Remarks on the Composition of the Large Ciliate Class Kinetofragmophora de Puytorac et al., 1974, and Recognition of Several New Taxa Therein, with Emphasis on the Primitive Order

J. PROTOZOOL. 21(2), 207-220 (1974). 207

Remarks on the Composition of the Large Ciliate Class Kinetofragmophora de Puytorac et al., 1974, and Recognition of Several New Taxa Therein, with

Emphasis on the Primitive Order Primociliatida N. Ord.* JOHN 0. CORLISS

Department of Zoology, University of Maryland, College Park, Maryland 20742 U S A .

SYNOPSIS. With the realization that new data (especially ultrastructural) and new ideas are making necessary a major revision of the scheme of classification of the Ciliophora, several groups of ciliatologists are preparing treatises on the subject. The present paper is concerned with the composition of the large new class of ciliates, Kinetofragmophora de Puy- torac et al., 1974, established very recently by the French group. Several new taxa, at ordinal and subordinal levels, are proposed for inclusion in that class, with special emphasis on the new order to contain the most primitive of extant species. Actions taken here are incorporated in a major review and revisory work of the author which is being published elsewhere.

The class Kinetofragmophora, by far the largest of the 3 classes now recognized as comprising the whole phylum Ciliophora, is itself considered to contain 4 sizeable subclasses and to embrace a total of 13 orders and 14 suborders. Twoorders and 6 suborders are named and described here as new, enumerated and briefly identified as follows: Order Primociliatida n. ord., for the most “primitive” of gymnostomes, with three new suborders-Homokaryotina n. subord., for the homokaryotic genus Stephanopogon; Karyorelictina n. subord., for a number of mostly interstitial ciliates which, though heterokaryotic, possess nondividing, diploid macronuclei (e.g. Trachelocerca, Trachelonema, and Tracheloraphis) ; and Prorodontina n. subord., for a group of relatively specialized formerly “rhabdophorine” gymnostomes such as Coleps, Placus, and Prorodon and order Haptorida n. ord., for rapacious carnivorous forms, formerly lumped with the preceding groups as “rhabdophorines,” many with oral toxicysts and well developed thigmotactic ciliature (e.g. Actinobolina, Didinium, Dilep- tus, Enchelys, Spathidium, and Trachelius) . All foregoing taxa are members of the 1st kinetofragmophoran subclass, the Gymnostomata. In the taxonomic conclusions drawn, new significance is placed on ultrastructural data, on macronuclear differences of evolutionary importance, and on habitat and behavior. A brief review of the literature on psammophilous ciliates is presented.

In the subclass Vestibulifera is now located the order Entodiniomorphida Reichenow, a group formerly considered to be a spirotrich taxon. A suborder, Blepharoeorythina n. subord., is proposed to contain the old “trichostome” family Blepharo- corythidae, species commensalistic in horses and ruminants and now-with their syncilia, etc.-considered ancestral to the ophryoscolecids and relatives.

In the subclass Hypostomata, order Nassulida, the suborder Paranassulina n. subord. is established to contain nassulids which appear more highly evolved than Nassula itself (e.g. Paranassula and Enneameron) in perioral ciliature, mode of stomatogenesis, etc. In the enigmatic and still vexatious order Rhynchodida, the suborder Aneistroeomina n. subord. is erected to embrace rhynchodid genera with an anteriorly located sucking tentacle (and other unique characteristics)-for example, Ancistrocoma, Crebricoma, Holocoma, and Sphenophrya. With the banishment of the bulk of the old “thigmotrichs” to the oligohymenophoran order Scuticociliatida, the ancistrocomines are left with the family Hypocomidae (and relatives) in the order Rhynchodida. It is not yet clear, however, how closely related the 2 suborders of rhynchodids should be considered.

Index Key Words: Kinetofragmophora, class of ciliates; Primociliatida n. ord., with Homokaryotina n. subord., Karyo- relietina n. subord., Prorodontina n. subord.; Haptorida n. ord.; Blepharocorythina n. subord. ; Paranassulina n. subord. ; Aneistrocominn n. subord. ; systematics of ciliate Protozoa ; higher taxa ; taxonomic revision.

Special nomenclatural problems are also involved.

OR several years (e.g. see 37), there has been widespread F realization among ciliatologists that a rather drastic revision of the scheme of classification for the ciliate Protozoa must be made if we are to take into account the recent accumula- tion of impressive new data and the appearance of several new hypotheses published within the past decade on possible phylogenetic relationships among the major groups of these organisms. The classifications of FaurC-Fremiet (64), Cheissin & Poljansky (32) , Corliss (33, 36), Honigberg et al. (105), Poljansky & Cheissin (137), and Raabe (148, 149) are simply inadequate today, even apart from such criticisms as those launched by Canella (26, 27). The schemes of Jankowski (108, 110) also have not met or do not seem destined to meet with general acceptance, in spite of their considerable value (see below). Thus, in my opinion, nowhere is there available- in published form at the time of preparation of the present paper-an overall classification scheme to meet the modern needs of ciliate systematics.

* Support of National Science Foundation grant GB-27050-X3 is gratefully cited. I wish also to acknowledge the critical comments of numerous colleagues in America and Europe, with special thanks to Messrs Gilbert Deroux, Denis H. Lynn, and Eugene B. Small.

SOME OVERALL CONSIDERATIONS I t is fully appropriate and in the best interests of all con-

cerned to mention here that a t least 3 large works on the Ciliophora are currently approaching final stages of preparation, and that the authors involved have classification schemes pretty clearly in mind, if not in hand, at this date. These treatises are : the long-awaited “FaurC-Fremiet volume,” Tome 2, of the T r a i t i de Zoologie, now to be a multi-authored monograph under the directorship or supervision of the editor, P.-P. GrassC; the section on the ciliates being developed in extenso by Eugene B. Small and selected colleagues and students for inclusion in the long-planned Illustrated Guide to the S tudy of the Protozoa, a future publication of the Society of Protozoologists under the editorship of John J. Lee; and a 2nd edition of my own book (36) , T h e Ciliated Protozoa, a work now at least 13 years out of date. I t cannot be expected that there will be complete harmony among the schemes of higher- level ciliate classification eventually appearing in these 3 potentially authoritative publications. But, by correspondence, personal contact, and first-hand involvement, I can state that there is presently a “consensus” of agreement regarding not only the need for change but also the general kinds of changes

208 NEW TAXA IN CILIATE CLASS KINETOFRAGMOPHORA

envisioned and, often, even the exact form the changes themselves are to take.

Imminent though the appearance of one or more of the above-mentioned treatises may be, it seems reasonablein- deed, inevitable-to expect several “advance” publications, works which, although much shorter, may permit greater detail or explanation of specific taxonomic situations involving one or more of the major groups requiring ultimate revision. Also, space restrictions in the “final” works may prevent inclusion of material of value from an historical point of view. Then, too, book or monograph writers often prefer publishing new taxonomic names in advance of their overall, comprehensive treatments, so that the larger revisory work can be freed from such nomenclatural considerations.

Three papers-to be in press at approximately the same time-come to mind in illustration of the comments made above. One is the invitational historical review by this writer (40) in which also a new overall classification for the “phylum Ciliophora” is proposed; but space did not permit much detail in treatment of the new or newly rearranged major groups considered. Any “new” group names used there were officially and formally proposed or established first in the 2nd and 3rd papers to be mentioned, although they were in only manuscript form at the time of writing of the “overview” essay. One of the 2 revisory works is the preliminary note by de Puytorac et al. (“et al.” includes this writer; see 142) which proposes a number of new high-level names and signifies several important shifts of categories in the hierarchy, conclusions drawn principally from stimulating discussions held at an international meeting in Aubi6re (Clermont-Ferrand) in May 1973. The other is represented by the present effort, in May 1973. The other is represented by the present effort, in which some discussion is offered, in English, of certain of the decisions tentatively reached by the group meeting a year ago in France and in which some additional new taxa are described and named, especially for groups of ciliates comprising the 1st subclass of the new class Kinetofragmophora de Puytorac et al., 1974, groups left aside or “undecided,” for one reason or another, by the (predominantly) French group of ciliatologists.

TRIBUTE T O JANKOWSKI As also pointed out in the other 2 publications (40, 142)

paralleling this one (see above), a new arrangement-at the very least-of the higher ciliate taxa is required if we are to appreciate and utilize the wealth of new information, especially ultrastructural data, now available to ciliate systematists. Quite recently, Jankowski ( 108), feeling essentially the same about the overall situation but perhaps not as appreciative of the findings of electron microscopy’, proposed a novel scheme of classification, though it lost much of its uniqueness in his own very recent revision (1 10). General discussion of his stimulating and well documented approach to solution of the problems the existence of which we have all recognized for some time is well beyond the scope of the present taxonomic note. But it is important to give proper credit to Jankowski and to admit that some of his ideas, particularly in his 1967 master- piece (108), have served as the basis or model for the schemes now being proposed or endorsed by the French workers (142) and by me (40, and present paper).

Jankowski (108), using the subclass level, established and named 3 major groups : the Fragmophora, the Tetrahymeno-

‘Many of which, however, were not available or not known to him at the time of preparation of his 1967 monograph.

phora, and the Polyhymenophora. These embraced all ciliate species except certain “lower” gymnostomes and the suctorians. (Unfortunately, in his 2nd systematic review, 110, the Russian ciliatologist has dropped the 2nd and 3rd taxa, or their names, and greatly reduced the significance of the 1st one: a “recan- tation” rather difficult to understand.) We (40, 142) are, in effect, reviving these 3 of Jankowski’s 1967 names or concepts, although expanding the boundaries of the 1st group and treating it as a class (with a larger, more exact, name, as well); restricting the coverage of his 2nd assemblage a bit, and renaming it, more appropriately, the Oligohymenophora; and retaining his Polyhymenophora, for the old “spirotrich” group, without significant change (except for elevating it to class rank).

In any complete revision of a major taxonomic group, there are at least 3 areas or kinds of contributions, interrelated to various degrees: assignment of new names to groups appearing to need them; rearrangement of high-level taxa, frequently with changes in comparative rank; and promulgation of new ideas or hypotheses concerning possible evolutionary or phylogenetic relationships, ideas often directly affecting the taxonomy of the group. Jankowski’s most important contribution has been in this last-mentioned category. His clearly expressed views will, I predict, have a long-lasting impact on the field.

Some further consideration is given to Jankowski’s valuable works in the area of ciliate systematics elsewhere (40). But even in the present note, the reader’s attention should also be called to his separate and magnificent monograph on the Chonotricha ( 11 1 ) , a work which demonstrates abundantly that these forms should now be recognized as a major and evolutionarily exciting assemblage of ciliophorans.

COMPOSITION O F KINETOFRAGMOPHORA The class Kinetofragmophora de Puytorac et al., 1974, in this

writer’s opinion, is comprised of 4 major subgroups or subclasses, a view paralleled in a general way by that of the French group (142)z.

Subclass Gymnostomata Biitschli, 1889 contains the allegedly more primitive families of the old gymnostomes sensu lato, including forms with “relict” macronuclei, species with essentially apical cytostomes, organisms with relatively noncomplex cytopharyngeal armature, and close relatives (including the old “pleurostome” group). Members of this subclass are treated in more detail below.

Subclass Vestibulifera de Puytorac et al., 1974 is comprised of the old trichostomes sensu lato (but with Colpodidn de Puytorac et al., 1974 now separated out by Dragesco and Grain as an independent order) plus the entodiniomorphids, an entire group removed from the “spirotrich” assemblage following the rather startling findings of Noirot-TimothCe, Wolska, and others in recent years (see literature citations and further discussion below).

Subclass Hypostomata Schewiakoff, 1896 emend., de Puy- torac et al., 1974, represents the most greatly expanded group within the old gymnostome framework, systematically the most controversial. It is here that data from investigations with electron microscopy have been most helpful, but possibly far more

‘Unfortunately, at the time of sending the present MS off to the Editor I have still not had an opportunity to view the MS by de Puytorac et al. (142) ; thus, making a detailed comparison of my paper, here, with the French contribution is clearly difficult, as I hope the reader will appreciate. Both papers, including, additionally, mine in Systematic Zoology (40), will presumably be “in press” at the same time.

NEW TAXA I N CILIATE CLASS KINETOFRAGMOPHORA 209

information is needed to arrive at enduring taxonomic conclusions: several of the outstanding problems are considered in subsequent sections of this paper.

Subclass Suctoria Claparede & Lachmann, 1858 seems straightforward enough, although 2 areas of major dispute continue: the level or rank of the whole group ( a number of ciliatologists, including Jankowski and Poljansky, would still like to see it elevated to a position equivalent to that of all of the other ciliates massed together); and the number of suprafamilial included groups. No further comments will be made on this subclass in the present work except to note that a number of suctorian ultrastructural characteristics are reminiscent, to a greater or lesser degree, of the situation found in certain other ciliophoran groups, a fact tending generally to support a phylogenetic closeness, rather than separateness, of Suctoria and other high-level ciliate taxa.

CHARACTERIZATION O F KINETOFRAGMOPHORA The name chosen for this class should receive brief considera-

tion. It is based on the apt term proposed by Jankowski (108), kinetofragmon, to describe his concept of one or more kinetids separated from or derived from a ‘‘regular” somatic kinety of a ciliate and forming a part of the “PCA” (preoral ciliary apparatus) at the anterior end of the organism. Either primarily or secondarily, such “kinetal fragments” are, in my opinion, involved in the structure and morphogenesis of the oral ciliature of all of the species assigned to the new class; thus they are kinetofragmon-bearing forms or “kinetofragmophorans.” Jankowski himself ( 108) created a new subclass Fragmophora, the name obviously based on the concept described above. But he used it for a far more restricted group of ciliates; and he reduced it still more in importance in his 2nd paper (110) on overall ciliate systematics. Thus, I believe that there is ample justification for the proposing of a new and more accurately descriptive name by de Puytorac et al. (142), even though it is, unfortunately, a longer word.

A convenient way to describe a major group succinctly is to list the principal features shown by all or most of its included species. In the present case, the class Kinetofragmophora may be considered to embrace those ciliates that show various com- binations of the following characteristics deemed to be of comparative taxonomic value. Detailed consideration is beyond the scope of this note. Such data, plus appropriate drawings, photomicrographs, and electron micrographs, will be appearing in the forthcoming monographs referred to a t the beginning of this paper; attention may also be directed to the figures in Corliss (40).

The oral ciliary apparatus, if present or if recognizable as such at all, is not associated with a definitive buccal cavity. The circumoral infraciliature is ultrastructurally unique and far less complex than that of the buccal infraciliature of the 2 “higher” classes, the Oligohymenophora and the Polyhymeno- phora.

2. Any oral cilia which are present are essentially in a perioral position with respect to the cytostome and cytopharynx and arise (with rare exception), both ontogenetically and phylogenetically, from somatic elements or kinetids or from isolated fragments of the somatic kineties. There is no true autonomy in the continuity of such oral ciliature and generally no oral kinetosomal field sensu stricto in stomatogenesis.

A telokinetal type of stomatogenesis is exhibited (following the terminology of my very recently revised treatment of patterns of stomatogcnesis: see 39) ; occasionally the type may

1.

3.

resemble the parakinetal mode seen in species of the “higher” classes of ciliates.

4. The cytopharyngeal apparatus, or “nasse,” is often a prominent feature. I t is composed of bundles of microtubules or nematodesmata, more complex in arrangement in some taxa than in others.

The macronucleus is relatively primitive, very primitive or even missing in certain groups, showing evolutionary stages in development generally believed to have preceded most of those typical today of the two “higher” classes.

Body ciliation is variable, ranging from complete absence in the mature stage of the life cycle (e.g. in the Suctoria) to a heavy and uniform covering.

Body form and size cover a broad range, related to many specializations both morphologic and physiologic in nature.

A wide diversity in habitats, feeding habits, and modes of life (free-living, epibiotic, parasitic, etc.) is shown. Species may be marine, freshwater, or brackish in their biotopes, with a large and rather specialized fauna (particularly in the 1st subclass) characterizable as psammophilous (see a subsequent section of this paper). Cysts are common (42) .

The Kinetofragmophora is by far the largest of the 3 classes of the Ciliophora. I t contains the greatest numbers of species and of higher taxa; not unexpectedly, it also shows the greatest diversity of included forms. A number of “evolutionary experi- ments” may be considered as having been tried within this vast and ancient assemblage, all the way from the intriguing case of homokaryotic species to the appearance of the distinctive suctorians. In spite of the immenese aid available today through electron microscopy, it will not be surprising if the “internal taxonomy” of this class remains in a somewhat un- settled condition for some years to come.

A detailed taxonomic revision of the class, complcte with lengthy descriptions or characterizations, nomenclatural syn- onymies, drawings, and micrographs, is well beyond the scope of the present paper, as noted above. I n order to identify and name several new high-level taxa, however, a t least a brief survey of all of the included suprafamilial taxa is necessary; often, but not always, my “skeletal scheme” will coincide with that to be found in the in-press treatment by de Puytorac et al. (142); and it has been incorporated in my review paper (40), also in press.

I envision the 1st subclass, Gymnostomata, as comprised of 3 orders, 2 newly erected here, with 4 suborders (all but 1 here described for the 1st time) assigned to the first one. Particular attention is given to the classification and general biology of members of the 1st order, Primociliatida n. ord., because that group, sizable though it is, is given relatively little attention in the other 2 works in press (40, 142) on overall revision of ciliate systematics.

5.

6.

7.

8.

ORDER 1. PRIMOCILIATIDA3 n. ord. Protozoologists have long assumed that some group among

the gymnostomes was most likely ancestral to all other ciliates and probably arose, itself, from some homokaryotic zooflagel- late stock ( 3 6 ) . Although factors of convergence (adaptation to particular habitats, etc.) may have played some role in causing certain present-day gymnostomes to seem relatively

3 A better choice for the name of this new taxon would be the word “Protociliatida.” Unfortunately, the widespread and well entrenched usage of this term for Metcalfs opalinid infusorians, although they may well have been effectively banished from the phylum Ciliophora altogether (see 105 and works cited therein), essentially precludes its adoption here.


“simple” in many of their features-their overall organization, their frequent lack of highly differentiated cortical or ciliary structures or organelles, their uncomplicated polarity and symmetry, their usually terminally located oral apparatus, their carnivorous feeding habit, etc.-it is not entirely unreasonable to postulate that their primitiveness also may be related to their possible location, in eons past, at the base of the ciliate phylogenetic tree (36, 38, 40, 64, 108). Today support for this idea has come from an unusual source, from comparative investigations of the nuclear apparatus, generally of little taxonomic significance in ciliatology at suprageneric levels (41 ) . Raikov’s ( 170) concept of “karyological relicts” (term first suggested by Grell, 95, 96) makes profitable the consideration of the existence of certain relict species and even relict higher taxonomic groups among gymnostome ciliates. It seems to me that such a phylogenetic postulation demands, or at least suggests, recognition of several new taxonomic groups; hence a major aim of this paper is to name and erect the assemblages that I believe are involved.

The Primociliatida n. ord. is characterized essentially by its 4 included suborders (see below). It is approximately the equivalent of the bulk of the old rhabdophorine gymnostomes of Fad-Fremiet and Corliss (see 33, 36), except for those now assigned to the more specialized, smaller groups comprising the following 2 orders. Some of the best-known of all ciliates are to be found here, and habitats range from marine biotopes, including the sands of intertidal zones (see next section), to brackish and freshwater habitats of all kinds and to the restricted niches afforded by the endoparasitic lives led by some of the included species.

Interstitial Ciliates

A brief digression from my strictly taxonomic survey seems in order, primarily because of its pertinence to my decision to give more prominent classificational recognition to certain groups of gymnostome ciliates but also because the topic has been so sadly neglected by ciliatologists in general to date. Thus a cursory review of our knowledge of the interstitial ciliate fauna is presented here.

The pioneering work on such forms dates back fewer than 50 years, and it was some years later before the investigations of men like Sauerbrey (181) and Kahl (112, 113) were followed by the comprehensive accounts of Bock (7-9) and FaurC-Fremiet (65-67). In the late 1950’s and 1960’s, research on psammophilous forms began to flourish (1-3, 12,

178) and “primociliates” figured prominently among the species found in this ancient and rich biotope, then being re- discovered.

Since 1970, significant taxonomic-ecologic works on interstitial ciliates have continued to appear (4, 5, 16, 17, 22, 23, 98). However, as discussed in subsequent sections of this paper, considerably greater interest has developed in the nuclear condition of many of the sand-dwelling “lower” gymnostomes, resulting in sophisticated RNA-DNA studies (62, 170, 176, 187), following particularly the extensive series of earlier cytochemical investigations carried out by Raikov (155, 157- 159, 162-164), Faurk-Fremiet (65-67), and Dragesco (49, 53-55 ) , and in ultrastructural research of considerable promise (e.g. 118, 171-175, 177).

Except for the interpretations briefly presented by FaurC- Fremiet (67, 69, 76), theoretical papers involving psammophilous forms and concerned with such significant matters as the origin of the kind of nuclear duality so characteristic of

14, 46-49, 51, 53-55, 69, 83-85, 130, 134-136, 160, 161, 165,

today’s ciliate Protozoa have emanated principally from the Poljansky-Raikov school in Leningrad (137, 138, 156, 166, 168, 170). Worthy of note, however, is the space devoted to the subject in 2 recent textbooks of protozoology by Grell (97) and Sleigh (182). The present paper represents the 1st serious attempt to apply such ideas to the systematics and classification of these “lower” or possibly “relict” ciliophoran species, clearly a difficult task in view of the still all too numerous gaps in our knowledge of these forms.

( 1 ) Homokaryotina n. subord.

From an evolutionary point of view, this group was possibly an extensive and certainly a long-lived one filled with the hypothetical “euciliate” ancestors, species with but one kind of nucleus-neither a micro- nor a macronucleus, as we know these chromatin-rich organelles today. Many, or one major, line(s) may have evolved from populations within the group, and many subgroups may have become extinct. Yet today we still find, in species of the genus Stephanopogon, persistence of the homokaryotic condition.

Therefore, I am erecting the Homokaryotina n. subord. as a kind of “relict” taxonomic group containing, today and at our present state of knowledge, the single genus Stephanopogon Entz. Its characterization rests mainly on its major and unique feature of possession of but a single type of nucleus (though occurring in the multiple condition), probably diploid and containing a prominent RNA-rich endosome or nucleolus. I am in full agreement with Raikov (170) that this condition was not likely to have arisen secondarily from the dual nuclear apparatus characteristic of the “higher,” heterokaryotic ciliates.

Stephanopogon is a small marine organism and appears primitive in other features as well as in its nuclear apparatus. It is sand-dwelling, flattened dorsoventrally with only the ventral surface ciliated; its oral area is apical, slit-like and unadorned; it appears to possess neither specialized nor unique cortical or cytoplasmic organelles; its feeding behavior is alleged to be simple. I t undergoes multiple, but uncomplicated, division within a reproductive cyst. For modem studies and references to the rather scant older literature on this most interesting organism, see Borror (13), Dragesco (53), and Raikov (170).

If diligent searches are carried out in appropriate ecologic niches, I predict discovery of additional surviving homokaryotic ciliates. Such an effort would be a most worthwhile under- taking.

(2 ) Karyorelictina n. subord.

This group, which I admit is potentially the most artificial of the several suborders here recognized, is erected to establish a taxon for the ciliates which surely must have existed at one time to bridge the gap between homokaryotic forms and the “typical” heterokaryotic, and now predominating, species with diploid micronuclei and polyploid (or, better, polygenomic ) macronuclei. Even today, thanks especially to the thorough investigations of Dragesco, FaurC-Fremiet, and Raikov (see references cited on a preceding page of this paper), such a group of heterokaryotic ciliates is known to exist, a group still manifesting this intermediate or “in-between” condition.

The micronuclei are diploid, are rich in DNA, divide mitot- ically, undergo meiosis during occurrence of sexual phenomena, etc.; but the macronuclei are also (only) diploid, are rich in RNA, never divide, but are replaced periodically by differen- tial transformation of certain of the division products of the

NEW TAXA IN CILIATE CLASS KINETOFRAGMOPHORA 21 1

micronuclei. Both kinds of nuclei are generally small in size, spherical or ovoid in shape, sometimes occurring in great numbers throughout the cytoplasm, and sometimes present in one or more nuclear complexes or clusters. As suggested for Stephanopogon in the preceding section, here, too, I am in accord with Russian workers (e.g. see 170) that the curious ‘‘relict’’ macronuclei known in such surviving species could hardly have been derived evolutionarily by “degeneration” of the polygenomic macronucleus typical of the majority of extant ciliates.

The present-day f o r m s s o m e 100 species described to date- in which macronuclear polyploidization has not yet evolved may be considered, from the po in t of view of nuclear development, to be remnants of a once much wider spectrum of forms. I t may be of significance that thcse species are not all found in the same habitat today (although the majority are marine sand-dwelling forms4) and that they do not all appear to be members of the same ordinal or subordinal group, consider- ing such taxa as those available in existing schemes of classification, when compared in other taxonomic characteristics. At our current state of imperfect knowledge, then, we appear to have 4 options, taxonomically speaking. (A) We can consider various groups of extant forms with diploid macronuclei to represent now separate, un-named lines or suborders evolved with the passage of time from a single group such as the one here named Karyorelictina n. subord., leaving it as perhaps an empty taxon. (B) We can argue that, in spite of subsequent evolution in other characteristics, retention of the ancestral feature of a primitive macronucleus is significant enough to require keeping all forms still showing it in a single taxonomic group, the one named here as new. ( C ) We can minimize the importance of the nuclear state and disperse all species manifesting the primitive condition into separate existing taxa on the basis of other characteristics they possess, making totally unnecessary the creation of any new high-level taxa and suggesting that polyploidization of the macronucleus occurred repeatedly in the early evolution of ciliates ( a real possibility). ( D ) We can suggest a kind of compromise solution: recognize the likelihood of a single ancestral taxon (the Karyorelictina n. subord.) and assign most species with diploid macronuclei, especially those still found in the psammophilous biotope, to it; yet, at the same time, realize 2 important further possibilities-that the taxonomic affinities of some such extant species need not be considered particularly close5 (even if

’ It might be well to note here that species belonging to the 2 “higher” classes of the Ciliophora, the Oligohymenophora and the Polyhymenophora, are also found in the sands of the intertidal zone (e.g. see Dragesco, 49, 51, 54, 55; and various papers by others whose works are cited in a preceding section of this paper). They, too, by convergent evolution, show certain characteristics, apparently selected for survival in such a particular habitat, held in common with kinetofragmophoran species living in the same biotope. It is a fact of significance, however, that none of the “higher” forms exhibits a macronucleus arrested at the diploid stage of its evolutionary history. Incidentally, mention must be made, even if only in passing, that students of multicellu- lar organisms comprising the intersitial fauna are faced with similar exciting evolutionary and ecologic problems, since intertidal sands are widely believed to have been-and to be-a habitat favoring survival of ancient or primitive forms among the Meta- zoa. For classical early papers on the ecology and general taxonomy of the micro-metazoan forms of psammophilous inclination, the reader is referred to the following works, by recognized lead- ers in the field, and to the citations in their bibliographies: Pen- na i (133), Remane ( 1 7 9 ) , and Swedmark (186).

In a way, I am subscribing to the “grade” concept here, suggesting that the forms under consideration represent a certain evolutionary level in their organization and that classifying them

the forms are grouped together at a high level) and that perhaps a few forms (such as the freshwater, detritus-inhabiting Loxodes spp.) are better classified today in such an order as the Pleurostomatida (see below), a group which then would contain organisms with both diploid and polygenomic macronuclei. I t is this 4th option, ( D ) , which I am provisionally adopting, here and elsewhere (40) .

In the present note, further speculative discussion of this fascinating topic is not appropriate. I t is also obvious that more data relevant to it must be collected to aid future systematic ciliatologists in making the most judicious taxonomic decisions concerning the organisms involved. But, even at this time, I would definitely assign to the Karyorelictina n. subord. the formerly rhabdophorine genera of Trachelocerca Ehrenberg, Trachelonema Dragesco, and Tracheloraphis Dragesco; and possibly Ciliofaurea Dragesco, Cryptopharynx Kahl, Geleia Kahl, Kentrophoros Sauerbrey, and Remanella Kahl (or should some of these, with Loxodes Ehrenberg, be assigned elsewhere?).

A convenient indirect source of much of the literature on members, or potential members, of this new suborder is the excellent review paper by Raikov ( 170). All of the principal works published to date on the ciliate interstitial fauna have been cited in a preceding section of the present paper; many of them contain data on karyorelictine species. Papers concerned with the infraciliature, morphogenesis, and electron microscopy of the taxonomically enigmatic genus Loxodes also are becoming quite numerous (for some recent examples, see 6, 58, 59, 61, 67, 76, 129, 145, 158, 170, 173, 193; and con- sult the literature review by Canella, 27) .

(3) Prostomatina Schewiakoff, 1896

For this restricted group of the large assemblage formerly known as the rhabdophorine gymnostomes, I have resurrected the old tribal name of Schewiakoff which emphasizes nicely one of the principal taxonomic characters exhibited by the included species : possession of a relatively simple, externally unadorned oral area located at the apical pole of the organism (elsewhere with rare exception). The cytostome is oval or rounded. The cytopharyngeal complex is composed of bundles of microtubules, the nematodesmata, but does not sliow the ultrastructural elaboration so typical of members of the subclass Hypostomata. Toxicysts are absent from the region of the mouth.

The macronucleus is polyploid or polygenomic in all species included here, but otherwise it is relatively simple and not highly evolved. The micronucleus is usually single in number.

Holophrya Ehrenberg may be considered to be a representative genus. Buetschlia Schuberg and neighboring genera (all distinguished by possession of so-called concretion vacuoles and by occurrence as commensals in the digestive tract of such herbivores as camels and horses) also are members of this relatively “lowly” group of gymnostomes. Urotricha Clapar2de & Lachmann, or a Urotricha-like ancestral form, may have been the gymnostome involved in the origin of the hypostome subclass within the class Kinetofragmophora; and, similarly, Platyophrya Kahl may have been ancestral to the vestibuliferan assemblage (the old trichostome group, expanded; see 142).

Many of the species assignable to this suborder are well known from the literature. Recently, ultrastructural investiga-

together may be somewhat more of a taxonomic convenience than a proven or provable matter of reality. Further discussion of this interesting possibility, however, is well beyond the scope of this short paper.


tions have been carried out on some of them (e.g. see 90, 94, 140, 143).

(4) Prorodontina n. subord. This is another specialized group split off from the over-

large, over-extended former rhabdophorine gymnostome group of Faurt-Fremiet and Corliss. Based particularly on details of the infraciliature and the ultrastructural findings reported by half a dozen skilled French protozoologists (56-59, 78, 79, 94, 114, 139, 144, 146), the suborder may be characterized by the presence of a more complex “nasse” or cytopharyngeal complex of nematodesmata than is typical of such prostomatine ciliates as Holophrya, although not as elaborate as those of members of the subclass Hypostomata, and by exhibition of a mode (or subtype) of stomatogenesis which also is postulated to be more highly evolved than that known in the Prostomatina.

The oral apparatus is still terminally, if not apically, located; and the cytostome is oval or rounded and essentially on the surface of the organism; but body symmetry at the anterior end is seldom radial. There are no toxicysts in the immediate area of the mouth. Except for their possession of polygenomic macronuclei and a few important ultrastructural specializations, the species assignable here remain representative of relatively primitive forms when viewed from the perspective of the evolution of the entire phylum Ciliophora. I t seems justifiable, however, to recognize their separation from the larger preceding suborder.

Among the genera to be included within the Prorodontina n. subord. are Coleps Nitzsch, Placus Cohn, Prorodon Ehren- berg, and various related forms.

ORDER 2. HAPTORIDA n. ord. In recognition of the accumulating ultrastructural differences

of taxonomic significance now being found in gymnostome species formerly lumped together in the single suborder Rhabdo- phorina, it appears sensible to set off a number of genera into still another separate high-level taxon. Therefore, I am here erecting a 2nd new order, named the Haptorida n. ord. because of the “seizing” behavior, in feeding, of the rapacious carnivorous forms included in it. Future workers may wish to divide this new group into 2 (or even more), based on fine- structural differences in organization of the cytopharyngeal apparatus, for example, but no such attempt is made here.

The haptorids are gymnostomes which may be distinguished by their terminally located cytostome (of slit-like shape in some forms) and often their unique display of toxicysts in the oral region. Areas of thigmotactic or clavate ciliature may be present. Their feeding behavior, as mentioned above, also represents a unique characteristic.

The group includes some of the longest- and best-known genera in the entire phylum Ciliophora; for example, to list but a dozen or so: Actinobolina Strand, Chaenea Quennentedt, Cyclotrichium Meunier, Didinium Stein, Dileptus Dujardin, Enchelyomorpha Kahl, Enchelys 0. F. Muller, Lacrymaria Bory, Monodinium Fabre-Domergue, Paradileptus Wenrich, Spathidium Dujardin, Teutophrys Chatton & Beauchamp, Tra- chelius Schrank.

Of these, species of Didinium and Dileptus have been the most intensively studied in recent years and with the most modern cytologic technics (50, 52, 60, 63, 88, 89, 93, 94, 101, 180, 184, 194-196, 203). But investigations with electron microscopy have also now been carried out on various species of other genera in the order (10, 11, 45, 77, 78, 80, 81, 92, 94, 102-104, 116, 117).

Although the Haptorida n. ord. are being split off from other gymnostomes, the observation should be made that the diversity within the new group is still immense, further justification for the recognition of the new taxon at the ordinal level. The former lumping together of all “rhabdophorine” forms- we can now say, in retrospect-was taxonomically naive. Once again, it is the advances made with the aid of electron microscopy that, as I have stressed elsewhere (40, 41), today allow such an improved understanding of both the separateness and the interrelationships of the major ciliate taxa.

ORDER 3. PLEUROSTOMATIDA Schewiakoff, 1896 Schewiakoff’s old tribe of essentially the same name and

composition is here retained, elevated to ordinal rank, for the group of gymnostome ciliates so aptly described by the appella- tion: that is, forms with “lateral mouths.”

The curious ciliates comprising this 3rd order are often of relatively large body size; are frequently elongate and laterally compressed; and all possess a slit-like but nonapically located cytostome. They may well have evolved from the suborder Karyorelictina n. subord. (of the order Primociliatida n. ord., described above), although the “typical” species now found here possess polygenomic macronuclei.

Four representative genera are Amphileptus Ehrenberg, Bryo- phyllum Kahl, Litonotus Wrzesniowski, and Loxophyllum Du- jardin. Beyond the scope of the present note is discussion of the advisability of including here (perhaps as a separate suborder) such sand-dwelling forms with diploid macronuclei as Geleia Kahl and Loxodes Ehrenberg and alleged relatives, a taxonomic-phylogenetic problem touched on in a preceding section ( Karyorelictina n. subord. ) but still requiring additional data before reasonable resolution will be possible.

Although pleurostome species are numerous and well known (6, 13, 15, 17, 25, 49, 55, 56, 58, 61, 112), they have been little studied with the electron microscope. However, several very recent ultrastructural investigations (94, 99, 100) are providing some interesting data of potential use in future more refined study of their comparative taxonomy.

SUBCLASS VESTIBULIFERA The 2nd subclass of the class Kinetofragmophora, Vestibuli-

fera de Puytorac et al., 1974, needs to be mentioned briefly here because I am proposing a new suborder in one of its 3 orders.

While agreeing fully with FaurC-Fremiet (72) that the old order Trichostomatida could be made much more rnonophyletic by removal of several families traditionally included there, the assemblage also needs to be expanded today in order to embrace the “spirotrichous” Entodiniomorphida, a group which may be recognized as having close affinities with the trichostomes sensu stricto. The bases for making the latter major taxonomic shift are to be found in the heuristic papers of Grain (90, 91), Noirot-Timothte (131, 132), and Wolska (197- 202). Jankowski (110) was the first “synthesizer” to make the actual change in the ciliate classification scheme, although he put the entodiniomorphids in a subclass position paralleling the “holotrichs” (which, like the “spirotrichs” and several other groups, were given the rank of subclass in his scheme). Canella (27) discussed the question, covering the literature on the subject up to 1970; via personal communications, he was able to “forecast” Jankowski’s moves. The “companion” papers of the present communication (40, 142) place the organisms in the position also endorsed here: one of 3 orders comprising the subclass Vestibulifera.

NEW TAXA I N CILIATE CLASS KINETOFRAGMOPHORA 213

A comment may be in order here concerning the name Vestibulifera, but characterization of the group and defense of the concepts involved are subjects totally beyond the scope of the present paper (but see 142, and the several monographs in preparation). We thought originally (following the May meeting in France) that “Vestibulata” would be most apt as a name. But that designation has been used in the past (86, and see 34 for its history) for quite a different group (including the genus Paramecium, now a hona fide hymenostome ciliate), so it became inappropriate.

The 3 orders included in the new subclass are: Trichostoma- tida Biitschli, 1889; Colpodida de Puytorac et al., 1974; and Entodiniomorphida Reichenow in Doflein & Reichenow, 1929. Nothing more will be said here about these orders (but see 40, 142), except to discuss the new suborder, within the last order, erected here as new.

BLEPHAROCORYTHINA N. SUBORD. In addition to moving the “true”-thanks primarily to the

observations of Grain and Wolska cited above-entodiniomorphids into the vestibuliferan fold, we may recognize today their possible progenitors among some little-known trichostomes which include the family Blepharocorythidae. Thus I believe that the classically known ophryoscolecids and relatives should comprise the nominate suborder, the Entodiniomorphina, with that name credited to Reichenow again, and that the allegedly ancestral group should be grouped in a suborder called Blepharo- corythina n. subord.

Members of the suborder have the characteristics of the family which comprises it, the Blepharocorythidae Hsiung, 1929 (see 106, 107), updated by the infraciliary and ultrastructural findings of Grain and Wolska. The cytostome is anteriorly located, in a “vestibulum” of sorts. Somatic ciliature is reduced, and tufts of “syncilia” (to use an appropriate term originally suggested for ophryoscolecids by von Gelei, years ago; see 87) appear at various locations on the body, ciliary organelles whose fine structure is strikingly like that shown by the similar ciliary organelles (often called membranelles) found in the ento- diniomorphines. The species, belonging to several genera, occur in the digestive tract of horses and ruminants; but their similarity to the entodiniomorphids sensu stricto is not considered to be due solely to convergence.

A particularly full and well illustrated account of the group, including new ultrastructural data, may be expected in the section of the forthcoming Trait6 volume being prepared by Jean Grain.

SUBCLASS HYPOSTOMATA Even as Kinetofragmophora is the largest class in the “new

new systematics” of the phylum Ciliophora (40) , Hypostomata Schewiakoff, 1896 emend. de Puytorac et al., 1974 is very likely the largest and most diverse kinetofragmophoran subclass. As I envision it (which, as pointed out below, is not identical to the French view), the Hypostomata is comprised of 6 orders, 3 with suborders. Two suborders are described as new in the present paper (see following sections).

To me, the great problem in the taxonomy of this subclass rests in the proper comparative interpretation of the rapidly accumulating ultrastructural data, some not yet even published (and many more still to be discovered). Therefore, I must predict that our views of the “proper” intragroup classification will change as our knowledge grows. The dilemma is that we can hardly leave the group to one side in our current

overall revision of the ciliates. In my opinion, the world‘s expert on the systematics of the group is Gilbert Deroux; but until his scheme of classification is available in full and published form, we have to make our own deductions. Thus my present views will differ, in part, from those of the French school (e.g. see 142); however, tentative though they may be, I should like to expose mine here and offer justification for establishing 2 new suborders, groups also adopted in my major paper in press (40). As has been true throughout the present note, fuller explanations, figures, etc. must be sought and/or awaited in other or subsequent publications.

The 6 orders that I place in the subclass are as follows: Cyrtophorida FaurC-Fremiet in Corliss, 1956; Nassulida Jan- kowski, 1967 (with 2 suborders: see below); Microthoracida Jankowski, 1967; Rhynchodida Chatton & Lwoff, 1939 (with 2 suborders; see below) ; Chonotrichida Wallengren, 1895 (with suborders Exogemmina Jankowski, 1972, and Endogemmina Jankowski, 1972); and Apostomatida Chatton & Lwoff, 1928. Although, as is apparent from the authorships and dates, most of these groups have been known for some years, they have never before been placed in the taxonomic juxtaposition indicated here (and in Corliss, 40) . The chonotrichs have recently been thoroughly and beautifully monographed by Jan- kowski ( l l l ) , and older papers of importance are cited in Corliss (36); attention should also be drawn to Mohr’s (128) most recent contribution. The apostomes, so magnificently treated in Chatton & Lwoff‘s classic (28), remain taxonomically enigmatic, though precise ultrastructural work such as that carried out by Bradbury (18-21) eventually should help solve the riddles surrounding their phylogenetics.

PARANASSULINA N. SUBORD. The order Nassulida, now separated from the cyrtophorids

and relatives primarily on the basis of new knowledge of dif- ferential value available from ultrastructural studies and pro- targol silver investigations (43, 44, 68, 70, 71, 73-75, 78, 94, 126, 147, 167, 169, 185, 188-192), embraces a number of families of hypostome ciliates which themselves are turning out to be so different (i.e. taxonomically separable at a high level) that creation of several suborders seems indicated. How many suborders and what they should be named remain open to some question, and I do not profess to know the best answers at this time. Jankowski (108, 110), in effect, included the Micro- thoracida, a separate order in my scheme (40), as another nassulid suborder. Other workers may wish to follow his lead, but I have long been impressed (25) with the unique phylogenetic significance of such microthoracid fo rm as Pseudo- microthorax.

I am suggesting 2 nassulid suborders; Nassulina Jankowski, 1967, the nominate suborder; and Paranassulina n. subord., as the 2nd suborder. The new group is not characterized here in detail; such treatment will be appearing elsewhere. But the principal differences which allow-or, in my view, require-the separation of the paranassulines from the nassulines should be pointed out. Most significant, perhaps, is the con- traction (in size) or lessening of the area occupied by the “pseudomembranelles” of the “frange” until they are truly in a restricted perioral position-and located only on the left- with respect to the nasse. In fact, these kinetal fragments of oral ciliature, ranging from 3 to several in number depend- ing on the genus, even come to lie in a kind of “vestibulum” or “buccal cavity” in several species. The pre-oral suture at the apical pole has become more extensive, and a large number of ciliary meridians or kineties terminate on it, losing a degree

214 NEW TAXA IN CILIATE CLASS KNETOFRACMOPHORA

of their primitive bipolarity. The process of stomatogenesis thus also becomes more localized and easily distinguishable from that known for species in the suborder Nassulina and in other neighboring orders and suborders.

Among the genera included within the Paranassulina n. subord. are Paranassula and Enneameron; species of these genera are easily separable, today, from species of such nassuline genera as Nassula, Nassulopsis, Orthodonella, and Scaphidiodon by the appearance of their infraciliature (structurally and ultrastructurally ) and by details of their stomatogenesis. How- ever, a number of species known only from the older literature were misidentified in original descriptions; thus a great deal of taxonomic-nomenclatural confusion exists. But discussion of such problems, important though they are, is beyond the scope of the present note. Incidentally, with typical vision, FaurC-Fremiet (68, 70, 71, 74, 75) laid much of the ground- work for our modem understanding of these fascinating and not-at-all-simple ciliates.

ENIGMATIC ORDER RHYNCHODIDA For some years a number of ciliatologists have realized that

the bulk of Chatton & Lwoff‘s “thigmotrichs,” the so-called arhynchodine or mouthed forms, must lose their independent identity at the level of order or suborder and become absorbed by the pleuronematine scuticociliates (37, 40, 183). What should be the fate, then, of the few remaining “thigmotrichs,” the rhynchodine (literally, rostrate) or suckered forms? Here the problem is complicated by recently discovered differences within the group itself, leading to various hypotheses concerning affinities which, with a measure of despair, Canella (27) terms ‘‘th6ses diamttralement oppostes” in his recent review of the controversy. In spite of still morc recent discoveries via electron microscopy, the taxonomic problems involved have not (yet), in my opinion, become easier of resolution. It is beyond thc scope of this note to reconsider all of the data and ideas involved; but it is necessary to attempt a justification for the tentative stand which I have developed for exposure here (and see 40), fully aware of the likelihood of change as additional pertinent data become available.

Briefly, the 4 acceptable families of the old “thigmotrich” suborder Rhynchodina-the Ancistrocomidae, Crateristomatidae, Hypocomidae, and Sphenophryidae-themselves appear to fall into 2 quite diverse taxonomic groupings. The 2 larger families, the Ancistrocomidae and the Sphenophryidae, possess a true rostrum and arc alike in other characteristics; on the other hand, the Hypocomidae and Jankowski’s (109) recent mono- generic family Crateristomatidae have a cytopharyngeal apparatus, no rostrum, and show ultrastructural characteristics strongly reminiscent of members of the order Cyrtophorida.

My provisional resolution of the difficulties briefly exposed above is to recognize as an order the former “rhynchodine group,” as a whole; this view has been maintained in Corliss (40), also. Then the 4 controversial families are arbitrarily assigned to either a nominate or a 2nd suborder. The Hypo- comidae group goes into the suborder Rhynchodina Chatton & Lwoff, 1939; and the Ancistrocomidae group, into a new suborder, Ancistrocomina n. subord. The Rhynchodina so defined are considered to have arisen phylogenetically from the hypostome order Cyrtophorida; the origin or exact affinities of the Ancistrocomina n. subord. remain problematical, although the old Chatton-Lwoff idea (30, 31) that they arose from the Hemispeiridae assemblage of the former arhynchodine “thigmotrichs” (now pleuronematines in the oligohymenophoran subclass Hymenostomata) remains a viable possibility.

Part of the “rhynchodid debate” currently in progress between the French school (see 142) and myself (40, and the present paper) is, unfortunately, related to the least important part of the overall matter, the choice of nomenclature for the 2 suborders. I must admit that my assigning the nonrostrate hypocomids to the suborder Rhynchodina is etymologically disturb- ing; and leaving the 2 groups within the same order may turn out to be unwise. To have allocated the hypocomids to a suborder with a name such as “Hypocomina,” however, was im- possible because of preoccupation: that is, there is a genus Hypocomina in the contrasting family Ancistrocomidae, as fate would have it! The name “Arhynchodina” would not do for the hypocomids because it was used for some time to identify the bulk of the now vanquished “thigmotrichs” proper.

Therefore the French workers, particularly Gilbert Deroux, and I are not really so far apart in our current understanding of a taxonomic mess which is not truly resolvable until more data are available. We both agree that the hypocomids-call them what you wi l l should be considered close to the cyrtophorids; and that the ancistrocomids are different, to an un- known degree, from the hypocomids and may deserve a more appropriate taxonomic location (than one alongside the hypocomids) when we can figure out what that should be. In the meantime, the provisional allocations made in this paper may be considered to represent improvement over the classificational arrangements of the past when even fewer data were available for comparative study.

COMPOSITION OF SUBORDERS

Characterizations of the 2 rhynchodid suborders, controversial though their names may be, have been covered to some extent in the preceding section. But an additional word about genera assigned to them and about recent publications 011

both groups is in order here, as well as a few other remarks on their identification and classification, although they must be made briefly.

Rhynchodina Chatton & Lwoff, 1939 If this group is to continue to be restricted to inclusion of the

Hypocomidae and the Crateristomatidae, the small total number of genera can be named with ease: Crateristoma, Heterocoma, Hypocoma, and Parahypocoma. The infraciliary plan of species studied carefully is strikingly like that of certain dysteriid cyrtophorids (for latest strong support of this view, see 43) . The ventrally placed cytopharyngeal tube (ending in a “sucker”) and the posteriorly located adhesive organelle are also reminiscent of cyrtophorids, the most likely ancestral group. Stomato- genesis appears to involve a “kinetofragmon” always found in the upper right sector of the ventral surface.

Species of the genera Heterocoma, Hypocoma, and Para- hypocoma are parasites in or on tunicates or peritrich protozoa and have been little studied since their monographic treatment by Chatton & Lwoff (29, 31).6 However, very recently a note has appeared on a new species of Parahypocoma found in solitary ascidians from the Pacific Northwest (24). The single species comprising the genus Crateristoma is parasitic in the mantle cavity of Balanus (109).

Unfortunately, for students interested in an up-to-date account of the group, they are omitted from the recent monographic series on the “thigmotrichs” by Raabe, for the Polish authority judiciously considered them to be unrelated to any members of his order Thigmotricha-including the family Ancistrocomidae, which he did retain in the order (see 150 and 152, for his reason- ing).

NEW TAXA IN CILIATE CLASS KINETOFRACMOPHORA 2 15

Aneistrocomina n. subord.

Although these are the truly rostrate “rhynchodids” whose final taxonomic location may be out of the order ( a kind of etymologic contradiction, admittedly), they represent a significant assemblage of species. We owe knowledge of their existence primarily to 3 sets of investigators: the team of Chatton & Lwoff (29-31); Raabe, alone or sometimes with co-workers such as Jarocki (150- 154, and references therein to Raabe’s earlier publications) ; and Kozloff alone or, recently, with Lom (119-125, 127a, b ) . Practically the complete literature may be found in the combined bibliographies of the selected references just cited; but also see Canella (27) and Fenchel (82) . Recent ultrastructural works are 115, 127a, b, 141.

A sucking tentacle is located anteriorly in the ancistrocomines, and thigmotactic ciliature is often confined to the anterior part of essentially the ventral surface or is missing entirely from mature forms. There is characteristically a short trans- verse row of cilia in the lower right sector of the ventral surface (contrast this with condition in rhynchodines, described above).

More than 2 dozen genera have been described in the families Ancistrocomidae and Sphenophryidae. The species occur predominantly in or on bivalve molluscs; a few are associated with polychaete worms. Some of the better known ones belong to the following genera : Ancistrocoma, Crebricoma, Gargarius, Heterocineta, Holocoma, Hypocornella, Hypocomina, Lwoffia, Raabella, Sphenophrya, and Syringopharynx.

NOTES ADDED IN PRESS

After this paper was in press, the Editor kindly allowed insertion of 2 “Addenda” which may help clarify some of the complex problems pertinent to the present work, to other works still in press (e.g. 40, 142), and to the forthcoming treatises or monographs predicted in the introduction. The information in these addenda was not available a t the time of 1st preparation of the manuscript resulting in the present paper. Among other advantages to the reader is the opportunity to note the relation- ship of the class treated in preceding pages to the other 2 classes of the entire phylum (see 40) Ciliophora.

FIRST ADDENDUM

Although the multi-authored French paper ( 142) referred to in the present work allegedly preceded this one of mine, I did not see the manuscript while it was in preparation. Now that it is in press, I have had such an opportunity and wish to include here as Table 1 the overall scheme of classification suggested in it. I have made slight stylistic changes in order to set it up for easy comparison with my Table 2 (see below). Taxa named as new in the French work (142) are properly given here with “de Puytorac et al., 1974” as the authorship and date.

Space simply dors not permit further discussion of the text of de Puytorac et al. ( 142) ; many of its salient features have indeed already been mentioned or referred to in the text of this paper.

SECOND ADDENDUM

For the sake of comparison, I am including as Table 2 thc classification scheme suggested in my own major paper (40) in press (which is likely to turn out to precede both the present paper and the French work, 142, in actual date of appearance). One principal difference may be noted between Table 2 here and the Table 5 of Corliss (40) . Here I am listing as new taxa (or, at least, the names are new) those groups so treated in

TABLE 1 . Scheme of ciliate classification according to de Puytorac et al . (142).*

PHYLUM C I L I 0 P H 0 R A Doflein, 1901 Class I. KINETOFRAGMOPHORA de Puytorac et al., 1974

Subclass 1. Gymnostomata Biitschli, 1889 Order 1 ) PROSTOMATIDA Schewiakoff, 1896

Suborder (1) Archistomatina de Puytorac et al., 1974 ( 2 ) Prostomatina Schewiakoff, 1896

2) PLEUROSTOMATIDA Schewiakoff, 1896 Vestibulifera de Puytorac et al., 1974 Subclass 2.

Order 1 ) TRICHOSTOMATIDA Biitschli, 1889 2) COLPODIDA de Puytorac et al., 1974 3) ENTODINIOMORPHIDA Reichenow in Dof-

Subclass 3. Hypostomata Schewiakoff, 1896 emend. de

Nassulidea Jankowski, 1967

lein & Reichenow, 1929

Puytorac et al., 1974 Superorder [ 11

Order 1) SYNHYMENIDA de Puytorac et al., 1974 Suborder (1) NassuloDsina de Puvtorac et al.. 1974

( 2 ) Synhymenina de Pdytorac et al.: 1974 2) NASSULIDA Jankowski, 1967

Suborder ( 1 ) Nassulina Tankowski. 1967 ( 2 ) Microthoracina Jankowski, 1967

CYRTOPHORIDA FaurC-Fremiet in Corliss, 1956 2) CHONOTRICHIDA Wallengren, 1895

( 2 ) Endogemmina Jankowski, 1972 [3] Rhynchodea Chatton & Lwoff, 1939 RHYNCHODIDA Chatton & Lwoff, 1939

[4] Suctoridea Clapartde & Lachmann, 1858 SUCTORIDA Clapartde & Lachmann, 1858

[5] Apostomatidea Chatton & Lwoff, 1928 APOSTOMATIDA Chatton & Lwoff, 1928

[2] Phyllopharyngidea de Puytorac et al., 1974 Order 1 )

Suborder ( 1 ) Exogemmina Jankowski, 1972

Order 1 )

Order 1 )

Order 1 )

Class 11. OLIGOHYMENOPHORA de Puytorac et al., 1974 Subclass 1 . Hymenostomata Delage & HCrouard, 1896

Order 1 ) HYMENOSTOMATIDA Delage & HCrouard,

Suborder ( 1 ) Tetrahymenina FaurC-Fremiet in Corliss,

emend. de Puytorac et al., 1974

1896

1956 ( 2 ) O&-yoglenina Canella & Rocchi-Canella,

1964 ( 3 ) Peniculina FaurC-Fremiet in Corliss, 1956

( 1 ) Philasterina Small. 1967 2 ) SCUTICOCILIATIDA Small, 1967

( 2 ) Pleurostomatina FaurC-Fremiet in Corliss, 1956

3) ASTOMATIDA Schewiakoff, 1896 Subclass 2. Peritrieha Stein, 1859

Order 1) PERITRICHIDA Stein, 1859 Suborder ( 1 ) Sessilina Kahl, 1933

( 2 ) Mobilina Kahl, 1933

Class 111. POLYHYMENOPHORA Jankowski, 1967 Order 1) HETEROTRICHIDA Stein, 1859

Suborder ( 1 ) Heterotrichina Stein, 1859 f 2 ) Armoohorina Tankowski. 1964 ( 3 ) Coliphorina Jankowski, i967 ( 4 ) Licnophorina Corliss, 1957

2 ) ODONTOSTOMATIDA Sawaya, 1940 3) HYPOTRICHIDA Stein, 1859

( 1 ) Stichotrichina FaurC-Fremiet, 1961 ( 2 ) Sporadotrichina FaurC-Fremiet, 1961

( 1 1 Olieotrichina Biitschli. 1887 4 ) OLIGOTRICHIDA Biitschli, 1887

( 2 ) Tisinnina Kofoid &‘ Campbell, 1929

*With appropriate indication of names proposed as new in that work.

216 NEW TAXA IN CILIATE CLASS KINETOFRACMOPHORA

TABLE 2. Scheme of ciliate classification according t o Corliss (40) .*

PHYLUM C I L I 0 P H 0 R A Doflein, 1901 Class I. KINETOFRAGMOPHORA de Puytorac et al., 1974

Subclass 1. Gymnostomata Biitschli, 1889 Order 1) PRIMOCILIATIDA n. ord.

Suborder ( 1 ) Homokaryotina n. subord. (2) Karyorelictina n. subord. (3) Prostomatina Schewiakoff, 1896 ( 4 ) Prorodontina n. subord.

2) HAFTORIDA n. ord. 3) PLEUROSTOMATIDA Schewiakoff, 1896

Subclass 2. Vestibulifera de Puytorac et al., 1974 Order 1 ) TRICHOSTOMATIDA Biitschli, 1889

2) COLPODIDA de Puytorac et al., 1974 3) ENTODINIOMORPHIDA Reichenow in Dof-

lein & Reichenow, 1929 Suborder ( 1 ) Blepharocorythina n. subord.

& Reichenow, 1929 ( 2 ) Entodiniomorphina Reichenow in Doflein

Subclass 3. Hypostomata Schewiakoff, 1896 emend. de

Order 1 ) CYRTOPHORIDA Faur6-Fremiet in Corliss, Puytorac et al., 1974

1956 2) NASSULIDA Jankowski, 1967

Suborder (1) Nassulina Jankowski, 1967 ( 2 ) Paranassulina n. subord.

3) MICROTHORACIDA Jankowski, 1967 4) RHYNCHODIDA Chatton & Lwoff. 1939 ’

(1) Rhynchodina Chatton & Lwoff,’ 1939 ( 2 ) Ancistrocomina n. subord.

5) CHONOTRICHIDA Wallengren, 1895 (1) Exogemmina Jankowski, 1972 ( 2 ) Endogemmina Jankowski, 1972

6) APOSTOMATIDA Chatton & Lwoff, 1928

( 2 ) Ancistrocomina n. subord. 5) CHONOTRICHIDA Wallengren, 1895

(1) Exogemmina Jankowski, 1972 ( 2 ) Endogemmina Jankowski, 1972

6) APOSTOMATIDA Chatton & Lwoff, 1928

Subclass 4. Suctoria Clapar&de & Lachmann, 1858 Order 1) SUCTORIDA Claparede & Lachmann, 1858

( 2 ) Endogenina Collin, 19 12

Subclass 1. Hymenostomata Delage & HBrouard, 1896

Order 1 ) HYMENOSTOMATIDA Delage & HCrouard,

Suborder ( 1 ) Tetrahymenina Fa&-Fremiet in Corliss,

Suborder (1) Exogenina Collin, 1912

Class 11. OLIGOHYMENOPHORA de Puytorac et al., 1974

emend. de Puytorac et al., 1974

1896

1956 *---

( 2 ) Ophryoglenina Canella & Rocchi-Canella, 1964

3) ASTOMATIDA Schewiakoff, 1896

Subclass 2. Peritricha Stein, 1859 Order 1) PERITRICHIDA Stein, 1859

Suborder (1) Sessilina Kahl, 1933 (2) Mobilina Kahl, 1933

Class 111. POLYHYMENOPHORA Jankowski, 1967

Order 1) HETEROTRICHIDA Stein, 1859 Subclass 1. Spirotricha Biitschli, 1889

Suborder ( 1 ) Heterotrichina Stein, 1859 (2) Coliphorina Jankowski, 1967 (3) A.mophorina Jankowski, 1964 (4) Licnophorina Corliss, 1957

2) ODONTOSTOMATIDA Sawaya, 1940 3) OLIGOTRICHIDA Biitschli, 1887

(1) O!igotrichina Biitschli, 1887 ( 2 ) Tintinnina Kofoid & Campbell, 1929

4) HYPOTRICHIDA Stein, 1859

*With appropriate indication of names proposed as new in the present paper.

the present paper; whereas in Corliss (40), these will, of course, bear the authorship and year (“Corliss, 1974”) of the present note. The work by de Puytorac et al. (l42), and the present work as well, are thus to be considered, nomenclaturally, to have preceded the publication of my major paper (40) in Systematic Zoology. Without this word of clarification, the situation might have become historically confusing in the future, since all 3 works are appearing in the year 1974.

As is the case described in the preceding addendum, it is not appropriate here to discuss, or defend, conclusions drawn in my separate paper, even though it has supplied the data for the Table 2 of the present note. I t is also too late to insert references to Table 2 in preceding pages, helpful though this might have been for the reader. Fortunately, both tables represent self-contained units and are clear without specific comment in the text of the present paper.

Comparison of Tables 1 and 2 may be instructive. Areas of continuing controversy are easily identifiable, as are the many areas of essentially complete agreement.

REFERENCES 1. Agamaliev, F. G. 1966. New species of psammobiotic ciliates

of the western coast of the Caspian Sea [in Russian, English summary]. Acta Protozool. 4, 169-86.

2. - 1967. Faune des ciliCs mbsopsammiques de la c8te ouest de la mer Caspienne. Cah. Biol. Mar . 8, 359-402.

3 . - 1968. Materials on morphology of some psammophilic ciliates of the Caspian Sea. Acta Protozool. 6, 225-43.

4. - 1971. Complements to the fauna of psammophilic ciliates of the western coast of the Caspian Sea [in Russian, English summary]. Acta Protorool. 8, 379-404.

5. __ 1972. Ciliates from microbenthos of the islands of Apgeronskij and Bakinskij archipelagos of the Caspian Sea [in Russian, English summary]. Acta Protorool. 10, 1-27.

6. Bick, H. 1972. Ciliated Protozoa: A n Illustrated Guide to the Species Used As Biological Indicators in Freshwater Biology. World Health Organization, Geneva.

7. Bock, K. J. 1952. Uber einige holo- und spirotriche Ciliaten aus den marinen Sandgebieten der Kieler Bucht. Zool. Anz.

8. ___ 1952. Zur Ukologie der Ciliaten des marinen Sand- grundes der Kieler Bucht I. Kieler Meeresforsch. 9, 77-89.

9. - 1953. Zur Ukologie der Ciliaten des marinen Sand- grundes der Kieler Bucht 11. Kieler Meeresforsch. 9, 252-6.

10. Bohatier, J. 1970. Structure et ultrastructure de Lacry- maria olor (0. F. M. 1786). Protistologica 6, 331-42.

11. - 1972. Etude cytologique de la rCgCnCration de la rCgion buccale chez le ciliC Lacrymaria olor (0. F. M. 1786). Protistologica 8, 439-59.

12. Borror, A. C. 1963. Morphology and ecology of the benthic ciliated protozoa of Alligator Harbor, Florida. Arch. Pro- tistenk. 106, 465-534.

13. - 1965. New and little-known tidal marsh ciliates. Trans. Amer. Micros. SOC. 84, 550-65.

14. - 1968. Ecology of interstitial ciliates. Trans. Amer. Micros. SOC. 87, 233-43.

15. - 1972. Tidal marsh ciliates (Protozoa) : morphology, ecology, systematics. Acta Protorool. 10, 29-71.

16. - 1973. Tracheloraphis haloetes sp. n. (Ciliophora, Gymnostomatida) : description and a key to species of the genus Tracheloraphis. J. Protorool. 20, 554-8.

17. ~ 1973. Marine flora and fauna of the northeastern United States. Protozoa: Ciliophora. NOAA Tech. Rep. NMFS Circ. No. 378.

18. Bradbury, P. C. 1966. The fine structure of the mature tomite of Hyalophysa. J . Protorool. 13, 591-607.

19. - 1973. The fine structure of the cytostome of the apostomatous ciliate Hyalophysa chattoni. J . Protozool. 20, 405- 14.

20. - 1974. The fine structure of the phoront of the apostomatous ciliate, Hyalophysa chattoni. J . Protorool. 21, 1 12- 20.

21. Bradbury, P. C. & Pitelka, D. R. 1965. Observations on kinetosome formation in an apostome ciliate. J . Micros. 4, 805- 10.

149, 107-15.

NEW TAXA IN CILIATE CLASS KINETOFRAGMOPHORA 217

22. Burkovsky, I. V. 1970. The ciliates of the mesopsammon of the Kandalaksha Gulf (White Sea) I. [in Russian, English summary]. Acta Protozool. 7, 475-90.

23. - 1970. The ciliates of the rnesopsammon of the Kandalaksha Gulf (White Sea). 11. Acta Protozool. 8, 47-66.

24. Burreson, E. M. 1973. Symbiotic ciliates from solitary ascidians in the Pacific northwest, with a description of Parahy- pocoma rhamphisokarya n. sp. Trans. Amer. Micros. SOC. 92, 5 17-22.

25. Canella, M. F. 1960. Contributo ad una revisione dei generi Amphileptus, Hemiophrys e Lionotus. Ann. Univ. Ferrara (N. S. , Sect. 111) 2, 47-95.

26. - 1964. Strutture bucrali, infraciliature, filogenesi e sistematica dei ciliofora. Fatti, ipotesi, speculazioni. Ann. Univ. Ferrara (N. S. , Sect. 111) 2, 119-88.

27. __ 1971. Sur les organelles ciliaires de I’appareil buccal des hymbnostomes et autres ciliCs. Ann. Univ. Ferrara (N. S. , Sect. 111) 3 (Suppl.) (year 1970), 1-235.

28. Chatton, E. & Lwoff, A. 1935. Les ciliCs apostomes. I. Aperqu historique et gCnCral ; Ctude monographique des genres et des esp6ces. Arch. Zool. Exp. G i n . 77, 1-453.

29. ~ 1939. Sur la systCmatique de la tribu des thigmotriches rhynchoydes. Les deux familles des Hypocomidae Butschli et des Ancistrocomidae n. fam. Les deux genres nouveaux, Heterocoma et Parahypocoma. C . R . Acad. Sci. Paris 209, 429- 31.

30. ~ 1949. Recherches sur les ciliCs thigmotriches. I. Arch. Zool. Exp. Gkn. 86, 169-253.

31. ~ 1950. Recherches sur les ciliCs thigmotriches. 11. Arch 2001. Exp. Gkn. 86, 393-485.

32. Cheissin, E. M. & Poljansky, G. I. 1963. On the taxonomic system of Protozoa. Acta Protorool. 1, 327-52.

33. Corliss, J. 0. 1956. On the evolution and systematics of ciliated protozoa. Syst. Zool. 5, 68-91; 121-40.

34. - 1957. Nomenclatural history of the higher taxa in the subphylum Ciliophora. Arch. Protistenk. 102, 113-46.

35. - 1958. The phylogenetic significance of the genus Pseudomicrothorax in the evolution of the holotrichous ciliates. Acta Biol. Acad. Sci. Hung. 8, 367-88.

36. - 1961. The Ciliated Protozoa: Characterization, Classification, and Guide to the Literature. Pergamon Press, Lon- don and New York.

37. - 1969. The changing world of ciliate systematics. (Abstr.) 3rd Int. Cow. Protozool.. Lenincrad. Tulv 1969. Proe-

I . I , , I

;ess in Protozoology, p< 364-65. 38. __ 1972. The ciliate Protozoa and other organisms:

some unresolved questions of major phylogenetic significance. Amer. Zool. 12, 739-53.

39. - 1973. Evolutionary trends in patterns of stomatogenesis in the ciliate Protozoa. (Abstr.) J. Protozool. 20, 506.

40. - 1974. The changing world of ciliate systematics: historical analysis of past efforts and a newly proposed phylogenetic scheme of classification for the protistan phylum Cilioph- ora. Syst. Zool. 23, in press.

41. __ 1974. Time for evolutionary biologists to take more interest in protozoan phylogenetics?

42. Corliss, J. 0. & Esser, S. C. 1974. Comments on the role of the cvst in the life cvcle and survival of free-living Drotozoa. Trans.

’

Taxon 23, in press.

A m . Micros. SOC. 93, in press. 43. Deroux, G. 1970. La sCrie “chlamydonellienne” chez les

Chlamvdodontidae (holotriches. Cvrtouhorina FaurC-Fremiet) . I , .

Protistologica 6, 155-82. 44. Deroux, G. & Dragesco, J. 1968. Nouvelles donnCes sur

quelques ciliCs holotriches crytophores B ciliature ventrale. Pro- tistologica 4, 365-403.

45. Didier, P. & Bohatier, .I. 1970. Structures microfibrillaires chez le ciliC Lacrymaria olor (0. F. Miiller). C. R . Soc. Biol.

46. Dragesco, J. 1954. Diagnoses prkliminaires de quelques ciliCs psammophiles nouveaux. Bull. SOC. Zool. Fr. 74, 57-62.

47. ___ 1954. Diagnoses prCliminaires de quelques ciliCs nouveaux des sables. Bull. SOC. Zool. Fr. 74, 62-70.

48. ~ 1954. Diagnoses preliminaires de quelques cilids nouveaux des sables de Banyuls-sur-Mer. Vie et Milieu 4, 633-7.

49. -- 1960. CiliCs mksopsammiques littoraux. SystCma- tique, morphologie, Ccologie. Trav . Sta. Biol. Roscoff (N. S . )

50. - 1962. Capture et ingestion des proies chez les in-

51. __ 1962. On the biology of sand-dwelling ciliates.

164, 317-9.

12, 1-356.

fusoires ciliCs. Bull. Biol. Fr. Belg. 46, 123-67.

Sci. Progress 50, 353-63.

52. - 1963. RCvision du genre Dileptus, Dujardin 1871 (Ciliata Holotricha) (systkmatique, cytologie, biologie) . Bull. Btol. Fr. Belg. 97, 103-45.

53. - 1963. ComplCments B la connaissance des ciliks mCsopsammiques de Roscoff. I. Holotriches. Cah. Biol. Mar . 4, 91-1 19.

54. - 1963. ComulCments A la connaissance des ciliCs mesopsammiques de Roscoff. 11. HCtCrotriches. 111. Hypotriches. Cah. Biol. Mar . 4, 251-75.

55. - 1965. CiliCs mCsopsammiques d’Afrique Noire. Cah. Biol. Mar . 6. 357-99.

56. - 1966. Observations sur quelques ciliCs libres. Arch.

57. - 1966. Quelques ciliCs libres du Gabon. Biol. Gabon. Protistenk. 109, 155-206.

- - 2, 91-117.

58. - 1966. CiliCs libres de Thonon et ses environs. Protistologica 2, 59-95.

59. - 1970. Ciliks libres du Cameroun. Ann. Fac. Sci. YaoundC, YaoundC.

60. Dragesco, J., Auderset, G. & Baumann, M. 1965. Observa- tions sur la structure et la genkse des trichocystes toxiques et des protrichocystes de Dileptus (ciliCs holotriches) . Protistologica 1

61. Dragesco, J. & NjinC, T. 1971. Complements B la connaissance des ciliCs libres du Cameroun. Ann. Fac. Sci. Cameroun,

62. Dragesco, J. & Raikov, I. B. 1966. L’appareil nucldaire, la division et quelques stades de la conjugaison de Tracheloraphis margaritatus (Kahl) et T . caudatus sp. nov. (Ciliata, Holotricha). Arch. Protistenk. 109, 99-1 13.

63. Dumont, J. N. 1 9 6 1 . Observations on the fine structure of the ciliate Dileptus anser. J. Protozool. 8, 392-402.

64. Fad-Fremiet, E. 1950. Morphologie comparCe et sys- tCmatique des ciliCs. Bull. SOC. Zool. Fr. 75, 109-22.

65. - 1950. Ecologie des ciliCs psammophiles littoraux. Bull. Biol. Fr. Belg. 84, 35-75.

66. ~ 195 1. The marine sand-dwelling ciliates of Cape Cod. Biol. Bull. 100, 59-70.

67. - 1954. Reorganisation du type endomixique chez les Loxodidae ciliCs ct chez les Centrophorella. J . Protozool. 1,

68. - 1959. La famille des Nassulidae (Ciliata Gymnos- tomatida) et le genre Nassulopsis n. gen. C . R . Acad. Sci. Paris

69. - 196 1. Quelques considCrations sur les ciliCs mesopsammiques B propos d’un recent travail de J. Dragesco. Cah. Biol. Mar. 2, 177-86.

70. - 1961. Les ciliCs Cyrtophorina et leur diversifica- tion morphologique. C . R . Acad. Sci. Paris 252, 3912-6.

7 1 . - 1962. Le genre Paranassula Kahl (Ciliata Cyrto- phorina). Cah. Biol. Mar. 3, 61-77.

72, - 1963. Remarques sur la morphologie comparke des ciliks Trichostomatida. C . R . Acad. Sci. Paris 257, 3089-94.

73. - 1965. Morphologie des Dysteriidae (Ciliata Cyrto- phorina) . C. R . Acad. Sci. Paris 260, 6679-84.

74. - 1967. La frange ciliaire des Nassulidae (Ciliata Cyrtophorina) et ses possibilitCs Cvolutives. C. R . Acad. Sci. Paris

75. - 1967. Le genre Cyclogramma Perty 1852. J . Pro-

76. - 1970. A propos de la note de M. Thomas NjinC sur le ciliC Loxodes magnus. C. R . Acad. Sci. Paris 270, 523-4.

7 7 . Faurk-Fremiet, E. & Andre, J. 1965. L’organisation du ciliC gymnostome Plagiocampa ovata Gelei. Arch. Zool. Exp. G i n . 105, 361-8.

78. - 1968. L’organisation corticale des Ciliata. C. R . Acad. Sci. Paris 266, 487-90.

79. FaurC-Fremiet, E., AndrC, J. & Ganier, M.-C. 1968. Cal- cification tCgumentaire chez les ciliCs du genre Coleps Nitzsch. J. Micros. 7, 693-702.

80. Fad-Fremiet, E. & Ganier, M.-C. 1969. Structure fine de la zone buccale chez le ciliC Chaenea vorax. C . R . Acad. Sci. Paris 268, 813-5.

81. - 1969. Morphologie et structure fine du ciliC Chaenea vorax Quenn. Protistologica 5, 353-61.

82. Fenchel, T. 1965. Ciliates from Scandinavian molluscs. Ophelia 2, 71-174.

83. ~ 1968. The ecology of marine microbenthos. 11. The food of marine benthic ciliates. Ophelia 5, 73-121.

84. ~ 1969. The ecology of marine microbenthos. IV. Structure and function of the benthic ecosystem, its chemical

( 2 ) , 81-90.

NO. 7-8, 97-140.

20-7.

249, 1429-33.

264, 68-72.

tozool. 14, 456-64.


and physical factors and the microfauna communities with special reference to the ciliated protozoa. Ophelia 6, 1-182.

85. Fjeld, P. 1955. On some marine psammobiotic ciliates from Drebak (Norway). Nytt Mag. Zool. 3, 5-65.

86. Gelei, J. von 1952. Einiges iiber die Systematik der Un- terordnung Trichostornata der Ciliaten [in Hungarian, German summaryl. Ann. Biol. Univ. Hung. 1, 351-60.

87. Gelei, J. von & SebestyCn, 0. 1932. Einige Bernerkungen zum Bau und Funktion der Syncilien bei den Darmciliaten, be- sonders der Entodiniomorpha. Acta Biol. Szeged. 2, 141-61.

88. Golihska, K. 1972. Studies on stomatogenesis in Dileptus (Ciliata, Holotricha) in the course of division processes. Acta Protozool. 9, 283-97.

89. Golihska, K. & Grain, J. 1969. Observations sur les modi- fications ultrastructurales lors de la rCgCnCration chez Dileptus cygnus Clap. et Lach., 1859, ciliC holotriche gymnostome. Pro- tistologica 5, 447-64.

90. Grain, J. 1966. Etude cytologique de quelques ciliCs holotriches endocommensaux des ruminants et des CquidCs. Protisto- logica 2(1), 59-141; 2 ( 2 ) , 5-51.

91. - 1969. Le cinktosome et ses dCrivCs chez les ciliCs. Ann. Biol. 8, 53-97.

92. - 1970. Structure et ultrastructure de Lagynophrya fusidens Kahl, 1927. Protistologica 6, 37-51.

93. Grain, J. & Golihska, K. 1969. Structure et ultrastructure de Dileptus cygnus Claparhde et Lachmann, 1859, ciliC holotriche gymnostome. Protistologica 5, 269-90.

94. Grain, J., Puytorac, P. de & Bohatier, J. 1973. Essai de systCmatique des ciliCs gymnostomes fondCe sur les caractkristiques de I’infraciliature circumorale. C . R . Acad. Sci. Paris 277, 69-72.

95. Grell, K. G. 1962. Morphologie und Fortpflanzung der Protozoen (einschliesslich Entwicklungsphysiologie und Genetik) . Fortsch. Zool. 14, 1-85.

96. ~ 1964. The protozoan nucleus, in Brachet, J. & Mirsky, A. E., eds., T h e Cell, Academic Press, New York and London, 6, 1-79.

97. ~ 1973. Protozoology. Springer-Verlag, Berlin, Heidel- berg, and New York.

98. Hartwig, E. 1973. Die Ciliaten des Gezeiten-Sandstrandes der Nordseeinsel Sylt. I. Systematik. Akad. Wiss. Li t . (Mainz) Math. Naturwiss. K1. Mikrofauna Meeresbodens 18, 387-453.

99. Hausmann, E. & Hausmann, K. 1973. Cytologische Studien an Trichocysten. VII. Die Feinstruktur der Trichocystenwanen von Loxophyllum meleagris. Protistologica 9, 139-47.

100. Hausmann, K. & Wohlfarth-Bottermann, K. E. 1974. Cytologische Studien an Trichocysten. VIII. Feinstruktur und Funktionsweise der Toxicysten von Loxophyllum meleagris und Prorodon teres. Protistologica 10, in press.

101. Holt, P. A. & Chapman, G. B. 1971. The fine structure of the cyst wall of the ciliated protozoon Didinium nasutum. J. Protozool. 18, 604-14.

102. Holt, P. A. & Corliss, J. 0. 1973. Pattern variability in microtubular arrays associated with the tentacles of Actinobolina (Ciliata: Gymnostomatida). J . Cell Biol. 58, 21 3-9.

103. Holt, P. A., Lynn, D. H. & Corliss, J. 0. 1974. An ultrastructural study of the tentacle-bearing ciliate Actinobolina smalli n. sp. and its systematic and phylogenetic implications. Protistologica 9 (year 1973), in press.

104. Holt, P. A., Maugel, T. K. & Corliss, J. 0. 1973. Micro- tubular substructure in a ciliated protozoon. Trans. Amer. Micros. SOC. 92, 503-7.

105. 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.

106. Hsiung, T. S. 1929. A survey of the protozoan fauna of the large intestine of the horse. (Abstr.) J. Parasit. 16, 99.

107. __ 1930. A monograph on the protozoa of the large intestine of the horse. Iowa St . Coll. 1. Sci. 4, 356-423.

108. Jankowski, A. W. 1967. A new system of ciliate Pro- tozoa (Ciliophora) [in Russian]. Akad. Nauk SSSR, Trudy Zool. Inst. 43, 3-54.

109. - 1968. Morphology and phylogeny of Crateristoma (Ciliophora: Rynchodida) [in Russian, English summaryl. Hy- drobiol. J . 4, 33-41.

110. - 1973. Taxonomic revision of subphylum Cilioph- ora Doflein, 1901 [in Russian, English summaryl. Zool. Zh.

111. - 1973. Fauna of the USSR: Infusoria subclass Chonotricha [in Russian], Vol. 2, No. 1. USSR Akad. Nauk, Nauka Publishing House, Leningrad. 355 pp.

52, 165-75.

112. Kahl, A. 1930-1935. Urtiere oder Protozoa: I: Wimper- tiere oder Ciliata (Infusoria) , eine Bearbeitung der freilebenden und ectocommensalen Infusorien der Erde, unter Ausschluss der marinen Tintinnidae, in Dahl, F., ed., Die Tierwelt Deutschlands, G. Fischer, Jena, parts 18(year 1930), 21(1931), 25(1932),

113. ~ 1933. Ciliata libera et ectocommensalia, in Grimpe, G. & Wagler, E., Die Tierwelt der Nord- und Ostsee, Lief. 23 (Teil 11, ca), Leipzig, pp. 29-146.

114. Kattar, M. R. 1972. Quelques aspects de I’ultrastructure du cilik Pseudoprorodon arenicola Kahl, 1930. Protistologica 8,

115. Khan. M. A. 1969. Fine structure of Ancistrocoma bel-

30( 1935), pp. 1-886.

135-41.

seneeri (Chatton et Lwoff) , a rhynchodine thigmotrichid cil&te. Acta Protozool. 7, 29-47.

116. Kink, J. 1972. Observations on morDholoav and oral morphogeneds during regeneration of ciliate Lacrymaria olor (0. F.M. 1786) (Holotricha, Gymnostomatida) . Acta Protozool.

117. - 1973. The organization of fibrillar structures in the trophic and encysted Dileptus visscheri (Ciliata, Rhabdo- phorina). Acta Protozool. 12, 173-94.

118. KovalEva, V. G. & Raikov, I. B. 1972. Saccules Cnig- matiques en soucoupes” et leur relation avec les protrichocystes chez le ciliC holotriche Trachelonema sulcata Kovaleva. Protis- tologica 8, 413-25.

119. Kozloff, E. N. 1946. Studies on ciliates of the family Ancistrocomidae Chatton and Lwoff (order Holotricha, suborder Thigmotricha). I. Hypocomina tegularum sp. nov. and Cre- bricoma gen. nov. Biol. Bull. 90, 1-7.

120. - 1946. Studies on ciliates of the family Ancistro- comidae Chatton and Lwoff (order Holotricha, suborder Thig- motricha). 11. Hypocomides mytili Chatton and Lwoff, Hypo- comides botulae sp. nov., Hyqocomides parva sp. nov., Hypoco- mides kelliae sp. nov., and Instgnicoma venusta gen. nov., sp. nov. Biol. Bull. 90, 200-12.

121. - 1946. Studies on ciliates of the family Ancistro- comidae Chatton and Lwoff (order Holotricha, suborder Thig- motricha) . 111. Ancistrocoma pelseneeri Chatton and Lwoff, Ancistrocoma dissimilis sp. nov., and Hypocomagalma pholadidis sp. nov. Biol Bull. 91, 189-99.

122. - 1946. Studies on ciliates of the family Ancistro- comidae Chatton and Lwoff (order Holotricha, suborder Thig- motricha) . IV. Heterocineta janickii Jarocki, Heterocineta goni- obasidis sp. nov., Heterocineta fluminicolae sp nov., and Enerthe- coma properam Jarocki. Biol. Bull. 91, 200-9.

123. - 1955. Lwoffia cilifera gen. nov., sp. nov., a ciliated member of the family Sphenophryidae (Holotricha: Thig- motricha). Biol. Bull. 108, 283-9.

124. - 1961. A new genus and two new species of ancistrocomid ciliates (Holotricha: Thigmotricha) from sabellid polychaetes and from a chiton. 1. Protozool. 8, 60-3.

125. - 1965. Colligocineta furax gen. nov., sp. nov., an ancistrocomid ciliate (Holotricha : Thigmotricha) from the sabellid polychaete Laonome kroeyeri Malmgren. J . Protozool. 12, 333-4.

126. Lom, J. & Corliss, J. 0. 1971. Morphogenesis and cortical ultrastructure of Brooklynella hostilis, a dysteriid ciliate ecto- parasitic on marine fishes. J . Protozool. 18, 261-81.

127a. Lom, J. & Kozloff, E. N. 1968. Observations on the ultrastructure of the suctorial tube of ancistrocomid ciliates. Folia Parasit. (Praha) 15, 291-308.

127b. - 1969. Ultrastructure of the cortical regions of ancistrocomid ciliates. Protistologica 5 , 173-92.

128. Mohr, J. L., Matsudo, H. & Leung, Y.-M. 1970. The ciliate taxon Chonotricha. Oceanogr. Mar. Biol. Ann. Rev. 8,

129. NjinC, T. 1970. Structure et morphogknbe buccales chez le ciliC holotriche Loxodes magnus Stokes 1887. C . R. Acad. Sci. Paris 270, 519-28.

130. Nobili, R. 1957. Contributo all’ecologia dei ciliati psam- mofili del Golfo di Napoli. Boll. Zool. 24, 211-25.

131. Noirot-Timothke, C. 1960. Etude d’une famille de ciliks: les Ophryoscolecidae. Structures et ultrastructures. Ann. Sci. Nut. Zool. (sCr. 12) 2, 527-718.

132. - 1969. Les affinitks des Entodiniomorphida (Cili- ata). (Abstr.) 3rd Int. Cong. Protozool., Leningrad, July 1969, Progress in Protozoology, p. 371.

133. Pennak, R. W. 1951. Comparative ecology of the interstitial fauna of freshwater and marine branches. Ann. Biol. 27,

10, 205-13.

415-56.

449-80.

NEW TAXA IN CILIATE CLASS KINETOFRAGMOPHORA 219

134. Petran, A. 1963. Contributii la cunoasterea microfaunei de ciliate psamofile din marea Neagra (littoralul romlnesc) . Stud. Cercet. Biol. ( A n i m . ) 15, 187-97.

135. __ 1967. Cercetzri asupra faunei de ciliate psamo- bionte la plajele din sudul litoralului romlnesc el Mlrri Negrc. Ecol. Mar. 2, 169-91.

136. ~- 1968. Les ciliCs mCsopsammiques de Mangalia et quelques considCrations sur la faune infusorienne des sables du littoral roumain de la mer Noire. Rapp. Comm. Znt. Mer. Medit .

137. Poljansky, G. I. & Cheissin, E. M. 1965. Dogiel‘s General Protozoology. Oxford University Press, London and New York.

138. Poljansky, G. I. & Raikov, I. B. 1961. Nature et origine du dualisme nuclCaire chez les infusoires ciliCs. Bull. SOG. 2001. Fr. 86, 402-1 1 .

139. Puytorac, P. de 1964. Quelques aspects de I’ultrastructure du ciliC: Prorodon viridis Ehrbg. Kahl. Acta Protozool. 2,

140. ~ 1965. Sur I’ultrastructure du complexe cytostome-cytopharynx chez le ciliC Holophrya vesiculosa Kahl. C . R . Soc. Biol. 159, 661-3.

141. ~ 1969. Remaroues 2 DroDos de I’ultrastructure du

19, 175-7.

147-52.

sucoir d’Ancistrocoma myae (K. et B.),‘ ciliC Rhynchodea. C . R . Acad. Sci. Paris 268, 820-2.

142. Puytorac, P. de, Batisse, A., Bohatier, J., Corliss, J. O., Deroux, G., Didier, P., Dragesco, J., Fryd-Versavel, G., Grain, J., Grolikre, C. A,, Hovasse, R., Iftode, F., Laval, M., Roque, M., Savoie, A. & Tuffrau, M. 1974. Proposition d’une classification des ciliCs. C . R . Acad. Sci. Paris 276. in press.

143. Puytorac, P. de & Grain, 1: 1972. Bactkries intramito- chondriales et particularitCs de I’ulirastructure cytosomo-pharyn- gienne chez le cilik Urotricha ovata Kahl. C . R . SOG. Biol. 166, 604-15.

144. Puvtorac. P. de & Kattar. M. R. 1969. Observations sur I’ultrastruiture du ciliC Helicoprorodon multinucleatum Dragesco 1960. Protistologica 5, 549-60.

145. Puytorac, P. de & NjinC, T. 1970. Sur I’ultrastructure des Loxodes (ciliCs holotriches trichostomes) . Protistologica 6, 427-44.

146. Puytorac, P. de & Savoie, A. 1968. Observations cytologi- ques et biologiques sur Prorodon palustsic nov. sp. Protistologica 4, 53-60.

147. Pyne, C. K. & Tuffrau, M. 1970. Structure et ultrastructure de I’appareil cytopharyngien et des tubules complexes en relation avec celui-ci chez le ciliC gymnostome Chilodonella unci- nata Ehrbg. 1. Micros. 9, 503-16.

148. Raabe, Z. 1964. Remarks on the principles and outline of the system of Protozoa. Acta Protozool. 2, 1-18.

149. ~ 1964. Zarys Protoroologii [in Polish]. Polish Sci- entific Publs., Warsaw.

150. ~ 1967. Ordo Thigmotricha (Ciliata-Holotricha) . I. Acta Protozool. 5, 1-36.

151. - 1970. Ordo Thigmotricha (Ciliata-Holotricha) . 11. Familia Hemispeiridae. Acta Protozool. 7, 117-80.

152. ~ 1970. Ordo Thigmotricha (Ciliata-Holotricha) . 111. Familiae Ancistrocomidae et Sphenophryidae. Acta Prototool.

153. ___ 197 1 . Ordo Thigmotricha (Ciliata-Holotricha) . IV. Familia Thigmophryidae. Acta Protozool. 9, 121-70.

154. - 1972. Ordo Thigmotricha (Ciliata-Holotricha) . V. Familiae Hysterocinetidae et Protoanoplophryidae. Acta Proto-

155. Raikov, I. B. 1957. Nuclear apparatus and its reorganization during the fission cycle in the infusoria Trachelocerca mar- garitata (Kahl) and T . dogieli, sp. n. (Holotricha) [in Russian, English summary]. Zool. Zh. 36, 344-59.

156. ~ 1957. Reorganization of the nuclear apparatus in ciliates and the problem of the origin of their binuclearity [in Russian]. Vest. Leningrad Univ. , No. 15, 21-37.

157. -~ 1958. Der Formweschsel des Kernapparates eini- ger niederer Ciliaten. I. Die Gattung Trachelocerca. Arch. Pro- tistenk. 103, 129-92.

158. - 1959. Der Formweschsel des Kernapparates eini- ger niederer Ciliaten. 11. Die Gattung Loxodes. Arch. Protis- tenk. 1M, 1-42.

159. - 1959. Cytological and cytochemical peculiarities of the nuclear apparatus and division in the holotrichous ciliate Geleia nigriceps Kahl [in Russian]. Tsitologia 1, 566-79.

160. - 1960. The interstitial infusoria of the sandy littoral regions of the Dal’Nezelenetskaya Bay (eastern Muran). Murmanskogo Morskogo Biol. 2 , 172-85.

7, 385-462.

2001. 10, 115-84.

161. - 1962. Les ciliCs mksopsammiques du littoral de la Mer Blanche (U.R.S.S.) avec une description de quelques esptkes nouvelles ou peu connues. Cah. Biol. Mar . 3, 325-61.

162. ~ 1963. Nuclear apparatus and division of Rema- nella granulosa Kahl and R . rugosa Kahl (Holotricha, Gymnosto- matida) [in Russian], in Poljansky, G. I., Sukhanova, K. M. & Cheissin, E. M., eds., Morphology and Physiology of Protozoa. Moscow and Leningrad, pp. 20-34.

163. - 1963. The nuclear apparatus of Ramanella mul- tinucleata Kahl (Ciliata, Holotricha). Acta Biol. 14, 221-9.

164. ~ 1963. The nuclear apparatus of the holotrichous ciliates Geleia orbis FaurC-Fremiet and G . murmanica Raikov [in Russian, English summary]. Acta Protozool. 1, 21-9.

165. ___ 1963. Ciliates of the mesopsammon of the Ussuri Gulf (Japan Sea) [in Russian, English summaryl. Zool. Z h . 42,

166. ___ 1963. On the origin of nuclear dualism in ciliates. 1st Int. Cong. Protozool., Prague, Aug. 1961, Progress in

167. - 1964. DNA content of the nuclei and nature of macronuclear chromatin strands of the ciliate Nassulopsis elegans (Ehrbg.). Acta Prototool. 2 , 339-57.

168. ~ 1965. New data on polyploid nuclei in Protozoa [in Russian], in Polyploidy i Selekcija CProc. Conf. Polyploidy Leningrad, 19631, Moscow, Leningrad, pp. 134-42.

169. - 1966. Elektronemikroskopische Untersuchung des Kernapparates von Nassula ornata Ehrbg. (Ciliata, Holotricha) . Arch. Protistenk. 109, 71-98.

170. ~ 1969. The macronucleus of ciliates, in Chen, T.- T., ed., Research in Protozoology, Pergamon Press, London and New York, 3, 1-128.

1 7 1 . ~ 1972. The nuclear apparatus of the psammophilic ciliate Kentrophoros fistulosum FaurC-Fremiet : structure, divisional reorganization and ultrastructure. Acta Protorool. 10,

172. - 1972. Ultrastructures macronuclCaires et micro- nuclCaires de Tracheloraphis dogieli, ciliC marin B macronoyaux diploi‘des. C . R . Soc. Biol. 166, 608-13.

173. ___ 1973. Mitose intranuclCaire acentrique du micro- noyau de Loxodes magnus, ciliC holotriche. Etude ultrastructurale. C . R . Acad. Sci. Paris 276, 2385-8.

174. - 1973. Ultrastructures cytoplasmiques de Kentro- phoros latum Raikov, ciliC holotriche psammophile: organisation corticale, endoplasme et trichocystes. Ann. Stat. Biol. Besse-en- Chandesse 8, 21-53.

175. - 1974. Fine structure of the nuclear apparatus of a lower psammobiotic ciliate, Tracheloraphis dogieli Raikov. d c t a Protozool. 13, in press.

176. Raikov, I. B., Cheissin, E. M. & Buze, E. G. 1963. A photometric study of DNA content of macro- and micronuclei in Paramecium caudatum, Nassula ornata, and Loxodes magnus. Acta Protozool. 1, 285-300.

177. Raikov, I. B. & Dragesco, ,I. 1969. Ultrastructure des noyaux et de quelques organites cytoplasmiques du ciliC Trache- loraphis caudatus Dragesco et Raikov (Holotricha, Gymnostoma- tida) . Protistologica 5, 193-208.

178. Raikov, I. B. & Kovaleva, V. G. 1968. Complements to the fauna of psammobiotic ciliates of the Japan Sea (Posjet Gulf). Acta Protozool. 6, 309-33.

179. Remane, A. 1952. Die Besiedelung des Sandbodens im Meere und die Bedeutung der Lebensformtypen fur die Okologie. Verh. Deutsch. Zool. Ges., pp. 327-59.

180. Rieder, N. 197 1. Elektronenoptische Untersuchungen an Didinium nasutum 0. F. Muller (Ciliata, Gymnostomata) in In- terphase und Teilung. Forma et Functio 4, 46-86.

181. Sauerbrey, E. 1928. Beobachtungen uber einiae neue oder wenig bekannte marine Ciliaten. Arch Protistenk. 62, 355- 407.

182. Sleigh, M. A. 1973. T h e Biology of Protozoa. Edward Arnold, London.

183. Small, E. B. 1967. The Scuticociliatida, a new order of the class Ciliatea (phylum Protozoa, subphylum Ciliophora). Trans A m . Micros. SOG. 86, 345-70.

184. Small, E. B., Antipa, G. A. & Marszalek, D. S . 1972. The cytological sequence of events in the binary fission of Didin- ium nasutum. Acta Protorool. 9, 275-81.

185. Soltyhska, M. S. 1971. Morphology and fine structure of Chilodonella cucullulus (O.F.M. ) . Cortex and cytopharyngeal apparatus. Acta Protozool. 9, 49-82.

186. Swedmark, B. 1964. The interstitial fauna of marine sand. Biol. R e v . 39, 1-42.

1753-67.

Protozoology, pp. 253-8.

227-47.


187. Torch. R. 1964. Autoradioarauhic studies of nucleic acid synthesis in a- gymnostome ciliate, pracheloraphis sp. (Abstr.) J . Cell Biol. 23, 98A.

188. Tucker, J. B. 1968. Fine structure and function of the cytopharyngeal basket in the ciliate Nassula. J . Cell Sci. 3, 493- 514.

189. - 1970. Morphogenesis of a large microtubular organelle and its association with basal bodies in the ciliate Nassula. J . Cell Sci. 6, 385-429.

190. - 1970. Initiation and differentiation of microtubule patterns in the ciliate Nassula. J . Cell Sci. 7 , 793-821.

191. - 1971. Development and deployment of cilia, basal bodies, and other microtubular organelles in the cortex of the ciliate, Nassula. J . Cell Sci. 9, 539-67.

192. - 1972. Microtubule-arms and propulsion of food particles inside a large feeding organelle in the ciliate Phascolo- don vorticella. J . Cell Sci. 10, 883-903.

193. Tuffrau, M. 1963. Les structures infraciliaires et la sto- matogknbse chez les Loxodes. (Abstr.) 1st Int. Gong. Protozool., Prague, Aug. 1961, Progress in Protozoology, pp. 278-80.

194. Vinnikova, N. V. 1974. Fine structural changes of the macronuclei of Dileptus anser during conjugation. Acta Proto- zool. 13, in press.

195. Wessenberg, H. & Antipa, G. A. 1968. Studies on Didin- ium nasutum. I. Structure and ultrastructure. Protistologica 4, 427-47.

196. - 1970. Capture and ingestion of Paramecium by Didinium nasutum. J . Protozool. 17, 250-70.

197. Wolska, M. 1966. Study on the family Blepharocory- thidae Hsiung. I. Preliminary remarks. Acta pro to zoo^. 4,

198. - 1967. Study on the family Blepharocorythidae Hsiung. 11. Charonina uentriculi (Jameson). Acta Protozool.

199. - 1967. Study on the family Blepharocorythidae Hsiung. 111. Raabena bella gen. n., sp. n. from the intestine of the Indian elephant. Acta Protozool. 4, 285-90.

200. - 1968. Study on the family Blepharocorythidae Hsiung. IV. Pararaabena dentata gen. n., sp. n. from the intestine of Indian elephant. Acta Protozool. 5, 219-24.

201. - 1971. Studies on the family Blepharocorythidae Hsiung. V. A review of genera and species. Acta Protozool. 9,

202. ~ 197 1. Studies on the family Blepharocorythidae Hsiung. VI. Phylogenesis of the family and the description of the new genus Circodinium gen. n. with the species C. minimum (Gassovsky, 1918). Acta Protozool. 9, 171-94.

203. Yagiu, R. & Shigenaka, Y. 1965. Electron microscopy of the ectoplasm and the proboscis in Didiniurn nasutum. J . Proto-

97-104.

4, 279-84.

23-39.

~ 0 0 1 . 12, 363-81.

BOOK REVIEW. . . Grell, K. G. 1973. Protozoology. Springer-Verlag, Berlin and Heidelberg, Germany. viii + 554 pp. $43.90.

This volume started to be an English translation from the German of the 2nd edition of Grell’s Protozoologie, but 5 years had elapsed since that book appeared so enlargement was unavoidable. Many figures have been changed or added, but the beautiful illustrations of the previous editions remain.

The book will repay anyone who reads it by its fine general discussions. Taxonomy is not stressed. There are chapters on the subject nature (10 pages), structure (110 pages), reproduc- tion (31 pages), fertilization and sexuality (76 pages), altema- tion of generations ( 3 pages), heredity (45 pages), motility (36 pages), behavior (15 pages), nutrition (28 pages), and parasitism and symbiosis (15 pages). These are followed by a taxonomic survey 103 pages long. There is a short list of 14 “summary presentations,” followed by 1845 references to separate works and works from associated fields. Most useful is a list of 57 films on protozoa. The great majority are in German and some are in French. None is in English, but the pictures are more important than the words, so little would be lost by

an English-speaking audience. There is a 5%-page subject index and a 6%-page list of genera and species.

Grell is not a taxonomist or a parasitologist, and his discussions in these areas leave something to be desired. The classification that he uses is his own and sadly dated. He does not even mention the modem Society of Protozoologists’ classification, and he completely omits important genera, such as Sarco- cystis and Leucocytozoon. He includes the trypanosomes in the order Protomonadina, not mentioning the Kinetoplastida and ignoring the Hoare-Wallace names for the various stages. He lumps all trichomonads together as Trichomonas. He says of Toxoplasma, “Electron microscopic investigations suggest that Toxoplasma gondii is a coccidian. There is some-though not yet enough-evidence that it is related to Isospora.”

These strictures sound formidable, but the book should not be denigrated as severely as they might suggest. For the dis- criminating protozoologist it contains a great deal of valuable information, while for the nonprotozoologist it is an enticing introduction to the phylum. I cannot, however, resist mentioning that, at $43.90, it costs 7.9 cents a page.-NORMAN D. LEVINE, College of Veterinary Medicine, Univ. of Illinois, Urbana, IL 61801, USA.

Documents

Remarks on the Composition of the Large Ciliate Class Kinetofragmophora de Puytorac et al., 1974, and Recognition of Several New Taxa Therein, with Emphasis on the Primitive Order