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Characterization and Role of Major Deep-Sea Pennatulacean Corals in the Bathyal Zone of Eastern Canada

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Characterization and Role of Major Deep-Sea Pennatulacean Corals in the Bathyal Zone of Eastern Canada Characterization and role of major deep-sea pennatulacean corals in the bathyal zone of Eastern Canada by ©Sandrine Baillon A thesis submitted to the School of Graduate Studies in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Ocean Sciences and Department of Biology, Memorial University of Newfoundland, St. John’s, Newfoundland and Labrador, Canada October 2014 i A mes parents, ii Abstract Pennatulaceans (Octocorallia: Pennatulacea), commonly called sea pens, are colonial corals that typically anchor themselves into soft sediment (mud, sand), allowing them to colonize large areas of the sea floor from the intertidal zone down to the abyssal plain. Sea pens can occur sparsely or form large aggregations, suggesting that they may provide an important structural habitat to other organisms by increasing the complexity of the muddy seabed. The investigation of the three most common sea pens (Anthoptilum grandiflorum, Halipteris finmarchica and Pennatula aculeata) of the continental slope of Newfoundland and Labrador (Northwest Atlantic) showed different morphologies and adaptations to environmental parameters. Variations in their morphology were visible along bathymetric and latitudinal gradients, following food availability. This study identified different feeding strategies according to stable isotope signatures and macromorphologies (polyp diameter, colony length, shape). Different defense strategies were also identified based on the presence and localisation of sclerites in the tissues. Reproductive strategies were determined for A. grandiflorum and H. finmarchica, with both species presenting a seasonal spawning between April (Southern Newfoundland) and July/August (Labrador). The latitudinal shift in spawning followed the development of the surface phytoplankton bloom (i.e. sinking of phytodetritus). Low fecundity at the polyp level (13 and 6 oocytes polyp-1, respectively) yielded similar whole-colony fecundity in the two species (500-6000 oocytes colony-1). The measure of fecundity is discussed to highlight the importance of standardizing metrics (mature oocyte just before spawning) to avoid an overestimation of the fecundity and to allow comparison among iii species. Only ~20% of the oocyte matured in both species. The remaining oocytes in A. grandiflorum disappeared, indicating that oogenesis develops and culminates over 12 months. In H. finmarchica the persistence of the small oocytes indicates protracted oogenesis (>12 months). Finally, an analysis of the associated biodiversity showed that sea pens have relatively few associates but that they play an important role in their life history. Some associates are obligate (e.g. parasitic copepods) and others facultative symbionts (the sea anemone Stephanauge nexilis). The seasonal (April-May) presence of fish larvae (Sebastes spp.) and shrimp larvae (Pandalus borealis, Pasiphae multidentata and Acanthephyra pelagica) emphasize the role of sea pens as nursery habitat, and provides an argument to recognize them as an essential fish habitat. iv Acknowledgements I would like to thank Annie Mercier and Jean-François Hamel for believing in me in 2008 at the beginning of my Masters and then for offering me the opportunity to pursue this Ph.D. Thanks for all your support during these last 6 years. It has been an amazing time with you.Thanks Annie for your availability to discuss the project. Thanks Jean- François for the long hours on the phone which have always been helpful. I also would like to thank all the scientific staff of Fisheries and Oceans Canada and the Canadian Coast Guard for helping us with sampling on board of the CCGS Teleost. Thanks to Vonda Wareham and Kent Gilkinson for access to the samples. Thanks to Iliana Dimitrove and Katrin Zipperlen from the School of Medical Studies for their assistance with histological processing. Thanks to the different students who helped me in the acquisition of the data, especially Matthew English and Christa Spurrell. Thanks to my supervisory committee members, Suzanne Dufour and Igor Eeckhaut, for advice during the last three years. Thanks to the members of my examination committee Gary Williams, Robert Hooper and Iain McGaw for their constructive comments. Thanks to the students in the Mercier Lab (Zhao, David and Katie G.) for your time, discussing about sciences and other topics. Thanks to all the new students (Bruno, Matt, Katie K., Emy, Jess) who have given me breaks in my writing to help in the lab during these last few months. v Thanks to my friends (Emily, Jen, Lindsey, Yannick, Kim and Marie) and roommates (Gen, Laura, Desta) for listening to me and for their understanding during my bad days. Thanks to my family, Vincent, Poy, Luca and Loïc. Last but not least I want to thank my parents for their support during these last 29 years. Nothing would have been possible without you. “Ne crois pas que tu t’es trompé de route quand tu n’es pas allé assez loin” Claude Aveline vi Table of contents Abstract ________________________________________________________ iii Acknowledgements ________________________________________________ v Table of contents ________________________________________________ vii List of tables ____________________________________________________ xi List of figures __________________________________________________ xiii List of appendices ______________________________________________ xviii Co-authorship statement __________________________________________ xx Chapter 1 : General introduction __________________________________ 1-1 1. Corals ________________________________________________________ 1-2 1.1 Coral morphology _____________________________________________________ 1-2 1.2 Reproduction _________________________________________________________ 1-4 2 Cold-water corals _______________________________________________ 1-6 2.1 Overview ____________________________________________________________ 1-6 2.2 Pennatulacean corals ___________________________________________________ 1-7 3. Study areas and focal species _____________________________________ 1-9 4. Objectives and chapter structure ________________________________ 1-10 References _____________________________________________________ 1-13 Figures ________________________________________________________ 1-20 Chapter 2 : Comparative biometry and isotopy of three dominant pennatulacean corals in the Northwest Atlantic _____________________ 2-1 Abstract ________________________________________________________ 2-2 1. Introduction ___________________________________________________ 2-3 2. Material and methods ___________________________________________ 2-7 2.1 Sampling ____________________________________________________________ 2-7 2.2 Biometrics ___________________________________________________________ 2-7 2.2.1 Colonies and polyps ______________________________________________________ 2-7 2.2.2 Sclerites ________________________________________________________________ 2-8 2.3 Isotopic analysis _____________________________________________________ 2-10 2.4 Data analysis ________________________________________________________ 2-11 3. Results ______________________________________________________ 2-12 3.1 Colony metrics ______________________________________________________ 2-12 3.1.1 Colony length, wet weight and weight/length ratio ______________________________ 2-12 3.1.2 Peduncle length _________________________________________________________ 2-15 3.2 Polyp diameter and density _____________________________________________ 2-16 vii 3.3 Sclerite metrics ______________________________________________________ 2-18 3.3.1 Description ____________________________________________________________ 2-18 3.3.2 Influence of colony length, depth and region on sclerite metrics ___________________ 2-20 3.4 Isotopic analysis _____________________________________________________ 2-22 4. Discussion ____________________________________________________ 2-23 4.1 Variations and similarities in macrometrics ________________________________ 2-24 4.2 Diet, trophic level and macrometrics _____________________________________ 2-25 4.3 Spatial and temporal trends in macrometrics _______________________________ 2-27 4.4 Description and role of sclerites _________________________________________ 2-30 4.5 Variations in sclerite metrics ____________________________________________ 2-32 Acknowledgements ______________________________________________ 2-33 References _____________________________________________________ 2-34 Tables _________________________________________________________ 2-42 Figures ________________________________________________________ 2-46 Supplementary materials _________________________________________ 2-56 Chapter 3 : Seasonality in reproduction of the deep-water pennatulacean coral Anthoptilum grandiflorum ______________________________________ 3-1 Abstract ________________________________________________________ 3-2 1. Introduction ___________________________________________________ 3-3 2. Material and methods ___________________________________________ 3-5 2.1 Sampling and handling _________________________________________________ 3-5 2.2 Sample processing _____________________________________________________ 3-7 2.3 Fecundity ____________________________________________________________ 3-7 2.4 Gametogenic stages ____________________________________________________ 3-8 2.5 Maturity stage index and oocyte and spermatocyst size distribution ______________ 3-9 2.6 Data analysis ________________________________________________________
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