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& Kahan, 1986 
Incertae sedis in Family Platyophryidae 
 – Ottowphrya Foissner, Agatha, & Berger, 2002 *
 Family SAGITTARIIDAE Grandori & Grandori, 
 (for Proshymenidae ) 
 Size, small; shape, ovoid; free-swimming, but 
some species in mucous sheaths; somatic ciliation, 
holotrichous, but sparse and with caudal cilium; oral 
region on rounded anterior end of cell with paro-
ral in one segment and serial left oral polykinetid 
zone of the same length ; macronucleus, globular 
to ellipsoid; micronucleus, in perinuclear space (?); 
contractile vacuole, present; cytoproct (?); bacte-
rivorous; in terrestrial habitats; one genus. 
 – Sagittaria Grandori & Grandori, 1934 
17.3 The Ciliate Taxa to Genus 405
 Family WOODRUFFIIDAE von Gelei, 1954 
 (syn. Woodruffidae ) 
 Size, small to large; shape, broadly ovoid; free-
swimming; somatic ciliation, holotrichous, with 
anterior ends of left somatic kineties typically 
forming several paratene-like rows bordering the 
serial left oral polykinetids ; oral region, subapical, 
slanted, on right side of cell; right paroral dikinetid 
not segmented; serial left oral polykinetids, in a 
more or less distinctive adoral zone extending 
into preoral suture, thus longer than paroral ; sto-
matogenesis, pleurotelokinetal with parental paroral 
probably reorganized during the process; macronu-
cleus, globular to ellipsoid; micronucleus, in perinu-
clear space; contractile vacuole, present; cytoproct, 
present; feeding on cyanobacteria, fungal spores, and 
smaller protists, including other ciliates; in marine, 
freshwater, and terrestrial habitats; six genera. 
 – Etoschophrya Foissner, Agatha, & Berger, 2002 *
 – Kuklikophrya Njiné, 1979 
 – Rostrophrya Foissner, 1993 
 – Rostrophryides Foissner, 1987 
 – Woodruffia Kahl, 1931 
 – Woodruffides Foissner, 1987 
Incertae sedis in Order Cyrtolophosidida 
kamp, Song, & Wilbert, 1989 
 Size, small; shape, ovoid; free-swimming; somatic 
ciliation as kineties of dikinetids, restricted to right 
(= ventral) surface with a single somatic kinety of 
dikinetids on the anterior left ; oral region, equato-
rial, with an uninterrupted paroral on right and several 
rectangular oral polykinetids on left; distinct cyr-
tos-like cytopharyngeal apparatus ; macronucleus, 
globular, may be in two nodules; micronucleus, in 
perinuclear space (?); contractile vacuole, present; 
cytoproct (?); feeding on microalgae, especially dia-
toms; in freshwater habitats, particularly spring-fed 
margins of rivers; two genera. 
 – Hackenbergia Foissner, 1997 
 – Pseudochlamydonella Buitkamp, Song, & 
Wilbert, 1989 
 Order Sorogenida Foissner, 1985 
 Size, small; shape, elongate, ovoid; free-swim-
ming; somatic ciliation, holotrichous; oral region 
at anterior pole of cell with paroral of dikinetids 
on right and several serial left oral polykinetids, 
together forming almost a closed circle around 
the cytostome ; stomatogenesis, pleurotelokinetal 
with parental structures retained during the proc-
ess; macronucleus, globular to slightly ellipsoid; 
micronucleus, present; contractile vacuole, present; 
cytoproct, present; feeding on smaller Colpoda
species; only in terrestrial habitats, especially 
on submerged leaves, where it may aggregate to 
form aerial sorocarps, like some myxomycetes, 
enclosing “spores” (= resting cysts) ; one family. 
 Family SOROGENIDAE Bradbury & Olive, 1980 
 With characteristics of the order; one genus. 
 – Sorogena Bradbury & Olive, 1980 
Incertae sedis in Class COLPODEA 
 – Rhyposophrya Kahl, 1933 
 Class PROSTOMATEA Schewiakoff, 1896 
 (syns. Apicostomata p.p ., Paramastigina p.p ., 
 Prostomata p.p ., Prostomina p.p ., Telostomata p.p .) 
 Size, small to large; shape, ovoid to cylin-
droid; free-swimming with some sessile forms 
in loricae; alveoli, often well-developed; somatic 
ciliation, holotrichous, but with reduced ciliation 
in posterior in some taxa; somatic kineties often 
arranged in complete circumferential paratenes 
with dikinetids at their anterior extremities ; 
somatic monokinetids usual, with radial transverse 
ribbon, slightly convergent postciliary ribbon, and 
anteriorly directed kinetodesmal fibril that does not 
overlap those of other kinetids; somatic dikinetid, 
when present, with posterior kinetosome derived 
from monokinetid and anterior kinetosome bearing 
only a tangential transverse ribbon; oral region, 
apical, subapical, or lateral with cytostome 
at or near body surface; circumoral ciliation 
with oral dikinetids (a paroral homologue?), 
radial to tangential to perimeter of oral area; 
oral dikinetid postciliary ribbons extending 
laterally from each dikinetid, overlapping one 
another, and, in some species, forming a circu-
lar microtubular band that supports the walls 
of a shallow atrium ; cytopharyngeal apparatus, 
cyrtos-like, with nematodesmata originating from 
the bases of circumoral dikinetids; stomatogenesis, 
406 17. The Ciliate Taxa Including Families and Genera
 merotelokinetal, with migration of ventral anlagen 
subapically to apically; macronucleus, homomer-
ous; bacterivorous, algivorous, carnivorous, and 
histophagous, including Cryptocaryon as a para-
site of marine fishes ; in freshwater, terrestrial, 
and marine habitats, widely distributed with some 
planktonic species; two orders and one family 
incertae sedis . 
 Order Prostomatida Schewiakoff, 1896 
 (syn. Vasicolina p.p .) 
 Size, small to medium; shape, cylindroid; free-
swimming; loricae produced by several species; 
somatic ciliation, holotrichous, clearly “radi-
ally symmetrical” ; paratenes, typically conspicu-
ous; oral region, apical, surrounded by circumoral 
dikinetids; brosse, absent ; toxicysts, absent ; sto-
matogenesis, not described; microphagous; two 
 Family APSIKTRATIDAE Foissner, Berger, & 
Kohmann, 1994 
 (syn. Enchelyidae p.p ., Holophryidae p.p .) 
 Size, small to medium; shape, cylindroid; free-
swimming; somatic kineties, mostly bipolar, 
but some inserted on an indistinct suture as 
short segments between bipolar ones ; paratenes, 
present but not conspicuous; oral region, apical, 
surrounded by simple oral dikinetids; macronu-
cleus, globular; micronucleus, present; contractile 
vacuole, present; cytoproct, present; feeding on 
cyanobacteria and microalgae; in freshwater habi-
tats; one genus. 
NOTE : Foissner, Berger, and Kohmann (1994) 
discovered problems with the nomenclature of 
Holophrya , Prorodon , and Pseudoprorodon , which 
necessitated the establishment of the new genus 
Apsiktrata with the type species Urotricha graci-
lis Penard, 1922. This genus presents the form of 
what was previously known as Holophrya (e.g. in 
Corliss, 1979; Small & Lynn, 1985). 
 – Apsiktrata Foissner, Berger, & Kohmann, 1994 
 Family METACYSTIDAE Kahl, 1926 
 Size, small to large; shape, cylindroid; free-
swimming, but living in a pseudochitinous or 
gelatinous lorica ; somatic kineties, bipolar ; 
paratenes conspicuous ; caudal cilia, present; oral
region, apical, with perioral ciliature composed 
of several rings of ciliature, surrounding the 
circumoral ciliature ; macronucleus, globular to 
ellipsoid; micronucleus, present; contractile vacu-
ole, present; cytoproct (?); bacterivorous (?); in 
marine, brackish, and freshwater habitats; three 
 – Metacystis Cohn, 1866 
 – Pelatractus Kahl, 1930 
 – Vasicola Tatem, 1869 
 Order Prorodontida Corliss, 1974 
 (syns. Bursellopsida p.p ., Colepina, Placina ) 
 Size, small to large; shape, ovoid to cylindroid; 
free-swimming; alveoli, well-developed, including 
calcium carbonate concretions as skeletal plates 
in the Family Colepidae; somatic ciliation, hol-
otrichous, but may be reduced in posterior half 
of cell, which typically bears one to many cau-
dal cilia; somatic extrusomes as mucocysts; oral
extrusomes as toxicysts, may be in oral palps or

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