C 55S. 
 
 5 d\R(L ./§ 
 
 NOAA TR NMFS CIRC-378 
 
 A UNITED STATES 
 DEPARTMENT OF 
 
 COMMERCE 
 
 PUBLICATION 
 
 NOAA Technical Report NMFS CIRC-378 
 
 A^°'^ 
 
 
 U.S. DEPARTMENT OF COMMERCE 
 
 National Oceanic and Atmospheric Administration 
 
 National Marine Fisheries Service 
 
 
 Marine Flora and Fauna of 
 
 
 the Northeastern United States. 
 
 
 Protozoa: Ciliophora 
 
 
 ARTHUR C. BORROR 
 
 
 I §§f§W§ § 
 
 SEATTLE, WA 
 September 1973 
 
 
 
 
NOAA TECHNICAL REPORTS 
 
 National Marine Fisheries Service, Circulars 
 
 The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the 
 abundance and geographic distribution of fishery resources, to understand and predict fluctuations in the quan- 
 tity and distribution of these resources, and to establish levels for optimum use of the resources. NMFS is also 
 charged with the development and implementation of policies for managing national fishing grounds, develop- 
 ment and enforcement of domestic fisheries regulations, surveillance of foreign fishing off United States coastal 
 waters, and the development and enforcement of international fishery agreements and policies. NMFS also 
 assists the fishing industry through marketing service and economic analysis programs, and mortgage insurance 
 and vessel construction subsidies. It collects, analyses, and publishes statistics on various phases of the industry. 
 
 The NOAA Technical Report NMFS CIRC series continues a series that has been in existence since 1941. The 
 Circulars are technical publications of general interest intended to aid conservation and management. Publica- 
 tions that review in considerable detail and at a high technical level certain broad areas of research appear in 
 this series. Technical papers originating in economics studies and from management investigations appear in 
 the Circular series. 
 
 NOAA Technical Reports NMFS CIRC are available free in limited numbers to governmental agencies, both 
 Federal and State. They are also available in exchange for other scientific and technical publications in the ma- 
 rine sciences. Individual copies may be obtained (unless otherwise noted) from NOAA Publications Section, Rock- 
 ville, Md. 20852. Recent Circulars are: 
 
 315. Synopsis of biological data on the chum salmon, 
 Oncorhynchus keta (Walbaum) 1792. By Rich- 
 ard G. Bakkala. March 1970, iii + 89 pp., 15 
 figs., 51 tables. 
 
 319. Bureau of Commercial Fisheries Great Lakes 
 Fishery Laboratory, Ann Arbor, Michigan. By 
 Bureau of Commercial Fisheries. March 1970, 
 8 pp., 7 figs. 
 
 330. EASTROPAC Atlas: Vols. 4, 2. Catalog No. 
 
 1 49.4:330/(vol.) 11 vols. ($4.75 each). Avail- 
 able from the Superintendent of Documents, 
 Washington, D.C. 20402. 
 
 331. Guidelines for the processing of hot-smoked chub. 
 By H. L. Seagran, J. T. Graikoski, and J. A. 
 Emerson. January 1970, iv + 23 pp., 8 figs., 
 
 2 tables. 
 
 332. Pacific hake. (12 articles by 20 authors.) March 
 1970, iii + 152 pp., 72 figs., 47 tables. 
 
 333. Recommended practices for vessel sanitation and 
 fish handling. By Edgar W. Bowman and Alfred 
 Larsen. March 1970, iv + 27 pp., 6 figs. 
 
 335. Progress report of the Bureau of Commercial 
 Fisheries Center for Estuarine and Menhaden 
 Research, Pesticide Field Station, Gulf Breeze, 
 Fla., fiscal year 1969. By the Laboratory staff. 
 August 1970, iii + 33 pp., 29 figs., 12 tables. 
 
 336. The northern fur seal. By Ralph C. Baker, Ford 
 Wilke, and C. Howard Baltzo. April 1970, iii + 
 19 pp., 13 figs. 
 
 337. Program of Division of Economic Research, 
 Bureau of Commerecial Fisheries, fiscal year 
 
 1969. By Division of Economic Research. April 
 
 1970, iii + 29 pp., 12 figs., 7 tables. 
 
 338. Bureau of Commercial Fisheries Biological Lab- 
 oratory, Auke Bay, Alaska. By Bureau of Com- 
 mercial Fisheries. June 1970, 8 pp., 6 figs. 
 
 339. Salmon research at Ice Harbor Dam. By Wesley 
 J. Ebel. April 1970, 6 pp., 4 figs. 
 
 340. Bureau of Commercial Fisheries Technological 
 Laboratory, Gloucester, Massachusetts. By Bu- 
 reau of Commercial Fisheries. June 1970, 8 pp., 
 8 figs. 
 
 341. Report of the Bureau of Commercial Fisheries 
 Biological Laboratory, Beaufort, N.C., for the 
 fiscal year ending June 30, 1968. By the Lab- 
 oratory staff. August 1970, iii + 24 pp., 11 figs., 
 16 tables. 
 
 342. Report of the Bureau of Commercial Fisheries 
 Biological Laboratory, St. Petersburg Beach, 
 Florida, fiscal year 1969. By the Laboratory staff. 
 August 1970, iii + 22 pp.," 20 figs., 8 tables. 
 
 343. Report of the Bureau of Commercial Fisheries 
 Biological Laboratory, Galveston, Texas, fiscal 
 year 1969. By the Laboratory staff. August 
 1970, iii + 39 pp., 28 figs., 9 tables. 
 
 344. Bureau of Commercial Fisheries Tropical Atlan- 
 tic Biological Laboratory progress in research 
 1965-69, Miami, Florida. By Ann Weeks. Oc- 
 tober 1970, iv + 65 pp., 53 figs. 
 
 346. Sportsman's guide to handling, smoking, and pre- 
 serving Great Lakes coho salmon. By Shearon 
 Dudley, J. T. Graikoski, H. L. Seagran, and Paul 
 M. Earl. September 1970, iii + 28 pp., 15 figs. 
 
 347. Synopsis of biological data on Pacific ocean perch, 
 Sebastodes alutus. By Richard L. Major and 
 Herbert H. Shippen. December 1970, iii + 38 
 pp., 31 figs., 11 tables. 
 
 Continued on inside back cover. 
 
^OWMOSP 
 
 r ^IENJ Of 
 
 U.S. DEPARTMENT OF COMMERCE 
 Frederick B. Dent, Secretary 
 
 NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION 
 Robert M. White, Administrator 
 
 NATIONAL MARINE FISHERIES SERVICE 
 Robert W. Schoning, Director 
 
 NOAA Technical Report NMFS CIRC-378 
 
 Marine Flora and Fauna of 
 the Northeastern United States. 
 Protozoa: Ciliophora 
 
 ARTHUR C. BORROR 
 
 
 
 o 
 
 1 
 
 a. 
 a 
 
 o 
 
 
 uS 
 
 
 =5 
 
 
 
 SEATTLE, WA 
 
 
 September 1973 
 
 For sale by the Superintendent of Documents, U.S. Government Printing Office 
 Washington, D.C. 20402 - Price 65 cents 
 
FOREWORD 
 
 This issue of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna 
 of the Northeastern United States." This subseries will consist of original, illustrated, 
 modern manuals on the identification, classification, and general biology of the estuarine 
 and coastal marine plants and animals of the Northeastern United States. Manuals 
 will be published at irregular intervals on as many taxa of the region as there are 
 specialists willing to collaborate in their preparation. 
 
 The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates 
 of the Woods Hole Region," edited by R. I. Smith, published in 1964, and produced under 
 the auspices of the Systematics-Ecology Program, Marine Biological Laboratory, Woods 
 Hole, Mass. Instead of revising the "Woods Hole Keys," the staff of the Systematics- 
 Ecology Program decided to expand the geographic coverage and bathymetric range and 
 produce the keys in an entirely new set of expanded publications. 
 
 The "Marine Flora and Fauna of the Northeastern United States" is being prepared 
 in collaboration with systematic specialists in the United States and abroad. Each man- 
 ual will be based primarily on recent and ongoing revisionary systematic research and 
 a fresh examination of the plants and animals. Each major taxon, treated in a separate 
 manual, will include an introduction, illustrated glossary, uniform originally illustrated 
 keys, annotated check list with information when available on distribution, habitat, life 
 history, and related biology, references to the major literature of the group, and a system- 
 atic index. 
 
 These manuals are intended for use by biology students, biologists, biological ocean- 
 ographers, informed laymen, and others wishing to identify coastal organisms for this 
 region. In many instances the manuals will serve as a guide to additional information 
 about the species or the group. 
 
 Geographic coverage of the "Marine Flora and Fauna of the Northeastern United 
 States" is planned to include organisms from the headwaters of estuaries seaward to 
 approximately the 200-m depth on the continental shelf from Maine to Virginia, but 
 may vary somewhat with each major taxon and the interests of collaborators. When- 
 ever possible representative specimens dealt with in the manuals will be deposited in 
 reference collections of the Gray Museum, Marine Biological Laboratory, and other 
 universities and research laboratories in the region. 
 
 After a sufficient number of manuals of related taxonomic groups have been pub- 
 lished, the manuals will be revised, grouped, and issued as special volumes. These vol- 
 umes will thus consist of compilations of individual manuals within phyla such as the 
 Coelenterata, Arthropoda, and Mollusca, or of groups of phyla. 
 
CONTENTS 
 
 Page 
 
 Introduction 1 
 
 Glossary 2 
 
 Techniques 4 
 
 Key primarily to families of the Ciliophora 6 
 
 Annotated systematic list 44 
 
 Selected bibliography 51 
 
 Systematic index 57 
 
 Acknowledgments 62 
 
 Coordinator's comments 62 
 
 The National Marine Fisheries Service (NMFS) does not approve, rec- 
 ommend or endorse any proprietary product or proprietary material 
 mentioned in this publication. No reference shall be made to NMFS, or 
 to this publication furnished by NMFS, in any advertising or sales pro- 
 motion which would indicate or imply that NMFS approves, recommends 
 or endorses any proprietary product or proprietary material mentioned 
 herein, or which has as its purpose an intent to cause directly or indirectly 
 the advertised product to be used or purchased because of this NMFS 
 publication. 
 
 Ill 
 
MARINE FLORA AND FAUNA OF THE 
 
 NORTHEASTERN UNITED STATES. 
 
 Protozoa: Ciliophora 
 
 ARTHUR C. BORROR 1 
 
 ABSTRACT 
 
 This manual includes an introduction on the general biology, an illustrated key, an an- 
 notated systematic list, a selected bibliography, and an index to the marine ciliated Protozoa 
 of coastal and estuarine waters of New England. The key facilitates identification to family 
 of nonencysted, nondividing marine ciliates at any stage in the life cycle. 
 
 INTRODUCTION 
 
 This manual is a guide to the families of ma- 
 rine ciliated Protozoa of coastal and estuarine 
 waters of New England. It includes ciliates to 
 be encountered not only in areas of high salinity 
 but also in the upper reaches of estuaries where 
 the water is brackish. It is intended for use by 
 a broad audience, hence is artificial (order not 
 necessarily parallel to phylogenetic sequence), 
 and is stripped of unnecessary technical terms. 
 The key facilitates identification to family of 
 nonencysted, <nondividing marine ciliates at any 
 stage in the life cycle. 
 
 Some families are omitted from this key be- 
 cause they are represented only by ciliates living 
 in fresh water or in soil. These include Actino- 
 bolinidae, Amphibothrellidae, Astylozoidae, Bur- 
 sariidae, Dendrocometidae, Discomorphellidae, 
 Glaucomidae, Marynidae, Microthoracidae, Oph- 
 rydiidae, Reichenowellidae, Spirozonidae, Trich- 
 ospiridae, and Urocentridae. Families that are 
 represented solely by ciliates endozoic in nonma- 
 rine hosts are also omitted. These include 
 Buetschiliidae, Clevelandellidae, Cyathodiniidae, 
 Gyrocorythidae, Isotrichidae, Maupasellidae, 
 Paraisotrichidae, Plagiotomidae, Pycnotrichi- 
 dae, and Thigmocomidae. 
 
 1 Department of Zoology, Spaulding Building Uni- 
 versity of New Hampshire, Durham, NH 03824. 
 
 In general, the families recognized in the key 
 and genera included in each are those recognized 
 by Corliss (1961); numerous exceptions based 
 on more recent studies are noted in the Anno- 
 tated Systematic List. Each family and most 
 couplets are illustrated by line drawings. 
 
 Since a few characteristics (i.e., presence or 
 absence of an undulating membrane) may be 
 difficult to observe, some families key out in two 
 different places. In such instances, the other 
 couplet in which the family is named is indi- 
 cated. When different genera within the same 
 family key out in different places, the family 
 name is followed by the words "in part." 
 
 Members of the subphylum Ciliophora, usually 
 known as ciliates, are usually single celled with 
 membrane-bound nuclei. Branching colonies 
 occur in the Vorticellidae and Epistylididae 
 (Peritrichida) and in the Dendrosomatidae and 
 Ophryodendridae (Suctorida). Colonies in lin- 
 ear series occur in the Intoshellinidae, Hoplito- 
 phryidae, and Opalinopsidae. 
 
 Most ciliates have two kinds of nuclei, that is, 
 with one to many micronuclei and one to many 
 macronuclei. However, members of the family 
 Stephanopogonidae have only one type of nu- 
 cleus. 
 
 The micronucleus is usually diploid, less than 
 5 [im in diameter, and usually visible only in 
 stained preparations. Ciliates in strains of some 
 species are amicronucleate. 
 
 The macronucleus is usually hyperpolyploid 
 
(contains many sets of chromosomes). It may 
 be single (compact, ribbonlike, beaded, or 
 branched) or separated into membrane-bound 
 structures that typically reconsolidate prior to 
 fission. Macronuclei of members of the Trache- 
 locercidae, Loxodidae, and Geleiidae have exten- 
 sive Feulgen-negative areas, cannot replicate, 
 and, in some cases, have been shown to be diploid. 
 
 Within the Ciliophora there is wide variation 
 in the arrangement of the cilia, and this forms 
 the basis for classification. Cilia are absent at 
 some stages of the life cycle of all members of 
 the order Suctorida, many members of the 
 family Sphenophryidae, and members of the 
 genus Conidiophrys. Cilia may occur singly, 
 in rows, bands, or ribbons, or may be grouped 
 as compound structures (cirri, undulating mem- 
 branes, and membranelles). Ciliates that lack 
 cilia during much of their life cycle, however, 
 possess other characteristics of the group. 
 
 Except for members of the Suctorida, Asto- 
 matida, and some members of the order Thig- 
 motrichida, all ciliates possess a permanent open- 
 ing, the cytostome, through which food is 
 ingested. 
 
 All ciliates are heterotrophic and do not ex- 
 hibit ameboid locomotion. In binary fission, the 
 division plane bisects the ciliary rows, dividing 
 the cell into anterior and posterior daughter 
 cells. 
 
 GLOSSARY 
 
 The following terms are used in this key to 
 describe the anatomy of marine ciliates. Com- 
 monly used terms are illustrated in Figures 1-5. 
 More complete definitions of structures are given 
 by Corliss (1959). 
 
 Aboral Surface opposite that on which the 
 cytostome or buccal cavity occurs. 
 
 Buccal cavity A depression in the surface of 
 the cell containing cilia that are differentiated 
 from those of the general body surface and 
 that are used in feeding. This cavity contains 
 the cell mouth or cytostome (see Fig. 1). 
 
 Buccal cilia Cilia in the buccal cavity used in 
 feeding. These cilia usually are organized 
 into compound organelles, such as an undulat- 
 ing membrane or membranelles. 
 
 Ciliary row A linear series, usually longitudi- 
 nal, of simple cilia (see Fig. 2, 3). 
 
 Cirrus A compound ciliary organelle polygonal 
 in cross section, composed of several to dozens 
 of individual cilia. These usually occur on the 
 ventral surface of members of the order Hy- 
 potrichida and are used in locomotion (see 
 Fig. 1). 
 
 Compound ciliary organelle Any structure such 
 as a cirrus, undulating membrane or membra- 
 nelle composed of many functionally united 
 cilia. 
 
 Contractile vacuole A clear membrane-bound 
 vesicle functioning in expulsion of water and 
 accompanied by rhythmic changes in its di- 
 ameter. 
 
 Cytopharyngeal rod Stiff fibrous structures 
 lining the cytopharynx, often in a conical ar- 
 rangement (see Fig. 3). 
 
 Cytopharynx That part of the cytoplasm im- 
 mediately adjacent and internal to the cyto- 
 stome and not ciliated (see Fig. 3). 
 
 Cytostome The cell mouth. This may be on 
 the surface, in a vestibulum ciliated by the 
 cilia of the general cell surface, or in a buccal 
 cavity (see Fig. 5). 
 
 Fronto-ventral cirri Cirri arranged in scat- 
 tered groups anterior to the transverse cirri. 
 
 Lorica A secreted chamber open at least at one 
 end in which some ciliates live. 
 
 Marginal cirri Cirri arranged in longitudinal 
 rows to the right and left of the ventral mid- 
 line (see Fig. 1). 
 
 Membranelle A compound ciliary organelle! 
 composed of several closely set and parallel 
 rows of cilia. They are associated with a buc- 
 cal cavity and may function both in feeding 
 and in locomotion (see Fig. 1). 
 
 Motile Capable of independent locomotion on 
 or off the substratum. 
 
 Penictdus A compound ciliary organelle of the 
 buccal cavity of some members of the order 
 Hymenostomatida. It may consist of a band 
 typically four cilia wide but many cilia in 
 length, very compactly arranged (see Fig. 4). 
 
 Psarnmolittoral Living in interstices among 
 sand grains. 
 
 Scopula A surface organelle that secretes the 
 stalk in many sessile ciliates. Usually at one 
 apex of the cell. 
 
 Sessile Generally incapable of locomotion. 
 Usually attached to a substratum by a stalk or 
 flattened sole, or wrapped around the bristle 
 of some invertebrate. 
 
buccal cavity 
 membranelle 
 
 undulating membrane 
 
 cirrus 
 
 ^-marginal cirri 
 midventral cirri 
 
 transverse cirri 
 
 cytopharyngeal 
 
 ciliary 
 row 
 
 lit 
 
 somatic 
 cilium 
 
 stylus 
 
 somatic cilium 
 
 peniculus 
 
 vestibulum 
 
 Figure 1. — Holosticha diademata, ventral aspect. 
 Figure 2. — Uronema sp., oral aspect. 
 
 Figure 3. — Dysteria sp., ventral aspect. Ciliary rows indicated by dash lines. 
 Figure 4.- — Frontonia sp., oral aspect with region of buccal cavity enlarged in the dia- 
 gram to the right. 
 Figure 5. — Diagram of a ciliate with a lateral vestibulum. 
 
 Simple cilium A cilium that emerges singly 
 
 from the cell surface and is morphologically 
 
 and functionally a unit. 
 Somatic cilia All kinds of simple and compound 
 
 cilia of the general body surface, usually used 
 
 in locomotion (see Fig. 2). 
 Stylus A posterior cirruslike extension of the 
 
 cell surface in members of the Dysteriidae, 
 resembling the posterior projection of some 
 rotifers. It is a secretory organelle for tem- 
 porary attachment (see Fig. 3). 
 Thigmotactic Capable of crawling along sur- 
 faces, and resisting removal from such sur- 
 faces. 
 
Tomite The stage in the life cycle of some cil- 
 iates which results from repeated divisions 
 of a larger cell without any cell growth, such 
 that the eventual cell is considerably smaller 
 than the original cell. 
 
 Transverse cirri A group of five to many rel- 
 atively large cirri near the posterior end of the 
 bod}' on the ventral midline (see Fig. 1). 
 
 Trophont The normal feeding adult stage of 
 the life cycle of the ciliate. 
 
 Undulating membrane A compound ciliary or- 
 ganelle consisting of one or several closely 
 set rows of cilia usually placed longitudinally 
 along the right edge of the buccal cavity where 
 it functions in feeding (see Fig. 1). 
 
 Vestibulum A depression of the cell surface 
 bearing simple cilia used in feeding (see Fig. 
 5). 
 
 TECHNIQUES 
 
 Ciliated Protozoa occur in a wide variety of 
 environments, wherever there are sufficient nu- 
 trients, moisture, and appropriate microhabitats. 
 Ciliates can be observed easily by examining col- 
 lections from tidal pools on rocky shores and 
 marshes. They are particularly abundant both 
 intertidally and subtidally in interstices among 
 sand grains. Ciliates also reach great numbers 
 among fine filaments of some green algae (i.e., 
 Cladophora and Vaucheria) . Wide-mouthed 
 glass jars (100-2,000 cm 3 ) make excellent con- 
 tainers for nonquantitative collecting of ciliates 
 from such habitats. Add enough substratum to 
 fill the container about one-third and an equal 
 amount of water. 
 
 As soon as you can do so conveniently, distrib- 
 ute some of the material among several finger- 
 bowls with varying proportions of substratum 
 and water, to increase the variety of possible 
 ciliates. 
 
 Ciliates may be concentrated from field sam- 
 ples by filtration, moderate centrifugation, and 
 other separation methods such as electromigra- 
 tion (Hairston and Kellermann, 1964) and ex- 
 traction with seawater ice (Uhlig, 1965). 
 
 To remove ciliates from interstices of mats of 
 vegetation, filter them through two-ply cheese- 
 cloth, then concentrate them with Whatman #1 
 filter paper. As this will retain all but the small- 
 est organisms (less than 15 /j.m width), the con- 
 tents of the filter can be removed to a petri dish 
 
 for examination. Series of filters containing 
 nylon gauze of known mesh size (Nytex) allow 
 separation of most metazoans and larger frag- 
 ments of debris from the water containing cil- 
 iates. 
 
 Moderate centrifugation (5-20 min at 2,500 
 rpm) can also help to concentrate many ciliates. 
 The speed or duration of centrifugation can be 
 varied to correspond with the size and fragility 
 of the organism. 
 
 Although most ciliates undergo sexual pro- 
 cesses, they normally reproduce asexually and 
 hence can be cultured to obtain additional spec- 
 imens for further study. Cultures can be main- 
 tained in suitable covered containers. Addition 
 of simple nutrients (i.e., rice grains, wheat 
 grains, split peas, short dried lengths of grass 
 stems) will increase the growth rate of the mi- 
 croflora on which many ciliates feed. 
 
 In addition to these simple methods there are 
 many inorganic solutions and complex culture 
 media available for maintaining mixed cultures 
 of ciliates (see Kudo, 1966). 
 
 The couplets in this key are based on living 
 material and assume sufficient microscopic reso- 
 lution to allow visibility of the structures de- 
 scribed. As the organisms are small, they must 
 be examined with great care, and since living 
 ciliates often move too rapidly to allow identi- 
 fication, it is necessary to slow them down. This 
 is accomplished by removing sufficient water 
 from the edge of the preparation to squeeze the 
 animal gently between the slide and coverslip. 
 As this may cause distortion and death of the 
 cell, studies on living specimens should be aug- 
 mented with stained preparations. 
 
 Observations of living individuals of ciliates 
 can often be supplemented by appropriate fix- 
 ation and staining procedures that allow reso- 
 lution and differentiation of features difficult to 
 see in life (i.e., nuclei, details of the cilia in the 
 region of the cytostome). The simplest cyto- 
 logical methods easily employed in marine cil- 
 iates are variations of the iron hematoxylin 
 method, the Feulgen nucleal reaction (both de- 
 scribed by Kudo, 1966), and the nigrosin-mer- 
 curic chloride-Formalin technique (NMF meth- 
 od) (see Borror, 1968a, 1969). 
 
 The NMF method can be applied to fresh ma- 
 terial rapidly, giving permanently stained cells 
 in 10 or 15 min. The stain-affixative is mixed 
 
as needed and can be used repeatedly for at least 
 a month if kept cool: 
 
 HgCk, saturated aqueous, 10 ml 
 
 Glacial acetic acid, 2 ml 
 
 Formalin, cone, 2 ml 
 
 formol-nigrosin, 1-2 ml (depending upon 
 
 intensity desired) 
 t-butanol, 10 ml. 
 
 The formol-nigrosin is mixed as follows: 
 
 Formalin, cone, 20 ml 
 nigrosin, soluble, 4 g 
 distilled water, 100 ml. 
 
 The procedure for the NMF method is: Place 
 a drop of a concentrated suspension of organ- 
 isms on a clean slide and pipette onto it from 
 a height of 2-3 cm a drop of stain-fixative. After 
 a few seconds wash the culture fluid to the ends 
 of the slide by additional drops of the stain- 
 fixative. Practically all the specimens will be 
 fixed, stained, and attached to the slide, and after 
 about 15 sec the preparation may be dehydrated, 
 cleared, and covered. Ciliary organelles gener- 
 ally will appear black against a gray background, 
 cytopharyngeal rods, isolated dorsal cilia, gran- 
 ules between ciliary rows, compound buccal cilia, 
 surface ridges, and general surface morphology 
 are demonstrated (Borror, 1968a). Highly con- 
 tractile ciliates sometimes can be prepared by 
 this method following their relaxation in 8% 
 MgCh; extremely fragile ciliates often can be 
 prepared by this method following preliminary 
 fixation in Champy's fluid (Borror, 1969). 
 
 In addition, most ciliate specialists use one or 
 both of the following methods: the Chatton- 
 Lwoff technique (sometimes called the French 
 silver method or the wet silver method) and the 
 protargol (or silver proteinate) method (Corliss, 
 1953; Tuffrau, 1967a). The body form is par- 
 ticularly well preserved with the latter methods. 
 Both methods require several hours for prepa- 
 ration and careful attention to the variables of 
 time and temperature, and are part of the rep- 
 ertoire of the specialist. Although not neces- 
 sary for the use of this key generally, the Chat- 
 ton-Lwoff technique is particularly desirable for 
 identification of members of the Hymenostoma- 
 tida, whose buccal structures are difficult to dis- 
 cern in life. This technique also demonstrates 
 basal bodies of cilia and other cortical features 
 such as cytostome, pore of the contractile vac- 
 uole, cell anus, and fiberlike "silver lines" that 
 apparently are places of contiguity of cortical 
 membrane systems. The protargol method stains 
 cilia and associated fibrillar systems as well as 
 nuclei and, in association with the Chatton-Lwoff 
 technique, is particularly useful in developmental 
 studies. 
 
 Further information on the use of the micro- 
 scope for examination of ciliates, preparation of 
 slides with both fresh and permanent material, 
 as well as additional cytological techniques, can 
 be found in Kudo (1966). In the hands of a 
 nonspecialist, a small amount of 10% Methocel 
 in seawater greatly facilitates study of living 
 ciliates and does not prevent subsequent staining 
 with acidified methyl green. 
 
KEY PRIMARILY TO FAMILIES OF THE CILIOPHORA 
 
 Cilia distributed uniformly, except at the 
 anterior or posterior ends of the cell, or at 
 the vestibulum or buccal cavity. Cilia in 
 rows. Cytostome present or absent 
 
 Cilia reduced in number, 
 often nonuniform in dis- 
 tribution, leaving noncil- 
 iated surfaces. Cyto- 
 stome present or absent. 
 
 54 
 
 Cilia completely absent; no cytostome 
 
 2 (i) 
 
 2 (1) 
 
 Cilia all simple, not 
 forming compound, 
 membranous, or syn- 
 chronous units. (Note: 
 membranellelike 
 groups of cilia occur 
 on the general cell 
 surface of the ciliates 
 Placus socialis, Proro- 
 don mimeticus, and 
 member of the Family 
 NASSULIDAE) 
 
 Compound cilia present as cirri, undulating 
 membranes, peniculi, or membranelles, func- 
 tioning synchronously, in feeding and/or 
 locomotion 
 
 29 
 
3 (2) Cytostome apical, lateral, or ventral on cell surface 
 or in depression. Free-living or endocommensal. . . 
 
 3 (2) Cytostome absent. Cell oval-elongate, living in various 
 invertebrates 
 
 24 
 
 4 (3) Cytostome apical; circular or slitlike 5 
 
 4 (3) Cytostome lateral or ventral 11 
 
 5 (-4) Cytostome at base of ciliated apical vestibulum. Family 
 COELOSOMIDIDAE. 
 
 5 (4) Cytostome on general cell surface or at 
 base of nonciliated depression 
 
6 (5) Some ciliary rows terminating anteriorly against a neighbor- 
 ing ciliary row. Posterior to the cytostome along this zone 
 is a ribbon of 2-3 closely set rows of short cilia 
 
 6 (5) No such ribbon of short cilia present 8 
 
 7 (6) Cytostome oval or circular, surrounded by cilia. 
 Family PRORODONTIDAE. 
 
 7 (6) Cytostome slitlike on a nonciliated ridge offset by a groove 
 along which cilia are dense; anterior end obliquely trun- 
 cate. Family SPATHIDIIDAE. 
 
 8 (6) Cell firm, noncontractile, barrel-shaped, with armored stavelike 
 plates in cortex, usually with spines. Several cilia emerging 
 between adjacent staves. Family COLEPIDAE. 
 
 8 (6) Surface not as above. If firm, then only 1 cilium on each plate 9 
 
9 (8) Clear vacuole filling posterior end of cell. Cilia closely set, apparently in 
 whorls. Vase-shaped lorica present or absent. Surface firm, with rectan- 
 gular alveoli. Family METACYSTIDAE. 
 
 9 (5) Clear vacuole not filling posterior end of cell. Never with lorica 10 
 
 10 (9) Ciliary rows spiraled when cell contracts. Macronucleus var- 
 iously shaped with normal Feulgen reaction. Shapes various; 
 behavior generally nonthigmotactic. Family ENCHELYIDAE. 
 
 10 (9) No spiral contraction. Nuclei largely Feulgen negative. Animal generally 
 highly thigmotactic, contractile, fragile, and snakelike in shape. Family 
 TRACHELOCERCIDAE in part (also see couplet 95). 
 
 11 (A) Cytostome not in vestibulum , 12 
 
 11 U) Cytostome in vestibulum 14 
 
12 (11 ) Cilia in longitudinal rows. Free-living 13 
 
 12 (11) Ciliary rows spiraled. Cytostome radially compartmented, rosette- 
 shaped. Commensal in or on various invertebrates, especially ar- 
 thropods. Family FOETTINGERIIDAE. 
 
 13 (12) Cytostome at base of proboscis. Three bipolar aboral closely set rows of short 
 cilia. Family TRACHELIIDAE. 
 
 13 (12) Proboscis absent. Cytopharyngeal rods conspicuous. Triple dorsal rows 
 of short cilia lacking. Family NASSULIDAE. 
 
 14 (11) Some ciliary rows terminating along edge of left 
 
 half of vestibulum 15 
 
 14 (11) Ciliary rows parallel to left edge of vestibulum and 
 not terminating against its edge 
 
 18 
 
 10 
 
15 (lJf) Cell dorsoventrally flattened. Longitudinal nonciliated stripe ex- 
 tending from right corner of vestibulum at least one-half cell length 
 ventrally. Vestibulum transverse. Family PLAGIOPYLIDAE. 
 
 15 (14) Cell circular in cross section. Vestibulum oblique, triangular, or 
 circular. No such unciliated stripe 
 
 16 (15) Vestibulum triangular, anteroventral, all ciliary rows terminating against 
 it. Living in various invertebrates. Family BALANTIDIIDAE. 
 
 16 (15) Vestibulum otherwise. Free-living 17 
 
 17 (16) Vestibulum circular; ciliary rows symmetrical. Cilia in left part 
 of vestibulum in closely set radial rows independent of ciliary rows 
 on general cell surface. Family CLATHROSTOMATIDAE. 
 
 17 (16) Vestibulum generally narrow, oblique ; ciliary rows asymmetrical. 
 Cilia in left side of vestibulum apparently the modified anterior 
 part of ciliary rows to left of vestibulum. Family COLPODIDAE. 
 
 11 
 
18 (14) Ciliary rows spiraled. Fingerlike projections near cytostome. Endocommen- 
 sal in ctenophores. Pericaryon cesticola, order Trichostomatida (see Anno- 
 tated Systematic List) . 
 
 18 (14) Not as above 19 
 
 19 (18) Vestibulum elongate; right edge of vestibulum with a narrow dense bor- 
 der of cilia. Closely set longitudinal rows of cilia along left edge of ves- 
 tibulum. Macronucleus extensively Feulgen negative. Family GELEI- 
 IDAE. 
 
 19 (18) Vestibulum and its cilia otherwise 20 
 
 20 (19) Vestibulum longitudinal, narrow, about one-third to one-half body length, 
 with one row of cilia. Endocommensal in sea urchins. Schizocaryum 
 dogieli, Family PLAGIOPYLIDAE. 
 
 20 (19) Vestibulum otherwise 21 
 
 21 (20) Vestibulum a pocket in posterior half of body 22 
 
 21 (20) Vestibulum in anterior half of body 
 
 23 
 
 12 
 
22 (21 ) Vestibular ciliary rows contiguous with somatic ciliary rows to right of 
 cytostome. Endozoic in bivalves and snails. Family CONCHOPHTHIR- 
 IDAE (also see couplet 53) . 
 
 1/ 'fit&Z 1 
 
 
 22 (21) Vestibular ciliary rows contiguous with somatic ciliary rows to left of 
 cytostome. Ectocommensal on gills of bivalve molluscs. Family THIG- 
 MOPHRYIDAE (also see couplet 53). 
 
 23 (21) Cell oval. Vestibulum one-third body length from anterior end and rel- 
 atively small. Macronucleus oval. Endocommensal in bivalves. Gullmarel- 
 la faurei, order Trichostomatida (see Annotated Systematic List). 
 
 23 (21) Cell vermiform. Vestibulum large, triangular, near anterior end. Macronucleus 
 elongate. Free-living. Conchostoma longissimum, order Trichostomatida (see An- 
 notated Systematic List). 
 
 24 (3) Cell vermiform, attached by apex to renal epithelium of cephalopods. Fam- 
 ily OPALINOPSIDAE. 
 
 24 (3) Not as above. 
 
 25 
 
 13 
 
25 (24) Anterior end with holdfast apparatus of skeletal 
 elements and areas of thigmotactic cilia 
 
 26 
 
 25 (24) Holdfast apparatus, if present, without skeletal elements 27 
 
 26 (25) One or two hooks in holdfast apparatus under thigmotactic area. Cell about 
 2!/2 times longer than wide. Endocommensal in turbellarians. 
 Family HAPTOPHRYIDAE. 
 
 26 (25) No hooks in holdfast apparatus. Cell length more than 21/2 times width, 
 docommensal in gut of polychaetes. Family HOPLITOPHRYIDAE. 
 
 En- 
 
 27 (25) Macronuclei scattered. Cell pyriform. 
 Endozoic in polychaetes. Cyrtocaryiim 
 halosydnae, Family OPALINOPSIDAE. 
 
 27 (25) Macronucleus single, central. Cell variously shaped. 
 
 28 
 
 28 (27) Anterior suctorial tentacle present, variously shaped. Ectocommensal on gills 
 of bivalve molluscs. Macronucleus variably shaped. Family ANCISTRO- 
 COMIDAE in part (also see couplets 98 and 110). 
 
 14 
 
28 (27) No anterior suctorial tentacle present. Macronucleus 
 long, axial. Endocommensal in gut of worms, particu- 
 larly polychaetes. Family ANOPLOPHRYIDAE. 
 
 29 (2) A zone of 6 or more (commonly 25 
 to more than 100) membranelles 
 anterior to and to the the left of the 
 cytostome 
 
 30 
 
 29 (2) Never more than 4 membranelles present. 
 
 3(5 
 
 30 (29) Locomotory cilia organized in helical rows of cirri. Family SPIROFILIDAE 
 (also see couplet 99). 
 
 30 (29) Locomotory cilia not as above 31 
 
 31 (30) Cell loricate, sessile. Family FOLLICULINIDAE. 
 
 31 (30) Cell aloricate, generally free-swimming. 
 
 15 
 
32 (31) Conspicuous unciliated buccal cavity between zone of mem- 
 
 branelles and large undulating membrane 33 
 
 32 (31) Essentially no unciliated surface between right border of zone of 
 membranelles and adjacent ciliary rows 
 
 33 (32) Buccal cavity triangular. Macronucleus usually beaded. Family CON- 
 DYLOSTOMATIDAE. 
 
 33 (32) Buccal cavity circular. Macronucleus ribbonlike. Family CHATTONI- 
 DIIDAE. 
 
 34 (32) Zone of membranelles apical, spiraled, enclosing ciliated field. Family 
 STENTORIDAE. 
 
 34 (32) Zone of membranelles longitudinal, or spiraled, and not apical 35 
 
 16 
 
35 (3 %) Zone of membranelles longitudinal. Ciliary rows to right of zone of mem- 
 branelles uniformly spaced. Family SPIROSTOMATIDAE. 
 
 35 (34) Zone of membranelles spiraling 90°-180° to the right, paralleled anteriorly by 
 4-5 closely set rows of cilia. Family METOPIDAE. 
 
 36 (29) Cytostome in anterior two-thirds of cell, usually in central one-third of body. 
 
 Free-living or endocommensal 37 
 
 36 (29) Cytostome in posterior one-fifth of cell. Endocommensal in various invertebrates. . . 49 
 
 37 (36) One to three peniculi along left side of buccal cavity 
 or winding across floor of buccal cavity. Undulating 
 membrane inconspicuous. Cell densely ciliated, often 
 over 150 fim long, with sharply delimited buccal cavity. 
 
 38 
 
 37 (36) No peniculi present. Undulating membrane present 
 along right edge of buccal cavity. Remainder of buc- 
 cal cilia variously arranged. Cell of various sizes. . . . 
 
 40 
 
 17 
 
38 (37) Three peniculi winding across buccal cavity floor 39 
 
 m 
 
 38 (37) One to three short longitudinal peniculi on left side of buccal cavity. 
 Family FRONTONIIDAE. 
 
 39 (38) Buccal cavity semispherical. Family OPHRYOGLENIDAE. 
 
 39 (38) Buccal cavity tubular, preceded by ciliated vestibulum. 
 CIIDAE. 
 
 Family PARAME- 
 
 40 (37) Postbuccal ciliary rows present. Buccal cavity round to triangu- 
 lar, in anterior one-half of body. Undulating membrane L- 
 shaped. Three buccal membranelles perpendicular to long axis 
 of body 41 
 
 40 (37) No postbuccal ciliary rows. Buccal cavity elongate; cytostome 
 often at or posterior to middle of body. Undulating membrane 
 various. Other buccal cilia in various groups different from 
 above 42 
 
 18 
 
41 (40) Cilia to right of undulating membrane appear as transverse rows. 
 Contractile vacuole midventral and central. Family LOXOCEPHALI- 
 DAE. 
 
 41 (40) No transverse rows apparent at level of buccal cavity, 
 posterior. Family TETRAHYMENIDAE. 
 
 Contractile vacuole 
 
 42 (40) Undulating membrane long, saillike. Other buccal cilia variously posi- 
 tioned but not immediately adjacent to undulating membrane. Family 
 PLEURONEMATIDAE in part (also see couplet 52). 
 
 42 (40) Undulating membrane short, linear, or C-shaped, not extending beyond cyto- 
 stome. Other buccal cilia closely adjacent to undulating membrane 
 
 43 
 
 43 (42) Buccal cilia appearing as 2 dense parallel longitudinal 
 
 series, neither more than 2 ciliary rows wide 44 
 
 43 (42) Buccal cilia appearing as a single right-hand undulating mem- 
 brane and 2 to several membranelles usually 2 cilia or more wide. 
 
 i i 
 
 % 
 
 o 
 
 45 
 
 19 
 
44 (43) Buccal cavity narrow; left-hand longitudinal series part of first ciliary row to left 
 of cytostome. Family COHNILEMBIDAE. 
 
 44 (43) Buccal cavity wide. Left portion of buccal cilia a membrane 1-2 cilia wide, 
 variously segmented into 1-3 linear parts, distinct from somatic ciliary rows. 
 Family PSEUDOCOHNILEMBIDAE. 
 
 45 (A3) Cell flat on one side with caudal projection. Endozoic in sea urchins. 
 
 45 (43) Cell circular in cross section, without caudal projection. Including free-living 
 and endocommensal forms 
 
 46 
 
 48 
 
 46 (45) Anterior end truncate. Buccal cavity on small side, near anterior end, over- 
 hung by frontal lobe. Entorhipidium, Family ENTORHIPIDIIDAE. 
 
 46 (45) Anterior end rounded. Buccal cavity variously located. 
 
 47 (46) Ciliary rows longitudinal, equidistant. Buccal cavity small in anterior one- 
 fourth of cell. Caudal projection small with 1 caudal cilium. Entodiscus, 
 order Scuticociliatida (see Annotated Systematic List). 
 
 47 (46) Ciliary rows asymmetrical. Buccal cavity elongate, variously located. 
 Caudal prolongation varying in size, with one to many cilia. Crypto- 
 chilum, Cryptochilidium, order Scuticociliatida (see Annotated System- 
 atic List). 
 
 20 
 
48 (45) Cell length variable, commonly over 70 /urn. Anterior end tapered 
 to blunt point. Right dorsolateral surface typically concave. Mem- 
 branelles massive; cytostome anterior to equator. Family PHIL- 
 ASTERIDAE. 
 
 48 (45) Cell usually under 50 fim in length. Anterior end truncated, con- 
 spicuously naked. Right dorsolateral surface typically convex. 
 Membranelles inconspicuous. Cytostome often subequatorial. 
 Family URONEMATIDAE. 
 
 49 (36) Anterior end with nonciliated V-shaped sucker. Endozoic in oligo- 
 chaetes. Family HYSTEROCINETIDAE. 
 
 49 (36) No such anterior nonciliated sucker 50 
 
 50 (49) Cell flattened, with distinct caudal projection. Endozoic in sea urchins. Big- 
 garia (see Annotated Systematic List). 
 
 50 (49) Cell oval or circular in cross section. Posterior end rounded 51 
 
 21 
 
51 (50) Undulating membrane long, extending in a spiral along the periphery 
 of the shallow buccal cavity 
 
 52 
 
 51 (50) Undulating membrane short, linear, inconspicuous, in an oblique 
 vestibulum near posterior end 
 
 53 
 
 52 (51) Undulating membrane closely paralleled by ribbon of cilia, 
 2-3 cilia wide. Cytostome at or near posterior pole. On 
 gills of bivalve molluscs or endozoic in echinoderms. Family 
 HEMISPEIRIDAE in part (also see couplet 105). 
 
 52 (51) Buccal cilia do not closely parallel undulating membrane; 
 some arranged as transverse membranelles in forms of V 
 or Y near cytostome. Commensal on hydroids and mus- 
 sels. PLEURONEMATIDAE, in part, (Pleurocoptes, 
 Peniculistoma) (also see couplet 42). 
 
 53 (51) Ciliary rows in vestibulum contiguous with somatic rows 
 to right of cytostome. Endozoic in mussels. Family CON- 
 CHOPHTHIRIDAE (also see couplet 22). 
 
 53 (51) Ciliary rows in vestibulum contiguous with somatic cilium 
 rows to left of cytostome. On gills of bivalve molluscs. 
 Family THIGMOPHRYIDAE (also see couplet 22). 
 
 22 
 
54 (1) 
 
 54 (1) 
 
 Cilia mostly anteri- 
 or or apical. Circ- 
 lets of simple cilia 
 sometimes occur- 
 ring at the aboral 
 pole or near the 
 middle of the body. 
 Cytostome present. . 
 
 55 
 
 Cilia restricted to one surface, or in circlets, parallel oblique rows, isolated 
 groups of rows, or some other arrangement. Cytostome present or absent. . , 
 
 83 
 
 55 (5-4) Cell loricate 56 
 
 55 (54) Cell without lorica 72 
 
 56 (55) Free-swimming, typically planktonic 57 
 
 56 (55) Sessile; lorica attached to substratum 71 
 
 w 
 
 57 (56) Lorica with particles of foreign or self-constructed 
 
 material on surface or within its layers 58 
 
 57 (56) Lorica without particles, but with a superficial reticulated pattern 
 
 or polygonal network 62 
 
 57 (56) Lorica clear; ringed, striated, or simple, but not granulated 
 
 or reticulated 66 
 
 23 
 
58 (57) Collar present, hyaline, with ring or spiral structure. Lorica top- 
 shaped; aboral end of lorica rounded to pointed; wall of bowl 
 much thicker than that of collar. Family CODONELLOPSIDAE. 
 
 58 (57) Collar without ring or spiral structures, or absent. 
 
 59 
 
 59 (58) Lorica with small prisms and gelatinous mantle with scattered inclusions. 
 Three longitudinal folds on posterior half of lorica. Collar present. Family 
 TINTINNIDAE, in part (Brandtiella) (also see couplet 70). 
 
 59 (58) Not as above. Lorica without longitudinal folds. 
 
 60 
 
 60 (59) Posterior narrowed horn or tail. No collar. Family COXLIELLIDAE, in 
 part (Poroecus) (also see couplet 66). 
 
 60 (59) Not as above. Lorica without tail 61 
 
 61 (60) Lorica tubular and sacular, soft, gelatinous, thickly beset with particles. 
 Lorica straight sided or flaring. Family TINTINNIDIIDAE. 
 
 24 
 
61 (60) Lorica firm, with or without collar. Aboral end may have a horn or 
 be pointed. Family CODONELLIDAE. 
 
 62 (57) Collar with 1 or 2 rows of holes. Lorica semispherical or oval. 
 Family DICTYOCYSTIDAE. 
 
 62 (57) Collar without holes. Lorica sometimes pointed posteriorly. 
 
 63 
 
 63 (62) Reticulum ending anteriorly in free, clearly differentiated lines. 
 Lorica acorn-shaped, with blunt, pointed or horned aboral end. 
 Wall thin and hyaline, deeply reticulated. Family EPIPLO- 
 CYLID1DAE. 
 
 63 (62) Anteriorly just as reticulated as rest of lorica 64 
 
 64 (63) Collar flared, set off by constriction. Oral edge slightly toothed or wavy. 
 Lorica acorn-shaped, with posterior point. Family CYTTAROCYLIDIDAE. 
 
 64 (63) No flared collar. Oral edge with or without teeth. Lorica kettle-shaped or 
 chalice-shaped 
 
 65 
 
 25 
 
65 (64) One or two anterior suboral swellings. Lorica Kettle-shaped; bowl elongate 
 with suddenly pointed posterior. Family PTYCHOCYLIDIDAE, in part 
 (Ptychocylis) (also see couplet 69). 
 
 65 (64) No anterior suboral swellings. Lorica elongate, chalice-shaped, 
 with long, narrow aboral pedicle. Family XYSTONELLIDAE. 
 
 66 (57) A spiral band over at least oral one-half of lorica. Family 
 COXLIELLIDAE in part (also see couplet 60). 
 
 66 (57) Lorica simple, ribbed or ringed, but without spiral structures. 
 
 67 
 
 67 (66) Lorica short, cup- or goblet-shaped 68 
 
 67 (66) Lorica long, bell- to trumpet-shaped 69 
 
 26 
 
68 (67) Oral margin built into outer collar and inner shelf. 
 Lorica clear. Family PETALOTRICHIDAE. 
 
 68 (67) Oral margin simple, without collar and shelf , and narrowed. 
 Lorica simple or with rings. Family UNDELLIDAE. 
 
 69 (67) Wall trilaminate. Oral margin not flared, and without ribs or striae. Lorica 
 bell- or kettle-shaped; bowl typically cylindrical, elongate, with posterior point 
 or horn. Family PTYCHOCYLIDIDAE, in part (Favella) (also see couplet 
 65). 
 
 69 (67) Wall simple, two layered. Oral margin flared. Ribs or striae often present. 
 Lorica cup- to spindle-shaped 
 
 70 
 
 70 (69) Oral aperture slightly flared, with broad, grooved rim. Longitudinal ribs 
 or striae always present. Lorica cone- or chalice-shaped. Aboral end 
 closed. Family RHABDONELLIDAE. 
 
 n 
 
 70 (69) Oral aperture flared, simple. Longitudinal ribs sometimes 
 present. Lorica elongate, usually vase-shaped. Aboral end 
 open or closed. Family TINTINNIDAE in part (also see coup- 
 let 59). 
 
 27 
 
71 (56) Cell attached to lorica only at mouth aperture; only buccal cilia 
 capable of being extended through opening of lorica. Stalk absent. 
 Family LAGENOPHRYIDAE. 
 
 71 (56) Cell not attached to oral end of aperture; entire oral end of body extendable 
 beyond opening of lorica. Stalk present, often short. Family VAGINICO- 
 LIDAE. 
 
 72 (55) Cell motile 73 
 
 72 (55) Cell sessile 77 
 
 73 (72) Aboral circlet (s) of cilia present. 
 
 . 74 
 
 73 (72) No aboral circlet of cilia. Sometimes an equatorial circlet of cilia. . . 76 
 
 74 (73) Aboral adhesive disc strengthened by skeletal ring. 
 Family URCEOLARIIDAE 
 
 75 
 
 28 
 
74 (73) No aboral adhesive disc. Migratory motile stages of ciliates of order Peri- 
 trichida, suborder Sessilina. 
 
 75 (74) Skeletal ring with 20-50 smooth links. Subfamily Urceolarinae. 
 
 75 (7 A) Skeletal ring of hooked or spiked links. Subfamily 
 Trichodininae. 
 
 76 (73) Cytostome on ventral surface, not surrounded by zone of membranelles. 
 Family HALTERIIDAE. 
 
 76 (73) Cytostome apical, enclosed within spiral zone of membranelles. Family 
 STROBILIDIIDAE. 
 
 77 (72) Zone of membranelles winding at least 360° to anteroventral cytostome. 
 Family LICNOPHORIDAE. 
 
 77 (72) Buccal cilia not in a clockwise zone of membranelles 78 
 
 29 
 
78 (77) Buccal cilia in apical counterclockwise groups that descend 
 helically down a deep, tubular buccal cavity. Animal cup- or 
 bell-shaped, solitary or colonial 79 
 
 78 (77) Buccal cilia inconspicuous, in groups within apical cone or funnel. 
 
 Animal cylindrical or spindle-shaped, with flaring apex, noncolonial 81 
 
 79 (78) A differentiated stalk present 80 
 
 79 (78) Stalk absent. Family SCYPHIDIIDAE. 
 
 80 (79) Stalk contractile. Family VORTICELLIDAE. 
 
 80 (79) Stalk not contractile. Family EPISTYLIDIDAE. 
 
 30 
 
81 (78) Stalk at least as long as body. Family CHILODOCHONIDAE. 
 
 81 (78) Stalk less than one-fourth length of body or absent 82 
 
 82 (81) Apical funnel in 3 spirals. Family SPIROCHONIDAE. 
 
 82 (81) Apical funnel single or double, and not spiraled. Family STYLOCHONIDAE. 
 
 83 (54) Cilia restricted to ventral surface. Dorsal surface non- 
 ciliated, with cilia only at borders, or with sparse rows 
 or groups of cilia not used in locomotion. (These dia- 
 grams show cross sections of animals) 
 
 84 
 
 83 (5 A) Cilia not restricted to one surface, in cir- 
 clets, parallel oblique rows, isolated groups 
 of rows, or some other arrangement 
 
 104 
 
 31 
 
84 (83) Locomotory cilia simple, in rows 85 
 
 84 (83) Locomotory cilia organized as cirri in rows/or groups. 
 
 99 
 
 85 (84) Cytostome ventral. Body seldom twice as long as wide 86 
 
 85 (84-) Cytostome anterior, lateral on narrow edge, or 
 
 absent. Body of various shapes. 92 
 
 86 (85) Cytopharyngeal rods present 87 
 
 86 (85) Cytopharyngeal rods absent 88 
 
 32 
 
87 (86) Posteroventral stylus or secretory organelle of attachment pres- 
 ent. Usually 2 cytopharyngeal rods. Family DYSTERIIDAE. 
 
 87 (86) No such stylus present. Cytopharyngeal rods in conical group. Family 
 CHLAMYDODONTIDAE. 
 
 88 (86) Zone of membranelles present. 
 
 89 
 
 88 (86) Zone of membranelles absent 91 
 
 89 (88) Cell firm, grooved, buccal cavity midventral, with 3 membranelles. Family 
 CINETOCHILIDAE. 
 
 89 (88) Cell soft, flexible, buccal cavity otherwise. 
 
 90 
 
 33 
 
90 (89) Zone of membranelles bordering anterior end. Length over 70 fim. 
 Family PERITROMIDAE. 
 
 90 (89) Zone of membranelles reduced, limited to buccal cavity. Length under 70 pm 
 Protocruzia, order Heterotrichida (see Annotated Systematic List). 
 
 91 (88) Cilia ventral in dense curving band. Motile larval stages in life cycle of 
 members of the order Chonotrichida. 
 
 91 (88) Cilia in C-shaped rows along borders of cell. Askoella, 
 Conidiophrys (tomite only), and Phthorophyra. Family 
 ASKOELLIDAE (also see couplet 112). 
 
 92 (85) Cytostome apical or lateral often inconspicuous 93 
 
 92 (85) Cytostome absent. 
 
 / 
 97 
 
 93 (92) Cytostome anterior 
 
 93 (92) Cytostome a lateral slit. 
 
 94 
 96 
 
 34 
 
94 (93) At least some of cilia in bipolar rows. Macronuclei small, vesicular 95 
 
 94 (93) Ciliary rows in a ventral band about one-half cell width, extending about 
 four-fifths cell length. Macronucleus single, large, C-shaped. Family 
 CRATERISTOMATIDAE. 
 
 95 (9U) Body contractile and snakelike in shape. Cytostome circular. Family TRA- 
 CHELOCERCIDAE, in part (also see couplet 10). 
 
 95 (9b) Cell length less than 50 ju,m, humped dorsally. Cytostome anterior 
 and slitlike. Family STEPHANOPOGONIDAE. 
 
 96 (93) Cytostome on concave left edge of body, preceded by hooklike lateral ex- 
 tension. Nuclei vesicular. No contractile vacuole. Family LOXODIDAE. 
 
 96 (93) Cytostome on convex left edge. Macronuclei one to several, not vesic- 
 ular. One to several contractile vacuoles. Family AMPHILEPTIDAE. 
 
 35 
 
97 (92) Anterior suctorial tentacle or sucking disc present. . 98 
 
 97 (92) No anterior suctorial tentacle 
 Annotated Systematic List). 
 
 Silenella ovoidea, order Suctorida (see 
 
 98 (97) Cilia restricted to one flattened surface. A short (about 8 cilia) 
 ciliary row at anterolateral margin. Commensal on hydroids, 
 and in branchial cavity of ascidians. Family HYPOCOMIDAE. 
 
 98 (97) Cell ovoid to pyriform in shape. No such short ciliary row, or ciliary 
 rows more extensive. Ectocommensal on gills of bivalve molluscs. Family 
 ANCISTROCOMIDAE (also see couplets 28 and 110). 
 
 99 (84) Helical torsion reduces aboral surface. Ventral cirri in helical rows. 
 Family SPIROFILIDAE (also see couplet 30). 
 
 99 (84) Aboral surface not so reduced, usually bearing sparse, widely spaced rows of 
 simple (often very short) cilia not used in locomotion 
 
 100 
 
 100 (99) Cirri of midventral axis arranged in a closely set double row — actually a 
 single zigzag series. Family HOLOSTICHIDAE. 
 
 36 
 
 i 
 
100 (99) Cirri of midventral axis in single rows or isolated groups, but not in a zigzag 
 
 series 101 
 
 101 (100) Practically all cirri arranged in parallel rows 102 
 
 101 (100) Cirri of midventral surface arranged in groups. Marginal cirrus rows some- 
 times present 103 
 
 102 (101) Cell oval, with large buccal cavity. Rows of cirri curved, paralleling 
 edges of buccal cavity. Undulating membrane well developed. Fam- 
 ily PSILOTRICHIDAE. 
 
 102 (101) Cell more elongate, cirri in parallel rows that extend well beyond the buccal 
 cavity. Family UROSTYLIDAE. 
 
 103 (101) At least 2 rows of cirri present, each with more than 5 (usually 20 or more) 
 cirri in addition to groups of cirri. Family OXYTRICHIDAE. 
 
 103 (101) No such marginal rows present, or at least 
 1 row reduced to fewer than 5 cirri; remain- 
 ing ventral cirri in groups. Family EUPLO- 
 TIDAE. 
 
 37 
 
104 (83) 
 
 104 (83) 
 
 Cytostome apical, 
 whorls 
 
 Cilia in circlets or spiraling 
 
 Cytostome nonapical or ab- 
 sent. Cilia in parallel oblique 
 rows, isolated groups of rows 
 or some other arrangement. . 
 
 106 
 
 105 (104) Cytostome on cell surface, often on a conical 
 prominence. Retractile seizing organelles in 
 some species. Cilia in circlets. Family 
 DIDINIIDAE. 
 
 105 (10^) Cytostome in apical vestibulum. 
 Family TRIMYEMIDAE. 
 
 Cilia in spiraling whorls 
 
 105 (104) Cytostome at posterior pole. Prominent spiraling undulating mem- 
 brane present. Anterior group of thigmotactic cilia present. Ecto- 
 commensal on gills of bivalve molluscs. Family HEMISPEIRIDAE, 
 in part (also see couplet 52). 
 
 106 (104) Cell laterally compressed, with rigid pellicle. Cytostome 
 
 and compound cilia present 107 
 
 106 (104) Cell without rigid pellicle or cytostome. All cilia simple 109 
 
 38 
 
107 (106) Zone of membranelles in spiraling buccal cavity, paralleled anteriorly 
 by 5 closely set rows of cilia. One to three posterior spines present. 
 Family CAENOMORPHIDAE. 
 
 107 (106) A rectangular group of 9-12 membranelles in small buccal cavity. Seven 
 rows of prebuccal cilia. No posterior spines present 
 
 108 
 
 108 (107) All somatic cilia simple. Family EPALXELLIDAE. 
 
 108 (107) One to two posterior cirri present. Family MYLESTOMATIDAE. 
 
 109 (106) Apical suctorial tentacle or sucking disc present. Ciliary rows 
 straight or spiraled, usually only one-third to two-thirds cell 
 length, terminating near suctorial tentacle. Ectocommensal on 
 gills of bivalve molluscs 
 
 110 
 
 109 (106) No suctorial tentacle or sucking disc. Cilia in 1-3 
 fields, including a ribbon of 4-15 parallel oblique 
 ciliary rows encircling cell like a neck scarf. Cili- 
 ated larvae, order Suctorida. (Families separated 
 by adult morphology, couplets 112-121.) 
 
 39 
 
110 (109) Ciliary rows equidistant, usually about two-thirds cell length. Family 
 ANCISTROCOMIDAE (also see couplets 28 and 98). 
 
 110 (109) Ciliary rows in 2 longitudinal groups of 4-6 short ciliary rows each. . Ill 
 
 111 (110) Cell pyriform. Tomite of member of family SPHENOPHRYIDAE (also 
 see couplet 113). 
 
 Ill (110) Cell laterally compressed. Ridge separating the 2 ciliated surfaces. 
 Family SPHENOPHRYIDAE (Lwoffia cilifera). 
 
 112 (1) One to many sucking tentacles present 113 
 
 112 (1) No sucking tentacles present. Attached to exoskeletal hairs of crustaceans. 
 Family ASKOELLIDAE (Conidiophrys) (also see couplet 91). 
 
 40 
 
113 (112) One short sucker present. Attached to gills of bivalve molluscs 
 in part by large, flat sole. Family SPHENOPHRYIDAE (also 
 see couplet 111). 
 
 113 (112) Several tentacles present. In various habitats 114 
 
 ; 
 
 114 (113) Cell with lorica 115 
 
 0^ 
 
 114 (113) Cell without lorica 117 
 
 115 (114) Tentacles of two definite types. Family EPHELOTIDAE, in part 
 (Actinocyathus, Podocyathns) (also see couplet 118). 
 
 115 (114) Tentacles all similar 116 
 
 116 (115) Tentacles with enlarged tips, in clusters. Family ACINETIDAE 
 
 116 (115) Tentacles not clustered. Family PODOPHRYIDAE, in part (Paracineta, 
 Metacineta) (also see couplet 120). 
 
 41 
 
117 (11A) Stalk present by which cell is attached to substratum 118 
 
 117 (lib) Stalk absent. Cell attached to substratum by broad flat- 
 tened surface 
 
 121 
 
 118 (117) Tentacles of two definite types. Family EPHELOTIDAE (also see 
 couplet 115). 
 
 118 (117) Tentacles all similar 119 
 
 119 (118) Cell beyond stalk elongate, at least twice as long as broad. Stalk usu- 
 ally short. Family OPHRYODENDRIDAE. 
 
 119 (118) Cell beyond stalk less than twice as long as broad, often 
 globular. Stalk often as long as rest of cell 
 
 120 
 
 42 
 
120 (119) Bud developing within cavity of parental suctorian. Family 
 DISCOPHRYIDAE. 
 
 120 (119) Bud developing externally on parental suctorian. 
 Family PODOPHRYIDAE (also see couplet 116). 
 
 121 (117) Tentacles minute, on surface next to substratum, acting only as 
 organelles of attachment. On appendages of polychaetes. Family 
 PHALACROCLEPTIDAE. 
 
 121 (117) Tentacles used in feeding, and in clusters on superior surface. 
 Family DENDROSOMATIDAE. 
 
 43 
 
ANNOTATED 
 SYSTEMATIC LIST 
 
 The following list is arranged according to the 
 classification used in The Ciliated Protozoa (Cor- 
 liss, 1961), with modifications based on more 
 recent revisions. No ranges are given since 
 most species are cosmopolitan, their distribu- 
 tions depending more on microhabitat than on 
 macrodistributional factors. Notes on system- 
 atics and ecology are given when they are im- 
 portant in identification of the taxa. Important 
 papers on a given family are also mentioned 
 (see Selected Bibliography). 
 
 Class CILIATA 
 
 Order Gymnostomatida 
 
 Family AMPHILEPTIDAE Butschli, 1889. 
 Four genera with marine representatives: Am- 
 phileptus, Hemiophrys, Litonotus, and Loxophyl- 
 lum; all free-living and benthic. See Kahl 
 (1931) and Canella (1960). 
 
 Family CHLAMYDODONTIDAE Stein, 1859. 
 At least eight genera with marine representa- 
 tives: Atopochilodon, Chilodonatella, Chilodo- 
 nella, Chlamydodon, Chlamydonella, Lynchella, 
 Odontochlamys, and Parachilodonella. Free-liv- 
 ing and benthic. See Dobrzanska-Kaczanowska 
 (1963), Dragesco (1966, 1967), and Deroux 
 (1970). 
 
 Family COLEPIDAE Ehrenberg, 1838. Free- 
 living, widespread. Two genera: Coleps and 
 Tiarina; both marine. See Kahl (1930). 
 
 Family DIDINIIDAE Poche, 1913. Ten gen- 
 era described, mostly marine: Acropisthium, 
 Askenasia, Choanostoma, Ctenoctophrys, Cyclo- 
 trichium, Didinium, Liliimorpha, Mesodinium, 
 Monodinium, and Zonotrichium. Mesodinium 
 widespread and ubiquitous; planktonic, but also 
 occurs near substrata. See Kahl (1930). 
 
 Family DYSTERIIDAE Claparede and Lach- 
 mann, 1858. Free-living and benthic, or ecto- 
 commensal on crustaceans and fish. At least 
 eight genera with marine representatives: 
 Brooklynella, Dysteria, Hartmannula, Mirodys- 
 teria, Paratrochilia, Sigmocineta, Trochilia, and 
 Trochilioides. See Jankowski (1967a). 
 
 Family ENCHELYIDAE Ehrenberg, 1838. 
 Free-living, widespread. This is a large, loosely 
 defined family in need of serious systematic re- 
 vision. Genera with marine species: Chaenea, 
 Chilophrya, Enchelyodon, Enchelys, Helicopro- 
 rodon, Holophrya, Ileonema, Lacrymaria, Lagy- 
 nurus, Longitricha, Microregma, Nannophrya, 
 Pithothorax, Placus, Plagiocampa, Platyophrya, 
 Rhopalophrya, and Spasmostoma. See Faure- 
 Fremiet (1961c). 
 
 Family LOXODIDAE Butschli, 1889. Four 
 genera with marine representatives: Ciliofau- 
 rea, Kentrophoros, Loxodes, and Remanella; all 
 free-living, benthic, usually among sand grains. 
 Kentrophoros is a ribbonlike psammolittoral 
 ciliate usually included in this family, but it is 
 of uncertain affinities since its cytostome has yet 
 to be described. Cryptopharynx, usually placed 
 in the family CHLAMYDODONTIDAE also 
 keys out here. See Dragesco (1960). 
 
 Family METACYSTIDAE Kahl, 1926. Three 
 genera: Metacystis, Pelatractus, and Vasicola. 
 Metacystis occurs regularly in tidal marshes. 
 See Kahl (1930). 
 
 Family NASSULIDAE de Fromentel, 1874. 
 Five genera with marine species: Chilodontop- 
 sis, Nassula, Nassulopsis, Orthodonella, and 
 Stomatophrya. Free-living. See Agamaliev 
 (1967) and Dragesco (1960). 
 
 Family PRORODONTIDAE Faure-Fremiet, 
 1961. This family, split from the family 
 ENCHELYIDAE by Faure-Fremiet in 1961 
 (1961c), contains at least the following genera, 
 all with marine species: Lagynophrya, Pro- 
 rodon, Trachelophyllum, and Urotricha. Free- 
 living, widespread. 
 
 Family SPATHIDIIDAE Kahl in Doflein and 
 Reichenow, 1929. At least ten genera (listed 
 in Corliss, 1961). Free-living and benthic. See 
 Kahl (1930). 
 
 Family STEPHANOPOGONIDAE Corliss, 
 1961. One genus, Stephanopogon, free-living in 
 marine sands and algal mats. See Dragesco 
 (1963b) and Borror (1965b). 
 
 Family TRACHELIIDAE Ehrenberg, 1838. 
 At least three marine genera: Dileptus, Para- 
 dileptus, and Trachelitis; all free-living. See 
 Dragesco (1963a). 
 
 44 
 
Family TRACHELOCERCIDAE Kent, 1880. 
 Six genera: Gruvelina, Nephrocerca, Protrieh- 
 ophora, Trachelocerca, Tracheldnema, and Trw- 
 cheloraphis; all free-living, marine, and char- 
 acteristic of sandy substrata. See Dragesco 
 (1960). 
 
 > 
 
 Order Trichostomatida 
 
 Family BALANTIDIIDAE Reichenow, 1929. 
 One genus, Balantidium; free-living and in the 
 gut of polychaetes and amphipods and the gas- 
 trovascular cavity of medusae. See Kahl (1934a) 
 and Faure-Fremiet (1955). 
 
 Family CLATHROSTOMATIDAE Kahl, 
 1926. This family (also known as PARANAS- 
 SULIDAE Faure-Fremiet, 1961) is placed here 
 rather than in the Gymnostomatida as the ves- 
 tibular ciliature used in feeding is distinct from 
 the somatic cilia. As some of these ciliates may 
 possess structures interpretable as peniculi, their 
 inclusion in the Trichostomatida is artificial. 
 One genus with marine species, Paranassula,. 
 Free-living. See Faure-Fremiet (1962a). Gull- 
 marella faurei Fenchel, 1964, endocommensal 
 in bivalves, is of uncertain position in this order 
 since little is understood of its morphogenesis, 
 but may also belong in this family. See Fenchel 
 (1964). 
 
 Family COELOSOMIDIDAE Corliss, 1961. 
 Three genera with marine representatives: 
 Coelosomides, Paraspathidium, and Pseudopro- 
 rodon; free-living. Conchostoma longissimum 
 Faure-Fremiet, 1963, is a species of uncertain 
 taxonomic position in this order, but may belong 
 in this family. 
 
 Family COLPODIDAE Ehrenberg, 1838. 
 Contains Woodruffia (in estuaries) and Colpoda 
 (typically a soil ciliate, but sometimes found in 
 tidal marshes). Free-living. See Kahl (1931) 
 and Prelle (1963). 
 
 Family GELEIIDAE Kahl, 1933. Two gen- 
 era: Corlissia and Geleia; free-living in ma- 
 rine sands and algal mats. The genus Corlissia 
 is of questionable systematic position, since little 
 is understood of the cilia in the region of the 
 cytostome. See Dragesco (1960). 
 
 Family PLAGIOPYLIDAE Schewiakoff, 1896. 
 At least four genera with marine species: Lech- 
 
 riopyla, Plagiopyla, Plagiopyliella, and Sonderia. 
 Free-living or endocommensal in sea urchins. 
 See Kahl (1933). Schizocaryum dogieli Pol- 
 jansky and Golikova, 1957, endocommensal in 
 sea urchins, may also be a member of the family 
 PLAGIOPYLIDAE. See Berger (1961d). 
 
 Family TRIMYEMIDAE Kahl, 1933. One 
 genus, Trimyema. Free-living. See review by 
 Faure-Fremiet (1962b). 
 
 Pericaryon cesticola Chatton, 1911 was de- 
 scribed as endocommensal in the ctenophore, 
 Venus' girdle, Cestus veneris. This species has 
 been regarded as being in the family FOET- 
 TINGERIIDAE (order Apostomatida), but 
 possesses a vestibulum and lacks a cytostomal 
 rosette. See Chatton and Lwoff (1935). 
 
 Order Chonotrichida 
 
 Family CHILODOCHONIDAE Wallengren, 
 1895. One genus, Chilodochona; ectocommensal 
 on the exoskeleton of Ebalia and Portunus. See 
 Faure-Fremiet, Rouiller, and Gauchery (1956). 
 
 Family SPIROCHONIDAE Stein, 1854. One 
 genus, Spirochona; ectocommensal on pleopodal 
 bristles of gammaridean amphipods. See Ma- 
 tsudo and Mohr (1968). 
 
 Family STYLOCHONIDAE Mohr, 1948. Five 
 genera with marine species: Heliochona, Ken- 
 trochona, Lobochona, Oenophorachona, and Sty- 
 lochona; ectocommensal on crustaceans (includ- 
 ing Nebalia and Limnoria). See Matsudo and 
 Mohr (1965) and Mohr, Levague, and Matsudo 
 (1963). 
 
 Order Suctorida 
 
 Family ACINETIDAE Stein, 1859. At least 
 eight genera with marine species: Acineta, 
 Acinetopsis, Dactylophrya, Endosphaera, Pot- 
 tsiocles, Pseudo gemma, Tachyblaston, and The- 
 cacineta. Free-living and ectocommensal. Mi- 
 gratory stages formed by internal (endogenous) 
 budding. See Kahl (1934b). 
 
 Family DENDROSOMATIDAE Fraipont, 
 1878. A large family of at least 13 genera, of 
 which only Lernaeophrya and Trichophrya have 
 marine representatives. Budding endogenous. 
 
 45 
 
Family DISCOPHRYIDAE Collin, 1912. At 
 least 14 genera, of which but Corynophrya and 
 Thaumatophin/a occur in marine environments 
 
 Family EPHELOTIDAE Kent, 1880. Four 
 genera with marine species: Actinocyathus, 
 Ephelota, Metephelota, and Podocyathus. Bud- 
 ding exogenous, apical, and multiple. Free- 
 living and ectocommensal on hydroids. See Kahl 
 (1934b). 
 
 Family OPHRYODENDRIDAE Stein, 1867. 
 Five genera with marine species: Collin ophry a, 
 Dendrosomides, Ophry odendron, Rhabdophrya, 
 and Trophogemma. Free-living and ectocom- 
 mensal. Budding exogenous. See Jankowski 
 (1970). 
 
 Family PHALACROCLEPTIDAE Kozloff, 
 1966. One species, Phalacrocleptes verrucifor- 
 mis, originally considered a member of the order 
 Thigmotrichida. Ectocommensal on the poly- 
 chaete Schizobranchia. See Kozloff (1966). 
 
 Family PODOPHRYIDAE Haekel, 1866. At 
 least five genera with marine species: Lecano- 
 phrya, Ophry ocephalus, Paracineta, Parapodo- 
 phrya, and Podophrya. Free-living and ectocom- 
 mensal. Migratory stages formed by external 
 (exogenous) budding. Development of stalk 
 may vary within a genus. 
 
 Silenella ovoidea Fenchel, 1965 (couplet 97) 
 may be a ciliated motile stage of an unidentified 
 suctorian. Described as ectocommensal on an 
 amphipod. See Fenchel (1965b). 
 
 Order Apostomatida 
 
 Family ASKOELLIDAE Jankowski, 1967. 
 Three genera: Askoella, Conidiophrys, and 
 Phthorophrya. Members of the genus Askoella 
 are commensal on barnacles and amphipods 
 (Fenchel, 1965b; Jankowski, 1967a) and accord- 
 ing to Jankowski represent neotenic tomites of 
 the order Apostomatida with a secondary cyto- 
 stome. Conidiophrys has been considered to be 
 a trichostome (Chatton and Lwoff, 1936). The 
 trophont bears no cilia and is attached to hairs 
 and papillae of appendages of amphipods and 
 isopods. According to Jankowski (1967b) the 
 ciliated stages resemble members of the order 
 Apostomatida but with a significantly different 
 life cycle. Consequently, Jankowski (1967b) 
 
 placed this animal in a suborder by itself. Non- 
 ciliated adult stages of Conidiophrys are known 
 to occur on Corovhium, Gammarus, and Ido- 
 thea. 
 
 Family FOETTINGERIIDAE Chatton, 1911. 
 At least 14 genera represented in marine habi- 
 tats. Ciliates found in body fluids of copepods, 
 amphipods, crabs, hermit crabs, and sea anem- 
 ones. Life cycle complex, polymorphic, with 
 encysted stages. See Chatton and Lwoff (1935) . 
 
 Family OPALINOPSIDAE Hartog, 1906. 
 Three genera: Chromidina, Cyrtocaryum, and 
 Opalinopsis; endocommensal in cephalopods, 
 coelenterates, ctenophores, and polychaetes. 
 Chromidina occurs in cephalopods, Sepia, Loligo, 
 and Illex. See Chatton and Lwoff (1935). Cyr- 
 tocaryum occurs in the lateral caeca of the di- 
 gestive tube of the polychaete Halosydna. The 
 life cycle includes a tomite with a group of five 
 to six fine, short ciliary rows once division allows 
 size to fall below 40 pxa. Tomites are liberated 
 into free seawater; they are asymmetrical, with 
 a long caudal cilium. See Faure-Fremiet and 
 Mugard (1949). 
 
 Order Astomatida 
 
 Family ANOPLOPHRYIDAE Cepede, 1910. 
 At least 12 genera, of which five have marine 
 representatives. Members of the genera Ano- 
 plophryopsis, Herpetophrya, Herpinella, and 
 Rhizocaryum occur in polychaetes including 
 Cirratulus and Polydora. Members of the genus 
 Perseia occur in Phascolosoma. See Cepede 
 (1910), Kahl (1933), and Puytorac (1954). 
 
 Family HAPTOPHRYIDAE Cepede, 1923. 
 About eight genera, of which Lachmannella and 
 Steinella have marine representatives in turbel- 
 larians. See Sikora (1963) and Corliss, Puy- 
 torac, and Lorn (1965). 
 
 Family HOPLITOPHRYIDAE Cheissin, 1830. 
 Includes over 30 genera (including those some- 
 times placed in the family INTOSHELLINIDAE 
 Cepede, 1910) , of which at least five have marine 
 species: Buetschliella, Durchoniella, Hovassiel- 
 la, Helella, and Spirobuetschliella. Marine mem- 
 bers of the family are endocommensal in the gut 
 of polychaetes, Ophelia, Cirratulus, Polydora, 
 and Potamoceros. 
 
 46 
 
Order Hymenostomatida 
 
 Family CINETOCHILIDAE Perty 1852. Two 
 genera, of which one, Cinetochilam, with one ma- 
 rine species. Free-living. See Gelei (1940). 
 
 Family COHNILEMBIDAE Kahl, 1933. In- 
 cludes only the genus Cohnilembus. Free-living. 
 See Borror (1961, 1963). 
 
 Family ENTORHIPIDIIDAE Madsen, 1931. 
 Contains at least the genus Entorhipidium, oc- 
 curring in the gut of sea urchins. See Berger 
 (1961b). 
 
 Family FRONTONIIDAE Kahl, 1926. Eight 
 genera; two with marine species: Frontonia 
 and Frontoniella. Widespread, free-living. See 
 Roque (1961). 
 
 Family LOXOCEPHALIDAE Jankowski, 
 1964. At least two genera with marine species, 
 Dexiotricha and Loxocephalus. Free-living. 
 Cardiostomatella may belong here also. See 
 Jankowski (1964a) and Faure-Fremiet (1968). 
 
 Family OPHRYOGLENIDAE Kent, 1880. 
 Possibly four genera; only one, Ophryoglena, 
 in the marine habitat. Free-living in brackish 
 waters; some species histophagic. See Roque 
 (1961). 
 
 Family PARAMECIIDAE Dujardin, 1841. 
 One genus, Paramecium, with many species, of 
 which P. woodruff i and P. calkinsi occur in 
 brackish water. Free-living. See Wichterman 
 (1953). 
 
 Family PHILASTERIDAE Kahl, 1931. The 
 boundaries between this family and the URONE- 
 MATIDAE are unclear. The members of the 
 genera Philaster, Philasterides, and Porpostoma 
 form a natural group within the PHILASTERI- 
 DAE. Anophrys salmacida may belong here also 
 (Small, 1967). Additionally, members of the 
 genera Glauconema, Helicostoma, Paranophrys, 
 Parauronema, and Potomacus, most of which 
 have been placed in the URONEMATIDAE, may 
 be members of the PHILASTERIDAE. Free- 
 living and endocommensal in sea urchins, some- 
 times found in association with hydroids and 
 anemones, some histophagic. 
 
 Family PLEURONEMATIDAE Kent, 1880. 
 Genera with marine members are Cristigera, 
 
 Cyclidium, Histiobalantium, Pleurocoptes, Pleu- 
 ronema, and Schizocalyptera. The genus Cycli- 
 dium is sometimes placed in the family CYCLI- 
 DIIDAE Ehrenberg, 1838. The genus Penicu- 
 listoma, sometimes considered in the order 
 Thigmotrichida, may also belong in this family. 
 Free-living and commensal on mussels and sea 
 urchins. See Faure-Fremiet (1961a), Dragesco 
 (1968), and Borror (1963). 
 
 Family PSEUDOCOHNILEMBIDAE Evans 
 and Thompson, 1964. On the basis of similari- 
 ties in morphogenesis, Pseudocohnilembus spp. 
 and Anophrys sarcophaga belong here (Evans 
 and Thompson, 1964; Small, 1967) . On the basis 
 of interphase morphology, Paralembus and 
 Cryptochilidium cuenoti may also belong here. 
 Free-living and endocommensal in sea urchins 
 (C. cuenoti occurs in the esophagus of the sipun- 
 culid Phascolosoma) . See Berger (1961a, 1965) 
 and Mugard (1949). 
 
 Family TETRAHYMENIDAE Corliss, 1952. 
 Includes at least six genera; two with marine 
 species: Paratetrahymena and Platynematum. 
 Recently Corliss (1961) split several genera 
 from this family and placed them in the family 
 GLAUCOMIDAE. See Borror (1962), Corliss 
 (1961), Czapik (1968), and Thompson (1963). 
 
 Family URONEMATIDAE Thompson, 1964. 
 The boundaries between this family and the 
 PHILASTERIDAE are unclear, but members 
 of the genera Uronema and Uropedalium form 
 a well-defined group within this family. In ad- 
 dition, members of the genera Glauconema, Mi- 
 amiensis, Parauronema, and Potomacus have 
 been placed here. Free-living, sometimes found 
 in association with hydroids and anemones. See 
 Thompson (1964a, b, 1966) and Thompson and 
 Berger (1965). 
 
 In addition to the hymenostomes assigned to 
 the families above, members of several genera 
 (Biggaria, Cryptochilum, Entodiscus, and Mad- 
 senia, all endocommensals of sea urchins) are 
 considered here as hymenostomes with no as- 
 signed familial status (Berger, 1961c). 
 
 Recently, Small erected a new order Scutico- 
 ciliatida for certain hymenostome and thigmot- 
 rich families apparently related by particular 
 features of stomatogenesis (Small, 1967). 
 
 47 
 
Order Thigmotrichida 
 
 Family ANCISTROCOMIDAE Chatton and 
 Lwoff, 1939. About 23 genera; 12 with marine 
 representatives: Colligocineta, Crebricoma, 
 Goniocoma, Heterocineta, Holoeoma, Hypocoma- 
 galma, Hypocomella, Hypocomides, Hypocomina, 
 Insignicoma, Isoeomides, and Raabella. Typi- 
 cally ectocommensal on gills of bivalves. See 
 Fenchel (1965a), Kozloff (1965 and earlier pa- 
 pers), and Raabe (1970b). 
 
 Family CONCHOPHTHIRIDAE Kahl, 1934. 
 At least four genera; one, Cochliophilus, with a 
 marine representative endocommensal in Phytia, 
 a tidal marsh pulmonate snail of the west coast. 
 See Kahl (1934a) and Raabe (1963b). 
 
 Family CRATERISTOMATIDAE Jankowski, 
 
 1967. Contains Crateristoma kindi, a carnivore 
 on prostome ciliates found in association with 
 barnacles. See Jankowski (1968). 
 
 Family HEMISPEIRIDAE Konig, 1894. Con- 
 tains at least 17 genera, of which nine have ma- 
 rine members: Ancistrospira, Ancistrum, Bo- 
 veria, Hemispeira, Isselina, Orchitophrya, Pla- 
 giospira, Proboveria, and Protophyra. Most are 
 ectocommensal on gills of bivalves; one is en- 
 dozoic in echinoderms. Nucleocorbula adherens 
 Santhakumari and Balakrishnan Nair, 1970, for 
 which the authors erected the family NUCLEO- 
 CORBULIDAE, keys out here. See Lorn, Corliss, 
 and Noirot-Timothee (1968), Fenchel (1965a), 
 and Raabe (1970a). 
 
 Family HYPOCOMIDAE Biitschli, 1889. 
 Three genera ; two, Heterocoma and Parahypo- 
 coma, with marine representatives; ectocom- 
 mensal on solitary and colonial peritrichs, and in 
 branchial cavity of ascidians. See Chatton and 
 Lwoff (1949). 
 
 Family HYSTEROCINETIDAE Diesing, 
 1866. Members of this family (at least 10 gen- 
 era) are typically endocommensal in terrestrial 
 and freshwater oligochaetes. See Raabe (1949) 
 and Kozloff (1960). 
 
 Family SPHENOPHRYIDAE Chatton and 
 Lwoff, 1921. Four genera with marine repre- 
 sentatives: Gargarius, Lwoff ia, Pelecophrya, 
 and Sphenophrya; ectocommensal on gills of bi- 
 valves. Mature form with no cilia (except in 
 genus Lwoffia), but budded larval forms with 
 
 several rows of cilia. See Kozloff (1955), Do- 
 brzanska (1961), and Raabe (1970b). 
 
 Family THIGMOPHRYIDAE Chatton and 
 Lwoff, 1923. Two genera, Conchophyllum and 
 Thigmophyra, with marine representatives; 
 ectocommensal on gills of bivalves. Undulating 
 membrane difficult to observe in life. See Fen- 
 chel (1965a). 
 
 The genus Peniculistoma, sometimes consid- 
 ered to be a member of this order, is included 
 here in the family PLEURONEMATIDAE (or- 
 der Hymenostomatida) . Recently, Small erected 
 a new order Scuticociliatida for certain hyme- 
 nostome and thigmotrich families apparently 
 related by particular features of stomatogene- 
 sis. 
 
 Order Peritrichida 
 
 Family EPISTYLIDIDAE Kahl, 1933. Four- 
 teen genera, of which at least three have marine 
 representatives: Epistylis, Opisthostyla, and 
 Rhabdostyla. They occur attached to algae or 
 the outer surfaces of copepods and barnacles, 
 (Epistylis), on stalks of ascidians (Opistho- 
 styla) , and on polychaetes, midge larvae, and 
 sea cucumbers (Rhabdostyla). Individuals soli- 
 tary and colonial. See Kahl (1935); Lorn 
 (1964). 
 
 Family LAGENOPHRYIDAE Biitschli, 1889. 
 At least two genera; one, Lagenophrys, with ma- 
 rine representatives, ectocommensal on crusta- 
 ceans. See Couch (1967). 
 
 Family SCYPHIDIIDAE Kahl, 1933. Ten 
 genera, of which three have marine representa- 
 tives. Members of the genus Ellobiophrya occur 
 on Donax; Paravorticella occurs on polychaetes; 
 Scyphidia occurs on snails and polychaetes. See 
 Kahl (1935). 
 
 Family URCEOLARIIDAE Dujardin, 1941. 
 Subfamily Urceolarinae contains the genera 
 Urceolaria, Leiotricha, Trichodinopsis, and Poly- 
 cycla that occur as ectocommensals on marine 
 invertebrates, on gills and respiratory surfaces 
 of marine invertebrates, in the gut of snails, and 
 in the gut of sea cucumbers, respectively. The 
 subfamily Trichodininae contains eight genera, 
 of which seven have marine representatives. 
 Members of the genera DipartieUa, Trichodinel- 
 la, and Tripartiella occur attached to fish gills; 
 
 48 
 
members of the genera Paravauchomia, Poljans- 
 kina, Tripartiella, and Vauchomia occur in the 
 urinary bladder of fish; and members of the 
 genus Trichodina are found in association with 
 a wide variety of hosts. See review by Raabe 
 (1963a). 
 
 Family VAGINICOLIDAE de Fromentel, 
 1874. Ten genera, of which six have marine rep- 
 resentatives. Members of the genera Caulicola 
 and Thuricola are free-living. Cothurnia, 
 Platycola, Pyxicola, and Vaginicola occur at- 
 tached to red or green algae, as well as the 
 outer surfaces of various invertebrates, includ- 
 ing crustaceans, polychaetes, hydroids, and 
 snails. See Kahl (1935). 
 
 Family VORTICELLIDAE Ehrenberg, 1838. 
 Eight genera; three with marine representa- 
 tives. Members of the genera Carchesium and 
 Vorticella are free-living. Members of the genus 
 Zoothamnium are free-living and ectocommensal 
 on crustaceans and hydroids. See Kahl (1935). 
 Myonemes in colonial forms usually continuous, 
 except in the genus Carchesium. 
 
 Family METOPIDAE Kahl, 1932. Six gen- 
 era; one, Metopus, with marine species. Free- 
 living in areas of low oxygen concentration. 
 See Jankowski (1964b). 
 
 Family PERITROMIDAE Stein, 1867. Two 
 genera; one, Peritromus, with marine species. 
 Free-living. See Borror (1963) and Tuffrau 
 (1967b). 
 
 Family SPIROSTOMATIDAE Stein, 1867. 
 At least eight genera, of which four have ma- 
 rine representatives: Blepharisma, Gruberia, 
 Parablepharisma, and Anigsteinia. Large, 
 elongate, free-living animals. See Isquith 
 (1968). The genus Protocruzia, usually placed 
 in this family, is sufficiently aberrant to warrant 
 transfer to a different family. They are free- 
 living, benthic, small, and fragile. See Ammer- 
 mann (1968). 
 
 Family STENTORIDAE Carus, 1863. Four 
 genera; three with marine species: Climacos- 
 tomum, Fabrea, and Stentor. They occur pri- 
 marily in areas of low salinity. See Kahl ( 1932) . 
 
 Order Heterotrichida 
 
 Family CAENOMORPHIDAE Poche, 1913. 
 Three genera; one, Caenomorpha, with marine 
 representatives. Occur in regions of low oxygen 
 concentration. Free-living. See Jankowski 
 (1964b). 
 
 Family CHATTONIDIIDAE Villeneuve- 
 Brachon, 1940. One genus, Chattonidium, free- 
 living. See Villeneuve-Brachon (1940). 
 
 Family CONDYLOSTOMATIDAE Kahl in 
 Doflein and Reichenow, 1929. One genus, Con- 
 dylostoma, widespread and free-living. See 
 Borror (1963). 
 
 Family FOLLICULINIDAE Dons, 1914. 
 Members of this family are grouped into at least 
 30 genera (listed by Corliss, 1961), all marine. 
 They are ectocommensal on various inverte- 
 brates. 
 
 Family LICNOPHORIDAE Biitschli, 1887. 
 One genus, Licnophora, ectocommensal on snails, 
 mussels, sea cucumbers, marine worms, and 
 other invertebrates. See Villeneuve-Brachon 
 (1940). 
 
 Order Odontostomatida 
 
 Family EPALXELLIDAE Corliss, 1960. 
 Three genera; one, Epalxella, with marine spe- 
 cies. Free-living, in areas of low oxygen con- 
 centration. See Jankowski (1964b). 
 
 Family MYLESTOMATIDAE Kahl, 1932. 
 Two genera; one, Mylestoma, with marine spe- 
 cies. Free-living, in areas of low oxygen con- 
 centration. See Jankowski (1964b). 
 
 Order Oligotrichida 
 
 Family HALTERIIDAE Claparede and Lach- 
 mann, 1858. Seven genera; four with marine 
 representatives: Cephalotrichidium, Meta- 
 strombidium, Strombidium, and Tontonia. 
 These are typically pelagic. Members of the 
 genus Strombidium are widespread, ubiquitous, 
 and in some cases benthic. See Kahl (1932). 
 Recently members of this family with a girdle 
 of trichites and polygonal cortical platelets, in- 
 cluding Strombidium and Tontonia, were placed 
 in the family STROMBIDIIDAE. See Faure- 
 Fremiet (1969). 
 
 49 
 
Family STROBILIDIIDAE Kahl in Doflein 
 and Reichenow, 1929. Six genera, of which 
 three have marine species: Ciliospira, Lohman- 
 niella. and Strobilidium. Typically pelagic, 
 planktonic. See Kahl (1932). 
 
 Order Tintinnida 
 
 Families in this order are differentiated on 
 the basis of the morphology of the lorica. See 
 Kofoid and Campbell (1929), Schwarz (1964), 
 Loeblich and Tappan (1968), and Tappan and 
 Loeblich (1968). Members of this order gen- 
 erally are pelagic, planktonic. Many genera are 
 known only as fossils. 
 
 Family CODONELLIDAE Kent, 1882. At 
 least five marine genera: Codonaria, Codonella, 
 Codonopsis, Rhizodomus, and Tintinnopsis. 
 
 Family CODONELLOPSIDAE Kofoid and 
 Campbell, 1929. Eight genera; at least three 
 with marine species: Codonellopsis, Laackman- 
 niella, and Stenosemella. 
 
 Family COXLIELLIDAE Kofoid and Camp- 
 bell, 1929. Eight genera; seven with recent ma- 
 rine representatives: Climacocylis, Coxliella, 
 Helicostomella, Metacylis, Stylicauda, Favel- 
 loides, and Poroecus. 
 
 Family CYTTAROCYLIDIDAE Kofoid and 
 Campbell, 1929. Only Cyttarocylis. 
 
 Family DICTYOCYSTIDAE Kent, 1881. 
 Three genera with marine species: Dictyocysta, 
 Luminella, and Wangiella. 
 
 Family EPIPLOCYLIDIDAE Kofoid and 
 Campbell, 1929. Three genera with marine rep- 
 resentatives: Epicancella, Epiplocylis, and 
 Epiplocyloides. 
 
 Family PETALOTRICHIDAE Kofoid and 
 Campbell, 1929. Six genera with marine rep- 
 resentatives: Acanthostomella, Ascampbelliella, 
 Luxiella, Niemarshallia, Petalotricha, and Wai- 
 lesia. 
 
 Family PTYCHOCYLIDIDAE Kofoid and 
 Campbell, 1929. Four genera with marine rep- 
 resentatives: Cymatocylis, Favella, Protocymw- 
 tocylis, and Ptychocylis. 
 
 Family RHABDONELLIDAE Kofoid and 
 Campbell, 1929. Four marine genera: Epirhab- 
 donella, Protorhabdonella, Rhabdonella, and 
 Rhabdonellopsis. 
 
 Family TINTINNIDAE Claparede and Lach- 
 mann, 1858. About 23 genera with marine rep- 
 resentatives (listed by Tappan and Loeblich, 
 1968). 
 
 Family TINTINNIDIIDAE Kofoid and 
 Campbell, 1929. Two marine genera: Leprotin- 
 tinnus and Tintinnidium. 
 
 Family UNDELLIDAE Kofoid and Campbell, 
 1929 . Seven genera with marine species: 
 Amplectella, Amplectellopsis, Cricimdella, Mi- 
 crundella, Proplectella, Undella, and Undellopsis. 
 
 Family XYSTONELLIDAE Kofoid and 
 Campbell, 1929. Four genera with marine spe- 
 cies: Parafavella, Parundella, Xystonella, and 
 Xystonellopsis. 
 
 Order Hypotrichida 
 
 Family EUPLOTIDAE Ehrenberg, 1838. 
 Eleven genera, each with marine representa- 
 tives: Aspidisca, Certesia, Diophrys, Discoceph- 
 alus, Euplotaspis, Euplotes, Euplotidium, Gas- 
 trocirrhus, Paraeuplotes, Swedmarkia, and 
 Uronychia. Most are benthic and free-living. 
 See Borror (1963, 1965a, 1968b, and 1972) and 
 Deroux and Tuffrau (1965). 
 
 Family HOLOSTICHIDAE Faure-Fremiet, 
 1961. At least six genera with marine species: 
 Holosticha, Keronopsis, Paraholosticha, Trich- 
 otaxis, Uroleptopsis, and Uroleptus. Free-living. 
 See Borror (1972a). 
 
 Family OXYTRICHIDAE Ehrenberg, 1838. 
 Marine genera include: Ancystropodium, Gas- 
 trostyla, Histriculus, Laurentia, Onychodromus, 
 Oxytricha, Parastylonychia, Stylonychia, Tach- 
 ysoma, and Trachelostyla. Free-living and ben- 
 thic. See Borror (1972a) and Faure-Fremiet 
 (1961b). 
 
 Family PSILOTRICHIDAE Butschli, 1889. 
 Three genera: Caryotricha, Kiitricha, and 
 Psilotricha, all marine. , Free-living. See Kahl 
 (1932). 
 
 50 
 
Family SPIROFILIDAE Gelei, 1929. Seven 
 genera; three with marine species: Sticho- 
 tricha, Strongylidium, and Urostrongylum. 
 Free-living and benthic. 
 
 Family UROSTYLIDAE Butschli, 1889. At 
 least seven genera with marine representatives: 
 Amphisiella, Balladyna, Banyulsella, Epiclintes, 
 Lacazea, Paraurostyla, and Urostyla. Members 
 free-living and benthic. See Borror (1972a). 
 
 SELECTED BIBLIOGRAPHY 
 
 AGAMALIEV, F. G. 
 
 1967. Faune des cilies mesopsammiques 
 de la cote ouest de la Mer Caspienne. 
 Cahiers Biol. Mar. 8: 359-402. [Morphol- 
 ogy of many species; new species de- 
 scribed.] 
 
 AMMERMANN, VON D. 
 
 1968. Die Kernverhaltnisse des Ciliaten 
 Protocrucia depressa n. sp. Arch. Pro- 
 tistenkd. 110:434-438. [Includes careful 
 description of morphology.] 
 
 BERGER, J. 
 
 1961a. The comparative buccal morphol- 
 ogy of certain hymenostome ciliates en- 
 tocommensal in echinoids. Proc. First 
 Int. Cong. Protozool. p. 86-88. [Describes 
 ciliature of buccal cavity in Cryptochilum 
 and Biggaria spp.] 
 
 1961b. Stomatogenic evidence for the hy- 
 menostome affinities of the genus Ento- 
 rhipidium Lynch. J. Protozool. 8 
 (Suppl.): 11. [Morphology of this ciliate 
 from sea urchins.] 
 
 1961c. Systematic reassignment, based on 
 buccal morphology of two species of cili- 
 ates entocommensal in echinoids. J. Pro- 
 tozool. 8 (Suppl.) : 11. [Transfers Ento- 
 discus and Madsenia from the Trichos- 
 tomatida to the Hymenostomatida.] 
 
 1961d. Morphology and systematic posi- 
 tion of Schizocaryum dogieli, a ciliate en- 
 tocommensal in strongylocentrotid echi- 
 noids (Ciliata: Trichostomatida). J. 
 Protozool. 8:363-369. [Places it in the 
 trichostome family PLAGIOPYLIDAE.] 
 
 1965. Infraciliary morphology of a little- 
 known echinophilous hymenostome ciliate 
 from the Indo-West Pacific. Proc. Sec- 
 
 ond Int. Conf. Protozool. No. 255. [Dis- 
 cusses morphology of Cryptochilidium 
 polynucleata.1 
 BORROR, A. C. 
 
 1961. Feeding apparatus of the ciliate 
 Cohnilembus verminus (Muller). Bull. 
 Assoc. Southeastern Biol. 8:21. [De- 
 scription of buccal and somatic ciliature.] 
 
 1962. Ciliate Protozoa of the Gulf of Mex- 
 ico. Bull. Mar. Sci. Gulf Caribb. 12: 
 333-349. [List of recorded species and 
 suggested synonymies.] 
 
 1963. Morphology and ecology of the ben- 
 thic ciliated Protozoa of Alligator Harbor. 
 Florida. Arch. Protistenkd. 106: 465-534. 
 [Morphology of many species; four new 
 species described.] 
 
 1965a. Morphological comparison of Di- 
 ophrys scutum (Dujardin, 1841) and 
 Diophrys peloetes n. sp. (Hypotrichida, 
 Ciliophora). J. Protozool. 12:60-66. 
 [Intraspecific variations tabulated.] 
 
 1965b. New and little-known tidal marsh 
 ciliates. Trans. Am. Microsc. Soc. 84: 
 550-565. 
 
 1968a. Nigrosin-HgCl2-Formalin; a stain- 
 fixative for ciliates (Protozoa, Cilioph- 
 ora). Stain Technol. 43:293-295. [Form- 
 ula for a method that stains cilia and 
 cortical sculpturing black against a white 
 background.] 
 
 1968b. Systematics of Euplotes (Cilioph- 
 ora, Hypotrichida) ; toward union of the 
 old and the new. J. Protozool. 15:802- 
 808. [Discussion of morphology of four 
 species, showing importance of cortical 
 features (viewed prior to dependence up- 
 on silver line system) in identification of 
 species.] 
 
 1969. Application of the stain-fixative Ni- 
 grosin-HgCl2-Formalin to fragile or con- 
 tractile ciliates. Trans. Am. Microsc. Soc. 
 88: 454-458. [Modification of the fixation 
 and relaxation methods applied to above 
 method.] 
 
 1972a. Revision of the order Hypotrichida 
 (Ciliophora, Protozoa). J. Protozool. 
 19:1-23. [Lists 310 species distributed 
 among 51 genera and 6 families. All 
 families and genera are diagnosed and 
 illustrated and synonymies are listed for 
 each species.] 
 
 51 
 
BORROR— Cont. 
 
 1972b. Ciliates of New Hampshire tidal 
 
 marshes. Acta Protozool. 10:29-71. 
 
 [About 100 species listed; morphology 
 
 and autecology of over 30 spp. discussed; 
 
 4 new species.] 
 CANELLA, M. F. 
 
 1960. Contributi alia conscenza dei Ciliati. 
 III. Contribute ad una revisione dei ge- 
 neri Amphileptus, Hemiophrys e Liono- 
 tus (Ciliata. Holotricha, Gymnostomata) . 
 Ann. Univ. Ferrara, Sez. 3, Biol. Anim. 
 2:47-95. [Systematics of the Amphilep- 
 tidae; a lengthy treatise.] 
 
 CEPEDE, C. 
 
 1910. Recherches sur les infusoires astom- 
 es. Anatomie, biologie, ethologie para- 
 sitaire, systematique. Arch. Zool. Exp. 
 Gen. (Ser. 5) 3:341-609. [An early but 
 important work describing many species 
 of the order Astomatida.] 
 
 CHATTON, E., and A. LWOFF. 
 
 1935. Les cilies apostomes. Premiere 
 partie. Apergu historique et general. 
 Etude monographique des genres et des 
 especes. Arch. Zool. Exp. Gen. 77:1-453. 
 [A classic monograph on the Apostoma- 
 tida.] 
 
 1936. Les Pilisuctoridae Chatton et Lwoff. 
 Cilies parasites des poils secheteurs des 
 Crustaces Edriophthalmes. Polarite, ori- 
 entation, et desmodexie chez les Infu- 
 soires. Bull. Biol. Fr. Belg. 70:86-144. 
 [A description of the life cycle of the non- 
 ciliated sessile Conidiophrys.~] 
 
 1949. Recherches sur les cilies thigmo- 
 triches. Arch. Zool. Exp. Gen. 86:169- 
 253, 393-495. [An important monograph 
 on the members of the order Thigmotrich- 
 ida.] 
 CORLISS, J. O. 
 
 1953. Silver impregnation of ciliated pro- 
 tozoa by the Chatton-Lwoff technic. 
 Stain Technol. 28:97-100. [A redescrip- 
 tion of the classical French silver meth- 
 od.] 
 
 1959. An illustrated key to the higher 
 groups of the ciliated Protozoa, with def- 
 inition of terms. J. Protozool. 6:265-281. 
 
 1961. The ciliated Protozoa; characteriza- 
 tion, classification, and guide to the liter- 
 ature. Pergamon Press, Inc., N.Y. 310 p. 
 
 [Contains important coverage of history, 
 systematics, and bibliographic sources 
 for the entire class.] 
 
 CORLISS, J. O., P. de PUYTORAC, and J. LOM. 
 1965. Resolution of persistant taxonomic 
 and nonmenclatural problems involving 
 ciliate Protozoa assignable to the astome 
 family Haptophryidae Cepede, 1923. J. 
 Protozool. 12:265-273. [Morphology of 
 Steinella and Lachma?iella discussed.] 
 
 COUCH, J. A. 
 
 1967. A new species of Lagenophrys (Cil- 
 iatea: Peritrichida: Lagenophryidae) 
 from a marine crab, Callinectes sapidus. 
 Trans. Am. Microsc. Soc. 86:204-211. 
 [Morphology of cell and lorica discussed.] 
 
 CZAPIK, A. 
 
 1968. La famille Tetrahymenidae et son 
 importance dans la systematique et revo- 
 lution des cilies. Acta Protozool. 5:315- 
 358. [A recent review of the family 
 Tetrahymenidae.] 
 
 DEROUX, G. 
 
 1970. La serie "Chlamydonellienne" chez 
 les Chlamydodontidae (Holotriches, 
 CYRTOPHORINA Faure-Fremiet) . Pro- 
 tistology 6:155-182. 
 
 DEROUX, G., and M. TUFFRAU. 
 
 1965. Aspidisca orthopogon n. sp. Revi- 
 sion des certains mecanismes de la mor- 
 phogenese a l'aide d'une modification de 
 la technique au Protargol. Cahiers Biol. 
 Mar. 6: 293-310. [Important descriptions 
 of morphogenesis in this hypotrich.] 
 
 DOBRZANSKA, J. 
 
 1961. Further study on Sphenophrya 
 dreissenae Dobrzanska 1958 (Ciliata, 
 Thigmotricha). Acta Parasitol. Pol. 9: 
 117-140. [Comparative morphology of 
 many thigmotrichs, including ciliated 
 tomites.] 
 
 DOBRZANSKA-KACZANOWSKA, J. 
 
 1963. Comparaison de la morphogenese des 
 Cilies: ChUodonella uncinata (Ehrbg.), 
 Allosphaeriiim paraconvexa sp. n. et Heli- 
 ochona scheuteri (Stein). Acta Protozool. 
 1 : 353-394. [On comparative morphology 
 of the gymnostones and the chonotrichs.] 
 
 DRAGESCO, J. 
 
 1960. Les cilies mesopsammiques littoraux 
 (Systematique, morphologie, ecologie). 
 Trav. Stn. Biol. Roscoff 12 (Nov. Ser.): 
 
 52 
 
1-356. [A lengthy treatise on sand 
 dwelling ciliates, with descriptions of 
 many new species.] 
 
 1963a. Revision du genre Dileptus, Du- 
 jardin 1871 (Ciliata: Holotricha) Sys- 
 tematique, Cytologie, Biologie. Bull. Biol. 
 Fr. Belg. 97:103-145. [A recent review 
 of the genus.] 
 
 1963b. Complements a la connaissance des 
 Cilies mesopsammiques de Roscoff. 1. 
 Holotriches. Cahiers Biol. Mar. 4:91-119. 
 [Morphology: descriptions of new spe- 
 cies.] 
 
 1966. Observations sur quelques cilies 
 libres. Arch. Protistenkd. 109:155-206. 
 [Morphology: descriptions of new spe- 
 cies.] 
 
 1967. Armature fibrilaire interne chez 
 Hartmannula acrobates Entz (Cilie, hol- 
 otriche gymnostome). Protistologica 3: 
 61-65. [Studied this gymnostome with 
 protargol method.] 
 
 1968. Les genres Pleuronema Dujardin, 
 Schizocalyptera nov. gen. et Histobalant- 
 ium Stokes (Cilies holotriches hymeno- 
 stomes). Protistologica 4:85-106. [Dis- 
 cussion of intraspecific variation, and 
 descriptions of new species.] 
 
 EVANS, F. R., and J. C. THOMPSON, JR. 
 
 1964. Pseudocohnilembidae n. fam., a 
 hymen stome ciliate family containing one 
 genus, Pseudocohnilembus n.g., with 
 three new species. J. Protozool. 11:344- 
 352. 
 FAURE-FREMIET, E. 
 
 1955. La position systematique due genre 
 Balantidium. J. Protozool. 2:54-58. 
 [Considers it a trichostome.] 
 
 1961a. Pleurocoptes hydractinae (Ciliata 
 Pleuronematina) et son adaptation struc- 
 tural a la vie epizoique. Cahiers Biol. 
 Mar. 2:447-52. [Shows its Pleuronema- 
 like buccal ciliature.] 
 
 1961b. Protozoologie. — Remarques sur la 
 morphologie comparee et la systematique 
 des Ciliata Hypotrichida, C. R. Acad. 
 Sci., Paris 252: 3515-3519. [Brief review 
 of the order, with erection of new fam- 
 ilies.] 
 
 1961c. Protozoologie. — Les Cilies Cyrto- 
 phorina et leur diversification morpholog- 
 ique. C. R. Acad. Sci., Paris 252:3912- 
 
 3916. [Distinguishes between the En- 
 chelyidae and the Prorodontidae and dis- 
 cusses other gymnostomes.] 
 
 1962a. Le genre Paranassula Kahl (Cili- 
 ata, Cyrtophorina) . Cahiers Biol. Mar. 
 3:61-77. [Illustrates its vestibular cili- 
 ature.] 
 
 1962b. Le genre Trimyema Lackey et les 
 caracteristiques structurales des cilies 
 Trichostomatida. J. Protozool. 9:146- 
 149. [Describes somatic and vestibular 
 ciliature.] 
 
 1963. Conchostoma longissimum n.g. n. sp., 
 cilie trichostome psammobie. Cahiers 
 Biol. Mar. 4:193-199. [Describes this 
 extremely elongate form from sand.] 
 
 1968. Les genres Dexiotricha Stokes et 
 Loxocephalus Eberhard dans leurs rela- 
 tions auxomorphiques. Protistologica 4: 
 115-125. 
 
 1969. Remarques sur la systematique des 
 cilies Oligotrichida. Protistologica 5: 
 345-352. [A review of the order ; a new 
 family erected.] 
 
 FAURE-FREMIET, E., and H. MUGARD. 
 1949. Zoologie. — Un infusoire apostome 
 parasite d'un polychete: Cyrtocaryum 
 halosydnae, n. gen., n. sp. C. R. Acad. 
 Sci., Paris 228: 1753-1755. [Morphology, 
 life cycle, and systematics.] 
 FAURE-FREMIET, E., C. ROUILLER, and 
 M. GAUCHERY. 
 
 1956. Structure et origine du pedoncule 
 chez Chilodochona. J. Protozool. 3:188- 
 193, [Morphology of this member of the 
 Chonotrichida.] 
 FENCHEL, T. 
 
 1964. Gullmarella faurei n. g., n. sp., a hol- 
 otrichous ciliate from the intestine of 
 Lamellibranchs. Cahiers Biol. Mar. 5: 
 311-318. [Morphology of this peculiar 
 trichostome.] 
 
 1965a. Ciliates from Scandinavian mol- 
 luscs. Ophelia 2:71-174. [An important 
 treatise containing morphological details 
 on the many thigmotrichs, and descrip- 
 tions of new species.] 
 
 1965b. On the ciliate fauna associated with 
 the marine species of the amphipod genus 
 Gammariis J. G. Fabricius. Ophelia 2: 
 281-303. [Morphology; Silenella de- 
 scribed.] 
 
 53 
 
GELEI. G. 
 
 1940. Cinetoehilum und sein Neuronemen- 
 system. Arch. Protistenkd. 94:57-79. 
 [Shows the buccal apparatus to be Tetra- 
 hymena-Wke.] 
 
 HAIRSTON, N. G., and S. L. KELLERMANN. 
 1964. Paramecium ecology; electromigra- 
 tion for field samples and observations 
 on density. Ecology 45:373-376. [De- 
 scribes a method for isolating paramecia 
 from bottom sediments based on their lo- 
 comotion in a weak electrical current.] 
 
 ISQUITH, I. R. 
 
 1968. Anigsteinia n. g., a member of the 
 family Spirostomatidae. J. Protozool. 
 15(Suppl.) :26. [Splits this genus from 
 Blepharisma."] 
 JANKOWSKI, A. W. 
 
 1964a. Morphology and evolution of Cili- 
 ophora. IV. Sapropelebionts of the fam- 
 ily Loxocephalidae fam. nova, their tax- 
 onomy and evolutionary history. Acta 
 Protozool. 2:33-58. [Discusses morphol- 
 ogy of Dexiotricha and Loxocephalus.] 
 
 1964b. Morphology and evolution of cili- 
 ophora. III. Diagnoses and phylogenesis 
 of 53 Sapropelebionts, mainly of the order 
 Heterotrichida. Arch. Protistenkd. 107: 
 185-294. [An important revision c-f the 
 Odontostomatida and the Metopus-hke 
 heterotrichs.] 
 
 1967a. Ciliates from the mantle cavity of 
 Balanus in Barents Sea. Parasitology 
 1:82-93. [Morphology and systematics, 
 including descriptions of several new spe- 
 cies.] 
 
 1967b. A new system of the Ciliate Proto- 
 zoa (Ciliophora). Acad. Sci. USSR. 
 Trav. Zool. Inst. 43:1-52. [A lengthy 
 important treatise defending a reclassi- 
 fication of the ciliates, emphazing a new 
 realignment particularly of many holo- 
 trichs.] 
 
 1968. Morphology and phylogeny of Cra- 
 teristoma (Ciliophora: Rhynchodida). 
 Hydrobiol. J. 4:33-41. [Morphological 
 details added to previous description.] 
 
 1970. T?*op ho gemma poljansky gen. et 
 sp. n. (Ciliophora, Suctoria) from the 
 shrimp's antennae. Hydrobiol. J. 6: 
 454-457. [Morphology.] 
 
 KAHL, A. 
 
 1930-1935. Urtiere oder Protozoa. I: 
 Wimpertiere oder Ciliata (Infusoria). 
 In F. Dahl, Die Tierwelt Deutschlands, 
 18(1930), 21(1931), 25(1932), 30 
 (1935), p. 1-886. [Most important 
 treatise on the morphology and system- 
 atics of the ciliates. Attempts to deal 
 with every species, except for marine 
 tintinnids and endozoic ciliates.] 
 
 1933. Ciliata libera et ectocommensalia. 
 In G. Grimpe and E. Wagler (Begrun- 
 det) , Die Tierwelt der Nord- und Ostsee 
 23:29-146. [This monograph, published 
 in another series, overlaps significantly 
 with the above, but it contains much im- 
 portant morphological and systematics 
 information.] 
 
 1934a. Ciliata entocommensalia et para- 
 sitica. In G. Grimpe and E. Wagler 
 (Begrundet) , Die Tierwelt der Nord- und 
 Ostsee 26:147-183. [Complements the 
 monograph cited above, with coverage on 
 parasitic forms.] 
 
 1934b. Suctoria. In G. Grimpe and E. 
 Wagler (Begrundet), Die Tierwelt der 
 Nord- und Ostsee 26: 184-226. [An im- 
 portant monograph on the order Suctor- 
 ida.] 
 KOFOID, C. A., and A. S. CAMPBELL. 
 
 1929. A conspectus of the marine and 
 fresh water Ciliata belonging to the sub- 
 order Tintinnoinea, with descriptions of 
 new species principally from the Agassiz 
 expedition to the eastern tropical Pacific, 
 1904-1905. Univ. Calif. Publ. Zool. 34: 1- 
 403. [A classical monograph on these 
 planktonic ciliates.] 
 KOZLOFF, E. N. 
 
 1955. Lwoffia cilifera gen. nov., sp. nov., 
 a ciliated member of the family Sphen- 
 ophryidae (Holotricha: Thigmotricha). 
 Biol. Bull. 108:283-289. [Morphology of 
 a ciliate ectocommensal on mussels.] 
 
 1960. Morphological studies on holotrich- 
 ous ciliates of the family Hysterocinetidae 
 I. Hysterocineta eiseniae Beers and 
 Ptychostomum campelomae sp. nov. J. 
 Protozool. 6: 41-50. [Morphology of mem- 
 bers of the Astomatida.] 
 
 1965. Colligocineta furax gen. nov., sp. 
 nov., an Ancistrocomid cilliate (Holo- 
 
 54 
 
tricha: Thigmotricha) from the Sabellid 
 polychaete Laonome kroyeri Malmgren. 
 J. Protozool. 12:333-334. [The most re- 
 cent of a long series of papers by this 
 author on the ciliates that occur in or on 
 marine invertebrates.] 
 1966. Phalacrocleptes vemiciformis gen. 
 nov., sp. nov., an unciliated ciliate from 
 the sabellid polychaete Schizobranchia 
 insignis Bush. Biol. Bull. 130:202-210. 
 [A small bizarre protozoon with short 
 tentacles used for attachment to the host.] 
 
 KUDO, R. R. 
 
 1966. Protozoology. C. C. Thomas, Spring- 
 field, 1174 p. [The latest edition of this 
 important textbook.] 
 
 LOEBLICH, A. R., JR., and H. TAPPAN. 
 
 1968. Annotated index to genera, subgen- 
 era, and suprageneric taxa of the ciliate 
 order Tintinnida. J. Protozool. 15:185- 
 192. [An important source of informa- 
 tion of the systematics of this order. See 
 also Tappan and Loeblich, 1968.] 
 
 LOM, J. 
 
 1964. The morphology and morphogenesis 
 of the buccal ciliary organelles in some 
 peritrichous ciliates. Arch. Protistenkd. 
 107:131-162. [Applied to protargol 
 method to members of many genera.] 
 
 LOM, J., J. 0. CORLISS, and C. NOIROT- 
 TIMOTHEE. 
 
 1968. Observations on the ultrastructure 
 of the buccal apparatus in thigmotrich 
 ciliates and their bearing on thigmotrich- 
 peritrich affinities. J. Protozool. 15:824- 
 840. [Members of these two orders close- 
 ly related due to comparable arrangement 
 of ciliary rows in the buccal cavity.] 
 
 MATSUDO, J., and J. L. MOHR. 
 
 1965. A new species of Heliochona, H. 
 psychra n. sp. (Ciliata: Chonotrichida), 
 from Nuwuk Lake, Alaska. J. Protozool. 
 12:523-527. [Morphological study.] 
 
 1968. Oenophorachona ectenolaemus n. 
 gen., n. sp. and Spirochona halophila n. 
 sp., two new marine chonotrichous cili- 
 ates. J. Protozool. 15:280-284. [Mor- 
 phological study.] 
 MOHR, J. L., J. A. LEVAGUE, and 
 H. MATSUDO. 
 
 1963. On a new collar ciliate of a gribble: 
 Lobochona prorates n. sp. on Limnoria 
 
 tripuncta. J. Protozool. 10:226-233. 
 [Morphological study.] 
 
 MUGARD, H. 
 
 1949. Contribution a l'etude des infusoires 
 hymenostomes histiophages. Ann. Sci. 
 Nat., Zool. Biol. Anim. (Ser. 11) 10:171- 
 268. [An important paper illustrating 
 patterns of cilia on several members of 
 the Uronematidae and Philasteridae.] 
 
 PRELLE, A. 
 
 1963. Observations sur le cilie holotriche 
 Woodruff ia rostrata (Johnson et Larson 
 1938). Arch. Zool. Exp. Gen. 102:239- 
 244. [Discussion of this trichostome and 
 its relationship to the Colpodidae.] 
 
 PUYTORAC, P. de. 
 
 1954. Contribution a l'etude cytologique et 
 taxonomique des Infusoires astomes. 
 Ann. Sci. Nat. Zool. Biol. Anim. 16:85- 
 270. [Important recent monograph on 
 the order Astomatida.] 
 RAABE, Z. 
 
 1949. Studies on the family Hysterocineti- 
 dae Kiesing. Ann. Mus. Zool. Polon. 14: 
 21-68. [Important study on the Asto- 
 matida.] 
 
 1963a. Systematics of the family Urceo- 
 lariidae Dujardin 1841. Acta Protozool. 
 1:121-138. [Revision of this family of 
 peritrichs.] 
 
 1963b. La stomatogenese chez Conchoph- 
 thirus (Ciliata, Thigmotricha). Acta 
 Protozool. 1:139-146. [Morphology and 
 life cycle of this thigmotrich.] 
 
 1970a. Ordo Thigmotricha (Ciliata - Hol- 
 otricha). II. Family Hemispeiridae. 
 Acta Protozool. 7:117-180. 
 
 1970b. Ordo Thigmotricha (Ciliata - Hol- 
 otricha). III. Familiae Ancistrocomidae 
 et Sphenophryidae. Acta Protozool. 7: 
 385-463. [These two papers are an im- 
 portant recent revision of this order.] 
 ROQUE, M. 
 
 1961. Recherches sur les infusoires cilies; 
 les hymenostomes peniculiens. Bull. Biol. 
 Fr. Belg. 95:431-519. [An important 
 monograph.] 
 SCHWARZ, S. 
 
 1964. Die Tintinnoidea. Hydrobiol. 23: 
 213-245. [A brief but concise and illus- 
 trated set of diagnoses to family and 
 genus.] 
 
 55 
 
SIKORA. J. 
 
 1963. Study on the parasitic ciliate Stei- 
 nella uncinata (Schultze). Acta Proto- 
 zool. 1: 13-20. [Morphology of this mem- 
 ber of the family Haptophryidae (order 
 Astomatida).] 
 
 SMALL, E. B. 
 
 1967. The Scuticociliatida, a new order of 
 the class Ciliatea (Phylum Protozoa, sub- 
 phylum Ciliophora). Trans. Am. Mi- 
 crosc. Soc. 86:345-370. [A rearrange- 
 ment of families and genera previously 
 placed in the order Hymenostomatida or 
 the order Thigmotrichida, on the basis of 
 differences in morphogenesis.] 
 
 TAPPAN, H., and A. R. LOEBLICH, JR. 
 
 1968. Lorica composition of modern and 
 fossil Tintinnida (Ciliata, Protozoa), sys- 
 tematics, geological distribution, and 
 some new tertiary taxa. J. Paleontol. 42: 
 1378-1394. 
 
 THOMPSON, J. C, JR. 
 
 1963. The generic significance of the buc- 
 cal infra-ciliature of the family Tetrahy- 
 menidae and a proposed new genus and 
 species, Paratetrahymena wassi. Vir- 
 ginia J. Sci. 14 (New Ser.) : 126-135. 
 [Brief paper on morphology.] 
 
 1964a. A redescription of Uronema mar- 
 inum, and a proposed new family Uro- 
 nematidae. Virginia J. Sci. 15:80-87. 
 [Morphological study, with photomicro- 
 graphs.] 
 
 1964b. Miamiensis avidus n.g., n. sp., a 
 marine facultative parasite in the ciliate 
 order Hymenostomatida. J. Protozool. 
 1 1 : 378-380. [A species apparently close- 
 ly allied to Uronema, but differing in ex- 
 act arrangement of buccal ciliature.] 
 
 1966. Glauconema trihymene n.g., n. sp., 
 a hymenostome ciliate from the Virginia 
 
 coast. J. Protozool. 12:393-394. [Mor- 
 phology of a Uronema-like ciliate.] 
 
 THOMPSON, J. C, and J. BERGER. 
 
 1965. Paranophrys marina n.g., n. sp., a 
 new ciliate associated with a hydroid 
 from the Northeast Pacific (Ciliata: 
 Hymenostomatida). J. Protozol. 12: 
 527-531. [Morphological study.] ' 
 
 TUFFRAU, M. 
 
 1967a. Prefectionnements et practique de 
 la technique d'impregnation au Protargol 
 des infusoires cilies. Protistologica 3:91- 
 98. [A discussion of the use of the pro- 
 targol or silver proteinate method in 
 staining of ciliates.] 
 1967b. Les structure fibrillaires soma- 
 tiques et buccales chez les cilies hetero- 
 triches. Protistologica 3:369-394. [Use 
 of the protargol technique in elucidating 
 the cilia of the buccal cavity and their 
 associated fibillar systems in various 
 heterotrichs.] 
 
 UHLIG, G. 
 
 1965. Untersuchungen zur Extraktion der 
 vagilen Mikrofauna aus marinen Sedi- 
 menten. Verh. Deutsch. Zool. Ges. 10: 
 151-157. [Describes use of melt water 
 from seawater ice to drive small meta- 
 zoans and protozoons from samples of 
 sand.] 
 
 VILLENEUVE-BRACHON, S. 
 
 1940. Recherches sur les cilies hetero- 
 triches, cinetome, argyrome, myonemes. 
 Formes nouvelles ou peu connus. Arch. 
 Zool. Exp. Gen. 82: 1-180. [An important 
 monograph on this order.] 
 
 WICHTERMAN, R. 
 
 1953. The biology of Paramecium. Blak- 
 iston Co., Inc., N.Y., 527 p. [An impor- 
 tant source of information on this widely 
 known ciliate.] 
 
 56 
 
SYSTEMATIC INDEX 
 
 Acanthostomella 50 
 
 Acineta 45 
 
 Acinetidae 41, 45 
 
 Acinetopsis 45 
 
 Acropisthium 44 
 
 Actinocyathtis 41, 46 
 
 Amphileptidae 35, 44 
 
 Amphileptus 44 
 
 Amphisiella 51 
 
 Amplectella 50 
 
 Amplectellopsis 50 
 
 Ancistrocomidae 14, 36, 40, 48 
 
 Ancistrospira 48 
 
 Ancistrum 48 
 
 Ancystropodium 50 
 
 Anigsteinia 49 
 
 Anophrys 
 
 salmacida 47 
 
 sarcophaga 47 
 
 Anoplophryidae 15, 46 
 
 Anoplophryopsis 46 
 
 Apostomatida 45, 46 
 
 Ascampbelliella 50 
 
 Askenasia 44 
 
 Askoella 34, 46 
 
 Askoellidae 34, 40, 46 
 
 Aspidisca 50 
 
 Astomatida 2, 46 
 
 Atopochilodon 44 
 
 Balantidiidae 11, 45 
 
 Balantidium 45 
 
 Balladyna 51 
 
 Banyidsella 51 
 
 Biggaria 21, 47 
 
 Blepharisma 49 
 
 Boveria 48 
 
 Brandtiella 24 
 
 Brooklynella 44 
 
 Buetschliella 46 
 
 Caenomorpha 49 
 
 Caenomorphidae 39, 49 
 
 Carchesium 49 
 
 Cardiostomatella 47 
 
 Caryotricha 50 
 
 Caulicola 49 
 
 Cephalotrichidium 49 
 
 Certesia 50 
 
 Cestus veneris 45 
 
 Chaenea 44 
 
 Chattonidiidae 16, 49 
 
 Chattonidium 49 
 
 Chilodochona 45 
 
 Chilodochonidae 31, 45 
 
 Chilodonatella 44 
 
 Chilodonella 44 
 
 Chilodontopsis 44 
 
 Chilophrya 44 
 
 Chlamydodon 44 
 
 Chlamydodontidae 33, 44 
 
 Chlamydonella 44 
 
 Choanostoma 44 
 
 Chonotrichida 34, 45 
 
 Chromidina 44 
 
 Ciliofaurea 44 
 
 Ciliospira 50 
 
 Cinetochilidae 33, 47 
 
 Cinetochihim 47 
 
 Cirratulus 46 
 
 Clathrostomatidae 11, 45 
 
 Climacocylis 50 
 
 Climacostomum 49 
 
 Cochliophilus 48 
 
 Codonaria 50 
 
 Codonella 50 
 
 Codonellidae 25, 50 
 
 Codonellopsidae 24, 50 
 
 Codonellopsis 50 
 
 Codonopsis 50 
 
 Coelosomides 45 
 
 Coelosomididae 7, 45 
 
 Cohnilembidae 20, 47 
 
 Cohnilembus 47 
 
 Colepidae 8, 44 
 
 Coleps 44 
 
 Colligocineta 48 
 
 Collinophrya 46 
 
 Colpoda 45 
 
 Colpodidae 11, 45 
 
 Conchophthiridae 13, 22, 48 
 
 Conchophyllum 48 
 
 Conchostoma longissimum 13, 45 
 
 Condylo stoma 49 
 
 Condylostomatidae 16, 49 
 
 Conidiophrys 2, 34, 40, 46 
 
 Corlissia 45 
 
 Corophium 46 
 
 Corynophrya 46 
 
 Cothurnia 49 
 
 Coxliella 50 
 
 Coxliellidae 24, 26, 50 
 
 57 
 
Crateristoma klndi 48 
 
 Crateristomatidae 35, 48 
 
 Crebriscoma 48 
 
 Cricundella 50 
 
 Cristigera 47 
 
 Cryptochilidium 20 
 
 cuenoti 47 
 
 Cryptochilum 20, 47 
 
 Cryptopharynx 44 
 
 Ctenoctophrys 44 
 
 Cyclidiidae 47 
 
 Cyclidmm 47 
 
 Cyclotrichium 44 
 
 Cymatocylis 50 
 
 Cyrtocaryum 46 
 
 halosydnae . 14 
 
 Cyttarocylididae 25, 50 
 
 Cyttarocylis 50 
 
 Dactylophrya 45 
 
 Dendrosomatidae 1, 43, 45 
 
 Dendrosomides 46 
 
 Dexiotricha 47 
 
 Dktyocysta 50 
 
 Dictyocystidae 25, 50 
 
 Didiniidae 38, 44 
 
 Didinium 44 
 
 Dileptus 44 
 
 Diophrys 50 
 
 Dipartiella 48 
 
 Discocephalus 50 
 
 Discophryidae 43, 46 
 
 Donax 48 
 
 Durchoniella 46 
 
 Dysteria 44 
 
 Dysteriidae 3, 33, 44 
 
 Ebalia 45 
 
 Ellobiophrya 48 
 
 Enchelyidae 9, 44 
 
 Enchelyodon 44 
 
 Enchelys 44 
 
 Endosphaera 45 
 
 Entodiscus 20, 47 
 
 Entorhipidiidae 20, 47 
 
 Entorhipidium 20, 47 
 
 Epalxella 49 
 
 Epalxellidae 39, 49 
 
 Ephelota 46 
 
 Ephelotidae 41, 42, 46 
 
 Epicancella 50 
 
 Epiclintes 51 
 
 Epiplocylididae 25, 50 
 
 Epiplocylis 50 
 
 Epiplocyloides 50 
 
 Epirhabdonella 50 
 
 Epistylis 48 
 
 Epistylididae 1, 30, 48 
 
 Euplotaspis 50 
 
 Euplotes 50 
 
 Euplotidae 37, 50 
 
 Euplotidium 50 
 
 Eabrea 49 
 
 Favella 27, 50 
 
 Favelloides 50 
 
 Foettingeriidae 10, 45, 46 
 
 Folliculinidae 15, 49 
 
 Frontonia 47 
 
 Frontoniella 47 
 
 Frontoniidae 18, 47 
 
 Gammarus 46 
 
 Gargarius 48 
 
 Gastrocirrhus 50 
 
 Gastrostyla 50 
 
 Geleia 45 
 
 Geleiidae 2, 12, 45 
 
 Glaucomidae 47 
 
 Glauconema 47 
 
 Goniocoma 48 
 
 Gruberia 49 
 
 Gruvelina 45 
 
 Gullmarella faurei 13, 45 
 
 Gymnostomatida . 44, 45 
 
 Halosydna 46 
 
 Halteriidae 29, 49 
 
 Haptophryidae 14, 46 
 
 Hartmannula 44 
 
 Helella 46 
 
 Helicoprorodon 44 
 
 Helicostoma 47 
 
 Helicostomella 50 
 
 Heliochona 45 
 
 Hemiophrys 44 
 
 Hemispeira 48 
 
 Hemispeiridae 22, 38, 48 
 
 Herpetophrya 46 
 
 Herpinella 46 
 
 Heterocineta 48 
 
 Heterocoma 48 
 
 Heterotrichida 34, 49 
 
 Histiobalantium 47 
 
 Histrimdus 50 
 
 Holocoma 48 
 
 Holophrya 44 
 
 Holosticha 50 
 
 Holostichidae 36, 50 
 
 58 
 
Hoplitophryidae 1, 14, 46 
 
 Hovasseiella 46 
 
 Hymenostomatida 2, 5, 47, 48 
 
 Hypocomagalma 48 
 
 Hypocomella 48 
 
 Hypocomidae 36, 48 
 
 Hypocomides 48 
 
 Hypocomina 48 
 
 Hypotrichida 50 
 
 Hysterocinetidae 21, 48 
 
 Idothea 46 
 
 Ileonema 44 
 
 Illex 46 
 
 Insignicoma 48 
 
 Intoshellinidae 1, 46 
 
 Isocomides 48 
 
 Isselina 48 
 
 Kentrochona 45 
 
 Kentrophoros 44 
 
 Keronopsis 50 
 
 Kiitricha 50 
 
 Laackmanniella 50 
 
 Lacazea 51 
 
 Lachmannella 46 
 
 Lacrymaria 44 
 
 Lagenophryidae 28, 48 
 
 Lagenophrys 48 
 
 Lagynophrya 44 
 
 Lagynurus 44 
 
 Laurentia 50 
 
 Lecanophrya 46 
 
 Lechriopyla 45 
 
 Leiotricha 48 
 
 Leprotintinnus 50 
 
 Lernaeophrya 45 
 
 Licnophora 49 
 
 Licnophoridae 29, 49 
 
 Liliimorpha 44 
 
 Limnoria 45 
 
 Litonotus 44 
 
 Lobochona 45 
 
 Lohmanniella 50 
 
 Loligo 46 
 
 Longitricha 44 
 
 Loxocephalidae 19, 47 
 
 Loxocephalus 47 
 
 Loxodes 44 
 
 Loxodidae 2, 33, 44 
 
 Loxophyllum 44 
 
 Luminella 50 
 
 Luxiella 50 
 
 Lwoffia 48 
 
 cilifera 40 
 
 Lynchella 44 
 
 Madsenia 47 
 
 Mesodinium 44 
 
 Metacineta 41 
 
 Metacylis 50 
 
 Metacystidae 9, 44 
 
 Metaeystis 44 
 
 Metastrombidium 49 
 
 Metephelota 46 
 
 Metopidae 17, 49 
 
 Metopus 49 
 
 Miamiensis 47 
 
 Microregma 44 
 
 Micrundella 50 
 
 Mirodysteria 44 
 
 Monodinium 44 
 
 Mylestoma 49 
 
 Mylestomatidae 39, 49 
 
 Nannophrya 44 
 
 Nassida 44 
 
 Nassulidae 6, 10, 44 
 
 Nassulopsis 44 
 
 Nebalia 45 
 
 Nephrocerca 45 
 
 Niemarshallia 50 
 
 Nucleocorbula adherens 48 
 
 Nucleocorbulidae 48 
 
 Odo7itochlamys 44 
 
 Odontostomatida 49 
 
 Oenophorachona 45 
 
 Oligotrichida 49 
 
 Onychodromus 50 
 
 Opalinopsidae 1, 13, 14, 46 
 
 Opalinopsis 46 
 
 Ophelia 46 
 
 Ophryocephalus 46 
 
 Ophryodendridae 1, 42, 46 
 
 Ophryodendron 46 
 
 Ophryoglena 47 
 
 Ophryoglenidae 18, 47 
 
 Opisthostyla 48 
 
 Orchitophyra 48 
 
 Orthodonella 44 
 
 Oxytricha 50 
 
 Oxytrichidae 37, 50 
 
 Parablepharisma 49 
 
 Parachilodonella 44 
 
 Paracineta 41, 46 
 
 Paradileptus 44 
 
 Paraeicplotes 50 
 
 59 
 
Parafavella 50 
 
 Paraholosticha 50 
 
 Parahypocoma 48 
 
 Paralembus 47 
 
 Parameciidae 18, 47 
 
 Paramecium 47 
 
 woodruff I 47 
 
 calkinsi 47 
 
 Pa)-anassula 45 
 
 Paranassulidae 45 
 
 Paranophrys 47 
 
 Parapodophyra 46 
 
 Pa raspathidium 45 
 
 Parastylonychia 50 
 
 Paratetrahymena 47 
 
 ParatrocMlia 44 
 
 Parauronema 47 
 
 Paraurostyla 51 
 
 Paravauchomia 49 
 
 Paravorticella 48 
 
 Parundella 50 
 
 Pelatraetus 44 
 
 Pelecophrya 48 
 
 Peniculi stoma 22, 47, 48 
 
 Perikaryon cesticola . . 12, 45 
 
 Peritrichida 1, 29, 48 
 
 Peritromidae 34, 49 
 
 Peritromus 49 
 
 Perseia 46 
 
 Petalotricha 50 
 
 Petalotrichidae 27, 50 
 
 Phalacrocleptes verruciformis 46 
 
 Phalacrocleptidae 43, 46 
 
 Phascolosoma 46, 47 
 
 Philaster 47 
 
 Philasteridae 21, 47 
 
 Philasterides 47 
 
 Phthorophrya 34, 46 
 
 Phytia 48 
 
 Pithothorax 44 
 
 Placus 44 
 
 socialis 6 
 
 Plagiocampa 44 
 
 Plagiopyla 45 
 
 Plagiopylidae 11, 12, 45 
 
 Plagiopyliella 45 
 
 Pkigiospira 48 
 
 Platycola 49 
 
 Platynematum 47 
 
 Platyophrya 44 
 
 Pleurocoptes 22, 47 
 
 Pleuronema 47 
 
 Pleuronematidae 19, 22, 47, 48 
 
 Podocyathus 41, 46 
 
 Podophrya 46 
 
 Podophryidae 41, 43, 46 
 
 Poljanskina 49 
 
 Polycycla 48 
 
 Polydora 46 
 
 Poroecus 24, 50 
 
 Porpostoma 47 
 
 Portunus 45 
 
 Potamoceros 46 
 
 Potomacus 47 
 
 Pottsiocles 45 
 
 Proboveria 48 
 
 Proplectella 50 
 
 Prorodon 44 
 
 mimeticus 6 
 
 Prorodontidae 8, 44 
 
 Protocruzia 34, 49 
 
 Protocymatocylis 50 
 
 Protophrya 48 
 
 Protorhabdonella 50 
 
 Protrichophora 45 
 
 Pseudocohnilembidae 20, 47 
 
 Pseudocohnilembus 47 
 
 Pseudogemma 45 
 
 Pseudoprorodon 45 
 
 Psilotricha 50 
 
 Psilotrichidae 37, 50 
 
 Ptychocylididae 26, 27, 50 
 
 Ptychocylis 26, 50 
 
 Pyxicola 49 
 
 Raabella 48 
 
 Remanella 44 
 
 Rhabdonella 50 
 
 Rhabdonellidae 27, 50 
 
 Rhabdonellopsis 50 
 
 Rhabdophrya 46 
 
 Rhabdostyla 48 
 
 Rhizocaryum 46 
 
 Rhizodomus 50 
 
 Rhopalophrya 44 
 
 Schizobranchia 46 
 
 Schizocalyptra 47 
 
 Schizocaryum dogieli 12, 45 
 
 Scuticociliatida 20, 47, 48 
 
 Scyphidia , 48 
 
 Scyphidiidae 30, 48 
 
 Sepia 46 
 
 Sessilina 29 
 
 Sigmocineta 44 
 
 Silenella ovoidea 36, 46 
 
 60 
 
Sonderia 45 
 
 Spasmostoma 44 
 
 Spathidiidae 8, 44 
 
 Sphenophrya 48 
 
 Sphenophryidae 2, 40, 41, 48 
 
 Spirobuetschliella 46 
 
 Spirochona 45 
 
 Spirochonidae 31, 45 
 
 Spirofilidae 15, 36, 51 
 
 Spirostomatidae 17, 49 
 
 Steinella 46 
 
 Stenosemella 50 
 
 Stentor 49 
 
 Stentoridae 16, 49 
 
 Stephanopogon 44 
 
 Stephanopogonidae 1, 35, 44 
 
 Stichotricha 51 
 
 Stomatophrya 44 
 
 Strobilidiidae 29, 50 
 
 Strobilidium 49 
 
 Strombidiidae 49 
 
 Strombidium 49, 50 
 
 Strongylidiam 51 
 
 Stylicauda 50 
 
 Stylochona 45 
 
 Stylochonidae 31, 45 
 
 Stylonychia 50 
 
 Suctorida 1, 2, 36, 39, 45 
 
 Swedmarkia 50 
 
 Tachyblaston 45 
 
 Tachysoma 50 
 
 Tetrahymenidae 19, 47 
 
 Thaumatophrya 46 
 
 Thecacineta 45 
 
 Thigmophyra 48 
 
 Thigmophryidae 13, 22, 48 
 
 Thigmotrichida 2, 46, 47, 48 
 
 Thuricola 49 
 
 Tiarina 44 
 
 Tintinnida 50 
 
 Tintinnidae 24, 27, 50 
 
 Tintinnidiidae 24, 50 
 
 Tintinnidium 50 
 
 Tintinnopsis 50 
 
 Tontonia 49 
 
 Tracheliidae 10, 44 
 
 Trachelius 44 
 
 Trachelocerca 45 
 
 Trachelocercidae 2, 9, 35, 45 
 
 Trachelonema 45 
 
 Trachelophyllum 44 
 
 Trachelor aphis 45 
 
 Trachelostyla 50 
 
 Trichodina 49 
 
 Trichodinella 44 
 
 Trichodininae 29, 48 
 
 Trichodinopsis 48 
 
 Trichophrya 45 
 
 Trichostomatida 12, 13, 45 
 
 Trichotaxis 50 
 
 Trimyema 45 
 
 Trimyemidae 38, 45 
 
 Tripartiella 48, 49 
 
 Trochilia 44 
 
 Trochilioides 44 
 
 Tropho gemma 46 
 
 Undella 50 
 
 Undellidae 27, 50 
 
 Undellopsis 50 
 
 Urceolaria 48 
 
 Urceolariidae 28, 48 
 
 Urceolarinae 29, 48 
 
 Uroleptopsis 50 
 
 Uroleptus 50 
 
 Uronema 47 
 
 Uronematidae 21, 47 
 
 Uronychia 50 
 
 Uropedalium 47 
 
 Urostrongylum 51 
 
 Urostyla 51 
 
 Urostylidae 37, 51 
 
 Urotricha 44 
 
 Vaginicola 49 
 
 Vaginicolidae 28, 49 
 
 Vasicola 44 
 
 Vauchomia 49 
 
 Vorticella 49 
 
 Vorticellidae 1, 30, 49 
 
 Wailesia 50 
 
 Wangiella 50 
 
 Woodruffia 45 
 
 Xystonella 50 
 
 Xystonellidae 26, 50 
 
 Xystonellopsis 50 
 
 Zonotrichium 44 
 
 Zoothamnium 49 
 
 61 
 
ACKNOWLEDGMENTS 
 
 Preparation of the "Marine Flora and Fauna of the 
 Northeastern United States" is being coordinated by 
 the following Board: 
 
 Coordinator: Melbourne R. Carriker, Systematics- 
 Ecology Program, Marine Biological 
 Laboratory, Woods Hole, Mass. 
 
 Advisers: Marie B. Abbott, Systematics-Ecology 
 Program. 
 
 Arthur G. Humes, Boston University Ma- 
 rine Program, and Systematics-Ecology 
 Program. 
 
 Wesley N. Tiffney, Department of Biology, 
 Boston University, and Systematics- 
 Ecology Program. 
 
 Ruth D. Turner, Museum of Comparative 
 Zoology, Harvard University, and Syste- 
 matics-Ecology Program. 
 
 Roland L. Wigley, National Marine Fish- 
 eries Service, Woods Hole, Mass. 
 
 Robert T. Wilce, Department of Botany, 
 University of Massachusetts, and Syste- 
 matics-Ecology Program. 
 
 The Board established the format for the "Marine 
 Flora and Fauna of the Northeastern United States," 
 invites systematists to collaborate in the preparation of 
 manuals, reviews manuscripts, and advises the Scientific 
 Editor of the National Marine Fisheries Service. 
 
 John 0. Corliss, Helen T. Loeblich, Jesse C. Thompson, 
 Jr., and Eugene B. Small submitted criticisms, sugges- 
 tions, and helpful additions for construction of the key. 
 Illustrations were penciled by Arthur C. Borror and 
 were inked by Susan P. Heller. Eva S. Montiero typed 
 the manuscript. Preparation of the manual was sup- 
 ported in part by Project No. 18050 FBW, Federal Water 
 Pollution Control Administration, to Borror, and by 
 Grant GB-24,832 from the National Science Foundation 
 to the Systematics-Ecology Program. A contribution of 
 the Jackson Estuarine Laboratory. 
 
 COORDINATOR'S COMMENTS 
 
 Publication of the "Marine Flora and Fauna of the 
 Northeastern United States" is most timely in view of 
 the growing universal emphasis on environmental work 
 and the urgent need for more precise and complete ident- 
 ification of coastal organisms than has been available. 
 It is mandatory, wherever possible, that organisms be 
 identified accurately to species. Accurate scientific names 
 unlock the great quantities of biological information 
 storec in libraries, obviate duplication of research al- 
 ready done, and make possible prediction of attributes 
 of organisms that have been inadequately studied. 
 
 Dr. Arthur C. Borror began his systematic investi- 
 gation of the ciliates of the east coast of the United 
 States in 1959 while pursuing graduate studies at Flor- 
 ida State University, Tallahassee. He moved to the 
 University of New Hampshire in 1961, where he has 
 continued his research on the morphology and ecology 
 of marine ciliates. 
 
 Manuals are available for purchase from the Super- 
 intendent of Documents, U.S. Government Printing Of- 
 fice, Washington, D.C. 20402. The manuals so far pub- 
 lished in the series and their cost per copy are listed 
 below. 
 
 COOK, DAVID G., and RALPH O. BRINKHURST. Marine flora and fauna of the Northeastern 
 
 United States. Annelida : Oligochaeta $0.35 
 
 BORROR, ARTHUR C. Marine flora and fauna of the Northeastern United States. Protozoa: 
 
 Ciliophora 0.65 
 
 62 
 
349. Use of abstracts and summaries as communica- 
 tion devices in technical articles. By F. Bruce 
 Sanford. February 1971, iii + 11 PP-, 1 fig. 
 
 350. Research in fiscal year 1969 at the Bureau of 
 Commercial Fisheries Biological Laboratory, 
 Beaufort, N.C. By the Laboratory staff. No- 
 vember 1970, ii + 49 pp., 21 figs., 17 tables. 
 
 351. Bureau of Commercial Fisheries Exploratory 
 Fishing and Gear Research Base, Pascagoula, 
 Mississippi, July 1, 1967 to June 30, 1969. By 
 Harvey R. Bullis, Jr., and John R. Thompson. 
 November 1970, iv + 29 pp., 29 figs., 1 table. 
 
 352. Upstream passage of anadromous fish through 
 
 navigation locks and use of the stream for spawn- 
 ing and nursery habitat, Cape Fear River N C 
 1962-66. By Paul R. Nichols and Darrell E. 
 Louder. October 1970, iv + 12 pp., 9 figs., 4 
 tables. 
 
 356. Floating laboratory for study of aquatic organ- 
 isms and their environment. By George R. 
 Snyder, Theodore H. Blahm, and Robert J. Mc- 
 Connell. May 1971, iii + 16 pp., 11 figs'. 
 
 361. Regional and other related aspects of shellfish 
 consumption — some preliminary findings from 
 the 1969 Consumer Panel Survey. By Morton 
 M. Miller and Darrel A. Nash. June 1971, iv + 
 18 pp., 19 figs., 3 tables, 10 apps. 
 
UNITED STATES PENN state university libraries 
 
 DEPARTMENT OF COMMERCE "' 
 
 NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION 
 
 NATIONAL MARINE FISHERIES SERVICE 
 
 SCIENTIFIC PUBLICATIONS STAFF 
 
 ROOM 450 
 
 FOURTH CLASS 
 
 A00007201flbQb 
 
 1 107 N.E 45TH ST ..S.MAIL 
 
 SEATTLE. WA 98105 
 OFFICIAL BUSINESS 
 
 UH