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432 Pages·1990·30.83 MB·English
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Evolutionary Biogeography of the Marine Algae of the North Atlantic NATO ASI Series Advanced Science Institutes Series A series presenting the results of activities sponsored by the NATO Science Committee, which aims at the dissemination of advanced scientific and technological knowledge, with a view to strengthening links between scientific communities. The Series is published by an international board of publishers in conjunction with the NATO Scientific Affairs Division A Life Sciences Plenum Publishing Corporation B Physics London and New York C Mathematical and Kluwer Academic Publishers Physical Sciences Dordrecht, Boston and London D Behavioural and Social Sciences E Applied Sciences F Computer and Springer-Verlag Systems Sciences Berlin Heidelberg New York G Ecological Sciences London Paris Tokyo Hong Kong H Cell Biology Barcelona Series G: Ecological Sciences Vol. 22 Evolutionary Biogeography of the Marine Algae of the North Atlantic Edited by David J. Garbary Department of Biology St. Francis Xavier University Antigonish, Nova Scotia Canada B2G 1C O G. Robin South Huntsman Marine Science Centre St. Andrews, New Brunswick Canada EOG 2XO Springer-Verlag Berlin Heidelberg New York London Paris Tokyo Hong Kong Barcelona Published in cooperation with NATO Scientific Affairs Division Proceedings of the NATO Advanced Research Workshop on Evolutionary Biogeography of the Marine Algae of the North Atlantic held in St. Andrews, N.B., Canada, September 18-22, 1989 Library of Congress Cataloging-in-Publication Data NATO Advanced Research Workshop on Evolutionary Biography of Marine Algae of the North Atlantic (1989 : Sl. Andrews, N. B.) Evolutionary biogeography of the marine algae of the North Atlantic 1 edited by David J. Garbary, G. Robin South. p. cm.-(NATO ASI series. Series G, Ecological sciences; vol. 22) "Proceedings of the NATO Advanced Research Workshop on Evolutionary Biogeography of the Marine Algae of the North Atlantic. held in SI. Andrews, N. B., Canada, September 18-22, 1989"-T.p. verso. "Published in cooperation with NATO Scientific Affairs Division." ISBN-13: 978-3-642-75117-2 e-ISBN-13: 978-3-642-75115-8 001: 10.1007/978-3-642-75115-8 1. Marine algae-North Atlantic Ocean-Geographical distribution-Congresses. 2. Marine algae-North Atlantic Ocean-Evolution-Congresses.1. Garbary, David J. II. South, G. Robin. III. North Atlantic Treaty v. Organization. Scientific Affairs Division. IV. Title. Series: NATO ASI series. Series G, Ecological sciences; no. 22. QK572.5.N67N381990 589.392'11-dc20 90-43825 This work is subjectto copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, re-use of illustrations, recitation, broadcasting, reproduction on microfilms or in other ways, and storage in data banks. Duplication of this publication or parts thereof is only permitted under the provisions of the German Copyright Law of September 9,1965, in its current version, and a copyright fee must always be paid. Violations fall under the prosecution act of the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1990 Softcover reprint of the hardcover 1st edition 1990 2131/3140-543210-Printed on acid-free-paper PREFACE Evolutionary biogeography of the marine algae of the North Atlantic was the primary theme of the Fourth Seaweed Biogeography Worksho.p. This five day symposium was held in St. Andrews, New Brunswick, in September 1989, and was organized under the auspices of the International Working Group on Seaweed Biogeography. Previous workshops in this series were held at Hamburg (1982), Groningen (1984) and Helgoland (1986), and were published in Helg.oliinder Meeresunters (38: 225-417 and 41: 233-383). The Organizing Committee for the conference comprised the two of us, and C. van den Hoek, who formulated the scientific program, and identified the primary participants. The primary financial support for the meeting was provided by a grant from NATO (ARW 931/88), under their Advanced Research Workshop program. Primm The Theme This meeting was based on the notion that the current status of algal biogeography, and its advance, could be best addressed by a multidisciplinary approach to a single theme: THE EVOLUTIONARY BIOGEOGRAPHY OF THE MARINE ALGAE OF THE NORTH ATLANTIC. Participants were asked to focus their presentations on this theme, or to provide philosophical and/or methodological discussions of biogeography. In addition to strictly algal biogeographers, people in related fields of geology, theoretical biogeography and marine ecology were invited, so that their theoretical and philosophical concerns, as well as their methods, could be better evaluated by th~ more traditional 'phycologists', many of whom undertake biogeography as a supplement to their primary systematic, physiological, or ecological interests. Biogeography, is by its nature, interdisciplinary, and the St Andrews meeting was a reflection of this. VI The St. Andrews Symposium: A NATO Advanced Research Workshop The meeting was attended by about 30 scientists from eight countries, and included a series of 20 formal paper presentations, of which 17 are included in this volume. The presentations not in this volume are: 1) M. Ragan, C.J. Bird and E.L. Rice "Nuclear small-subunit ribosomal ribonucleic acid sequences and molecular relationships within the Gracilariales"; 2) S. Srivastava "Evolution of the North Atlantic -Jurassic to present"; and 3) J. Cabioch "Adaptive strategies and distribution of Corallinales". Two presentations of computer sofware for biogeography and phylogenetic analysis were also made (p. Legendre, D. Garbary). At the conclusion of the meeting, a panel discussion was used as a venue for discussing theoretical issues, progress that had been made in understanding the overall problem that was the theme of the meeting, and how algal biogeographers might redirect their approach so that biogeographic hypotheses might be more rigorously evaluated. Throughout the meeting, informal discussions were held in Anderson House, before, during and after meals, and long into the evenings. These discussions were suitably lubricated from the fridge in the basement of Anderson House. The overall tone of the five day meeting was, at the same time, both intense and informal, and highly productive. This volume identifies the formal contributions of the symposium; however, it cannot capture the excitement and general atmosphere of the symposium. Organization of Entries The papers in this volume have been put into a particular order to reflect some of the dominant themes of the conference, as well as seaweed biogeography in general. None of the sections outlined below are mutually exclusive, and some papers might easily fit into other sections. The first section treats conceptual and methodological issues and includes two papers. The opening paper by I. Ball is a discussion of some of the conceptual/philosophical issues in contemporary biogeographic theory, and his paper provides a brief introduction to this literature for a phycological audience. This is followed by a complementary paper by P. Legendre describing quantitative methods in biogeography, providing a list of the appropriate computer programs for implementing these methods. VII The next section includes two papers, by D. Schnitker & J. Jorgensen and C. van den Hoek & A. M. Breeman, who examine the effects of Pleistocene glaciation on algal distributions and the changes that occurred after retreat of the ice. This is examined on a local scale (phytoplankton in the Bay of Fundy), and on a wider scale (seaweeds in the North Atlantic). The next section treats concepts of dispersal and adaptation in an ecological and evolutionary context; in the first instance dealing with barnacle evolution in eastern Canada (E. Bourget et at.), and the second dealing with evolution of herbivory on coralline algae in the North Atlantic and North Pacific (R. Steneck). Processes of speciation were a major theme of the meeting and the next three papers by P.V.M. Bot & C. van den Hoek, B. Egan et at., and B. Stache provide insight into seaweed speciation and genetics. They use a variety of different approaches, including hybridization studies, morphometric analysis and DNA/DNA hybridization. Two papers dealing with evolution of Laminariales were presented, by K. Liining & I. tom Dieck and L. Druehl, which applied morphological and physiological features, as well as molecular biology to the problem of kelp phylogeny. These papers illustrate, on the one hand, the extent to which phylogenetic relationships can be hypothesized using more traditional features; and on the other, the exciting potential for resolving phylogenetic problems using molecular techniques. The relationship between systematics and biogeography in Rhodophyta is discussed in three papers by A. Athanasiadis, C. Maggs, and M.D. Guiry & D. Garbary studying a diversity of red algal taxa. These papers emphasize the importance of taxonomy in dealing with historical biogeography. The division of the North Atlantic into biogeographic regions is an ongoing theme in seaweed biogegraphy; this is treated by Tittley et at. using a variety of quantitative methods, including an innovative use of cladistic analysis. The final section of these proceedings includes two highly synthetic papers discussing the evolutionary biogeography the of entire seaweed flora of the Arctic (Wilce) and the red algae of the North Atlantic (M. Hommersand). Both of these papers emphasize the importance of earth history and systematics in developing an understanding of evolutionary biogeography of marine algae. Preparation of articles for publication Authors were asked to provide copies of manuscripts and computer disk versions of the papers at the St.Andrewsmeeting. Manuscripts were reviewed, generally by two referees, at least one of whom was not a participant at the meeting. The editors are especially grateful to the reviewers who read and critically evaluated the various manuscripts. The individuals who did not participate in the conference, but who provided reviews included: M. Alvarez, M. VIII Blouw, M. Cormaci, P.W. Gabrielson, R.G. Hooper, S. Lindstrom, J.L. Olsen, C. Plowright, G. Russell, E. Theriot, and A. Whittick. After review and revisions, manuscripts were copy edited, checked for style, and were then formatted using Microsoft Word (version 4) (Copyright Microsoft Corp.), implemented on a Macintosh SE30 computer. Manuscripts were set in Times, and camera-ready copy was produced using an Apple laser printer. Dedication to "Bud" Brinkhuis and "Barty" Egan After this conference was organized, and during the preparation of this volume, two of the contributing authors, P.F. "Barry" Egan and B. "Bud" Brinkhuis, passed away. It is with considerll:ble sadness that we mark the deaths of these friends and fellow phycologists. The deaths, of an established colleague, an~ a young scientist of considerable promise, make this a particular tragedy. This volume is dedicated, in part, to these two phycologists. Acknowledgments This volume, and the symposium on which it is based, would not have come to fruition without the help and dedication of a wide variety of people and organizations. First, we thank the Staff (especially Irene Pohle)and Directors of the Huntsman Marine Science Centre, and the Department of Fisheries and Oceans, Biological Station, St. Andrews for the high standard of general organization of the meeting, and of the facilities that were made available. All of the participants were housed in Anderson House, whose Staff are complimented for the quality of meals, and the fine surroundings in which many of the discussions were held. Barbara South is to be congratulated for the excellent banquet that she hosted. We are most grateful to the external and internal reviewers who evaluated the manuscripts. Mr. J. Hendra and Mr. A. M. Garbary provided major assistance in editing and preparation of the camera-ready copy. We would also like to thank the authors for their patience and c9Qperation during the production of these proceedings. Finally we would like to thank NATO who provided the primary financial assistance towards travel and accomodation for the symposium contributors, and the Natural Sciences and Engineering Research Council of Canada who provided additional funds, through a Conference Grant, for support of particular individuals. David J. Garbary G. Robin South May, 1990 TABLE OF CONTENTS Part 1. Philosgphical and Methodological Concerns The framing of biogeographic hypotheses 1. Ball. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . .. 1 Quantitative methods and biogeographic analysis Pierre Legendre. . ... . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Part. 2. The Pleistocene and its Aftennath Late glacial and Holocene diatom successions in the Gulf of Maine: response to climatologic and oceanographic change D. Schnitker and J. B. Jorgensen. .......................... 35 Seaweed biogeography of the North Atlantic: where are we now? C. van den Hoek and A. M. Breeman. . . . . . . . . . . . . . .55 Part 3. Evolutionary Ecology and Adaptation Behavioural, morphological and genetic changes in some North Atlantic populations of the barnacle Semibalanus balanoides E. Bourget, N. Martel, L. Lapointe, and D. Bussieres. ................ 87 Herbivory and evolution of nongeniculate coralline algae (Rhodophyta,Corallinales) in the North Atlantic and North Pacific R. S. Steneck ....................................1 07 Part 4. Approaches to Speciation The application of DNA-DNA hybri~zation in seaweeds and DNA divergence between some amphi-Atlantic Cladophora (Cladophorales, Chlorophyta) species P.V.M. Bot and C. van den Hoek. . . . .......................1 31 Genetics of morphology and growth in Laminaria from the North Atlantic Ocean -implications for biogeography B. Egan, Z. Garcia-Ezquivel, B. H. Brinkhuis and C. Yarish ..............1 47 Sexual compatability and species concept in Ectocarpus siliculosus (Ectocarpales, Phaeophyceae) from Italy, North Carolina, Chile, and New Zealand B. Stache ......................................1 73 x Part 5. Kelp Evolution The distribution and evolution of the Laminariales: North Pacific -Atlantic relationships Klaus LUning and Inka tom Dieck. ..........................1 87 Molecular evolution in the Laminariales: a review L. D. Druehl .................................... 205 Part 6. Taxonomy and Biogeography Evolutionary biogeography of the North Atlantic Antitharnnioid algae A. Athanasiadis . . . . . . . . . . . . . . . . . . . . . . . . . . 219 Distribution and evolution of non-coralline crustose red algae in the North Atlantic Christine A. Maggs. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . 241 A preliminary phylogenetic analysis of the Phyllophoraceae, Gigartinaceae and Petrocelidaceae (Rhodophyta) in the North Atlantic and North Pacific M. D. Guiry and D. J. Garbary . . . . . . . . . . . . . . . . . . . . . . . . . . . 265 Part 7. Biogeographic Provinces Algal provinces in the North Atlantic -do they exist? I. Tittley, G.L.J. Paterson, P.J.D. Lambshead and G. R. South. . . . . . ..... 291 Part 8. Evolution of Floras in the Arctic and Atlantic Oceans Role of the Arctic Ocean as a bridge between the Atlantic and Pacific Oceans: fact and hypothesis R. T. Wilce . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323 Biogeography of the marine red algae of the North Atlantic Ocean M. H. Hommersand . . . . . . . . . . . . . . . . . . . . . . .349 Taxonomic Index • • • 411 Subject Index • . 423

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Algal systematists, geologists and evolutionary biologists provide a synthesis of the evolutionary biogeography of red, brown, and green algae of the North Atlantic Ocean also considering their relationships with species and genera in the Arctic and Pacific Oceans as well as other subtropical and tr
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