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of dikinetids on the right (= ventral?) 
surface and a single left lateral kinety border-
ing the left (= dorsal?) surface; extrusomes, 
not reported; oral region, circular to ovoid, 
ringed by inner and outer dikinetid files with 
17.3 The Ciliate Taxa to Genus 343
an intrabuccal kinety, which may be short or 
long ; nuclei in clusters, typically two macronu-
clei and one micronucleus; contractile vacuole, 
absent; cytoproct (?); feeding on bacteria and 
smaller protists; in marine interstitial habitats; 
two genera. 
 – Apocryptopharynx Foissner, 1996 
 – Cryptopharynx Kahl, 1928 
 Family LOXODIDAE Bütschli, 1889 
 (syns. Ciliofaureidae , Drepanostom(at)idea ) 
 Size, medium to very large; shape, long, flat, 
with beak-like anterior rostrum, which inter-
rupts perioral kineties at the anterior end ; free-
swimming; somatic ciliature as files of dikinetids 
on the right (= ventral?) surface and a single left 
lateral kinety bordering the left (= dorsal?) surface; 
extrusomes as somatic cnidocyst-like organelles 
in Remanella ; Müllerian vesicles in the endo-
plasm, containing strontium ( Remanella ) and 
barium ( Loxodes ) salts ; oral area in long ven-
tral groove behind rostrum, with inner and 
outer dikinetid files and a long, rectilinear 
intrabuccal kinety ; nuclei in clusters with one or 
two macronuclei and a single micronucleus; con-
tractile vacuole, in freshwater species; cytoproct 
(?); feeding on bacteria and other protists, such as 
microalgae; in freshwater ( Loxodes ) and marine 
(Remanella ) habitats, typically in sediments but 
ranging into the water column when oxygen levels 
decline; two genera. 
 – Loxodes Ehrenberg, 1830 
 – Remanella Foissner, 1996 
 Order Protoheterotrichida Nouzarède, 1977 
 Size, large; shape, elongate, highly contractile; 
often pigmented, appearing black in transmitted 
microscopic light; free-swimming; somatic cili-
ation, holotrichous; oral region, ventral, with 
an elaborate oral ciliature including a series 
of transverse rows of monokinetids, conspicu-
ous either on the left or the right side of the 
oral region, depending upon the genus ; sto-
matogenesis, not described; nuclei in clusters, 
typically two macronuclei and one micronu-
cleus; contractile vacuole, absent; cytoproct (?); 
feeding on microalgae and other protists; in 
marine or brackish coastal sands and gravels; 
one family. 
 Family GELEIIDAE Kahl, 1933 
 (syn. Aveliidae ) 
 With characteristics of the order; four genera. 
 – Avelia Nouzarède, 1977 
 – Geleia Kahl in Foissner, 1998 
 – Gellertia Dragesco, 1999 
 – Parduczia Dragesco, 1999 
Incertae sedis in Class KARYORELICTEA 
 – Ciliofaurea Dragesco, 1960 
 – Corlissia Dragesco, 1954 
 Class HETEROTRICHEA Stein, 1859 
 (syns. Heterotricha , Heterotrichida , Hetero-
trichorina , Membranellata p.p ., Membranellophora 
p.p ., Spirotricha p.p , Spirotrichophora p.p.)
 Size, medium to large; shape, variable, from 
compressed to conical, often elongate and con-
tractile; free-swimming, but some species are 
temporarily or permanently sessile; some species 
are pigmented, often brightly, with pigment in 
specialized pigmentocysts; somatic ciliation, hol-
otrichous; somatic kinetids as dikinetids with 
postciliodesmata whose major microtubular 
ribbons are typically separated by 1 + 2 micro-
tubules ; extrusomes, sometimes as mucocysts, but 
pigmentocysts can also be secreted; left serial oral 
polykinetids conspi cuous, typically paramem-
branelles, encircling the anterior end clockwise 
before plunging into the oral cavity ; one or 
more “parorals” on the right side of the oral cav-
ity; stomatogenesis, parakinetal; macronucleus, 
highly polyploid, divided by extra-macronu-
clear microtubules ; micronuclei, typically multi-
ple; conjugation, temporary; contractile vacuole, 
often conspicuous, with long collecting canals; 
cytoproct, present; distributed widely in marine, 
freshwater, and terrestrial habitats; one order. 
 Order Heterotrichida Stein, 1859 
 (syns. Blepharismina p.p ., Coliphorida p.p ., Colipho-
rina p.p ., Condylostomatina p.p ., Peritromida 
p.p ., Stentorina p.p .) 
 With characteristics of the class; nine families. 
344 17. The Ciliate Taxa Including Families and Genera
 Family BLEPHARISMIDAE Jankowski in Small 
& Lynn, 1985 
 Size, medium to large; shape, pyriform or ellipsoid, 
somewhat narrowed anteriorly, laterally compressed; 
free-swimming; pigmentocysts, common, filled with 
the pigment blepharismin; somatic ciliation, holot-
richous; extrusomes as sacculate mucocysts; oral 
polykinetids extending along the left margin of the 
oral region and circling counter-clockwise into a 
shallow posterior oral cavity; paroral dikinetids 
anterior of cytostome (precytostomal) ; macro-
nucleus, ovoid to sometimes nodular; micronucleus, 
may be multiple; contractile vacuole, present; cyto-
proct, present; feeding on bacteria, microalgae, and 
other protists, including ciliates, with some species 
becoming cannibalistic; in marine, freshwater, and 
terrestrial habitats; four genera. 
 – Anigsteinia Isquith, 1968 
 – Blepharisma Perty, 1849 
 – Parablepharisma Kahl, 1932 [nomen nudum] 
 – Pseudoblepharisma Kahl, 1927 
 Family CHATTONIDIIDAE Villeneuve-Brachon, 
1940
 Size, large; shape, somewhat rotund, contractile, 
with gently pointed posterior end; free-swimming; 
somatic ciliation, holotrichous; a unique somatic 
“posteroaxial cavity,” containing 6–7 ciliary 
organelles, opening at posterior pole ; oral cavity 
opening apically and bearing full circle of strong 
oral polykinetids; macronucleus, very long, with 
loops; micronucleus, present; contractile vacuole 
(?); cytoproct (?); feeding on bacteria, microalgae, 
and other protists, including ciliates; in hypereu-
trophic brackish waters; one genus. 
 – Chattonidium Villeneuve, 1937 
 Family CLIMACOSTOMIDAE Repak, 1972 
 (syn. Pediostomatidae ) 
 Size, medium to large; shape, ovoid, often anteri-
orly pointed; free-swimming; somatic ciliation, holot-
richous; extrusomes as sacculate somatic mucocysts; 
oral region anterior, with very prominent, ciliated 
peristomial field occupying much of the anterior 
part of the body, bordered at least on the posterior 
and right margins by the serial adoral polyki-
netids ; paroral ciliature, inconspicuous; macronu-
cleus, usually as a thick ribbon, often coiled and 
lengthy; micronucleus, present; contractile vacuole, 
at least in freshwater forms; cytoproct (?); symbiotic 
zoochlorellae in some species; feeding on bacteria, 
microalgae, and other protists, including ciliates; in 
marine and freshwater habitats; three genera. 
 – Climacostomum Stein, 1859 
 – Fabrea Henneguy, 1890 
 – Pediostomum Kahl, 1932 
 Family CONDYLOSTOMATIDAE Kahl in Doflein 
& Reichenow, 1929 
 (syns. Condylostomidae , Condylostentoridae ) 
 Size, medium to large; shape, very elongate in some 
forms, nearly ellipsoidal in others; free-swimming; 
somatic ciliation, holotrichous; extrusomes as saccu-
late somatic mucocysts in some species; oral region 
expansive; oral ciliature, including adoral polyki-
netids and a prominent paroral membrane ; per-
istomial field absent ; macronucleus, typically long 
and moniliform; micronucleus, typically multiple; 
contractile vacuole often with long collecting canal; 
cytoproct, present; feeding on bacteria, microalgae, 
and other protists; in marine, freshwater, and terrestrial 
habitats; seven genera and one genus incertae sedis . 
 – Condylostoma Bory de St. Vincent, 1824 
 – Condylostomides Da Silva Neto, 1994 
 – Copemetopus Villeneuve-Brachon, 1940 
 – Electostoma Jankowski, 1979 
 – Linostomella Aescht in Foissner, Berger, & 
Schaumberg, 1999 
 – Predurostyla Jankowski, 1978 
 – Procondylostoma Jankowski, 1979 
Incertae sedis in Family Condylostomatidae 
 – Dellochus Corliss, 1960 
 Family FOLLICULINIDAE Dons, 1914 
 (syn. Coliphorida , Coliphorina )

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