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tracts of diverse hosts, such 
as echinoids, arthropods, and some vertebrates; 
two orders. 
NOTE : Lynn (2004) established this as one 
of the two “ riboclasses ” within the phylum as 
representatives from the Orders Armophorida 
and Clevelandellida are strongly associated based 
on small subunit rRNA gene sequences (Affa’a, 
Hickey, Strüder-Kypke, & Lynn, 2004; van Hoek, 
van Alen, Sprakel, Hackstein, & Vogels, 1998). 
The odontostomatids , which have been historically 
364 17. The Ciliate Taxa Including Families and Genera
associated with the armophorids (see Jankowski, 
1964) are now tentatively removed to the Class 
 PLAGIOPYLEA (Stoeck, Foissner, & Lynn, 
 Order Armophorida Jankowksi, 1964 
 (syn. Metopina ) 
 Size, generally small to medium; shape, top-
like, usually twisted to left, often much so; free-
 swimming; somatic ciliature, holotrichous, but 
may be absent except for caudal tuft and several 
anteriorly located cirri; oral region spiralled, 
with series of 3–5 perioral or perizonal somatic 
kineties along its anterior edge ; oral polykinetids 
as paramembranelles, extending into an oral cav-
ity with cytostome near the antapical pole in some 
forms; paroral as diplostichomonad; typically in 
marine and freshwater benthic anaerobic habitats 
(i.e., sapropel), but some are endosymbionts of 
 echinoids ; two families. 
 Family CAENOMORPHIDAE Poche, 1913 
 (syns. Gyrocoridae , Gyrocorycidae , Gyrocoryth-
idae , Ludiidae , Ludioidae ) 
 Size, small; shape, round or conical, rigid, twisted left 
less prominently than Metopidae (see below); free-
swimming; somatic cilia as small kineties or cirrus-
like tufts ; oral polykinetids, several, in a small 
oral cavity in the posterior half of the cell; paroral, 
not described; macronucleus, globular to ellipsoid; 
micronucleus, present; contractile vacuole, present; 
cytoproct (?); feeding on bacteria; in brackish and 
freshwater anaerobic habitats; four genera. 
 – Caenomorpha Perty, 1852 
 – Cirranter Jankowski, 1964 
 – Ludio Penard, 1922 
 – Sulfonecta Jankowski, 1978 
 Family METOPIDAE Kahl, 1927 
 Size, small to medium; shape, contorted with 
anterior part of body uniquely twisted to left, 
and posterior part sometimes tailed and/or 
bearing a tuft of longer caudal cilia ; free-
swimming; somatic ciliation, holotrichous; oral 
polykinetids, multiple, sometimes extending out 
of a more posterior oral cavity onto a broader 
peristomial region; macronucleus, globular to 
ellipsoid; micronucleus, present; contractile 
vacuole, present; cytoproct (?); feeding on bacteria;
in marine and freshwater anaerobic habitats; nine 
 – Bothrostoma Stokes, 1887 
 – Brachonella Jankowski, 1964 
 – Eometopus Small & Lynn, 1985 
 – Metopus Claparède & Lachmann, 1858 
 – Palmarella Jankowski, 1975 
 – Parametopidium Aescht, 2001 
 – Spirorhynchus da Cunha, 1915 (subj. syn. Metopus ) 
 – Tesnospira Jankowski, 1964 (subj. syn. Metopus ) 
 – Tropidoatractus Levander, 1894 
 Order Clevelandellida de Puytorac & Grain, 1976 
 (syns. Clevelandellidia , Nyctotherina p.p ., Paranyc-
totherina p.p .) 
 Size, medium to large, often > 150 µm; shape, 
typically flattened; free-swimming; somatic cili-
ature, holotrichous, dense, with somatic kineties 
forming a variety of sutures or complex secant 
systems, which are used, in part, to distinguish 
families and genera; somatic dikinetids with 
non-microtubular retrodesmal and cathetodes-
mal fibrils ; sometimes conspicuous dorsoanterior 
sucker region; oral structures as many left serial 
oral polykinetids or heteromembranelles, not 
usually conspicuous, arranged in a long peris-
tomial groove that precedes a well- developed 
infundibulum ; paroral as diplostichomonad; 
macronucleus anchored in a karyophore in many 
species; conjugation often synchronized with 
reproductive life cycle of the host; contractile 
vacuole, present; cytoproct in several forms lined 
with cilia; feeding on bacteria and organic detritus; 
in marine, freshwater, and terrestrial habitats as 
endocommensals in the digestive tracts of oligo-
chaetes , insects , centipedes , millipedes , molluscs , 
and some vertebrates; five families. 
 Family CLEVELANDELLIDAE Kidder, 1938 
 (for Clevelandiidae ) 
 Size, small to medium; shape, basically ovoid 
or elongate-ovoid, flattened, with posterior 
pole as oddly shaped projection, which bears 
the inconspicuous opening of the oral cavity 
17.3 The Ciliate Taxa to Genus 365
and its infundibular opening ; free-swimming; 
somatic ciliation, holotrichous, often with very 
developed preoral secant system; oral cavity, 
extending from the posterior opening anteriorly 
into the body, with oral polykinetids and paroral 
and endoral; macronucleus, globular to ellipsoid, 
supported by karyophore; micronucleus, present; 
contractile vacuole, may be present; cytoproct 
(?); feeding on bacteria and organic detritus; 
in terrestrial habitats in the digestive tracts of 
 termites and wood-feeding roaches only; three 
 – Clevelandella Kidder, 1938 
 – Metaclevelandella Uttangi & Desai, 1963 
 – Paraclevelandia Kidder, 1937 
 (syns. Nathellidae , Nathelliidae ) 
 Size, small to medium; shape, roughly ovoid, 
flattened, but distorted somewhat by huge sucker 
on right side at anterior end ; free-swimming; 
somatic ciliation, holotrichous, with one right 
caudal secant system, one left caudal secant 
system, and one right transverse secant system ; 
oral region as extensive peristome bearing oral 
polykinetids that extend anteriorly out onto the 
body surface from the infundibular opening at 
the truncate posterior pole ; macronucleus, ellip-
soid, may be supported by a karyophore; micro-
nucleus, present; contractile vacuole (?); cytoproct 
(?); feeding (?); in freshwater habitats in the intes-
tine of certain fishes ; three genera. 
 – Ichthyonyctus Jankowski, 1974 
 – Inferostoma Ky, 1971 
 – Nathella Singh, 1953 [nomen nudum] 
 Family NEONYCTOTHERIDAE Affa’a, 1987 
 Size, small to medium; shape, ovoid, flattened; 
polysaccharide elements forming a reticulated 
subpellicular system under the entire cortex ; 
free-swimming; somatic ciliation, holotrichous, 
with one preoral secant system and one apical 
right secant system ; oral cavity, inconspicuous, 
but with oral polykinetids and paroral and endoral; 
macronucleus, globular to ellipsoid; micronucleus 
(?); contractile vacuole, present; cytoproct, quite 
long, opening near posterior end; feeding (?); in 
freshwater and terrestrial habitats in the digestive 
tract of amphibians; one genus. 
 – Neonyctotherus Affa’a, 1983 
 Family NYCTOTHERIDAE Amaro, 1972 
 (syn. Paranyctotherida p.p .) 
 Size, small to large; shape, ovoid to slightly 
reniform, plump; free-swimming; somatic cilia-
tion, holotrichous, with secant systems, varying 
significantly with included genera, but never a 
transverse secant system; no skeletal appara-
tus beneath concave surface and “sucker” not 
obvious ; oral ciliature running from near-apical to 
sub-equatorial position, in a sigmoid-like curve as 
it enters conspicuous infundibulum; macronucleus, 
ellipsoid, large, compact, in anterior half of body, 
supported by more or less well-developed karyo-
phore; micronucleus, present; contractile vacuole 
(?); cytoproct (?); feeding (?); in marine, freshwa-
ter, and terrestrial habitats as endocommensals in 
wide variety of hosts from oligochaetes , insects 
( cockroach ), and myriapods ( centipede , millipede ) 
to molluscs ( shipworm ), fish , amphibians ( frog , 
 toad ), and reptiles ; 15 genera. 
 – Cameronyctus Jankowski, 1986 [nomen nudum] 
 – Cichlidotherus Affa’a, 1989 
 – Cryptonyctus Jankowski, 1978 
 – Falconyctus Jankowski, 1978 
 – Indonyctus Jankowski, 1978 
 – Metanyctotherus Albaret, 1970 
 – Micronyctus Jankowski, 1978 
 – Nyctositum Affa’a, 1979 
 – Nyctotheroides Grassé,