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Thesis) Submitted to the School of Graduate Studies In GROWTH IN COLD-WATER OCTOCORALS: RATES, MORPHOLOGY AND ENVIRONMENTAL CONTROLS by © Bárbara de Moura Neves (Thesis) submitted to the School of Graduate Studies in partial fulfillment of the requirements for the degree of PhD in Biology/Faculty of Sciences Memorial University of Newfoundland September 2016 St. John’s Newfoundland and Labrador i Abstract Deep-water corals can have high longevities and slow growth rates. While most studies have been focusing on growth rates of gorgonians, few studies have examined rates and patterns in other octocorals, such as sea pens. Here I studied growth rates and patterns in the sea pens Halipteris finmarchica, Umbellula encrinus, and Pennatula grandis, and in the gorgonians Primnoa pacifica, Primnoa resedaeformis, and Keratoisis sp. Longevity and growth rates were estimated by examining growth rings in their skeletons (axes), whose morphology and composition were also explored. Growth patterns were investigated by examining relationships between colony and axis characteristics. Relationships between colony growth rates and environmental factors were also assessed. The morphology of the axis in sea pens varies among taxa and within colonies, and their carbonate portion is composed in average of 65-85% of magnesian calcite (Ca, Mg)CO3. Sea pens can have longevities reaching ~70 years in U. encrinus. Diametric growth rates in H. finmarchica, U. encrinus, and P. grandis were comparable in magnitude, ranging between 0.13-0.14 mm∙yr-1. Using new and published data, diametric growth rates in P. pacifica and P. resedaeformis averaged 0.43 mm∙yr-1 and were not statistically different. In the Arctic populations of Keratoisis sp. studied here radial growth rates ranged between 0.010-0.018 mm∙yr-1, much slower than in most other populations of this coral. Relationships between growth rates and environmental variables were statistically significant for bottom temperature (U. encrinus), chlorophyll a (P. grandis and Primnoa spp.), and depth (Primnoa spp.). Geochemical analyses yielded a number of trace element peaks comparable to the number of growth rings in sea pens. The 14C curves for the ii studied sea pens follow the same patterns seen in the calcite of other octocorals in the region, in which rings are formed annually. Growth rate and longevity data can support conservation efforts for these vulnerable species. iii Acknowledgments I am deeply grateful to my supervisor Dr. Evan Edinger, who provided the opportunity and the resources that allowed me to explore a whole new coral world. I also thank my committee members Dr. Graham Layne, Dr. Annie Mercier, and Dr. Kent Gilkinson for the guidance and valuable comments through the years. The examiner committee members Dr. Robert Hooper, Dr. Stewart Fallon, and Dr. Gary Williams are also thanked for their suggestions and comments. I thank Dr. Sally Leys (chief scientist), Jackson Chu, Vince Lecours, CCGS Vector captain and crew, and the ROPOS team for making possible the 2011 ROV cruise in the Strait of Georgia. I also thank CCGS Amundsen captains and crew, chief scientists Dr. Louis Fortier, Dr. Maurice Levasseur, Dr. Philippe Archambault, ROV pilots Ian Murdock, Peter Lockhart, Trevor Shepherd, and Vincent Auger for the 2013-2015 cruises to Baffin Bay. Vonda Wareham (DFO) provided a great number of the samples used in this thesis, and shared her knowledge and enthusiasm, which were a great incentive to the progress of this work. Dr. Margaret Treble and Dr. Kevin Hedges (DFO), Cindy Grant, Alexis Burt, Brynn Devine, and Laura Wheeland also provided valuable sea pen samples included in this study. Dr. Pål Buhl-Mortensen provided raw unpublished data on Primnoa resedaeformis, and Victoria Howse provided Primnoa samples from Labrador. The use of museum specimens was only possible thanks to museum curators and technicians who kindly allowed me to study precious specimens. I sincerely thank Dr. Stephen Cairns and Tim Coffer (Smithsonian Institution) for the Primnoa samples, Dr. iv Jean-Marc Gagnon (Canadian Museum of Nature) for the Pennatula grandis samples, Nathalie Djan-Chékar (The Rooms, St. John’s) for access to the collection at The Rooms, and Andrew Cabrinovic (Natural History Museum, London) for access to the Pennatula collection. Several people from the CREAIT Network team (Memorial University of Newfoundland) contributed to logistical/methodological aspects of this thesis. Glenn Piercey and Brian Loveridge kindly gave me access to the isomet and rock saws all the times I needed. Glenn also patiently guided me to use the microdrill, performed the trace elements analyses, and also contributed with interesting questions and comments about my study. Michael Shaffer helped with the SEM, David Grant and Dylan Goudie with the polishing machine, and Wanda Aylward provided the XRD analyses. Vince Lecours and Dave Mercer helped me in my first steps in the world of ArcGIS, and Charlie Conway helped with his advice on mapping and cartography. I also thank Dr. Patrick Gagnon for allowing access and use of equipment in his lab, and the staff at the Biology department for facilitating everything, especially Gary Collins, Cherie Hussey, Alvin Kenny, Shirley Kenny, and also Pam Murphy from the Geography department. I thank Dr. Ted Miller for his great incentive and support along these years. My research was funded by the Canadian Healthy Oceans Network (CHONe) and by a Doctoral scholarship from the Natural Sciences and Engineering Research Council of Canada (NSERC), which made this project possible. Finally, my husband David Bélanger could not have been more patient with my incessant biology talk, and I am extremely grateful for his tolerance and support. I also v thank my mom Lúcia and my sisters Débora and Gabriela for the immense support and love. vi Table of Contents Abstract ........................................................................................................................... ii Acknowledgments ........................................................................................................... iv Table of Contents........................................................................................................... vii List of figures ................................................................................................................ xvi List of tables ................................................................................................................. xxx Abbreviations ............................................................................................................ xxxiv Co-authorship Statement ............................................................................................ xxxv Appendices .............................................................................................................. xxxvii Publications arising ................................................................................................... xxxix 1. Introduction .............................................................................................................. 1-1 1.1 Octocoral morphology, skeletal composition, and growth patterns ....................... 1-3 1.1.1 Growth patterns in alcyonaceans (order Alcyonacea) ..................................... 1-5 1.1.2 Growth patterns in sea pens (order Pennatulacea) ......................................... 1-10 1.1.3 Growth rings in octocorals ............................................................................ 1-14 1.2 Growth rates ......................................................................................................... 1-18 1.2.1 Growth rates in relation to environmental variables ..................................... 1-19 1.3 Methods for studying growth rates in octocorals ................................................. 1-20 1.3.1 Aquaria/mesocosm ......................................................................................... 1-21 1.3.2 Ring counting ................................................................................................ 1-22 1.3.3 Validation techniques ..................................................................................... 1-24 1.4 Limiting factors to octocoral growth .................................................................... 1-28 vii 1.5 Conservation perspective ..................................................................................... 1-29 1.6 Thesis outline ....................................................................................................... 1-30 1.7 References ............................................................................................................ 1-33 2. Decadal longevity and slow growth rates in the deep-water sea pen Halipteris finmarchica (Sars, 1851) (Octocorallia: Pennatulacea): Implications for vulnerability and recovery from anthropogenic disturbance ....................................................................... 2-1 Abstract ........................................................................................................................ 2-1 2.1 Introduction ............................................................................................................ 2-3 2.2 Materials and Methods ........................................................................................... 2-6 2.2.1 Study area and sampling .................................................................................. 2-6 2.2.2 Ring counting and growth rate estimation ......................................................
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