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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

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315. Synopsis of biological data on the chum salmon,
Oncorhynchus keta (Walbaum) 1792. By Rich-
ard G. Bakkala. March 1970, iii + 89 pp., 15
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319. Bureau of Commercial Fisheries Great Lakes
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330. EASTROPAC Atlas: Vols. 4, 2. Catalog No.

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331. Guidelines for the processing of hot-smoked chub.
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Emerson. January 1970, iv + 23 pp., 8 figs.,

2 tables.

332. Pacific hake. (12 articles by 20 authors.) March
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333. Recommended practices for vessel sanitation and
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335. Progress report of the Bureau of Commercial
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Research, Pesticide Field Station, Gulf Breeze,
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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.

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1969. By Division of Economic Research. April

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342. Report of the Bureau of Commercial Fisheries
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344. Bureau of Commercial Fisheries Tropical Atlan-
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346. Sportsman's guide to handling, smoking, and pre-
serving Great Lakes coho salmon. By Shearon
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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
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NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION
Robert M. White, Administrator

NATIONAL MARINE FISHERIES SERVICE
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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. BORROR1

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 cm3) 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&Z1

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.

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AMMERMANN, VON D.

1968. Die Kernverhaltnisse des Ciliaten
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description of morphology.]

BERGER, J.

1961a. The comparative buccal morphol-
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1961b. Stomatogenic evidence for the hy-
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1961c. Systematic reassignment, based on
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1961d. Morphology and systematic posi-
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1965. Infraciliary morphology of a little-
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ond Int. Conf. Protozool. No. 255. [Dis-
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BORROR, A. C.

1961. Feeding apparatus of the ciliate
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1962. Ciliate Protozoa of the Gulf of Mex-
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1963. Morphology and ecology of the ben-
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[Morphology of many species; four new
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1965a. Morphological comparison of Di-
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[Intraspecific variations tabulated.]

1965b. New and little-known tidal marsh
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1968a. Nigrosin-HgCl2-Formalin; a stain-
fixative for ciliates (Protozoa, Cilioph-
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background.]

1968b. Systematics of Euplotes (Cilioph-
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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
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families and genera are diagnosed and
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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.
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CEPEDE, C.

1910. Recherches sur les infusoires astom-
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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-
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[A description of the life cycle of the non-
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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.

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