Exochorda: Fivespecie So Rone ? Cip-Datakoninklijk Ebibliotheek ,De Nhaa G
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EXOCHORDA: FIVESPECIE SO RONE ? CIP-DATAKONINKLIJK EBIBLIOTHEEK ,DE NHAA G GaoFangyo u (F.Y.Gao ) Exochorda: five species or one? A biosystematic study of the Rosaceous genus Exochorda. Thesis,Wageninge n Agricultural University In English, with summaries in Dutch and Chinese. With references. 111.: W. Wessel- brand, author. ISBN 90-5485-825-7 Key words: Anatomy, biosystematics, cytology, distribution, embryology, Exochorda, morphology, multivariate analysis, pollination biology, populations, Rosaceae, taxonomy. Cover: author. Exochorda racemosa (Lindl.)Rehde rsubsp .racemosa, population 01 in QixiaSha n Mountain,Nanjing ,Jiangs u province,China . Printed by:Cop y Master, Netherlands B.V. Copyright © 1998b yWageninge n Agricultural University,th eNetherlands . II ,JNOF2OI,23<KS GaoFangyo u Exochorda:five specie so rone ? Abiosystemati c study of theRosaceou s genus Exochorda Proefschrift ter verkrijging vand egraa dva ndocto r opgeza gva nd erecto r magnificus vand eLandbouwuniversitei tWageninge n dr.C.M .Karsse n inhe t openbaart everdedige n op2 8januar i 1998 desnamiddag st e vieruu ri nd eAul a ra ^v) ^S^1°l Promotor: Prof.Dr . ir.L.J.G . van derMaese n Hoogleraar ind e plantentaxonomie BIBLIOTHEEK IV LANDBOUWUNIVERSITEIT WAGENINGEN pjy^zo», 23£ 5 Propositions 1. Exochorda populations aremorphologicall y very similar, and appear tobelon g toth e same species,althoug h theirdistribution s areclearl y disjunct. this thesis 2. Thepheneti c study doesno t supportfive distinc t taxa(i.e .five species ) inth e Exochordacomplex .Instead ,thre ebroadl ydefine d subspecific taxaca nb e recognized.Thes ecoincid e inpar t withth e geographical distribution inth egenus . thisthesi s (page: 83) 3. Exochorda originated before theemergenc e of theTaklamaka n Desert. Itprobabl y onceoccupie d alarg eare a inth enorther n hemisphere of Asia.Th e disjunct distribution pattern represents therelic t of aforme r widean d continuous distribution pattern, the intervening areas havingbee n depopulated. thisthesi s (page: 106) 4. Thegenu sExochorda hasbee n neglected for quite alon gtim eespeciall y in China. Itsmerit s shouldbecom e more and more appreciated for gardening. author Chinai son eo f therichest region s of plant diversity contributing manyimportan t crops,particularl y fruit trees.Othe r important crops areBrassica campestris and related species,Camellia sinensis, Colocasia esculenta, Corchorussinensis, Glycine max,Panicum milaceum, Raphanus sativus, Setariaitalica etc . A.C.Zeve n &P.M .Zhukovsky . 1975.Dictionar yo f cultivated plantsan dthei rcentre so fdiversity :27 . 6. Knowledge andexperienc ed ono tnecessaril y speakth esam elanguage . Taoistphilosoph y 7. Education isa lifelon g process,a proces stha tstart s longbefor e theforma l starto f schooleducation , andon etha t should bea nintegra lpar to fone' sentir e life. 8. Learning without thinkingi suseless ;thinkin g without learningi sperilous . Confucius (551-479 B.C.) 9.Fait h isth ebir d thatfeel s theligh twhe nth edaw n isstil l dark. Rabindranath Tagore (1816-1941) 10.Wha teveryon e wants from life iscontinuou s andgenuin e happiness.Happines s is therationa l understanding of life andth e world. Baruch Spinoza (1631-1677) Stellingen, horend bij hetproefschrif t van GaoFangyou : Exochorda: five specieso rone ? Abiosystemati c study ofth eRosaceou sgenu s Exochorda Wageningen, 28januar i 1997 Naturebring sm esunshin eanckmusi c inm ylife . Author ToProf .Dr .L.J.G . van derMaese n Tom yparent s Tom yhusban d Contents Page Preface X Summaries (inEnglish ,Dutc h and Chinese) XI 1.Introductio n and history of thegenu sExochorda 1 2.Lea f morphology, anatomy andseedlin g morphology 6 2.1. Introduction 6 2.2.Material s andmethod s 6 2.3. Results 7 2.3.1.Lea f morphology 7 2.3.2.Epidermi s morphology andstomata l type 8 2.3.3.Transvers e section of thelea fblad e 8 2.3.4.Petiol eanatom y 9 2.3.5.Seedlin g morphology 9 2.4.Discussio n 9 3.Woo d anatomy 14 3.1. Introduction 14 3.2.Material s andmethod s 14 3.3.Result s 16 3.4.Discussio n andconclusion s 20 3.4.1. Speciesdelimitatio n andrelationshi p withothe rgener a in 20 Rosaceae 3.4.2.Ecologica l trendsi n wood ofExochorda inChin a 20 3.4.2.1. Mutual relationshipbetwee n macroclimate and 23 moistureavailabilit y 3.4.2.2.Woo d anatomical characters inrelationshi p to 23 ecology 4. Pollen morphology 28 4.1. Introduction 28 4.2.Material s andmethod s 28 4.2.1. LM 29 4.2.2. SEM 30 4.2.3. TEM 30 4.2.4.Measurement s 30 4.3. Results 30 4.3.1. General morphology 30 4.3.2. Apertures 31 4.3.3. P/Evalu ean d shapeo f grain 33 4.3.4. A/Evalu e andlengt h ofcolp i 33 4.3.5. Exinearchitectur e 34 VII 4.3.5.1. Stratification 34 4.3.5.2. Ornamentation 34 4.4. Discussion and conclusions 34 5. Cytotaxonomy 39 5.1. Introduction 39 5.2. Materials andmethod s 39 5.3. Results anddiscussio n 40 6.Embryolog y ofExochorda 43 6.1. Introduction 43 6.2.Material san dmethod s 43 6.3. Results 43 6.3.1. Pistil 43 6.3.2.Microsporangiu m and microsporogenesis 44 6.3.3.Megasporogenesi s and megagametogenesis 45 6.4. Discussion 45 7.Pollinatio n ofExochorda 51 7.1. Introduction 51 7.2. Materials and methods 52 7.2.1. Site 52 7.2.2. Plant species andflora l biology 52 7.2.3. Pollinators 53 7.2.4. Fruit set 54 7.3. Results 54 7.3.1. Plant-animal interaction 54 7.3.2.Pollinator s andthei r behaviour 56 7.3.3.Frui t set 60 7.4.Discussio n andconclusion s 60 8.Geneti c segregation ofrando mamplifie d polymorphic DNAi n 63 somegener ao f Rosaceae 8.1. Introduction 64 8.2. Materials andmethod s 64 8.2.1. Plantmaterial s 64 8.2.2.DN Aisolatio n 65 8.2.3.Primer s 66 8.2.4.PC R analysisan d electrophoresis 66 8.2.5. Dataanalysi s 66 8.3. Results anddiscussio n 67 9. Multivariate analysis ofmorphologica l variation inth eExochorda 70 complex 9.1. Introduction 70 9.2.Material s andmethod s 70 VIII 9.3. Results 73 9.3.1. Character analysis 73 9.3.2. Principal component analysis(PCA ) 79 9.3.3. Pheneticanalysi s 81 9.4. Discussion 83 10.Revisio n 85 10.1.Descriptio n of the genus 85 10.2. Description of the species 86 11.Geographica l distribution andevolutionar y trends 98 11.1. Introduction 98 11.2.Presen tgeographica l distribution of the genus 98 11.2.1.Th etype so f distribution 99 11.2.2.Centr eo f diversity 101 11.3.Tim ean dplac eo f origin 101 11.4.Dispersa l andevolutionar y trends 103 11.5.Discussio n andconclusion s 106 References 108 Acknowledgments 116 Curriculum vitae 119 Appendix 120 Indexo f plant names 132 IX Preface The present dissertation has three objectives: first, to solve the question: how many species exist in the Rosaceous genusExochorda: five o r one?;t o describe and analyze the kinds of adaptive radiations that could have given rise to morphological and anatomical differences to distinguish species in Exochorda; second: to compare Exochorda with other genera in Rosaceae, with respect to leaves, wood, pollen and chromosome, these data give us a record of taxonomie information; third: to explore the origin, dispersal and present geographical distribution of the genus and ecological factors thatinfluenc e diversification. Chapter 2 to 5provid e information for thetaxonom y of thegenus .I n chapter 6, the embryology of Exochorda istreated . Chapter 7 discusses thepollinatio n of thespecies . In chapter 8, molecular methods were used to attempt to detect differences between populations. In chapter 9, numerical taxonomy of Exochorda isperformed . In chapter 10, the taxonomie revision is given. Chapter 11present s the postulates of the origin, dispersal andpresen t geographical distribution of thegenus . GaoFangyo u Wageningen, theNetherland s December, 1997 X Summary The Rosaceous genus Exochorda consisted sofar of five accepted species and is distributed in East Asia and Central Asia. Morphologically the taxa in theExochorda complex have similar characters, hence the classification in the complex was questionable. A further problem is the relationship between the genus Exochordaan d other generai nRosaceae .I nthi sdissertation , based onwil d material of 22 populations in China, cultivated accessions in the botanical gardens of Wageningen Agricultural University, the Netherlands, and herbarium material, a biosystematic study of this genusi sreported .Th eresult s aresummarize d asfollows : Leaf anatomy: The epidermis consists of one layer of cells and the shape of epidermis cells is irregular in outline as seen on both surface. Epidermal cell shape in surface view isquit e uniform inExochorda. Stomata are always confined to the lower leaf surface. An anomocyticpatter n isfoun d inal lsamples . Wood anatomy: growth rings are distinct, boundaries are marked by differences in vessel diameter between earlywood and latewood, andb y rows of flattened late wood fibres. The wood is ring-porous to semi-porous. Quantitative anatomical characters express continuous variation within the "species", wood of Exochorda shows a single wood structural type similar totha to f thePrunoidea e than toth e Spiraeoideae. Pollen morphology and exine structure show little variation in this genus. Exochorda pollen grains are isopolar, three-colporate, small or medium-sized, grain shape isspheroidal .P = 20.9-30. 5 (19.6-36.9) urn,E = 22.1-30.7 (17.9-35.7) um, A= 5.2-9.5 (3.6-15.5) um, P/E = 0.88-1.36 (0.81-1.73). Ornamentation is striate.