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Hierarchical Classification and Characterization of the Protists NENCKI INSTITUTE OF EXPERIMENTAL BIOLOGY VOLUME 33 NUMBER 1 WARSAWhttp://rcin.org.pl, POLAND 1994 ISSN 0065-1583 Polish Academy of Sciences Nencki Institute of Experimental Biology ACTA PROTOZOOLOGICA International Journal on Protistology Editor in Chief Jerzy SIKORA Editors Hanna FABCZAK and Anna WASIK Managing Editor Małgorzata WORONOWICZ Editorial Board Andre ADOUTTE, Paris Stanisław L. KAZUBSKI, Warszawa Christian F. BARDELE, Tübingen Leszek KUZNICKI, Warszawa, Chairman Magdolna Cs. BERECZKY, Göd John J. LEE, New York Jacques BERGER, Toronto Jiri LOM, Ćeske Budejovice Y.-Z. CHEN, Beijing Pierangelo LUPORINI, Camerino Jean COHEN, Gif-Sur-Yvette Hans MACHEMER, Bochum John O. CORLISS, Albuquerque Jean-Pierre MIGNOT, Aubiere Gyorgy CSABA, Budapest Yutaka NAITOH, Tsukuba Isabelle DESPORTES-LIVAGE, Paris Jytte R. NILSSON, Copenhagen Stanisław DRYL, Warszawa Eduardo ORIAS, Santa Barbara Tom FENCHEL, Helsing0r Dimitrii V. OSSIPOV, St. Petersburg Wilhelm FOISSNER, Salsburg Igor B. RAIKOV, St. Petersburg Vassil GOLEMANSKY, Sofia Leif RASMUSSEN, Odense Andrzej GRĘBECKI, Warszawa, Vice-Chairman Michael SLEIGH, Southampton Lucyna GRĘBECKA, Warszawa Ksenia M. SUKHANOVA, St. Petersburg Donat-Peter HÄDER, Erlangen Jiri VAVRA, Praha Janina KACZANOWSKA, Warszawa Patricia L. WALNE, Knoxville Witold KASPRZAK, Poznań ACTA PROTOZOOLOGICA appears quarterly. © NENCKI INSTITUTE OF EXPERIMENTAL BIOLOGY, POLISH ACADEMY OF SCIENCES Printed at the MARBIS, 60 Kombatantów Str., 05-070 Sulejówek, Poland Front cover: Vauchomia nephritica Corliss, 1979. In: The Ciliated Protozoa. 2ed. Pergamon Press, London. http://rcin.org.pl ACTA Acta Protozoologica (1994) 33: 1 - 51 PROTOZOOLOGICA An Interim Utilitarian ("User-friendly") Hierarchical Classification and Characterization of the Protists John O. CORLISS Albuquerque, New Mexico, USA Summary. Continuing studies on the ultrastructure and the molecular biology of numerous species of protists are producing data of importance in better understanding the phylogenetic interrelationships of the many morphologically and genetically diverse groups involved. Such information, in turn, makes possible the production of new systems of classification, which are sorely needed as the older schemes become obsolete. Although it has been clear for several years that a Kingdom PROTISTA can no longer be justified, no one has offered a single and compact hierarchical classification and description of all high-level taxa of protists as widely scattered members of the entire eukaryotic assemblage of organisms. Such a macrosystem is proposed here, recognizing Cavalier-Smith's six kingdoms of eukaryotes, five of which contain species of protists. Some 34 phyla and 83 classes are described, with mention of included orders and with listings of many representative genera. An attempt is made, principally through use of well-known names and authorships of the described taxa, to relate this new classification to past systematic treatments of protists. At the same time, the system will provide a bridge to the more refined phylogenetically based arrangements expected by the turn of the century as future data (particularly molecular) make them possible. The present interim scheme should be useful to students and teachers, information retrieval systems, and general biologists, as well as to the many professional phycologists, mycologists, protozoologists, and cell and evolutionary biologists who are engaged in research on diverse groups of the protists, those fascinating "lower" eukaryotes that (with important exceptions) are mainly microscopic in size and unicellular in structure. Key words. Protists, algae, fungi, protozoa; macrosystematics of the Eukaryota: kingdoms, phyla, classes, orders, and representative genera. INTRODUCTION eukaryotic algae, and "lower" fungi) have been in a state of great activity, with an increasing number of biologists During the past 20 years, studies on the systematics becoming interested in such research. As our knowledge and evolution of the protists (essentially all protozoa, has grown concerning the cytoarchitecture and the phylogenetic interrelationships of the large number of Address for correspondence: J. O. Corliss, P. O. Box 53008, species (and their higher taxa) involved, so has our Albuquerque. New Mexico 87153, USA. understanding with respect to the most "natural" scheme This paper is dedicated to the memory of Zdzisław Raabe of classification to employ for these ubiquitous and (1909-1972) who, 30 years ago, perceptively foresaw the prob- cosmopolitan eukaryotic - generally microscopic and lems involved in constructing a macrosystem for the Protozoa based on evolutionary principles (Raabe 1964a) and who, in the often unicellular - organisms. In recent years, with the same year, published a comprehensive protozoological textbook development of molecular chronometric techniques (e.g. ! (Raabe 1964b) and, in the preceding year, had founded the international journal Acta Protozoologica. ribosomal RNA sequencing: see Christen 1992), com- http://rcin.org.pl 2 J. O. Corliss bined with ultrastructural investigations and the applica- (Wiley 1981). But such assemblages, so identified, are tion of sophisticated cladistic analyses, the outlook is sometimes used here. I am in sympathy with Raabe's even more auspicious for our learning enough about the (1964a) observation of 30 years ago that it is "not evolution of protistan groups to be able to propose a necessary nor possible to follow the rules of a strict robust classification system that will withstand the test purism as to the monophyletism of groups in of such phylogenetic principles as monophyly and can protozoological systematics". thus be expected to endure for a reasonable number of Scattered controversial matters (ever present in taxo- years. nomy !) are arbitrarily resolved in place, sometimes pure- At the present time, however, we are taxonomically ly by intuition, but with due attention to priority, in a state of flux. We are frustratingly trapped between common sense, courtesy, and stability (Corliss 1972), as existing classifications of protists that are recognized to well as to (my interpretation of) the facts available in be faulty and some future scheme(s) not yet available. the case. I agree with Silva (1984) that a considerable The latter, hopefully closer to the ideal natural system degree of subjectivity is inevitable when anyone at- long awaited, probably will not be ready for at least tempts to construct a macrosystem for a large and several years yet, perhaps not until the turn of the diverse group of organisms, many poorly known, no century. matter what approaches or principles are used nor how This paper represents an earnest attempt to fill in that conscientiously they are followed. long time-gap between available classifications. I In the present effort, I acknowledge my dependence believe that there is a pressing need now for a use- on the insightful analyses of Cavalier-Smith (1981, ful/usable interim system treating the protists overall in 1986, 1989b, 1991b, 1993a-c). He has provided the a manner understandable to the general protozoolo- evolutionary framework for much of the classification gist/phycologist/mycologist and the myriads of cell and presented on the following pages. But I make no attempt evolutionary biologists, biochemists, and general to utilize most of his proposed intermediate-level group- biologists (including students and teachers as well as the ings, his frequently interposed sub-, infra-, and supra- many professional researchers) who use or talk about taxa at kingdom, phyletic, class, and ordinal levels, these fascinating yet often neglected eukaryotes the although I appreciate their value to him. My system also described species of which may have already reached differs from his in other ways (e.g. I have fewer the respectable number of 200,000. I am presenting a protozoan but more chromist phyla, and I cover the compact "user-friendly" taxonomic scheme that is built fungal and plant protists as well); and I offer more along traditional lines but also incorporates the latest detailed comparative descriptions of groups and list ultrastructural and molecular data that are available. many more representative genera for each major taxon A major aim here is to offer the higher protistan taxa covered. As a born "splitter", Cavalier-Smith may indeed in a standard hierarchical arrangement, even if a measure have been guilty of a degree of "taxonomic inflation" in of speculation or presumption concerning relationships his classifications. I have tried to avoid this myself; and must sometimes be invoked to do this. The desired result it may be noted that no taxa are erected as new in this should be a system conveniently understandable to more paper. As our knowledge increases, however, newly than just a few specialists. Finally, in order to link the discovered significant differences between organisms or present with both the past and the future, a deliberate groups of organisms often require their greater, even effort is made to preserve groups and names of groups much greater, separation taxonomically than was pre- either familiar from past classifications or potentially viously given to them. In my scheme, for example, I familiar (or at least palatable!) in cases of certain newer have endorsed far more phyla and classes than appeared groups or names that I have adopted from research in the Levine Report of 14 years ago (Levine et al. 1980). papers of current workers in fields that impact on protis- For details,
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