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Size, medium to large; shape, typically elongate, 
contractile; often pigmented; mature forms, sessile 
and sedentary, always residing in a lorica; body, 
especially in neck region in some species, with 
conspicuous pair of “peristomial wings” bearing 
the prominent oral ciliature ; at division, complex 
morphogenesis with vermiform migratory larval 
stage that “recapitulates” a typical heterotrich 
17.3 The Ciliate Taxa to Genus 345
with reduced spiralling of the adoral zone ; macro-
nucleus, variable, as single ellipsoid to multiple and 
moniliform; micronucleus, single to multiple; contrac-
tile vacuole, present at least in freshwater forms; cyto-
proct (?); feeding on bacteria, microalgae, and other 
protists; widely distributed in marine habitats, but a 
few species in freshwater, attached to algae, higher 
aquatic plants, or integument or shells of invertebrates 
(e.g., molluscs, various crustaceans, bryozoa, coelen-
terates); 30 genera including some fossil forms. 
 – Ampullofolliculina Hadzi, 1951 
 – Ascobius Henneguy, 1884 
 – Atriofolliculina Hadzi, 1951 [nomen nudum] 
 – Aulofolliculina Hadzi, 1951 
 – Botticula Dioni, 1972 
 – Claustrofolliculina Hadzi, 1951 
 – Diafolliculina Hadzi, 1951 [nomen nudum] 
 – Echinofolliculina Dons, 1934 
 – Epifolliculina Hadzi, 1951 
 – Eufolliculina Hadzi, 1951 
 – Folliculina Lamarck, 1816 
 – Folliculinopsis Fauré-Fremiet, 1936 [nomen 
nudum]
 – Halofolliculina Hadzi, 1951 [nomen nudum] 
 – Lagotia Wright, 1857 
 – Latifolliculina Hadzi, 1951 
 – Magnifolliculina Uhlig, 1964 [nomen nudum] 
 – Metafolliculina Dons, 1924 
 – Mirofolliculina Dons, 1928 
 – Pachyfolliculina Hadzi, 1951 
 – Parafolliculina Dons, 1914 
 – Pebrilla Giard, 1888 
 – Perifolliculina Hadzi, 1951 
 – Planifolliculina Hadzi, 1951 
 – Platyfolliculina Hadzi, 1938 
 – Priscofolliculina Deflandre & Deunff, 1957 (fossil) 
 – Pseudofolliculina Dons, 1914 
 – Pseudoparafolliculina Andrews & Nelson, 1942 
 – Splitofolliculina Hadzi, 1951 [nomen nudum] 
 – Stentofolliculina Hadzi, 1938 
 – Valletofolliculina Andrews, 1953 
 Family MARISTENTORIDAE Miao, Simpson, 
Fu, & Lobban, 2005 
 Size, large, majestic when fully extended; shape, 
trumpet-shaped with apical area expanded into a 
bilobed “cap” divided by a ventral indentation, 
very contractile ; free-swimming, but typically tem-
porarily attached to substrate; pigmented, due both 
to blood-red pigmentocysts and to the presence of 
symbiotic zooxanthellae; somatic ciliation, holotri-
chous; oral ciliature of over 300 oral polykinetids, 
spirals around flared-out anterior end, encircling 
an anterior peristomial field with scattered cili-
ate dikinetids, not arranged in kineties ; paroral, 
very reduced; macronucleus, ellipsoid; micronuclei, 
multiple; contractile vacuole, absent; cytoproct, not 
reported; feeding on bacteria, microalgae, and other 
protists; in marine habitats, at least associated with 
coral reefs; one genus. 
 – Maristentor Lobban, Schefter, Simpson, Pochon, 
Pawlowski, & Foissner, 2002 *
 Family PERITROMIDAE Stein, 1867 
 Size, medium; shape, ellipsoidal, dorsoventrally 
flattened, contractile; free-swimming; somatic cili-
ation primarily on ventral (= right?) surface; single
somatic kinety on slightly convex dorsal (= left?) 
surface; spine-like cilia of dorsal kinetids emerg-
ing from wart-like papillae ; extrusomes, not 
reported; oral region expansive with left serial 
oral polykinetids extending anteriorly from 
equatorial oral cavity along anterior border 
of cell and paroral extending parallel to serial 
oral polykinetids ; macronucleus, typically in two 
lobes; micronucleus, present; contractile vacuole, 
present; cytoproct, present; feeding on bacteria, 
microalgae, and other protists; generally in marine 
habitats, including salt marshes; one genus. 
 – Peritromus Stein, 1863 
 Family SPIROSTOMIDAE Stein, 1867
(syn. Spirostomatidae ) 
 Size, medium to large; shape, often elongate, cylin-
drical and very contractile; some pigmented forms; 
free-swimming; somatic ciliation, holotrichous; extru-
somes, not reported; oral region in anterior half; 
peristomial field, long, narrow, not ciliated; oral 
ciliature sometimes relatively inconspicuous, but 
still with many serial oral polykinetids, extend-
ing 1/3–1/2 body length ; paroral may extend almost 
entire length of oral region, paralleling the adoral zone; 
macronucleus, moniliform; micronucleus, present; 
contractile vacuole posterior, frequently large, and 
may have lengthy collecting canal; cytoproct, present; 
346 17. The Ciliate Taxa Including Families and Genera
feeding on bacteria, microalgae, and other protists; 
predominantly in freshwater habitats; two genera and 
two genera incertae sedis . 
 – Gruberia Kahl, 1932 
 – Spirostomum Ehrenberg, 1834 
Incertae sedis in Family Spirostomidae 
 – Diplogmus Mansfeld, 1923 
 – Propygocirrus Mansfeld, 1923 
 Family STENTORIDAE Carus, 1863 
 Size, medium to large, becoming majestic in 
size and movement; shape, trumpet-shaped with 
apical area not bilobed, very contractile ; often 
pigmented due to pigmentocysts with the pig-
ment stentorin and/or with symbiotic zoochlo-
rellae; free-swimming, but typically temporarily 
attached to the substrate, with a few species residing 
in mucilaginous loricae; somatic ciliation, holotri-
chous, with posterior end having thigmotactic cilia 
permitting temporary attachment to substrate; oral 
ciliature spirals nearly 360° around flared-out 
anterior end, encircling an anterior peristomial 
field that is covered by ciliated kinetids arranged 
in ordered kineties; paroral accompanying entire 
length of adoral zone of polykinetids ; macro-
nucleus, ellipsoid to ribbon-like and moniliform; 
micronucleus, one to many; contractile vacuole, 
may be multiple; cytoproct, present; feeding on bac-
teria, microalgae, and other protists, including other 
ciliates; typically in freshwater habitats only; one 
genus and three genera incertae sedis . 
 – Stentor Oken, 1815 
Incertae sedis in Family Stentoridae 
 – Heterostentor Song & Wilbert, 2002 *
 – Parastentor Vuxanovici, 1961 
 – Stentoropsis Dogiel & Bychowsky, 1934 
 Subphylum INTRAMACRONUCLEATA Lynn, 1996 
 ( Ciliostomatophora p.p ., Homoiotricha p.p ., Homo-
tricha p.p ., Kinetodesmatophora p.p ., Postcilio-
nematophora p.p ., Tubulicorticata + Filicorticata 
+ Epiplasmata + Membranellophora p.p ., Trans-
versonematophora p.p .) 
 Size, small to large; shape, variable, from globular 
to ellipsoid to elongate; free-swimming or sessile; 
cortical alveolar system typically well-developed; 
somatic ciliation, holotrichous, but forms with girdles 
and strips, and even non-ciliated taxa are known; 
parasomal sacs, present; extrusomes as somatic 
mucocysts and trichocysts and oral and somatic 
toxicysts; oral structures, variable, minimally with 
oral dikinetids, either encircling the cytostome or on 
the right side as a paroral, but some forms also with 
several to many oral polykinetids in an adoral zone 
and other forms astomatous or with oralized somatic 
kinetids; stomatogenesis, variable from telokinetal 
to buccokinetal; fission, typically isotomic, rarely 
anisotomic and multiple; macronuclear genome 
typically differentiated by fragmentation of micro-
nuclear chromosomes during anlage development; 
polyploid macronucleus dividing by intramacro-
nuclear microtubules ; micronucleus, present; con-
jugation, typically temporary and isogamontic, but 
some forms showing complete conjugation with ani-
sogamonty; feeding habits, diverse, including several 
major classes and subclasses as obligate symbionts, 
sometimes parasitic; widely distributed in marine, 
freshwater, and terrestrial habitats; nine classes. 
NOTE : Lynn (1996a) suggested that the rapid 
radiation within this subphylum arose from a 
fundamentally different property of ciliate cellular 
organization, perhaps