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Oral Morphometrics, of Homostyly from Distyly In ORAL MORPHOMETRICS,DEVELOPMENT AND EVOLUTION OF HOMOSTYLYFROM DISTYLYIN AMSINCKI. (BORAGINACEAE) Ping Li Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy Dalhousie University Halifax, Nova Scotia August 2001 O Copyright by Ping Li, 2001 National Library Bibliothèque nationale 1+1 ,canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395. rue WeUington OnawaON KlAOW OttawaON KlAON4 Canada Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loaq distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microfonn, vendre des copies de cette thèse sous paper or electronic formats. la foxme de microfichelfilm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright ir; this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fkom it Ni la thèse ni des extraits substantiels may be printed or othenvise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. .. Sigiiature page ...............................................................................................................u... Copyright agreement page ............................................................................................ru Table of contents ...........................................................................................................iv . List of figures ................................................................................................................vil List of tables ..................................................................................................................x .. Abstract .........................................................................................................................xi1... List of abbreviations and symbols ................................................................................xiil Acknowledgernents ....................................................................................................... xvi Chapter 1. Generd Introduction .................... ......................................................... 1 1.1. Introduction ....................................................................................................I 1.2. Research objectives ........................................................................................ 3 Chapter 2. Heterochrony in Plant Evolutionary Studies ................. ..................... 7 2.1. Abstract ................................. ... ..................................................................7 2.2. Lntroduction ......................................... ........................................................8 2.3. Heterochrony. Evolution and Development .................................................. 11 2.4. Types of Heterochrony ...................................................................................12 2.5. Problems and Solutions............................................................................ 17 2.5.1. Atomizing development .... ....................................................................17 2.5.2. Homology .............................................................................................18 2.5.3. Developmentd reference points ...................................... .. ................... 19 2.5.4. Absolute versus relative timing ............................................................ 19 2-55 Phylogenies ...........................................................................................20 2.6. Allometry. a Tool Complementary to Heterochronic Study ....................... ... 2 1 2.7. Applicability of Heterochrony to Plant Studies ............................................. 23 2.8. Heterochrony in Fossil Plants ........................................................................24 2.9. Heterochrony in Flowenng Plants ................................................................. 25 2.9.1. Heterochrony and timing of flowering ............................................... 25 2-92 Heterochrony and floral morphology ................................................... 28 2.10. Heterochrony at the Cellular and Tissue Levels ....................................... 34 2.1 1. Heterochrony at the Molecular Level ........................................................ 35 2.12. Homeosis ...................................................................................................... 39 2.13. Heterotopy .................................................................................................... 42 2.14. Conclusions ..................... .... ................................................................... 44 Chapter 3. Study Species ...................................................................................... 47 3.1 Introduction and Species .....-.............. ... ................................................... 47 3.2 Inflorescence and Fiower Morphology ..........................................................47 3.3 Mating Systems of Amsinckia .................... .... ............ .... 56 3.4 Phylogeny of Amsinckia ................ .............................. .... 57 Chapter 4. Comparative Floral Morphometria of Distyly and Homostyly in Three Evolutionary Lineages of Amsinckio (Borenaceae) .......... 60 4.1, Abstract ...*,.*......... ..*........................................................*..............................60 4.2. Introduction ........................................................................................ 6 1 4.3. Matenal and Methods .................................................................................... 63 4.3.1. Species and floral morphs ..................................................................... 63 4.3.2. Measurements .......................................................................................65 4.3.3. StaUstical andysis .................................................................................69 4.4. Results ............................................................................................................70 4.4.1. The size of floral traits in different lineages and morphs ...................... 70 4.4.2. A comparison of pin vs. thrurn .............................................................79 4.4.3. A comparison of distyly vs. homostyly ................................................ 86 4.4.4. A comparison of pin vs. thrum vs. homostyle ......................................87 4.5. Discussion ................ ,. ....................................................................................88 4.5.1. Distyly vs . homostyly ...........................................................................88 4.5.2. Evolution of homostyly ......................................................................... 91 4.5.3. Pinvs. thrum ......................................................................................... 92 4.5.3.1. Relative reciprocity ratios ..................... .. ...................... ............... 93 4.5.3.2. Papillae. style and stigma ............................................................. 98 4.5.3.3. Anther and filament ................... ... ......................................... 100 4.5.3.4. Pollen ..................... ................................................................... 101 4.5.3.5. Differences among lineages ......................................................... 102 Chapter 5. Discriminant Analyses of Distyly and Homostyly in Three Evolutionary Lineages of Amsinck .................... .......................... 105 5.1. Absmct ......................... ,. ............................................................................ 105 5.2. Introduction .................................................................................................... 106 5.3. Materials and Methods ...................................................................................109 5.3.1. Species and fioral morph types ............................................................ 109 5.3.2. Measurements ................................................................................... 111 5.3.3. Statistical analysis ................................................................................. 111 5.4. Results ............................................................................................................ 116 5.4.1. The most important traits differentiating distyly and homostyly .......... 116 5.4.2. The most important traits differentiating pin. thmm and homostyle .... 117 5.4.3. The most important traits differentiating pin. thrum. large homostyle and small homostyle in A . spectabilis .........*...............*....... 130 5.4.4. The most important traits differentiating three evolutionary lineages .. 137 5.5. Discussion ......................................................................................................144 5.5.1. Differentiating distyly from homostyly ................................................ 144 5.5.2. Differentiating pin. thmm and homostyle ............... .. ............................ 148 5.5.3. Differentiating pin, thmm, large homostyle. and small homostyle in A . specrabilis
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