Cap 17
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Cap 17


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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 \u201c riboclasses \u201d within the phylum as 
representatives from the Orders Armophorida 
and Clevelandellida are strongly associated based 
on small subunit rRNA gene sequences (Affa\u2019a, 
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, 
2007).
 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\u20135 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. 
 \u2013 Caenomorpha Perty, 1852 
 \u2013 Cirranter Jankowski, 1964 
 \u2013 Ludio Penard, 1922 
 \u2013 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 
genera. 
 \u2013 Bothrostoma Stokes, 1887 
 \u2013 Brachonella Jankowski, 1964 
 \u2013 Eometopus Small & Lynn, 1985 
 \u2013 Metopus Claparède & Lachmann, 1858 
 \u2013 Palmarella Jankowski, 1975 
 \u2013 Parametopidium Aescht, 2001 
 \u2013 Spirorhynchus da Cunha, 1915 (subj. syn. Metopus ) 
 \u2013 Tesnospira Jankowski, 1964 (subj. syn. Metopus ) 
 \u2013 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 
genera. 
 \u2013 Clevelandella Kidder, 1938 
 \u2013 Metaclevelandella Uttangi & Desai, 1963 
 \u2013 Paraclevelandia Kidder, 1937 
 Family INFEROSTOMATIDAE Ky, 1971 
 (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. 
 \u2013 Ichthyonyctus Jankowski, 1974 
 \u2013 Inferostoma Ky, 1971 
 \u2013 Nathella Singh, 1953 [nomen nudum] 
 Family NEONYCTOTHERIDAE Affa\u2019a, 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. 
 \u2013 Neonyctotherus Affa\u2019a, 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 \u201csucker\u201d 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. 
 \u2013 Cameronyctus Jankowski, 1986 [nomen nudum] 
 \u2013 Cichlidotherus Affa\u2019a, 1989 
 \u2013 Cryptonyctus Jankowski, 1978 
 \u2013 Falconyctus Jankowski, 1978 
 \u2013 Indonyctus Jankowski, 1978 
 \u2013 Metanyctotherus Albaret, 1970 
 \u2013 Micronyctus Jankowski, 1978 
 \u2013 Nyctositum Affa\u2019a, 1979 
 \u2013 Nyctotheroides Grassé,