Cap 17

extra-oral, near kinetids of “brosse” ; oral region, 
apical to subapical, surrounded by circumoral diki-
netids; brosse (homologous to the oral polyki-
netids of oligohymenophoreans?), typically of 
three or more dikinetidal rows bearing clavate 
cilia, varying from parallel to perpendicular to 
body axis, and developing on parental ventral 
surface ; cytostome, round or elliptical, sometimes 
in shallow atrium, which is lined by oral ridges 
supported by two unequal rows of microtubules 
(homologous to the oral ribs of oligohymenophore-
ans?); most species carnivores or scavengers, a few 
algivorous; widely distributed in freshwater and 
marine habitats; eight families. 
 Family BALANIONIDAE Small & Lynn, 1985 
 Size, small; shape, ovoid with truncate, flat-
tened anterior pole ; free-swimming; somatic 
ciliation, restricted to an apparent girdle encir-
cling the anterior 1/2–3/4 of the cell ; caudal 
cilium, present; brosse, inconspicuous, with its 
units internal to oral dikinetids of circumoral 
ciliature ; oral region, on flattened anterior end, 
with very elongate oral “palps” bearing toxicysts, 
surrounding cytostome; macronucleus, globular 
to ellipsoid; micronucleus, present; contractile 
17.3 The Ciliate Taxa to Genus 407
vacuole, present; cytoproct (?); feeding on bacte-
ria, small algae, and heterotrophic flagellates; in 
marine and freshwater habitats, in plankton; one 
genus.
 – Balanion Wulff, 1919 
 Family COLEPIDAE Ehrenberg, 1838 
 Size, small to large; shape, barrel-like, with 
prominent anterior and caudal spines often present; 
free-swimming; alveoli as cuirass of longitudinal 
rows of armored calcium carbonate plates with 
small lateral teeth ; somatic ciliation, holotrichous, 
with caudal cilium, typically long; oral extrusomes 
as toxicysts; brosse as three, short, inconspicuous 
files of dikinetids; oral region, apical, surrounded 
by oral dikinetids whose circle may be broken 
by the intrusion of the brosse kinetids; macronu-
cleus, globular to elongate ellipsoid; micronucleus, 
present; contractile vacuole, present; cytoproct (?); 
scavengers and histophages; in marine and fresh-
water habitats; eight genera. 
 – Baikalocoleps Obolkina, 1995 
 – Coleps Nitzsch, 1827 
 – Kotinia Obolkina in Aescht, 2001 
 – Macrocoleps Obolkina, 1995 
 – Plagiopogon Stein, 1859 
 – Planicoleps Dragesco & Dragesco-Kernéis, 1991 
 – Tiarina Bergh, 1881 
 – Tiarinella Obolkina, 1995 
 Family HOLOPHRYIDAE Perty, 1852 
 (syn. Cryptocaryonidae ) 
 Size, small to large; shape, ovoid to cylindroid; 
free-swimming; somatic ciliation, holotrichous, 
often dense, sometimes with caudal cilia; oral 
region, apical or subapical, surrounded by circu-
moral ciliature; somatic extrusomes as mucocysts 
and oral extrusomes as toxicysts; brosse as several 
to many kinetofragments, with somatic kine-
ties parallel to the brosse (i.e., aklitoloph) or 
abutting against the brosse region (i.e., euklito-
loph) ; oral region, apical to subapical, surrounded 
by circumoral dikinetids; macronucleus, globular 
to ellipsoid to elongate and even ribbon-like; 
micronucleus, present; contractile vacuole, present; 
cytoproct (?); scavengers and histophages with 
Cryptocaryon as a parasite of marine fishes ; in 
marine and freshwater habitats; five genera and one 
genus incertae sedis . 
NOTE : Foissner, Berger, and Kohmann (1994) 
discovered problems with the nomenclature of 
Holophrya , Prorodon , and Pseudoprorodon , 
which necessitated the redescription of the genus 
Holophrya with Holophrya ovum Ehrenberg, 1831 
as type. This genus presents the form of what was 
previously known as Prorodon (e.g. in Corliss, 
1979; Small & Lynn, 1985). 
 – Cryptocaryon Brown, 1951 
 – Holophrya Ehrenberg, 1831 
 – Paraprorodon Foissner, 1983 
 – Pelagothrix Foissner, Berger, & Schaumberg, 
1999 
 – Pleurofragma Jankowski, 1976 
Incertae sedis in Family Holophryidae 
 – Fundenia Vuxanovici, 1962 
 Family LAGYNIDAE Sola, Guinea, Longas, & 
Fernández-Galiano, 1990 
 Size, small; shape, pyriform when alive; free-
swimming; somatic ciliation as perioral ciliature 
or a girdle, completely encircling cell apex as 
“trikinetids” ; oral region, apical, with brosse as 
3–4 inconspicuous rows between perioral cili-
ature and circumoral dikinetids ; macronucleus, 
globular to reniform; micronucleus, may be mul-
tiple; contractile vacuole, present; cytoproct (?); 
feeding on flagellates and other smaller protists; in 
freshwater habitats; one genus. 
 – Lagynus Quennerstedt, 1867 
 Family PLACIDAE Small & Lynn, 1985 
 Size, small to medium; shape, ovoid, slightly 
flattened; free-swimming; somatic ciliation, 
holotrichous, having slightly spiralling kine-
ties with striae between ; somatic extrusomes as 
mucocysts; “oral” extrusomes as toxicysts dis-
persed along the brosse or in a lateral pocket 
in Spathidiopsis ; brosse as a single dikinetid 
file, extending posteriorly from the circumoral 
dikinetid ring ; oral region, subapical, a slightly 
elongate groove; macronucleus, ellipsoid to elon-
gate ellipsoid; micronucleus, present; contractile 
vacuole, present; cytoproct (?); feeding on flagel-
408 17. The Ciliate Taxa Including Families and Genera
lates and smaller ciliates; in marine and brackish 
habitats; two genera. 
 – Placus Cohn, 1866 
 – Spathidiopsis Fabre-Domergue, 1889 
 Family PLAGIOCAMPIDAE Kahl, 1926 
 Size, small; shape, ovoid; free-swimming; 
somatic ciliation, holotrichous; caudal cilia, com-
mon; oral extrusomes as toxicysts; brosse of three 
units, on posterior right of oral area, opposite oral 
dikinetids; oral region, subapical with oral diki-
netids on dorsal-right, as semicircle accompa-
nied by extensible “palps” or lappets in which 
toxicysts reside ; macronucleus, globular to ellip-
soid; micronucleus, may be very small; contractile 
vacuole, present; cytoproct (?); feeding on bacteria, 
dinoflagellates, microalgae, and other ciliates; in 
marine, freshwater, and terrestrial habitats; three 
genera and one genus incertae sedis . 
 – Chilophrya Kahl, 1930 
 – Paraurotricha Foissner, 1983 
 – Plagiocampa Schewiakoff, 1893 
Incertae sedis in Family Plagiocampidae
 – Plagiocampides Foissner, Agatha, & Berger, 2002 *
 Family PRORODONTIDAE Kent, 1881 
 (syn. Amphibot[h]rellidae [for Amphibothridae ]) 
 Size, medium to large; shape, ovoid; free-
swimming; somatic ciliation, holotrichous; “ bro-
sse” as an extension of the unclosed circumoral 
ciliature ; oral region, subapical, as an elongate 
groove; macronucleus, ribbon-like; micronucleus, 
may be multiple; contractile vacuole, present; 
cytoproct (?); feeding on other protists, even cili-
ates and small metazoans; in freshwater habitats; 
one genus. 
NOTE : Foissner, Berger, and Kohmann (1994) 
discovered problems with the nomenclature of 
Holophrya , Prorodon , and Pseudoprorodon , which 
necessitated applying the name Prorodon to cili-
ates similar to the type Prorodon niveus Ehrenberg, 
1834. This genus presents the form of what was 
previously known as Pseudoprorodon (e.g. in 
Corliss, 1979; Small & Lynn, 1985), which is an 
objective synonym of Prorodon . 
 – Prorodon Ehrenberg, 1834 
 Family UROTRICHIDAE Small & Lynn, 1985 
 (syn. Bursellopsidae ) 
 Size, small to large; shape, ovoid to cylindroid; 
free-swimming; somatic ciliation, with posterior 
1/4–1/3 of body non-ciliated and rest of body 
evenly ciliated ; caudal cilia, typical; brosse, two 
or three units, outside circumoral dikinetids, and 
quite elongate in Longifragma ; oral region, apical 
to subapical, can be quite expansive, almost cavity-
like; macronucleus, globular to elongate ellipsoid; 
micronucleus, present; contractile vacuole, present; 
cytoproct (?); feeding on bacteria, microalgae, and 
other smaller protists; in marine and freshwater 
habitats, often planktonic; six genera. 
 – Bursellopsis Corliss, 1960 
 – Dissothigma Jankowski, 1976 
 – Longifragma Foissner,