Cap 17

yidae, Phtorophryida , Phtorophryinae , Polyspiri-
dae , Spirophryinae , Synophryinae , Terebrospirinae , 
and possibly Kofoidellidae and Perezellidae ) 
 Size, small in tomite stage, to large in feeding tro-
phont; shape, ovoid to spherical; free-swimming;
somatic kineties, right-spiralled, ranging from 
about 9 to < 22 ; oral apparatus, as for subclass 
and with rosette ; macronucleus, globular to 
extremely elongate and ribbon-like, depending 
upon the life cycle stage; micronucleus, present; 
contractile vacuole, present; exuviotrophic, 
but one genus ( Phtorophrya ) hyperparasite on 
Gymnodinioides species and another ( Synophrya ) 
with sanguicolous stage; in marine, rarely brack-
ish and freshwater (?), habitats as symbionts 
(parasites?) on marine crustacea , such as hermit 
crabs , shrimp , and copepods ; 17 genera and three 
genera incertae sedis . 
 – Calospira Chatton & Lwoff, 1935 
 – Foettingeria Caullery & Mesnil, 1903 
 – Gymnodinioides Minkiewicz, 1912 
 – Hyalophysa Bradbury, 1966 
 – Hyalospira Miyashita, 1933 (subj. syn. Gymno-
dinioides ) 
 – Metaphrya Ikeda, 1917 
 – Ophiuraespira Chatton & Lwoff, 1930 
 – Pericaryon Chatton, 1911 
 – Phoretophrya Chatton, A. & Μ. Lwoff, 1930 
 – Phtorophrya Chatton, A. & Μ. Lwoff, 1930 
 – Polyspira Minkiewicz, 1912 
 – Rosea de Puytorac, 1994 [nomen nudum] 
428 17. The Ciliate Taxa Including Families and Genera
 – Spirophrya Chatton & Lwoff, 1924 
 – Synophrya Chatton & Lwoff, 1926 
 – Terebrospira Debaisieux, 1960 
 – Traumatiophtora Chatton & Lwoff, 1931 
 – Vampyrophrya Chatton & Lwoff, 1931 
Incertae sedis in Family Foettingeriidae 
 – Jeppsia Corliss, 1960 
 – Kofoidella Cépède, 1910 
 – Perezella Cépède, 1910 
 Order Astomatophorida Jankowski, 1966 
 (syns. Astomophorina , Nephrocolida , Neph-
rocolina ) 
 Size, small to medium; shape of trophont, long, 
vermiform; free-swimming, but trophont attached 
by its anterior end to host tissue; somatic ciliation, 
holotrichous, kineties much spiralled and somatic 
ciliature markedly thigmotactic; no cytostome 
(in stages of life cycles known to date), but 
remnants of oral ciliature; fission of tomont-
trophont by sequential formation of tomites 
(catenulation) or by multiple transverse fission 
with tomites remaining connected ; macronu-
cleus, very variable, from fragmented nodules to 
an irregular network; micronucleus, present but 
obscure; contractile vacuole (?); in marine habi-
tats as parasites, for example, in the internal 
organs, such as liver, kidney, and gonad of
cephalopods ; one family. 
 Family OPALINOPSIDAE Hartog, 1906 
 (syns. Chromidinida , Chromidinidae ) 
 With characteristics of the order; two genera. 
 – Chromidina Gonder, 1905 
 – Opalinopsis Foettinger, 1881 
 Order Pilisuctorida Jankowski, 1966 
 Size, small to large; shape, ovoid to elongate; free-
swimming but attached to host in the feeding state; 
body with ventral adhesive organelle ; species
of most genera permanently in so-called “neo-
tenic” tomite stage; somatic kineties of tomite 
arched, following rim of flattened ventral sur-
face ; mature trophonts (e.g., Conidophrys ), non-
ciliated, immobile, characteristically attached to 
seta or cuticle of host, and a migrating tomite, 
which is ciliated but lacks a cytostome; tomites 
produced by synchrony or strobilation ; macro-
nucleus, elongate ellipsoid, irregularly shaped to 
band-form and ribbon-like; micronucleus, present; 
contractile vacuole, present; feeding on tissue flu-
ids (i.e., hemolymph); in marine habitats on the 
cuticular processes of amphipods , isopods , deca-
pods , and cirripeds , and possibly a terrestrial mite ; 
single host life cycle; one family. 
NOTE : De Puytorac (1994h) recognized three 
monotypic families in this order, each includ-
ing one of the three genera here included in the 
single family of this monotypic order. Perhaps 
gene sequence data will confirm that the genetic 
diversity of these forms warrants this higher order 
taxonomy. 
 Family CONIDOPHRYIDAE Kirby, 1941 
 (for Pilisuctoridae ; syns. Ascophryidae , Askoellida , 
 Askoellidae , Conidophryidae ) 
 With characteristics of the order; three genera. 
 – Ascophrys Campillo & Deroux, 1974 
 – Askoella Fenchel, 1965 
 – Conidophrys Chatton & Lwoff, 1934 
 Subclass Peritrichia Stein, 1859 
 (syns. Cyclohymenophora , Dexiotricha , Peritri-
chasina , Peritrichidea , Peritrichorida , Stomatoda ) 
 Size of zooids, small to medium, rarely large, 
but colonial forms can be macroscopic; shape, 
characteristically inverted bell- or goblet-shaped 
or conical-cylindrical; sessile and sedentary except 
as dispersive telotrochs or swarmers, although sev-
eral taxa are always free-swimming; prominent 
holdfast derivatives or scopula, which secretes 
the stalk of sessile species and includes a field 
of thigmotactic cilia, at aboral pole ; alveoli, 
well-developed, with pellicle perforated by pores 
(= parasomal sacs?); somatic ciliature, reduced 
to subequatorial locomotor fringe, the trochal 
band, which is permanently ciliated on mobile 
species and temporarily ciliated on the dis-
persal stage or telotroch of sessile species ; oral 
region, often retractable, encircling apical pole as 
peristome, bordered by a more or less prominent 
collarette; oral ciliature, conspicuous, winding 
counterclockwise around the border of the 
prominent peristome, from its outer terminus 
17.3 The Ciliate Taxa to Genus 429
on an elevated central, often extensible episto-
mial disk, with peristomial part of the paroral 
(haplokinety, stichodyad) on the outside and 
oral polykinetid (polykinety) 1 on the inside, but 
both actually originating in an oral cavity, the 
infundibulum, at the terminus of which is the 
cytostome ; oral ciliature of infundibulum includ-
ing inner parts of paroral and oral polykinetid 1, 
which is accompanied at its inner terminus by 
infundibular polykinetids 2 and 3 (formerly called 
peniculi); stomatogenesis, ophryobuccokinetal, 
with involvement of germinal field or row of 
kinetosomes ; fission, with its plane parallel to 
the major body axis, suggesting that the apical-
 antapical axis is a secondary adaptation to sessility; 
conjugation, invariably total, typically involving 
fusion of a migratory microconjugant with a 
stationary macroconjugant ; contractile vacuole 
and cytoproct, emptying into infundibulum ; 
bacterivorous and microphagous, with symbiotic 
species ingesting detritus and tissue debris of 
host; in marine, freshwater, and rarely terrestrial 
habitats, very widespread with species generally 
free-living but many occurring as commensals or 
even parasites on or in diverse hosts, ranging from 
protozoa to vertebrates; two orders. 
NOTE : Gong et al. (2006) demonstrated that 
the mobilid peritrichs are a separate lineage from 
the sessilid peritrichs based on small subunit rRNA 
gene sequences, but the two lineages are apparently 
not sister clades. We remain conservative in our 
assignment of both sessilids and mobilids to the 
Subclass Peritrichia until sampling of additional 
genes and taxa refutes this assignment. 
 Order Sessilida Kahl, 1933 
 (syns. Aloricata + Loricata , Astylozo(on)ina p.p ., 
 Cothurniina p.p ., Epistylina p.p ., Fibrodiscida p.p ., 
 Lagenophryina p.p ., Loricina p.p ., Natantina p.p ., 
 Operculariina p.p ., Opisthonectina p.p ., Ophrydiina 
p.p ., Rovinjellina p.p ., Scyphidiina p.p ., Sedentaria , 
 Sessilia , Sessili[i]da , Stylophorina , Syncyathellina 
p.p ., Thigmodiscina , Vorticellina p.p .) 
 Size of zooids, small to medium, rarely large; 
shape, inverted bell- or goblet-shaped or coni-
cal-cylindrical; zooids, dimorphic, with mature 
zooids or trophonts, sedentary or sessile, com-
monly stalked or with inconspicuous adhesive 
disc, attached to substrate by scopula, but a 
few species presumed to be secondarily mobile, 
and dispersal