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NOTE TO USERS This reproduction is the best copy available. SYSTEMATKS AND BIOGEOGRAPHY OF THE RED ALGAL ORDER HLLDENBRANDIALES (RHODOPHYTA) A Thesis Presented to The Faculty of Graduate Studies of The University of Guelph by ALISON RUTH SHERWOOD In partial fulfihent of requirements for the degree of Doctor of Philosophy December, 2000 O Aiison R. Sherwood, 2000 National Library Bibliothèque nationale du Canada Acquisitions and Acquisitions et Bibliog raphic Services services bi bIiograp hiques 395 Wellington Street 395. rue Wellington Ottawa ON KIA ON4 Ottawa ON K1A ON4 Canada Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant à la National Lïbrary of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire' prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la foxme de microfiche/nlm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts from it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. ABSTRACT SYSTEMATICS AND BIOGEOGRAPHY OF THE RED ALGAL ORDER HILDENBRANDMES (RHODOPHYTA) Alison Ruth Shercvood Advisor: University of Guelph, 2000 Professor R-G, Sheath The genetic variability of the genus Hildenbrandia throughout its distributional range and the taxonomie implications of this variation were examined using a combination of DNA sequence analyses (rbcL and 18s rRNA genes, and ITS regions), othzr rno lecular marker techniques (ISSR analyses) and morpliometric analyses. The phylogenetic relationship of Hildenbrandia to the second genus of the Hildenbrandiales, Apophlaea, was also exarnined using these techniques. Analyses of North American Hildenbrandia demonstrated high sequence divergence values within and arnong marine and fieshwater forms (225.8% [rbcL]; ~9.7%[18S rRNA]); although these levels were comparable in marine European rnembers of the genus (524.9% [rbcL];55.8% [18S &NA]), they were much lower for fieshwater European members (r 1.9% [rbcL]; ~3.6% [18S &NA]). The rbcL gene was transitionally saturated within the North Amencan and global level comparisons, but the phylogenetic signal for the 18s rRNA gene was strong even among global collections. Biogeographic comparisons of marine and fkeshwater Hildenbrandia collections fiom a small geographical region (southem Sweden) revealed different relationships among the samples, but comparisons to other samples fiom Europe indiczted a close phylogenetic relatiooship among al1 European freshwater Hildenbrandia. Some trends revealed by morphometric analysis correlated with those yielded by phylogenetic analyses, especiaily at a global level. Except for the common marine species, rubra, samples with similar morphologies fonned clades in phylogenetic trees. Based on these analyses, as well as morphornetric examinations of the qpe specimens and global representatives, taxonomie revision is recommended to reduce the number of currently accepted marine Hildenbuandia taxa fiom 12 to seven, with no synonymies recommended for the fieshwater taxa Inclusion of Apophlaea in the analyses indicated that although Apophlaea is a monophy Ietic genus, Hildenbrundia is not. A rnicroscopic examination of the freshwater species, H. angolemis, and its unique form of asemal reproduction demonstrated that gemrnae develop within the algal thallus and are released from the thallus surface. Histochemical cornparisons between gemma and thallus cells revealed large amounts of starch in the gemmae, but not in thallus cells, Completing this degree would not have been possible without help and assistance fkom a nuniber of people and organizations. Most importantly, 1 would like to thank my advisor, Dr. Robert Sheath, for his guidance and support since my undergraduate days. 1 am gratefül for the multitude of research, travel and teaching opportunities that he has provided over the years. The other members of my advisory cornmittee (Dr. Joe Gerrath, Dr. Denis Lynn and Dr. Larry Peterson) have also helped in the design of my projects and provided advice in their areas of expertise. 1 have shared the lab with a number of people at various points during my degree, and I would like to thank them al1 for making the thne fun and interestùig: Lesley Campbell, Dana Couture, Laura Kline, Kirsten Müller, Tara Rintoul, Troina Shea and Stacey Thompson. Thanks aiso to my fiends in both the botany and zoology departments. 1have received a large amount of help with my thesis projects fiom other people. Many phycologists around the world have provided me with material for this thesis, and to them 1 owe a large "thank you" for giving me the opportunity to examine specimens with such a wide geographical scope. Troina Shea did rnuch of the PCR work for Chapter 4 and Stacey Thompson helped with the light microscopy sectioning for Chapter 5. Dr. Paul Silva (UC Berkeley Herbarium) has kindly sorted through my taxonomie and nomenclatural quenes and Dr. Ten Crease (Dept. of Zoology, Guelph) hm spent a good deal of time helping me with my phylogenetic analyses. Angela Holiiss (Guelph Molecular Supercentre) has sequenced innumerab2e templates for me over the past four years- Many thanks to Ron Deckert, who has been a wonderfil field assistant, source of conimon sense, general botany guru, listener and friend. Also to my parents, John and Glenda Shenvood, who have always been supportive of my academic endeavors. 1would like to acknowledge financial assistance fkom NSERC in the form of two post-graduate schola. awards. Research costs have been covered through NSERC grant OGP 0 183 503 to Dr. Robert Sheath, the University of Guelph and a PSA grant-in-aid of research. *.. 111 TABLE OF CONTENTS Page Acknowledgments i-ii Table of Contents z-vii List of Tables viii-ix List of Figures x-xiv Abbreviations xv-XVi Chapter 1: General Introduction 1-21 1.1. Morphology and distribution of the Hildenbrandides 1.2, Taxonomy of the Hildenbrandides 1-3. Phylogenetic positioning of the Hildenbrandiales 1.4. Biogeographic study of the Hiidenbrandiales 1S. The origins of fieshwater Hildenbrandia 1.6. Asexual propagation by gernmae in Hildenbrandia 1-7. Research objectives 1-8. Literature cited Chapter 2: Biogeography and systematics of Hildenbrandia in North America: inferences from morphometrics and sequence analysis of the rbcL and 18s rRNA genes 2.1. Introduction 2.2, Materials and Methods 2.2.1. Collection and identification of materials and DNA extraction 2.2.2. Morphometric analysis 2.2.3. Amplification and sequencing of the rbcL and 18s rRNA genes 2.2.4. rbcL and 185 rRNA gene analyses TABLE OF CONTENTS cont. 2.3. Results 2.3.1. Morphometric analysis 34 2.3 -2. Analysis of transitional saturation of the rbcL and 18s rRNA 39 genes 2.3 -3. rbcL gene distance, parsimony and quartet puzzIing analyses 47 2.3 -4. 18s rRNA gene distance and parsimony analyses 52 2.4. Discussion 57 2-5. Literature Cited 62 Chapter 3: Biogeography and systematics of Hildenbrandia in Europe: 67-101 inferences from morphometrics and sequence analysis of the rbcL and 18s rRNA genes 3.1. Introduction 3 -2. Materials and Methods 3.2.1. Sample collection and DNA extraction 3 -2.2. Morphometric analysis 3-2.3. Amplification and sequencing of the rbcL and 18s rRNA genes 3.2.4. rbcL and 18s rRNA gene sequence analyses 3 -3. Results 3 -3.1. Morphometric analyses 3 -3-2. Analysis of transitional saturation of the rbcL and 18S rRNA genes 3 -3-3. Parsirnony, distance and quartet puzzling analyses of the rbcL gene 3 -3-4. Parsimony, distance and quartet puzzling analyses of the 18s rRNA gene TABLE OF CONTENTS cont. 3 -4. Discussion 3 -5. Literature Cited Chapter 4: The relationship between marine and freshwater Hiïdenbrandia along an historical salinity gradient 4.1. Introduction 4.2. Materials and Methods 4.2.1. Collection of materials and morphologicd examination 4.2.2. DNA extraction, rbcL and 18s rRNA gene amplification, sequencing and gene sequence analyses 4.2.3. ITS 1 and ITS2 amplification and analyses 4.2.4. ISSR amplification and analyses 4.2.5. Determination of time since isolation f7om the Baltic Sea 4.3. Results 4.3.1. Morphological examination of collections 113 4.3 -2. Analyses of the ITS 1 and ITS2 regions 114 4.3 -3. ISSR-PCR results 221 4.3.4. rbcL and 18s rRNA analyses of representative collections 125 4-4. Discussion 131 4.5. Literature Cited 135 Chapter 5: Microscopic analysis and seasonality of gemma production 140-164 in Hilden brandia angolensis 5.1. Introduction 5.2. Materials and Methods 5.2.1. Collection of HiZdenbrundia angolensis 5.2.2. Microscopical and histochemical techniques 5.2.3. Seasonality of gemma production TABLE OF CONTlENTS cont. 5-3. Results 5-3.1. Gemma morphology and anatomy 144 5.3 -2. Histochemistry and X-ray microanalysis 145 5-3 -3. Gernma development 149 5 -3-4. Seasonality of gemma production 154 5.4. Discussion 157 5.5. Literature Cited 260 Chapter 6: Analysis of global collections and the type specimens of 165-140 Hildenbrandia 6.1. Introduction 6.2. Materiais and Methods 6.2.1. Type specimens, historicaliy signifTcant specimens and global collections analyzed 6.2.2. Morphometric analyses 6.2.3. rbcL and 18s rRNA gene sequence analyses 6.3. Results 6.3.1. Morphometric analyses 179 6.3-2. Analysis of transitional saturation of the rbcL and 18s rRNA 203 genes 6.3.3. 18s rRNA gene sequence analyses - parsimony analysis 203 6.3-4. 18 S rFWA gene sequence analyses - distance analy sis 209 6.3.5. 18s rRNA gene sequence analyses - quartet puzzling analysis 214 6.3.6.

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