Cap 17
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Cap 17


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round or elliptical; macronucleus, globular to elon-
gate ellipsoid; micronucleus, present; contractile 
vacuole (?); cytoproct (?); feeding on various heter-
otrophic protists; in freshwater and terrestrial habi-
tats; three genera and one genus incertae sedis . 
 \u2013 Ovalorhabdos Foissner, 1984 
 \u2013 Paraenchelys Foissner, 1983 
 \u2013 Pseudoholophrya Berger, Foissner, & Adam, 1984 
Incertae sedis in Family Pseudoholophryidae 
 \u2013 Songophrya Foissner, 2003 *
 Family PSEUDOTRACHELOCERCIDAE Song, 
1990 
 Size, medium; shape, cylindrical or bottle-
shaped; free-swimming; somatic ciliation, hol-
otrichous, having bipolar kineties with anterior 
kinetids more densely packed; kinetids of brosse 
kinety irregularly arranged and continuous 
with only one somatic kinety ; extrusomes as oral 
toxicysts; oral region, apical; macronucleus, elon-
gate band; micronucleus (?); contractile vacuole, 
present; cytoproct (?); feeding possibly on bacteria; 
in marine habitats; one genus. 
 \u2013 Pseudotrachelocerca Song, 1990 
 Family SPATHIDIIDAE Kahl in Doflein & 
Reichenow, 1929 
 (syns. Arcuospathidiidae , Bryophyllidae , 
 Myriokaryonidae , Paraspathi diidae , Perispiri-
dae , Protospathidiidae , Spathidiina p.p ., Teutho-
phryidae ) 
370 17. The Ciliate Taxa Including Families and Genera
 Size, small to very large; shape, ovoid to elon-
gate, often flask- or sack-shaped, flattened, with 
obliquely truncate anterior end; free-swimming; 
somatic ciliation, holotrichous; brosse kineties, 
2\u20134; extrusomes as somatic mucocysts and sev-
eral types of oral toxicysts; oral region, anterior, 
flattened, usually elongate dorsoventrally (except 
Protospathidium ) with slit-like cytostome, gener-
ally located apically on non-ciliated ridge of body, 
facilitating ingestion of large prey; circumoral 
dikinetids as proliferated anterior fragments of 
somatic kineties, which may exceed the number 
of somatic kineties, and which may remain as 
separated groups after stomatogenesis ; macro-
nucleus, extremely variable, from single ellipsoid, 
to multiple, to ribbon-like and moniliform; micro-
nucleus, present; contractile vacuole, present; cyto-
proct (?); feeding on flagellates and other ciliates; 
in marine, freshwater, and terrestrial habitats; 29 
genera and three genera incertae sedis . 
NOTE : Foissner, Berger, and Schaumburg 
(1999) redescribed Teuthophrys and suggested 
that it was related to the spathidiids based on its 
infraciliary pattern and extrusomes, and this was 
confirmed by Strüder-Kypke et al. (2006) using 
its small subunit rRNA gene sequence. Foissner 
(2003) established the Family Myriokaryonidae 
to include the genera Myriokaryon , Bergophrya , 
Cephalospatula , and Kahlophrya , but admitted 
that this family was difficult to separate from the 
Family Spathidiidae . Until gene sequence data 
demonstrate its monophyly, we have retained these 
genera in the Family Spathidiidae . 
 \u2013 Apobryophyllum Foissner, 1998 
 \u2013 Arcuospathidium Foissner, 1984 
 \u2013 Armatospathula Foissner & Xu, 2006 *
 \u2013 Bergophrya Foissner, 2003 *
 \u2013 Bryophyllum Kahl, 1931 
 \u2013 Cephalospatula Foissner, 2003 *
 \u2013 Cranotheridium Schewiakoff, 1893 
 \u2013 Cultellothrix Foissner, 2003 *
 \u2013 Diceratula Corliss, 1960 (subj. syn. Spathidium ) 
 \u2013 Edaphospathula Foissner & Xu, 2006 *
 \u2013 Epispathidium Foissner, 1984 
 \u2013 Kahlophrya Foissner, 2003 *
 \u2013 Lacerus Jankowski, 1967 
 \u2013 Latispathidium Foissner, Berger, & Zechmeister-
Boltenstern, 2005 
 \u2013 Micromidas Delphy, 1938 
 \u2013 Myriokaryon Jankowski, 1973 
 \u2013 Neobryophyllum Foissner in Foissner & Lei, 
2004*
 \u2013 Paraspathidium Noland, 1937 
 \u2013 Penardiella Kahl, 1930 
 \u2013 Perispira Stein, 1859 
 \u2013 Protospathidium Dragesco & Dragesco-Kernéis 
in Foissner, 1984 
 \u2013 Semispathidium Foissner, Agatha, & Berger, 
2002*
 \u2013 Spathidiodes Kahl, 1926 
 \u2013 Spathidioides Brodsky, 1925 
 \u2013 Spathidiosus Gajewskaja, 1933 
 \u2013 Spathidium Dujardin, 1841 
 \u2013 Supraspathidium Foissner & Didier, 1982 
 \u2013 Teuthophrys Chatton & de Beauchamp, 1923 
 \u2013 Thysanomorpha Jankowski, 1967 
Incertae sedis in Family Spathidiidae
 \u2013 Apospathidium Foissner, Agatha, & Berger, 
2002*
 \u2013 Enchelaria Foissner, Agatha & Berger, 2002 *
 \u2013 Proboscidium Meunier, 1910 
 Family TRACHELIIDAE Ehrenberg, 1838 
 (syns. Branchioecetidae , Dileptidae , Dileptina ) 
 Size, medium to large; shape, flask-shaped 
with dorsal proboscis of varying relative 
length ; free-swimming; somatic ciliation, holot-
richous; extrusomes as somatic mucocysts and 
several types of oral toxicysts; brosse kineties, 
three or more; oral region, circular or elliptical, 
possibly with permanent cytostome, distant 
from extreme anterior end of body at base of 
proboscis, but with oral kinetids extending 
along the borders of the ventral surface of the 
proboscis and with toxicysts in this ventral 
band or distributed around the cytostome ; oral 
nematodesmata, lengthy, prominent, supporting 
the cytopharynx, typically in two rings, the outer 
one associated with the circumoral dikinetids; 
macronucleus, very variable in shape, from single 
globular to band-shaped to multiple globular and 
even moniliform; micronucleus, present, may 
be multiple; contractile vacuole, at least present 
in freshwater forms; cytoproct (?); feeding on 
flagellates, microalgae, and other ciliates; in 
marine, freshwater, and terrestrial habitats, with 
some forms planktonic; ten genera and one genus 
incertae sedis . 
17.3 The Ciliate Taxa to Genus 371
 \u2013 Branchioecetes Kahl, 1931 
 \u2013 Dileptus Dujardin, 1841 
 \u2013 Dimacrocaryon Jankowski, 1967 
 \u2013 Micruncus Delphy, 1938 
 \u2013 Monilicaryon Jankowski, 1967 
 \u2013 Paradileptus Wenrich, 1929 
 \u2013 Pelagodileptus Foissner, Berger, & Schaumberg, 
1999
 \u2013 Pseudomonilicaryon Foissner, 1997 
 \u2013 Rimaleptus Foissner, 1984 
 \u2013 Trachelius Schrank, 1803 
Incertae sedis in Family Tracheliidae 
 \u2013 Ctenoctophrys Weill, 1946 [not listed in Aescht] 
 Family TRACHELOPHYLLIDAE Kent, 1882 
 (syns. Enchelyodontidae p.p ., Lagynophryidae , 
 Trachelophyllina p.p .) 
 Size, small to medium, rarely large; shape, 
long-ovoid or flask-shaped, slightly flattened; free-
swimming; somatic ciliation, holotrichous; brosse 
kineties, two to many in Acaryophrya ; extrusomes 
as somatic mucocysts and lepidosomes (e.g., in 
Lepidotrachelophyllum and Spetazoon ) and some-
times several types of oral toxicysts; oral region 
circular to elliptical, sometimes forming an obvi-
ously pointed dome; circumoral dikinetids typi-
cally at anterior end of bipolar somatic kineties, 
not exceeding the number of somatic kineties 
(except some Enchelyodon species) ; macronu-
cleus, quite variable, from single ellipsoid to paired 
ellipsoid to band-form or multiple and moniliform; 
micronucleus, present; contractile vacuole, present; 
cytoproct, present; feeding on flagellates and other 
ciliates, even rotifers in large forms; in marine, 
freshwater, and terrestrial habitats; 12 genera. 
NOTE : The synapomorphy for this family is not 
a strong one, but subdivision on characters such as 
the lepidosomes (see Foissner et al., 2002) seems 
premature at this time. 
 \u2013 Acaryophrya André, 1915 
 \u2013 Bilamellophrya Foissner, Agatha, & Berger, 
2002*
 \u2013 Enchelyodon Claparède & Lachmann, 1859 
 \u2013 Enchelyotricha Foissner, 1987 
 \u2013 Epitholiolus Foissner, Agatha, & Berger, 2002 *
 \u2013 Foissnerides Song & Wilbert, 1989 
 \u2013 Lagynophrya Kahl, 1927 
 \u2013 Lepidotrachelophyllum Nicholls & Lynn, 1984 
 \u2013 Luporinophrys Foissner, 2005 *
 \u2013 Sleighophrys Foissner, 2005 *
 \u2013 Spetazoon Foissner, 1994 
 \u2013 Trachelophyllum Claparède & Lachmann, 1859 
 Order Pleurostomatida Schewiakoff, 1896 
 (syns. Amphileptida p.p ., Amphileptina p.p ., 
 Litonotina p.p ., Pleurostomata , Pleurostom[at]ina , 
 Scaphotrichina , Thysanophorina ) 
 Size, medium to large; shape, leaf-like or later-
ally compressed, sometimes with lengthy, attenu-
ated