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Protozoan Taxonomy and Secondary article Systematics John O Corliss, University of Maryland, College Park, Maryland, USA Taxonomy

and systematics of major groups of the Protozoa, an assemblage of so-ca lled lower eukaryotes, refer basically to the classification of such groups, that is, their arrangement into a hierarchy of evolutionary interrelated groups (taxa) of scien tifically named phyla, classes, orders, etc. Article Contents .Introduction .Historical Considerations .Major Groups of Protozoa .The Old Phylum Protozoa .Modern Options Concerning the Place of Protozoa in the Biotic World .The New Kingdom Protozoa Introduction The classification of protozoa and other microorganisms above the organizational level of the bacteria has always been dependent on microscopy because the body sizes involved generally range from only one micrometre to one or

two millimetres in length. Any structures these species possess, useful in comparative studies of their morphology and thus their taxonomy and systematics, are at the cellular and subcellular levels, and invisible to the naked eye. The physiological properties of protozoa (and the neighbouring algae)have also played a role in the classification of these ubiquitous eukaryotic microorganisms; and of growing significance

are the findings made possible by research using molecular biological approaches. Because we are continuing to learn more and more about such minute organisms, protozoan systematics that is, the taxonomy (classification)and the evolutionary interrelationships of major groups of protozoa remains a topic of debate and change, still today. Some of the rather large and unwieldy taxonomic

Paradoxically.groupings of past years are particularly subject to revision with expansion of and refinement in our knowledge about the members of those and related assemblages. Thus. the protozoa themselves are becoming more difficult to define with precision as our information about them and other microbial assemblages increases. presenting a single satisfactory circumscribed definition for them is not .

As a . retaining their firstanimal definition dating from 150 years earlier. Historical Considerations Until well beyond the middle of the twentieth century. It is attempted towards the end of this article. for sake of easy task. protozoa were widely treated taxonomically as a (mere) subset of the kingdom Animalia. further background information is first supplied. but.

no photosynthetic pigments present. yet. and capable of independent locomotion.e. phagotrophic. a goodly number of chlorophyll-possessing algal groups were included in the phylum). incongruently.phylum of unicellular animals. It is now abundantly clear that this classical definition of protozoa is at best misleading and incomplete. . they were thought to exhibit major characteristics typical of that kingdom: colourless (i.

sometimes even the same species. by the botanists (phycologists and mycologists). Thus. and they often worked with taxonomic disregard for studies of what might actually be the same group.and that it requires considerable refinement. unfortunately. zoologists were the principal investigators and namers and claimers of such microorganisms. In former times. territoriality and authoritarianism also played .

As early as a century ago. Advances in our knowledge of the protozoa in general have followed progress in microscopy. algae and lower fungi (Corliss. 1986). novel improvements in methods of light microscopy and related techniques of fixing and staining were already beginning to . as mentioned above.major roles in determining the early systematic status of the protozoa.

Then. at mid-twentieth century. a myriad of . During the Age of Ultrastructure (as it has been called). the use of electron microscopical approaches in cell biology opened up a whole new epoch of exploration in protozoology and related fields.make possible the revelation of morphological cellular characteristics that would have remained totally unrecognized before the appearance of such technological advances.

has offered a refinement in taxonomic investigations that . Even more recently. large and small. the exciting development of molecular biological approaches. particularly in study of genealogical and phylogenetic relationships within all groups of organisms.previously unknown subcellular structures were revealed that became of immense value in the comparative taxonomy of the lower eukaryotes (especially protozoa and algae). plant or animal.

and from more modern apENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / .is without parallel in past decades. conventional classifications of these minute organisms.els. Here an attempt is made to review and understand the systematics of protozoa both from the more traditional point of view. keeping in mind that many biologists are still familiar mostly with older.

the reader is referred particularly to comprehensive papers by Cavalier-Smith (1993. algae and some other eukaryotic microorganisms together as Protista and by their usage of more sophisticated data. 1998) . For more detailed information concerning protozoan protistan overlapping relationships.Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics proaches. The latter are complicated by the relatively recent advent of the protistological perspective lumping protozoa. 1998)and Corliss (1994.

and to the insightful textbooks by Sleigh (1989)and Hausmann and Hu¨ lsmann (1996). as has been done in numerous textbooks of biology and . it has been convenient to assign species known as protozoa to just four major categories or assemblages. published in the 1880s). Major Groups of Protozoa For many decades (commencing with the perceptive schemes of Otto Bu¨ tschli.

in a broad sense). to arrange protozoa taxonomically in such a way.zoology as well as protozoology. including groups . for sake of discussion under diverse circumstances. Despite advances and improvements in our knowledge of eukaryotic microorganisms in general. it is still often rather helpful. The categories generally recognized are: (1)the amoeboid forms (the Sarcodina. (2) the flagellated forms (the Mastigophora.

(3) the ciliated forms (the Ciliophora. once assigned to a very broad group called the Sporozoa. some highly pathogenic to their hosts. and (4) the various totally symbiotic or parasitic forms (primarily spore-forming species that are typically endoparasites.of autotrophic or photosynthetic as well as heterotrophic species). . the most stable and perhaps most circumscribed of all protozoan assemblages).

1986). especially during the busy second half of the twentieth century (even before the protist revolution : see Corliss.a high-level taxon that subsequently became divided into the Sporozoa and the Cnidosporidia). while generally still . classes. orders)of protozoa. One of the pedagogically oft-regretted but inevitable changes. was the tremendous expansion in the total numbers of high-level taxonomic groups (subphyla.

recognizing the four major top divisions mentioned above. the classification scheme of . students and researchers alike. The Old Phylum Protozoa Basically. The discovery of new and unique differentiating characteristics useful in classification and in evolutionary and phylogenetic studies as well required such a multiplication and fragmentation of taxa even though it placed/places greater demands on (the memories of)teachers.

Although this popular arrangement in comparison with the long-followed classical one of Bu¨ tschli of 80 years earlier contained nearly four times the number of taxonomic units above the level of family. it showed .the widely accepted Honigberg Report of 1964 may be used to illustrate the conventional situation concerning the systematics of the protozoa (Table 1).

For an outstanding example of its conservatism. the Levine Report of 1980. recognized seven separate phyla (in a subkingdom Protozoa)and increased the .few novelties of great significance. all treated as comprising a single class. of flagellated protozoa (the Mastigophora). the Phytomastigophorea. the classification still retained several major groups of algae. A subsequent revision supported by the international Society of Protozoologists.

it followed the arrangement of its predecessor (Table 1). By then. the protist revolution had already been going strong for a full ..number of taxa above the familial level to 229 (Table 2). with its classification scheme mainly that of the Levine Report. Five years later. But. fundamentally. the well-known and widely used Illustrated Guide to the Protozoa (Lee et al. 1985)appeared.

besides the fact that those classifications of 20 40 years ago .decade (see accounts in Corliss. 1986. One reason for including the above discussion here. 1998)and many findings were indicating the pressing need for a fresh look at the old persisting problems of how to treat the systematics of the conventional/ traditional protozoan. algal and fungal assemblages of microorganisms.

is to help bridge the gap between such neoclassical systems and the suggested recent arrangements of the 1990s (see below). primarily . but our ideas concerning their most likely interrelationships at the higher taxonomic levels have done so. have not changed over the decades.are still accepted by many biologists. The organisms involved. and often their common group names as well.

by fresh analyses of the continuing accumulation of data of high phylogenetic and evolutionary significance from precise ultrastructural and molecular biological investigations (Coombs et al. Several of . 1998).. Modern Options Concerning the Place of Protozoa in the Biotic World There are a number of ways in which modern biologists are viewing the overall placement of the protozoa with respect to other major groups of organisms.

or considering protozoan groups as representing simply an evolutionary grade or level of cellular organization between the anucleate prokaryotes (essentially the bacteria)and the well-known higher (nucleated)eukaryotic forms of life (particularly the multicellular and multitissued plants and animals). arrangement of species of protozoa based solely on their nutritional preferences or requirements.these are not truly taxonomic in their approach: for example. .

net .And popular today.els. especially among workers employing cladistic approaches in their investigation of ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.

Ciliatea Class 3. Piroplasmea Class 1. (Modified and abbreviated from Corliss. Cnidospora (2) Filosia Class 1. Actinopodea Subclass (1) Holotrichia Subclass (1) Radiolaria (2) Peritrichia (2) Acantharia (3) Suctoria (3) Heliozoia (4) Spirotrichia (4) Proteomyxidia phylogenetic interrelationships of life forms. Teleosporea Class 1. nearly four times the number in vogue a mong earlier protozoologists well into the twentieth century. Phytomastigophorea Subclass (1) Gregarinia Class 2. Ciliophora Class 2. Sarcodina Class 3. Haplosporea Class 1. Zoomastigophorea (2) Coccidia Superclass 2. Rhizopodea Subclass (1) Lobosia Subphylum III. distinct evolutionary lines or lineages but ones often with unknown or nonpostulated possible taxonomic relationships with other lines of organisms . Mastigophora Class 1. 1998. The total number of suprafamilial categories reached 140.Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics Table 1 Taxonomic classification of the single phylum Protozoa according to the Report published by Honigberg et al. is the idea of clades. Sporozoa Superclass 1. (1964). but omitting ordinal and subordinal names. Myxosporidea (3) Granuloreticulosia Class 2. Sarcomastigophora Subphylum II. Microsporidea (4) Mycetozoia (5) Labyrinthulia Subphylum IV. Opalinata Class 2. Toxoplasmea Superclass 3.) Subphylum I.

Recognition of a single kingdom . The only true taxonomic options today are the inclusion of the protozoa with other eukaryotic microorganisms in a single kingdom called Protista or the recognition of a number of kingdoms (one called the Protozoa)within the overall assemblage of the Eukaryota. These two approaches require further discussion. 1999).(microorganisms or macroorganisms)(see Patterson.

Haeckel and others. still popular today among a number of biologists. the . divisions and classes of organisms formerly representing the protozoa. first voiced more than 120 years ago by E. In this contain all eukaryotic microorganisms (and some of their larger but yet basically unicellular or single-tissued relatives)is a choice. a neoHaeckelian kingdom Protista (or Protoctista )contains commingled phyla.

Plantae and Animalia. and of the (other) eukaryotic moieties known as the Fungi. on the one hand. on the other hand (Margulis et al. 1998). It is given equal standing with the neighbouring kingdoms of the prokaryotic Bacteria.algae and taxa of lower fungi. This five-kingdom arrangement represents a pragmatic one that is quite . Margulis and Schwartz. 1990..

attractive pedagogically (easy to teach at all levels in the educational hierarchy)and is convenient for general nonspecialist usage and for information retrieval systems. the single kingdom Protista hypothesis lumps together lines of microscopic organisms that are proving to be not very closely related evolutionarily and . But it is becoming evident that. as our store of relevant data continues to expand.

overall.includes still other lines that are being shown to have closer affinities with forms assigned to one of the other three eukaryotic kingdoms. The second major alternative available to us today is to consider the protists. And the protozoa are not sufficiently given their own identity as an independent high-level taxonomic assemblage. as best assignable to a .

Plantae. 2000) is considered below . 1998. Corliss. 1998. 1994). Corliss. 1993. the Chromista (taxonomic home for most lines of algae).number of separate kingdoms. The composition of this protozoan assemblage distinctly separated from the four other eukaryotic assemblages or kingdoms recognized. one of which has now been labelled with the familiar name Protozoa (Cavalier-Smith. Fungi and Animalia (see Cavalier-Smith.

(1)A formal high-level taxonomic category called Protista is no longer needed or ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / .in further detail. Two points need emphasis here.els.

Ciliophora (4) Heliozoea Class (1) Kinetofragminophorea (2) Oligohymenophorea (3) Polyhymenophorea recognized. but here omitting all categories below the ra nk of class. Ascetospora (5) Plasmodiophorea Class (1) Stellatosporea (6) Filosea (2) Paramyxea (7) Granuloreticulosea (8) Xenophyophorea Phylum VI. (Modified and abbreviated from Corliss. Microspora Class (1) Lobosea Class (1) Rudimicrosporea (2) Acarpomyxea (2) Microsporea (3) Acrasea (4) Eumycetozoea Phylum V.Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics Table 2 Taxonomic classification of the animal subkingdom Protozoa according to the Report published by Levine et al.) Phylum I. Apicomplexa Subphylum 2. 1998. Actinopoda Class (1) Myxosporea Class (1) Acantharea (2) Actinosporea (2) Polycystinea (3) Phaeodarea Phylum VII. nearly double the number endorse d in the preceding Honigberg Report of 1964 (see Table 1). Rhizopoda Phylum IV. (1980). Opalinata Class (1) Perkinsea Class (1) Opalinatea (2) Sporozoea Subphylum 3. Mastigophora Class (1) Labyrinthulea Class (1) Phytomastigophorea (2) Zoomastigophorea Phylum III. Myxozoa Superclass 2. The total number of suprafamilial categories reached 229. Sarcomastigophora Phylum II. all former protists (spelled with a lower case p )can be reassigned to various of the . Labyrinthomorpha Subphylum 1. Sarcodina Superclass 1.

(2)The (new)kingdom Protozoa. while containing many conventional groups of protozoa. see also Honigberg et al.. see preceding sections of this article. 1964. . is purged of others and thus is not identical in overall taxonomic composition to the old phylum Protozoa embraced in very popular classifications of both distant and more recent past years (for example.five kingdoms mentioned immediately above.

The New Kingdom Protozoa Of the two options briefly described above. 1980.. the concept of dividing up major taxa of former protozoa (and of algae and lower fungi.and Levine et al. with new assignments to several different kingdoms. and contrast the contents of Table 1 and Table 2 with that of Table 3). many biologists now favour the second one. one of which . too).

Cavalier-Smith. 1993. The kingdom Protozoa may be briefly described (after Corliss. plasmodial or colonial protists (eukaryotic microorganisms) that are . The kingdom comprises predominantly unicellular. starting in the 1980s (see full discussions in Cavalier-Smith. 1998). 1994. The first person to do this systematically and in some detail was identified as the kingdom Protozoa. 2000)as follows.

when mitochondria are absent. Nearly universally present are tubular mitochondrial cristae. no stored starch. lacking cellulosic cell walls.mostly phagotrophic or osmotrophic. colourless. and surrounded by three membranes. they are typically replaced by hydrogenosomes. Included species that are capable of photosynthesis (but some are nutritionally mixotrophic) typically possess unique cytosolic plastids with stacked thylakoids. and microscopic in body size. Golgi .

Flagellar mastigonemes. As one of the two (of five)eukaryotic kingdoms ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.bodies and peroxisomes are widely present. Free-living (typically independently motile via pseudopodia. or cilia) and symbiotic species are numerous from a great variety of habitats. if . flagella.els. are never rigid or tubular.

with endosymbiotic . 1818. 1994.Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics Table 3 The 14 phyla. amitochondriate. 2000) Archamoebae Cavalier-Smith. including authorships and dates of their names. 1983 Large. benthic. microaerobic amoebae. allegedly primitive forms. comprising the kingdom Protozoa Goldfuss. 1998. with an indication of the kinds and numbers of protists included in each phyletic taxon (based on abbreviated characterization data from Corliss.

marine heterotrophic flagellates and amoeboflagellates. naked or with tests or thecae. free-living. all free-living in fresh water Neomonada Cavalier-Smith. some flagellated forms. small group.bacteria. 1845 Typically amoeboid. 1997 Often small. species few in number. fresh-or saltwater habitats . with differing kinds of pseudopodia. 4 5000 species found in soil. still ill-defined Rhizopoda von Siebold.

850 species. with alternation of haploid sexual and diploid asexual generations.Mycetozoa de Bary. shells of extinct species reach 15 cm in diameter. some very large. c. a few symbiotic forms Foraminifera d Orbigny. 1859 Plasmodial slime moulds (cellular and acellular). mostly in decaying vegetation. aerial (stalked) fruiting bodies produce spores. reticulate . 1826 Amoeboid forms in tests (usually calcareous).

with some 45 000 species (largest phylum in the kingdom)of which c. c. with slender radiating axopodial type of pseudopodia used in food capture. spherical forms. . 90% are fossil forms Heliozoa Haeckel. 1987 Marine.pseudopodia for feeding and locomotion. 100 species. many stalked Radiozoa Cavalier-Smith. 1866 Mostly freshwater group of the classical actinopod sarcodinids . mostly marine.

some poorly known Euglenozoa Cavalier-Smith. second largest phylum in the kingdom Percolozoa Cavalier-Smith. 1981 The old Euglenophyta . 100 species. with total of nearly 12 000 species (c. freshwater phytoflagellates . mainly free-living. 4 .typically planktonic. 1991 Small heterotrophic flagellates or amoeboflagellates. c. often with elaborate symmetrical shell pierced by stiff axopodia. three major subgroups. 65% fossil forms).

plus Kinetoplastidea (parasitic trypanosomes plus free-living bodonids). 4500. commonly with discoidal mitochondrial cristae and a paraxial rod in their main flagellum Dinozoa Cavalier-Smith. total species c. some thecate. with some 100 described . mostly marine planktonic. about half found as fossils. unique biflagellated protists.1000 species. 1981 Dinoflagellates. 4 600 species. cortical alveoli. one-half pigmented forms. a few colonial.

as parasites. typically digestive tract parasites (insects to humans). c. 1970 Essentially the Sporozoa . hydrogenosomes in place of mitochondria Parabasala Honigberg. 1952 Biflagellated to multiflagellated forms. in intestines of woodroaches to humans Apicomplexa Levine. 1973 Parasitic multiflagellated forms. 300 species. 4 400 species. many orders Metamonada Grasse´ . amitochondriate and with prominent parabasal (Golgi)apparatus.

4 5000 species in three major classes Ciliophora Doflein. phagotrophic.g. in birds. livestock. unique complex of apical organelles. others symbiotic or epibiotic. usually multiciliate. malaria). relatively large protists found mostly free-living in diverse fresh-/saltwater/soil habitats. all symbiotic. mostly in . 1901 All heterokaryotic (micro-and macronuclei). cortical alveoli. with many minute species as harmful endoparasites (e.of old. humans: outstandingly.

third largest protozoan phylum: c. many exhibit sexual phenomenon of conjugation. asexual reproduction by transverse fission. cortical alveoli. the refined kingdom Protozoa. many orders composed purely of protistan forms. counting fossil as well as extant species embrace nearly . often complex oral ciliature. the Protozoa As characterized above.or on invertebrate hosts. 8000 species in 8 10 classes.

els.large though it is. . is considerably more discriminating and 40% of all protists (numbering some 213 000 species) more restricted in its boundaries than was the old phylum described to date. The latter. Protozoa. contained a number of ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.

and most recently. All species of these taxa fail to possess major distinguishing features of the kingdom Protozoa listed above and/or . the curious labyrinthomorphids. the microsporidia. For example. the choanozoa. the myxozoa. at class and ordinal levels)that have now been eliminated.Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics major taxa (e.g. the opalinids. purged from the new kingdom have been half a dozen algal groups.

1998.g. Supported by recent molecular (e. about 60% of .have unique morphological. these other protists have been placed in one or another of the neighbouring kingdoms Chromista. Fungi. physiological or molecular characteristics of their own beyond discussion here. Plantae or Animalia (Corliss. The 14 phyla (containing nearly 83 000 species. 2000). rRNA sequencing information)as well as morphological and biochemical findings.

in Table 3. with abbreviated descriptions. but names of many such genera are included in the long papers by Cavalier-Smith (1993)and Corliss (1994). for sake of brevity within the table. Representative genera are not mentioned. Several additional observations might be helpful concerning some .which are fossil and 12% symbiotic forms)that then remain in the refined kingdom Protozoa are listed.

of the major differences between this new classification of the Protozoa and the older. but they have strikingly different characteristics (including presence of three. more conventional ones discussed on preceding pages. not two. plastidic membranes and frequent exhibition of nonphotosynthetic modes of . Two distinctive eukaryotic algal lines the euglenids and the dinoflagellates are retained as Protozoa.

chlorophytes (with such long-claimed protozoa genera as Volvox and Chlamydomonas). haptomonads. phaeophytes. ulvophytes.nutrition. the euglenophytes of old are phylogenetically closely related to the colourless kinetoplastideans (bodonids and .g.g. the prasinophytes. among others)from other major assemblages of algae now appearing in the separate kingdoms Chromista (e. In fact. charophytes. the chrysophytes. rhodophytes and glaucophytes). cryptomonads and lesser heterokontic groups)and Plantae (e.

parasitic trypanosomatids). a character not particularly known in other phyla of protists. are linked with the ciliates (Ciliophora)and the sporozoa (Apicomplexa) in a supraphyletic assemblage often designated as the Alveolata because of their common possession of cortical alveoli in their pellicles. the combining single phyletic name now being the Euglenozoa. The dinoflagellates (Dinozoa). furthermore. Mostly on .

the longlinked protozoan-cnidosporidian groups. along with chytrids (but not with oomycetes and hyphochytriomycetes. turn out to be not only not closely related to each other but also not members of the kingdom Protozoa. the enigmatic microsporidia and myxosporidia. The first. the Microspora.the basis of very recent rRNA data. but also supported by other differentiating characteristics. which are now .

are probably primitive or degenerate protistan members of the kingdom Fungi. Until rather recently. while the complex Myxozoa are better classified in the Animalia (perhaps close to the cnidarians). Entamoeba)were considered as very primitive protists in a kingdom . the Archamoebae (with genus Pelomyxa and possibly others such as the important parasite of the Chromista).

of their own. Latest studies (see references in Cavalier-Smith, 1998; Corliss, 2000) suggest that, while probably composed of very ancient species (now without mitochondria or hydrogenosomes), the phylum belongs more appropriately in the revised kingdom Protozoa, as seen in Table 3. The unique Choanozoa, collar-celled, loricate, colourless flagellates with flattened mitochondrial cristae, presumably at the crossroads of origins of

both kingdoms Animalia and Fungi, have been removed from the Protozoa and tentatively placed in the Animalia (Corliss, 2000), where another protistan group and former protozoan phylum, the Myxozoa, is also now located (see above). References Cavalier-Smith T (1993)Kingdom Protozoa and its 18 phyla. Microbiological Reviews 57: 953 994. Cavalier-Smith T (1998)A revised six-kingdom system of life. Biological

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Hutner SH and Bovee EC (eds)(1985)An Illustrated Guide to the Protozoa. Lawrence. Levine ND. 3rd edn.JJ. KS: Society of Protozoologists. Margulis L and Schwartz KV (1998) Five Kingdoms: an Illustrated Guide to the Phyla of Life on Earth. New York: WH . Corliss JO. (1980)A newly revised classification of the Protozoa. Journal of Protozoology 27: 37 58. Cox FEG et al.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www. Corliss . Margulis L. Boston: Jones and Bartlett. Patterson DJ (1999)The diversity of eukaryotes. American Naturalist 154 (Supplement): 96 124.Freeman. Melkonian M and Chapman DJ (eds)(1990) Handbook of Protoctista. 2nd edn. London: Arnold. Sleigh MA (1989) Protozoa and Other Protists.els.

pp.Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics Further Reading Andersen RA (1992)Diversity of eukaryotic algae. Leadbeater BSC and Diver WL (eds) The Chromophyte Algae: Problems and Perspectives. In: Green JC. Biodiversity and Conservation 1: 267 292. 381 407. Cavalier-Smith T (1995)Evolutionary protistology comes of age: biodiversity and molecular cell biology. Archiv fu¨r Protistenkunde . Cavalier-Smith T (1989)The kingdom Chromista. Oxford: Clarendon Press.

Margulis L (1996)Archaeal eubacterial mergers in the origin of Eukarya: phylogenetic classification of life. Corliss JO (1998)Haeckel s kingdom Protista and current concepts in systematic protistology. Stapfia 56: 85 104. Kudo RR (1966) Protozoology. Springfield. Proceedings of the National Academy of Sciences of the .145: 145 154. Berlin: Springer-Verlag. Grell KG (1973) Protozoology. 5th edn. IL: Charles C Thomas.

USA 93: 1071 1076. July August 1993). Stuttgart: G Fischer. In: Hausmann K and Hu¨ lsmann N (eds) Progress in Protistology (Proceedings of the IX International Congress of Protozoology. Vickerman K. Berlin. In: Coombs GH. Patterson DJ (1994)Protozoa: evolution and systematics. 1 14. Sleigh MA and Warren A (eds) Evolutionary Relationships Among . pp. Vickerman K (1998)Revolution among the Protozoa.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.Protozoa. Dordrecht: Kluwer Academic Publishers. 1 . pp.