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Ranunculaceae) , Inclusive of Cultonomic Aspects Promotor: Dr.Ir L.J.G Meclatisi n Clematis: Yellowflowering Clematis species Systematic studies in Clematis L.(Ranunculaceae) , inclusive of cultonomic aspects Promotor: dr.ir L.J.G. van der Maesen 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 other crosses 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 Isozyme polymorphism 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.
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