Acta Protozoologica (1994) 33: 1 - 51

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An Interim Utilitarian (MUser-friendlyM) Hierarchical Classification

and Characterization of the Protists

John O. CORLISS
Albuquerque, N ew M exico, 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.

ÏNTRODUCTION 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
Souquerque, New Mexico 87153, USA. understanding with respect to the most "natural" scheme
fiónos PaPer 1S dedicated to the memory of Zdzislaw Raabe of classification to employ for these ubiquitous and
L r 1972) who, 30 years ago, perceptively foresaw the prob­ cosmopolitan eukaryotic - generally microscopic and
us involved in constructing a macrosystem for the Protozoa
ased on evolutionary principles (Raabe 1964a) and who, in the often unicellular - organisms. In recent years, with the
*ne year, published a comprehensive protozoological textbook development of molecular Chronometrie techniques (e.g.
. aabe 1964b) and, in the preceding year, had founded the
^national journal Acta Protozoologica. ribosomal RNA sequencing: see Christen 1992), com­
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, I have depended heavily on the literature.
tological systematics. An INDEX of Taxonomic Names Monographs (and shorter papers as well) by phycologi'
is supplied to aid the reader in locating genera or higher cal, protozoological, and mycological specialists on dif
groups of special interest to him or her. Some of my taxa ferent taxa have been indispensable for fliy
may not be identical with putative evolutionary lines of understanding of the composition and taxonomic boufi'
the very recent literature; that is, they may not be daries of such groups: some of the major works amon?
indisputably monophyletic in nature. In fact, several are these are cited in the immediately following section. The
likely paraphyletic and a few perhaps polyphyletic authoritative chapters in four recent treatises (Harris^1
 "User-friendly" Classification of the Protists 3

oni Corliss 1991, Lee et al. 1985, Margulis et al. 1990, (1993), Leipe and Hausmann (1993), Levine (1988),
Parker 1982) deserve special mention; in general, how­ Lipscomb (1985, 1991), Lom(1990), Lom and Dyková
ever (to save space), these individual specialist-contribu- (1992), Lynn (1981), Lynn and Corliss (1991), Margulis
(jQijs - like many of the other numerous scattered (1970, 1981, 1993), Margulis et al. (1990), Margulis et
ta x o n o m ic papers consulted - are not directly cited in al. (1993), Margulis et al. (1984), Mattox and Stewart
this paper. (1984), Melkonian (1984), Melkonian et al. (1991),
Mishler and Churchill (1985), Moestrup (1982, 1991),
Möhn (1984), Moss (1991), Müller (1992), Mylnikov
FURTHER b a c k g r o u n d in f o r m a t io n (1991), O’Kelly (1992, 1993a,b), O ’Kelly and Floyd
(1984), Olive (1975), Page and Blanton (1985), Page
and Siemensma (1991), Parker (1982), Patterson
The protist literature has become so vast, with a (1989a ,b), Patterson and Fenchel (1985), Patterson and
continuing avalanche of papers since my reviews of 8-10 Larsen (1991), Patterson et al. (1989), Patterson and
years ago (Corliss 1984, 1986a), that no attempt can be Sogin (1993), Patterson and Zölffel (1991), Perkins
made here to cite all works of some relevance to my (1991), Powers (1993), Preisig (1989), Preisig et al.
present broad topic. The reader is referred to the follow­ (1991), de Puytorac et al. (1974, 1987, 1993), Ragan
ing publications (which include reviews and overviews (1988), Ragan and Chapman (1978), Raikov (1982),
containing bibliographic sections rich in references to Rothschild (1989), Rothschild and Heywood (1987),
hundreds of significant individual papers) that are most­ Round (1984), Round et al. (1990), Schlegel (1991),
ly concerned with recent ultrastructural or molecular Silva (1980), Sleigh (1989), Sluiman (1985), Small and
researches directly bearing on protist systematics: Lynn (1985), Smith and Patterson (1986), Sogin (1991),
Alexopoulos and Mims (1979), Andersen (1989, Sogin et al. (1989), Sprague (1977), Sprague et al.
1991, 1992), Andersen et al. (1993), Anderson (1983), (1992), Stewart and Mattox (1980), Tappan (1980),
Bárdele (1987), Baroin et al. (1988), Baroin- Taylor (1978, 1987), van den Hoek et al. (1993), Vick-
Tourancheau et al. (1992), Barr (1992), Bold and Wynne erman (1992), Vickerman et al. (1991), Vossbrinck et al.
(1985), Bowman et al. (1992), Bremer (1985), Bremer (1987), W ainrightet al. (1993), Whittaker (1969, 1977),
et al. (1989), Bovee (1991), Broers et al. (1990), Whittaker and Margulis (1978), Woese (1987), Woese
Brugerolle (1991a,b), Canning and Lom (1986), et al. (1990), Wolters (1991).
Cavalier-Smith (1986, 1987, 1989a,b, 1991a,b, 1993a- For nomenclatural help, the original literature has,
c), Chapman and Buchheim (1991), Christensen (1980, once again, been indispensable. But several comprehen­
1989, 1990), Cole and Sheath (1990), Corliss (1979, sive works deserve special mention: Bütschli (1880-
1984,1986a, 1987,1989,1991a), Copeland (1956), Cox 1889), Cavalier-Smith (1993c), Chrétiennot-Dinet et al.
(1980), Davidson (1982), Dodge (1979), Douglas et al. (1993), Copeland (1956), Karpov (1990), Krylov and
(1991), Dragesco and Dragesco-Kemeis (1986), Farmer Starobogatov (1980), Levine et al. (1980), Poche (1913),
(1993), Felsenstein (1988), Fenchel (1987), Fenchel and de Puytorac et al. (1987), and Silva (1980).
Finlay (1991),Fensome et al. (1993), Fleury et al. (1992), In "pre-protist" days and before the advent of
Foissner and Foissner (1993), Foissner (1987, 1993), widespread usage of electron microscopy in study of
Foissner et al. (1988), Gajadhar et al. (1991), Grain microorganisms ("Age of Ultrastructure": Corliss,
(1986), Green et al. (1989), Grell (1991a,b), Grell et al. 1974), biologists did not find it too difficult to recognize
(1990), Hanson (1977), Hasegawa et al. (1993), Haus­ or classify the several major groups of algae ("mini­
mann et al. (1985), Hawksworth et al. (1983), Hibberd plants") and protozoa ("mini-animals"). Algae were
and Norris (1984), Hori and Osawa (1987), Hülsmann predominantly photosynthetic organisms, often non-
(1992), Irvine and John (1984), Karpov (1990), Karpov mo tile, mainly unicellular or filamentous in organiza­
and Mylnikov (1989), Kendrick (1985), Kivic and tion; protozoa were mostly phagotrophic, motile, and
Walne (1984), Knoll (1992), Kreier (1977-1978), Kreier unicellular. The bases for separation of groups of species
and Baker (1991), Kristiansen and Andersen (1986), at higher taxonomic levels included differences in life
Krylov (1981), Krylov and Starobogatov (1980), cycles and pigmentation for algae (e.g. green, red,
Kuznicki and Walne (1993), Larsen and Patterson brown, golden-brown) and variation in kinds and num­
(1990), Larsson (1986), Lee et al. (1985), Lee and bers of locomotory structures (e.g. pseudopodia, flagel­
Kugrens (1992), Leedale (1974, 1980), Leipe et al. la, cilia) and other specialized organelles in the case of
4 J. O. Corliss

protozoa. Ecological characters (e.g. free-living vs revision; although the writer is a past chairman of the
parasitic, marine vs fresh-water, sessile vs free-swim­ committee, the present paper in no way is to be con.
ming, and nutritional proclivities) were also often in­ sidered the outcome of the committee’s current delibera,
voked. Botanists studied the algae; zoologists, the tions, which are continuing without my participation
protozoa. Also, independent of the Society, de Puytorac et al
A few years after the great evolutionary discovery of (1987) and Sleigh (1989), in editions of their well,
(the concept of) the prokaryotes and the eukaryotes, the known textbooks on the protozoa and other protists, have
’’protist perspective" began to be adopted when consider­ used taxonomic arrangements mostly of their own
ing the "lower" eukaryotes (see historical review in making, although largely based on taxonomic w orks
Corliss 1986a). At about the same time, growing accep­ from the literature.
tance of the Serial Endosymbiosis Hypothesis explained In addition to the classifications mentioned above that
the likely endosymbiotic origins of plastids and involve protists, the most outstanding recent attempt to
mitochondria, answering some previously inexplicable bring these organisms together under a single taxonomic
questions while raising some new ones concerning the heading - viz. the Pro(toc)tista - has been that of Mar­
evolution of organisms possessing such organelles. gulis and colleagues, commencing with such seminal
Most botanists have long accepted traditional views papers as those by Whittaker (1969, 1977), Margulis
in setting up hierarchical classifications for their (1974), and Whittaker and Margulis (1978) and cul­
divisions and classes of algae and "lower" fungi, follow­ minating in book form in Margulis et al. (1990). Many
ing such great authorities of the past as the Agardhs, workers, including the present author (e.g. note my
Blochmann, Chodat, Dangeard, de Bary, Fritsch, enthusiasm for the idea in Corliss 1984, 1986a,b, 1991a)
Kjellman, Klebs, Kützing, Lamouroux, Lemmermann, and numerous teachers and textbook writers around the
Lister, Luther, Pascher, Rabenhorst, Smith, West, Wei­ world, have adopted this appealing way of viewing the
tstem, Wille, Winter, and Zopf. See the widely accepted living world as divisible taxonomically into five con­
systems, with minor changes, adopted in many botanical venient kingdoms, among them the Protista. Some 27-45
and phycological (e.g. Bold and Wynne, 1985; van den phyla are generally assigned to the protist kingdom, in
Hoek et al. 1993) textbooks and in numerous recognition of the great diversity (supported today by
monographs as well. The zoologists have acted essen­ hundreds of ultrastructural observations) found among
tially the same with respect to the protozoa, relying on its numerous members (Barnes 1984; Corliss 1984;
the works of such leaders as Alexeieff, Balbiani, Karpov 1990; Margulis and Schwartz 1982, 1988).
Bütschli, Calkins, Cash, Cépède, Chatton, Deflandre, Other scattered proposals of multikingdom systems for
Doflein, Dogiel, Dujardin, Ehrenberg, Entz (Jr. and Sr.), the eukaryotes have not linked the various taxa com­
Grassé, Haeckel, Hartmann, Hertwig, Kahl, Kent, posed purely of protists together in a single kingdom
Kofoid, Kudo, Lankester, Laveran and Mesnil, Léger, (see the pioneering paper by Leedale 1974; Möhn 1984;
Leuckart, O.F. and J. Müller, Penard, Prowazek, and reviews in Corliss 1986a and Lipscomb 1991).
Schaudinn, Schewiakoff, Stein, and Wenyon. For ex­ Phylogeneticists have made invaluable contributions
ample, see the highly popular and authoritative to our understanding of the probable origins of various
protozoological volumes by Grell (1973) and Kudo major monophyletic lines of protists, but they have
(1966). uniformly been reluctant - to date - to suggest definitive,
In more recent years, the Society of Protozoologists hierarchical arrangements of ranked taxa of the principal
has established a special committee to produce "up­ assemblages of eukaryotes that involve species of
dated" systems of protozoan classification: see the protists (e.g. Lipscomb 1991; Patterson 1988; Patterson
reports of Honigberg et al. (1964) and Levine et al. and Sogin 1993). On the other hand, cell and evolution­
(1980). The resulting schemes have often been con­ ary biologist Cavalier-Smith has published a series of
sidered authoritative for sortie years following their heuristic papers during the past dozen years (e.g*
promulgation: for example, the popular "Illustrated Cavalier-Smith 1981, 1983, 1986, 1987, 1989b, 1991b,
Guide to the Protozoa," edited by Lee et al. (1985), in 1993a-c) in which he has boldly presented novel
large measure endorsed the Levine Report. These "con­ schemes of eukaryotic classification, naming and rank'
sensus" classifications have indeed, in many ways, rep­ ing all implicated major groups and generally distribut­
resented improvements over previous systems. The ing the protists among all but one (the Animalia) of his
Society now has a new committee working on a fresh several kingdoms.
 "User-friendly" Classification of the Protists 5

<j0? within the past 20-30 years, we have had, first, a Multiple kingdoms of eukaryotes have been sug­
clinging to the conventional macrosystems of algae and gested in the past, as mentioned on a preceding page.
r0tozoa originally set up essentially on the basis of The number has most commonly been four, of which
Morphological data obtainable by use of light micros­ often the protists have represented one (except in the
copy* Then we have witnessed the "protist revolution," works of Cavalier-Smith, who some years ago foresaw
wjth its emphasis on removing the taxonomic barriers the necessity for the demise of Protista as such: see
of old (often including rejection of formal names that references to his relevant papers above and below).
had become misleading or meaningless: e.g. Unusual was M ohris (1984) proposal of 16 kingdoms,
-phvtoflagellata" and "Zoomastigophora") on the basis with protists alone comprising 10 of them.
of abundant and more precise ultrastructural and If one recognizes kingdoms within the great
ï that jnolecular information. This integrated protistan ap­ eukaryotic group, and - for that matter - among the
pt to proach so permeated our thinking that there was an prokaryotes as well, then a name of still higher
omic exuberant and intensive drive to lump all protists taxonomic rank must be found for those two "super"
Mar- together into a single kingdom (although with many assemblages. There is considerable controversy over the
ninal separate evolutionary lines within that great as­ most appropriate appellation; I am going to employ the
gulis semblage), as discussed above. Today, the prevailing term "empire" without strong feelings either pro or con.
cul- view among leading protistological researchers is that Thus, the kingdoms described below comprise the em­
dany these "lower" eukaryotes can no longer be properly pire EUKARYOTA.
: my restricted to a single taxonomic kingdom, although few Here, as in following sections, I am presenting a
>9la) investigators have stated so directly and even fewer - classification in the conventional manner: naming the
d the aside from Cavalier-Smith (1981 et seq.), the most taxon (with authorship and date) and offering a very
g the notable exception - have relieved the problem in a brief diagnosis, description, or characterization, fol­
con- constructive way, by proposing explicit hierarchical lowed by mention of major embraced sub-taxa. I am
>7-45 classifications to contain the multiple kingdoms and essentially endorsing the six (a reasonable number)
m, in phyla assignable to the eukaryotic assemblage overall. eukaryotic kingdoms of Cavalier-Smith (1989a): three
ty by Thus a single kingdom Protista as such must be laid are comprised solely of protist species, one includes
nong to rest, but long live the protists themselves in all their many, another has only a few; and his sixth, Animalia,
1984; awesome diversity! has none at all, in my view. It may be noted that the
988). great bulk of the photosynthetic or "algal" protists, with
is for the important exceptions of the euglenoids and the
com- dinoflagellates, falls into two kingdoms, the Chromista
gdom MAJOR GROUPS OF EUKARYOTES and the Plantae; the largely heterotrophic or "protozoan"
1984; protists dominate two kingdoms also, the Archezoa and
)• The kingdoms the Protozoa. Mixotrophic species occur mainly in the
itions Chromista (except, again, for numerous members of the
inous Although all the desired information is far from being protozoan phyla Euglenozoa and Dinozoa). The single
have available, one may find it useful to consider what choice phylum of protists in the kingdom Fungi is composed
litive, and what number of kingdoms may best represent the solely of osmotrophic forms; and the heterotrophic,
icipal Principal groups of eukaryotes as they are known to date. multicellular, multitissued kingdom Animalia is con­
es of Certainly modem evolutionary studies have supplied sidered to be without species of protists.
erson sufficient data to demonstrate the tremendous A brief nomenclatural note regarding authorities for
ition- Phylogenetically significant diversity among the protists the names of all kingdoms but the (modem) Chromista
Les of alone to warrant unequivocably the demise of a single is needed here. I am crediting early workers with the
(e.g. kingdom Protista. Furthermore, the nature of the several names Archezoa, Protozoa, Plantae, Fungi, and
991b, high-level groups of these organisms makes clear what Animalia, while aware of the substantial changes over
novel the writer and others have often been reluctant to admit the past 100-240 years in the concepts, circumscription,
rank- ^ the past, viz., that some protists are more closely and composition applied to these top-level groups of
ribut' related to members of other long-accepted kingdoms or eukaryotic organisms. In the case of the Archezoa, I am
of his asseblages (e.g. Plantae, Fungi, Animalia) than they are disagreeing with both Cavalier-Smith and other workers
t° each other. by assigning the authorship to Haeckel who, I am con­

ti
6 J. O. Corliss

vinced, has been misunderstood with respect to his use Phylum 4. OPALOZOA Cavalier-Smith, 1991
and concept of the term. For the Protozoa, I am crediting Class (1) Proterozoea Cavalier-Smith, 1981
Class (2) Opalinatea Wenyon, 1926
Goldfuss with the name and the general concept, as is Class (3) Kinetomonadea Cavalier-Smith, 1993
conventionally done. Finally, in the case of the "big Class (4) Hemimastigophorea Foissner et al., 1988
three" multicellular "higher" eukaryotic assemblages, I
Phylum 5. MYCETOZOA de Bary, 1859
am honoring Linnaeus with all the names (and as of the
Class (1) Protostelea Olive & Stoianovitch, 1966
date 1753), although the plants were originally estab­ Class (2) Myxogastrea Fries, 1829
lished by him as comprising a kingdom Class (3) Dictyostelea Lister, 1909
VEGETABILIA.
Phylum 6. CHOANOZOA Cavalier-Smith, 1989
As is evident, much has happened taxonomically and Class Choanoflagellatea Kent, 1980
nomenclaturally in the 10-year period since my last
attempt to review the high-level status of the protists Phylum 7. DINOZOA Cavalier-Smith, 1981
Class (1) Protalveolatea Cavalier-Smith, 1991
(Corliss 1984). Yet, as will be clear from even a cursory
Class (2) Dinoflagellatea Bütschli, 1885
glance at the classification of the 34 phyla and 83 classes
detailed on the following pages (and see Table 1), the Phylum 8. CILIOPHORA Doflein, 1901
Class (1) Karyorelictea Corliss, 1974
majority of my formerly proposed 18 "supraphyletic
Class (2) Polyhymenophorea Jankowski, 1967
assemblages" and most of the 45 phyla suggested at that (= Heterotrichea Stein, 1859
time have survived in one form or at one level or another. + Spirotrichea Bütschli, 1889)
Some significant changes/interpretations have neverthe­ Class (3) Colpodea Small & Lynn, 1981
less been made - an indication of the impact of new data Class (4) Phyllopharyngea de Puytorac et al., 1974
Class (5) Nassophorea Small & Lynn, 1981
on postulated phylogenetic interrelationships among the Class (6) Oligohymenophorea de Puytorac et al., 1974
diverse groups of protists and, thus, on their macrosys- Class (7) Prostomatea Schewiakoff, 1896
tematics. And far more information is made available in Class (8) Litostomatea Small & Lynn, 1981
the present - and considerably longer - paper, including
Phylum 9. APICOMPLEXA Levine, 1970
a helpful INDEX of taxonomic names. Class (1) Perkinsidea Levine, 1978
Class (2) Gregarinidea Dufour, 1828
_________________________ Table_1_________________________
Class (3) Coccidea Leuckart, 1879
The taxonomic assignment of 34 phyla and 83 classes of protists Class (4) Haematozoea Vivier, 1982
to kingdoms of the EUKARYOTA
Phylum 10. RHIZOPODA von Siebold, 1845
Kingdom I. A R C H E Z O A Haeckel, 1894
Class (1) Lobosea Carpenter, 1861
Phylum 1. ARCHAMOEBAE Cavalier-Smith, 1983 Class (2) Entamoebidea Cavalier-Smith, 1991
Class Pelobiontea Page, 1976 Class (3) Filosea Leidy, 1879
Class (4) Granuloreticulosea de Saedeleer, 1934
Phylum 2. METAMONADA Grassé, 1952
(= mostly Foraminiferea d’Orbigny, 1826)
Class (1) Trepomonadea Cavalier-Smith, 1993
Class (5) Xenophyophorea Schulze, 1904
Class (2) Retortamonadea Grassé, 1952
Class (3) Oxymonadea Grassé, 1952 Phylum 11. HELIOZOA Haeckel, 1866
Phylum 3. MICROSPORA Sprague, 1977 Class (1) Actinophryidea Hartmann, 1913
Class (1) Rudimicrosporea Sprague, 1977 Class (2) Centrohelidea Kühn, 1926
Class (2) Microsporea Delphy, 1963 Class (3) Desmothoracidea Hertwig & Lesser, 1874
Class (4) Taxopodea Fol, 1883
Kingdom II. P R O T O Z O A Goldfuss, 1818
Phylum 1. PERCOLOZOA Cavalier-Smith, 1991 Phylum 12. RADIOZOA Cavalier-Smith, 1987
Class (1) Percolomonadea Cavalier-Smith, 1993
Class (2) Heterolobosea Page & Blanton, 1985 Subphylum - 1- ACANTHARIA Haeckel, 1881
Class (3) Lyromonadea Cavalier-Smith, 1993 Class Acantharea Haeckel, 1881
Class (4) Pseudociliatea Corliss & Lipscomb, 1982
Subphy\um-2- RADIOLARIA J. Müller, 1858
Phylum 2. PARABASALA Honigberg, 1973 Class (1) Polycystinea Ehrenberg, 1838
Class ( 1) Trichomonadea Kirby, 1947 Class (2) Phaeodarea Haeckel, 1879
Class (2) Hypermastigotea Grassi & Foà, 1911
Phylum 13. MYXOZOA Grassé, 1970
Phylum 3. EUGLENOZOA Cavalier-Smith, 1981 Class Myxosporea Bütschli, 1881
Class (1) Diplonematea Cavalier-Smith, 1993
Class (2) Euglenoidea Bütschli, 1884 Phylum 14. ASCETOSPORA Sprague, 1978
Class (3) Kinetoplastidea Honigberg, 1963 Class Haplosporidea Caullery & Mesnil, 1899
 "User-friendly" Classification of the Protists 7

Kingdom m. C H R O M I S T A Cavalier-Smith, 1981 Phylum 4. CHAROPHYTA Rabenhorst, 1863

Class (1) Charophyceae Rabenhorst, 1863
5W
¿kingdom (I) H E T E R O K O N T A Luther, 1899
Class (2) Conjugatophyceae Engler, 1892
Phylum 1. BICOSOECAE Cavalier-Smith, 1989
Subkingdom (13) B I L I P H Y T A Cavalier-Smith, 1981
Class Bicosoecidea Grassé & Deflandre, 1952
Phylum 1. RHODOPHYTA Rabenhorst, 1863
Phylum 2. LABYRINTHOMORPHA Page in Levine Class (1) Bangiophyceae Wettstein, 1901
et ah, 1980 Class (2) Florideophyceae Warming, 1884
Class (1) Labyrinthulea Cienkowski, 1867
Class (2) Thraustochytriacea Sparrow, 1943 Phylum 2. GLAUCOPHYTA Bohlin, 1901
Class Glaucophyceae Bohlin, 1901
Phylum 3. DICTYOCHAE Haeckel, 1894
Class (1) Silicoflagellatea Borgert, 1891 Kingdom V. F U N G I Linnaeus, 1753
Class (2) Pedinellea Kristiansen, 1990
Phylum CHYTRIDIOMYCOTA Sparrow, 1959
Phylum 4. RAPHEDOPHYTA Chadefaud, 1950 Class Chytridiomycetes Sparrow, 1959
Class Raphidomonadea Chadefaud, 1950 Kingdom VI. A N I M A L I A Linnaeus, 1753
Phylum 5. PHAEOPHYTA Weitstem, 1901 (no taxa of protists here)
Class (1) Phaeophyceae Kjellman, 1891
Class (2) Chrysophyceae Pascher, 1914
Class (3) Synurophyceae Andersen, 1987
Class (4) Pelagophyceae Andersen & Saunders, 1993 Empire E U KA R Y O T A
Class (5) Eustigmatophyceae Hibberd & Leedale, 1970
Class (6) Xanthophyceae Allorge in Fritsch, 1935 Kingdom I. ARCHEZOA Haeckel, 1894
Phylum 6. DIATOMAE Agardh, 1824
Class (1) Coscinodiscophyceae Round & Crawford, 1990 Unicellular protists that (allegedly) primitively lack
Class (2) Fragilariophyceae Round, 1990 mitochondria, plastids, typical Golgi bodies,
Class (3) Bacillariophyceae Haeckel, 1878
hydrogenosomes, and peroxisomes, while manifesting
Phylum 7. PSEUDOFUNGI Cavalier-Smith, 1986 various prokaryotic features in their ribosomes and their
Class (1) Oomycetes Winter in Rabenhorst, 1879 rRNAs. Energy produced by anaerobic glycolysis. In­
Class (2) Hyphochytriomycetes Sparrow, 1959 cluded species are amoeboid or flagellated (with low
Subkingdom (II) H A P T O P H Y T A Christensen, 1962 number of flagella), or have no means of independent
Phylum HAPTOMONADA Cavalier-Smith, 1989 locomotion. Some free-living, majority symbiotic in
Class (1) Pavlovea Cavalier-Smith, 1986 variety of hosts and of small to very small body size.
Class (2) Patelliferea Cavalier-Smith, 1993 Entire group is possibly polyphyletic; yet some workers
Swbkingdom (HI) C R Y P T O P H Y T A Pascher, 1914 would add the protozoan parabasalans to it as well.
Contains three phyla and several classes and orders.
Phylum CRYPTOMONADA Ehrenberg, 1838
Class (1) Goniomonadea Cavalier-Smith, 1993
Class (2) Cryptomonadea Stein, 1878

Subkingdom (IV) C H L O R A R A C H N I O P H Y T A
Hibberd & Norris, 1984 Kingdom II. PROTOZOA Goldfuss, 1818
Phylum CHLORARACHNIOPHYTA
Hibberd & Norris, 1984- Predominantly unicellular, plasmodial, or colonial
Class Chlorarachniophyceae Hibberd & Norris, 1984 phagotrophic, colorless protists, wall-less in the trophic
Kingdom IV. P L A N T A E Linnaeus, 1753 state. Included species that are capable of photosynthesis
(some mixotrophic) typically have cytosolic
Subkingdom (I) V I R I D I P L A N T A E
Cavalier-Smith, 1981 chloroplasts with stacked thylakoids, lacking starch, and
Phylum 1. PRASINOPHYTA Christensen, 1962
usually surrounded by three membranes. Nearly univer­
Class (1) Pedinophyceae Moestrup, 1991 sally present are tubular (with a few notable exceptions)
Class (2) Prasinophyceae Christensen, 1962 cristate mitochondria (when absent, replaced by
hydrogenosomes), Golgi bodies, and peroxisomes.
Phylum 2. CHLOROPHYTA Pascher, 1914
Class Chlorophyceae Wille in Warming, 1884 Flagellar mastigonemes, if present, never rigid and
Phylum 3. ULVOPHYTA Stewart & Mattox, 1978 tubular. Numerous free-living (typically independently
Class Ulvophyceae Stewart & Mattox, 1978 motile) and symbiotic species, commonly microscopic
8 J. O. Corliss

in size. As alleged progenitors of the following four Kingdom V. FUNGI Linnaeus, 1753
kingdoms, the protozoa - not unexpectedly - exhibit the
greatest morphological, physiological, and genetic Eukaryotic organisms without plastids 0r
diversity of all. The assemblage, although still broad and phagotrophy (osmotrophic/absorptive nutrition instead)
large and very likely paraphyletic, represents a and possessing cell walls containing chitin and ß-giu^
taxonomic refinement over the classically known cans. Mitochondria (with flattened cristae) and
"phylum Protozoa". peroxisomes nearly always present; Golgi bodies or
Contains numerous phyla, classes, and orders. individual cisternae present. Contains one phylum of
flagellated unicellular (occasionally filamentous)
protists; all of the supra-protistan groups have multicef
Kingdom III. CHROMISTA Cavalier-Smith, 1981 lular mycelia composed of hyphae and are completely
without pseudopodia, flagella, or even centrioles. Many
Predominantly unicellular, filamentous, or colonial symbiotic species but also many free-living, with the
phototrophic protists. Chloroplasts, located in lumen of latter often macroscopic in size.
rough endoplasmic reticulum, lack starch and Contains four phyla, only one of which is composed
phycobilisomes and have a two-membraned envelope of protists.
inside a periplastid membrane (all within the rough,
occasionally smooth, ER). Mitochondria (generally with Kingdom VI. ANIMALIA Linnaeus, 1753
tubular cristae), Golgi bodies, and peroxisomes always
present. When flagella present, at least one bears rigid, Multicellular, non-photosynthetic, usually phago-
tubular, and usually tripartite flagellar hairs or mas- trophic eukaryotes exhibiting a triploblastic body or­
tigonemes (most notable exception, the haptophytes). ganization with collagenous connective tissue
The relatively few species without plastids share other sandwiched between two dissimilar epithelia. Mito­
features in common with majority of forms embraced chondria (with flattened or rarely tubular cristae), Golgi
here. Mostly free-living (but some groups not inde­ bodies, and peroxisomes always present. Multiple tis­
pendently motile); many microscopic in size, with some sues and organ systems, and typically with complex
major exceptions (e.g. brown algae). embryological development during ontogeny. Com­
Contains several phyla with numerous classes and monly macroscopic in size. Mostly free-living and
orders. motile but with some symbiotic groups, latter usually
exhibiting osmotrophic nutrition.
Contains numerous phyla, classes, and orders, none
Kingdom IV. PLANTAE Linnaeus, 1753 of which includes any protists.

Unicellular, colonial, or multicellular phototrophic The phyla and their major lesser taxa
protists and multicellular photosynthetic "higher"
eukaryotes, all typically (but not universally) with cel- In recent years, the phylum (or, often, in the case of
lulosic cell walls in trophic stages. Cytosolic plastids, algae, the division) has become the principal high-level
enveloped by two membranes, usually contain starch or taxonomic rank in considering both phylogeny and sys­
phycobilisomes. Mitochondria (with flattened cristae), tematics of protists, a fact to which many papers in the
Golgi bodies, and peroxisomes always present. Green literature attest (see reviews in Corliss 1984, 1986a,
species have stacked thylakoids with chlorophylls a and 1993). And it is widely admitted that the number of phyla
b; red algae, totally without flagella, have single unstack­ of these "lower" eukaryotes must be a large one, not
ed thylakoids covered with phycobilisomes, with surprising in consideration of their great morphological
cytosolic starch. Predominantly free-living, and non- and genetic diversity.
motile in trophic stages. The "higher" plants, commonly In view of relatively meager molecular data, the
macroscopic in size, develop from embryos and are cladistic protistologist faces a difficult problem not only
mostly terrestrial and vascular forms with alternation of in identifying these phyla (or appreciating that s u c h a
haploid and diploid generations. rank is appropriate for them) but in interrelating then1
Contains several phyla with quite a number of classes phylogenetically on the protistan sensu lato "tree". Some
and orders. evolutionary biologists, on the other hand, feei a need
 "User-friendly" Classification of the Protists 9

such groups into still higher packages (e.g.

to a s s e m b l e to my DISCUSSION section for lengthier consideration
superphyla, infra- and subkingdoms, and kingdoms) and of some major evolutionary or taxonomic problems.
or ; ¿so to subdivide them unmercifully at lower ranks, For a convenient summary of the classification
ead) -j interposing between phyla and classes such ranks as sub- presented in this paper, the reader is referred to Table 1
•gla- and infraphyla and superclasses (not to mention inter­
and • mediate taxa at ordinal, familial, and generic levels as Phyla of Kingdom ARCHEZOA
•s or well)* I can understand the rationale involved in the latter
n of « practices, for they allow the user of the resulting In addition to the three quite disparate phyla
tous) schemes to appreciate the clustering of certain taxa that described below, some workers have suggested the
:icel- share key characters. Yet, for the general user, the inclusion here, also, of the parabasalan flagellates
etely teach er and the student, and the person interested in (considered as the second phylum of the kingdom
dany having a classification helpful in information retrieval, PROTOZOA in the present work) and/or of the
i the too much detail is not a desirable feature, in my opinion. family Entamoebidae (in a class of the protozoan
Also, the introduction of intermediate ranks often re­ phylum Rhizopoda here). Other taxonomists conser­
losed quires new names, adding to the memory burden of the vatively feei that this entire kingdom itself should be
non-specialist. Therefore, in the present treatment of the subsumed by the PROTOZOA.
protists, I am usually omitting reference to many of the
"intermediate'' ranks employed, for example, in the Phylum 1. ARCHAMOEBAE Cavalier-Smith, 1983
longer works of Cavalier-Smith (e.g. 1993c).
îago- Whenever possible, I have adopted phyletic names Amoeboid or amoeboflagellated (generally a poorly
y or- familiar from the literature, as long as the composition motile single flagellum) protists with characteristics
:issue and the rank of the taxon have not been too drastically of the kingdom (q.v.), although presumed presence
Vlito- changed over time (see DISCUSSION). With respect to of 70s ribosomes not yet confirmed. Microaerobic,
3olgi the spelling of names of protist phyla, especially the with symbiotic bacteria; free-living, mostly fresh­
e tis- prefixes and suffixes, I have - once again - tried to water habitats.
nplex maintain traditional forms of the words. In particular, I
Com- have retained the phylum/di vision ending "-a" (excep­ Class Pelobiontea Page, 1976
^ and tionally, -"ae" and one time "-i") so long used in both (syn. Karyoblastea Margulis, 1974 p.p.)
mally botanical and zoological taxonomic literature. For clas­ Single class, thus with characters of phylum. Two
ses, I have used the traditional protozoological and orders recognized by some workers: Mastigamoebida
none botanical suffixes of "-ea" and "-phyceae" (occasional­ Frenzel, 1892; and Phreatamoebida Cavalier-Smith,
ly, "-cetes"); and for subclasses, "-ia". My orders end in 1991, latter solely for the genus Phreatamoeba.
I "-ida"; but, in the interest of saving space, not many Mastigamoeba, M astigella, M astigina, Pelomyxa,
j orders are included on the following pages. Phreatamoeba
I As an aid to readers of various backgrounds with
ase of I respect to protistological high-level systematics, names Phylum 2. METAMONADA Grassé, 1952
-level °f a goodly number of representative or common or
dsys- familiar genera (plus some new within the past several Bi-, quadri-, octo- (or occasionally more) flagellated
in the years) are given under the lowest-ranked taxa listed protists, mostly symbiotic species, with characters of
986a, within a specific phylum. The INDEX will serve as a the kingdom (q.v.), including 70s ribosomes and (e.g.
phyla convenient guide to the page-locations of these generic Giardia) 16s rRNA. Some free-living, majority intes­
e, not names. tinal symbionts of various hosts. Parabasalans (see
ogical My ordering of phyla within a kingdom is basically second phylum of kingdom Protozoa) are here tenta­
(meant to be) phylogenetic. An alphabetical arrangement tively excluded from the present phylum.
a, the would be of little value. On the other hand, to make up
>t only f°r our ignorance of "true" evolutionary interrelation­ Class (1) Trepomonadea Cavalier-Smith, 1993
juch a ' s , a measure of "educated guess-work" (= healthy One or two karyomastigonts, each with 1-4 flagella;
¡ them speculation?) is required in quite a number of instances. contractile axostyle absent; cytostomal-cytopharyn-
Some : Quite often, brief comments are offered in place in geal apparatus present; few cell-surface cortical
i n eed
c°ntroversial situations; otherwise, the reader is referred microtubules. Free-living or symbiotic.
10 J. O. Corliss

Order 1. Diplomonadida Wenyon, 1926 Order 1. Minisporida Sprague, 1972

Brugerolleia, Giardia, Hexamita, Octomitus, Burkea, Buxtehudea, Chytridiopsis, Hessea
Spironucleus, Trepomonas, Trigonomonas Order 2. Microsporida Balbiani, 1882
Order 2. Enteromonadida Brugerolle, 1975 Culicospora, Encephalitozoon, Endoreticulatu$
Caviomonas, Enteromonas, Trimitus Enterocytozoon, Glugea, Gurleya, Loma, MicrofilUrn
Mrazekia, Nosema, Perezia, Pleistophora, Spraguea
Class (2) Retortamonadea Grassé, 1952
Stempellia, Tardivesicula, Telomyxa, Thelohania
Generally with characteristics given for the first class
Tricornia, Tuzetia, Unikaryon, Vairimorpha
(above), but with cortical microtubules over entire
body surface. Mostly intestinal symbionts (e.g. of
Phyla of Kingdom PROTOZOA
insects and mammals). Single order.
Order Retortamonadida Grassé, 1952
Phylum 1. PERCOLOZOA Cavalier-Smith, 1991
Chilomastix, Retortamonas
Unicellular, non-pigmented protozoa allegedly primi­
Class (3) Oxymonadea Grassé, 1952
tively lacking Golgi bodies; peroxisomes usually
One or more karyomastigonts, each with 4 flagella;
present; mitochondria or, more rarely, hydrogeno­
basal bodies of flagellar pairs are connected by a
somes present; mitochondrial cristae flat, sometimes
paracrystalline paraxostyle in which are embedded
discoidal, atypical of kingdom. Flagella usually
anterior ends of axostylar microtubules; axostyles
present, 1-4 (occasionally more), without mas-
typically contractile; cytopharynx absent. Intestinal
tigonemes; some species are amoeboflagellates,
symbionts of insects. Single order.
several of which never have flagella; fresh-water and
Order Oxymonadida Grassé, 1952
marine habitats.
Monocercomonoides, Notila, Oxymonas, Pyrsonym­
pha, Saccinobaculus Class (1) Percolomonadea Cavalier-Smith, 1993
Non-amoeboid, quadriflagellated forms; striated
Phylum 3. MICROSPORA Sprague, 1977 rootlets absent. Exhibiting the most primitive char­
acters of the phylum and comprising only a single
Minute, unicellular, obligate intracellular symbionts, genus, species here have been assigned by past
with characteristics of the kingdom (including 70s workers to the following much larger class.
ribosomes); sporoplasm uni- or binucleate; no flagel­ Percolomonas
lated stage in life cycle; underdeveloped Golgi(?)
Class (2) Heterolobosea Page & Blanton, 1985
bodies; resistant spores contain complex extrusome,
Monopodial amoeboid trophic form; transitory'
with polar tube and cap; one layer of thick spore wall
flagellated stage, sometimes missing altogether;
chitinous. Commonly in variety of cells of diverse
striated rootlets present; close association of rough
fresh-water, marine, or terrestrial hosts, mainly
endoplasmic reticulum with mitochondria. Fruiting
arthropods (especially insects) and fishes, but includ­
bodies present in the first order, absent in the second.
ing even other protists. Name of the phylum is,
Is the genus Fonticula here or in Rhizopoda?
unfortunately, identical to the generic name of a green
Order 1. Acrasida Schröter, 1886
algal protist in the kingdom Plantae.
Acrasis, Pocheina
Order 2. Schizopyrenida Singh, 1952
Class (1) Rudimicrosporea Sprague, 1977
Adelphamoeba, Gruberella, Heteramoeba, Naeg-
Extrusion apparatus rudimentary, with thick
leria, Paratetramitus, Pernina, Pseudovahlkampfi&
(manubroid) non-spiraled polar tube and no
Singhamoeba, Tetramastigamoeba, Tetramitus,
polaroplast or posterior vacuole. Single order.
Vahlkampfia
Order Metchnikovellida Vivier, 1975
Amphiacantha, Desportesia, Metchnikovella Class (3) Lyromonadea Cavalier-Smith, 1993
Anaerobic flagellates with hydrogenosomes and no
Class (2) Microsporea Delphy, 1963 peroxisomes; harp-shaped structure of microtubule^
Complex extrusion apparatus, with coiled polar tube, two pair anterior flagella, 1-4 nuclei. Only two
polaroplast and posterior vacuole typically present. genera.
Many species in second order. Lyromonas, P s alteromonas
 "User-friendly" Classification of the Protists 11

Class (4) Pseudociliatea Corliss & Lipscomb, 1982 with persistent nucleolus; many fresh-water free-
IVlultiflagellated, multinucleate forms with living forms, but also number of important symbiotic
mitochondria (with rigid discoid cristae) and species (e.g. human blood parasites); some species
peroxisomes. Single genus, with several species. photosynthetic, with chloroplasts in cytosol with
Some workers have appended these protists to the chlorophylls a and b and enveloped in three
phylum Euglenozoa (below). membranes but lacking starch. Three classes; a pos­
Stephanopogon sible fourth, Pseudociliatea, now appears in phylum
1, Percolozoa (above), and a possible fifth, Hemimas-
phylum 2. PARABASALA Honigberg, 1973 tigophorea, is placed tentatively in phylum 4,
Opalozoa (below). The bodonids plus the
Unicellular, almost exclusively symbiotic, flagellates trypanosomatids might deserve taxonomic separation
with mastigont system typically with multiple flagella from the euglenoids proper at the level of subphylum.
and one or more nuclei; 70s ribosomes; no
mitochondria but hydrogenosomes in a double en­ Class (1) Diplonematea Cavalier-Smith, 1993
velope; characteristic complex parabasal body ap­ Phagotrophic flagellates (two equal flagella) lacking
paratus (= Golgi body). Wide range of hosts, includ­ chloroplasts, pellicular plates, kinetoplasts, paraxial
ing humans, but many species in termites and wood- rods; plate-like mitochondrial cristae; feeding ap­
feeding roaches. Some workers place this phylum in paratus with vanes and two supporting rods. Single
the kingdom Archezoa. genus.
Diplonema (syn. Isonema)
Class (1) Trichomonadea
Typically 4-6 flagella; pelta and non-contractile axos- Class (2) Euglenoidea Bütschli, 1884
tyle part of each mastigont (one exception); trophic Unicellular or colonial, bi- (rarely more) flagellated
form in one genus (Dientamoeba) permanently forms; phagotrophic, photosynthetic (with paramylon
amoeboid with no flagella. storage product), osmotrophic, or mixotrophic; all
Order Trichomonadida Kirby, 1947 pigmented species with stigma (eyespot) containing
Bullanympha, Calonympha, Devescovina, Dien­ ß-carotene derivatives and other carotenoid pig­
tamoeba, Ditrichomonas, Hexamastix, Histomonas, ments. Numerous species. The exact taxonomic
Monocercomonas, Pseudotrichomonas, Snyderella, placement within the class of a number of species
Trichomonas symbiotic in copepods (especially members of such
genera, themselves of doubtful validity, as Con­
Class (2) Hypermastigotea Grassi & Foà, 1911 radinema, Paradistigma, Parastasia, and others)
Mastigont system with numerous flagella and multi­ must await further study.
ple Golgi bodies; basal bodies of flagella often ar­ Order 1. Euglenida Bütschli, 1884
ranged in closely packed longitudinal or spiral rows; Astasia, Colacium, Distigma, Euglena, Eutreptia,
single nucleus. Khawkinea, Phacus, Trachelomonas
Order 1. Lophomonadida Light, 1927 Order 2. Euglenamorphida Leedale, 1967
Joenia, Lophomonas, Mesojoenia, Microjoenia Euglenamorpha, Hegneria
Order 2. Trichonymphida Poche, 1913 Order 3. Rhabdomonadida Leedale, 1967
Barbulanympha, Deltotrichonympha, Holomas- M enoidium, Rhabdomonas
dgotoides, Hoplonympha, Kofoidia, Macrospironym- Order 4. Heteronematida Leedale, 1967
pha, Spiro trichonympha, Teranympha, Trichonympha Entosiphon, Heteronema, Peranema, Petalomonas,
Ploeotia, Sphenomonas
Phylum 3. EUGLENOZOA Cavalier-Smith, 1981
Class (3) Kinetoplastidea Honigberg, 1963
nd no Forms with 1-4 flagella, with paraxial rods and non- Small, colorless flagellates with 1-2 flagella (arising
bules; tubular mastigonemes; with peroxisomes and com­ from pocket and possessing paraxial rod) and
y two monly discoidal mitochondrial cristae, latter atypical prominent kinetoplast (distinctive body of massed
of kingdom; cytoskeleton of microtubules reinforcing DNA) within single mitochondrion, latter typically
cortex; Golgi bodies well developed; nuclear division extending length of body; numerous peroxisomes
12 J. O. Corliss

(known as glycosomes) present; free-living or sym­ Four (in first order) to many flagella, apical or ar­
biotic, with latter (including blood parasites, often ranged in oblique longitudinal rows; typically one
highly pathogenic) exhibiting elaborate life cycles (first order) or either two or many nuclei; no
frequently involving two hosts. peroxisomes; osmotrophic nutrition; all species sy^.
Order 1. Bodonida Hollande, 1952 biotic, endocommensals principally in amphibian
Bodo, Cephalothamnium, Cryptobia, Ichthyobodo, hosts. Differences between members of first and
Procryptobia, Rhynchomonas second order, all not listed here, may require greater
Order 2. Trypanosomatida Kent, 1880 taxonomic separation in the future. The entire class
Blastocrithidia, Crithidia, Endotrypanum, ifer- is rather atypical of the phylum v.v. Patterson’s
petomonas, Leishmania, Leptomonas, Phytomonas, (1986a) recent taxon Slopalinida also embraced
Trypanosoma Proteromonas, which I have placed in the preceding
class, Proterozoea.
Phylum 4. OPALOZOA Cavalier-Smith, 1991 Order 1. Karotomorphida Cavalier-Smith, 1993
Karotomorpha
Predominantly small, free-living, unicellular, Order 2. Opalinida Poche, 1913
uninucleate, biflagellated protozoa with tubular Cepedea, Opalina, Protoopalina, Protozelleriella,
mitochondrial cristae and totally lacking chloroplasts, Zelleriella
cortical alveoli, and rigid tubular mastigonemes.
Likely a paraphyletic assemblage, many of its Class (3) Kinetomonadea Cavalier-Smith, 1993
species have not yet been well studied by modern Free-living uninucleate forms with 2-4 flagella,
techniques. Cavalier-Smith (1993b, c) assigns some peroxisomes, usually unique extrusomes (kineto-
20 generally small orders here, many new, only a few cysts); mitochondrial cristae flat or with branched
are considered below. Among problematical groups tubules; some species with axopodial axonemes
possibly in this phylum, mostly in my class 1, are nucleated by axoplast associated with exceptionally
various proteomyxids s.L, Ebria, Phagodinium, long centrioles. Possibly multiple orders in this class,
Phagomyxa and the fungus-like Nephromyces. which includes some of the "helioflagellates" of the
literature.
Class (1) Proterozoea Cavalier-Smith, 1981
Ancyromonas, Dimorpha, Heliomonas, Histiona,
Generally with characters of phylum; rarely, with
Re dinomonas, Tetradimorpha
flattened mitochondrial cristae; a few groups contain
symbiotic forms. Many minute, little-studied free-
Class (4) Hemimastigophorea Foissner, Blatterer &
living marine, fresh-water, and soil flagellates or
Foissner, 1988
amoeboflagellates may belong here (some included
Small, colorless, multiflagellated, phagotrophic (but
in partially tentative lists of genera given below).
no permanent cytostome) protists with "infracilia-
Order 1. Heteromitida Cavalier-Smith, 1993
ture" reminiscent of ciliates; mitochondrial cristae
Amastigomonas, Anisomonas, Apusomonas, Cer­
saccular-tubular; no paraxial rods, no mastigonemes;
comonas, Diphyllea, Discocelis, Heteromita, Jakoba,
two microtubule- bearing pellicular plates; complex
Leucodictyon, Massisteria, Proteromonas, Pseudo­
extrusomes. Found primarily in soils. Some workers
spora, Thaumatomastix
consider this class as possibly a separate phylum, and
Order 2. Cyathobodonida Cavalier-Smith, 1993
even closer to the phylum Euglenozoa (above) than
Cyathobodo, Kathablepharis, Leucocryptos, Phalan­
indicated here. Single order, three genera.
sterium, Platychilomonas, Pseudodendromonas,
Order Hemimastigida Foissner et al., 1988
Spongomonas
Hemimastix, Spironema, Stereonema
Order 3. Plasmodiophorida Cook, 1928
(name better credited to Zopf, 1885?)
Phylum 5. MYCETOZOA de Bary, 1859
Octomyxa, Plasmodiophora, Polymyxa, Sorodiscus,
(syns. ± EUMYCETOZOA Zopf, 1885, and
Spongospora, Tetramyxa, Woronina
MYXOMYCETES & MYXOMYCOTA m cttpp)
Class (2) Opalinatea Wenyon, 1926
(syns. Protociliata Metcalf, 1918, Paraflagellata Free-living, unicellular or syncytial p l a s m o d i ^
Corliss, 1955, Slopalinida Patterson, 1986 p.p.) forms, non-flagellated in their uni- or multinucleate
 "User-friendly” Classification of the Protists 13

phagotrophic stages; mitochondrial cristae tubular; species may have quite complex basket-like loricae
uni- or multicellular aerial fruiting bodies of siliceous costae arranged longitudinally. These
(sporophores or sorocarps) bearing one to many widely distributed protists (especially in marine
spores with cellulosic or chitinous walls; spore ger­ habitats), known familiarly as the collar flagellates,
mination produces amoeboid or uni- or biflagellated have also been called choanoflagellates, choanomo-
cells. Widely distributed in decaying vegetation. Per­ nads, craspedomonads, craspedophyceans and even
haps the characterization of the whole phylum should craspedomonadophyceans. Cavaher-Smith’s phyletic
be expanded to include many of the taxonomically name, originally published as "Choanociliata",
enigmatic marine plasmodial protists with emended by him in 1989. Single class with single
reticulopodia studied by Grell (e.g. 1985,1991b) and order: names both credited to Kent here.
placed by him in an order Promycetozoida Grell,
1985: for example, Corallomyxa, Megamoebomyxa, Class Choanoflagellatea Kent, 1880
and Thalassomyxa. But relationships of such genera (syn. Craspedophyceae Chadefaud, 1960)
to the Rhizopoda (Lobosea or maybe the athalamid Order Choanoflagellida Kent, 1880
Granuloreticulosea), or even to certain heterokonts Acanthoeca, Acanthoecopsis, Bicosta, Calliacantha,
Iella, (in the kingdom Chromista), remain unresolved pos­ Codosiga, Conion, Diaphanoeca, M onosiga, Par­
sibilities. Phylum contains both cellular and "acel- vicorbicula, Pleurasiga, Proterospongia, Salpin­
lular" slime molds. goeca, Stephanoeca

Gass (1) Protostelea Olive & Stoianovitch, 1966

Single amoeboid cells, with filóse pseudopodia, give Phylum 7. DINOZOA Cavalier-Smith, 1981
rise to simple sorocarps (= sporocarps) of one to few
spores on delicate, narrow stalk. Several species Biflagellated, uninucleate protozoa with amphiesmal
(amoeboflagellates, in effect) have a flagellated vesicles or cortical alveoli (containing cellulosic
stage in their life cycle. plates in some groups), tubular (sometimes ampul­
Cavostelium, Ceratiomyxa, Protostelium liforme mitochondrial cristae, and peroxisomes; one
flagellum typically with paraxial rod; ca. 50% of
wna, Gass (2) Myxogastrea Fries, 1829 extant species pigmented, with chloroplasts contain­
Generally with characters of phylum, as largest group ing chlorophylls a and c, enveloped by three (rarely
of the plasmodial (or "acellular") slime molds. Mul­ two) membranes, lacking phycobilisomes, and lo­
tiple orders recognized. cated in cytosol; non-pigmented and some colored
Badhamia, Comatricha, Cribraria, Didymium, species phagotrophic; nucleus haploid, typically with
: (but Echinostelium, Fuligo, Licea, Lycogala, Physarum, distinctive chromosomes consisting primarily of non­
icilia- Stemonitis, Trichia, Tubulina protein complexed DNA. Assemblage at one time
ristae called the "Mesokaryota" because of its alleged pos­
^mes; Class (3) Dictyostelea Lister, 1909 session of a combination of pro- and eukaryotic
nplex Cellular slime molds from soil with triphasic life characters. First of three phyla known collectively as
>rkers cycle: unicellular amoeboid microphage, pseudoplas­ the "Alveolata", a super-category designated a
i, and modium (slug) formed by aligned aggregating parvkingdom by Cavalier-Smith (1993c). Possibly
) than myxamoebae, and multicellular sorocarp on branched assignable somewhere here is the puzzling "giant
or unbranched stalks. protist" Hochbergia (a cephalopod symbiont measur­
Acytostelium, Dictyostelium, Polysphondylium ing 1-2 mm in length: Shinn and McLean, 1989).

% lum 6. CHOANOZOA Cavalier-Smith, 1989 Class (1) Protalveolatea Cavalier-Smith, 1991

Atypical dinozoa; closed mitosis, but mitotic spindle
and Free-living, uniflagellated, colorless, unicellular or intranuclear; chromatin of normal eukaryotic form;
tt. pp) colonial forms with non-discoid flattened (atypical of all non-photosynthetic species, mostly free-living
kingdom) mitochondrial cristae; single flagellum phagotrophs but some marine forms totally symbiotic
lodiai surrounded by collar of microvilli (aetin filaments (osmotrophic); typically unicellular and uninucleate,
cleate internally) used in microphagous feeding; marine but symbiotic forms may be branched and multi-
14 J. O. Corliss

nucleate. Several quite distinctive orders may be terrestrial habitats, but symbiotic and symphoriontic
justifiable here. Some of the included genera require species associated with great variety of host or
further study to be certain of their taxonomic assign­ ganisms. Sometimes known as the ÍNFUSORU
ment. auctt. (a very old and vague term) or by the more ant
Colponema, Ellobiopsis, Oxyrrhis, Thalassomyces name of HETEROKARYOTA Hickson, 1903, the
cilioprotists comprise one of the largest protist phyia
Class (2) Dinoflagellatea Bütschli, 1885 with 8-10 classes and many orders. Second member
With characters of phylum, as its major class; mostly of "Alveolata" group.
unicellular species, but some form colonies (especial­
ly catenoid); often free-living, auto- or phagotrophic Class (1) Karyorelictea Corliss, 1974
or both (mixotrophic), but some groups symbiotic Flattened body, often ribbon-like and contractile’
(osmotrophic); members of a major endosymbiotic habitat commonly interstitial niches of marine sands
sub-taxon have no pellicular alveoli and very low (one major genus fresh-water); two to many non­
number of chromosomes with histones clearly dividing (and essentially diploid) macronuclei
present, but dinospores prove their position here. formed anew at organism’s fission from dividing
Widely distributed forms, mostly planktonic. Multi­ diploid micronuclei; postciliodesmata charac­
ple orders in literature, with half of contained species teristically present; generally without cortical alveoli;
represented by fossil forms (but only genera with many species without definitive mouth but
living species are listed below). Class described here phagotrophic via non-ciliated area of ventral surface.
is essentially the equivalent of botanists’ Pyr- The group, considered primitive by a number of
rhophyta Pascher, 1914, and Dinophyceae Fritsch, workers, is divisible into two orders. But the class
1927. itself should perhaps be reduced to a subordinate
Alexandrium, Amoebophrya, Amphidinium, Blas­ taxon of (part of) the next class, below: see comments
todinium, Ceratium, Chytriodinium, Cryp­ there.
thecodinium, Cystodinium, Dinophysis, Duboscq­ Kentrophoros, Loxodes, Remanella, Trachelocerca,
uella, Erythropsidinium, Glenodinium, Gleodinium, Trachelonema, Tracheloraphis
Gonyaulax, Gymnodinium, Gyrodinium, Haplozoon,
Kofoidinium, Noctiluca, Oodinium, Oxytoxum, Class (2) Polyhymenophorea Jankowski, 1967
Peridinium, Polykrikos, Prorocentrum, Protoperi­ (syns. Heterotrichea Stein, 1859 plus Spirotrichea
dinium, Ptychodiscus, Pyrocystis, Pyrophacus, Bütschli, 1889, in effect; and Postciliodes­
Rhizodinium, Roscoffia, Symbiodinium, Syndinium, matophora Gerassimova & Seravin, 1976 p.p.)
Thoracosphaera, Zooanthella Diverse body shapes and sizes (some quite large), of
both free-living (marine and fresh-water) and sym­
Phylum 8. CILIOPHORA Doflein, 1901 biotic species; some groups with postciliodesmata;
characteristically with many conspicuous buccal
Commonly with numerous longitudinal rows of cilia membranelies, plus compound somatic ciliature
(single or paired), with perkinetal fission (cirri) in some groups. Based on recent rRNA
(homothetogenic, as opposed to the symmetrogenic analyses, supported by ultrastructural observations,
of flagellates), and distinctive kinetidal infraciliature; some workers would split this huge and probably
many have complex oral ciliature; cortical alveoli paraphyletic assemblage into at least two classes
characteristic of most groups; mitochondrial cristae (thus elevating Polyhymenophorea to some supra-
tubular, often curved; in anaerobic species, level or even eliminating it), with my first class
mitochondria may be missing or replaced by Karyorelictea embraced by the heterotrich moiety
hydrogenosomes; nuclear apparatus heterokaryotic, and the remainder of my polyhymenophoreans as­
typically with one or more diploid micronuclei and signed to a spirotrich group. Here, each such major
one or more polyploid macronuclei; sexual section is conservatively treated as a subclass. Each
phenomenon of conjugation; heterotrophs nutrition­ has many orders (the first one would also have two
ally, but some species have photosynthetic algal subclasses of its own, the karyorelictians and the
protists as endosymbionts. Widely distributed, often heterotrichians s.s., if it were elevated to independent
conspicuous forms, mostly free-living in aquatic and class status).
 "User-friendly" Classification of the Protists 15

Subclass 1. H eterotrichia Stein, 1859 ly surrounded, in the old cyrtophorids , by nematodes-

The classically known heterotrichs s.L plus the newer mata (= cytopharyngeal basket or cyrtos); macro­
protoheterotrichs and possibly (as discussed above) nucleus characteristically heteromerous in many (but
the karyorelictids (see my class 1). not suctorian) species; suctorians also atypical in
Anigsteinia, Ascobius, Avelia, Blepharisma, other ways: polystomic with sucking tentacles, non-
Brachonella, Caenomorpha, Clevelandella, Climaco­ ciliated trophic stage carnivorous, commonly
stomum, Condylostoma, Epalxella, Fabrea, FoZ- stalked, reproduction by budding; chonotrichs
liculina, Geleia, Lagotia, Licnophora, Metopus, likewise specialized: heteromerous macronucleus
Mylestoma, Nyctotheroides, Nyctotherus, Paracichli- but no nematodesmata, sessile forms (ectosymbionts
dotherus, Peritromus, Phacodinium, Protocruzia, on crustaceans), limited ciliation, reproduction by
Reichenowella, Saprodinium, Sicuophora, Spiros­ budding. Three subclasses with several orders.
tomum, Stentor, Transitella Acineta, Ancistrocoma, Brooklynella, Chilodochona,
Chilodonella, Chlamydodon, Cyathodinium,
Subclass 2. S pirotrichia Bütschli, 1889 Dendrocometes, Dendrosoma, Dysteria, Endos­
The classically known oligotrichs s.Z. (i.e. oligotrichs phaera, Ephelota, Hartmannula, Heliochona, Helio-
s.s. + tintinnids s.Z. = today’s choreotrichs) and the phrya, Hypochona, Isochona, Lobochona, Lorico­
old hypotrichs s.Z. (i.e the pre-1980 stichotrichs and phrya, Lwoffia, Lynchella, Ophryodendron, Paraci­
sporadotrichs). neta,, Phalacrocleptes, Phascolodon, Podophrya,
Amphisiella, Aspidisca, Australothrix, Bakuella, CZr- Raabella, Rhabdophrya, Sphenophrya, Spirochona,
rhogaster, Codonella, Cyrtostrombidium, Diophry­ Stylochona, Tachyblaston, Thecacineta, Tokophrya,
opsis, Diophrys, Discocephalus, Euplotes, Favella, Trichochona, Trichophrya, Trochilia, Vasichona
Gastrostyla, Halteria, Kahliella, Kerona, Kiitricha,
Laboea, Lamtostyla, Leegaardiella, Lohmanniella, Class (5) Nassophorea Small & Lynn, 1981
Nolaclusilis, Onychodromus, Oxytricha, Pelagohal­ Characterized by common possession of highly dis­
teria, Pelagostrombidium, Plagiotoma, Strobilidium, tinctive "nasse" or cyrtos in cytopharyngeal area;
Strombidium, Stylonychia, Territricha, Tintinnopsis, hypostomial frange prominent or reduced to few
Tontonia, Tricoronella, Undella, Uronychia, pseudomembranelles; fibrous trichocysts; mostly
Urosomoides, Urostyla, Wallackia, Xystonellopsis, free-living, fresh-water forms. Several orders, but
Yvonniellina peniculines (e.g. Paramecium) are excluded (see 6th
subclass of next class, below), leaving only (some of)
Class (3) Colpodea Small & Lynn, 1981
the old cyrtophorids here.
Somatic dikinetids, reticulate silverline system, and
Furgasonia, Leptothorax, Microthorax, Nassula,
somatic stomatogenesis; posterior kinetosome has
Nassulopsis, Pseudomicrothorax, Scaphidiodon,
well developed transverse microtubular ribbon ex­
Zosterodasys
tending posteriorly, forming LKm fiber by paralleling
and overlapping with ribbons from more anterior
Class (6) Oligohymenophorea de Puytorac et al., 1974
dikinetids; oral ciliature consists of right and left
Somatic kineties, unless entirely absent, often com­
ciliary fields; mainly terrestrial or edaphic forms.
posed of monokinetids; buccal apparatus, when
Two subclasses, second one solely for five small
present, consists basically of paroral (formerly un­
families; great bulk of the species are in the nominate
dulating membrane, UM) dikinetid on right and
subclass, which contains half a dozen orders.
several membranelles or polykinetids (AZM) on left;
Aristerostoma, Bresslaua, Bryometopus, Bryophrya,
distinct, overlapping kinetodesmata; mucocysts com­
Bursaria, Bursaridium, Colpoda, Cosmocolpoda,
mon, with explosive trichocysts in some species. Six
Cyrtolophosis, Grandoria, Grossglockneria, Haus-
quite diverse subclasses warrant separate descriptions
manniella, Kreyella, Maryna, Mycterothrix,
here.
Platyophrya, Pseudo glaucoma, Sorogena, Thylaki-
dium, Trihymena, Woodruffia
Subclass 1. Hymenostomatia Delage & Hérouard, 1896
\ Class (4) Phyllopharyngea de Puytorac et al., 1974 Somatic monokinetids; right-most postoral kinety
Cytopharynx lined with radially arranged leaf-like stomatogenic; buccal ciliature tetrahymenal (UM +
microtubular ribbons (= phyllae), themselves typical­ AZM). Two orders, second - the ophryoglenids - with
 T
16 J. O. Corliss

a unique watchglass organelle and a complex life Opercularia, Ophrydium, Opisthonema, Orboper
cycle as obligate histophagous symbionts. cularia, Pallitrichodina, Platycola, Polycycia
Bursostoma, Colpidium, Curimostoma, Espejoia, Propyxidium, Rhabdostyla, Scyphidia, Semi
Glaucoma, Ichthyophthirius, Jaocorlissia, Lambor- trichodina, Trichodina, Trichodinopsis, Urceolaria
nella, Monochilum, Ophryoglena, Tetrahymena, Vaginicola, Vorticella, Zoothamnium
Turaniella

Subclass 2. Scuticociliatia Small, 1967 Subclass 5. Apostomatia Chatton & Lwoff, 1928
Paroral dikinetid in three distinct segments, with Ciliary rows typically spiraled, widely spaced or
stomatogenesis via third and/or scutico-vestige; cilia- sometimes entirely missing; cytostome incon­
tion usually sparse, with thigmotactic area anteriorly, spicuous (or absent), usually associated with unique
caudal cilium posteriorly; mitochondria long and rosette; well-developed kinetodesmata; often poly­
sometimes fused into huge chondriome. Convention­ morphic life cycle, with most species ectosymbionts
ally, three orders: the philasterids, the pleuro- (phoronis) on marine crustaceans. Three orders.
nematids, and the totally symbiotic thigmotrichs. Ascophrys, Askoella, Chromidina, Collinia, Con-
Ancistrum, Ancistrumina, Boveria, Cinetochilum, idiophrys, Cyrtocaryum, Foettingeria, Gym­
Cohnilembus, Conchophthirus, Cyclidium, Dexio- nodinioides, Hyalophysa, Opalinopsis Ophiurae-
tricha, Dragescoa, Entodiscus, Fenchelia, Hemi­ spira, Phtorophrya, Vampyrophrya
speira, Histiobalantium, Hysterocineta, Loxocepha­
lus, M iamiensis, Myxophthirus, Parauronema, Subclass 6. Peniculinia Fauré-Fremiet in Corliss, 1956
Paurotricha, Peniculistoma, Philaster, Pleurocoptes, Buccal cavity contains paroral membrane, peniculi,
Pleuronema, Proboveria, Ptychostomum, Schizo­ and quadrulus; oral nematodesmata also present;
calyptra, Thigmocoma, Thigmophrya, Uronema, somatic dikinetids; cortical alveoli distinct; explosive
Urozona trichocysts; predominantly monomorphic, free-
living, fresh-water microphagous forms. Assigned to
Subclass 3. Astomatia Schewiakoff, 1896 the class Nassophorea (above) by some taxonomic
Mouthless forms, endosymbionts mostly in annelids ciliatologists.
(usually, but not exclusively, terrestrial oligochaetes) Clathrostoma, Disematostoma, Frontonia, Lem­
but one group in amphibians and turbellarians; fre­ badion, Marituja, Neobursaridium, Paramecium,
quently with well developed cortical endoskeleton, Stokesia, Urocentrum, Wenrichia
often with elaboration of some kind of holdfast or­
ganelle at anterior end of body. Two or three orders. Class (7) Prostomatea Schewiakoff, 1896
Anoplophrya, Buetschliella, Cepedietta, Clausilo- Mouth at or near anterior end of body, with relatively
cola, Contophrya, Durchoniella, Haptophrya, Hopli- simple oral ciliature; usually somatic monokinetids;
tophrya, Intoshellina, Lomiella, Maupasella, Radio- nematodesmata form rhabdos; toxicysts common; a
phrya, Steinella brosse characteristic of most species. Two orders,
Prostomatida Schewiakoff, 1896, and the much larger
Subclass 4. Peritrichia Stein, 1859 Prorodontida Corliss, 1974. Many of the old rhab-
Prominent oral ciliary field; somatic ciliature reduced dophorids are here, but some are in the following
to telotrochal band; widely distributed forms, many class a's well. The taxonomic place of a few genera
stalked and sedentary (though others mobile), some is controversial.
colonial, some loricate, all with aboral scopula; dis­ Bursellopsis, Coleps, Helicoprorodon, Holophrya
persal typically by migratory larval form (= telo- Metacystis, Nolandia, Placus, Plagiocampa
troch); often with strongly contractile myonemes, Planicoleps, Prorodon, Pseudobalanion, Pseudo-
body and/or stalk; fusion of micro- and macrocon- prorodon, Spathidiopsis, Tiarina, Urotricha, Vasicola
jugants. Symbiotic mobiline species have distinctive
denticulate ring on aboral surface of the body. Two
orders. Class (8) Litostomatea Small & Lynn, 1981
Apiosoma, Astylozoon, Carchesium, Cothurnia, E l­ Relatively inconspicuous or non-specialized oral
lobiophrya, Epistylis, Haplocaulus, Lagenophrys, ciliature; somatic monokinetids with two transversi
 "User-friendly" Classification of the Protists 17

jnicrotubular ribbons; kinetodesmata short, non-over­ Class (1) Perkinsidea Levine, 1978
lapping; oral ciliature derived from adjacent somatic Flagellated forms, typically with two unequal flagel­
kinetids with transverse microtubular ribbons sup­ la, but with most of apical complex organelles and
porting cytopharynx, latter (in many species) sur­ with cortical alveoli; large posterior vacuole with
rounded by nematodesmata (= rhabdos). Some diverse inclusions; some species with dinoflagellate-
species with toxicysts. Three or four subclasses with like trichocysts, some with contractile vacuoles;
number of orders, including groups classically known parasites of oysters or predators on various other
as haptorids, vestibuliferans/trichostomes, pleuros- protists. Two orders, one for each included genus.
tomes, and entodiniomorphids. Balantidium is the Colpodella (syn. Spiromonas), Perkinsus
only ciliate parasitic in humans. See also the com­
ments under Prostomatea, above. Class (2) Gregarinidea Dufour, 1828
Actinobolina, Alloiozona, Amphileptus, Arach­ Mature gamonts large, extracellular, exhibiting
nodinium,, Askenasia, Balantidium, Blepharocorys, syzygy, with production of essentially isogamous
Bryophyllum, Chaenea, Cycloposthium, Cyclotri­ gametes, but male gametes may be flagellated (with
chium, Didesmis, Didinium, Dileptus, Enchelys, basal body of nine singlet instead of usual eukaryotic
todinium, Gorillophilus, Isotricha, Lacrymaria, triplet micro tubules); zygotes undergo meiosis and
Lagynophrya, Lepidotrachelophyllum, Litonotus, sporogony within gametocystic membrane; trophonis
Loxophyllum, Mesodinium, O p hioscolex, Para- with mucron or epimerite; all species in digestive
bundleia, Paraisotricha, Phialinides, Plagiopyla, tract or body cavity of invertebrates or lower chor-
Pseudotrachelocerca, Pycnothrix, Quasillagilis, dates. Three or four orders recognized.
Rhabdoaskenasia, Rhinozeta, Sonderia, Spathidium, Actinocephalus, Ancora, Caulleryella, Cos-
Triadinium, Trichospira, Troglodytella, Vestibulon­ metophilus, Diplocystis, Doliospora, Gonospora,
gum Gregarina, Lankesteria, Lecudina, Monocystis,
Ophryocystis, Porospora, Rhynchocystis, Schizocys­
tis, Selenidioides, Selenidium, Siedleckia, Stepha-
Phylum 9. APICOMPLEXA Levine, 1970 nospora, S tylocephalus, Uradiophora, Zygocystis

Unicellular endosymbionts or predators characterized Class (3) Coccidea Leuckart, 1879

by having, at some stage in life cycle, an apical Gamonts typically intracellular; female gamont be­
complex typically composed of polar rings, rhoptries, comes macrogamete without division, syzygy
micronemes, and usually a conoid; highly com­ generally absent, microgametes many and with two
pressed smooth-membraned cistemae (= alveoli) or three flagella having basal bodies ultimately with
usually present in cell cortex of infective stage; sub- typical nine triplet microtubules; within oocystic
pellicular microtubules and micropores common; membrane, zygote produces sporoblasts which, in
flagella restricted (except in first class) to own membranes, produce two or more sporozoites;
microgametes or missing entirely; mitochondrial cris­ infective sporozoite invades host cell and, charac­
tae tubular, much reduced, or even absent. Except for teristically, grows and divides to form multiple
recent (and still bit disputed) addition of Perkinsus to merozoites capable of invading other host cells; even­
phylum, the whole assemblage remains essentially tually, some merozoites develop into gamonts, repeat­
identical to SPOROZOA Leuckart, 1879, the name ing cycle; highly resistant oocyst can survive outside
now treated by many workers as a synonym of host body (e.g. in soil) for long time before ingestion
APICOMPLEXA. Four classes, somewhat con­ and continued development and invasion, typically,
troversial, recognized here. Two classes proposed by of host’s gut epithelial cells. Most species
parasitologists (see Levine 1988) were subsequently monoxenous. Three orders (Eimeriida Léger, 1911 by
used by Corliss (1991c) as "group" names: Conoidea far the largest) commonly recognized.
for classes 2 and 3 (below), and Aconoidea for 4; to Adelea, Aggregata, Besnoitia, Caryospora,
which he added Zoosporea for class 1. This is the Coelotropha, Cryptosporidium, Cyclospora, Diplo­
third (and last: but see GLAUCOPHYTA?) phylum spora,, Dobellia, Dorisiella, Eimeria, Lrenkelia,
of the "Alveolata" assemblage (others are the Goussia, Grellia, Haemogregarina, Hepatozoon,
dinoflagellates and the ciliates: see above). Isospora, Karyolysus, Klossia, Klossiella,
18 J. O. Corliss

Lankesterelia, Legerella, Sarcocystis, Schellackia, belong here, as well as the baffling symbiotic Blas,
Selysina, Toxoplasma, Tyzzeria, Wenyonella tocystis (see comments on this genus under FUNGI)
and the curious apseudopodial Luffisphaera.
Class (4) Haematozoea Vivier, 1982 Acanthamoeba, Amoeba, Arcella, Balamuthia
Apical complex without conoid or conoidal rings and Cashia, Centropyxis, Chaos, Cochliopodium, Cucur-
rudimentary in other features; mitochondria simple bitella, Difflugia, Llabellula, Hartmannella, Hydra-
or absent entirely; motile zygote (= ookinete) moeba, Leptomyxa, Lesquereusia, Mayorella
penetrates vector-host’s gut wall, producing Nebela, Netzelia, Paramoeba, Platyamoeba, R0s-
numerous "naked" sporozoites which migrate to cuius, Saccamoeba, Stereomyxa, Thecamoeba
lumen of salivary glands, ready for transmission to Trichamoeba, Trichosphaerium, Vannella, Vexillifera
next definitive host; in formation of gametes, basal
bodies contain nine singlet microtubules but single Class (2) Entamoebidea Cavalier-Smith, 1991
flagellum produced exhibits typical 9 + 2 pattem. All With lobose pseudopodia, single nucleus, etc., but
species heteroxenous: merogony and formation of totally lacking mitochondria, peroxisomes, and
gamonts in blood cells of vertebrates; maturation of hydrogenosomes; small, if any, Golgi bodies; no
gametes, fertilization, and sporogony in gut of blood­ flagella; intranuclear centrosome present only during
sucking arthropods. mitotic prophase. Some workers have suggested that
Order 1. Haemosporida Danilewsky, 1885 this seemingly primitive group of symbiotic amoebae
Haemoproteus, Hepatocystis, Leucocytozoon, Plas­ may (better) belong in the kingdom Archezoa. It is
modium, Saurocytozoon not clear whether genera allegedly related to En­
Order 2. Piroplasmida Wenyon, 1926 tamoeba (e.g. Endamoeba, Endolimax, Iodamoeba)
Anthemosoma, Babesia, Dactylosoma, Echinozoon, should be assigned here or in subclass 1, above.
Theileria Dientamoeba - long placed in the family En-
tamoebidae - is now known, of course, to be a flagel-
Phylum 10. RHIZOPODA von Siebold, 1845 la-less member of the flagellate phylum
PARABASALA (above).
Non-flagellated (except for gametes of class 4), Entamoeba (plus any other genera?)
unicellular or plasmodial phagotrophs lacking aerial
sporangia; typically, pseudopodia serve in both Class (3) Filosea Leidy, 1879
locomotion and feeding; all non-photosynthetic Hyaline, filiform pseudopodia, sometimes branched
forms, except for groups with endosymbiotic algae; and occasionally anastomosing; some species naked,
Golgi bodies and mitochondria (generally with many with bottle-shaped tests. Several orders.
tubular cristae) always present except in class 2 where Amphorellopsis, Centropyxiella, Chardezia, Chlamy-
mitochondrial absence considered secondary; species dophrys, Cyphoderia, Euglypha, Gromia, Latero-
typically uninucleate (some exceptions) and free- myxa, Nuclearia, Ogdeniella, Paulinella, Penardia,
living (except for totally endosymbiotic forms of Pseudodifflugia, Sphenoderia, Trinema, Vampyrella
small class 2 and very few scattered other species).
Classically, phyla 10-12 sensu lato were combined Class (4) Granuloreticulosea de Saedeleer, 1934
under a super-taxon called the "Sarcodina". Granular, delicate, reticulate pseudopodia forming
anastomosing networks; few species naked, others in
Class (1) Lobosea Carpenter, 1861 single-chambered organic or calcareous test with no
Pseudopodia lobose or somewhat filiform; body often alternation of generations, but great majority in tests
naked, but also groups with tests (composed of or­ (organic, agglutinated, or calcareous) of one to many
ganic and/or inorganic materials, with single aper­ chambers with reticulopodia protruding from apon
ture). Predominantly free-living forms in soil, fresh­ tures and/or test wall perforations and with alternation
water, or marine habitats, widely distributed. Two of haploid sexual and diploid asexual generations,
subclasses and number of orders; but the class may known gametes uni- or biflagellated or a m o e b o i d ,
be a polyphyletic assemblage. The genera Copro- uni- or multinucleate forms, with asexual generation
myxa and Guttulinopsis (and some other former of some groups possessing dimorphic nuclei - one of
mycetozoa proving difficult to assign) might also more larger somatic nuclei and usually numerous
 >:til
"User-friendly" Classification of the Protists 19

'¡as- small generative nuclei, a heterokaryotic condition ally placed here, have been removed: the dimorphids
GD, ; reminiscent of that of ciliates; species phagotrophic to the Opalozoa (see above) and the ciliophryids to
(although some with endosymbiotic algae) and prac­ the Dictyochae (in kingdom Chromista, below). Class
thia, tically all marine, benthic forms; many more fossil 4 contains but one genus of unusual, small, biflagel­
CllT-
than contemporary genera described; class essentially lated marine forms possibly more closely related to
dra- composed of the foraminifers (Foraminiferea d’Or­ certain members of the next phylum, RADIOZOA.
ella, bigny, 1826: see genera below), with additionally Familiar name and conventional understanding of
Ros- included separate, but dubiously distinct, orders "the Heliozoa" maintained, although considerably
'eba, Athalamida Haeckel, 1862 (e.g. Arachnula and refined over classical usage; still, some workers
tfera Biomyxa) and Monothalamida Haeckel, 1862 (e.g. might prefer to elevate all or some of my classes to
Amphitrema, Lieberkuehnia, Micro gromia). Several independent phyletic status. Classically, the heliozoa
orders, numerous families, and many hundreds of s.I. and the radiozoa were combined under a super-
>îÿt genera (a few with living species listed below) of taxon called the "Actinopoda", based on their com­
and forams. Does Komokia (still) belong here? mon possession of axopodial pseudopodia.
; no Allogromia, Ammodiscus, Ammonia, Boderia,
Class (1) Actinophryidea Hartmann, 1913
Jring Bolivina, Carterina, Discorbis, Elphidium, Glabra­
Actinophrys, Actinosphaerium, Camptonema
Ithat tella, Globigerinella, Guttulina, Hastigerina,
»ebae Heterotheca, Iridia, Metarotaliella, Microglabratel­ Class (2) Centrohelidea Kühn, 1926
It is la, Myxotheca, Nonion, Ovammina, Patellinella, Acanthocystis, Actinocoryne, Cienkowskya, Gymnos­
En­ Planorbulina, Polystomella, Quinqueloculina, phaera, Hedraiophrys, Heterophrys, Raphidiophrys
jeba) Rhizammina, Rosalina, Rotaliella, Saccammina,
Class (3) Desmothoracidea Hertwig & Lesser, 1874
3ove. Schizammina, Schwagerina, Selenita, Sorites, Spiril­
Clathrulina, Hedriocystis, Orbulinella
En- lina, Spiroloculina, Textularia, Triloculina, Uvigerina
agel- Class (4) Taxopodea Fol, 1883
ylum Class (5) Xenophyophorea Schulze, 1904 Sticholonche
Relatively huge (up to 25 cm in diameter, although
only ca. one mm in thickness) but little studied marine Phylum 12. RADIOZOA Cavalier-Smith, 1987
benthic protists with multinucleate plasmodial stage
enclosed in branched-tube system which, in turn, is Typically spherical marine planktonic organisms,
iched within agglutinated test; presumably with filóse or often of large body size, characteristically having
aked, reticulose pseudopodia and biflagellated gametes. central capsule with pores; stiff axopodial
Some 36 species described from dozen genera; exact microtubules never in spiral pattern; endoskeleton
lami­ rank of taxon and its placement among other protists either siliceous or of strontium sulfate, in latter case
nero- remains uncertain. with radially arranged spicules; unicellular, oc­
ardin Galatheammina, Psammetta, Stannophyllum casionally colonial; single nucleus in early vegetative
relia stage, organisms often becoming multinucleate sub­
sequently; some species produce biflagellated
Phylum 11. HELIOZOA Haeckel, 1866 swarmer cells, not to be confused with symbiotic
rming dinoflagellates often present. Based on numerous
Lers in Unicellular phagotrophs with axopodia containing differences within the assemblage, phylum probably
ithno rigid microtubular axonemes; micro tubules typically best divided into two subphyla, with two classes in
i tests arrayed hexagonally, often nucleating on envelope of the second much larger and more familiar group.
many nucleus; kinetocysts common; mitochondrial cristae
aper- typically tubular; trophic stage usually without flagel­ Subphylum 1. Acantharia Haeckel, 1881
nation la; short filopodia in some species; several groups
itions; with stalks, one with perforated shell or test; mostly Class Acantharea Haeckel, 1881
sboid; fresh-water, some marine. Controversial whether four Acanthochiasma, Acantholithium, Acanthometra,
nation classes named below are closely enough interrelated Amphilonche, Astrolonche, Astrolophus, Haliom-
one or to warrant being clustered into one phylum; two other matidium, Lithoptera, Pleuraspis, Pseudolithium,
íerous taxa (of once-called "helioflagellates"), convention­ Xiphacantha
 T
20 J. O. Corliss

Subphylum 2. Radiolaria J. Müller, 1858 session of "spores" with polar filaments inside. ge
cause of their pluricellular stages, and the similarity
Class (1) Polycystinea Ehrenberg, 1838 (in development) of their polar capsules with
Cenosphaera, Coccodiscus, Collosphaera, Col­ cnidarian nematocysts, the MYXOZOA are placed
lozoum,, Halosphaera, Octodendron, Plagiacantha, in the Animalia by some taxonomists. Single class
Rhizosphaera, Spongodrymus, Thalassicolla, 77za-
lassophysa Class Myxosporea Bütschli, 1881
Ceratomyxa, Chloromyxum, Fabespora, G/ofo-
Class (2) Phaeodarea Haeckel, 1879 spora, Henneguya, Hoferellus, Kudoa, Lomosporus
Atlanticella, Aulacantha, Aulosphaera, Aulotractus, Myxidium, Myxobolus, Myxoproteus, Ortholinea
Castanella, Challengeriae Challengeron, Coelo­ Parvicapsula, Sinuolinea, Sphaeromyxa, Sphaero­
dendrum, Conchopsis, Halocella, Medusetta, spora, Trilospora, Unicapsula, Unicauda, Ward/a
Phaeodina Zschokkella

Phylum 13. MYXOZOA Grassé, 1970 Phylum 14. ASCETOSPORA Sprague, 1978

Symbiotic forms with valved multicellular spores Endosymbionts of (mainly) marine invertebrates,
having polar capsules with extrusible filaments; spores unicellular or with production of cells
trophic stages amoeboid (binucleate sporoplasm) or (sporoplasms) within cells; no polar capsules or fila­
plasmodial (multinucleate); no flagellated stage; ments; no flagellated stage in life cycle;
mitochondria with tubular to irregular-shaped cristae; mitochondrial cristae tubulo-vesicular; unique
somatic and generative nuclei somewhat reminiscent haplosporosomes characteristic of most included
of condition in ciliates, some foraminifereans, and species. Small but perhaps polyphyletic assemblage
some radiolarians; commonly coelozoic or histozoic requiring more study; here it is tentatively considered
in marine and fresh-water fishes, but a number of to embrace one or both of the groups Paramyxidea
fresh-water species are found in body cavity or intes­ Chatton, 1911 and Marteiliidea Desportes &
tinal epithelium of aquatic oligochaetes, probably Ginsburger-Vogel, 1977 (as well as the haplo-
undergoing an alternate stage in full life cycle of sporidians proper), without giving them specific
fresh-water fish symbionts. Assemblage contains a ranks. Nephridiophaga no longer here (Lange 1993)?
number of orders; but conventional breakdown into
two major groups, Myxosporidia Bütschli, 1881 and Class Haplosporidea Caullery & Mesnil, 1899
Actinomyxidia Stoic, 1899, has recently become Haplosporidium, M arteilia, Minchinia, Paramar-
highly suspect because of findings, confirmed ex­ teilia, Paramyxa, Urosporidium
perimentally, that some forms formerly assigned to
each are only stages in the life cycle of single
myxosporidian species that seem to require two hosts. Phyla of Kingdom CHROMISTA
Tetractinomyxon sipunculid symbiont, may survive
by transfer to a myxosporidian order. No ac- Four subkingdoms are recognized here in apprecia­
tinomyxidian generic names are listed below since, tion of significant differences among members of
often being of more recent date than myxosporidian their included taxa. Some workers may prefer com­
ones and/or legally possibly only "collective" names, pletely independent status for these four groupings,
they may well be suppressed in future taxonomic but the last three are relatively very small and arc
works (Kent et al. 1994). Phylum contains several probably best treated as I have done below; at least,
orders; but I consider the enigmatic Helicosporidium they ought to be appended in some way to the
of the literature, sometimes placed here, to be a Chromista until/unless further comparative data
member of the kingdom Fungi. Classically, the clearly indicate otherwise. The general relationship
myxosporidians s.L and the microsporidians (see of the informal group of "stramenopiles" (Patterson
kingdom Archezoa) were lumped together under the 1989a) to this kingdom is considered in my DISCUS'
name "Cnidosporidia", based on their common pos­ SION section.
 "User-friendly" Classification of the Protists 21

Subkingdom (I) HETEROKONTA Luther, 1899 Phylum 3. DICTYOCHAE Haeckel, 1894

■Be-
anty Essentially with characters of the kingdom sensu Mixture of pigmented and non-pigmented
with stricto: organisms with chloroplasts (unless secon­ heterokonts, free-swimming or stalked, marine and
laced darily lost) located within rough endoplasmic fresh-water habitats; often with anteriorly directed
ass. reticulum instead of free in cytosol, and with mas­ tentacles; one apically inserted flagellum typically
tigonemes (unless flagella lost) as rigid, tripartite, with two rows of tripartite mastigonemes and some­
tubular hairs, on one or both flagella, functioning in times small scales; second flagellum often reduced to
lobo- thrust reversal. Bulk of the chromist species belong basal body; some species (silicoflagellates, mostly
orus,
to phyla of this well-established and nearly univer­ marine fossil forms) with complex basket-shaped
inea,
sally accepted major taxonomic assemblage of "the external siliceous skeleton. Two classes. The second,
ciero-
heterokonts": the name - in one form or another (e.g. whose members have long been studied by
irdia,
some workers call it the HETEROKON­ phycologists, is here credited to Kristiansen (1990),
TOPHYTA) - and the concept are both deserving of although Cavalier-Smith (1986) had also established
preservation. it as a class some four years earlier and Möhn (1984)
and Karpov (1990) have both independently con­
sidered it as a new class (and a new phylum as well!).
Phylum 1. BICOSOECAE Cavalier-Smith, 1989 The pedinellids s.L, which include some of the
rates,
"helioflagellates" (see Davidson 1982) of the litera­
cells
Small, free-living, fresh-water or (few) marine, ture, may be polyphyletic; Cavalier-Smith (1993c)
r fíla­
planktonic biflagellated (one flagellum with mas- has at least partially relieved that condition by remov­
mele;
: tigonemes), heterotrophic non-pigmented forms typi­ ing Oikomonas to a separate class of its own, although
nique
cally living in loricae (attached by the second, smooth I have not done so here. Some workers vernacularly
luded
blage flagellum), and feeding on bacteria; some stalked, refer to (some or all of) the pedinelleans (or pedinel-
dered some form colonies. lophyceans) as actinomonads.
xidea Class (1) Silicoflagellatea Borgert, 1891
ís & • Class Bicosoecidea Grassé & Deflandre, 1952
Dictyocha (only genus with living species)
laplo- j Bicosoeca, Cafeteria, Pseudobodo
ecific Class (2) Pedinellea Kristiansen, 1990
993)? Actinomonas, Ciliophrys, Oikomonas, Parapedinel­
Phylum 2. LABYRINTHOMORPHA la, Pedinella, Pseudopedinella, Pteridomonas
j Page in Levine et al., 1980
jmar- Phylum 4. RAPHIDOPHYTA Chadefaud, 1950
Non-pigmented protists, trophic stage with ectoplas-
mic network of spindle-shaped or spherical non- Biflagellated forms with or occasionally without
amoeboid cells that move by gliding within network; plastids, fresh-water and marine; motile or palmelloid
unique cytoplasmic organelles, bothrosomes (first unicells; Golgi bodies in ring, over anterior surface
described as sagenetosomes); tubular mitochondrial of nucleus; unique extrusome in many species.
recia- cristae; known zoospores biflagellated, one bearing Phylum is also known as "the chloromonads"; but this
ers of mastigonemes, other naked; in marine, generally is inappropriate, because Chloromonas is a genus of
com- coastal, waters, often associated with, or ectosym- green algal protists in the kingdom Plantae.
pings, bionts of, aquatic angiosperms and certain algal
id are protists. Class Raphidomonadea Chadefaud, 1950
least, Chattonella, Gonyostomum, Heterosigma, Mero­
o the Class (1) Labyrinthulea Cienkowski, 1867 tricha, Olisthodiscus, Vacuolaria
data I Labyrinthula
mship Phylum 5. PHAEOPHYTA Wettstein, 1901
terson Class (2) Thraustochytriacea Sparrow, 1943
cus- Aplanochytrium, Labyrinthuloides, Thraustochy­ Photosynthetic heterokonts predominantly with
trium chlorophylls a plus c and leucosin and fat (or para-
22 J. O. Corliss
1
mylon, glucose, or laminarin) as storage products; Class (6) Xanthophyceae Allorge in Fritsch, 1935
often siliceous scales covering body; charac­ (syns. Heterochloridea Pascher, 1912 p D
teristically, a pair of flagella with anteriorly project­ Tribophyceae Hibberd, 1981)
ing one bearing rigid tubular mastigonemes; some Botrydiopsis, Brachynema, Bumilleriopsis
loricate species; many fresh-water forms with distinc­ Gloeobotrys, Gloeopodium, Heterogloea, ]yja^
tive statospore; other groups almost entirely marine lodendron, Ophiocytium, Pleurochloris, Retículos
(e.g. brown algae); diverse morphological types: phaera, Tribonema, Vaucheria
unicellular (some amoeboid), colonial, filamentous
or thalloid (multicellular); sizes small to very large
(brown seaweeds, kelp, up to 60 meters in length). Phylum 6. DIATOMAE Agardh, 1824
Entire assemblage is essentially the golden-brown
(plus some yellow-green) algae of the literature Pigmented unicells (occasionally colonial) with
minus diatoms, silicoflagellates, and haptophytes but secreted silica frustule consisting of two valves and
including the browns and eustigmatophytes (although one or two girdle bands; non-flagellated except for
latter without chlorophyll c, usually possess but single posterior flagellum on microgametes of one
single flagellum, and eyespot independent of group (gametes of other groups amoeboid); yellow-
chloroplast). Conservatively, I include six major clas­ brown plastids; mainly planktonic forms widespread
ses, which contain a number of orders and numerous in fresh-water and especially marine habitats, with
genera and species; but additional classes may be numerous fossils; some species in moist soils. Many
justified (e.g. for Reticulosphaera and Vaucheria, in thousands of diatoms have been described, usually as
class 6 below). Zoologists have traditionally claimed belonging to the botanical division conventionally
a number of motile chrysomonads s.l. as members of known as BACILLARIOPHYTA Engler & Gild,
the "old" Protozoa, assigning them mostly to a single 1924, and assigned to two major distinct groups
order (Chrysomonadida Engler, 1898). (centric and pennate); but at least three classes (Cos­
cinodiscophyceae Round & Crawford, 1990;
Class (1) Phaeophyceae Kjellman, 1891
Fragilariophyceae Round, 1990; Bacillariophyceae
(syns. Melanophyceae Rabenhorst, 1863,
Haeckel, 1878) are now recognized, with numerous
Fucophyceae Warming, 1884)
subclasses and orders plus several hundred genera
Alaria, Arthrocladia, Chordaria, Costaria, Cys­
(Round et al. 1990).
toseira, Dictyota, Ectocarpus, Fucus, Giffordia,
Achnanthes, Amphipleura, Amphora, Auricula,
Hormosira, Laminaria, Litosiphon, Macrocystis,
Bacillaria, Bacteriastrum, Biddulphia, Coscinodis­
Myrionema, Sargassum, Scytosiphon, Sorocarpus,
cus, Cyclophora, Cyclotella, Cylindrotheca,
Sporochnus, Stilopsis, Streptophyllum, Utriculidium,
Cymatopleura, Cymbella, Diadesmis, Diatoma,
Xiphophora, Zonaria
Fragilaria, Frustulia, Grammatophora, Gyrosigma,
Class (2) Chrysophyceae Pascher, 1914 Hanzschia, Hemidiscus, Hydrosilicon, Lennoxia, Lic­
Anthophysa, Chromulina, Chrysamoeba, Chryso­ mophora, Lithodesmium, Lyrella, Melosira, Minuto­
capsa, Chrysococcus, Chrysodendron, Dermato­ cellus, Navicula, Nitzschia, Odontella, Pleurosigma,
chrysis, Dinobryon, Epipyxis, Hibberdia, Podosira, Rhizosolenia, Stephanodiscus, Sticho­
Microglena, Monochrysis, Ochromonas, Poterio­ chrysis, Thalassionema, Thalassiosira, Toxarium,
ochromonas, Rhizochromulina (maybe better in class Triceratium
4?), Sarcinochrysis, Spumella, Triparma, Uroglena
Class (3) Synurophyceae Andersen, 1987
Phylum 7. PSEUDOFUNGI Cavalier-Smith, 1986
Mallomonas, Mallomonopsis, Synura, Tesselaria
Class (4) Pelagophyceae Andersen & Saunders, 1993 Osmotrophic, minute symbionts on other protists and
Pelagococcus, Pelagomonas aquatic plants or in hosts ranging from grapes and
potatoes to fishes; in fresh-water (mostly) or marine
Class (5) Eustigmatophyceae Hibberd & Leedale, 1970 habitats or in soil; bi- or uniflagellated z o o s p o r e
Chlorobotrys, Eustigmatos, Monodopsis, Nan­ stage; uninucleate or coenocytic walled protoplast in
nochloropsis, Pseudocharaciopsis, Vischeria vegetative stage. Long conventionally considered a>
1

"User-friendly" Classification of the Protists 23

a class in the kingdom FUNGI along with the chytrid Phylum HAPTOMONADA Cavalier-Smith, 1989
protists, which do belong there. Synonymous names
for the group include OOMYCOTA Dick, 1990 and With characters of subkingdom. Essentially
PSEUDOMYCOTA Barr, 1992. The latter name, synonyms for the name of (most of) the assemblage
like PSEUDOFUNGI, is also attractive in emphasiz­ are COCCOLITHOPHORA Lemmermann, 1903,
ing the pseudo-fungal characters of these heterokont and PRYMNESIOPHYTA Casper, 1972 (latter
protists, which are quite unlike the so-called name generally credited to "Hibberd, 1976", where
"Eumycota" or true fungi. Two classes are recog­ first Latin diagnosis is to be found). Contains two
nized. classes: first (Pavlovea Cavalier-Smith, 1986) for the
allegedly primitive genus Pavlova\ second (Patel-
liferea Cavalier-Smith, 1993) for all other genera (a
Class (1) Oomycetes Winter in Rabenhorst, 1879 few of which are given below). If future ultrastruc­
Zoospores typically with two flagella, anterior one tural and molecular studies show still greater dif­
with two rows of rigid mastigonemes, posterior ferences compared with the Chromista proper, as­
smooth or with only fine flexuous hairs; cytoplasmic semblage can be redefined and elevated to separate
and nucleus-associated microtubules. kingdom or assigned elsewhere (e.g. to the Protozoa,
Achlya, Albugo, Brevilegnia, Lagenidium, Lep­ where it was at one time included in an order Coc-
tomitus, M yzocytium, Olpidiopsis, Peronos- colithophorida).
clerospora, Peronospora, Phytophthora, Pythium, Calciosolenia, Canistrolithus, Chrysidalis, Chryso­
Rhipidium, Saprolegnia, Sclerospora, Verrucalvus, chromulina, Coccolithus, Emiliania, Isochrysis,
Zoophagus Ophiaster, Phaeocystis, Pleurochrysis, Prymnesium,
Umbilicosphaera
Class (2) Hyphochytriomycetes Sparrow, 1959
Zoospores with single, anterior flagellum (with mas­
tigonemes); cytoplasmic and nucleus-associated Subkingdom (III) CRYPTOPHYTA Pascher, 1914
microtubules absent. Following recent convention, I
have dropped the "id" originally in the class name Group of protists mostly photosynthetic, unicellular,
between the "tr" and the "io". and motile (biflagellated, with bipartite mas­
Anisolpidium, Hyphochytrium, Rhizidiomyces tigonemes generally on both); usually paired
thylakoids, chlorophylls a and c, and two phycobilins;
unique features: nucleomorph, ejectosome, and
Subkingdom (II) HAPTOPHYTA Christensen, 1962 periplast; mitochondrial cristae flattened; distinct gul­
let; chloroplast endoplasmic reticulum typically
Typically photosynthetic unicellular biflagellated present; fresh-water and marine habitats. By
protists characterized principally by possession of a zoologists at one time considered an order, Cryp­
haptonema, unique filiform appendage located be­ tomonadida Senn, 1900, of the "old" Protozoa.
tween anteriorly arising flagella, often very long
(sometimes coiled) and containing 6-8 singlet Phylum CRYPTOMONADA Ehrenberg, 1838
microtubules; atypical of kingdom, neither flagellum
bears tubular mastigonemes; commonly two parietal With characters of subkingdom. Contains two clas­
plastids, each with single pyrenoid; chloroplast en­ ses: first (Goniomonadea Cavalier-Smith, 1993) for
doplasmic reticulum present; with rare exception, allegedly primitive forms (e.g. phagotrophic
body covered by layers of small organic scales in turn goniomonads); second (Cryptomonadea Stein,
often covered by large unmineralized scales (coc- 1878) for all others. As in the cases of subkingdoms
coliths) on which calcium carbonate crystallized as II (above) and IV (below), future molecular data may
calcite or aragonite; single Golgi body, fan-shaped indicate a different phylogenetic/taxonomic place­
near anterior end of cell; mitochondrial cristae ment for this assemblage.
tubular; mostly marine, few fresh-water; a few Chilomonas, Chroomonas, Cryptomonas, Gonio­
species form colonies, and a few exhibit phagotrophy; monas, Hemiselmis, Pyrenomonas, Rhinomonas,
many fossil forms. Rhodomonas
24 J. O. Corliss

Subkingdom (IV) CHLORARACHNIOPHYTA taxa of protists, the green algae s.l. (but excluding
Hibberd & Norris, 1984 entirely the unrelated euglenoids of the kingdom
Protozoa), are unicellular (generally biflagellated
Marine photosynthetic protists with amoeboid plas­ without tubular mastigonemes), colonial, or filamen­
modial vegetative stage, with individual cells linked tous (multicellular), many without motile vegetative
by fine filopodia; uniflagellated zoospore (thus = an stages; all with starch-containing plastids bounded by
amoeboflagellate?) with its flagellum, coiled helical­ an envelope of two membranes; unmineralized scales
ly around cell body, bearing delicate mastigonemes; on bodies or flagella of many species; found
mitochondrial cristae tubular; chlorophyll a and b but predominantly in fresh-water habitats, but some en­
no c nor phycobilins; outermost membrane around tire groups marine. Modern phycologists are of
chloroplast lacking ribosomes on cytosolic face; diverse opinions concerning the exact num­
complex extrusomes. bers/names of high-level taxa (phyla/divisions, clas­
ses/subclasses) to be included here: four phyla are
Phylum CHLORARACHNIOPHYTA endorsed below. As mentioned above, the non-protist
Hibberd & Norris, 1984 phyla of the VIRIDIPLANTAE (i.e. the "higher" or
"land" plants, bryophytes and tracheophytes) are
With characters of subkingdom. Contains single beyond consideration in this paper.
class, Chlorarachniophyceae Hibberd & Norris,
1984. Taxonomic position of the organism remains Phylum 1. PRASINOPHYTA Christensen, 1962
somewhat controversial, as well as the most ap­
propriate name, authorship, and date for it at the The "grass-green scaly algae" (and close relatives),
various supraordinal levels. Tentatively, I am using typically small biflagellated unicells, presumably
the identical name for the subkingdom and phylum most primitive group among plant protists; organic
here. Single species? And no closely related (other) scales, with rare exceptions, on body and/or flagella;
genera? generally no cell walls; often unique extrusomes.
Chlorarachnion
Class (1) Pedinophyceae Moestrup, 1991
Phyla of Kingdom PLANTAE No scales; second flagellum represented by only its
basal body. Two genera.
The non-protist plant phyla (viz. BRYOPHYTA, Pedinomonas, Resultor
PTERIDOPHYTA, SPERMATOPHYTA) are
beyond the scope of this paper, so they are not further Class (2) Prasinophyceae Christensen, 1962
considered here. The protists of the kingdom are (syn. Micromonadophyceae Mattox & Stewart,
divided into two groups at the high level of sub­ 1984)
kingdom, the first - and much larger - assemblage Essentially with general characters of phylum s.s.
containing the green algae of the literature (along Three orders recognized. In the past, some species
with the "higher" plants proper, which clearly evolved have been considered members of the "old" Protozoa
from them) and the second the taxonomically enig­ by zoologists.
matic red algae plus, possibly, the even more refrac­ Bathycoccus, Dolichomastix, M amiella, Mantoniella,
tive glaucophytes. Further research may yield addi­ Mesostigma, Micromonas, Nephroselmis, Pseudo­
tional data that will make untenable these proposed scourfieldia, Pterosperma, Pyramimonas, Scourfiel­
"taxonomic marriages"; in which case, appropriate dia, Tetraselmis
classificational alterations can easily be made.
Phylum 2. CHLOROPHYTA Pascher, 1914
Subkingdom (I) VIRIDIPLANTAE
Cavalier-Smith, 1981 The "green algae" s.s. of the literature; many non-
motile species; motile ones usually bi- or quadriflagd'
Typically photosynthetic organisms with chloro­ lated, walled or naked; morphological types include
phylls a and b and flattened mitochondrial cristae; unicellular or colonial, tetrasporal, coccal, sarcinoid>
cellulosic cell walls common. Species of included filamentous, and parenchymatous. Single class
 "User-friendly" Classification of the Protists 25

(Chlorophyceae Wille in Warming, 1884) with per­ bers of this first class possess phragmoplast similar
haps a dozen separate orders. Traditionally, zoologists to that of "higher" plants, and their cellulosic cell
have claimed a number of motile species (outstanding walls sometimes heavily calcified. Species of second
examples, Volvox and Chlamydomonas) as protozoa, class, with no flagellated stages in life cycle, show
assigning them to the order Volvocida Francé, 1894 unique conjugation between cells (alone or of closely
(replacing the highly inappropriate name appressed filaments), with fusion of amoeboid
phytomonadida Blochmann, 1895, since Phytomonas gametes; in the essentially mirror-image unicellular
is in the protozoan class Kinetoplastidea). desmids, a pair of large, complex plastids are joined
Aphanochaete, Botryococcus, Carteria, Chaeto­ at an isthmus that contains a single shared nucleus;
chloris, Chlamydomonas, Chlorella, Chlorococcum, cellulosic cell walls often slimy, organisms gliding
Chlorogonium, Chloromonas, Coccomyxa, Coela­ on the secreted mucilage. Evolutionarily, members
strum, Dunaliella, Eudorina, Fritschiella, Gloeocys­ of the first class are considered directly ancestral to
tis, Gonium, Haematococcus, Hydrodictyon, Micro­ "higher" plants. Some taxonomic phycologists have
spora, Nanochlorum, Nautococcus, Palmodictyon, separated the two groups named below at phylum
Pascherina, Pediastrum, Phacotus, Pleodorina, (division) rather than class level.
Pleurastrum, Polytomella, Prasiola, Protosiphon,
Scenedesmus, Schizomerus, Selenastrum, Sphaero­ Class (1) Charophyceae Rabenhorst, 1863
plea, Stephanosphaera, Tetraspora, Tetrasporidium, With characters given above for the first class.
Trebouxia, Trentepohlia, Treubaria, Trichophilus, Vb/- Chaetosphaeridium, Chara, Chlorokybus, Coleo­
var, Yamagishiella chaete, Klebsormidium, Nitella, Nitellopsis, Raphido­
nema, Stichococcus, Tolypella
Phylum 3. ULVOPHYTA Stewart & Mattox, 1978
Class (2) Conjugatophyceae Engler, 1892
Most species macroscopic seaweeds (including "sea With characters given above for the second class. The
lettuce") from tropical marine waters; sessile with larger group, mostly because of huge number o
walled vegetative cells, thalli typically coenocytic or described desmids. Essentially synonymous names
multicellular; bi- or quadriflagellated reproductive include "Conjugaphyceae", Gamophyceae, Zyg­
cells common; morphology ranges from sarcinoid nematophyceae, and Zygophyceae.
and blade-like to siphonous. Single class (Ul­ Ancylonema, Arthrodesmus, Closterium, Cosmarium,
vophyceae Stewart & Mattox, 1978) and five orders Cylindrocystis, Desmidium, Micrasterias, Oocar­
recognized. dium, Sirogonium, Spirogyra, Staurastrum, Xan­
Acetabularia, Acrosiphonia, Blidingia, Bryopsis, thidium, Zygnema
swart, Chaetosiphon, CladophoraClaclophora, Codium,
Cymopohlia, Dasycladus, Eugomontia, Halimeda, Subkingdom (II) BILIPHYTACavalier-Smith, 1981
Phaeophila, Rhizoclonium, Siphonocladus, Tricho­
sarcina, Ulothrix, [7/va, Valonia Essentially the "red algae" of the literature. Unlike
other members of the kingdom PLANTAE in many
Phylum 4. CHAROPHYTA Rabenhorst, 1863 respects, their species also show little similarity to
other taxa of protists. Whether they are "algal plants",
Some species multicellular, macroscopic, and found as considered here, or better treated as an independent
submerged in shallow fresh-water habitats, a few kingdom is a matter for the future when additional
terrestrial, but majority (including the ubiquitous des- relevant data become available. Mostly marine
mids) unicellular or filamentous in fresh waters protists, some unicellular, others of macroscopic size
everywhere; the larger species, some commonly (length) - latter, like many brown (and a few green)
known as stoneworts, have macroscopic thalli with algae, called seaweeds; particularly distinguished by
/ non- main axis erect plus regular whorls of lateral total absence of centrioles and flagella and by
flagel- branches, and with male and female sex organs presence of single thylakoids in their chlorophyll a
nclude reminiscent of those of land plants: these charophytes containing plastids with phycobilins as accessory
cinoid, have motile (flagellated) swarmers, never with photosynthetic pigments; mitochondria with flattened
class eyespot, typically covered with scales; many mem­ cristae; starch stored in cytosol; often complex life
 1

26 J. O. Corliss

histories. Some workers accept - as a second phylum parasitologists and medical clinicians without ques
of the BILIPHYTA, in addition to the RHODO­ tion for over half a century. Recently, it has been
PHYTA - the enigmatic and possibly non-mono- (re)classified as a sporozoon (Apicomplexa), a
phyletic GLAUCOPHYTA Bohlin, 1901 (single lobosean amoeba (Rhizopoda), an "uncertain protist”
class, Glaucophyceae Bohlin, 1901, with genera and a unique organism requiring a new phylurn
Cyanophora, Glaucocystis, Glaucosphaera, Gloeo­ (named as a "protozoan subphylum": Blastocysta
chaete). Glaucophytes are small, fresh-water, Jiang & He, 1993) of its own, but with some workers
cyanelle-containing protists commonly with a pair of still considering it a fungus. See Belova (1992)
flagella in their life cycle, cortical alveoli (curious­ Boreham and Stenzel (1993), Garavelli and Libanore
ly!), etc., sharing some characters with red algae (e.g. (1993), Jiang and He (1993), Johnson et al. (1989)
possession of phycobiliproteins). The cyanelles are Zierdt (1988, 1993), and references within those
presumably evolutionarily derived from blue-green papers. I favor assignment to, or near, the rhizopod
algae (i.e. endosymbiotic cyanobacterial prokaryotes) class Lobosea (q.v.), at an undetermined rank, until
on their way to becoming genuine plastids in more comparative data of phylogenetic significance
glaucophytes and probably in several other taxa of are available on this taxonomically defiant organism.
protists as well).
Phylum CHYTRIDIOMYCOTA Sparrow, 1959
Phylum RHODOPHYTA Rabenhorst, 1863
Protists with definite fungal affinities: non-pigmented
Essentially with characters of subkingdom, as given forms (some filamentous) with chitinous cell walls in
above. Some workers recognize two principal clas­ hyphal stage, flat mitochondrial cristae, absorptive
ses: the more primitive and much smaller group, the mode of nutrition, symbionts or saprobes in soil or
Bangiophyceae Wettstein, 1901 ; and the widespread, fresh-water habitats; atypical of (the majority of) the
multicellular, much larger group, the kingdom, however, are such characteristics as their
Florideophyceae Warming, 1884. Numerous orders motile stages in life cycle (most gametes and some
have been described. asexual zoospores) with posteriorly directed single
Audouinella, Bangia, Bangiopsis, Batrachospermum, (rarely multiple) flagellum (without mastigonemes or
Boldia, Callocolax, Capreolia, Chondrus, Com­ scales), frequent unicellularity, and possession of
psopogon, Cyanidium, Dilsea, Endocladia, Erythro­ unusual cytoplasmic structures in many species (e.g.
trichia, Gigartina, Goniotrichum, Gracilaria, distinctive flagellar root system and the curious rum-
Halymenia, Heteroderma, Hildenbrandia, Iridaea, posome in members of two orders). The chytrids s í
Lithophyllum, Mesophyllum, Minium, Naccaria, P^/- differ significantly from members of the phylum
maria, Phragmonema, Phyllophora, Porolithon, P 6>r- PSEUDOFUNGI (q.v.), heterokonts of the kingdom
phyridium, Rhodella, Rhodochaete, Rhodophyllis, Chromista.
Rhodospora, Sporolithon, Thorea, Zymurgia
Class Chytridiomycetes Sparrow, 1959
Phyla of Kingdom FUNGI With characters of phylum. Four orders recognized.
Allomyces, Blastocladiella, Callimastix, Catenaria,
The non-protist fungal phyla (viz. ASCOMYCOTA, Chytridium, Chytriomyces, Coelomomyces, Karlin­
BASIDIOMYCOTA, ZYGOMYCOTA) are gia, Monoblepharella, Neocallimastix, Olpidium,
beyond scope of this paper, so are not treated here. Physoderma, Rhizophydium, Spizellomyces, Synchy-
They embrace ’’typical" fungal forms, the Fungi Im­ trium
perfecti, unicellular yeasts, and probably also the
enigmatic "protozoon" Helicosporidium and the Phyla of Kingdom ANIMALIA
taxonomically notorious Pneumocystis (but see note
of caution by Frenkel et al. 1990). Blastocystis, a Since none of the many phyla of animals - in my
common intestinal symbiont of many vertebrates in­ opinion - contains any protist species, they are beyond
cluding humans, was first discovered and described our consideration here. But, from time to time, sporn
more than 75 years ago as a "vegetable organism" (= ges (if one classifies the choanozoa there) have rep'
fungus), a taxonomic conclusion accepted by resented a possible exception. And, very recently*
 "User-friendly" Classification of the Protists 27

Cavalier-Smith (e.g. 1993c) has suggested that the considerably elevated over the years (especially in
multicellular, ciliated MESOZOA, with (usually) recent decades), the numbers of their species have in­
tubular mitochondrial crisiae and lack of collagenous creased dramatically over time with advances in micros­
connective tissue, should be removed from the copy, and the contents and boundaries of the assemblage
Animalia to the protistan kingdom Protozoa. This have often changed with our increased understanding of
taxonomic shift has not been endorsed, however, in them.
the present paper. Some place the myxozoa here. Some 15-20 years ago (see historical account in
Corliss 1986a), the "protist revolution" began to per­
meate the thinking of the biological scientific com­
DISCUSSION munity. In due time, it became unfashionable to retain
a phylum or subkingdom (of "animals") called the
As inferred in the INTRODUCTION and manifest Protozoa in light of our new appreciation of inter­
throughout the preceding pages of classification, relationships among (former) algal and protozoan
biologists can no longer think of protozoa or protists as groups, an intermingling that finally forced the break­
conveniently divisible into separate taxa based on down of the old plant/animal barriers in taxonomy. A
general characteristics such as modes of locomotion or neoHaeckelian kingdom Protista helds way, as it does
types of nutrition. In other words, no longer can there be still today in many circles. Round (1980) perceptively
named high-level taxonomic groups containing, for ex­ realized that we were going too far in discarding genuine
ample, only forms with pseudopodia or with flagella or differences between many algal and protozoan taxa, but
with chloroplasts or with a totally symbiotic style of life. his warning was not heeded at the time.
Our attempts to erect "natural" systems of classification With the emphasis on broad phylogenetic lines and
have now moved far beyond that stage, thanks primarily the desire to break completely with the past, most protis-
to the availability of more sophisticated ways of studying tologists failed to realize what we know today (but some
the properties/characteristics of these generally unicel­ still find difficult to accept), that the Protista are too
lular and microscopic "lower" eukaryotes. diverse to remain as a single taxonomic entity and that
There is no need (nor space!) to discuss all parts of some older concepts, properly refined, need not remain
the preceding classification scheme in any detail here. discarded. The Protozoa represent an outstanding ex­
It is clear that the general basis or rationale for arranging ample of this. As Cavalier-Smith (1993c, in particular)
the taxa as I have done is the degree to which various has resurrected the group - as a kingdom PROTOZOA
groups do, or do not, share key characters in common, - it deserves (re)acceptance, in my opinion (although a
reflecting their phylogenetic affinities. Different ap­ year ago I myself raised some objection to his choice of
proaches or schools of thought have been mentioned in name for the new kingdom, while tacitly admitting that
the INTRODUCTION; I consider myself an evolution­ no better one came to mind: Corliss 1993). Its boun­
ary biologist sensu Mayr (1990). Comments regarding daries have been sharpened by removal of several taxa
various controversial taxonomic decisions have been formerly inappropriately assigned to it: for example,
made in place on preceding pages. Here, I wish to focus groups that now reside, quite properly, in other
attention on four matters that deserve additional ex­ kingdoms. The ARCHEZOA represent a perfect haven
planation or discussion: my choice of the names and for the primitive amitochondrial groups of certain
concepts for the kingdoms PROTOZOA and amoebae, symbiotic flagellates, and the unique
CHROMISTA; consideration of phylum and class microsporidians. The CHROMISTA embrace certain
flames in general (including authorships and dates); the algal protists (e.g. chrysophyceans s.L and haptophytes)
taxonomic category of "incertae sedis"; and my reasons totally different from the (former) algal groups of
for supplying so many examples (so evident in the euglenoids and dinoflagellates (sensibly treated as
INDEX as well as in the text) of included genera. protozoa now) but many of which were (also) once
classified as protozoa. And the PLANTAE are the
in my The Kingdom PROTOZOA proper place for the green algae, which harbor the
beyond ancestors of the "higher" plants but some of which were
sporn The Protozoa, united into a formal group of generally traditionally labeled, simultaneously, as protozoa (e.g.
ve rep' ^croscopic, unicellular, phagotrophic forms, have been Chlamydomonas, Volvox, and their close relatives: the
îcently» ^ound for 175 years. The rank accorded them has been "phytomonads" or, better, the volvocids).
28 J. O. Corliss

Thus cleansed or purged, the Protozoa have become identical to most members of large assemblages giVen
a much more homogeneous, if, admittedly, still possibly different names (and slightly different boundaries) in
paraphyletic, assemblage. They certainly resemble the the literature. The classically known heterokonts (Pas^
old "phylum Protozoa" sufficiently enough to be recog­ cher 1937-1939) represent one such group, and this
nizable as the protozoa; in fact, many of the included name has been used by me for the major subkingdom
taxa of old remain intact in the new kingdom Protozoa, of the Chromista. The Chromophyta of Bourrelly
which is clearly separated from the other five eukaryotic (1957), perhaps the basis for the recently appearing
kingdoms. That the group is a large and genetically names Chromobionta and Chromobiota, also includes
diverse one is no reason in itself to require that it must many of the same groups and has been popular as a
be broken up. As the evolutionary proving ground from contrast to the Chlorobionta (see Christensen 1966), a
which emerged the other eukaryotic kingdoms (except name applied essentially to the green algae.
for the primitive Archezoa), the Protozoa might be ex­ Recently, the "stramenopiles" (more properly spelled
pected to show greater variety and even to be taxonomi­ "straminopiles"? See Vprs 1993) of Patterson (1989a)
cally unwieldy in some respects. Not surprisingly, new have become, in effect, a rival nomenclatural candidate
data may in time oblige us to make substantial revisions for the chromists of Cavalier-Smith (1986, 1989b). In
among a number of its numerous subtaxa, outstanding both m en’s cases, the same strong synapomorphic char­
examples - in my view - being several of the classes and acter has been used: the tripartite rigid tubular hairs or
orders of two of Cavaher-Smith’s (1993a-c) newest mastigonemes found on (or postulated to have been lost
phyla, viz. Percolozoa and Opalozoa. from) the flagella of allegedly all species assignable to
A final argument in favor of (re)recognizing the the overall group. This phylogenetically important fea­
Protozoa as a major high-level taxonomic unit among ture, however, is missing from species comprising
the eukaryotes is the fact that the concept underlying it several (but different) taxa in both Cavalier-Smith’s and
continues to satisfy the needs of field and bench eco­ Patterson’s suggested classifications. I accept Cavalier-
logists, who have long defined the Protozoa as basically Smith’s arguments - explaining the absences from his
comprised of primarily heterotrophic and colorless (with several included taxa that are without them - as the more
a few exceptions), motile, unicellular, mostly free- cogent ones. Thus, in brief, I agree with the latter worker
living, microscopic protists widespread in a variety of in excluding Patterson’s proteromonads + opalinids (=
habitats. This is essentially the same general - and useful the "slopalinids") and the heliozoan actinophryids from
- definition found for the old phylum Protozoa in many the (mostly phototrophic, with plastids inside the rough
textbooks. And, as Cavalier-Smith (1993c) has pointed endoplasmic reticulum) chromistic assemblage, while
out, protozoologists need not restrict their studies, or including the haptophytes, cryptophytes, and several
even their textbooks, to members of this kingdom alone. other chromist taxa left out of the "stramenopiles" in
In fact, our knowledge concerning the archezoan groups, Patterson’s circumscription of his informal group.
and other more widely dispersed protists classically The CHROMISTA is an important non-protozoan,
thought of as "protozoa" and sometimes - at the same non-plant kingdom of algal protists. In contrast to the bulk
time - "algae", will benefit from attention by students of the PROTOZOA, its members are mostly autotrophic
and researchers working in any field of the biological (although many are capable of mixotrophy), unicellular
sciences. forms with unique mastigonemes and a unique placement
Unresolved taxonomic problems within the Protozoa of their chloroplasts. The chromists represent a major
exist mostly in areas involving groups of small-sized group of "the algae" of old; and their heterokontic moiety
free-living and symbiotic heterotrophic flagellates, looms large among the seven distinct phylogenetic algal
which abound in a great diversity of habitats (Patterson lineages described by Andersen (1992) as having arisen
and Larsen 1991), and various amoeboid and plasmodial independently during geological time.
protists, especially taxa of mycetozoa s.l. and
amoeboflagellates s.l. Names and Authorships of Higher Taxa

The Kingdom CHROMISTA In general, the various codes of biological nomencla­

ture do not have much control over the choice of names
In a number of important characteristics, the protists for suprafamilial taxa (Corliss 1984, 1993; Jeffrey 1990’
assignable to my kingdom Chromista are similar or even Ride and Younès 1986). This may be considered both
 "User-friendly" Classification of the Protists 29

livea "good" and "bad", but the unchecked promulgation of number of quite unusual organisms (e.g. a single family
■s) in new ranks and names in recent years has led us nearly or even a single genus or species), then I especially have
(Pas-. ¡ t0 the brink of chaos in protistology, nomenclaturally tended to use the original name and authority for the
this j peaking (see extended discussions in Corliss 1984, group even if the rank (usually upward) may have
idem ¡99O, 1991b, 1993; Patterson and Larsen 1992). Some changed drastically. The description, while refined in
'relly , ¿mo ago, I stressed the need for "common sense and light of new knowledge, is basically concerned with the
aring courtesy" in the area (Corliss 1972), and Silva (1980) same organism(s). However, there are a few defensible
ludes has urged that an overriding consideration should be exceptions to this.
as a "effectiveness of communication". A reasonable degree 3. In the preceding and still other situations, I have
’6), a of stability is another (overlapping) goal certainly wor­ taken the liberty of altering prefixes and/or suffixes
thy of achievement in these times of perhaps too much without necessarily changing the authorship/date infor­
Jelled emphasis on constant change. mation. In the case of a number of former botanical
389a) In the classification of the 34 phyla formally offered classes, I have - along with other workers - felt free to
lidate ' in this paper, I have been confronted by the same dilem­ elevate the group to phylum (= division) status, altering
b). In mas described in my earlier overview work (Corliss the suffix appropriately. Also, I have not been con­
char- 1984), and have thus been obliged to make a number of strained by the lack of a Latin diagnosis in the first (or
irs or auite subjective decisions in choice of both names and subsequent) description to withhold credit from the
n lost authorities for the higher taxa included. Space does not original creator/proposer of a group or of its name. This
ble to permit a detailed discussion here. Suffice it to say, in latter decision has affected the date of authorship and
t fea- keeping with my objectives to present a "user-friendly" occasionally the authorities themselves in the case of
rising scheme of classification including all groups of protists, some botanically derived names, as will be apparent in
’s and I have adopted the guidelines given below, keeping in the text and in Table 1.
'alier- mind the practical observation made by Raabe (1964a) 4. According to conventional practices, a number of
m his some 30 years ago: "I am not an adherer of introducing group-names might well have been marked "emend",
more new names for old taxa, although they might stress better "sensu", "ex", "nom. nov.", or "stat. nov." (with or
'orker i their properties. It introduces confusion..." And Silva without additional author/date data), but, for the sake of
ids (= (1980) has sensibly suggested that a classification consistency and simplicity, I have not done this. I offer
. from should be one that is "familiar and acceptable to the apologies to nomenclatural purists and any offended
rough largest number of users". taxonomic specialists. Today, it is generally assumed
while 1. When possible/defensible, I have employed the by taxonomists that for descriptions of organisms or
everal oldest and/or most familiar name for a more or less groups of organisms one must go to the more recent
es" in conventional group of protists. Although the concept, rather than the older literature; in some ways, however,
i boundaries, composition, and even the rank-level may their nomenclature may be considered to be a separate
rzoan, I have changed somewhat over time, the name used may matter - primarily one more of historical interest.
e bulk be credited to the original author, using date of his 5 . 1 am not automatically opposed to all "new" names!
rophic creation of the name. As might be expected, sometimes For instance, I have endorsed/accepted some 20 of
ellular exceptions to this principle are advisable. Incidentally, Cavalier - Smith’s numerous high-level taxonomic / no­
ement to save space, I have not regularly listed all synonyms menclatural creations of the past dozen years. For a few
major : °f the names selected for the various high-level taxa examples of these neologisms (some of them altered
noiety endorsed on preceding pages. In a number of cases, slightly in this paper in rank or in spelling of the name):
; algal however, I have included a few for the benefit of readers his kingdom Chromista and subkingdoms Viridiplantae
arisen
who may have become more familiar with a name and Biliphyta; his phyla Archamoebae, Dinozoa,
different from the one chosen here for a particular Euglenozoa, Opalozoa, Percolozoa, and Radiozoa; his
(generally well-known) group, keeping in mind that classes Diplonematea, Entamoebidea, Protalveolatea,
persons coming from a botanical background, for ex- and Proterozoea. I am not necessarily rejecting his large
aniple, will have had a nomenclatural exposure likely number of new names for intermediate ranks. As stated
encla- in my IN TR O D U CTIO N , most of these have been
differing from that of students trained in zoological
nairies
^xonomy. omitted primarily to reduce the size of my own clas-
1990;
2- When both an original group and its later recogni- sificatory framework, making it more easily usable for
d both
; tlQn as a unique higher-level taxon involve a very small the many readers who are not taxonomic specialists and
30 J. O. Corliss

neither want nor need such details. But I am also not proper taxonomic homes. With respect to high-level
always convinced that our evidence to date requires the lineages widely recognized as truly monophyletic, I haVe
separation of so many genera at levels so far above that tried (on preceding pages) to show possible interrelated­
of the family. ness at phyletic (and lower) ranks, even if this process
has made demands on insight and intuition and involved
some healthy speculation. On more than one occasion
The Category ’’Incertae Sedis” I have deliberately (though often in a tentative way)
united paraphyletic groups under a single higher-level
I have not placed any major (or minor) taxa in the rank (as generally explained in the text in place).
convenient category of "uncertain status" for several
reasons. It is obvious that, as our knowledge continues
to grow, various species and higher groups as well will
need to be shifted about taxonomically. Systematics is Listing of Multiple Genera within Classes and Phyla
not a static science. Furthermore, from a puristic point
of view, we are really uncertain about a great many of I have offered far more than the usual number one
our ranks and group-interrelationships, frustrating sees of "representative genera" for each of the high-level
though this may be. It seems to me superfluous to mark taxa named on preceding pages (notable also in the
nearly everything as "incertae sedis" when we anticipate INDEX) because I should like to enable the readers -
changes based on fresh data of high phylogenetic/evolu­ no matter what their field of specialty - to find their
tionary (and thus taxonomic) value every year or so. It "favorites" and thus be able to relate them to the ranks
is to be expected that, at any given time, some groups above and to neighboring groups. All too often, it seems
are better known than others; but all deserve some place to me, papers that are concerned with phyla and classes
- even if it must be tenuous or tentative - in an overall fail to supply the reader with any clues as to the location
hierarchical classification, as I see it. of familiar genera within a newly proposed or newly
Patterson and colleagues (e.g. Brugerolle and Patter­ rearranged protist macrosystem (or portion thereof).
son 1990; Larsen and Patterson 1990; Patterson 1986a, Naturally, space restrictions preclude mention of all
1990; Patterson and Brugerolle 1988; Patterson and genera, which number in the thousands. But, among the
Zölffel 1991; V 0 rs 1988, 1993) favor labeling many 1100 that are included in this paper, I hope that I have
unique species as "incert. sed. protists," seemingly managed to select many of the better-known (as well as
without much desire to give them a (or place them in "representative") names of protists from the modern as
any preexisting) taxonomic rank above genus or family well as the classical literature. Perceptive
(with rare exception). Scores of exciting new protists are phycologists/protozoologists will note that certain
thus being more or less assigned to a vague "Anhängen" genera are no longer where they used to be in older,
position. By the same token, the Patterson school (e.g. conventional classifications. Ultrastructural studies, per­
see Patterson and Sogin 1993, and references therein), haps even more so than molecular biological data, have
and some other laboratories as well, determine necessitated such reassignments of sometimes familiar
monophyletic lineages of protists, excellent research to taxa. Consider cases of formerly (thought to be) "closely
carry out, while/but making no overt effort to interrelate related" genera or groups the members of which are now
these lines in a manner involving ranking and production so widely separated taxonomically from each other, such
of some kind of hierarchical system useful to the many as the following: Amoeba, Dientamoeba, Entamoeba,
people wanting and needing the overall classification and Pelomyxa; Proteromonas and Trypanosoma; Giar­
that would result. dia and Trichomonas; Acrasis and Dictyosteliutn',
Cavalier-Smith (e.g. 1993c, and references therein), Ciliophrys and Dimorpha; Stephanopogon or Opalina
among others of us, has attempted to find at least tem­ and the ciliates; microsporidians and myxosporidians;
porary or tentative homes for many "uncertain" protist dinoflagellates, chrysophyceans, and volvocines (a trio
genera (e.g. a number of those listed by Patterson and of taxa rather close in older zoological classifications);
Zölffel 1991 ), thus stimulating future workers to confirm and oomycetes plus hyphochytriomycetes and the
or disprove such allocations. Nevertheless, it is true that chytridiomycetes. More examples could be cited.
often protists poorly described in the older literature It has not been appropriate, here, to become involved
require rediscovery and restudy before they can be given in discussion/treatment of generic synonym^
 "User-friendly" Classification of the Protists 31

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36 J. O. Corliss

INDEX of Taxonomic Names

Names of top-level taxa (empire, kingdoms, phyla) are printed in boldface CAPS; those of classes and subclasses
in boldface upper- and lowercase. Ordinal (always ending in "-ida") and informal or vernacular names and generally
unused or discardable names (often in quotation marks)' appear in roman type. Representative genera are shown in
italics. Taxa that appear in Table 1. are so indicated by reference to that table following page number citations.

A Ammodiscus......................... .........19
Ammonia.............................. .........19
Acantham oeba....................... ...... 18 A m oeba............................... .........18, 30
ACANTHAREA.................. ...... 19, Table 1 Amoeboflagellates............. .........10, 12, 13, 28
ACANTHARIA................... .......19, Table 1 Am oebophrya..................... .........14
Acanthochiasma..................... ...... 19 Amphiacantha..................... .........10
Acanthocystis......................... ...... 19 Am phidinium ...................... .........14
Acanthoeca............................. ...... 13 Amphileptus........................ .........17
Acanthoecopsis...................... ...... 13 Amphilonche....................... .........19
Acantholithium....................... ...... 19 Am phipleura....................... .......... 22
Acanthometra........................ ...... 19 Amphisiella......................... ..........15
"Acellular" slime m olds..... .......13 Am phitrem a........................ ..........19
Acetabularia.......................... ...... 25 Am phora............................. .......... 22
A c h ly a .................................... ...... 23 Amphorellopsis.................. ..........18
Achnanthes............................ ...... 22 Ancistrocoma...................... ..........15
A cin eta ................................... ...... 15 Ancistrum............................ ..........16
A conoidea............................. .......17 Ancistrum ina...................... ..........16
A crasida................................ ...... 10 A ncora ................................ ..........17
Acrasis.................................... ....... 10, 30 Ancylonem a........................ ..........25
Acrosiphonia......................... ....... 25 Ancyromonas...................... .......... 12
Actinobolina.......................... ....... 17 Anigsteinia.......................... ..........15
Actinocephalus..................... ........17 ANIM ALIA....................... .......... 4, 5, 8 ,2 0 ,2 6 ,
Actinocoryne......................... ....... 19 Table 1
A ctinom onads....................... ....... 21 Anisolpidium....................... ..........23
Actinomonas.......................... ........21 Anisom onas........................ .......... 12
Actinom yxidia.................... ....... 20 Anoplophrya....................... ..........16
Actinophryidea................... ....... 19, Table 1 Anthem osom a..................... ..........18
Actinophryids........................ ........28 Anthophysa......................... .......... 22
Actinophrys........................... ....... 19 Aphanochaete................... ..........25
"Actinopoda"......................... ....... 19 A PIC O M PLEX A........... ..........17, 26, 31, Table 1
Actinosphaerium.................. ........19 Apiosom a........................... ..........16
Acytostelium .......................... ....... 13 Aplanochytrium................ .......... 21
A d e le a .................................... ....... 17 Apostom atia..................... ..........16
Adelpham oeba..................... ........10 Apusom onas...................... .......... 12
Aggregata.............................. ....... 17 Arachnodinium................. ..........17
A la ria ..................................... ....... 22 Arachnula.......................... ..........19
A lbugo.................................... ........23 A rcella ............................... ..........18
Alexandrium .......................... ....... 14 A R CH AM O EBAE......... .......... 9, 29, Table 1
A llogrom ia............................ ....... 19 ARCHEZOA.................... .......... 5, 7, 9, 11, 18,20,
Alloiozona............................. ....... 17 27, 28, Table 1
Allom yces.............................. ....... 26 Aristerostom a................... ...........15
"Alveolata"........................... ....... 13, 14, 17 Arthrocladia...................... ........... 22
Amastigomonas................... ........12 Arthrodesmus.................... ...........25
 "User-friendly" Classification of the Protists 37

as c e t o s p o r a ..... 20, Table 1 ............................................ Blastocrithidia................ ............ 12

Ascobius ................................. ..... 15 B lastocysta...................... ............ 26
aSc o m y c o t a .................. ..... 26 - Blastocystis...................... ............ 18, :
Ascophrys................................ 16 .............. Blastodinium ............ 14
..................................................

Askenasia 17
............................................................................... .............. Blepharism a 15
................................................. ...............................

Askoella 16
....................................................................................... .............. Blepharocorys ............ 17
.........................................

Aspidisca 15
.................................................................................. .............. Blidingia 25
.................................................................. .................................

Astasia 11
............................................................................................ .............. Blue-green alg ae............ ............. 26
Astomatia.............................. ..... 16 Boderia............................ ............. 19
Astrolonche............................ .......19 Bodo................................. ............. 12
Astrolophus............................ .......19 B odonida......................... 12 .................................

Astylozoon 16
............................................................................ ................. Bodonids 11
.................................................................. .................................

Athalamida 19
.......................................................................... ................. B oldia 26
............................................................................ .................................

Atlanticella 20
.......................................................................... ................. B olivina 19
.................................................................... .................................

Audouinella 26
....................................................................... ................. Botrydiopsis ............. 22
....................................................

Aulacantha............................. .......2 0 Botryococcus.................. ............. 25

Aulosphaera........................... .......20 B overia............................ ............. 16
Aulotractus............................. .......20 Brachonella.................... ............. 15
Auricula 22
....................................................................................... ................. Brachynema.................... 22 .................................

Australothrix 15
.................................................................. ................. Bresslaua 15
............................................................ .................................

Avelia ...... 15
.................................................................................................. Brevilegnia 23
.................................................... .................................

Brooklynella 15
............................................... ......................... ..

Brown algae ............. 8 ,

...............................................

B Brugerolleia................... 10 .................................

Bryometopus.................. ............. 15
Babesia................................... .......18 Bryophrya...................... ............. 15
Bacillaria.............................. ...... 22 Bryophyllum.................. 17 ....................................

Bacillariophyceae 22, Table 1 ....................................... ................. BRYOPHYTA 24

.................................... ....................................

BACILLARIOPHYTA 22 ............... .................... B ryophytes 24

.................................................... ....................................

Bacteriastrum 22
............................................................ ................. Bryopsis 25
................................................................. ....................................

Badhamia 13
............................................................................. .................... Buetschliella .............. 16
...............................................

Bakuella................................ ........15 Bullanympha.................. .............. 11

Balamuthia............................ ........18 Bumilleriopsis................ .............. 22
Balantidium........................... ........17 B urkea............................ 10 ....................................

Bangia 26
.......................................................................................... .................... Bursaria 15
.................................................................. ....................................

Bangiophyceae 26, Table 1

.................................................... .................... Bursaridium 15
................................................. ....................................

Bangiopsis 26
.......................................................................... .................... Bursellopsis 16
................................................. ....................................

Barbulanympha 11
.................................................... .................... Bursostoma .............. 16
....................................................

BASIDIOMYCOTA.......... ........26 Buxtehudea.................... 10 ....................................

Bathycoccus.......................... ....... 24
Batrachospermum................ ........26
Besnoitia............................... ........17 C
, Bicosoeca 21
............................................................................. ....................

BlCOSOECAE 21, Table 1

............................................... .................... Caenomorpha 15
....................................... ....................................

Bicosoecidea 21, Table 1

............................................................ .................... Cafeteria 21
............................................................ ....................................

Bicosta... 13 ................... Calciosolenia 23

......................................... ....................................

Biddulphia ........22
....................................................................... Calliacantha................. .............. 13
b i l i p h y t a ....................... ........25, 26, 29, Table 1 Callimastix.................... 26 .......................................

B iom yxa 19
............................................................................... ................... Callocolax 26
.................................................... .......................................

Biastocladiella.................... 26 .................... Calonympha 11

............................................ .......................................
38 J. O. Corliss

Camptonema...................................19 Chlorobotrys............................ 22
Canistrolithus................................. 23 Chlorococcum......................... 25
C apreolia........................................26 Chlorogonium.......................... 25
Carchesium.................................... 16 Chlorokybus............................. 25
Carteria........................................... 25 "Chloromonads"...................... 21
Carterina........................................ 19 Chloromonas........................... 21, 25
Caryospora....................................17 Chloromyxum.......................... 20
C ashia.............................................18 Chlorophyceae....................... 25, Table 1
Castanella.............................. 20 CHLOROPHYTA................ 24, Table 1
Catenaria........................................26 "Choanociliata"....................... 13
Caulleryella....................................17 Choanoflagellatea................. 13, Table 1
Caviom onas....................................10 Choanoflagellates................... 13
C avostelium ...................................13 Choanoflagellida.................... 13
Cellular slime molds.....................13 Choanomonads........................ 13
Cenosphaera.................................. 20 CHO ANO ZO A..................... 13, Table 1
Centrohelidea.............................. 19, Table 1 Choanozoa............................... 26
Centropyxiella.................................18 Chondrus................................. 26
Centropyxis.....................................18 Chonotrichs............................. 15
Cepedea........................................... 12 Chordaria................................ 22
Cepedietta....................................... 16 Choreotrichs............................ 15
Cephalothamnium.......................... 12 Chromidina............................. 16
Ceratiomyxa....................................13 CHROM ISTA....................... 5, 8,13,19,20,23,
C eratium ......................................... 14 26 - 29, Table 1
Ceratomyxa.................................... 20 Chromobionta......................... 28
Cercomonas....................................12 Chromobiota........................... 28
Chaenea.......................................... 17 Chromophy ta .......................... 28
Chaetochloris................................. 25 Chromulina............................. 22
Chaetosiphon................................. 25 Chroomonas........................... 23
Chaetosphaeridium....................... 25 Chrysamoeba.......................... 22
Challengeria.................................. 20 Chrysidalis.............................. 23
Challengeron.................................. 20 Chrysocapsa........................... 22
C haos...............................................18 Chrysochromulina................. 23
C hara.............................................. 25 Chrysococcus ......................... 22
Chardezia........................................ 18 Chrysodendron....................... 22
Charophyceae............................... 25, Table 1 Chrysomonadida..................... 22
CHAROPHYTA .............25, Table 1 C hrysom onads....................... 22
Charophytes................................... 25 Chrysophyceae...................... 22, Table 1
Chattonella......................................21 Chrysophyceans..................... 27, 30
Chilodochona..................................15 Chytridiomycetes................. 26, Table 1
Chilodonella....................................15 Chytridiomycetes.................. 31
Chilomastix.....................................10 CHYTRIDIOMYCOTA 26, Table 1
Chilomonas.................................... 23 Chytridiopsis.......................... 10
Chlamydodon..................................15 Chytridium.............................. 26
Chlamydom onas............................25, 27 C hytrids................................... 23, 26
Chlamydophrys.............................. 18 Chytriodinium......................... 14
Chlorarachnion..............................24 Chytriomyces.......................... 26
Chlorarachniophyceae............. 24, Table 1 Cienkowskya........................... 19
CHLORARACHNIOPHYTA . 24, Table 1 Ciliates..................................... 17, 30
Chlorella.........................................25 CILIOPHORA..................... 14, Table 1
C hlorobionta..................................28 Ciliophryids............................ 19
 "User-friendly" Classification of the Protists 39

Ciliophrys........................................21, 30 Contophrya..................................... 16
Cilioprotists.....................................14 Coprom yxa..................................... 18
Cinetochilum...................................16 Corallomyxa................................... 13
Cirrogaster....................................15 Coscinodiscophyceae...................22, Table 1
Cladophora.....................................25 Coscinodiscus.................................22
Clathrostoma...................................16 Cosm arium.....................................25
Clathrulina...................................... 19 Cosmetophilus................................17
Clausilocola....................................16 Cosmocolpoda................................15
Clevelandella........................,........ 15 Costaria...........................................22
Climacostomum............................. 15 Cothurnia.........................................16
Closterium.......................................25 Craspedomonadophyceans 36
"Cnidosporidia".............................20 Craspedom onads............................13
Coccidea......................................... 17, Table 1 Craspedophyceae..........................13
Coelodiscus................................... 20 Cribraria.........................................13
COCCOLITHOPHORA............23 C rithidia..........................................12
Coccolithophorida..........................23 Crypthecodinium............................14
Coccolithus.................................... 23 Cryptobia.........................................12
Coccomyxa.................................... .25 CRYPTOM ONADA...................23, Table 1
Cochliopodium............................... 18 Cryptom onadea........................... 23, Table 1
Codium.................................. 25 Cryptomonadida............................ 23
Codonella ..........................15 Cryptomonas..................................23
Codosiga........................................ 13 CRYPTOPHYTA.................... 23, Table 1
Coelastrum .......................... .25 Cryptophytes..................................28
Coelodendrum................................ 20 Cryptosporidium............................17
Coelomomyces............................... 26 Cucurbitella.................................... 18
Coelotropha..................... ..............17 Culicospora.................................... 10
Cohnilembus .................... .16 Curimostoma.................................. 16
Colacium........................................ 11 Cyanidium ...................................... 26
Coleochaete.................................. 25 Cyanophora....................................26
Coleps..............................................16 Cyathobodo................... .............. 12
Collar flagellates............................ 13 Cyathobodonida ......................... 12
Collinia........................................... 16 Cyathodinium................................. 15
Collosphaera.................................. 20 Cyclidium.........................................16
Collozoum ........................... 20 Cyclophora.............................. 22
Colpidium........................................ 16 Cycloposthium................................17
Colpoda........................................... 15 Cyclospora..................................... 17
Colpodea...................................... 15, Table 1 Cyclotella........................................ 22
Colpodella....................................... 17 Cyclotrichium................................. 17
Colponema......................................14 Cylindrocystis.................................25
Comatricha.....................................13 Cylindrotheca ......................22
Compsopogon ...........................26 Cymatopleura ....................22
Conchophthirus.............................. 16 Cymbella......................................... 22
Conchopsis......................................20 Cym opohlia....................................25
Condylostoma................................ 15 Cyphoderia..................................... 18
Conidiophrys...................................16 Cyrtocaryum................................... 16
Conion..............................................13 Cyrtolophosis................................. 15
Conjugaphyceae.............................25 Cyrtophorids .................... 15
Conjugatophyceae....................... 25, Table 1 Cyrtostrombidium......................... 15
A noidea......................................... 17 Cystodinium.................................... 14
Conradinema..................................11 Cystoseira....................................... 22
40 J. O. Corliss

D Disematostoma.................. 16
D istigm a............................. .......... 11
D actylosom a...................... ...........18 Ditrichomonas.................. .......... 11
Dasycladus......................... ...........25 Dobellia............................. ..........17
Deltotrichonympha........... ........... 11 Dolichomastix................... ..........24
D endrocometes................. ...........15 Doliospora......................... ..........17
Dendrosoma....................... ...........15 Dorisiella........................... ..........17
Dermatochrysis................ ........... 22 D ragescoa......................... ..........16
D esm idium ........................ ...........25 Duboscquella..................... ..........14
Desmids............................. ...........25 Dunaliella.......................... ..........25
D esm othoracidea............ ...........19, Table 1 Durchoniella...................... ..........16
D esportesia....................... ........... 10 D ysteria............................. ..........15
Devescovina..................... ........... 11
D exiotricha....................... ...........16
Diadesmis.......................... ........... 22 E
Diaphanoeca.................... ...........13
Diatoma............................. ........... 22 E bria................................... ........... 12
D IA T O M A E ................... ............22, Table 1 Echinostelium................... ...........13
D iatom s............................. ............ 22 Echinozoon........................ ...........18
D ictyocha.......................... ............ 21 Ectocarpus......................... ........... 22
D IC T Y O C H A E .............. ............19, 21, Table 1 Eim eria.............................. ...........17
D ictyostelea..................... ............13, Table 1 Eim eriida........................... ...........17
D ictyostelium ................... ............13, 30 Ellobiophrya...................... ...........16
Dictyota............................. ............ 22 Ellobiopsis......................... ...........14
Didesmis............................ ............17 Elphidium .......................... ...........19
D idinium ........................... ............17 Em iliania........................... ...........23
Didymium .......................... ............13 Encephalitozoon............... ........... 10
Dientamoeba.................... ............11, 18, 30 Enchelys............................. ...........17
D ifflugia............................ ............18 Endamoeba........................ ...........18
D ileptus............................. ............17 Endocladia......................... ...........26
D ilsea................................ ............26 Endo limax.......................... ...........18
D im orpha.......................... ............12, 30 Endoreticulatus................ ........... 10
D im orphids....................... ............19 Endosphaera...................... ...........15
D inobryon......................... ............ 22 Endotrypanum.................. ........... 12
Dinoflagellatea................ ............14, Table 1 Entamoeba......................... ...........18, 30
Dinoflagellates................. ............ 5, 17, 19, 27, 30 Entamoebidae.................. ........... 9, 18
D inophyceae................... ............14 Entam oebidea................ ...........18, 29, Table
Dinophysis......................... ............14 Enterocytozoon................ ........... 10
D IN O Z O A ....................... ............ 5, 13, 29, Table 1 Enterom onadida.............. ........... 10
Diophryopsis.................... ............15 Enteromonas..................... ........... 10
Diophrys............................ ............15 Entodiniomorphids.......... ...........17
Diphyllea.......................... ............ 12 Entodinium....................... ...........17
Diplocystis........................ ............17 Entodiscus........................ ...........16
Diplomonadida................ ............ 10 Entosiphon........................ ........... 11
Diplonema........................ ............ 11 Epalxella........................... ...........15
Diplonematea................. ............11, 29, Table 1 Ephelota............................ ...........15
D iplospora....................... ............17 E pipyxis............................ ........... 22
Discocelis......................... ............ 12 Epistylis............................ ...........16
Discocephalus................. ............15 Erythropsidinium............. ...........14
D iscorbis.......................... ............19 Erythrotrichia.................. ...........26
▼
"User-friendly" Classification of the Protists 41

16 G
25
E uglena. 11 -Galatheammina...................... ..... 19
11 Gamophyceae........................ ..... 25
11 Gastrostyla............................. ..... 15
11 G eleia...................................... ..... 15
11, Table 1 G iardia.................................... ..... 9, 10, 30
Euglenoids .. 5, 11, 24, 27, 30 Giffordia.................................. ..... 22
eugleno; 5, 11, 12, 29, Table 1 G igartina................................ ..... 26
Euglypha..... .18 G labratella............................. ..... 19
Eugomontia. .25 Glaucocystis........................... ..... 26
5, 7, Table 1 Glaucom a.............................. ..... 16
.1 2 Glaucophyceae..................... ..... 26, Table 1
"Eumycota".......... .23 GLAUCOPHYTA.............. ..... 17, 26, Table 1
Euplotes................ .15 Glaucophytes......................... ..... 24, 26
Eustigmatophyce: .22, Table 1 Glaucosphaera...................... ..... 26
Eustigmatophytes. .2 2 Glenodinium.......................... ..... 14
.2 2 Gleodinium............................ ...... 14
.11 G lobigerinella .................................... .......19
Globospora............................ ...... 20
Gloeobotrys........................... ...... 22
Gloeochaete........................... ....... 26
Gloeocystis............................ .......... 25
.2 0 Gloeopodium......................... .......22
Fabrea..... ..15 G lugea.................................... .......... 10
Favella.... ..15 Golden-brown algae............. .......22
Fenchelia , ..16 Goniomonadea..................... .......23, Table 1
.. 18, Table 1 G oniom onads....................... .......23
..18 G oniom onas......................... .......23
..26, Table 1 Goniotrichum....................... .......26
Foettingeria ..16 G onium ................................. .......25
Folliculina... ..15 Gonospora............................ .......17
..10 Gonyaulax............................ .......14
..19, Table 1 Gonyostomum .................................... ............ 21
F o ra m in ife re a n s. ..2 0 Gorillophilus........................ ........17
..19 Goussia................................. ........17
Foram s.... ..19 Gracilaria............................. ........26
F ragilaria. ..2 2 Grammatophora.................. ........ 22
..22, Table 1 Grandoria............................. ........15
..17 Granuloreticulosea............ ........13, 18, Table 1
Fritschiella. ...25 Grass-green scaly alg ae ......... ............ 24
Frontonia ... ..16 Green a lg a e .......................... ........24 - 28
Frustulia .... ...2 2 Gregarina............................. ........17
...2 2 Gregarinidea....................... ........17, 31, Table 1
Fucus.... 22 Grellia................................... ............ 17
Fuligo... ...13 Gromia ...................................................... ........ 18
f u n g i. ... 5. 8 , 18, ?0, 23, 26 Gross glockneria ........................... ............ 15
Table 1 Gruberella .......................................... ............ 10
...26 Gurleya................................ ............ 10
p
Urgasonia. ...15 Guttulina.............................. ........19
42 J.O . Corliss

Guttulinopsis........................ .........18 H em iselm is............................ ...... 23

Gymnodinioides.................. .........16 H em ispeira............................ .......16
Gymnodinium....................... .........14 Henneguya............................. ...... 20
Gymnosphaera.................... .........19 H epatocystis.......................... .......18
Gyrodinium.......................... .........14 Hepatozoon............................ ...... 17
Gyrosigma............................ .........22 H erpetom onas....................... ...... 12
H essea.................................... .......10
H eteram oeba......................... ...... 10
H Heterochloridea.................. ...... 22
Heteroderma.......................... .......26
Haematococcus................... .........25 Heterogloea........................... ...... 22
H aem atozoea...................... .........18, Table 1 HETEROKARYOTA...... .......14
Haemogregarina................. .........17 Heterokont............................ ...... 22
Haemoproteus...................... .........18 HETEROKONTA............. .......21, Table 1
Haem osporida...................... .........18 Heterokontophyta.............. .......21
Halimeda.............................. .........25 H eterokonts.......................... ...... 13, 21 - 23, 26, 28
Haliommatidium................. .........19 Heterolobosea...................... ...... 10, Table 1
Halocella.............................. .........20 Heteromita............................ .......12
Halosphaera......................... .........20 Heteromitida......................... ...... 12
Halteria................................ .........15 Heteronem a.......................... .......11
Halymenia............................ .........26 Heteronem atida................... ...... 11
Hanzschia............................ .........22 Heterophrys.......................... ....... 19
Haplocaulus........................ .........16 Heterosigma......................... ........21
H aplosporidea.................. .........20, Table 1 H eterotheca.......................... ........19
Haplosporidians................. .........20 H eterotrichea...................... ........14, Table 1
Haplosporidium................. .........20 Heterotrichia....................... ....... 15
H aplozoon........................... .........14 Heterotrichians.................... ....... 14
HAPTOM ONADA........... .........23, Table 1 H eterotrichs.......................... ....... 14, 15
H aptophrya................. ....... .........16 H examastix........................... ....... 11
H APTOPH YTA............... .........23, Table 1 H exam ita.............................. ........10
Haptophytes........................ ......... 8 , 22, 27, 28 Hibberdia.............................. ........22
Haptorids............................. .........17 H ildenbrandia...................... ........26
Hartmannella...................... .........18 Histiobalantium................... ....... 16
Hartmannula....................... .........15 H istiona................................ ........12
H astigerina......................... ..........19 Histomonas........................... ........11
Hausmanniella................... ..........15 Hochbergia........................... ....... 13
H edraiophrys..................... ..........19 Hoferellus............................. ........20
H edriocystis........................ ..........19 Holomastigotoides............... ........11
H egneria............................. ..........11 Holophrya............................. ........16
Helicoprorodon.................. ..........16 Hom osira.............................. ........22
Helicosporidium................. ..........2 0 , 26 H oplitophrya........................ ........16
Heliochona.......................... ..........15 Hoplonympha....................... ........11
"Helioflagellates"............... ..........12, 19, 21 Hyalophysa........................... ....... 16
H eliom onas......................... ..........12 Hydramoeba......................... ........18
H eliophrya.......................... ..........15 Hydrodictyon........................ ....... 25
HELIOZOA....................... ..........19, Table 1 Hydrosilicon......................... ........22
Hemidiscus.......................... ..........22 Hymenostomatia................ ........15
Hem im astigida................... ..........12 H yperm astigotea............... ........11, Table 1
Hemimastigophorea......... ..........11, 12, Table 1 Hyphochytriom ycetes...... ........23, Table 1
H em im astix......................... .......... 12 Hyphochytriom ycetes......... ........30
 "User-friendly" Classification of the Protists 43

¡{yphochytrium .................. 23 Kreyella...........................................15

Uypochona.......................................... 15 K udoa..............................................20
jfypotrichs.......................................15
H ysterocineta..................................... 16

I
Laboea........................................ 15
Ichthyobodo....... 12 LA BY R IN TH O M O R PH A 21, Table 1
Ichthyophthirius. 16 Labyrinthula.............................. 21
i n f u s o r ia ..... 14 L a b y rin th u le a .......................... 21, Table 1
Intoshellina....... 16 Labyrinthuloides........................ 21
lodamoeba......... 18 Lacrymaria................................ 17
Iridaea................ 26 Lagenidium................................ 23
Iridia.................. 19 Lagenophrys.............................. 16
Isochona............. 15 Lagotia....................................... . 15
Isochrysis........... 23 Lagynophrya............................. 17
Isonema.............. 11 Lambornella............................. 16
Isospora............. 17 Lam inaria................................. 22
17 Lam tostyla................................ 15
Lankesterella............................ 18
Lankesteria............................... 17
Lateromyxa.............................. . 18
Jakoba......... 12 Lecudina.................................... 17
Jaocorlissia. 16 Leegaardiella............................ 15
11 Legere Ila.................................... 18
Leishm ania............................... 12
K Lembadion................................ 16
Lennoxia.................................... 22
Kahliella.......... 15 Lepidotrachelophyllum........... 17
Karlingia......... 26 Leptomitus................................ 23
Karotomorpha. 12 Leptom onas.............................. 12
12 Leptom yxa................................ 18
9 Leptothorax.............................. 15
Karyolysus 17 Lesquereusia............................. 18
14, Table 1 Leucocryptos........................... . 12
14 Leucocytozoon......................... 18
15 Leucodictyon........................... 12
Kathablepharis. 12 Licea......................................... 13
Kentrophoros ... 14 Licm ophora............................. 22
Kerona . 15 Licnophora .............................. 15
K ha w kin ea ...... 11 Lieberkuehnia.......................... 19
K iitricha ........... 15 Lithodesmium.......................... 22
12, Table 1 Lithophyllum............................ 26
11, 25, Table 1 Lithoptera................................ 19
Klebsorm idium . 25 Litonotus................................... 17
Klossia . 17 Litosiphon................................ 22
17 L ito sto m a tea .......................... 16, Table 1
11 Lobochona............................... 15
14 Lobosea.................................... 13, 18, 26, Table 1
Komokia 19 Lohmanniella........................... 15
 1

44 J. O. Corliss

L o m a ................................................10 M etacystis......................................16
Lom iella.......................................... 16 METAMONADA 9, Table 1
Lomosporus.................................... 20 Metarotaliella............................... 19
Lophom onadida............................. 11 M etchnikovella............................. 10
Lophomonas....................................11 Metchnikovellida..........................10
Loricophrya....................................15 M etopus......................................... 15
Loxocephalus..................................16 Miamiensis..................................... 16
Loxodes........................................... 14 Micrasterias.................................... 25
Loxophyllum....................................17 M icrofilum..................................... 10
Luffisphaera....................................18 M icroglabratella.......................... 19
L w offia................... 15 M icroglena....................................22
Ly cogala.......................................... 13 Micro gromia................................. 19
Lynchella......................................... 15 M icrojoenia.................................. 11
Lyrella............................................. 22 M icromonadophyceae............... 24
Lyromonadea................................ 10, Table 1 Micromonas...................................24
Lyromonas....................................... 10 M ICRO SPO RA..........................10, Table 1
M icrospora....................................25
M icrosporea.................................. 10, Table 1
M M icrosporida................................ 10
Microsporidians............................. 20, 27, 30
M acrocystis.................................... 22 Microthorax.................................. 15
Macrospironympha........................11 Minchinia.......................................20
Mallodendron................................. 22 M inisporida .................... 10
M allom onas................................... 22 M inium........................................... 26
Mallomonopsis............................... 22 M inutocellus................................. 22
M am iella.........................................24 M onoblepharella..........................26
M antoniella.................................... 24 M onocercomonas.........................11
M arituja.......................................... 16 M onocercomonoides................... 10
M arteilia.........................................20 Monochilum.................................. 16
M arteiliidea.................................. 20 M onochiysis..................................22
M aryna............................................ 15 Monocystis..................................... 17
M assisteria......................................12 M onodopsis...................................22
M astigamoeba............................... 9 M onosiga....................................... 13
M astigamoebida............................ 9 Monothal amida............................. 19
M astigella...................................... 9 M razekia........................................ 10
M astigina....................................... 9 M YCETO ZO A........................... 12, Table 1
Maupasella......................................16 M ycetozoa..................................... 18, 28
M ayorella........................................ 18 M ycterothrix.............. ...................15
M edusetta........................................20 M ylestom a.....................................15
M egamoebomyxa........................... 13 Myrionema.....................................22
M elanophyceae.............................22 M yxidium....................................... 20
M elosira..........................................22 M yxobolus.....................................20
M enoidium ................................... ..11 M yxogastrea.................................. 13, Table 1
M erotricha......................................21 M YXOM YCETES.................... 12
M esodinium.....................................17 MYXOMYCOTA 12
M esojoenia......................................11 Myxophthirus................................ 16
"M esokaryota"............................... 13 M yxoproteus................................. 20
M esophyllum.................................. 26 M yxosporea................................. 20, Table 1
M esostigm a.................................... 24 M yxosporidia.............................. 20
M ESO ZO A................................... 27 M yxosporidians............................20, 30
T

"User-friendly" Classification of the Protists 45

Myxotheca......................................... 19 Onychodromus................................ 15
MYXOZOA..................................... 20,Table 1 Oocardium...................................... 25
¡yfyxozoa...........................................27 Oodinium................. 14
yiyzocytium.......................................23 O om ycetes...................................... 23, Table 1
Oomycetes...................................... 30
N OOMYCOTA................................23
Opalina.............................................12, 30
Naccaria.......................................... 26 O palinatea...................................... 12, Table 1
Naegleria........................................ 10 O palinida.........................................12
Nannochloropsis............................ 22 O palinids........................................ 28
Nanochlorum................................... 25 Opalinopsis............................. 16
Nassophorea..................................15, 16, Table 1 O PALOZOA................................ 11, 12,19, 28, 29,
Nassula................................... 15 Table 1
Nassulopsis..................................... 15 Opercularia ........................16
Nautococcus....................................25 Ophiaster........................................ 23
Navicula........................ 22 Ophiocytium................................... 22
Nebela...........................................,.18 Ophiuraespira.............................. 16
Neobursaridium............................. 16 Ophrydium.....................................16
Neocallimastix............................... 26 Ophryocystis................................. 17
Nephridiophaga............................. 20 Ophryodendron .........................15
Nephromyces............................... 12 Ophryoglena................................... 16
Nephroselmis...................................24 Ophryoglenids.......................... 15
Netzelia........................................... 18 Ophryoscolex.................................. 17
Nitella.............................................. 25 Opisthonecta................................... 16
Nitellopsis.......................................25 Orbopercularia............................... 16
Nitzschia............................ 22 Orbulinella..................................... 19
Noctiluca........................... ............ 14 O rtholinea...................................... 20
Nolaclusilis............................... .....15 Ovam m ina.......................................19
Nolandia..........................................16 Oxym onadea................................. 10, Table 1
Nonion.............................................19 Oxym onadida................................. 10
Nosema............................................ 10 Oxymonas ...........................10
Notila............................................... 10 Oxyrrhis........................................... 14
Nuclearia....................................... 18 Oxytoxum.........................................14
Nyctotheroides...............................15 Oxytricha........................ 15
Nyctotherus....................................15
P
O
Pallitrichodina............................... 16
Ochromonas................................... 22 Palmaria......................................... 26
Octodendron ......................19 Palmodictyon..................................25
Octomitus....................................... 10 PARABASALA............................ 11, 18, Table 1
Octomyxa................ ...................... 12 Parabasalans................................... 7
Odontella........................................ 22 Parabundleia.................................. 17
Ogdeniella......................................18 Paracichlidotherus........................ 15
Oikomonas.................................... 21 Paracineta...................................... 15
Oligohymenophorea.................. 15, Table 1 Paradistigma..................................11
Oligotrichs.................................... 15 Paraflagellata............................... 12
Olisthodiscus................................. 21 Paraíso tri cha..................................17
Olpidiopsis.................................... 23 Paramarteilia ....................20
Olpidium........................................26 Param ecium .................................. 15, 16
46 J. O. Corliss

Paramoeba......................... ...........18 Phacotus................................ ..... 25

Paramyxa........................... ........... 20 P hacus.................................... ..... 11
P aram y x id ea..................... ........... 20 Phaeocystis............................ ..... 23
Parapedinella..................... ........... 21 P h a e o d a re a........................... ..... 20
Parastasia.......................... ........... 11 Phaeodina.............................. ..... 20
Paratetramitus.................. ........... 10 Phaeophila............................. ..... 25
Parauronema..................... ...........16 Phaeophyceae....................... ...... 22, Table 1
Parvicapsula...................... ........... 20 PH A E O PH Y T A .................. ...... 21, Table 1
Parvicorbicula.................. ...........13 Phagodinium ......................... .......12
Pascherina......................... ...........25 Phagomyxa............................ ...... 12
P atelliferea........................ ...........23, Table 1 Phalacrocleptes..................... .......15
Patellinella......................... ...........19 Phalansterium....................... .......12
Paulinella........................... ...........18 Phascolodon.......................... ...... 15
Paurotricha........................ ...........16 Phialinides............................. ...... 17
P avlova.............................. ...........23 P hilaster................................ .......16
P av lo v ea............................ ...........23, Table 1 Philasterids............................ ...... 16
Pediastrum......................... ...........25 Phragm onem a....................... .......26
Pedinella........................... ........... 21 Phreatam oeba....................... ...... 9
P edinellea.......................... ...........21, Table 1 Phreatamoebida..................... ...... 9
Pedinelleans..................... ........... 21 Phtorophrya........................... .......16
Pedinellids......................... ........... 21 Phyllopharyngea................. ...... 15, Table 1
Pedinellophyceans........... ........... 21 Phyllophora........................... ...... 26
Pedinomonas.................... ...........24 Physarum............................... ...... 13
Pedinophyceae................ ...........24, Table 1 Phvsoderma........................... ...... 26
Pelagococcus................... ........... 22 "Phytoflagellata".................. ...... 5
Pelagohalteria................. ...........15 Phytomonadida...................... .......25
Pelagomonas.................... ........... 22 "Phytomonads"...................... .......27
P elagophyceae................ ...........22, Table 1 Phytomonas........................... ...... 12, 25
Pelagostrombidium.......... ...........15 Phytophthora......................... ...... 23
P e lo b io n tea ...................... ........... 9, Table 1 Piroplasmida......................... ...... 18
Pelomyxa........................... ............ 9 ,3 0 P lacus.................................... ...... 16
Penardia............................ ...........18 Plagiacantha........................ .......20
Peniculines........................ ...........15 Plagiocampa......................... .......16
P e n icu lin ia....................... ............16 Plagiopyla............................ .......17
Peniculistoma................... ............16 Plagiotom a........................... .......15
Peranem a.......................... ............ 11 Planicoleps........................... .......16
P ercolom onadea............. ............10, Table 1 Planorbulina......................... .......19
Percolom onas.................. ............ 10 P L A N T A E ........................... ....... 5, 8 , 10, 21,
P E R C O L O Z O A ............. ............10, 11,28, 29, Table 1 27, 31, Table 1
P erezia.............................. ............ 10 Plasmodial slime m olds..... .......13
Peridinium......................... ............14 Plasmodiophora.................. ...... 12
P e ritric h ia ........................ ............16 Plasmodiophorida................ .......12
Peritromus......................... ............15 Plasmodium.......................... .......18
P e rk in sid e a ..................... ............17, Table 1 Platyamoeba......................... .......18
Perkinsus........................... ............17 Platychilomonas.................. .......12
Pernina.............................. ............ 10 Platycola............................... ...... 16
Peronosclerospora........... ............23 Platyophrya.......................... ........15
Peronospora..................... ............23 Pleistophora......................... ........10
Petalomonas..................... ............ 11 Pleodorina............................ ........25
Phacodinium..................... ............15 Pleurasiga............................ ........13
 "User-friendly" Classification of the Protists 47

¡¡¡¿uraspis............................... ...... 19 Protoheterotrichs.................. ...... 15

pleurastrum.................................. 25 Protoopalina.......................... ...... 12
porochloris.......................... ...... 22 Protoperidinium................... ...... 14
pleurochrysis......................... .......23 Protosiphon........................... ...... 25
pleurocoptes.......................... ...... 16 Protostelea ............................ ...... 13, Table 1
pleuronema............................ .......16 Protostelium .......................... ...... 13
pleuronematids....................... ...... 16 Protozelleriella...................... ...... 12
• pleurosigma........................... ...... 22 PROTOZOA............................... 5 - 7, 9, 10, 22 - 28,
; pieurostomes.......................... .......17 Table 1
ploeotia.................................. .......11 PRYMNESIOPHYTA....... ........23
Pneumocystis......................... .......26 Prymnesium.................................. 23
Pocheina................................ ...... 10 Psalteriomonas..................... ....... 10
j Podophrya.............................. .......15 Psam m etta.................................... 19
Podosira.................................. .......22 Pseudobalanion.................... ....... 16
Polycycla................................ ....... 16 Pseudobodo.......................... ....... 21
Polycystinea......................... .......20, Table 1 Pseudocharaciopsis............. ....... 22
] Polyhymenophora.................... 14, Table 1 Pseudociliatea ..................... ....... 11, Table 1
i Polyhymenophoreans................... 15 Pseudodendromonas........... ....... 12
Polykrikos.............................. ....... 15 Pseudodifflugia..................... ....... 18
Polvmyxa............................... ........ 12 PSEUDOFUNGI................ ....... 23, 26, Table 1
Polysphondylium.................. ....... 13 Pseudo glaucom a................. ....... 15
Polystomella.......................... ....... 19 Pseudolithium....................... .... . 19
Polytomella........................... ........25 Pseudomicrothorax............. ........15
Porolithon............................. ........26 PSEUDOM YCOTA.......... ....... 23
Porospora............................. ....... 17 Pseudopedinella.................. ........21
Porphyridium........................ ........26 Pseudoprorodon .................. ........ 16
! Postciliodesmatophora..... ........14 Pseudoscourfieldia.............. ........ 24
! Poterioochromonas.............. ........22 Pseudospora......................... ....... 12
Prasinophyceae.................. ........24, Table 1 Pseudotrachelocerca................... 17
PRASINOPHYTA............. ....... 24, Table 1 Pseudotrichomonas..................... 11
Prasiola.................................. ........24 Pseudovahlkampfia..................... 10
Proboveria............................ ....... 16 Pteridom onas...................... ........ 21
Procryptobia......................... ....... 12 PTERIDOPHYTA..................... 24
Promycetozoida................... ........ 13 Pterosperma........................ .........24
Propyxidium.................................. 16 Ptychodiscus........................ .........14
Prorocentrum........................ .... 14 Ptychostomum .................... .........16
Prorodon............................... ........ 16 Pycnothrix........................... .........17
! Prorodontida.................................. 16 Pyramimonas....................... .........24
1Prostomatea........................ ........ 16, 17, Table 1 Pyrenomonas....................... .........23
Prostomatida........................ ........ 16 Pyrocystis............................ .........14
Protalveolatea..................... .........13, 29, Table 1 Pyrophacus.......................... .........14
1 Proteomyxids....................... .........12 Pyrrhophy ta ...................... .........14
I Pfoteromonads..................... .........28 Pyrsonympha....................... .........10
Pterom onas....................... .........12, 30 Pythium................................ .........23
Pterospongia..................... .........13
Proterozoea........... .........12, 29, Table 1
i p r o t is t a ......................... ......... 4, 5,2 7 Q
’ Protociliata......................... .........12
ïr°tocruzia......................................15 Quasillagilis.................................... 17
^ O T O C T IS T A .......................... 4 Quinqueloculina..............................19
 1

48 J. O. Corliss

R Rotaliella.........................................19
Rudim icrosporea......................... 10, Table 1
R aabella.......................................... 15
R adiolaria..................................... 20, Table 1 S
Radiolarians................................... 20
Radiophrya...................................... 16 Saccam m ina................................... 19
R A D IO ZO A ..................................19, 29, Table 1 Saccamoeba.................................. 18
Raphidiophrys................................ 19 Saccinobaculus...............................10
Raphidomonadea.........................21, Table 1 Salpingoeca.................................... 13
Raphidonema................................. 25 Saprodinium ................................... 15
RAPHIDOPHYTA...................... 21, Table 1 Saprolegnia..................................... 23
Reclinomonas..................................12 Sarcinochrysis................................22
Red a lg a e ........................................2 4 - 2 6 Sarcocystis...................................... 18
Reichenowella................................ 15 "Sarcodina"..................................... 18
Remanella ...................................... 14 Sargassum....................................... 22
Resultor...........................................24 Saurocytozoon................................ 18
Reticulosphaera.............. 22 Scaphidiodon.................................. 15
Retortamonadea........................... 10, Table 1 Scenedesmus...................................24
Retortamonadida............................ 10 Schellackia.......................................18
Retortamonas..................................10 Schizammina................................... 19
Rhabdoaskenasia........................... 17 Schizocalyptra................................ 16
Rhabdom onadida........................... 11 Schizocystis......................................17
Rhabdomonas..................................11 Schizomerus....................................25
Rhabdophorids............................... 16 Schizopyrenida............................... 10
Rhabdophrya...................................15 Schwagerina................................... 19
Rhabdostyla....................................16 Sclerospora..................................... 23
Rhinom onas................................... 23 Scourfieldia....................................24
Rhinozeta......................................... 17 Scuticociliatia................................ 16
Rhipidium........................................23 Scyphidia.........................................16
R hizam m ina....................................19 Scytosiphon..................................... 22
Rhizidiom yces................................ 23 Sea lettuce...................................... 25
Rhizochromulina............................22 Sea w eeds....................................... 22, 25, 26
Rhizoclonium ................................. 25 Selenastrum....................................25
Rhizodinium ....................................14 Selenidioides................................... 17
Rhizophydium................................. 26 Selenidium .......................................17
RHIZOPODA.............................. 9, 10, 13, 18, 26, Selenita............................................ 19
Table 1 Selysina....................... 18
Rhizosolenia................................... 22 Sem itrichodina............................... 16
Rhizosphaera................................. 20 Sicuophora.......................................15
R hodella..........................................26 Siedleckia.........................................17
Rhodochaete................................... 26 Silicoflagellatea............................ 21, Table 1
Rhodomonas................................... 23 Silicoflagellates............................. 21, 22
R hodophyllis.................................. 26 Singhamoeba.................................. 10
RH O DO PHYTA..........................26, Table 1 Sinuolinea....................................... 20
Rhodospora.................................... 26 Siphonocladus................................25
Rhynchocystis..................................17 Sirogonium ..................................... 25
Rhynchomonas............................... 12 Slopalinida............ 12
R osalina........................ 19 "Slopalinids"...................................28
R oscoffia......................................... 14 Sny de relia........................................11
Rosculus.......................................... 18 Sonderia......................................... 17
 "User-friendly" Classification of the Protists 49

Sorites..............................................19 Stichotricha..................................... 15
Sorocarpus......................................22 Stilopsis...........................................22
Sorodiscus....................................... 12 - Stokesia............................................ 16
Sorogena......................................... 15 Stoneworts...................................... 25
j spathidiopsis.............................. . 16 "Stramenopiles"............................. 20, 28
Spathidium...................................... 17 "Straminopiles"..............................28
SPERMATOPHYTA.................. 24 Streptophyllum................................22
Sphaeromyxa...................................20 Strobilidium.................................... 15
Sphaeroplea....................................25 Strombidium................................... 15
Sphaerospora................................. 20 Stylocephalus.................................. 17
Sphenoderia............... 18 Stylochona.......................................15
Sphenomonas..................................11 Stylonychia..................................... 15
Sphenophrya........................... 15 Suctorians........................................15
: Spirillina......................................... 19 Symbiodinium................................ 14
Spirochona............. 15 Synchytrium................................... 26
, Spirogyra......................................... 25 Syndinium..................................... 14
Spiroloculina...................................19 Synura.............................................22
Spiromonas..................................... 17 Synurophyceae..............................22, Table 1
Spironema....................................... 12
Spironucleus....................................10 T
Spirostomum...................................15
j Spirotrichea...................................14, Table 1 Tachyblaston.............................. 15
Spirotrichia....................................15 Tardivesicula.................................10
; Spirotrichonympha...................... 11 Taxopodea..................................... 19, Table 1
Spizellomyces .............. 26 Telomyxa........................................ 10
Spongodrymus ....... 20 Teranympha...................................11
! Spongomonas.................................. 12 Territricha...................................... 15
Spongospora................................... 12 Tesselaria.......................................22
Sporadotrichs..................................15 Tetractinomyxon...........................20
Sporochnus..................... 22 Tetradimorpha.............................. 12
Sporolithon......................................26 Tetrahymena................................... 16
SPOROZOA................................ .17 Tetramastigamoeba....................... 10
Spraguea ............................ 10 Tetramitus...................... 10
Spumella.......................................... 22 Tetramyxa .............................. 12
^ Stannophyllum................................19 Tetraselmis...................................... 24
Staurastrum.....................................25 Tetraspora....................................... 25
Steinella............................................16 Tetrasporidium...............................25
Stemonitis........................................ 13 Textularia ................................ 19
Stempellia........................................ 10 Thalassicolla ...............................20
Stentor................................. 15 Thalassionema................................22
Stephanodiscus............................... 22 Thalassiosira.................................. 22
Stephanoeca....................................13 Thalassomyces............................. 14
Stephanopogon.................................11 , 30 Thalassomyxa................................. 13
Stephanosphaera............................25 Thalassophysa................................20
Stephanospora ............................... 17 Thaumatomastix.............................12
Stereomyxa...................................... 18 Thecacineta.................................... 15
; Stereonema...................................... 12 Thecamoeba................................... 18
Stichochrysis......................... 22 Theileria...................... 18
Stichococcus................................... 25 Thelohania.......................................10
, Sticholonche....................................19 Thigmocoma.............................. 16
50 J. O. Corliss

Thigmophrya...................................16 Triloculina.......................................19
Thigmotrichs...................................16 Trilospora....................................... 20
Thoracosphaera............................. 14 Trimitus............................................ 10
Thorea............................................. 26 Trinema............................................ 18
Thraustochytriacea..................... 21, Table 1 Triparma......................................... 22
Thraustochytrium...........................21 Trochilia...........................................15
Thylakidium..................................... 15 Troglodytella................................... 17
Tiarina............................................. 16 Trypanosoma.................................. 12, 30
Tintinnids........................................ 15 Trypanosomatida............................12
Tintinnopsis.....................................15 Trypanosom atids............................11
Tokophrya........................................ 15 Tubulina...........................................13
Tolypella..........................................25 Turaniella........................................16
Tontonia.......................................... 15 Tuzetia.............................................. 10
Toxarium.........................................22 Tyzzeria............................................ 18
Toxoplasma.....................................18
Trachelocerca................................ 14 U
Trachelomonas............................... 11
Trache Ionema..................................14 U lothrix.......................................... 25
Tracheloraphis............................... 14 U lva................................................. 25
Tracheophytes................................ 24 Ulvophyceae..................................25, 31, Table 1
Transitella....................................... 15 ULVOPHYTA...............................25, 31, Table 1
Trebouxia........................................25 Umbilicosphaera........................... 23
Trentepohlia................................... 25 Undella............................................ 15
Trepomonadea............................. 9, Table 1 Unicapsula...................................... 20
Trepomonas.....................................10 Unicauda........................................ 20
Treubaria.......................... 25 Unikaryon........................................10
Triadinium....................................... 17 Uradiophora................................... 17
Tribonema.......................................22 Urceolaria.......................................16
Tribophyceae................................ 22 Urocentrum..................................... 16
Triceratium..................................... 22 Uroglena......................................... 22
Trichamoeba...................................18 Uronema.......................................... 16
Trichia............................................. 13 Uronychia........................................15
Trichochona....................................15 Urosomoides................................... 15
Trichodina....................................... 16 Urospora......................................... 31
Trichodinopsis................................ 16 Urosporidium.................................20
Trichomonadea............................. 11, Table 1 Urostyla...........................................15
Trichomonadida............................. 11 Urotricha.........................................16
Trichomonas....................................11, 30 Urozo n a........................................... 16
Trichonympha................................ 11 Utriculidium...................................22
Trichonymphida............................. 11 Uvigerina.........................................19
Trichophilus................................... 25
Trichophrya....................... 15 V
Trichosarcina................................. 25
Trichosphaerium............................ 18 Vacuolaria...................................... 21
Trichospira ............................ 17 Vaginicola........................................16
Trichostomes...................................17 Vahlkampfia.................................... 10
Tricornia........................ .....10 Vairimorpha.................................... 10
Tricoronella.....................................15 Valonia.............................................25
Trigonomonas................................ 10 Vampyrella.......................................18
Trihym ena....................................... 15 Vampyrophrya................................ 16
 "User-friendly" Classification of the Protists 51

^annella............................................18 Xiphophora..................................... 22
Vasichona........................................ 15 Xystonellopsis................................. 15
Vasicola........................................... 16
Vaucheria........................................22 Y
Vegetabilia................ 6
Verruea lv u s .....................................23 Yamagishiella.................................25
Vestibuliferans................................17 Yellow-green algae........................ 22
Vestibulongum.................................17 Yvonniellina.................................... 15
Vexillifera........................................ 18
VIRIDIPLANTAE.......................24, 29, 31, Table 1 Z
Vischeria......................................... 22
Volvocida........................................25 Zelleriella........................................12
Volvocids......................................... 27 Zonaria............................................22
Volvocines....................................... 30 Zooanthella..................................... 14
Volvox.............................................. 25, 27 "Zoomastigophora"...................... 5
Vorticella......................................... 16 Zoophagus ......................... 23
Zoosporea........................................17
W Zoothamnium.................................. 16
Zosterodasys.................................. 15
Wallackia......................................... 15 Zschokkella..................................... 20
Wardia............................................. 20 Zygnem a......................................... 25
Wenrichia........................................ 16 Zygnematophyceae................... 25
Wenyonella...................................... 18 Zygocystis........................................17
Woodruffia....................... 15 ZYGOMYCOTA ............ 26
Woronina......................................... 12 Zygophyceae.................................25
Zymurgia......................................... 26

Xanthidium......................................25
Xanthophyceae.............................. 2 2 , Table 1
Xenophyophorea...........................19, Table 1
Xiphacantha ......................................19 Received on 2nd November, 1993