The Foliose Bangiales (Rhodophyta)
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University of New Hampshire University of New Hampshire Scholars' Repository Doctoral Dissertations Student Scholarship Winter 2014 The foliose Bangiales (Rhodophyta) in the northern part of the North Atlantic and the relationship with the North Pacific foliose Bangiales - diversity, distribution, phylogeny and phylogeography Agnes Mols-Mortensen University of New Hampshire, Durham Follow this and additional works at: https://scholars.unh.edu/dissertation Recommended Citation Mols-Mortensen, Agnes, "The foliose Bangiales (Rhodophyta) in the northern part of the North Atlantic and the relationship with the North Pacific foliose Bangiales - diversity, distribution, phylogeny and phylogeography" (2014). Doctoral Dissertations. 2163. https://scholars.unh.edu/dissertation/2163 This Dissertation is brought to you for free and open access by the Student Scholarship at University of New Hampshire Scholars' Repository. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of University of New Hampshire Scholars' Repository. For more information, please contact [email protected]. THE FOLIOSE BANGIALES (RHODOPHYTA) IN THE NORTHERN PART OF THE NORTH ATLANTIC AND THE RELATIONSHIP WITH THE NORTH PACIFIC FOLIOSE BANGIALES – DIVERSITY, DISTRIBUTION, PHYLOGENY AND PHYLOGEOGRAPHY BY AGNES MOLS–MORTENSEN BSc, University of the Faroe Islands, 2002 MSc, University of Copenhagen, 2007 DISSERTATION Submitted to the University of New Hampshire in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Plant Biology December, 2014 This dissertation has been examined and approved in partial fulfillment of the requirements for the degree of PhD in Plant Biology by: Dissertation Director, Dr. Christopher D. Neefus, Professor of Plant Biology and Biometrics. Prof. Juliet Brodie, Research Phycologist. Dr. Arthur C. Mathieson, Professor of Botany. Dr. Anita Klein, Associate Professor of Genetics. Dr. Janet Sullivan, Affiliate Associate Professor of Botany. On 12th of September 2014 Original approval signatures are on file with the University of New Hampshire Graduate School. ACKNOWLEDGEMENTS I would like to express my appreciation to my advisors and committee, Dr. Chris Neefus, Prof. Juliet Brodie, Dr. Arthur Mathieson, Dr. Janet Sullivan and Dr. Anita Klein for guidance and support through the doctoral program, and to my friends and laboratory fellows at UNH for good discussions and enjoyable times during my time in the USA. I am grateful for funding provided by SYNTHESYS, the Faroese Research Council, the UNH College of Life Science and Agriculture, the UNH Department of Biological Sciences, the UNH Marine Program, the UNH Graduate School and the Hatch Grant from the New Hampshire Agricultural Experiment Station that has enabled the research work to be carried out. I would like to thank the following herbaria for letting me use their material for DNA extraction: Natural History Museum, London, UK; Botanical Museum in Copenhagen, Denmark; Norwegian University of Science and Technology, Tromsø, Norway; New York Botanical Garden, USA; National Research Council of Canada, Nova Scotia, Canada; and the following people for sending me samples: Dr. Sandra Lindstrom, Dr. Gary W. Saunders, Dr. Susse Wegeberg, Dr. Poul Møller Pedersen (deceased 17th of June 2012), Dr. Kjersti Sjøtun and Bernt-Gunnar Østerkløft. A special thank to Fiskaaling for providing laboratory and office space and to Heðin I. Abrahamsen for drawing maps. Finally, I would like to thank my close family, partner and friends for endless support throughout this long process. iii TABLE OF CONTENTS ACKNOWLEDGEMENTS………………………………………………………………………iii LIST OF TABLES………………………………………………………………………………..vi LIST OF FIGURES……………………………………………………………………………..viii ABSTRACT………………………………………………………………………………………ix CHAPTER I. INTRODUCTION…………………………………………………………………1 Biogeographic regions and effects of glaciation in the North Atlantic……………..3 Connection between the North Atlantic and the North Pacific Oceans……………4 The macroalgae…………………………………………………………………………5 The Bangiales…………………………………………………………………………...6 Diversity, distribution and molecular tools……………………………………………8 Aims………………………………………………………………………………………9 CHAPTER II. NEW INSIGHTS INTO THE BIODIVERSITY AND GENERIC RELATIONSHIPS OF FOLIOSE BANGIALES (RHODOPHYTA) IN ICELAND AND THE FAROE ISLANDS……………..........................................................................................12 Abstract…………………………………………………………………………………12 Introduction……………………………………………………………………………..14 Materials and methods………………………………………………………………..17 Results………………………………………………………………………………….22 Discussion……………………………………………………………………………...29 CHAPTER III. DIVERSITY AND DISTRIBUTION OF FOLIOSE BANGIALES (RHODOPHYTA) IN WEST GREENLAND: A LINK BETWEEN THE NORTH ATLANTIC AND NORTH PACIFIC…………………………………………………………………………41 Abstract………………………………………………………………………………....41 Introduction……………………………………………………………………………..42 Materials and methods………………………………………………………………..44 Results…………………………………………………………………………………..48 Discussion………………………………………………………………………………52 CHAPTER IV. DIVERSITY AND DISTRIBUTION OF FOLIOSE BANGIALES (RHODOPHYTA) SPECIES IN THE NORTHWEST ATLANTIC IN THE CONTEXT OF THE NORTH ATLANTIC ……………………………………………………………………...63 Abstract………………………………………………………………………………....63 Introduction……………………………………………………………………………..65 Materials and methods………………………………………………………………..68 Results…………………………………………………………………………………..70 Discussion………………………………………………………………………………81 CHAPTER V. WILDEMANIA AMPLISSIMA (BANGIALES, RHODOPHYTA) IN THE NORTH ATLANTIC AND NORTH PACIFIC: PRELIMINARY PHYLOGEOGRAPHIC OBSERVATIONS……………………………………………………………………………….91 Abstract…………………………………………………………………………………91 Introduction……………………………………………………………………………..93 iv Materials and methods………………………………………………………………..97 Results…………………………………………………………………………………100 Discussion…………………………………………………………………………….105 CONCLUSION………………………………………………………………………………...123 REFERENCES………………………………………………………………………………...127 APPENDICES…………………………………………………………………………………142 Appendix A……………………………………………………………………………143 Appendix B……………………………………………………………………………154 Appendix C……………………………………………………………………………166 Appendix D……………………………………………………………………………167 Appendix E……………………………………………………………………………189 Appendix F…………………………………………………………………………….190 v LIST OF TABLES CHAPTER II Table 2.1 History of foliose Bangiales collections in Iceland……………..…32 Table 2.2 History of foliose Bangiales collections in the Faroe Islands……33 Table 2.3 Pairwise distances between P. linearis, P. umbilicalis and P. dioica, based on cox1 (537 bp) and rbcL (367 bp at the 3ʼend) sequences…………………………………………………….34 Table 2.4 The 25 foliose Bangiales species recorded from the Northwest Atlantic (from Maine to Long Island; Pyropia njordii has also been observed in Nova Scotia, Canada), Northeast Atlantic, Iceland and the Faroe Islands…………………………….35 Table 2.5 Intraspecific sequence variation in the rbcL gene………………...36 CHAPTER III Table 3.1 Pairwise distances for the 3ʼ rbcL + 5ʼrbcL–S marker……………56 Table 3.2 The latitudinal distribution of foliose Bangiales species on the west coast of Greenland………………………………………….....57 Table 3.3 Foliose Bangiales species recorded from the North Atlantic, Iceland, Faroe Islands and Greenland……………………………..58 CHAPTER IV Table 4.1 Sites visited between Connecticut and Florida in May and June 2011……………………………………………………………..87 Table 4.2 Distribution of the foliose Bangiales species on the Northwest Atlantic coast……………………………………………………….....89 Table 4.3 Northeast Atlantic (including Iceland and the Faroe Islands) foliose bangiales species diversity and new distribution records based on DNA sequence identification…………………..90 CHAPTER V Table 5.1 A list of amplification and sequencing primers used in this study………………………………………………………………….112 Table 5.2 Clade I: Genetic variation in the rbcL gene based on Tamura-Nei genetic distance model and presented as patristic distances (sum of branch lengths) and differences in base pairs……………………………………………………………113 Table 5.3 Clade II: Genetic variation in the rbcL gene based on Tamura-Nei genetic distance model and presented as patristic distances (sum of branch lengths) and differences in base pairs……………………………………………………………114 vi Table 5.4 Clade III: Genetic variation in the rbcL gene based on Tamura-Nei genetic distance model and presented as patristic distances (sum of branch lengths) and differences in base pairs……………………………………………………………115 Table 5.5 Wildemania amplissima sampling locations……………………..116 Table 5.6 Geographic distribution of haplotypes…………………………….117 Table 5.7 Haplotype- and nucleotide diversity in the Northeast- and Northwest Atlantic, and the North Pacific………………………...118 vii LIST OF FIGURES CHAPTER II Figures 2.1- 2.11 Gross morphology of the Bangiales flora in Iceland……………...37 Figure 2.12 Maximum likelihood phylogram based on rbcL sequences, with specimens of foliose Bangiales species primarily from Iceland and the Faroe Islands………………………………………38 Figure 2.13 Maximum likelihood phylogram based on cox1 sequences, with specimens of foliose Bangiales species primarily from Iceland and the Faroe Islands………………………………………39 Figures 2.14- 2.20 Pyropia njordii sp. nov……………………………………………….40 CHAPTER III Figure 3.1 Map of collecting locations on the west coast of Greenland……………………………………………………………..59 Figure 3.2 Maximum likelihood phylogram based on rbcL sequences, placing foliose Bangiales from Greenland in a wider phylogenetic context…………………………………………………60 Figure 3.3 Distribution of the species on different substrata based on 75 specimens……………………………………………………………..61