Annotated Assignment of Families and Genera to the Orders and Classes Currently Comprising the Corlissian Scheme of Higher Classification for the Phylum Ciliophora

Annotated Assignment of Families and Genera to the Orders and Classes CurrentlyComprising the Corlissian Scheme of Higher Classification for the Phylum CiliophoraAuthor(s): John O. CorlissSource: Transactions of the American Microscopical Society, Vol. 96, No. 1 (Jan., 1977), pp.104-140Published by: Wiley on behalf of American Microscopical SocietyStable URL: http://www.jstor.org/stable/3225971 .

Accessed: 28/06/2014 10:50

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact [email protected].

.

Wiley and American Microscopical Society are collaborating with JSTOR to digitize, preserve and extendaccess to Transactions of the American Microscopical Society.

http://www.jstor.org

This content downloaded from 141.101.201.167 on Sat, 28 Jun 2014 10:50:36 AMAll use subject to JSTOR Terms and Conditions

http://www.jstor.org/action/showPublisher?publisherCode=black

http://www.jstor.org/action/showPublisher?publisherCode=amicros

http://www.jstor.org/stable/3225971?origin=JSTOR-pdf

http://www.jstor.org/page/info/about/policies/terms.jsp


ANNOTATED ASSIGNMENT OF FAMILIES AND GENERA TO THE ORDERS AND CLASSES CURRENTLY COMPRISING THE

CORLISSIAN SCHEME OF HIGHER CLASSIFICATION FOR THE PHYLUM CILIOPHORA1

JOHN O. CORLISS Department of Zoology, University of Maryland, College Park, Maryland 20742

Corliss, J. 0. 1977. Annotated assignment of families and genera to the orders and classes currently comprising the Corlissian scheme of higher classification for the phylum Ciliophora. Trans. Amer. Micros. Soc., 96: 104-140. Although there has been a rash of recent publications treating suprafamilial ciliate taxa in various revisions, all offering drastic and rather expansive changes over the basic Faurean classification so popular in the fifties, sixties, and early seventies, no workers have yet given detailed attention to the important problem of properly locating existing families and genera in the new schemes proposed. Following communication with many colleagues around the world, the present paper is being published as an attempt to succinctly meet that deficit. Some 1,123 genera and 204 families, considered bonafide taxa, are assigned to the orders and suborders (in their respective subclasses and classes) comprising the writer's classification system for the phylum Ciliophora. Authors' names are supplied for all taxa, some of which are placed in incertae sedis positions. Comments are included in place with respect to groups-at all levels- considered most controversial taxonomically. In explanatory introductory sections preceding the scheme itself, such problems as synonyms and homonyms, nomina nuda, relative abundance of species in the genera listed, and certain other related nomenclatural matters are treated briefly.

The "changing world of ciliate systematics" (Corliss, 1974a) is perhaps beginning to settle down now, but not without the anticipated pains of change, including groans from researchers, teachers, and students alike. The novel ideas- often involving drastic alterations in the much simpler classification system of the 1960's-propounded nearly simultaneously in very recent years by members of the French school of protozoologists, by Jankowski in Leningrad, and by Cor- liss and Small in America have sprung from an increased awareness of the inevitable necessity of reflecting new knowledge in an improved way taxonomically.

Most of the attempts referred to above have concentrated on revision (and, alas, multiplication) of the suprafamilial taxa only. This has been a requisite first step, perhaps, but it has left the student, particularly, in the awkward position of having hundreds of ciliate families and genera (not to mention thousands of species!)-groups precisely placed in most of the pre-1970 schemes of classification-with unknown assignment to suborders, orders, or even classes. Philoso- phizing over supposed phylogenetic interrelationships among both old and new orders, for example, has been a stimulating and presumably worthwhile exercise; but revising and relocating groups of the actual ciliates themselves-that is, at the "working level" of genera and families-seems, with rare exception, to have been either postponed or neglected altogether. Spottiness and overall paucity of needed data have been responsible for such understandable delay. But an attempt is made to make up the lingering deficiency here-to the extent possible

1 Support of National Science Foundation grant 76-19272 is gratefully acknowledged. Once again, I am also deeply appreciative of the many discussions with-or critical comments received from-colleagues in ciliatology around the world. Such aid during the past several years has been most helpful, but I hardly need to mention that I alone am responsible for the conclusions which I have ultimately drawn and presented in this paper. TRANS. AMER. MICROS. Soc., 96(1): 104-140. 1977.



CORLISS-FAMILIES AND GENERA OF CILIATES 105

in a single paper of restricted length and with realization of the handicap of our limited knowledge.

BRIEF BACKGROUND REVIEW

New "High-level" Approaches There is no need to repeat here a discussion of the new approaches to the

taxonomy of the phylum Ciliophora introduced in the recent papers of Corliss (1974a,b), Jankowski (1967, 1972, 1973a,b), and de Puytorac et al. (1974). These works are referred to in some detail in other still more recent publications, often rich in bibliographic references, by Canella & Rocchi-Canella (1976), Corliss (1975a, 1976), Hutner & Corliss (1976), de Puytorac & Grain (1976), and Small (1976). The data playing the major role in the various revised classifications have come from ultrastructural studies (e.g., see Grain, 1969; Grain et al., 1973; Pitelka, 1969; and works by many other cell biologists not cited directly here), as might be expected; but renewed attention to morphogenetic patterns (e.g., see Corliss, 1968, 1973; Frankel, 1974; Sonneborn, 1974; Tuffrau et al., 1974) and to nuclear (especially macronuclear) differences (e.g., see Corliss, 1975b; Raikov, 1969) has also been a significant factor in recognizing a greater degree of diversity among the ciliates than subscribed to in past decades. Ecological factors, too, are coming to be viewed as of increasing importance to our understanding of the ciliates (e.g., see-among scores of references which could be mentioned-Bick, 1972; Borror, 1973; Cairns, 1971; Corliss, 1973a; Corliss & Hartwig, 1977; Curds, 1969; Dragesco, 1974; Fenchel. 1969; Finley et al. 1974).

Many aspects of the emerging schemes of classification have, happilv, had much in common; and some of the earlier outlines have been modified in a possible "compromising" (in the right sense of the word) approach toward ultimately a single "new new" classification for universal usage. There are, admittedly, areas of remaining controversy; unfortunatelv-tempted though I am-the present paper is not the appropriate place for lengthy discussion or defense of my own proposed resolutions of such cases. Perhaps these matters will receive some needed attention at the forthcoming-June-Tuly 1977- Vth International Congress of Protozoology convening in New York City. Most of the outstanding ones, however- are indicated by the brief comments appearing in place throughout the classification section, below.

While the long-awaited, multi-authored tome 2 of Grasse's Traite de Zoologie, to be devoted (I believe) exclusively to the ciliated protozoa, has been languish- ing "in press" and while a monographic publication by Jankowski is also being anticipated (see abstract by Jankowski, 1975), I have been working indepen- dently and intensely for many months on production of a totally revised and greatly expanded second edition of my own book (Corliss, 1961) on the systematics and evolution of the ciliates. All of the efforts just mentioned will presumably include detailed treatment of these organisms at the lower taxonomic levels; but the exact date of appearance of none of them is known at the time of this writing, so none is specifically cited.

Genesis of Present Work

Nearly two years ago (see Corliss, 1975a), I published an expanded classification system which included succinct diagnoses of the suprafamilial taxa and indicated the location within these groups of some 426 named genera (with a few illustrated), noting that much of the information was derived from unpub- lished material destined to comprise the "middle section" of my forthcoming book. The present paper, concentrating on the lower taxa (but, for want of space, including no characterizations or figures), is based on the same (now up-



106 TRANS. AMER. MICROS. SOC., VOL. 96, NO. 1, JANUARY 1977

dated) source and includes a few significant changes or corrections-based on the availability of fresh data-in the "higher-taxa" information presented in the earlier work.2

A draft or "preprint" of the bulk of this paper was sent out last summer to nearly 100 ciliate specialists and teachers of protozoology around the world, with an earnest request for critical comments. I have been helped by the replies received and must particularly acknowledge the quite detailed remarks from colleagues Berger, Borror, Deroux, Dragesco, Grain, Laval, Lom, Small, and Tappan, while hastening to point out that I have not always agreed with nor followed all such advice received. My principal reason for publishing the following pages here, in advance of my book, has been the plea from many kind recipients of the drafted manuscript-including textbook writers and teachers responding on behalf of their students in protozoology-to "please give us in print as soon as possible information on the proper location of the families and genera of ciliates in your new scheme of classification of the higher taxa."

Rationale, familial characterizations, reasons for lower-level taxonomic shifts, full explanation of many vital nomenclatural decisions, dates of names, illustra- tions, and exhaustive references to the pertinent literature must still await appearance of my book. The following sections of further "preamble," however, are considered minimally essential for understanding the major section-an annotated outline of the classification itself-which comprises the essence of the present paper.

NOMENCLATURAL MATTERS

Although nomenclature serves only as a handmaiden to systematics, its in- volvement is great in any revisory work. After a taxonomic decision has been made, the rules of nomenclature must inevitably be followed-painful task though this may be for many workers (see detailed treatment by Corliss, 1962). Errors are replete in the ciliatological literature-incredibly so! Amalgamation or splitting of families or genera, for example, requires careful attention with respect to the proper (there is little choice!) names to employ for the resulting new groups. Precise rules also exist for treatment of cases of homonymy and synonymy. There is freedom for the person working at the suprafamilial levels, though even here one would hope for demonstration of a measure of "common sense and courtesy" (Corliss, 1972). Since the bulk of this paper is composed solely of names, several specific nomenclatural topics must receive at least brief consideration here; stylistic and other relevant matters are reserved for subsequent sections.

Synonyms and Homonyms In the classification section of this paper, names which I have reduced to the

status of (junior) synonyms or names which I (or someone else) have (has)

2 Influenced in particular by continued dialogue with French colleagues (and see de Puytorac & Grain, 1976), I have effected several taxonomic changes of some significance in the classification system presented here over that published in the first paper (Corliss, 1975a) of this pair of largely "pedagogical" expositions. Worthy of mention are the following four major emendations, which should be considered to supersede the pertinent parts of the earlier version of the so-called Corlissian scheme of classification: the suborder Blepharocorythina is removed from the order Entodiniomorphida and is treated as a suborder of the neighboring order Trichostomatida; the order Cyrtophorida is divided into three suborders (Chlamydodon- tina, Dysteriina, and Hypocomatina); the order Suctorida is given an additional (third) suborder, the Evaginogenina; and the order Heterotrichida is also expanded to include a new suborder, the Clevelandellina, among its total (now) of six suborders. Minor changes have primarily involved shifting of a number of genera, generally based on new knowledge or new interpretation of older data during the interim since April 1975.




discovered to be preoccupied (i.e., they are junior homonyms) are, with rare exception, excluded. (They will be included, of course, in my forthcoming book.) There are hundreds of such names, all told. A few of them are listed below- names which are particularly familiar to many researchers and/or teachers and might thus be looked for (in vain!) on subsequent pages. They have been obliged to fall either due to homonymy or because some worker has recently made the taxonomic decision of combining the species (or genera) of two former genera (or families) into one: in which cases, solely the older names can legally prevail. (Several, of special importance, are cited in place in my classification.)

A few instances of "disappearing" (invalid) names at the familial level are the following, chosen arbitrarily: Cyclogrammidae and Urceolariidae, which unfortunately require replacement (Furgasoniidae and Urceolariellidae, respec- tively) because of being based on preoccupied generic names; and Askoellidae, Calpionellidae, Colomiellidae, Hypotrichidiidae, Pleurotrichidae, Pseudomicro- thoracidae, Synhymeniidae, Spirozonidae, Uronychiidae, and Zoothamniidae, names falling as junior synonyms (among many many others!) in my classification.

Sample cases of legitimately "missing" (no longer bonafide) generic names would include-again being highly selective-the following, divided into the two groups indicated. Some names which are obliged to fall by virtue of being junior homonyms: Avela, Brachyspira, Cyclogramma, Cystophrys, Deflandrella, Dia- phanosoma, Discomorpha, Epalxis, Fatureia, Faurella, Heterocoma, Histrio, Kahlia, Laurentia, Leptodiscus, Lionotus, Micromitra, Oxychona, Palmarium, Ptyxidiella, Pyxidium, Saprophilus, Stammeriella, Torquatella, Trichopelma, Tri- chopus, and Urceolaria. And some generally well-known names which are treated by me as junior synonyms: Aegyria, Ancistruma, Balanophrya, Centrophorella, Clclochaeta, Dallasia, Dexiostoma, Gyrocorys, Kudoella, Leucophrys, Pigostyla, Plagionassula, Polianskina, Semifolliculina, Stichochaeta, and Symbionecta.

It is to be understood that, except for the cases of proven homonymy, any of these generic names or familial names (including hundreds not listed above) are eligible for resurrection by subseauent workers who taxonomically believe that the taxa involved should be (re)split into smaller groups than those currently endorsed by me. Examples of the all too plentiful instances of misspelled names -which I arbitrarily list under synonyms (sensu lato) in my book-are given in a subsequent section of this paper. Such names have no legal standing in taxonomy, but they remain an everlasting source of potential confusion.

Names for Groups Not Yet Described When a taxonomic name is published for a group not yet described, that

name must be considered a "naked name" or nomen nudum (or nudum nomen), plural nomina nuda. Taxonomists are, rightly, discouraged from the (premature) publishing of such names; it is highly appropriate to have a new name introduced only at the time of the simultaneous appearance of the description or characterization, ideally including figures, of the group of organisms involved. Recent rather unfortunate cases, though perhaps inadvertent, include Jankowski's (1975) nearly 100 new familial names and Levine's (1972) use of Berger's names for several undescribed genera of echinophilic ciliates.

Nevertheless, for a specific, pre-explained, and presumably good reason, names of not-yet-described groups may sometimes be published to advantage. And I consider the present paper (and my book) to represent such instances. In order to fill out some of the newer taxa, especially, and to call the readers' attention to the current specialists on the groups concerned, I am-with the knowledge and understanding of the workers involved-deliberately citing a number of familial and generic names of organisms the descriptions of which, at




the time of this writing, have not yet appeared in print (though often they are known to be somewhere in press). Some 18 families (also four suborders) and 45 genera are so treated. The future authors involved (sometimes myself) are properly recognized by including their names after the taxonomic name which, itself, is preceded by an asterisk (*) in this paper. A principal justification for my doing this is to alert ciliatologists and others to the "near-existence" of such often important new (or newly named) groups. Indeed, in a number of cases, the descriptions of the implicated organisms may well appear before or at the same time as this paper (or my book), so I am happy to be able to give proper, if slightly advance or in parallel, credit to the actual describers (or namers) of such ciliates.

A further note on this important, but possibly easily misunderstood, matter is in order. Generic "n. n." names are eventually to be credited, author and date, to the worker(s) ultimately publishing the formal description, wherever and whenever that may be; but "n. n." names of families (and similarly for the four suborders) so designated below (by the asterisk) are generally to be treated a little differently in the future. Unless the families involved are described by their authors before the appearance of my book, their names will have to be cited in full in the future as (for a hypothetical example) "X-idae Smythe in Corliss, 1977,3" because-unlike the situation for the genera mentioned above (whose names will be merely listed by me without diagnoses)-my book will include familial characterizations, descriptions thus appearing at the same time as the names themselves. "Smythe" will be receiving proper "credit," but necessarily "as of" or "in" my book (which will presumably bear a 1977 date). The possible nomenclatural complications arising from near-simultaneous publication (by "Smythe") elsewhere can easily be resolved by any subsequent taxonomic re- viewer of the case, who will be obliged by the rules to choose the earlier author- and-date, a policy with which, naturally, I am wholly in accord.

Taxa of Uncertain Position The more we learn about ciliates the less sure most of us become with

respect to their true taxonomic interrelationships! Yet, from both pragmatic and pedagogical points of view, we can hardly assign the bulk of these organisms to an uncertain position in some kind of appendix to the whole phylum Cili- ophora. At any given time, combining results of analyses of hard data with experienced intuition is justification enough for drawing tentatively reasonable conclusions concerning affinities and for reflecting this as ably as possible in a revised taxonomic scheme. One thing is certain: the "new new" classification, with all of its possible shortcomings, is a decided improvement over older schemes with respect to likely intergroup affinities. On this there is wide agreement among the specialists concerned, despite differences on more or less minor points. Any revised classification is "premature," of course, in the sense that all of the pertinent data desirable will never be available!

There are several places in the ciliate taxonomic system where I am relatively much less sure of relationships than I am in others, so there seems to be no good alternative to the occasional usage of an incertae sedis category to handle the more highly perplexing cases. While deliberately invoking this alternative as sparingly as possible, I have nevertheless used it in scattered places throughout the classification which occupies the following pages-usually only at the generic

3 Unless the asterisk-marked name is being proposed by me in my forthcoming monograph. In such cases, the book's date will simply follow my name in any complete citation of the nominal taxa involved: for example, family Phacodiniidae Corliss, 1977, or genus Urceolariella Corliss, 1977. Types will be designated for all of my new (or newly named) taxa.




level but sometimes, in frustration and/or desperation, at the familial level as well. In some of these instances, further study (really needed for ca. 90% of all ciliate genera!) may show that the uncertain organism's name should fall as a synonym of that of a better-known genus (or family). Aware of the inadequacies of many descriptions, I have, nevertheless, always been generous in accepting new genera of other workers, until or unless subsequent studies have quite con- vincingly demonstrated the lack of uniqueness of such taxa.

Another way of my indicating either personal uncertainty or areas of continu- ing controversy among the specialists of the groups concerned is by the inclusion of occasional remarks or comments, in place, in the classification section of this paper. I have also sometimes supplied the name of a homonym which is being replaced, if it had been a widely known generic name. Such actions-along with the use of "incertae sedis" (see above) and giving authors' names and an indica- tion of numbers of species per genus (explained below)-may be considered to augment the critical assignment of the hundreds of known ciliate families and genera to the higher taxonomic groups which I endorse at this time.

Total Numbers of Taxonomic Names Involved As can be ascertained from the listings presented below, I am here accepting

some 1,123 generic names and 204 familial names, plus 70 names at the higher levels (including the phylum name itself), as valid. This is a significant increase over the figures in Corliss (1961), viz.: ca. 275 more genera, 75 more families, and 37 additional higher taxa. But it is far below the totals suggested in such a work as Jankowski (1975). I recognize some 7,200 bonafide species, an increase of nearly 1,500 over a count made by me in 1960. Approximate numbers of species are indicated in the present classification by "+" signs following generic names, as explained in a following section.

The total number of one-time published names for ciliate groups at all levels above that of species is truly astounding: it would raise my total of 1,397 (see above) to well over 2,300!! So, it would seem that-after all-I am more of a lumper than a splitter (see Corliss, 1976), despite my enthusiastic support of a classification (e.g., in this paper) representing a considerable expansion over the basic Faurean scheme (see Corliss, 1961; Faure-Fremiet, 1950; Honigberg et al., 1964; Jones, 1974; Kudo, 1966; Lep?i, 1965; Manwell, 1968; Sleigh, 1973; and the not too dissimilar classifications-with respect to numbers of taxa-of Grell, 1973; Poljansky & Cheissin, 1965; Raabe, 1964; and Westphal, 1974).

It should be noted that the grand total of unique nomenclatural "entities" proposed by workers to date (ca. 2,300) includes counting not only synonyms and homonyms but also variants in spelling, which have been numerous. I consider it important, in my book, to list many of these variants (under synonyms sensu lato) because such misspellings, once perpetrated, have frequently been perpetuated-via textbooks and the like-to the extent that students and researchers alike are often confused as to what the legitimate form of a given name actually is. A few examples of common misspellings among generic names-not appearing generally on the following pages where mainly the correct names are printed-are these: Chaenia, Circundella, Conchophthirius, Conidiophrys, Curinostoma, Intrastylum, Lichnophora, Metacyclis, Paramaecium, Potamacus, Protoanoplophrya, Protocrucia, Spiratella, Squalophrya, Telotrichidium, Teuto- phrys, and Thylacidium. Perhaps most notorious among familial names is the popularly written "Epistylidae," which should appear only in the proper form of Epistylididae. Many additional cases could be cited.

Also to be kept in mind is the fact that accents and diacritical marks (hy- phens, too) are simply not allowed by the Code in Latin or latinized names. So genera originally (or subsequently) called Biitschlia, Lemboides (for two




examples), etc. must be emended appropriately, remaining themselves as in- admissable orthographic variants. I do not include such "misspellings" as synonyms in my book.

NUMBERS OF CILIATE SPECIES

It is most difficult to estimate the number of bonafide species of ciliates, often even within quite well-known genera (unless the number is only one). Nearly all ciliate genera are urgently in need of revision, a task which cannot be tackled successfully by a single investigator even in a lifetime of work. Careful studies generally result in expansion of families or genera; yet some names also frequently become synonymized. Only specialists on a given restricted group can straighten out its contents: but, alas, there are both "lumpers" and "splitters" among the experts! Although seldom based, therefore, on personal in-depth investigation, estimated figures representing relative abundance of species per genus are provided in this paper according to the method of notation described in the following section. Such figures are, of course, subject to change as new, carefully carried out revisory studies become available in the literature.

The Largest Genera For protozoa in general, few genera are considered to be rich in species-

when compared to such animal groups as the insects, in particular. Thus it may be of more than passing interest, especially to evolutionarily- or ecologically- oriented students, to name the groups of ciliates which do contain quite a few species. First I should point out that nearly 50% of my 1,123 genera are mono- typic (one species only).

The following eight "popular" genera are considered to have more than 75 species each: Acineta, Cothurnia, Epistylis, Opercularia, Spathidium, Tintinnop- sis, Trichodina, and Vorticella. If we drop to the level of ca. 50 species per genus, these additional groups can be added to the preceding very short list: Balantidium, Codonellopsis, Cymatocylis, Entodinium, Euplotes, Lacrymaria, Lagenophrys, Metopus, Nyctotheroides, Nyctotherus, Orbopercularia, Oxytricha, Podophrya, Prorodon, Tokophrya, Vaginicola, and Zoothamnium.

The position of these genera in the overall scheme of classification can be ascertained by finding their names again in later pages of this paper. In general, they are well distributed among the classes and orders; and symbiotic as well as free-living forms are represented.

Distribution by Higher Taxa Considered by the higher taxonomic groups, the estimated numbers of species

per class is quite even, though with a few less in the middle class than in the others: Kinetofragminophora, with 2,600 species; Oligohymenophora, with 2,100; and Polyhymenophora, with 2,500. The first class, interestingly enough, contains nearly 50% of the genera comprising the whole phylum. At the ordinal level, the seven taxa with the greatest number of species, arranged in descending order, are the following: Oligotrichida, Peritrichida, Hypotrichida, Suctorida, Haptorida (embracing the bulk of the old category of "rhabdophorine gymnostomes"), Heterotrichida, and, perhaps surprisingly, the relatively new group Scutico- ciliatida.

Symbiotic vs. Free-Living Forms With respect to totally "free-living" versus "parasitic" species, it is difficult

to give precise figures because so many groups of ciliates (with one or two major exceptions) do not fit either extreme. It is clear, however, primarily because of




such numerous forms as the pelagic tintinnines, the (bulk of the) hypotrichs, and many heterotrichines and gymnostomes and vestibuliferans (including psammophilic and "terrestrial" species), that the majority of ciliates are non- symbiotic in their habitat preferences-probably ca. 65% of the 7,200 species I recognize as comprising the phylum.

The other 35% of ciliophoran species may be divided roughly equally between "true" commensals (with a few pathogenically parasitic in or on their hosts) and symphorionts, many of the latter (such as numerous sessile peritrichs, chonotrichs, suctorians, and folliculinids) using the integument of other organisms merely as a convenient substratum for attachment. The principal com- mensalistic groups are the well-known entodiniomorphids, the apostomes, some trichostomes sensu lato, the thigmotrichs (and parts of related neighboring scuticociliate groups), the symbiotic heterotrichs, some mobiline peritrichs, the archi- stomatines, and members of a few small hypostomate (or other) taxa.

FORMAT AND ABBREVIATIONS

Arrangement of Taxa Names of suprafamilial taxa are arranged essentially as they will appear in my

book, very similar to the style used in Corliss (1975a) though some changes have been made4 and authors' names (but not dates) are included here directly in the text. Subclass names are in boldface type; classes and orders, in caps. Superorders, superfamilies, subfamilies, and subgenera are not used below nor in my book: there is nothing illegitimate about such taxa, but I personally believe that there is little value in employing them for ciliates at this time.

Familial names are centered and given in capital letters. The families themselves are arranged phylogenetically, to the best of my ability; often few data are available to aid in making such decisions. Generic names considered by me to be valid are presented alphabetically, in columns, within their families. Synonyms are seldom supplied; but a few homonyms are given.

Genera (occasionally families) of uncertain taxonomic location are given following the abbreviated words "Incert. sed.5," if their names are associated with the immediately preceding family-group; if the position is uncertain within a higher taxon, more information is provided: for example, "Incertae sedis5 in order X-ida." The location or the uniqueness of such taxa may be considered unknown or in doubt with respect to the current state of our (or at least my) knowledge about them. Remarks are placed in parentheses (sometimes in a separate paragraph) following the group or groups to which they make reference.

Nomina Nuda Taxa As explained in a preceding section, some of the names included in this paper

are those of taxa not yet formally (at least under the new name) described in the literature. Legally, these must be considered nomina nuda; they are preceded by an asterisk (*) and are followed, as are all names, by the (in these cases, potential) author's name. The reasons for making an exception to the generally sound advice of avoiding premature publication of taxonomic names have been presented on an earlier page. There are, surely, still further new genera (not to mention families) in papers in preparation by other workers, with planned names concerning which I have had no advance knowledge (thus I was unable to ask for permission to include such additional taxa here).

4See footnote number 2, on a preceding page, for a brief listing of the major suprafamilial changes in this paper over the scheme of classification presented in my earlier paper (Corliss, 1975a).

5 These Latin words are purposely not italicized on the following pages to avoid possible confusion with generic names and to help break the monotony of so much italicization there.




Species per Genera The problems of determining the most likely number of valid, acceptable, or

bonafide species have been discussed above, albeit very briefly. The relative abundance of species in each genus, in my perhaps generous assessment, is indicated in this paper by use of the following arbitrary scale: no mark after a generic name indicates 1-4 included species; + = 5-12; ++ = 13-34; +++ = 35-75; and ++++ (rare!) = >75.

Treatment of Authors' Names

Partly for appearance and for space-saving but also because many ciliate taxonomists are well known, a number of authors' names are presented in an abbreviated style following the names of the taxa which they have created (see legend, below). The words "et al." are generally used here when three or more authors are involved, although such multiple authorships will always be given in full in my book. Whenever "de Puy. et al." is used, it refers to the 16-authored work of 1974 (see reference section) which resulted from the celebrated international "Reunion de Systematique" convened in Clermont-Ferrand in the spring of 1973. The dates of names are omitted6, but they will appear in my forthcoming book; they are available for suprafamilial taxa in Corliss (1975a).

The (generally lengthy) names of some 80 selected individuals or teams of workers are thus arbitrarily abbreviated according to the following legend:

Agam. & Alek. = Agamaliev & Alekperov

Alb. = Albaret Art. & Unti = Artigas & Unti Awer. = Awerinzew Awer. & Mutaf. = Awerinzew &

Mutafowa Bloch. = Blochmann Bory = Bory de St. Vincent But. = Bitschli Caul. = Caullery Cep. = Cepede Ch. = Chatton Ch. & Beauch. = Chatton &

de Beauchamp Ch. & Lw. = Chatton & Lwoff Ch. & Per. = Chatton & Perard Clap. & Lach. = Claparede &

Lachmann Coll. = Collin Conn in C. & Edm. = Conn in Conn &

Edmondson

Corl. = Corliss Cuvil. & Sacal = Cuvillier & Sacal da C. = da Cunha da C. & Mun. = da Cunha & Muniz Defl. = Deflandre Dog. & Furss. = Dogiel & Furssenko Drag. = Dragesco Duj. = Dujardin Ehr. = Ehrenberg Engel. = Engelmann Fab.-Dom. = Fabre-Domergue F-F = Faure-Fremiet Foiss. = Foissner Gaj. = Gajewskaja Gass. = Gassovsky Gelei = J. von Gelei Georg. = Georgevitch Good. & Jahn = Goodrich & Jahn Gr. & Roes. = Gourret & Roeser Heid. = Heidenreich Holl. & Bat. = Hollande & Batisse Jank. = Jankowski

The principal reason for leaving dates out here is lack of space to offer explanatory information in support of the choices which I have sometimes made. Such comments are often advisable because of the numerous errors in the literature with respect to the proper year of publication of works containing the first acceptable descriptions of many ciliate taxa, especially genera and families. (One example of a constant source of confusion occurs when the number of a journal containing important taxonomic-nomenclatural material appears in the year following the year that it bears as a cover date. Another problem arises when a new name is published as new more than once! Etc.) In controversial cases, the reader might become confused if I included here only what I believe to be the correct date, without explanation; hence such information has been reserved for my book.




Jar. = Jarocki Jir. = Jirovec Jorg. = Jorgensen K. & C. = Kofoid & Campbell K. & MacL. = Kofoid & MacLennan Kidd. & Summ. = Kidder & Summers Kopp. = Kopperi Koz. = Kozloff L. = Linnaeus Latt. = Latteur Laut. = Lauterborn L6p.-Och. = L6pez-Ochoterena MacL. & Conn. = MacLennan &

Connell N-T = Noirot-Timothee Naid. & Zaika = Naidenova & Zaika O.F.M. = 0. F. Muller Pen. = Penard Per. & Aim. = Pereira & Almeida Polj. = Poljansky Polj. & Gol. = Poljansky & Golikova de Puy. = de Puytorac

Quenn. = Quennerstedt Rad. = Radoicic Reich. = Reichenow Saed. & Tell. = Saedeleer & Tellier Santh. & Nair = Santhakumari & Nair Schew. = Schewiakoff Sr.-Husek = Sramek-Husek St. = Stein Strel. & Virk. = Strelkov & Virketis Swarcz. = Swarczewsky Tapp. & Loeb. = Tappan & Loeblich Thomp. = Thompson Thur. = Thurston Tuc. = Tucolesco Utt. & Desai = Uttangi & Desai Vill.-Brach. = Villeneuve-Brachon Voros. = V6orsvary Vux. = Vuxanovici Wallen. = Wallengren Wood. & Lodge = Woodcock & Lodge Wrzes. = Wrzesniowski

The full Corlissian classification scheme for the ciliates, including families and genera acceptable to me at this date and superseding all of my earlier attempts, follows (below), annotated with remarks in appropriate places.

Phylum C I L I O P H O R A Doflein

Class I. KINETOFRAGMINOPHORA de Puy. et al.

Subclass (1) Gymnostomata But.

Order 1. PRIMOCILIATIDA Corl.

Fam. STEPHANOPOGONIDAE Corl.

Stephanopogon Entz

Order 2. KARYORELICTIDA Corl.

Fam. TRACHELOCERCIDAE Kent

Trachelocerca Ehr. ++ Trachelonema Drag. +

Tracheloraphis Drag. +++

Incert. sed.: Gruvelina Delphy, Nephrocerca Delphy, Protrichophora Delphy

Fam. LOXODIDAE But.

Kentrophoros Sauerbrey ++ (syn. Centrophorella)

Loxodes Ehr. + Remanella Kahl ++

Fam. GELEIIDAE Kahl

*Avelia Nouzarede (for Avela) Geleia Kahl ++




Incertae sedis in order Karyorelictida:

Ciliofaurea Drag. +, Corlissia Drag., Cryptopharynx Kahl

(Remark: Uncertainties exist concerning overall taxonomic affinities of the two preceding orders.)

Order 3. PROSTOMATIDA Schew.

Suborder (1) Archistomatina de Puy. et al.

Fam. BUETSCHLIIDAE Poche

Alloiozona Hsiung Ampullacula Hsiung Amylophorus Per. & Aim. Blepharocodon Bundle Blepharoconus Gass. Blepharomonas Kopp. Blepharoplanum Kopp. Blepharoprosthium Bundle Blepharosphaera Bundle Blepharozoum Gass. Buetschlia Schuberg Buissonella da C. & Mun. Bundleia da C. & Mun. Cucurbella Thur. & Grain Didesmis Fiorentini

Hemiprorodon Strelkow Holophryoides Gass. Holophryozoon Jir. Kopperia Corl. Levanderella Kopp. Meiostoma Sandon Paraisotrichopsis Gass. Pingius Hsiung Polymorphella Corl. Prorodonopsis Gass. Protolutzia da C. & Mun. Pseudobuetschlia Jir. Sciurula Corl. Sulcoarcus Hsiung

Suborder (2) Prostomatina Schew.

Fam. HOLOPHRYIDAE Perty

Acaryophrya Andre (syn. Balanophrya) Holophrya Ehr. +

Incert. sed.: Balanion Wulff

Fam. METACYSTIDAE Kahl

Metacystis Cohn ++ Pelatractus Kahl

Vasicola Tatem +

Suborder (3) Prorodontina Corl.

Fam. PRORODONTIDAE Kent

Bursellopsis Corl. Helicoprorodon F-F Placus Cohn Plagiocampa Schew. ++ Prorodon Ehr. +++

Pseudoprorodon Bloch. ++ Rhagadostoma Kahl Spathidiopsis Fab.-Dom. + Spiroprorodon Fenchel & Lee Urotricha Clap. & Lach. ++

Incert. sed.: Amphibothrella R. & L. Grandori, Peridion Vux., Peridionella Vux.

Fam. COLEPIDAE Ehr.

Coleps Nitzsch ++ Tiarina Bergh



CORLISS-FAMILIES AND GENERA OF CILIATES

Order 4. HAPTORIDA Corl. Fam. ENCHELYIDAE Ehr.

Chaenea Quenn. ++ (syn. Chaenia) Microregma Kahl Chilophrya Kahl Nannophrya Kahl Crobylura Andre Pithothorax Kahl Enchelyodon Clap. & Lach. ++ Plagiopogon St. Enchelys O.F.M. ++ Quasillagilis Busch Gymnozoum Meunier Rhopalophrya Kahl ++ Haematophagus Wood. & Lodge Schewiakoffia Corl. Ileonema Stokes Spasmostoma Kahl Lacrymaria Bory +++ Sphaerobactrum Schmidt Lagynophrya Kahl ++ Thalassiomastix Busch Lagynurus Mansfeld Trachelophyllum Clap. & Lach. + Longitricha Gaj. Urochaenia Savi

Incert. sed.: Microcardiosoma Vux., Microchoanostoma Vux.

Fam. SPATHIDIIDAE Kahl in Doflein & Reich.

Bryophyllum Kahl ++ Cranotheridium Schew. Diceratula Corl. Enchelydium Kahl + Homalozoon Stokes + Lacerus Jank. Legendrea F-F Micromidas Delphy

Incert. sed.: Proboscidium Meunier

Penardiella Kahl Perispira St. Spathidiodes Kahl Spathidioides Brodsky Spathidiosus Gaj. Spathidium Duj. ++++ Thysanomorpha Jank.

(Remark: Exact composition of the two preceding families remains to be determined.)

Fam. TRACHELIIDAE Ehr. Branchioecetes Kahl Paradileptus Wenrich + Dileptus Duj. +++ Teuthophrys Ch. & Beauch Micruncus Delphy Teutophrys) Myriokaryon Jank. Trachelius Schrank +

Incert. sed.: Ctenoctophrys Weill

Acropisthium Perty Askenasia Bloch. Choanostoma Wang Cyclotrichium Meunier + Didinium St. +

Fam. DIDINIIDAE Poche

Dinophrya But. Liliimorpha Gaj. Mesodinium St. + Monodinium Fab.-Dom. + Zonotrichium Meunier

Fam. ACTINOBOLINIDAE Kahl Actinobolina Strand Dactylochlamys Laut. Belonophrya Andre Enchelyomorpha Kahl

Incertae sedis in preceding order(s) Prostomatida and/or Haptorida: Baznosanuia Tuc., Celeritia Tuc., Pseudoenchelys Tuc., Racovitzaia Tuc.; and fam. PROTOHALLIIDAE da C. & Mun., with Protohallia da C. & Mun.

particularly

.(syn.

115




Order 5. PLEUROSTOMATIDA Schew.

Fam. AMPHILEPTIDAE But.

Acineria Duj. Amphileptus Ehr. + Heminotus Kahl

Hemiophrys Wrzes. ++ Litonotus Wrzes. +++ Loxophyllum Duj. +++

(Remark: Some workers would reduce the number of separate orders and suborders from the eight which I endorse for my subclass Gymnosto- mata.)

Subclass (2) Vestibulifera de Puy. et al.

Order 1. TRICHOSTOMATIDA But.

Suborder (1) Trichostomatina But.

Fam. PLAGIOPYLIDAE Schew.

Lechriopyla Lynch Paraplagiopyla Thur. & Grain Parasonderia F-F

Plagiopyla St. + Sonderia Kahl + Sonderiella Kahl

Incert. sed.: Schizocaryum Polj. & Gol. (deserves family of its own?)

Fam. COELOSOMIDIDAE Corl. Coelosomides Strand Conchostoma F-F

Incert. sed.: Epimecophrya Kahl, Paraspathidium Noland (a gymnostome?)

Fam. TRICHOSPIRIDAE Kahl

Spirozona Kahl

Fam. TRIMYEMIDAE Kahl

Trimyema Lackey +

Maryna Gruber +

Fam. MARYNIDAE Poche

Mycterothrix Laut.

Fam. BALANTIDIIDAE Reich. in Doflein & Reich.

Balantidium Clap. & Lach. +++

Fam. PYCNOTRICHIDAE Poche

Buxtonella Jameson Collinina Ch. & Per. Infundibulorium Bozhenko Muniziella da Fonseca

Dasytricha Schuberg Isotricha St. +

Fam. ISOTRICHIDAE But.

Protoisotricha Kopp.

Trichospira Roux

Nicollella Ch. & Per. Pycnothrix Schubotz Taliaferria Hegner & Rees




Fam. PARAISOTRICHIDAE da C. Enterophrya Hasselmann Rhizotricha Wolska Paraisotricha Fiorentini

Incert. sed.: Helicozoster Latt.

Fam. *PROTOCAVIELLIDAE Grain

Hydrochoerella da C. & Mun. Protocaviella Kopp.

Suborder (2) Blepharocorythina Wolska

Fam. BLEPHAROCORYTHIDAE Hsiung Blepharocorys Bundle + Ochoterenaia Chavarria Charonina Strand Pararaabena Wolska Charonnautes Strelkow Raabena Wolska Circodinium Wolska Spirocorys Wolska

Order 2. ENTODINIOMORPHIDA Reich. in Doflein & Reich.

Fam. OPHRYOSCOLECIDAE St. Caloscolex Dogiel Campylodinium Jank. Cunhaia Hasselmann Diplodinium Schuberg ++ Diploplastron K. & MacL. Elytroplastron K. & MacL. Enoploplastron K. & MacL. + Entodinium St. +++ Eodinium K. & MacL. +

Epidinium Crawley + Epiplastron K. & MacL. Eremoplastron K. & MacL. ++ Eudiplodinium Dogiel ++ Metadinium Awer. & Mutaf. + Ophryoscolex St. Opisthotrichum Buisson Ostracodinium Dogiel ++ Polyplastron Dogiel +

Fam. CYCLOPOSTHIIDAE Poche Bertolinella Carpano Bozasella Buisson Cycloposthium Bundle + Diplolophus Holl. & Bat. Lavierella Buisson Monoposthium Thur. & N-T Parentodinium Thur. & N-T

Prototapirella da C. Rhabdothorax Latt. & Bousez Toxodinium da C. Tricaudalia Buisson Trifascicularia Strelkow Tripalmaria Gass. Triplumaria Hoare

Fam. POLYDINIELLIDAE Corl.

Elephantophilus Kofoid Pterodinium Lal Polydiniella Corl. Thoracodinium

Fam. SPIRODINIIDAE Strelkow

Cochliatoxum Gass. Spirodinium Fio

Ditoxum Gass. Tetratoxum Gass.

tt. & Dartevelle Latt.

rentini

Fam. DITOXIDAE Strelkow

Triadinium Fiorentini




Fam. TELAMODINIIDAE Latt. & Dufey

Megadinium Latt. & Dufey Teratodinium Latt. & Dufey Telamodinium Latt. & Dufey

Fam. *TROGLODYTELLIDAE Corl.

Troglodytella Brumpt & Joyeux

(Remark: There should perhaps be fewer separate families, and fewer bonafide genera within the larger families, in this order than are indicated above.)

Order 3. COLPODIDA de Puy. et al.

Fam. COLPODIDAE Ehr. Bresslaua Kahl Bursostoma Voros. Colpoda O.F.M. ++

Cirrophrya Gellert

Kalometopia Bramy Tillina Gruber +

Fam. WOODRUFFIIDAE Gelei

Woodruffia Kahl + Platyophrya Kahl +

Incert. sed.: Bryophrya Kahl, Rhyposophrya Kahl

(Remark: This family, perhaps along with the following one, is now being considered by some workers to be a member of the subclass Hypo- stomata, below.)

Fam. CYRTOLOPHOSIDIDAE Stokes

Cyrtolophosis Stokes +

Incertae sedis in preceding order(s) Trichostomatida and/or Colpodida: Grandoria Corl., Opisthostomatella Corl., Orcavia Tuc., Rigchostoma Vux., Sigalasia Delphy; and fam. SULCIGERIDAE Gaj., with Sulcigera Gaj.

Subclass (3) Hypostomata Schew.

Order 1. SYNHYMENIIDA de Puy. et al.

Fam. *NASSULOPSIDAE Deroux

Nassulopsis F-F +

Orthodonella Bhatia Synhymenia Jank.

*Phasmatopsis Deroux

Fam. ORTHODONELLIDAE Jank.

*Zosterodasys Deroux

Incert. sed.: Eucamptocerca da C.

Fam. *SCAPHIDIODONTIDAE Deroux

Chilodontopsis Bloch. Scaphidiodon St.




Order 2. NASSULIDA Jank. Suborder (1) Nassulina Jank.

Fam. NASSULIDAE de Fromentel

Enigmostoma Jank. Nassula Ehr. ++ Incert. sed.: Archinassula Kahl, Chilodina Sr.-Husek,

Das et al., Stomatophrya Kahl

Enneameron Jank.

Fam. PARANASSULIDAE F-F

Paranassula Kahl

Fam. *FURGASONIIDAE Corl.

Furgasonia Jank. (for Cyclogramma)

(Remark: This family is now being considered by some workers as a primitive group-perhaps a separate order-in the class Oligohymenophora, below.)

Suborder (2) Microthoracina Jank.

Fam. LEPTOPHARYNGIDAE Kahl

Leptopharynx Mermod + Pseudomicrothorax Mermod

Fam. MICROTHORACIDAE Wrzes.

Drepanomonas Fresenius + Hemicyclium Eberhard Kreyella Kahl

Microthorax Engel. ++ Trochiliopsis Pen.

Incert. sed.: Conchophrys Ch., Discotricha Tuffrau (deserves family of its own?), Hexotricha Conn in C. & Edm., Microdiaphanosoma Wenzel

Order 3. CYRTOPHORIDA F-F in Corl. Suborder (1) *Chlamydodontina Deroux

Fam. CHILODONELLIDAE Deroux

Chilodonella Strand +++ *Thigmogaster Deroux Phascolodon St. Trithigmostoma Jank.

Incert. sed.: Chilodonatella Drag., Odontochlamys Certes, Phyllotrichum Engel.

Fam. CHLAMYDODONTIDAE St.

Chlamydodon Ehr. +

Atopochilodon Kahl Chlamydonella Deroux *Coeloperix Deroux

Cyrtophoron Deroux

Fam. LYNCHELLIDAE Jank. Gastronauta Engel. Lynchella Kahl +

Incert. sed.: Lophophorina Pen.

Cyrtohymenostoma

Stammeridium Wenzel (for Stammeriella)




Suborder (2) *Dysteriina Deroux

Fam. *PLESIOTRICHOPIDAE Deroux

*Atelepithites Deroux Plesiotrichopus F-F Pithites Deroux & Drag. *Trochochilodon Deroux

Fam. HARTMANNULIDAE Poche

Aegyriana Deroux Allosphaerium Kidd. & Summ. + Brooklynella Lom & Nigrelli *Chlamydonyx Deroux Hartmannula Poche *Microxysma Deroux

*Orthotrochilia Deroux Parachilodonella Drag. Paratrochilia Kahl Sigmocineta Jank. Trichopodiella Corl. Trochilioides Kahl +

Incert. sed.: *Horocontus Deroux

(Remark: Does Allosphaerium deserve a separate family of its own?)

Fam. DYSTERIIDAE Clap. & Lach.

*Agnathodysteria Deroux Mirodysteria Kahl Dysteria Huxley +++ *Schedotrochilia Deroux Hartmannulopsis Deroux & Drag. Trochilia Duj. +

Suborder (3) *Hypocomatina Deroux

Fam. HYPOCOMIDAE But.

Hypocoma Gruber Rynchocoma Jank. (for Heterocoma) Parahypocoma Ch. & Lw.

Fam. CRATERISTOMATIDAE Jank. Crateristoma Jank.

Order 4. CHONOTRICHIDA Wallen.

Suborder (1) Exogemmina Jank.

Eleutherochona Jank. Lobochona Dons

Fam. LOBOCHONIDAE Jank.

Oenophorachona Toxochona Jank.

Matsudo & Mohr

Fam. FILICHONIDAE Jank. Aurichona Jank. Filichona Jank.

Fam. PHYLLOCHONIDAE Jank. Phyllochona Jank.

Fam. HELIOCHONIDAE Jank. Heliochona Plate Heterochona Jank. +

Fam. CHILODOCHONIDAE Wallen. Chilodochona Wallen. Vasichona Jank. Cryptochona Jank.




Cavichona Jank. ++ Serpentichona Jank.

Fam. SPIROCHONIDAE St. Spirochona St. +

Suborder (2) Endogemmina Jank.

*Cetochona Jank. *Fallainochona Jank. Isochona Jank. +

Actinichona Jank. Carinichona Jank. Crassichona Jank.

Armichona Jank. Ctenochona Jank. Cyamichona Jank. Dentichona Jank. Eriochona Jank. Flectichona Jank. Inermichona Jank.

Coronochona Jank. Echinichona Jank.

Ceratochona Jank. Chonosaurus Jank. Inversochona Jank.

Fam. ISOCHONIDAE Jank. *Paracetochona Jank. Trichochona Mohr

Fam. ACTINICHONIDAE Jank. Cristichona Jank. Kentrochonopsis Doflein Rhizochona Jank.

Fam. STYLOCHONIDAE Mohr *Oxychonina Corl. Paraoxychona Jank. Pterochona Jank. Spinichona Jank. Stylochona Kent + Thalassochona Jank.

Fam. ECHINICHONIDAE Jank. Eurychona Jank.

Fam. INVERSOCHONIDAE Jank. Kentrochona Rompel Pleochona Jank.

Order 5. RHYNCHODIDA Ch. & Lw.

Fam. ANCISTROCOMIDAE Ch. & Lw. Ancistrocoma Ch. & Lw. Hypocomella Ch. & Lw. ++ Anisocomides Ch. & Lw. Hypocomides Ch. & Lw. + Colligocineta Koz. Hypocomidium Raabe Crebricoma Koz. Hypocomina Ch. & Lw. Enerthecoma Jar. Ignotocoma Koz. Goniocoma Ch. & Lw. Insignicoma Koz. Heterocinetopsis Jar. Isocomides Ch. & Lw. Holocoma Ch. & Lw. Kozloffiella Raabe Hypocomagalma Jar. & Raabe Raabella Ch. & Lw. Hypocomatidium Jar. & Raabe Syringopharynx Coll.

Incert sed.: Cepedella Poyarkoff

Fam. SPHENOPHRYIDAE Ch. & Lw. Gargarius Ch. & Lw. Pelecyophrya Ch. & Lw. Lwoffia Koz. Sphenophrya Ch. & Lw. +




Order 6. APOSTOMATIDA Ch. & Lw. Suborder (1) Apostomatina Ch. & Lw.

Fam. FOETTINGERIIDAE Ch.

Calospira Ch. & Lw. Foettingeria Caul. & Mesnil Gymnodinioides Minkiewicz + Hyalophysa Bradbury Hyalospira Miyashita Kofoidella Cep. Metaphrya Ikeda Ophiuraespira Ch. & Lw. Perezella Cep. Pericaryon Ch.

Phoretophrya Ch., A. & M. Lw. Phtorophrya Ch., A. & M. Lw. + Polyspira Minkiewicz *Rosea de Puy. Spirophrya Ch. & Lw. Synophrya Ch. & Lw. Terebrospira Debaisieux Traumatiophtora Ch. & Lw. Vampyrophrya Ch. & Lw.

Incert. sed.: Jeppsia Corl.

Fam. *CYRTOCARYIDAE Corl.

Cyrtocaryum F-F & Mugard Incert. sed.: Spirobuetschliella Hovasse

(Remark: Is this family better placed in the following suborder?)

Suborder (2) Astomatophorina Jank.

Fam. OPALINOPSIDAE Hartog Chromidina Gonder Opalinopsis Foettinger

Fam. COLLINIIDAE Cep.

Collinia Cep. +

(Remark: Is this family better placed in the preceding suborder?)

Suborder (3) Pilisuctorina Jank.

Fam. CONIDOPHRYIDAE Kirby

Ascophrys Campillo & Deroux Askoella Fenchel

Conidophrys Ch. & Lw.

(Remark: Do the first two genera, above, deserve a separate family of their own?)

Subclass (4) Suctoria Clap. & Lach. Order SUCTORIDA Clap. & Lach.

Suborder (1) Exogenina Coll.

Fam. PODOPHRYIDAE Haeckel

Kystopus Jank. Lecanophrya Kahl Mucophrya Gaj.

Ophryocephalus Wailes Podophrya Ehr. +++ Sphaerophrya Clap. & Lach. +




Fam. PARAPODOPHRYIDAE Jank.

Parapodophrya Kahl +

Metacineta But. Paracineta Coll. ++

Actinocyathula Corl. Ephelota Wright ++ Metephelota Willis

Fam. URNULIDAE Fraipont Urnula Clap. & Lach.

Fam. EPHELOTIDAE Kent

Podocyathus Kent Tunicophrya Jank.

Fam. SPELAEOPHRYIDAE Jank. in Batisse

Spelaeophrya Stammer +

Fam. RHABDOPHRYIDAE Jank. Dendrosomides Coll. Rhabdophrya Ch. & Coll. +

Asterifer Jank.

Trophogemma Jank.

Fam. STYLOSTOMATIDAE Batisse

Stylostoma Milne

Fam. OPHRYODENDRIDAE St.

Loricodendron Jank. Ophryodendron Clap. & Lach. ++

Schizactinia Jank.

Fam. TACHYBLASTONIDAE Grell

Tachyblaston Martin

Fam. THECACINETIDAE Matthes

Canellana Jank. Praethecacineta Matthes Corynophrya Kahl + Thecacineta Coll. ++

Fam. PHALACROCLEPTIDAE Koz.

Phalacrocleptes Koz. (highly aberrant organism)

Suborder (2) Endogenina Coll.

Fam. ACINETIDAE St.

Acineta Ehr. +++ Acinetides Swarcz. Acinetopsis Robin + Arcosoma Jank. Cryptophrya Jank.

Cucumophrya Kunz Dactylophrya Coll. Dactylostoma Jank. Lecanodiscus Jank. Loricophrya Matthes




Multifasciculatum Good. & Jahn Plicophrya Jank. Pottsiocles Corl. Pseudogemma Coll. Pseudogemmides Kormos

Solenophrya Clap. & Lach. ++ Squalorophrya Good. & Jahn Suctorella Frenzel Tokophryella Jank. Trematosoma Batisse

Incert. sed.: Allantosoma Gass. + (perhaps not a suctorian?)

Fam. DENDROSOMATIDAE Fraipont Anarma Good. & Jahn Hypophrya L6p.-Och. Baikalodendron Swarcz. Lernaeophrya Perez Baikalophrya Swarcz. Rhyncheta Zenker Brachyosoma Batisse Staurophrya Zacharias Choanophrya Hartog Thaumatophrya Coll. Dendrosoma Ehr. Tokophrya But. +++ Erastophrya F-F Tokophryopsis Swarcz. Gorgonosoma Swarcz.

Fam. TRICHOPHRYIDAE Fraipont Actinobranchium Jank. Phagobranchium Jank. Asterophrya Awer. Tetraedrophrya Zykoff Capriniana Strand Trichophrya Clap. & Lach. ++ Marinecta Jank.

Fam. *ENDOSPHAERIDAE Jank. Endosphaera Engel. Physaliella Pen.

Suborder (3) *Evaginogenina Jank.

Fam. DISCOPHRYIDAE Coll. Caracatharina Kormos Peridiscophrya Nozawa Discophrya Lachmann + Prodiscophrya Kormos Echinophrya Swarcz. Rhynchophrya Coll. Periacineta Coll.

Fam. *HELIOPHRYIDAE Corl.

Cyclophrya Gonnert Platophrya Gonnert Heliophrya Saed. & Tell.

Fam. DENDROCOMETIDAE Haeckel Cometodendron Swarcz. ++ Dendrocometes St. + Dendrocometides Swarcz.

Discosomatella Corl. Stylocometes St. Stylophrya Swarcz.

Fam. CYATHODINIIDAE da C.

Cyathodinium da C. + (highly unusual organism)

(Remarks: Proper location of a number of suctorian families and genera remains in doubt; and some workers recognize a larger number of suborders than the three which I have endorsed above.)




Incertae sedis in class Kinetofragminophora:

Arachnidiopsis Pen. (perhaps not a ciliate?), Dysterioides Matthes, Euploia Lohmann, Silenella Fenchel

Class II. OLIGOHYMENOPHORA de Puy. et al.

Subclass (1) Hymenostomata Delage & Herouard

Order 1. HYMENOSTOMATIDA Delage & Herouard

Suborder (1) Tetrahymenina F-F in Corl.

Fam. TETRAHYMENIDAE Corl.

Colpidium St. + Stegochilum Schew. Deltopylum F-F & Mugard Tetrahymena Furgason + Lambornella Keilin (resurrected)

Incert. sed.: Blepharostoma Schew., Malacophrys Kahl

Fam. GLAUCOMIDAE Corl. Chasmatostoma Engel. *Glaucomella Groliere Dichilum Schew. Monochilum Schew. Epenardia Corl. Physalophrya Kahl Espejoia Burger Pseudoglaucoma Kahl Glaucoma Ehr. +

Incert. sed.: Dexiotrichides Kahl, Discozoon Vux., Pinchatia Shibuya, Pleuro- chilidium St.

Fam. CURIMOSTOMATIDAE Jank. Curimostoma Koz. Dogielella Polj.

Incertae sedis in suborder Tetrahymenina: Fam. SAGITTARIIDAE R. & L. Grandori, with Sagittaria R. & L. Grandori

Suborder (2) Ophryoglenina Canella

Fam. OPHRYOGLENIDAE Kent

Ophryoglena Ehr. ++ Protophryoglena Mugard

(Remark: Is Protophryoglena an independent genus?)

Fam. ICHTHYOPHTHIRIIDAE Kent

Cryptocaryon Brown Ichthyophthirius Fouquet Ichthyophthirioides Roque & de Puy.

(Remark: Is Ichthyophthirioides only a synonym of Ichthyophthirius?)

Suborder (3) Peniculina F-F in Corl.

Fam. PARAMECIIDAE Duj. Paramecium O.F.M. ++

Incert. sed.: Physanter Jank. (for Faurella)




Fam. FRONTONIIDAE Kahl Disematostoma Laut. + Frontoniella Wetzel Frontonia Ehr. ++ Wenrichia Jank.

Incert. sed.: Schistophrya Kahl, Sigmostomum Gulati

Fam. CLATHROSTOMATIDAE Kahl Clathrostoma Pen. +

Fam. UROCENTRIDAE Clap. & Lach. Urocentrum Nitzsch

Marituja Gaj. Parastokesia Jank.

Fam. STOKESIIDAE Roque Stokesia Wenrich

Fam. NEOBURSARIDIIDAE Drag. & Tuffrau Neobursaridium Balech

Fam. *LEMBADIONIDAE Jank. Lembadion Perty +

Fam. TURANIELLIDAE Didier Turaniella Corl.

Order 2. SCUTICOCILIATIDA Small

Suborder (1) Philasterina Small

Fam. PHILASTERIDAE Kahl Glauconema Thomp. Helicostoma Cohn Madsenia Kahl Miamiensis Thomp. & Moewus *Mugardia Small Paralembus Kahl +

Paranophrys Thomp. & Berger + Parauronema Thomp. Philaster Fab.-Dom. *Philasterella Evans Philasterides Kahl Potomacus Thomp.

Incert. sed.: Metanophrys de Puy., Groliere, Roque & Detcheva, Porpostoma Mobius

(Remarks: Some genera should very likely be removed from this hetero- geneous group; do Paralembus and Potomacus deserve separate families of their own?)

Homalogastra Kahl Pseuduronema Hoare * Urocyclon Small

Fam. URONEMATIDAE Thomp. Uronema Duj. +++ Uronemopsis Kahl Uropedalium Kahl

Fam. COHNILEMBIDAE Kahl Cohnilembus Kahl +




Fam. ENTORHIPIDIIDAE Madsen

Entorhipidium Lynch +

Cryptochilidium Entodiscus Made

Biggaria Kahl *Biggariella Proj Cryptochilum M *Metaxystomium

Plagiopyliella Po

Fam Schouteden

sen

i. ENTODISCIDAE Jank. Pectenita Jank.

Fam. *CRYPTOCHILIDAE Berger *Metoikos Berger & Thomp.

Fant *Tanystomium Berger aupas Thigmozoon Santh. & Nair Berger *Yagiua Profant

Fam. THYROPHYLACIDAE Berger in Corl. ?lj. Thyrophylax Berger in Corl.

Fam. LOXOCEPHALIDAE Jank. Dexiotricha Stokes + Loxocephalus Eberhard +

Paradexiotricha Groliere Paratetrahymena Thomp.

Incert. sed.: Cardiostomatella Corl.

Fam. CINETOCHILIDAE Perty Cinetochilum Perty + Pseudoplatynematum Bock Platynematum Kahl + Sathrophilus Corl. +

Incert. sed.: Colpodopsis Gr. & Roes.

Fam. UROZONIDAE Groliere Urozona Schew.

Fam. PSEUDOCOHNILEMBIDAE Evans & Thomp. Pseudocohnilembus Evans & Thomp. +

Incert. sed.: Anophrys Cohn

Fam. THIGMOPHRYIDAE Ch. & Lw. Cochliodomus Raabe Myxophyllum Raabe Cochliophilus Koz. Thigmophrya Ch. & Lw. + Conchophyllum Raabe

Incertae sedis in suborder Philasterina: Andreula Kahl, Aristerostoma Kahl, Balanonema Kahl, Bizonula Corl., Cryptostomina Fedele, Eurychilum Andre, Gullmarella Fenchel, Lem- badionella Kahl, Protocruzia de Faria et al., Ptyssostoma Hentschel, *Pusilloburius Corl., Rhinodisculus Mansfeld, Sertumia Tuc.

Suborder (2) Pleuronematina F-F in Corl.

Fam. PLEURONEMATIDAE Kent *Hippocomos Czapik & Jordan Pleuronema Duj. ++ Pleurocoptes Wallen. Schizocalyptra Drag.




*Abathostoma Berger Calyptotricha Phillips Ctedoctema Stokes

Fam. CYCLIDIIDAE Ehr.

Cyclidium O.F.M. +++ *Mesogymnus Berger

Incert. sed.: Cristigera Roux ++

Fam. *HISTIOBALANTIIDAE de Puy. & Corl.

Histiobalantium Stokes

Fam. PENICULISTOMATIDAE Fenchel

*Echinosociella Berger *Mytilophilus Antipa & Small

Peniculistoma Jank.

Fam. THIGMOCOMIDAE Kazubski

Thigmocoma Kazubski

Fam. CONCHOPHTHIRIDAE Kahl in Doflein & Reich.

Conchophthirus St. ++ Conchoscutum Raabe

Incertae sedis in suborder Pleuronematina:

Larvulina Pen.

Suborder (3) Thigmotrichina Ch. & Lw.

Fam. ANCISTRIDAE Issel

Ancistrella Cheissin Ancistrum Maupas Ancistrumina Raabe ++ Boveria Stevens Dragescoa Jank. Eupoterion MacL. & Conn.

Fenchelia Raabe Proboveria Ch. & Lw. Protophrya Kofoid Protophryopsis Raabe Semiboveria Raabe

Nomen inquirendum: Isselina Cep.

Fam. HEMISPEIRIDAE Konig

Ancistrospira Ch. & Lw. Cheissinia Ch. & Lw. Hemispeira Fab.-Dom.

Plagiospira Issel Protospira Raabe

Fam. HYSTEROCINETIDAE Diesing

Coelothigma de Puy. Cotylothigma Raabe Craticuloscuta Koz. Drilocineta Raabe Elliptothigma Meier Epicharocotyle Koz. Hysterocineta Diesing + Kozloffia de Puy.

Kysthothigma Raabe Preptychostomum de Puy. Protoptychostomum Raabe Ptychostomum St. ++ Puytoracia Raabe Taeniocineta Raabe Thurstonia de Puy.




Fam. PROTANOPLOPHRYIDAE Miyashita Protanoplophrya Miyashita (syn. Protoanoplophrya)

Fam. NUCLEOCORBULIDAE Santh. & Nair

Nucleocorbula Santh. & Nair

Incertae sedis in suborder Thigmotrichina:

*Syndaetor Berger (deserves family of its own?) Incertae sedis in order Scuticociliatida:

Agigea Lep?i (actually impossible to identify?)

Order 3. ASTOMATIDA Schew.

Fam. *ARCHIASTOMATIDAE Foiss. *Archiastomata Foiss. (a free-living form!)

Almophrya de Puy. Anoplophrya St. +d Corlissiella de Puy. Herpetophrya Siedl< Lomiella de Puy. Lubetiella de Puy.

Anoplophryopsis de Buetschliella Awer.

Akidodes Lom Anglasia Delphy Buetschliellopsis de Delphyella de Puy. Hoplitophrya St. +- Jirovecella Lom

Fam. ANOPLOPHRYIDAE Cep. & Drag. Metastomum Georg.

F+ Orchitophrya Cep. Paranoplophrya Rohrbach

ecki Perseia Rossolimo Prototravassosia Art. & Unti Sigmophrya de Puy.

Fam. BUETSCHLIELLIDAE de Puy. Puy. Herpinella de Puy. + Rhizocaryum Caul. & Mesnil

Fam. HOPLITOPHRYIDAE Cheissin

Juxtamesnilella de Puy. + Juxtaradiophrya de Puy. +

Puy. + Mesnilella Cep. Mixtophrya de Puy.

+ Protoradiophryopsis Georg. Radiophryoides Lom

Fam. RADIOPHRYIDAE de Puy. Acanthodiophrya de Puy. & Drag. Anthonyella Delphy Cheissinophrya de Puy. & Drag. Coelophrya de Puy. & Drag. Desmophrya Raabe Dicoelophrya de Puy. & Drag. Durchoniella de Puy. Eudrilophrya de Puy. Helella Kaczanowski

Hovasseiella de Puy. Metaracoelophrya de Puy. & Drag. Metaradiophrya Held. ++ Mimophrya de Puy. Mrazekiella Kijenskij Ochridanus Georg. Paracoelophrya de Puy. Radiophrya Rossolimo ++ Radiophryopsis Georg.

Fam. CONTOPHRYIDAE de Puy. Contophrya de Puy. & Drag. Dicontophrya de Puy. & Drag.




Acanthophrya Heid. Buchneriella Heid.

Intoshellina Cep. +

Fam. MAUPASELLIDAE Cep.

Georgevitchiella de Puy. Maupasella Cep. ++

Fam. INTOSHELLINIDAE Cep.

Monodontophrya Vejdovsky

Fam. HAPTOPHRYIDAE Cep.

Annelophrya Lom Cepedietta Kay + (in amphibians) Haptophrya St. + (in turbellarians)

Lachmannella Cep. Steinella Cep.

Fam. *CLAUSILOCOLIDAE de Puy.

Clausilocola Lom Haptophryopsis de Puy.

Proclausilocola Lom

(Remark: Do the two preceding families deserve separation at a higher level from the other astome groups?)

Subclass (2) Peritricha St.

Order PERITRICHIDA St.

Suborder (1) Sessilina Kahl

Fam. VORTICELLIDAE Ehr.

Carchesium Ehr. ++ Entziella Stiller Haplocaulus Precht ++ Intranstylum F-F + Myoschiston Precht + *Parazoothamnium Piesik

Pseudocarchesium Sommer + *Rugaecaulis Lom *Tucolesca Lom Vorticella L. ++++ Zoothamnium Bory +++

Incert. sed.: Craspedomyoschiston Precht, *Monintranstylum Banina, Pseudo- vorticella Foiss. & Schiffmann

Fam. ASTYLOZOIDAE Kahl

Astylozoon Engel. + (syn. Geleiella) Hastatella Erlanger

Fam. EPISTYLIDIDAE Kahl

Apiosoma Blanchard +++ Campanella Goldfuss Epistylis Ehr. ++++

Opisthostyla Stokes + Rhabdostyla Kent +++

Incert. sed.: Systylis Bresslau

Fam. *OPERCULARIIDAE F-F

Ballodora Dog. & Furss. Opercularia Goldfuss +++ Operculariella Stammer

Incert. sed.: Orsomia Baer

Orbopercularia Lust ++ *Propyxidium Corl. ++ (for Pyxidiella

[for Pyxidium])




Opisthonecta F-F

Fam. OPISTHONECTIDAE Foiss.

Telotrochidium Kent

Fam. SCYPHIDIIDAE Kahl

Ambiphrya Raabe Paravorticella Kahl Gonzeella Kufferath Scyphidia Duj. ++ Ophrydiopsis Pen.

Incert. sed.: Pachystomos Rudzinska

Fam. OPHRYDIIDAE Ehr.

Gerda Clap. & Lach. (resurrected) Ophrydium Bory

Fam. ELLOBIOPHRYIDAE Ch. & Lw.

Caliperia Laird Clausophrya Naid. & Zaika

Ellobiophrya Ch. & Lw.

Fam. *TERMITOPHRYIDAE Lom

Termitophrya Noirot & N-T

Fam. ROVINJELLIDAE Matthes

Rovinjella Matthes

Fam. VAGINICOLIDAE de Fromentel

Caulicola Stokes Cothurnia Ehr. ++++ Cothurniopsis Entz + Pachytrocha Kent Platycola Kent ++

Incert. sed.: Cyclodonta Matthes

Pseudothuricola Kahl Pyxicola Kent + Thuricola Kent + Vaginicola Lamarck +++

Fam. LAGENOPHRYIDAE But.

Lagenophrys St. +++ Operculigera Kane

Incert. sed.: *Eilymophrys Corl.

Stylohedra Kellicott

Suborder (2) Mobilina Kahl

Fam. *URCEOLARIELLIDAE Corl.

*Urceolariella Corl. + (for Urceolaria)

Fam. LEIOTROCHIDAE Johnston

Leiotrocha Fab.-Dom.

Fam. POLYCYCLIDAE Polj.

Polycycla Polj.




Fam. TRICHODINOPSIDAE Kent

Trichodinopsis Clap. & Lach.

Fam. TRICHODINIDAE Claus

Dipartiella G. Stein Paratrichodina Lom Semitrichodina Kazubski + Trichodina Ehr. ++++

Trichodinella Sr.-Husek Tripartiella Lom Vauchomia Mueller

Incert. sed.: Acyclochaeta Zick

(Remark: A couple of the preceding families may need to be combined.)

Class III. POLYHYMENOPHORA Jank. Subclass Spirotricha But.

Order 1. HETEROTRICHIDA St.

Suborder (1) Heterotrichina St.

Fam. SPIROSTOMIDAE St.

Anigsteinia Isquith + Blepharisma Perty ++ Gruberia Kahl

Parablepharisma Kahl Pseudoblepharisma Kahl Spirostomum Ehr. ++

Incert. sed.: Diplogmus Mansfeld, Propygocirrus Mansfeld (these two genera not to be confused with the scuticociliate Protocruzia?)

Fam. *PHACODINIIDAE Corl.

Phacodinium Prowazek

Fam. METOPIDAE Kahl

Bothrostoma Stokes Brachonella Jank. + Metopus Clap. & Lach. +++ Palmarella Jank. (for Palmarium)

Spirorhynchus da C. Tesnospira Jank. Tropidoatractus Levander

Fam. CONDYLOSTOMATIDAE Kahl in Doflein & Reich.

Bryometopus Kahl Condylostoma Bory ++

Incert. sed.: Dellochus Corl.

Balantidioides Kahl

Climacostomum St. Fabrea Henneguy

Copemetopus Vill.-Brach.

Fam. REICHENOWELLIDAE Kahl

Reichenowella Kahl

Fam. CLIMACOSTOMIDAE Repak Pediostomum Kahl

Transitella Gellert




Fam. STENTORIDAE Carus

Stentor Oken ++ Incert. sed.: Parastentor Vux., Stentoropsis Dogiel & Bychowsky (belongs in

following suborder?)

Fam. BURSARIIDAE Duj. Bursaria O.F.M. Bursaridium Laut.

Thylakidium Schew. (syn. Thylacidium)

Fam. CHATTONIDIIDAE Vill.-Brach.

Chattonidium Villeneuve

Fam. PERITROMIDAE St.

Peritromus St. +

Suborder (2) Clevelandellina de Puy. & Grain

Fam. NYCTOTHERIDAE Amaro

Metanyctotherus Alb. Nyctotheroides Grasse +++

Geimania Alb. Ichthyonyctus Jank. + Metasicuophora Alb.

Nyctotherus Leidy +++ Pronyctotherus Alb. & Njine

Fam. SICUOPHORIDAE Amaro

Parasicuophora Alb. Prosicuophora de Puy. & Oktem Sicuophora de Puy. & Grain +

Fam. CLEVELANDELLIDAE Kidder

Clevelandella Kidder ++ Metaclevelandella Utt. & Desai

Paraclevelandia Kidder

Fam. INFEROSTOMATIDAE Ky

Inferostoma Ky Incertae sedis in suborder Clevelandellina:

Fam. NATHELLIDAE Singh, with Nathella Singh

Suborder (3) Armophorina Jank.

Fam. CAENOMORPHIDAE Poche

Caenomorpha Perty ++ Ludio Pen. Cirranter Jank.

Suborder (4) Coliphorina Jank.

Fam. FOLLICULINIDAE Dons

Ampullofolliculina Hadzi Atriofolliculina Hadzi Ascobius Henneguy + Aulofolliculina Hadzi




Botticula Dioni Claustrofolliculina Hadzi Diafolliculina Hadzi Donsia Hadzi Echinofolliculina Dons Epifolliculina Hadzi Eufolliculina Hadzi Folliculina Lamarck Folliculinopsis F-F Halofolliculina Hadzi Lagotia Wright ++ Latifolliculina Hadzi Magnifolliculina Uhlig Metafolliculina Dons + Mirofolliculina Dons

Pachyfolliculina Hadzi Parafolliculina Dons + Pebrilla Giard Pedifolliculina Hadzi Perifolliculina Hadzi Planifolliculina Hadzi Patyfolliculina Hadzi Priscofolliculina Defl. & Deunff + Pseudofolliculina Dons Splitofolliculina Hadzi Stentofolliculina Hadzi Tapetofolliculina Andrews & Nelson in

Hadzi Valletofolliculina Andrews

Suborder (5) Plagiotomina Alb.

Fam. PLAGIOTOMIDAE But.

Plagiotoma Duj.

Suborder (6) Licnophorina Corl.

Fam. LICNOPHORIDAE But.

Licnophora Claparede +

Order 2. ODONTOSTOMATIDA Sawaya

Fam. EPALXELLIDAE Corl.

Epalxella Corl. + (for Epalxis) Pelodinium Laut.

Saprodinium Laut. +

Fam. DISCOMORPHELLIDAE Corl.

Discomorphella Corl. (for Discomorpha)

Fam. MYLESTOMATIDAE Kahl in Doflein & Reich.

Atopodinium Kahl Mylestoma Kahl +

Order 3. OLIGOTRICHIDA But.

Suborder (1) Oligotrichina But.

Fam. HALTERIIDAE Clap. & Lach.

Cephalotrichium Meunier Halteria Duj. + Halterioforma Horvath

Meseres Schew. Metastrombidium F-F Parastrombidium F-F

Insert. sed.: Jeannellia Tuc., Octocirrus Rao, Spelaeonecta Jank.




Buehringa Busch Laboea Lohmann

Fam. STROMBIDIIDAE F-F

Strombidium Clap. & Lach. +++ Tontonia F-F

Fam. STROBILIDIIDAE Kahl in Doflein & Reich.

*Patronella Corl. (for Torquatella) Strobilidium Schew. ++

Strombidinopsis Kent

Incert. sed.: Ciliospina Leegaard, Lohmanniella Leegaard, Sphaerotrichium Wulff

Suborder (2) Tintinnina K. & C.

Fam. TINTINNIDIIDAE K. & C.

Leprotintinnus Jorg. +

Fam. CODONELLIDAE Kent

Amphorellina Colom Chitinoidella Doben ++ Claretinella Keij Codonaria K. & C. + Codonella Haeckel +++ Codonopsis K. & C. Coxliellina Colom + Crassicollaria Remane + Dicloeopella Eicher Durandella Dragastan Lorenziella Knauer & Nagy Parachitinoidella Trejo Poroecus Cleve

Praetintinnopsella Borza Pseudarcella Spandel Remanellina Tapp. & Loeb. Remaniella Catalano Salpingellina Colom + Spinarcella Keij Spinophenia Szczechura Tintinnopsella Colom ++ Tintinnopsis St. ++++ Tytthocorys Tapp. & Loeb. Urnulella Szczechura Vautrinella Cuvil. & Sacal Yvonniellina Tapp. & Loeb.

Fam. CODONELLOPSIDAE K. & C.

Calpionella Lorenz + Calpionellites Colom + Calpionellopsella Trejo Calpionellopsis Colom Codonellopsis Jorg. +++ Colomiella Bonet

Deflandronella Trejo Laackmanniella K. & C. Praecalpionellopsis Borza Stenosemella Jorg. ++ Stenosemellopsis Colom +

Incert. sed.: Micropoculum Dons

Fam. COXLIELLIDAE K. & C.

Climacocylis Jorg. + Coxliella Brandt ++ Favelloides Colom Helicostomella Jorg. + Metacylis Jorg.

Incert. sed.: Stylicauda Balech

Pseudometacylis Balech Rhabdonelloides Colom + Rhizodomus Strel. & Virk. Spiroxystonellites Knauer

Tintinnidium Kent ++




Fam. EPIPLOCYLIDIDAE K. & C.

Epicancella K. & C. Epiplocylis Jorg. ++

Epiplocyloides Hada +

Fam. CYTTAROCYLIDIDAE K. & C.

Cyttarocylis Fol ++

Fam. PETALOTRICHIDAE K. & C.

Acanthostomella Jorg. + Niemarshallia Corl. Ascampbelliella Corl. + Parapetalotricha Hada Luxiella Lecal Petalotricha Kent +

Incert. sed.: Wailesia K. & C.

Fam. DICTYOCYSTIDAE Haeckel

Dictyocysta Ehr. +++ Luminella K. & C.

Wangiella Nie

Fam. PTYCHOCYLIDIDAE K. & C.

Cymatocylis Laackmann +++ Favella Jorg. ++

Protocymatocylis K. & C. Ptychocylis Brandt ++

Fam. RHABDONELLIDAE K. & C.

Epirhabdonella K. & C. Protohabdonella Jorg. +

Fam. XYSTONELLIDAE K. & C.

Parafavella K. & C. ++ Parafavelloides Defl. & Defl.-Rigaud Parundella Jorg. ++

Xystonella Brandt + Xystonellopsis Jorg. +++

Fam. UNDELLIDAE K. & C.

Amplectella K. & C. + Proplectella K. & C. ++ Amplectellopsis K. & C. Undella Daday ++ Cricundella K. & C. Undellopsis K. & C. ++ Micrundella Busch

Incert. sed.: Rotundocylis Kufferath

Fam. TINTINNIDAE Clap. & Lach.

Albatrossiella K. & C. Amphorellopsis K. & C. + Amphorides Strand + (for Amphorella) Brandtiella K. & C. Bursaopsis K. & C. + Buschiella Corl. Canthariella K. & C. Clevea Balech

Dadayiella K. & C. + Daturella K. & C. ++ Epicranella K. & C. + Eutintinnus K. & C. +++ Odontophorella K. & C. Ormosella K. & C. + Proamphorella K. & C. Prostelidiella K. & C.

Rhabdonella Brandt ++ Rhabdonellopsis K. & C. +




Rhabdosella K. & C. Steenstrupiella K. & C. + Salpingacantha K. & C. + Stelidiella K. & C. Salpingella Jorg. ++ Tintinnus Schrank + Salpingelloides Campbell

Incertae sedis in suborder Tintinnina (some are particularly dubious as ciliates): Bonetocardiella Dufour, Cadosina Wanner ++, Cadosinopsis Scheibner, Campbelliella Rad., Carpistomiosphaera Nowak, Colomis- phaera Nowak, Daturellina Rad. +, Hadziina Rad., Hemistomiosphaera Nowak, Metacyclina Rad. +, Parastomiosphaera Nowak, Patelloides Leischner, Stomiosphaera Wanner ++, Zetella Rad.

Order 4. HYPOTRICHIDA St.

Suborder (1) Stichotrichina F-F

Fam. SPIROFILIDAE Gelei

Atractos Voros. Hypotrichidium Ilowaisky + Kahliella Corl. + Plesiotricha Drag.

Spiretella Borror Spirofilopsis Corl. Stichotricha Perty + Urospinula Corl.

Incert. sed.: Kahliela Tuc.

Fam. STRONGYLIDIIDAE F-F

Chaetospira Lachmann + Cladotricha Gaj. + Klonostricha Vux.

Banyulsella Drag. Hemicycliostyla Stokes Isosticha Kiessel.

Amphisiella Gr. & Roes. ++ *Bakuella Agam. & Alek. Balladyna Kowalewski Balladynella Stiller Gonostomum Sterki Holosticha Wrzes. ++ Keronopsis Pen. ++ Laurentiella Drag. & Njine Paruroleptus Kahl + Parurosoma Gelei

Strongylidium Sterki + Urostrongylum Kahl

Fam. UROSTYLIDAE But.

Paraholosticha Kahl + Paraurostyla Borror Urostyla Ehr. ++

Fam. HOLOSTICHIDAE F-F

Psammomitra Borror Pseudourostyla Borror Trachelochaeta Sr.-Husek Trachelostyla Kahl + Trichotaxis Stokes + Uncinata Bullington Uroleptoides Wenzel Uroleptopsis Kahl + Uroleptus Ehr. + *Wallackia Foiss.

(Remarks: Are all of the above genera truly confamilial? Should Uroleptus [and close relatives] especially be removed?)

Fam. PSILOTRICHIDAE But.

Hemiholosticha Gelei Psilotricha St.




Caryotricha Kahl

Epiclintes St. Eschaneustyla Stokes +

Fam. KIITRICHIDAE Nozawa Kiitricha Nozawa

Fam. KERONIDAE Duj. Kerona Ehr. Lacazea Drag.

Incertae sedis in suborder Stichotrichina:

Balladinopsis Ghosh, Prooxytricha Poche, Psilotrix Gr. & Roes., Sty- lonethes Sterki

Suborder (2) Sporadotrichina F-F

Actinotricha Cohn Ancystropodium F-F Gastrostyla Engel. + Histriculus Corl. + Onychodromus St. Opisthotricha Kent ++ Oxytricha Bory +++

Fam. OXYTRICHIDAE Ehr. Parahistriculus Groliere Parastylonychia Drag. Pleurotricha St. + Steinia Diesing ++ Stylonychia Ehr. ++ Tachysoma Stokes + Urosoma Kowalewski +

Fam. ASPIDISCIDAE Ehr.

Aspidisca Ehr. +++ Onychaspis St. Euplotaspis Ch. & Seguela

Incert. sed.: Paraeuplotes Wichterman

Fam. EUPLOTIDAE Ehr.

Certesia Fab.-Dom. Cytharoides Tuffrau Diophrys Duj. ++

Euplotes Ehr. +++ Swedmarkia Drag. Uronychia St. +

Incert. sed.: Discocephalus Ehr. in Hemprich & Ehr. +

Fam. GASTROCIRRHIDAE F-F

Cirrhogaster Ozaki & Yagiu Gastrocirrhus Lep?i Euplotidium Noland

Incertae sedis in suborder Sporadotrichina: Allotricha Sterki, Cinetoconia Renault & Roche, Gruberella Corl., Ony- chodromopsis Stokes

(Remark: The proper distributional assignment of hypotrich genera to one or the other of the two suborders accepted here remains controversial in a number of cases.)

LITERATURE CITED7

BICK, H. 1972. Ciliated Protozoa: An Illustrated Guide to the Species Used as Biological Indicators in Freshwater Biology. World Health Organization, Geneva. 198 pp.

7The number of citations is purposely kept low here, since many of the hundreds of pertinent works have already been cited fully in the group of papers by me which are given below. Furthermore, well over 2,650 references will be painstakingly included in the bibliog- raphy of my forthcoming book, a work to appear, hopefully, before the end of the current year.



CORLISS FAMILIES AND GENERA OF CILIATES

BORROR, A. C. 1973. Marine flora and fauna of the northeastern United States. Protozoa:

Ciliophora. NOAA Tech. Rep. NMFS CIRC., No. 378. 62 pp. CAIRNS, J., JR., ED. 1971. The Structure and Function of Fresh-Water Microbial Communi-

ties. Virginia Polytechnic Institute and State University Press, Blacksburg, Virginia. 301 pp.

CANELLA, M. F. & ROCCHI-CANELLA, I. 1976. Biologie des Ophryoglenina (cilies hymeno- stomes histophages). Ann. Univ. Ferrara (N. S., Sect. III), 3(Suppl. 2): 1-510.

CORLISS, J. 0. 1961. The Ciliated Protozoa: Characterization, Classification, and Guide to the Literature. Pergamon Press, London and New York. 310 pp.

1962. Taxonomic-nomenclatural practices in protozoology and the new International Code of Zoological Nomenclature. J. Protozool., 9: 307-324.

1968. The value of ontogenetic data in reconstructing protozoan phylogenies. Trans. Amer. Micros. Soc., 87: 1-20.

1972. Common sense and courtesy in nomenclatural taxonomy. Syst. Zool., 21: 117-122. 1973a. Protozoan ecology: a note on its current status. Amer. Zool., 13: 145-148. 1973b. Evolutionary trends in patterns of stomatogenesis in the ciliate Protozoa. (Abstr.)

J. Protozool., 20: 506. 1974a. The changing world of ciliate systematics: historical analysis of past efforts and a

newly proposed phylogenetic scheme of classification for the protistan phylum Cili- ophora. Syst. Zool., 23: 91-138.

1974b. Remarks on the composition of the large ciliate class Kinetofragmophora de Puy- torac et al., 1974, and recognition of several new taxa therein, with emphasis on the primitive order Primociliatida n. ord. J. Protozool., 21: 207-220.

1975a. Taxonomic characterization of the suprafamilial groups in a revision of recently proposed schemes of classification for the phylum Ciliophora. Trans. Amer. Micros. Soc., 94: 224-267.

1975b. Nuclear characteristics and phylogeny in the protistan phylum Ciliophora. Bio- Systems, 7: 338-349.

1976. On lumpers and splitters of higher taxa in ciliate systematics. Trans. Amer. Micros. Soc., 95: 430-442.

CORLISS, J. 0. & HARTWIG, E. 1977. The "primitive" interstitial ciliates: their ecology, nuclear uniquenesses, and postulated place in the evolution and systematics of the phylum Ciliophora. Akad. Wiss. Lit. (Mainz) Math.-Naturwiss. Kl. Mikrofauna Meeresbodens, 61. (in press)

CURDS, C. R. 1969. An Illustrated Key to the British Freshwater Ciliated Protozoa Com- monly Found in Activated Sludge. Ministry of Technology, H.M.S.O., London. 90 pp.

DRAGESCO, J. 1974. Ecologie des protistes marins. In Puytorac, P. de & Grain, J., eds., Actualites Protozoologiques, Vol. 1, University of Clermont, France, pp. 219-228.

FAURE-FREMIET, E. 1950. Morphologie comparee et systematique des cilies. Bull. Soc. Zool. Fr., 75: 109-122.

FENCHEL, T. 1969. The ecology of marine microbenthos. IV. Structure and function of the benthic ecosystem, its chemical and physical factors and the microfauna communities with special reference to the ciliated protozoa. Ophelia, 6: 1-182.

FINLEY, H. E., BICK, H. & BROWN, E. M. 1974. Ecology of fresh-water protists. In Puy- torac, P. de & Grain, J., eds., Actualites Protozoologiques, Vol. 1, University of Cler- mont, France, pp. 229-237.

FRANKEL, J. 1974. Positional information in unicellular organisms. J. Theoret. Biol., 47: 439-481.

GRAIN, J. 1969. Le cinetosome et ses derives chez les cilies. Ann. Biol., 8: 53-97. GRAIN, J., PUYTORAC, P. DE & BOHATIER, J. 1973. Essai de systematique des cilies gymno-

stomes fondee sur les caracteristiques de l'infraciliature circumorale. Compt. Rend. Acad. Sci., 227: 69-72.

GRELL, K. G. 1973. Protozoology. Springer-Verlag, Berlin, Heidelberg, New York. 554 pp.

139




HONIGBERG, B. M., BALAMUTH, W., BOVEE, E. C., CORLISS, J. O., GOJDICS, M., HALL, R. P., KuDo, R. R., LEVINE, N. D., LOEBLICH, A. R., JR., WEISER, J. & WENRICH, D. H. 1964. A revised classification of the phylum Protozoa. J. Protozool., 11: 7-20.

HUTNER, S. H. & CORLISS, J. 0. 1976. Search for clues to the evolutionary meaning of ciliate phylogeny. J. Protozool., 23: 48-56.

JANKOWSKI, A. W. 1967. [A new system of ciliate Protozoa (Ciliophora).] Akad. Nauk SSSR, Trudy Zool. Inst., 43: 3-54. (in Russian)

1972. Recapitulation of phylogenesis in ciliate ontogeny. In Vorontsov, N. N., ed., Prob- lems of Evolution, Vol. 2, Akad. Nauk SSSR, Nauka, Novosibirsk, pp. 95-123. (in Russian with English summary)

1973a. Taxonomic revision of subphylum Ciliophora Doflein, 1901. Zool. Zh., 52: 165- 175. (in Russian with English summary)

1973b. [Fauna of the USSR: Infusoria Subclass Chonotricha.] Vol. 2, No. 1. Akad. Nauk SSSR, Nauka, Leningrad. 355 pp. (in Russian)

1975. [A conspectus of the new system of subphylum Ciliophora Doflein, 1901.] (Abstr.) In Balashov, U. S., ed. [Account of Scientific Sessions on Results of Scientific Work, Year 1974: Abstracts of Reports], Akad. Nauk SSSR, Zool. Inst. Leningrad, pp. 26-27. (in Russian)

JONES, A. R. 1974. The Ciliates. Hutchinson, London. 207 pp. KUDO, R. R. 1966. Protozoology. 5th ed. Charles C Thomas, Springfield, Illinois. 1174 pp. LEPSI, J. 1965. Protozoologie. Academia Republicii Socialiste Romania, Bucarest. 999 pp.

(in Roumanian) LEVINE, N. D. 1972. Relationships between certain protozoa and other animals. In Chen,

T.-T., ed., Research in Protozoology, Vol. 4, Pergamon Press, London and New York, pp. 291-350.

MANWELL, R. D. 1968. Introduction to Protozoology. 2nd ed. Dover, New York. 642 pp. PITELKA, D. R. 1969. Fibrillar systems in protozoa. In Chen, T.-T., ed., Research in Proto-

zoology, Vol. 3, Pergamon Press, London and New York, pp. 279-388.

POLJANSKY, G. I. & CHEISSIN, E. M. 1965. Dogiel's General Protozoology. Oxford Univer-

sity Press, London and New York. 747 pp. PUYTORAC, P. DE, BATISSE, A., BOHATIER, J., CORLISS, J. O., DEROUX, G., DIDIER, P., DRAGESCO,

J., FRYD-VERSAVEL, G., GRAIN, J., GROLIERE, C.-A., HOVASSE, R., IFTODE, F., LAVAL, M., ROQUE, M., SAVOIE, A. & TUFFRAU, M. 1974. Proposition d'une classification du

phylum Ciliophora Doflein, 1901 (reunion de systematique, Clermont-Ferrand). Compt. Rend. Acad. Sci., 278: 2799-2802.

PUYTORAC, P. DE & GRAIN, J. 1976. Ultrastructure du cortex buccal et evolution chez les cilies. Protistologica, 12: 49-67.

RAABE, Z. 1964. Zarys Protozoologii. Polish Scientific Publs., Warsaw. 283 pp. (in Polish) RAIKOV, I. B. 1969. The macronucleus of ciliates. In Chen, T.-T., ed., Research in Proto-

zoology, Vol. 3, Pergamon Press, London and New York, pp. 1-128.

SLEIGH, M. A. 1973. The Biology of Protozoa. Edward Arnold, London. 315 pp. SMALL, E. B. 1976. A proposed subphyletic division of the phylum Ciliophora Doflein,

1901. Trans. Amer. Micros. Soc., 95: 739-751. SONNEBORN, T. M. 1974. Ciliate morphogenesis and its bearing on general cellular morpho-

genesis. In Puytorac, P. de & Grain, J., eds., Actualites Protozoologiques, Vol. 1, Univer- sity of Clermont, France, pp. 327-355.

TUFFRAU, M., LOM, J. & HALLER, G. DE. 1974. La stomatogenese chez les cilies. In Puy- torac, P. de & Grain, J., eds., Actualities Protozoologiques, Vol. 1, University of Cler- mont, France, pp. 279-292.

WESTPHAL, A. 1974. Protozoen. Eugen Ulmer, Stuttgart. 244 pp.



Documents

Annotated Assignment of Families and Genera to the Orders and Classes Currently Comprising the Corlissian Scheme of Higher Classification for the Phylum Ciliophora