Establishment of a New Family (Glaucomidae n. fam.) in the Holotrich Hymenostome Ciliate Suborder Tetrahymenina, and Description of a New Genus (Epenardia n. g.) and a New Species

Establishment of a New Family (Glaucomidae n. fam.) in the Holotrich Hymenostome CiliateSuborder Tetrahymenina, and Description of a New Genus (Epenardia n. g.) and a New Species(Glaucoma dragescui n. sp.) Contained ThereinAuthor(s): John O. CorlissSource: Transactions of the American Microscopical Society, Vol. 90, No. 3 (Jul., 1971), pp.344-362Published by: Wiley on behalf of American Microscopical SocietyStable URL: http://www.jstor.org/stable/3225195 .

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ESTABLISHMENT OF A NEW FAMILY (GLAUCOMIDAE N. FAM.) IN THE HOLOTRICH HYMENOSTOME CILIATE SUBORDER

TETRAHYMENINA, AND DESCRIPTION OF A NEW GENUS (EPENARDIA N. G.) AND A NEW SPECIES (GLAUCOMA

DRAGESCUI N. SP.) CONTAINED THEREIN1

JOHN O. CORLISS

Department of Zoology, University of Maryland, College Park, Maryland 20742

CORLISS, J. 0. 1971. Establishment of a new family (Glaucomidae n. fam.) in the holotrich hymenostome ciliate suborder Tetrahymenina, and description of a new genus (Epenardia n. g.) and a new species (Glaucoma dragescui n. sp.) contained therein. Trans. Amer. Micros. Soc., 90: 344-362. As the result of accumulation of new data, generally resulting from application of new techniques, and appearance of new ideas concerning ciliate systematics in general, taxonomic revisions have often been necessitated at various levels within the subphylum. In the present case, greater and more precise knowledge about hymenostome holotrichs than was possible in the past has demanded revision of the tetrahymenine family Tetrahymenidae, rapidly becoming too heterogeneous. The part of the revision specifically treated here includes the establishment and characterization of the Glaucomidae n. fam.; of Epenardia n. g., with type-species E. myriophylli (Penard, 1922) n. comb.; and of Glaucoma dragescui n. sp., new member of Glaucoma, type-genus of the new family. The new family Glaucomidae is described as containing four bonafide genera (Glaucoma, the new genus Epenardia, and the taxonomically controversial Espejoia and Turaniella) and four genera of less certain status (Dichilum, Monochilum, Physalophrya, and Pinchatia). The genus Glaucoma, in revision, is reduced to eight "good" species: the type-species, G. scintillans; the new species, G. dragescui; and G. avellana, G. chattoni, G. frontata, G. intermedia, G. kirki, and G. macrostoma.

The combined impact of new ideas and new techniques has indeed, in very recent years, brought us into a "changing world of ciliate systematics" (Corliss, 1969). There is still much work to be done at the alpha taxonomic level (and the present paper represents a small example of this), but even in such a basic, purely descriptive type of investigation the influence of new or revised hypotheses, generally stemming from synthetic approaches to analyses of freshly accumulated data, is inevitable and, of course, desirable. In the case of the hymenostome holotrichs, to which the familial groups under present consideration belong, recent literature of particular theoretical importance must include, at the very least, the

following papers2: Canella (1964, 1970), Corliss (1968), Faure-Fremiet (1967), Jankowski (1967a), and Small (1967).

Following recognition of the pronounced heterogeneity of Kahl's (1926, 1931) hymenostome family Frontoniidae and the need for division of the order itself into at least three suborders (see Corliss, 1961), the family Tetrahymenidae Corliss, 1952, within the suborder Tetrahymenina, began to receive special taxo-

1 Support of National Science Foundation grant GB 27050X in preparation of this paper is gratefully acknowledged. Among several colleagues to whom I have been indebted over the years for exchange of both pertinent data and ideas, Dr. E. Faure-Fremiet, now Professor- Emeritus of the College de France, Paris, and my "Patron" ever since my acceptance of his kind invitation to spend a postdoctoral year in his laboratory in 1951-52, deserves my deepest gratitude and thanks for his continuous interest, aid, and inspiration during the past two decades.

2 Exclusive of additional papers, by the authors listed or still other workers, now known, by personal communication, to be in final stages of preparation for publication in various appropriate journals. TRANS. AMER. MICROS. Soc., 90(3): 344-362. 1971.



CORLISS-NEW FAMILY OF HYMENOSTOME CILIATES 345

nomic and nomenclatural attention (e.g., see the works of Corliss & Dougherty, 1967; Czapik, 1968; Jankowski, 1964, 1967b; Thompson, 1963). Most often, however, only individual genera or species (especially of the type-genus Tetrahymena: see review in Corliss, 1971) have been receiving consideration, in works generally related to other overall matters (e.g., see Dragesco, 1970, a book on the free-living ciliates of the Cameroons). Perhaps inevitably, the Tetrahymenidae began to encompass a growing number of superficially similar genera, becoming too heterogeneous and approaching the vast and unwieldy size of the Frontoniidae of the 1930's. Combining the genera from lists included in Corliss (1961), Czapik (1968), and Thompson (1963), and adding a number of "uncertain" or recently described new genera with reasonable claim to inclusion in the same familial group, the family might appear to be obliged to contain the following 27 genera: Blepharo- stoma, Cardiostomatella, Chasmatostoma, Cinetochilum, Colpidium, Curimostoma, Deltopylum, Dexiostoma, Dexiotricha, Dichilum, Dogielella, Epenardia (new in present paper), Espejoia, Glaucoma, Loxocephalus, Malacophrys, Monochilum, Paratetrahymena, Pinchatia, Platynematum, Pleurochilidium, Physalophrya, Pseu- doplatynematum, Sathrophilus, Stegochilum, Tetrahymena, and Turaniella.

Because of its direct bearing on the subject of the present paper, at least brief mention must be made here of what I consider to be reasonable disposition of the above genera, leaving, by elimination, only 11 in the Tetrahymenidae proper (with five-Blepharostoma, Chasmatostoma, Malacophrys, Pleurochilidium, and Pseudoplatynematum-of uncertain taxonomic status until rediscovery and redescription of their type-species). Cardiostomatella is possibly a cyrtophorine gymnostome, thus in a separate holotrich order; Cinetochilum has its own family, in a different suborder; Curimostoma has its own family; Dexiostoma falls as a synonym of Colpidium, in my judgment; Dexiotricha is in a separate family with Loxocephalus; Dichilum may well belong to the Glaucomidae, described in the present paper as a new family; Dogielella is in a separate family with Curimostoma; Epenardia, Espejoia, and Glaucoma are in the Glaucomidae; Loxocephalus has its own family, in a different suborder; Monochilum, Pinchatia, and Physalophrya may well belong to the Glaucomidae; Sathrophilus is in a separate family with Cinetochilum, in a different suborder; Turaniella is in the Glaucomidae.

Discussion of such significant taxonomic dispersal of alleged "tetrahymenid" genera is largely inappropriate here; but two important points may be made in passing. The family Tetrahymenidae is reduced in size to a quite compact group of closely related genera (bonafide members: Colpidium, Deltopylum, Paratetra- hymena, Platynematum, Stegochilum, Tetrahymena); and attention to new ideas and new techniques has required recognition of some former presumed members of this tetrahymenine group as members actually of families belonging to an en- tirely separate suborder of the hymenostomes: glaring evidence, it must be ad- mitted, of the primitiveness of the state of protozoan systematics.

The major aim of the present paper is restricted to relief of some of the heterogeneity of the well established family Tetrahymenidae by recognition and description of a new, closely related, yet separate, familial group, the Glaucomidae n. fam., with type-genus Glaucoma Ehrenberg, 1830.3 Since a new genus is involved, all genera must be treated; and since a new species of Glaucoma is described, all recognized species of that genus must be characterized in a compara-

3 I have been thinking about such a separation ever since my early studies, started over 20 years ago, on three of the "key" genera involved, Colpidium, Glaucoma, and Tetrahymena. Ex- cept for Tetrahymena (see Corliss, 1970 and 1971 for references to my papers on that genus), reports published to date on the results of such research are limited essentially to three brief abstracts (Corliss, 1953a, 1954a, 1959a) and a short paper on Glaucoma frontata (Corliss, 1954b). The genus Colpidium, considered as a good member of the family Tetrahymenidae, still awaits full taxonomic treatment.



346 TRANS. AMER. MICROS. SOC., VOL. 90, NO. 3, JULY 1971

tive way, in accordance with the latest provisions of the International Code of Zoological Nomenclature (1964).

MATERIALS AND METHODS

Most of the data used in the present paper were gathered long ago, especially during the stimulating year spent in the laboratory of Professor E. Faure-Fremiet, in Paris, 1951-52. Not only were living cultures of a number of the species described below established and studied at that time but also stained preparations from the extensive slide collection of Faure-Fremiet were available for comparative morphological analyses. In subsequent years additional information and additional silver impregnation and Feulgen nuclear preparations have been accumulated, filling out some gaps in the overall taxonomic picture. In the mean- time, also, new facts have been discovered and published by other workers; these have thus been incorporated in the present taxonomic revision.

Silver-impregnated material has been indispensable in this study: the "silverline system" remains the single best feature in the comparative morphological- taxonomic investigation of the systematics of most ciliate groups. While recognizing the often superior value of the protargol technique (Small, 1967; Tuffrau, 1967), especially for morphogenetic characteristics, the Chatton-Lwoff technique of silver impregnation (Corliss, 1953b) remains also an excellent method. Most of the slides examined in the present study were prepared by the latter technique (see Figs. 1-7); in fact, protargol had not yet come into fashion for ciliates during the times when most of the preparations I studied were made.

Most of my personal observations have been made using Zeiss or Leitz or Wild optics, frequently employing the oil immersion objective.

Many of the detailed measurements taken are not included in the present rather concise comparative taxonomic account: e.g., widths and lengths of the infraciliary bases of the buccal membranelles. Such data are available in my record books should anyone desire to see them. None of the generalizations or conclusions drawn in the following pages conflicts with such statistical "back-up" material.

CHARACTERIZATION OF NEW FAMILY

I recognize the holotrich order Hymenostomatida as comprised of five suborders today, as compared with the three generally accepted since the time of publication of the monograph of Corliss (1961). The two added to the well known Tetrahymenina, Peniculina, and Pleuronematina are the Philasterina Small, 1967 and the Ophryoglenina Canella, 1964.4 The Tetrahymenina may be considered to contain only three familial groups: a small family of mouthless, parasitic forms, the Curimostomatidae Jankowski, 1967; the Tetrahymenidae, best known through the widely distributed and widely studied genus Tetrahymena; and the Glauco- midae n. fam., here split off from the Tetrahymenidae in recognition of significant taxonomic differences, as described below.

Family GLAUCOMIDAE n. fam. Characterization. Contains species of variable body size, but typically length

ranges from 40 to 70 u (though occasionally up to 250 I or more), with form char- acteristically ovoid to elliptical. Ciliation uniform, dense, with no specialized

4 Characterizations and discussions of hymenostome ciliates generally beyond the scope of the present paper are included in contributions of mine, now in press, prepared by invitation for Volume II of P.-P. Grasse's Traite de Zoologie. The taxa which are treated in the present work are also included in that material, of course.




caudal cilium or cilia. Pre-oral suture bent, angled, or displaced slightly to the left of longitudinal axis of the body. Buccal overture elliptical or tear-shaped, oriented at a slight angle to the body axis, and much smaller than the buccal cavity. Pronounced "ectoplasmic lip" on right rim of the oral opening (at least in type-genus). Buccal cavity large, containing inconspicuous undulating membrane (UM) or haplokinety and three membranelles or polykineties of various sizes comprising the adoral zone of membranelles (AZM), the base of one of which (M2 or M3, depending on the genus) is especially conspicuous in its size. Multiple post-oral meridians (POM's), the right-most one stomatogenous in nature. Stoma- togenesis of the somatic-meridional type (Corliss, 1967), with distinctly recogniza- ble stages unique to the family, such as the elongate early buccal anlagen oriented parallel to the longitudinal axis of the body. Contractile vacuole (water expulsion vesicle) discharges through single pore (CVP), often located subequatorially rather than near posterior end of the body, on right side. Single, typically compact macronucleus, centrally located, with single, spherical micronucleus nearby. Generally monomorphic forms, though several species exhibit a non-obligatory dimorphic life cycle. Most species microphagous bacteria-feeders; strains of several have been established in axenic laboratory culture. Physiology and genetics little studied to date. All free-living, fresh-water forms.

Composition. Four genera, Glaucoma (the type-genus), Epenardia (new), Espejoia, and Turaniella, seem quite clearly assignable to this new family. Four others are possibly members of the family (Dichilum, Monochilum, Physalophrya, and Pinchatia), but more precise data are needed on their species before their inclusion can be made on a sound basis.

Comparison with Related Families. The only family with which comparison really needs to be made is the closely related Tetrahymenidae, especially since the type of the new family is Glaucoma, a genus formerly assigned there. Without presenting a detailed characterization of the "parental" family itself, eight principal differences may be noted by contrasting the following features of species of the Tetrahymenidae with the description of members of the Glaucomidae given above (or below). (1) Body and buccal overture shapes: pyriform. (2) Pattern of convergence of right-hand meridians onto pre-oral suture: sharper angle, less curved. (3) Width of infraciliary bases of AZM: always only 3-4 kinetosomes across; no one membranelle particularly more conspicuous than the others. (4) Anlagen of buccal organelles in early stomatogenesis: much shorter and at wider angle to axis of body. (5) Post-oral meridians (POM's): only 1-5 in number. (6) CVP's: one to several, typically near posterior end of body. (7) Greater variability with respect to body size, life cycle complexity, shape of macronucleus, presence or absence of caudal cilia. (8) Wider distribution in nature, greater range in habitats and ways of life.

Several of the key morphological familial differences may be noted in the semi- diagrammatic drawings of the type-species of the type-genera of the two taxa, Tetrahymena pyriformis and Glaucoma scintillans, respectively: cf. Figure 8 with 9, Figure 20 with 21.

CHARACTERIZATION OF INCLUDED GENERA

Of the eight genera possibly assignable to the family Glaucomidae, the four better known ones (Glaucoma, the type; Epenardia, new; Espejoia and Turaniella, taxonomically controversial forms) are described first. The last four (Dichilum, Monochilum, Physalophrya, Pinchatia) are characterized only very briefly, since their status must remain uncertain until we have more information about them, particularly about the composition of their buccal apparatus and their mode of stomatogenesis.




Type-Genus GLAUCOMA Ehrenberg, 1830 With characters of the family sensu stricto. Ciliary meridians or kineties from

the right side of the ventral surface converge onto the pre-oral suture in a pattern forming a much wider angle with the suture than that formed by those meridians converging from the left. Membranelle M2, brush-like with fused cilia, is hypertelically developed in some species. An "X" body is frequently associated with the anterior end of the base of M2. Several species common and widely dispersed, others rarely seen.

Some 58 species of ciliates have been described in the literature as having taxonomic affiliation, at one time or another, with this genus. Kahl (1931, 1935) reduced the number to 16, but at least four more have been described since his review. I recognize only eight, including one described below as new, with real- ization that rediscovery and redescription may "validate" a few others as well (perhaps four or five), though many of them are definitely not congeneric with the type-species G. scintillans (best known example of this is the "G. pyriformis" of the literature, which is really Tetrahymena pyriformis: see Corliss, 1953c, and Corliss & Dougherty, 1967, for reference to this nomenclaturally complex case). The eight are characterized and discussed, generally very succinctly, below, ar- ranged alphabetically after consideration of the type-species. It should be under- stood that common generic (or familial) features are, in general, not repeated in my treatment of the included species. Also minute "silverline" structures of little or no differential significance are deliberately omitted, though noted and recorded during my long-extended studies (e.g., secondary somatic meridians, intermeridio- nal connectives, intrameridional cross-fibrils, apical loops, circumoral rings, pre- oral suture, inter-membranellar connecting fibrils, oral ribs, cytoproct, contractile vacuole pores).

G. scintillans Ehrenberg, 1830. Type-species. Widely known. Body length 40-75 u; 33-40 ciliary meridians, with 7-9 POM's. Width of infraciliary bases of membranelles of AZM5, in kinetosomes: ca. 5-6 for M1, 8-10 for M2, 3 for M3. CVP subequatorial. See Figures 1,2,9,21.

Many ciliatologists have known this species well (actually the original description by Ehrenberg, 1830, is unusually good): the literature on it is too nu- merous to cite in full. Several key taxonomic-morphological references, works in which in general silver-impregnated specimens were used in the study, are the following: Bary (1950), Corliss (1953c, 1954a,b, 1959a,b), Czapik (1968), Dragesco (1970), Furgason (1940), von Gelei (1935, 1936), von Gelei & Horvath

5 A small detail, but one not without possible significance, concerning the exact composition of the infraciliary bases of the buccal organelles of species belonging especially to the genera Glaucoma and Epenardia should be made here, since it is purposely omitted in the text description of these species-partly because it is often not precisely known and partly for the sake of conciseness in their diagnoses. With the notable exception of the undulating membrane or haplokinety, whose zig-zag base is uniformly two basal bodies in width, the kinetosomal widths of the membranelles or polykineties may vary considerably at different places along their own lengths in specimens of a given strain. This fact has also not been particularly indicated in Figures 21-25, because of their semi-diagrammatic nature.

FIGURES 1-4 are all unretouched photomicrographs of the ventral surface of specimens of Glaucoma species impregnated with silver by the Chatton-Lwoff technique. Fig. 1. G. scintillans, type-species, individual from a strain of rather small size. Notice especially the "mouth- parts," the infraciliary bases of the three membranelles in the buccal cavity. Fig. 2. G. scintillans, individual from same culture starting to undergo new mouth-formation (stomatogenesis). Fig. 3. G. macrostoma, deep focus to show (bases of) buccal organelles. Fig. 4. G. macrostoma, same specimen, high focus to show ciliary meridians, outline of buccal overture, etc.




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(1931), Kahl (1931), Klein (1926, 1927), Parducz (1936). But the organisms labelled "G. scintillans" used in early researches of Chatton in France and in phys- iological work by Kidder in America are now considered to have been strains of a different species, G. chattoni (see below).

G. avellana Kahl, 1926. Little known, but differentiated by its distinctive "hazelnut" body shape (with acuminate anterior end), rather short M1, smallish macronucleus, and location of CVP in posterior quarter of the body. Body length 30-60 pL. See Kahl (1926, 1931) and Figure 11.

G. chattoni Corliss, 1959. Superficially resembles Tetrahymena pyriformis more than G. scintillans (with which it is, however, congeneric, of course) because of its tapered anterior end, smaller body size (often 45-50 p), less conspicuous bases of the AZM (though "X" body associated with M2 often easily distinguish- able), more posteriad location of the CVP, fewer rows of cilia (only 22-30), and fewer POM's (5-6). See Figures 6,10.

Although recognized only in recent years, G. chattoni may become more frequently studied than G. scintillans, partly because of the ease with which it may be grown in axenic culture in the laboratory. Since its first brief description as a new species (Corliss, 1959a), the species has been studied morphogenetically, corticotypically, and ultrastructurally (e.g., see Frankel, 1960a,b, 1961; Klug, 1968; Pitelka, 1961) and also physiologically and biochemically (e.g., see Dewey, 1967; Hall, 1967; Holz, 1964: and references therein).

G. dragescui n. sp. Long ago recognized as different from all other species of the genus (mentioned in my 1954a abstract as a species "to be described as new in a forthcoming paper"), based on accumulated unpublished data from study of strains in the Paris area, the same organism was very recently well described by Dragesco (1970) under the name "G. myriophylli." (Other workers may also have observed this organism in the past, but only Dragesco's account is detailed enough to make identification certain.) Thus, it is a pleasure to name the species in honor of my long-time colleague, friend, and superb protozoologist, Jean Dragesco. See Figures 14,24.

G. dragescui is the largest species of Glaucoma (though not the largest member of the family Glaucomidae) which I accept as a bonafide member of the genus. Length is commonly 100-175 tA; shape is broadly ovoid; there are 75-90 ciliary meridians, with 7-9 POM's. Single most distinctive differentiating character is the infraciliary base of M2 of the AZM: it is an enormous plaque of closely appressed kinetosomes (at least 15 rows across), far more prominent than the bases of M1 or M3 (Fig. 24). It has a clearly visible "X" body at its anterior end, in actuality probably a detached portion of M2 rather than an independent buccal organelle. The only other species of Glaucoma with a closely similar membranellar configuration (hypertelically developed M2 plus distinctive "X" body) is G. frontata (see below, and Corliss, 1954b). On this striking point alone, future taxonomists of the group may wish to consider erection of a separate genus, or at least a separate subgenus, for G. dragescui and G. frontata; but I am opposed to such a decision on the basis of the data available to date.

Superficially, there may be two ciliates which might be confused with this new species. One, the "G. myriophylli" of the literature, is treated below as type of a new genus: on close study, the only real similarity between G. dragescui and Epenardia myriophylli is in the large body size. The second ciliate is the "Glau-

FIGURES 5-7 are all unretouched photomicrographs of essentially the ventral surface of specimens of Glaucoma species impregnated with silver by the Chatton-Lwoff technique. Fig. 5. G. macrostoma, specimen in a view and focal plane to show interior of buccal cavity and even



CORLISS-NEW FAMILY OF HYMENOSTOME CILIATES

6

the cytopharynx. Fig. 6. G. chattoni, anterior end. Note obvious differences in comparison with preceding figures of G. scintillans and G. macrostoma (and see text). Fig. 7. G. frontata, group of individuals, showing general "silverline" features and variation in body shape (lower power of magnification than that of the preceding figures). Notice particularly the important generic characteristic of the 'looping over" of the right-hand meridians just anterior to the buccal overture.

351




coma gigantea" of R. & L. Grandori (1934), a soil form again reminiscent of G. dragescui in size (and shape) but so insufficiently described that specific identification is totally impossible. About this species Kahl (1935) himself stated that it can only be considered "eine weitere grosse Glaucoma," reexamination of which is desirable. Should the organism reported from Italy be rediscovered and properly redescribed and found to be identical with G. dragescui, then the name G. dragescui would have to fall as a junior synonym of G. gigantea R. & L. Grandori, 1934.

Known strains (from my previously unpublished data and from observations of Dragesco, 1970) of G. dragescui occur in fresh-water habitats in the environs of Paris, France and Yaounde, Cameroons, Africa. Whether or not Czapik (1968) saw or confused specimens of this species with Epenardia myriophylli, in France and in Poland, is discussed below (under the new genus Epenardia).

G. frontata (Stokes, 1886) da Cunha, 1913. Synonym G. bacillivorax von Gelei, 1936. Body spindle shaped, rounded anteriorly in typical Glaucoma-way, tapered posteriorly often to a distinctly pointed tail (Stokes' "microscopic shark"). Large, length often 100-140 /A; 50-60 ciliary rows, 7-9 POM's. Ham-shaped infraciliary base of M2 very wide, as described above for G. dragescui, and "X" segment prominent. Bacteria-feeder; found in America, Africa, and Europe, though not frequently. Described from study of silver-impregnated specimens by Corliss (1954b), Dragesco (1970), and von Gelei (1936). See Figures 7,16,23.

Although Corliss (1954b) was remiss in not realizing the existence of the excellent earlier description of this species by von Gelei (1936: in Hungarian), von Gelei himself did not seem to know of the scattered literature on Stokes' "Dallasia frontata" and thus unnecessarily but inadvertently named the species as new. Kahl would surely have caught this, but his completed monographic work was published (see sections on hymenostomes in Kahl, 1931, 1935) a full year before the appearance of von Gelei's (1936) paper. We are indebted to da Cunha (1913) for first recognizing the Glaucoma affinities of Stokes' (1886) curious ciliate.

The life cycle of G. frontata is puzzling (apparently complex, polymorphic: see Calkins & Bowling, 1928, 1929) and definitely requires restudy.

G. intermedia (Gajevskaja, 1927) Corliss, 1954. A species probably rather closely related (possibly identical) with G. kirki Bary, 1950 (see below); dat crucial to differential diagnosis are largely lacking in Gajevskaja's (1927, 1933) original and subsequent description of "Dichilum intermedium" and in the account by Kahl (1931); the latter author, however, recognized its Glaucoma affinities. Body length, 55-60 /; short, dense ciliation. Macronucleus even larger than in G. kirki and possibly fewer POM's and a smaller buccal cavity. Feeds on algae (Ulothrix) as well as bacteria; found in the cold waters of Lake Baikal. Species transferred to the genus Glaucoma by Corliss (1954a). See Figure 15.

G. kirki Bary, 1950. Small, average length ca. 35 u; typical Glaucoma-shape, but anterior end terminates in small, dome-like knob; ca. 30 ciliary meridians, with 5-6 POM's. Prominent ectoplasmic lip or flange on right of buccal overture. Width of bases of membranelles of AZM, in kinetosomes: 2 for M1, 4 for M2, and, uniquely, only 2 for Ma. CVP in posterior third of body. Macronucleus unusually large. Bacteria-feeder in stagnant ponds near Wellington, New Zealand. In conjugation, one member of the pair is much larger in body size. Well described by Bary (1950) in spite of lack of use of a silver technique; a 10% nigrosin solution, carefully applied, revealed important features of the somatic and buccal infraciliature which are usually discoverable only in silver-impregnated material. See Figures 12,22.

G. macrostoma Schewiakoff, 1889. Similar to G. scintillans and often considered identical to it by scattered workers (the name then falling as a synonym



CORLISS--NEW FAMILY OF HYMENOSTOME CILIATES 353

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FIGURES 8-13 are semi-diagrammatic outlines of the body form, buccal overture, and general pattern of ciliature on the ventral surface of various species of tetrahymenine hymenostomes, all drawn to the same scale (as indicated). Mouth parts and other details are intentionally omitted. Fig. 8. Tetrahymena pyriformis, type-species of the type-genus of the family Tetrahymenidae. Fig. 9. Glaucoma scintillans, type-species of the type-genus of the family Glaucomidae n. fam. Fig. 10. G. chattoni. Fig. 11. G. avellana. Fig. 12. G. kirki. Fig. 13. G. macrostoma.




of that of the earlier, Ehrenberg, species). But careful comparative study of data I collected on strains found near Paris, plus reexamination of the original description and figures of Schewiakoff (1889) and the comments of Kahl (1931), sup- ports the conclusion that they are separate, though very closely related, species. Body length 35-40 Ij; plump oval shape, with both ends equally rounded; buccal overture relatively huge (for the body size) and located in the posterior portion of the anterior half of the body; ca. 40 ciliary meridians, packed closely together with 7-9 POM's. AZM bases as in G. scintillans. CVP subequatorial. One of the smallest species of Glaucoma but with one of the largest, and certainly most conspicuous, oral areas. See Figures 3-5, 13.

New Genus EPENARDIA n. g. Large body (length up to 220 Iu, although some strains reach only 140 v); form

much as in Glaucoma, though often more elongate and even tapered posterior end, and with a distinctly different pattern of convergence of ciliary meridians onto the pre-oral suture: the kineties from both sides converge at an acute angle. Very densely ciliated; over 100 rows of cilia, with 9-10 POM's. Pellicle shows curious sculpturing or tessellation. Buccal overture more tear-shaped than oval or elliptical. CVP subequatorial. Stomatogenesis as in Glaucoma. Single most outstanding taxonomic character: huge size of the infraciliary base of membranelle Ms, some 15-20 kinetosomes in width (thus at least double the width of M2 here), hidden deep in the buccal cavity, and displaced far to the right (Fig. 25). Very distinct "X" body associated with M2. Genus named in honor of Eugene Penard, great ciliate-watcher of the early 20th century and discoverer of the type-species, E. myriophylli (Penard, 1922) n. comb.

E. myriophylli (Penard, 1922) n. comb. Type- and sole species; with characteristics of the genus. Average body size (length) generally between 150 and 200 /J: varies with the strain. Found in association with Myriophyllum by its discoverer, Penard (1922), but noted by others to feed on Beggiatoa or fungal mycelia. Quite widespread distribution, apparently. See Figures 17,25.

This is the celebrated "Glaucoma myriophylli" of Penard (1922) and Kahl (1931, 1935) and of the general literature, though often workers have not given enough data for us to be certain, today, that they were dealing with this organism of some other (such as Glaucoma dragescui: see discussion under that species, above). With customary care, Penard actually revealed the very characteristics which cause its removal from the genus Glaucoma6; but subsequent workers have not properly noted them all. In addition to describing such features as the pattern of meridional convergence on the pre-oral suture and the tessellated pellicle, the intrepid Geneva protozoologist also correctly depicted-in spite of his lack of silver-impregnated material and even of the magnification afforded by an oil immersion objective (which he had not)-the infraciliary bases of the membranelles of the tripartite AZM! Unfortunately, his observation was made from the dorsal surface (his figure correctly labelled "vue par transparence") and, thus, subsequent readers of his work may have failed to grasp the fact that he saw clearly the hypertelically developed M3. (In the genus Glaucoma, it may be re- called, it is M2 which, in several species, is similarly enlarged: cf. Figs. 23,24 with 25).

I have studied several strains collected in the area of Paris, France. The morphological data from silver-impregnated specimens extend the original observations of Penard (1922) and also support, or are confirmed by, the excellent description of "Glaucoma myriophylli" given by von Gelei (1936), who also used

6 Incidentally, none of Penard's eight "Glaucoma" species is currently retained in that genus!



CORLISS-NEW FAMILY OF HYMENOSTOME CILIATES

( I /

' 4 ' 4 15 ' 6 , -

l~~~~ojxu~~~ 20^no~~~~ 4c' 17 ' ' 18 19

FIGURES 14-19 are semi-diagrammatic outlines of the body form, buccal overture, and general pattern of ciliature on the ventral surface of various species in the tetrahymenine family Glaucomidae n. fam., drawn to the two scales indicated. Fig. 14. Glaucoma dragescui n. sp. Fig. 15. G. intermedia. Fig. 16. G. frontata. Fig. 17. Epenardia myriophylli (n. g., n. comb.). Fig. 18. Espejoia mucicola. Fig. 19. Turaniella vitrea.

silver techniques in his research on this interesting organism. The "G. myriophylli" of Czapik (1968) is also quite possibly E. myriophylli, though some confusion arises in careful examination of her paper. Either the bases of the AZM are in- correctly portrayed in her schematic Figure 12 (seemingly contradicted by several

355

_ 28s~~~~~~~~M




of her photomicrographs) or she was inadvertently studying a mixture of specimens from two different species: E. myriophylli and G. dragescui (not an unlikely possibility).

Dragesco (1970) seemed puzzled, in his account of his African "G. myriophylli," by the differences he was finding with the description by Czapik (1968) and with certain other characteristics mentioned in the literature on the species. Recalling my account of the new species G. dragescui, above, I believe that Dragesco's puzzlement is now quite understandable: he was dealing with a different "large Glaucoma," not Epenardia myriophylli at all! The immense M2, as contrasted with an immense M3, is the most striking differential characteristic; other important differences also exist, of course (see preceding pages).

Genus ESPEJOIA Burger, 1908 Controversial in its taxonomic allocation in recent years, this genus is tenta-

tively included in the family Glaucomidae until or unless new data or new sound reinterpretation of older data suggest that such a position is untenable. Body length 70-100 ,t; shape elongate-ellipsoid, with anterior end truncate on right side; ca. 40 ciliary meridians, with 6-7 POM's. Pattern of convergence of right-hand kineties onto pre-oral suture unlike that of Glaucoma. Dimorphic life cycle, with buccal overture very prominent in the macrostome form and highly reduced in size in the lean, rapidly swimming microstome form. Membranelles of AZM all four-rowed in width, at least superficially reminiscent of the peniculi7 of such peniculine hymenostomes as Paramecium: the base of M3, in particular, reminds one of a quadrulus; but convergent evolution might serve as a satisfactory ex- planation of such seeming homology. Species found in fresh-water habitats, always in association with the mucilaginous material surrounding eggs of aquatic insects and molluscs or the thalli of certain algae. Depending on possible synonymies (the nomenclatural problem involved is beyond the scope of the present paper), the name of the type-species is either E. culex (Smith, 1897) Kahl, 1931 or E. obtusa Biirger, 1908 (see Watson, 1940, for an indication of the complications involved). Burger (1908) and Kahl (1931) may be consulted for further characterization of the genus.

E. mucicola (Penard, 1922) Kahl, 1931,8 essentially with characteristics of the

7 The exact nature and definition of a "peniculus" still defy the very ciliatologists who are supposedly the experts on the species possessing such structures. Canella (1970) discusses this problem and related matters in extenso in his very recently published hefty critique which did not reach me until the present work was in press: thus I cannot consider his contributions here, appropriate though many of his points may be to topics treated in this paper of mine.

8 Unfortunately, this name-now rather well known-may itself fall as a junior synonym of the name (whatever it may be!) of the type-species, if future workers decide that the several nominal species involved are, in fact, identical at the species level. Penard (1922) considered the present ciliate as a new species in Schewiakoff's (1889) controversial genus Balantiophorus, but I consider that genus to be a relative of Pleuronema in my treatment of the hymenostome suborder Pleuronematina (for the Traite de Zoologie, in press).

FIGURES 20-27 are semi-diagrammatic drawings of the buccal apparatus, in ventral view, particularly showing the configuration of the bases of the membranelles comprising the AZM (as visualized through the opening of the buccal overture) of selected species of tetrahymenine hymenostomes, drawn to the scales indicated. Details which would be confusing are deliberately omitted (see text); location of the UM is portrayed in only the first three figures (on the viewer's far left). Membranelles at the AZM are known as M1, M2, Ms, reading from viewer's right to left. Fig. 20. Tetrahymena pyriformis, representative of the picture in the Tetrahymenidae. Fig. 21. Glaucoma scintillans, representative of the Glaucomidae n. fam. (although variations on the theme for this family appear in the following figures). Fig. 22. G. kirki. Fig. 23. G. frontata.




20

23

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Fig. 24. G. dragescui n. sp. Fig. 25. Epenardia myriophylli (n. g., n. comb.). Fig. 26. Espejoia mucicola (drawing based on Figure 3 in Faure-Fremiet & Mugard, 1949). Fig. 27. Turaniella vitrea (drawing based on Figure 2 in Iftode & Versavel, 1968).

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genus as outlined above, is the only species to be considered in the present paper. It has been studied in recent times by Faure-Fremiet & Mugard (1949), using silver-impregnated material and offering a modern account of the morphogenetics of its interesting dimorphic life cycle, an ontogeny reminiscent of that found in several species of the genus Tetrahymena (see Corliss, 1971, for an up-to-date review of the situation in T. patula, T. vorax, and T. paravorax). See Figures 18, 26.

The mode of stomatogenesis (somatic-meridional), the cortical picture, and the life cycle are the principal reasons for considering Espejoia to be a tetrahymenine rather than a peniculine ciliate-probably closer to Glaucoma than to Tetrahymena, however. Its alleged possession of "true" peniculi at one time influenced workers (e.g., Corliss, 1961) to place it near Paramecium. Very recently, several French investigators (e.g., see Didier, Iftode & Versavel, 1970) have considered it, along with the genus Turaniella (treated below), as possibly a member of a "connecting link" group between the hymenostome suborders Tetrahymenina and Peniculina.

Genus TURANIELLA Corliss, 1960 A genus as taxonomically controversial as Espejoia (above), and essentially

for the same reasons. Body length may reach sizes (e.g., 250 p) which would be among the greatest in the family; shape is elongate-ovoid, tapered posteriorly; there are ca. 65 ciliary meridians, with 6-8 POM's. Pattern of convergence of right-hand kineties onto pre-oral suture unlike that of Glaucoma. Buccal overture large, very prominent. Membranelles, or polykineties, of AZM show varying con- figurations: M1 is four rows of kinetosomes in width; M2, six; M3, four, but in very widely separated lines, extremely reminiscent (at least superficially) of a peniculine quadrulus (see Fig. 27). Stomatogenesis is very much like that found in species of Glaucoma. Fresh-water habitats, carnivorous on other ciliates, such as species of Colpidium. Single, type-species, T. vitrea (Brodsky, 1925) Corliss, 1960.

Turaniella is a replacement name for the preoccupied Turania Brodsky, 1925 (see Corliss, 1960). T. vitrea, with the characteristics of the genus, was originally described by Brodsky (1925) and included in Kahl's monographic works (Kahl, 1931) as a member of his large family Frontoniidae. Corliss (1961) listed the genus as an "unassigned tetrahymenine." Very recently, several French workers have studied the species with considerable care, employing electron microscopy as well as methods of silver impregnation in an attempt to learn about its ultrastructure and its morphogenesis and, incidentally, its possible phylogenetic affinities (see Didier et al., 1970; Iftode & Versavel, 1968; Iftode et al., 1969). They particularly compared it (favorably, taxonomically) with the genus Colpidium (would that they had chosen a Glaucoma for comparison!), but still remained influenced by its alleged possession of peniculi. See Figures 19,27.

Thus, as mentioned above under Espejoia, our French colleagues have sug- gested a "connecting link" sort of position for these curious ciliates with cortical and stomatogenic features so clearly tetrahymenine (specifically, Glaucoma-like) in nature and mouth-parts possibly peniculine in structure. But factors of convergent or even parallel evolution might account for the "penicular" appearance of the bases of the membranelles. And separation of M3 into four distinct rows need not, taken by itself, indicate (a) that it is a "true" quadrulus, or (b) that its possessor is thus a peniculine hymenostome. Further information on the precise ultrastruc- tural composition, configuration, and even interrelationship of polykinetal bases in the various taxonomic groups within the Hymenostomatida is desirable, with the reasonable hope that the similarities and/or differences found will throw useful light on the systematics of the species involved.



CORLISS NEW FAMILY OF HYMENOSTOME CILIATES

Four Uncertain Genera As mentioned above, I consider the possible inclusion of four additional genera

in the family Glaucomidae. The type-species of these groups, however, must be rediscovered and properly redescribed before firm taxonomic allocation can be made with any high degree of confidence. Thus the genera are characterized only briefly in the present paper.

Dichilum Schewiakoff, 1889. Ovoid to elliptical forms; length 40-140 u. Small oral area. Type-species, D. cuneiforme Schewiakoff, 1889. Very small, rapidly moving ciliate. Limited information appears in Schewiakoff (1889, 1896) and Kahl (1931); the latter includes references to other species possibly admissable to the genus.

Monochilum Schewiakoff, 1893. Ovoid, with tapering posterior end; length 60-180 t. Type-species, M. frontatum Schewiakoff, 1893. About 80 t in length and with anterior end more rounded than posterior, this species reminds one, at least superficially, of a small strain of Epenardia myriophylli. Unfortunately, precise data on mouth-parts are not included in the descriptions of Schewiakoff (1893, 1896) and Kahl (1931). Several species have been reported (see Kahl, 1931), but whether or not they are really congeneric with M. frontatum remains to be confirmed.

Physalophrya Kahl, 1931. Elongate, cylindrical shape; large forms, length 150-310 L. Oral ciliature possibly Glaucoma-like. Unique ribbon-like macronucleus. Type-species may be considered to be P. spumosa (Penard, 1922) Kahl, 1931. This and a second species, P. cylindrica, were both originally placed in the genus Glaucoma by Penard (1922) and both removed by Kahl (1931) to his genus Physalophrya near Paramecium (then considered a trichostome ciliate). More data needed. If the genus truly has affinities with the other genera included in the Glaucomidae in the present paper, then the characterization of the family itself may have to be expanded a little to properly accommodate its species.

Pinchatia Shibuya, 1931. Ovoid to pyriform in shape, with truncate appearance to anterior end (reminiscent of Espejoia); small forms, length only ca. 30 L. Buccal cavity relatively large. Single, type-species, P. chaetophorae (Penard, 1922) Shibuya, 1931. Found associated with the gelatinous coat of the fresh-water alga Chaetophora-again reminding one of Espejoia species. Penard (1922) described P. chaetophorae as a member of the genus Balantiophorus (which I now consider related to Pleuronema in the hymenostome suborder Pleuronematina); Kahl (1931) placed it in Glaucoma; and Shibuya (1931) thought it deserved separate generic status, although its description, by today's standards at least, leaves much to be desired. Watson (1940) has reviewed some of the problems associated with placing it properly taxonomically. If Pinchatia truly does possess characteristics relating it to both Glaucoma and Espejoia, its rediscovery and restudy might shed considerable light on systematic interrelationships, and their bases, within the entire family Glaucomidae.

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359




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Documents

Establishment of a New Family (Glaucomidae n. fam.) in the Holotrich Hymenostome Ciliate Suborder Tetrahymenina, and Description of a New Genus (Epenardia n. g.) and a New Species