,77, 82 ,9 4 Clematis'Mrs .Rober t Brydon' 61 Clematis ochroleuca 40 Clematis'Orang e Peel' 226,228-22 9 Clematis orientalis 7,8,9,11,12 , 55,59,60 , 61, 65,68 ,69 ,70 ,72 ,77, 86 ,96, 99,100 , 103, 104, 106, 117,120,121, 125, 128, 129,130-146, 147, 151,160 , 165,169 , 175, 182, 183,184,185, 186, 188, 191,192, 193,216 ,217 ,218 ,220 ,23 0 {Clematispamiralaica) 105,16 9 Clematispaniculata 186 {Clematisparvifolid) 14 7 Clematispatens 41, 55,59 ,60 ,61 ,65 ,68 ,83 , 188,190,21 6 Clematispitcheri40 ,59 ,60 ,65 ,6 8 {Clematispseudoorientalis) 156 Clematis recta5 ,6 ,7 ,55 ,59 ,60 ,65, 68 ,23 9 {Clematis sarezica) 151 {Clematis serrata)16 0 Clematis serratifolia 55,59 ,60 ,65 ,68 ,69 ,70 , 105, 106, 117, 720,121, 125, 128,129 , 160-163, 188, 191,192, 193,219 ,227 ,23 0 Clematis stans18 6 {Clematis tangutica) 55,59 ,60 ,61 ,62 ,68 ,69 ,70 , 105, 106, 117, 125, 169, 175, 219 {Clematis tenuifolia) 139 Clematis texensis82 , 186,21 6 Clematis tibetana105 , 106, 116, 117,120,121,122, 125, 128,164-178, 230 Clematistibetana subsp.tangutica 77,101,125,128,165, 169-174,176,178,182,183, 184,185, 188,221 ,222 ,225 ,226 ,227 ,23 0 Clematis tibetana subsp.tibetana 125, 128, 129,165-168, 775 Clematis tibetanasubsp . vernayi77,103,125, 128, 129,175-177, 778, 182, 183,184, 185, 188, 790,191,192,193,219 ,223 ,226 ,22 8 {Clematis vernayi) 59,60 ,65, 68 ,69 ,70 ,71 , 106, 117, 125, 175,21 9 Clematis viorna6 ,7 , 8, 186,21 6 Clematis virginiana 216 Clematis vitalba5 ,6 ,7 , 8, 11, 55,59 ,60 ,61 ,65 ,68 ,82 ,216 ,24 0 Clematis viticella 5,6 ,7 , 8,41 , 55,59 ,60 , 61,65 ,68 , 77,82 ,96 , 100,21 6 286 Clematis viticella'Hendersonii'19 9 Clematis'Walsall ' 230 (Clematis wilfordi)16 0 (Clematis zeylanica) 92 Naraveliazeylanica 9 287