Phylogeny, phylogeography and population connectivity of Lessonia (Phaeophyceae) Cover: Lessonia variegata in the lower intertidal of Moa Point, Wellington. Phylogeny, phylogeography and population connectivity of Lessonia (Phaeophyceae) by Peter Martin A dissertation submitted to Victoria University of Wellington in fulfilment of the requirements for the degree of Doctor of Philosophy in Ecology and Biodiversity Victoria University of Wellington Te Whare Wānanga o te Ūpoko o te Ika a Māui 2011 i Abstract The brown algal genus Lessonia is distributed in the Southern Hemisphere where it can form dominant kelp beds on the exposed rocky shores of New Zealand, South America and Tasmania. Its disjunct distribution within the West Wind Drift contrasts with the view that it is a poor disperser. Apart from studies in Chile, where it is an economically important genus, little is known about Lessonia and in some areas even the number of species is unknown. Using different genetic markers I examined the phylogeny, phylogeography, and the connectivity of populations in Lessonia. Using the literature, species affiliations and nomenclatural problems have been investigated. Combining the sequences of three mitochondrial, one chloroplast and two nuclear markers, a supermatrix approach was used to investigate the phylogenetic relationship and the timing of speciation for all known Lessonia species. The Australasian Lessonia species form a clade within a paraphyletic grouping of South American species. Radiation in Lessonia occurred about 5 Mya at the beginning of the Pliocene and rapid radiation took place in Australasia 3.5 Mya. The data also revealed cryptic species within a L. variegata species complex. Further analysis within the Australasian clade, using mitochondrial (atp8-sp) and chloroplast (rbc-sp) markers and wider sampling (469 individuals from 57 sample sites) supported four cryptic species and revealed localized distribution for all Australasian lineages. Genetic breaks between Lessonia lineages corresponded well to known biogeographic regions and could be correlated to the geographic structure of New Zealand at the end of the Pliocene. The Cook Strait region was analysed more closely with newly developed microsatellite markers to test the influence of geographic breaks (Cook Strait and Palliser Bay) on the connectivity of populations. ii The results suggested that connectivity depends on the width of unsuitable habitat, and within inner Cook Strait it is facilitated by sometimes strong tidal flows that create turbulences and unique current patterns. The results implied that rafting is an important mean of dispersal. The study of the early literature on Lessonia supported the new lectotypification of L. flavicans but revealed that L. frutescens and possibly L. ovata (supported by images of rediscovered herbarium material) are synonymous to L. searlesiana and as the older epithets they should have priority. Suggestions have been made for the lectotypification of L. fuscescens and L. ovata. In general Lessonia shows non-overlapping distribution in Australasia but overlapping distribution in South America. Despite being a poor disperser, indicated by fine scale genetic structure, Lessonia is also able to connect populations over wide areas of unsuitable habitats. iii für W altraud und Ludw ig iv Acknowledgements I have to mention red-hatted Jacques-Yves Cousteau as the person who planted an interest in marine biology into the mind of a very young Peter Martin. I am very grateful to my parents, who stood behind me and even showed excitement when I finally decided to follow my dreams and changed my professional career into an academic one with an uncertain future. Now at the end of my PhD I want to thank all the people I met along the way, who inspired me and helped me achieving my goal. These are my fellow students from the Universities in Halle, Rostock and the Victoria University in Wellington, the teachers and lecturers I enjoyed at these Institutions and my PhD supervisors at Victoria University. I also want to show gratitude to the three examiners of this thisis, Sharyn Goldstien, Ken Ryan, and John Bolton. I like and thank my best friends that have also been listeners and tellers of scientific fictions, Kai Ziervogel, Christian Bödeker, Rudi Schnitzler, Marc Hasenbank, and Elizabeth Heeg. Last but most important of all I want to thank the people I love and without their help nothing would have been possible. These are my mother, Waltraud, my father, Ludwig, my brother, Jochen, and my partner, Walli (and the apes Wumbaba and Wum). v Acknowledgements to Chapter 2 I am grateful to Ceridwen (Crid) Fraser, Cesar Cardenas, Christian Bödeker, Christopher Lane, Erasmo Macaya-Horta, Joe Buchanan, Loa Hunt, Robyn Dunmore, and Wendy Nelson for providing algal material. I also appreciate the efforts of Heroen Verbruggen and two anonymous reviewers for the Journal of Phycology for a critical reading of previous versions of this manuscript. This research was funded through a scholarship from the Victoria University of Wellington. Acknowledgements to Chapter 3 I am grateful to Christian Bödeker, Don Neale, Erasmo Macaya-Horta, Gert & Christine Vermeer, Joe Buchanan, Loa Hunt, Nick Shears, Robyn Dunmore, Waltraud Schiedermair and Wendy Nelson for helping with the sampling and/or providing algal material. I am thankful to the Wellington Botanical Society for grants to ease the cost of travelling. This research was partially funded by the Department of Conservation New Zealand and through a scholarship from the Victoria University of Wellington. Acknowledgements to Chapter 4 This project was partially funded by the Department of Conservation New Zealand and through a scholarship from the Victoria University of Wellington. I am grateful vi to Lara Shepherd, Joe Buchanan, and Erasmo Macaya-Horta for their help in microsatellite library construction. Acknowledgements to Chapter 5 The extensive samplings have only been possible with the help of self-giving Waltraud Schiedermair and also Erasmo Macaya-Horta. I want to thank the caretaker of the Fighting Bay Cable Station for granting excess to the bay and the management of Meridian Energy for providing escort to Cape Terawhiti. Acknowledgements to Chapter 6 The curators of the following herbaria have been helpful in providing specimens or high quality images of Lessonia collections: BM, PC, S, TCD, UC and US. I am especially grateful to Paul Silva and Kathy Ann Miller from UC who helped with taxonomic questions and provided images of anatomic sections, and to Bruno de Reviers who sent me an early manuscript of his research. I am also grateful to Phil Garnock-Jones for helping in dealing with the International Code of Botanical Nomenclature. vii Table of Contents Abstract ........................................................................................................................ i Acknowledgements ................................................................................................................. iv Table of Contents .................................................................................................................... vii List of Figures ...................................................................................................................... xi Supplementary Figures ........................................................................................................................ xix List of Tables .................................................................................................................... xxi Supplementary Tables........................................................................................................................ xxiv Chapter 1 Introduction & overview .................................. 1 1.1 Thesis structure ................................................................................................................1 1.2 Lessonia – First descriptions and biogeography ...............................................4 1.2.1 Distinguishing features of Lessonia species and their synonyms .......... 9 1.2.1.1 Australasian species ...................................................................... 9 1.2.1.2 South American species .............................................................. 11 1.2.2 Figures and Tables ............................................................................. 14 1.3 Lessonia – Biology and dispersal ........................................................................... 26 1.3.1 Figures............................................................................................... 32 1.4 Lessoniceae – New molecular taxonomy and phylogeny .......................... 34 1.4.1 Tables ................................................................................................ 38 1.5 References ........................................................................................................................ 41 Chapter 2 Molecular Phylogeny and timing of radiation in Lessonia (Phaeophyceae, Laminariales) .. 53 2.1 Abstract.............................................................................................................................. 53 2.2 Introduction..................................................................................................................... 55 2.3 Material and Methods ................................................................................................. 58 2.3.1 Sampling ..........................................................................................