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of Amphioxus ; 
seven genera and two genera incertae sedis . 
 – Apofrontonia Foissner & Song, 2002 *
 – Didieria Small & Lynn, 1985 
 – Disematostoma Lauterborn, 1894 
 – Frontonia Ehrenberg, 1838 
 – Frontoniella Wetzel, 1927 (subj. syn. 
Frontonia ) 
 – Paraclathrostoma Small & Lynn, 1985 
 – Wenrichia Jankowski, 1967 (subj. syn. 
Disematostoma ) 
Incertae sedis in Family Frontoniidae 
 – Schistophrya Kahl, 1933 
 – Sigmostomum Gulati, 1925 (subj. syn. Frontonia ) 
 Family LEMBADIONIDAE Jankowski in Corliss, 
 Size, small to medium; shape, broadly ovoid; 
free-swimming; somatic ciliation, holotrichous, 
with postoral suture; caudal cilia, long, forming 
a tuft; oral region, expansive, occupying nearly 
entire ventral surface; left oral cilia appearing 
as one long polykinetid (probably longitudinally 
fused peniculi) ; paroral, long, accompanied on 
its right by a long file of dikinetids (an ophry-
okinety?) ; macronucleus, elongate, ellipsoid; 
micronucleus, present; contractile vacuole, single; 
cytoproct (?); feeding on flagellates and other cili-
ates; in freshwater habitats, often planktonic; one 
 – Lembadion Perty, 1849 
 Family MARITUJIDAE Jankowski in Small & 
Lynn, 1985 
 Size, medium; shape, subspheroid, barrel-
shaped; free-swimming; somatic ciliation, holot-
richous, forming distinct transverse paratenes 
over the body; ophryokineties, numerous, encir-
cling “right” of oral region; oral region, on 
anterior surface of body ; oral structures as a 
paroral and three long polykinetids (i.e. peniculi); 
macronucleus, ribbon-like; micronucleus, present; 
contractile vacuole, present; cytoproct (?); feeding 
on bacteria and microalgae (?); in freshwater habi-
tats, planktonic; one genus. 
 – Marituja Gajewskaja, 1928 
 Family NEOBURSARIDIIDAE Dragesco & 
Tuffrau, 1967 
 Size, large, to over 700 µm in length; shape, 
elongate ovoid, typically rounded and broader at 
anterior end; free-swimming; somatic ciliation, 
holotrichous, dense, with “heterotrich-looking”, 
extensive, false adoral zone of polykinetids, 
actually formed by transverse paratenes of 
somatic kineties, lying in a prebuccal area as a 
much expanded preoral groove occupying the 
anterior half of the body ; oral region, midventral, 
expansive, with inner or right-most oral polyki-
netid of four widely spaced rows (i.e., a quadrulus); 
macronucleus, dumbbell-shaped; micronuclei, mul-
tiple; contractile vacuoles, two; cytoproct, present; 
17.3 The Ciliate Taxa to Genus 413
carnivorous on other ciliates ; in freshwater 
habitats, only pan-tropical ; one genus. 
 – Neobursaridium Balech, 1941 
 Family PARAMECIIDAE Dujardin, 1840 
 Size, medium; shape, elongate ovoid, rounded 
and/or pointed at either or both ends (the 
“slipper-shaped animalcules,” so long and 
well known); free-swimming; somatic ciliation, 
holotrichous, dense, with a prebuccal area 
as a preoral groove (formerly called a “ves-
tibulum”) not so much expanded, covered by 
paratenes, and leading to oral region ; oral 
cavity in anterior half to equatorial, with inner 
or right-most oral polykinetid of four widely 
spaced rows (i.e., a quadrulus) on dorsal wall 
of oral cavity; macronucleus, ellipsoid to elon-
gate ellipsoid; micronucleus, may be multiple; 
contractile vacuoles, typically two; cytoproct, 
present; some species containing zoochlorellae; 
feeding on bacteria and microalgae; in brackish 
and freshwater habitats; one genus and one genus 
incertae sedis . 
 – Paramecium O.F. Müller, 1773 
Incertae sedis in the Family Parameciidae 
 – Physanter Jankowski, 1975 
 Family STOKESIIDAE Roque, 1961 
 Size, medium; shape, distinctly cone- or 
heart-shaped, with flattened ventral surface 
and humped dorsal surface ; free-swimming; 
somatic ciliation, holotrichous; oral region, rela-
tively large, as a V-shaped ventral depression 
on a somewhat flattened ventral body surface ; 
oral region, midventral, with inner or right-most 
oral polykinetid of four widely spaced rows (i.e., 
a quadrulus); macronucleus, ellipsoid to lenticu-
lar; micronucleus, may be multiple; contractile 
vacuole, present; cytoproct, present; feeding on 
flagellates and microalgae, such as diatoms; in 
freshwater habitats, typically planktonic; two 
 – Parastokesia Jankowski, 1967 (subj. syn. 
Disematostoma ) 
 – Stokesia Wenrich, 1929 
 Order Urocentrida Jankowski, 1980 
 (syn. Urocentrina ) 
 Size, medium; shape, short, cylindrical, with 
larger, rounded anterior half ; free-swimming, 
but may be temporarily attached to the substra-
tum by a mucous thread; somatic ciliation as a 
distinct equatorial girdle; caudal cilia, forming 
a conspicuous tuft that is used for temporary 
attachment to substrates by a mucous thread; 
somatic kinetids only as monokinetids with 
broad, tangential transverse ribbon; somatic 
extrusomes as mucocysts ; no depressed preoral 
area; oral region, equatorial to subequatorial; oral 
structures as a paroral along the right margin of the 
oral opening and three oral polykinetids of three 
rows each along the dorsal-left wall; macronu-
cleus, band-like; micronucleus, present; contractile 
vacuole, single, with multiple collecting canals; 
cytoproct (?); feeding on bacteria and smaller 
protists; in freshwater habitats, typically in ponds 
where it attaches to the substrate and rotates on its 
mucous thread; one family. 
 Family UROCENTRIDAE Claparède & Lachmann, 
 (syn. Calceolidae ) 
 With characteristics of the order; one genus. 
 – Urocentrum Nitzsch, 1827 
 Subclass Scuticociliatia Small, 1967 
 (syns. Scuticostomata , Stichostomata p.p .) 
 Size, small to medium; shape, ovoid to elongate 
ovoid; mostly free-swimming, rarely restricted to 
a secreted lorica; alveoli, well-developed; somatic 
ciliation, holotrichous, though sometimes sparse, 
with thigmotactic field of somatic cilia in some 
symbiotic species; caudal cilia, often one or more; 
somatic dikinetids, predominant, both kineto-
somes ciliated over much of the body; extrusomes 
as somatic mucocysts; mitochondria, large, elon-
gate, cortically located, often-fused as a single 
cortical chondriome; oral region, quite variable 
in shape and extent, with right paroral as a 
file of dikinetids divided into a , b , and c seg-
ments, especially conspicuous during stoma-
togenesis, and typically three oral polykinetids, 
often as membranoids, on left; stomatogenesis, 
 scuticobuccokinetal, involving proliferation of 
414 17. The Ciliate Taxa Including Families and Genera
kinetosomes at least from paroral c segment or 
a scutica or scuticovestige, posterior or paral-
lel to paroral a and b segments ; macronucleus, 
typically single but fragmented in some species; 
micronuclei, one to many; commonly bacterivo-
rous, but facultatively (?) parasitic species, often 
histophagous on crustaceans and fish ; cysts wide-
spread; abundant in marine habitats, but also in 
some freshwater and terrestrial habitats, as free-
living forms or in symbiotic association primarily 
with invertebrates, such as molluscs , echinoids , 
and annelids ; three orders and one family incertae
NOTE : Lynn and Strüder-Kypke (2005) have 
demonstrated that this subclass is basically mono-
phyletic using small subunit rRNA (SSrRNA) gene 
sequences. However, the familial assignments of 
genera proposed in the classification below, which 
is based primarily on oral features, both morpho-
static and stomatogenetic, receives little support 
from SSrRNA gene sequences (Lynn & Strüder-
Kypke, 2005; Shang, Song, & Warren, 2003). 
We are currently maintaining this morphological 
classification for the purposes of stability, and until 
there are more gene sequence data on this subclass, 
both from more genes and a wider diversity of gen-
era and species. 
 Order Philasterida Small, 1967 
 (syns. Cinetochilina p.p ., Deuterostomatina 
p.p ., Loxocephalida p.p ., Loxocephalina

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