The Ciliated Protozoa The Ciliated Protozoa Characterization, Classification, and Guide to the Literature Third Edition Denis H. Lynn University of Guelph Canada Denis H. Lynn Department of Integrative Biology University of Guelph Guelph, Ontario Canada [email protected] Author’s notes: With the exception of the Glossary (Chapter 2) and The Ciliate Taxa (Chapter 17) all chapters have been rewritten for the third edition. The Glossary and The Ciliate Taxa, originally from the second edition by John O. Corliss, have been extensively revised and considerably expanded. Original illustrations of ciliate taxa drawn by Owen Lonsdale and reconstructions of the somatic cortex illustrated by Jennie Knopp are contained throughout the book, except in Chapter 2. The author acknowledges that there are instances where he was unable to trace or contact the copyright holder for permission to reproduce selected material in this volume. The author has included complete source references for all such material. If notifi ed, the publisher will be pleased to rectify any errors or omissions at the earliest opportunity. All figures are available for free viewing via the book’s homepage at www.springer.com ISBN 978-1-4020-8238-2 e-ISBN 978-1-4020-8239-9 DOI: 10.1007/978-1-4020-8239-9 Springer Dordrecht Heidelberg London New York Library of Congress Control Number: 2008923552 Originally published by Pergamon Press, New York, 1979 © Springer Science + Business Media B.V. 2008, Corrected printing 2010 No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfi lming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifi cally for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Cover illustration : A phylogenetic tree of the 11 classes of ciliates Cover image of marsh © 2007 JupiterImages Corporation Printed on acid-free paper 9 8 7 6 5 4 3 springer.com To my parents, Beverley and Blanche Lynn, whose faithful support and unfailing encouragement were only stopped by their tragic and untimely deaths. “Some come my friend be not afraid, we are so lightly here. It is in love that we are made, in love we disappear.” Leonard Cohen Acknowledgements To my three mentors: Jacques Berger, Eugene B. Small, and John O. Corliss Jacques Berger (1934–1995) Jacques was my “academic father”, who I met with his research crew in the summer of 1968 while I was investigating the autecology of mos- quito larvae in Newfoundland, working for Marshall Laird. Jacques’ sense of humor, his strong, well−founded opinions, his broad and deep academic interests in ciliates, baseball and golf, and in later years the history of science, attracted me to the academic life at a time when I was considering a career in medicine. Eugene B. Small Gene, who I consider my “academic uncle”, has been a colleague, with boundless enthusiasm for the ciliates and life, always willing to engage in discussion on topics as arcane as “the number of ciliated kinetids in Kinety X” to the spiritual elements of Zen. As a mentor, Gene generously shared both his deep intuitive insights into the taxo- nomic significance of the cortical characters of ciliates and his broad knowledge of ciliate diversity. Over the years, our collaboration has been inspirational and deeply rewarding. John O. Corliss John, my “academic grandfather” as Jacques Berger completed his Doctoral dissertation with John in 1964, has always been supportive of my career in ciliatology. John has been a professional role model by demonstrating outstanding devotion to students and colleagues both far and wide, and to one’s discipline. His willingness in 1997 to begin our collaboration on a Third Edition demonstrated his enthusiasm for “our wee beasties” and his confidence in our partnership. For his trust, encouragement, and big-hearted generosity, I am sincerely grateful. vii Preface to the Third Edition Soon after I began graduate studies in Protozoology proposed phylogenetic scheme of classification at the University of Toronto in September 1969, for the protistan phylum Ciliophora”. This was the Jacques Berger brought in his copy of the First “Age of Ultrastructure,” as John called it, but the Edition of “The Ciliated Protozoa” as required “newly proposed phylogenetic scheme” was only reading. This book synthesized the “state of knowl- moderately influenced by these new data. edge” of ciliate systematics at that time, and While in Maryland, Gene Small and I became it brought the formal study of ciliate diversity, deeply involved in discussing the implications of especially in its nomenclatural aspects, to a highly ultrastructural features, and these discussions lead professional level. In the following decade events to my publication of “the structural conservatism occurred that set me on the path to pursuing ciliate hypothesis” in 1976. Applying that idea, Gene and research. John Corliss, the author of that little-big I proposed a radically different macrosystem for ciliate book visited Jacques in Toronto, and I met the ciliates in 1981, which I supported by a major him. John suggested that I visit Gene Small at the review of the comparative ultrastructure of ciliate Department of Zoology, University of Maryland, kinetids, demonstrating the conservative nature USA. In 1971, I met Gene, whose enthusiasm for of these important cortical components. While “these wee bugs” was infectious, and whose intui- ultrastructural study still formed an element of my tive grasp of the systematic significance of particu- research program in the 1980s, Gene encouraged lar features was marvellous. I resolved to return to me to consider moving into molecular phylogenet- Maryland to work with Gene, taking a “sabbatical” ics to test the robustness of our ideas, which had leave from my doctoral thesis research to do so. now been slightly modified with publication of There was, of course, another wonderful reason for the First Edition of “An Illustrated Guide to the the move to Maryland in September 1972 – I had Protozoa”. In an ultimately productive sabbatical met Dr. Portia Holt who, at the time, was working year in 1986–1987, I worked with Mitch Sogin as a postdoctoral fellow with Dr. Corliss. So the at the National Jewish Hospital, Denver, to learn 1972–1973 period was a rich experience of immer- the techniques of cloning and sequencing. Mitch sion in ciliate systematics, coupled with immersion and I were finally able to provide one of the first in my developing relationship with my future wife, larger comparative datasets on genetic diversity Portia. During this time, John Corliss, Head of the of ciliates based on the small subunit rRNA gene Department of Zoology, provided financial assist- sequences, derived at that time by reverse tran- ance as well as academic support. At that time, scriptase sequencing. On the other side of the John was beginning preparations for the Second Atlantic, our colleagues in France, led by André Edition of “The Ciliated Protozoa”, having just Adoutte, were using the same approach with the co-authored a major revision of the ciliate mac- large subunit rRNA gene and generating an even rosystem with his colleagues in France. By 1974, larger dataset. Both approaches demonstrated two these co-authored and authored papers on the new things: firstly, confirmation that the ultrastructural macrosystem were published, including his paper approach informed by structural conservatism was entitled “The changing world of ciliate systemat- providing resolution of the major natural assem- ics: historical analysis of past efforts and a newly blages or clades of ciliates; and secondly, genetic ix x Preface to the Third Edition distances between groups of ciliates were as vast as significant hurdles to obtain copyright permissions the genetic distances between plants and animals for the over 1,000 required illustrations, and I put – THE major eukaryotic kingdoms at that time! the publication schedule ahead of this element. I continued to collaborate with Mitch, and in There are a number of significant illustrated guides 1991 my first “molecular” Magisterial student, to genera and species that have recently been pub- Spencer Greenwood, published an article estab- lished. References are made to these throughout lishing 1990 or thereabouts as the beginning of the book as sources that readers can consult for this the “Age of Refinement” – the period when gene aspect of ciliate diversity. A future project that I am sequencing techniques would deepen our under- contemplating is an illustrated guide to all the valid standing of the major lines of evolution within ciliate genera. the phylum. Nearing the end of that decade, I was This book has been a collaborative effort from fairly confident that we had resolved the major the beginning. In addition to my indebtedness to lines of evolution, mostly confirming the system John, I have appreciated the support provided by that Gene and I had proposed in the mid-1980s. We my new contacts at Springer – Dr. Paul Roos, published a revisionary paper in 1997 in the Revista Editorial Director, Environmental/Sciences, and as a tribute to our Mexican colleague, Eucario Betty van Herk, his Senior Assistant. Since the Lopez-Ochoterena. Third Edition depended heavily on several sec- As I look back on my correspondence, it was about tions from the Second Edition – notably the this time that I approached John regarding writing a Glossary and The Ciliate Taxa , I was helped Third Edition of “The Ciliated Protozoa”. While turn- immensely by the secretarial assistance of Lori ing “just” 75 in 1997, John enthusiastically embraced Ferguson, Felicia Giosa, Irene Teeter, and Carol the idea and we began collaborating on a book Tinga, who created electronic files of Chapters 2, proposal that travelled with several editors through 20, and 22 from the Second Edition.