Robert Riding (Ed.) Calcareous Algae and Stromatolites With 16 Plates and 202 Figures Springer-Verlag Berlin Heidelberg New York London Paris Tokyo Hong Kong Barcelona Dr. ROBERf RIDING University of Wales College of Cardiff Cardiff CFt 3YE United Kingdom ISBN 978-3-642-52337-3 ISBN 978-3-642-52335-9 (eBook) DOI 10.1007/978-3-642-52335-9 Library of Congress Cataloging in Publication Data. Calcareous algae and stromatolites / R. Riding [editor). p. cm. Includes articles originally presented at the 4th International Symposium on Fossil Algae, held in Cardiff, July 1987. Includes index.!. Algae, Fossil - Congresses. 2. Stromatolites - Congresses, I. Riding, Robert. II. International Symposium on Fossil Algae (4th: 1987: Cardiff, Wales) QE955.C338 1990 561'.93 - dc20 90-10112 This work is subject to copyright. 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Typesetting: International Typesetters Inc., Makati, Philippines 32/3145(3011)-543210 - Printed on acid-free paper Preface "Calcareous algae and stromatolites" is shorthand for a wider array of organisms and fabrics that also includes calcified cyanobacteria, plus thrombolites and other microbial carbonates. Composition is the link: these are all important components of CaC0 sediments, from 3 Archaean to present and from the ocean floor to streams and lakes. It is hardly possible to examine limestones of any age without en countering them. Simultaneously they are fossils, sediments, and en vironmental indicators. It is the range of significance, coupled with the breadth of their distribution in time and space, which compels their study. Modern calcareous marine algae mainly include reds (corallines, squamariaceans, and the nemalialean Galaxaura) and greens (dasy cladaleans, udoteaceans, halimedaceans). Blue-greens, of course, are cyanobacteria and not algae, and significantly, although they are largely responsible for Recent tidal flat stromatolites, they are not calcified in the same way that pre-Cenozoic marine blue-greens are. It is in the freshwater environment of calcareous streams and lakes that we find modern calcified cyanobacteria, and they are commonly associated with the only major group of non-marine calcareous algae, the charophytes. However, in the past, and especially in the Palaeozoic and Mesozoic, things look radically different. Mingling with the ancestors of the modern flora are distinct, and often problematic, organisms. Their organization could be consistent with an algal or cyanobacterial affinity, and they have been treated as such; but sometimes they hard ly seem to fit conceptions founded in the Recent. This is another world, and it informs and confuses at the same time. Most reported calcareous "algae" in the Cambrian are actually more likely to be cyanobacteria, and are especially important in reefs. Never again, except briefly in the Upper Devonian, do these forms assume such importance in marine environments. Dasycladaleans and halimedacean-like greens, strikingly similar to Recent forms, ap pear in the Ordovician. But any semblance of modernity is destroyed by other newcomers which raise possibilities and problems for algal evolution which are still unresolved. For the greens there are Ordovi cian receptaculitids and cyclocrinitids to be considered, together with Late Palaeozoic "green" phylloids, and also "erect udoteaceans", VI Preface which so often resemble cyanobacteria. For the reds there are Soleno poraceae and Moniliporellaceae to be taken into account in the Mid dle Palaeozoic, and these are augmented by "ancestral coralline" and squamariacean-like phylloid algae, as well as by Gymnocodiaceae, in the Late Palaeozoic. Along with these a host of more minor proble matica, which for better or worse have been, entrusted to palaeolago logists for study, create a systematists jungle in the Carboniferous and Permian. End-Palaeozoic extinctions failed to eliminate this disorder. Dur ing the Mesozoic, despite dasycladalean diversification and the ap pearance of coralline red algae, the undercurrent of problematic forms is still evident, with bizarre newcomers mingling with survivors from the Late Palaeozoic. No less do stromatolites present problems and show dramatic changes with time. The striking differences between Recent forms and those of the Precambrian are often glossed over. More than most other fossils, stromatolites exist at two levels in the minds of those who study them. On the one hand, there are the tangible deposits: laminated layers, domes, and columns which are fairly easy to discern in isolation but whose clotted, spongy, sometimes filamentous, micro fabrics also often, since the Early Cambrian, lurk obscurely in skeletal reefs. On the other hand, there are the shifting dogmas of ex planation: environment of origin (freshwater, intertidal, subtidal); mi crobial community (cyanobacterial, bacterial, algal); process of for mation (trapping and binding, calcification); reason for decline (ox ygen levels, invertebrate grazing, coarse bioclastic sediment); reason for survival (saline refuges and desiccated tidal flats). Descriptive em phases have also varied: geometric or Linnean classifications, macro structures or micro fabrics. The problem with stromatolites is not their complexity, but their simplicity. They offer so little to grasp. But, like Plautus's Pseudolus, we so often seem to have overlooked the certainties while seeking uncertainties. Stromatolites are just the tip of a microbial carbonate iceberg: lamination varies and may disappear; there are gradations between clotted and peloidal spongiostrome fabrics, porostromate fabrics, and macroscopically clotted thrombolite fabrics; calcification or cement encrustation of the microbes may radically alter the ap pearance of the deposit. The clues have always been there in the car bonate fabrics. The challenge is not only to correctly interpret them but also to objectively and accurately describe them. The incentive for this book came about because of the 4th Interna tional Symposium on Fossil Algae, held at Cardiff in July 1987. But I decided early on that there was much more need for an overview of the subject as a whole, than for a compilation of conference papers. Only just over half of the articles here were originally presented as talks at the meeting, so this is not strictly a symposium volume. Preface VII Nevertheless, were it not for the stimulus of the Symposium this book would not have appeared. The title of the Fossil Algae Symposium is an overstatement. The four meetings so far (Erlangen 1975, Paris 1979, Denver 1983, Cardiff 1987) have mainly attracted researchers interested in benthic calcified algae, cyanobacteria, stromatolites, and other microbial carbonates. But even within this much more limited area there is an enormous variety of information and it is very scattered. Calcareous algae and prokaryotes have yet to benefit from the kind of general synthetic treatment applied to invertebrate fossils, on the one hand, and to vas cular plant fossils, on the other. Precisely because of this, it is worth trying to draw together a review volume. I cannot claim to have achieved anything close to an even, comprehensive coverage. What has emerged, however, does demonstrate the breadth of the subject, the variety of approach, and the state of the science at present. Despite omissions due to con straints on length or non-availability of authors, this volume does in clude discussions of major groups of algae and cyanobacteria, stromatolites, and other microbial carbonates, of all ages, both extant and extinct, marine and non-marine and, where appropriate, benthic and planktonic. I am very grateful indeed to the following colleagues who gener ously helped by giving their time and expertise to referee manuscripts: Stanley M. Awramik, Jack A. Babcock, Michael A. Borowitzka, Can dace M. Brooke, Yvonne M. Chamberlain, Barrie Dale, Graham F. Elliott, Monique Feist, Erik Flugel, Richard A. Fortey, Patrick Genot, Nicole Grambast-Fessard, John R. Groves, Llewellya Hillis-Colin vaux, Hans 1. Hofmann, David M. John, Andrew R. Leitch, Bernard L. Mamet, Xi-nan Mu, Ernst Ott, Allan Pentecost, Katharina Perch Nielsen, Brian R. Pratt, Alain Roux, Daniel Vachard, Malcolm R. Walter, V. Paul Wright. It is a pleasure to thank the authors for their willingness to con tribute to this volume, and for their patience with my suggestions. The skill and care of the staff of the Springer Geosciences editorial office in Heidelberg brought the whole to fruition. Cardiff, August 1990 ROBERI' RIDING Contents I Introduction...................................... 1 1 Calcification Processes in Algae and Cyanobacteria A. Pentecost (with 4 Figures) ....................... 3 2 Classification of Microbial Carbonates R. Riding (with 11 Figures) ......................... 21 II Major Groups .................................... 53 3 Calcified Cyanobacteria R. Riding (with 15 Figures) ......................... 55 4 The Solenoporaceae: A General Point of View A.-F. Poignant .................................... 88 5 Coralline Algae: Mineralization, 'Thxonomy, and Palaeoecology D. W. J. Bosence (with 3 Figures) .................... 98 6 Cyclocrinitids S. C. Beadle (with 4 Figures) ........................ 114 7 Dasycladalean Algae of the Palaeozoic and Mesozoic G. F. Elliott ....................................... 125 8 Cenozoic and Recent Dasycladales P. Genot (with 43 Figures) .. . . ... . . . .. .. .... . . . . . . .. 131 9 Fossil Udoteaceae and Gymnocodiaceae x. Mu (with I Plate and 3 Figures) .................. 146 10 Recent Calcified Halimedaceae L. Hillis (with 12 Figures) .......................... 167 11 The Genus Concept in Charophyta: Evidence from Palaeozoic to Recent M. Feist and N. Grambast-Fessard (with 6 Figures) 189 12 Calcification of the Charophyte Oosporangium A. R. Leitch (with I Plate and 2 Figures) ............. 204 13 Calcareous Nannofossils S.D. Houghton (with 19 Figures) .................... 217 14 Fossil Calcareous Dinoflagellate Cysts H. Keupp (with 17 Figures) ......................... 267 x Contents III Algae and StromatoUtes Through Time .............. 287 15 Archaean and Proterozoic Stromatolites S. M. Awramik (with 7 Figures) ..................... 289 16 Cambrian Calcareous Cyanobacteria and Algae R. Riding (with 6 Figures) .......................... 305 17 Ordovician Algae and Global Thctonics A. Roux (with 5 Figures) ........................... 335 18 Ordovician to Devonian Marine Calcareous Algae A. Roux (with 6 Figures) ........................... 349 19 Carboniferous Calcareous Algae B. Mamet (with III Plates and 3 Figures) ............. 370 20 Permian Marine Calcareous Algae R. Riding and L. Guo (with 22 Figures) ............. , 452 21 Triassic and Jurassic Marine Calcareous Algae: A Critical Review E. Flugel (with 8 Figures) .......................... 481 22 Mesozoic and Cenozoic Marine Benthic Calcareous Algae with Particular Regard to Mesozoic Dasycladaleans F. Barattolo (with VII Plates and 6 Figures) .......... 504 23 Modern Stromatolites: A Review S. Golubic (with II Plates) .......................... 541 Index ................................................ 563 List of Contributors Awramik, S. M. 289 Hillis, L. 167 Barattolo, F. 504 Houghton, S. D. 217 Beadle, S. C. 114 Keupp, H. 267 Bosence, D. W. J. 98 Leitch, A. R. 204 Elliott, G. F. 125 Mamet, B. 370 Feist, M. 189 Mu X. 146 Flugel, E. 481 Pentecost, A. 3 Genot, P. 131 Poignant, A.-F. 88 Golubic, S. 541 Riding, R. 21, 55, 305, 452 Grambast-Fessard, N. 189 Roux, A. 335, 349 Guo L. 452 1 Introduction
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