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Systematics of the Compsopogonales (Rhodophyta) With SYSTEMATICS OF THE COMPSOPOGONALES (RHODOPHYTA) WITH EMPHASIS ON THE FRESHWATER FAMJLES IN NORTH AMERICA. A Thesis Presented to The Faculty of Graduate Studies of The University of Guelph by TARA RINTOUL In phal fulfilment of requirements for the degree of Master of Science August, 1998 0Tara Rintoul, 1998 National Library Bibliothèque nationale du Canada Acquisitions and Acquisitions et 8ibliographic Services services bibliographiques 395 Wellington Street 395, rue Wellingtori OttawaON KIAW Ottawa ON K1A ON4 Canacla Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence dowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or seil reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/film, 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 substmtial extracts fiom 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 The Compsopogonaies is supported as a valid taxonomic entity through phylogenetic analysis of gene sequence data (18s rRNA and rbcL genes). A well separated freshwater lineage was identified through these analyses and a close relationship was recognized between the Boldiaceae and the Cornpsopogonaceae. Biogeographical distribution of the Boldiaceae and Compsopogonaceae in North America was assessed through gene sequence data of the 5.8s rRNA gene and internally transcribed spacer regions ITS 1 and ITS2 of the rRNA gene. Analysis of the Boldiaceae indicated that introductions occurred fiom southem populations into northem areas. No obvious geographic trend was evident in the Compsopogonaceae. hdividuals in both families are thought to be dispersed through vector-assisted transport. The Compsopogonaceae is shown to be monotypic in North America, containhg the single species Compsopogon coenrletlr. Observations of developing monosporangia demonstrated that the fieshwater families share ultrastructurai characteristics with the marine Erythropeltidaceae, and that the method of monosporangia formation is a usefid taxonomic character to distinguish the order Compsopogonales. TABLE OF CONTENTS TABLE OF CONTENTS ACKNOWLEDGMENTS LIST OF TABLES LIST OF FIGURES AND PLATES CHAPTER 1 : INTRODUCTION AND LITERATURE REVIEW 1.1 Systematics of the Order Compsopogonales 1.2 Distribution and biology of the freshwater families Boldiaceae and Compsopogonaceae i) Boldiaceae ii) Compsopogonaceae 1.3 Taxonomy of the Cornpsopogonaceae 1.4 Ultrastructure of monospore production in the Compsopogonales CHAPTER 2: MATERIALS AND METHODS 2.1 DNA Extraction, Polymerase Chain Reaction and DNA Sequencing 2.2 DNA Sequence analysis 2.3 Microscopy i) Transmission electron microscopy ii) Scanning electron microscopy iii) Light microscopy CHAPTER 3: RESULTS 3.1 Phylogeny of the Compsopogonales i) Phylogeny of the Compsopogonales based on rbcL gene sequences. ii) 18s rRNA gene phylogeny of the Compsopogonales iii) Combined 18s rRNA and rbcL gene phylogeny of the Compsopogonales 3.2 Biogeography of the Boldiaceae 3.3 Biogeography of the Compsopogonaceae 3.4 Taxonomy of the Compsopogonaceae 3.5 Ultrastructure of monosporangia in the Boldiaceae and Compsopogonaceae i) Boldiaceae ü) Compsopogonaceae CHAPTER 4:DISCUSSION 4.1 Phylogeny of the Compsopogonales 4.2 Biogeography of the Boldiaceae and Compsopogonaceae in North Arnerica 4.3 Taxonomy of the Compsopogonaceae 4.4 Ultrastructure of monosporangia in the Boldiaceae and Compsopogonaceae SUMMARY LITElRATURE CITED ACKNOWEDGMENTS 1 thank Dr. Robert Sheath for the opportunity, the support and the ideas which were essential in the completion of this research. 1 thank members of my advisory codtteeDr. Joe Gerrath and Dr. Bnan Husband for insight and enthusiasm. 1 appreciate the training and energy that Morgan Vis-Chiasson shared with me at the beginning of my project. 1 am gratefbi to Angela Holliss for assistance in DNA sequence analysis and Joseline Beaulieu, P.G. Davison, Ray Holton and Bnan Oates for collecting and providing samples for this research. To the people 1 have shared space and experience with, Lesley Campbell, Kirsten Miiller and Alison Sherwood, 1 extend heartfelt thanks for times filled with ideas and mirth, and sympathetic support. Gord Lemon, my traveling cornpanion, translater and good fnend, shared my joumey through this process and 1 was glad to have him dong. 1 am also greatly indebted to the people who surround me in the world away fiom scientifïc endeavor, who maintain a delightful, enteriahhg and loving atmosphere for me to corne home to. 1 thank my family for ali of their support and belief in my abilities. And as the most kind, generous and understanding force in my life, I thank Robert Wouda for being with me and consta;ritly reminding me that I was capable of everything. LIST OF TABLES Table Title Pages number Collection and gene sequence information for simples 20-22 included in phylogenetic dysisof the order Compsopogonales. Collection information for samples included in North Amencan biogeographic shidy of the Boldiaceae and Compsopogonaceae. Primer sequences for phylogenetic and biogeographic analyses of the Compsopogonales. rbcL gene percent sequence divergence and Kimura 2 parameter distances of the Compsopogonales. 18s rWA gene percent sequence divergence and Kimura 2 parameter distances of the Compsopogonales. 5.8s rRNA gene percent sequence divergence and Kirnura 2 parameter distances of the Boldiaceae. ITS 1 percent sequence divergence and Kirnura 2 parameter distances of the Boldiaceae, ITS2 percent sequence divergence and Kimura 2 parameter distances of the Boldiaceae. 5.8s rRNA gene percent sequence divergence and Kimura 2 parameter distances of the Compsopogonaceae. 10 ITS 1 percent sequence divergence and Kirnura 2 parameter distances of the Compsopogonaceae. 11 ITS2 percent sequence divergence and Kimura 2 parameter distances of the Compsopogonaceae. 12 rbcL gene percent sequence divergence and Kimura 2 parameter distances of the Compsopogonaceae. 13 18s rRNA percent gene sequence divergence and Kimura 2 parameter distances of the Compsopogonaceae. 14 Summary of important characters reported for the 97-99 Compsopogonales. LIST OF FIGURES AND PLATES Figure Title Pages Number Distribution of Boldiaceae in North America as represented by reported collection sites. Distribution of Compsopogonaceae in North America as represented by reported collection sites. Parsimony analysis of rbcL gene sequence data of the Compsopogonaies and Bangiales. Neighbour joining distance tree of Kimura 2 Parameter distances of rbcL gene sequence data of the Compsopogonales and Bangiales. Parsimony analysis of 18s rRNA gene sequence data of the Compsopogonaies and Bangiales. Neighbour joining distance tree of Kimura 2 parameter distances fiom 18s rRNA gene sequence data of the Compsopogonales and Bangiales. Parsimony analysis of the combined 18s rRNA and rbcL gene sequence data of the Compsopogonales and Bangiales. Neighbour joinùig distance analysis of combined 18s rRNA and rbcL gene sequence data of the Compsopogonales and Bangides. Distribution of sequenced collections of Boldicl erythrosiphon fiom North Arnerica Parsimony analysis of combined sequence data of 5.8s rRNA 58-59 gene, ITS I and ITSZ of the Boldiaceae of North America Neighbour joining distance trees of 5.8s rRNA gene, ITS 1 and 60-6 1 ITS2 combined sequence data of the Boldiaceae of North America. Distribution of sequenced collections of Cornpsopogonaceae in 63-64 North America, Parsimony analysis of combined sequence data of 5.8s rRNA 69-70 gene, ITS 1 and ITS2 of the Compsopogonaceae in North America. Neighbour joining distance analysis of 5.8s rRNA gene, ITSl and 72-73 ITSZ combined sequence data of Compsopogonaceae in North America, Parsimony (A) and neighbour joining distance (B) anaiysis of 74-75 combined 5.8s rRNA gene, ITSl and ITS2 sequence data of the Compsopogonaceae in North America with the collections ALTR and SL2A removed. Parsimony analysis (A) and neighbour joining distance (B) 78-79 analysis of 5.8s rRNA gene, ITSl and ITSZ combined sequence data of Compsopogonaceae. Mature thalli of Boldia erythrosiphon Transverse sections of thalli of Boldia erythrosiphon Transverse sections of thalli and monosporangia of Boldia vii 20 Germlings, young plants and monosporaflgia of the 9 1-92 Compsopogonaceae. 21 Ce11 division and monosporangia of the Compsopogonaceae. 93-94 22 Gene tree of combined 18s rRNA and rbcL gene data of 100- Compsopogonales with morphological characters mapped onto 101 branches. CHAPTER 1: INTRODUC'MON AND LITERATURE REWW 1.1: Systematics of the Order Compsopogonales The order Compsopogonales currentiy consists of three families: the Compsopogonaceae, Boldiaceae and Erythropeltidaceae (Garbary et al. 198Oa). The first two families are typically fkshwater in their occurrence, though Compsopogon coeruleus has ken collected in brackish envhnments (Tomas et al. 1980).In contrast, the third family is composed of only marine representatives. The Compsopogonales was erected in 1939 by Skuja and included only the family Compsopogonaceae and the single genus Compsopogon
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