Cap 17
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Cap 17


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arc-like file that 
parallels the anterior left serial oral polyki-
netids and a frontoventral cirral zig-zag with 
362 17. The Ciliate Taxa Including Families and Genera
1\u20136 files to the right of left marginal cirral file 
and also with several cirral files to the left of the 
right marginal cirri; multiple \u201cmarginal files\u201d 
derive from unique anlagen during morpho-
genesis ; transverse cirri, present or absent; caudal 
cirri, absent; dorsal somatic ciliature as several 
files of bristle dikinetids; oral ciliature as for order 
with paroral and endoral; macronucleus, ellipsoid, 
multiple; micronucleus, present; contractile vacu-
ole, present; cytoproct, likely present; feeding on 
bacteria, algae, smaller protists, including testate 
amoebae and ciliates; in freshwater and terrestrial 
habitats; three genera. 
 \u2013 Hemicycliostyla Stokes, 1886 
 \u2013 Pseudourostyla Borror, 1972 
 \u2013 Trichotaxis Stokes, 1891 
 Family UROSTYLIDAE Bütschli, 1889 
 (syns. Bakuellidae , Bakuellinae , Erionellidae , 
 Holostichidae , Holostichina , Holostichinae , 
 Holostichoidea , Psammomitrinae Pseudoam-
phisiellidae , Urostylinae , Urostyloidea ) 
 Size, small to large; shape, elongate ovoid with 
some tailed forms; free-swimming; somatic ven-
tral ciliature with several frontal cirri some-
what larger than other frontoventral cirri, and 
with frontoventral cirri as a single zig-zag file 
of paired cirri or a series of shorter files off-
set at their anterior and posterior ends (e.g., 
Bakuella, Eschaneustyla ) and typically not with 
additional \u201cmarginal files\u201d on both sides of this 
zig-zag (cf. Pseudourostylidae) ; transverse cirri, 
may be numerous; caudal cirri, present or absent; 
dorsal somatic ciliature as three to many files of 
bristle dikinetids; oral ciliature as for order with 
paroral and endoral; during division morpho-
genesis, frontoventral cirri differentiate from a 
longitudinal field of more than five oblique cili-
ary streaks; macronucleus, ellipsoid, two to many 
nodules; micronucleus, present; contractile vacu-
ole, present; cytoproct, likely present; feeding on 
bacteria, algae, and smaller protists, including 
ciliates; in marine, freshwater, and terrestrial 
habitats; 24 genera. 
 \u2013 Afrothrix Foissner, 1999 
 \u2013 Anteholosticha Berger, 2003 *
 \u2013 Australothrix Blatterer & Foissner, 1988 
 \u2013 Bakuella Agamaliev & Alekperov, 1976 
 \u2013 Biholosticha Berger, 2003 *
 \u2013 Birojimia Berger & Foissner, 1989 
 \u2013 Caudiholosticha Berger, 2003 *
 \u2013 Diaxonella Jankowski, 1979 
 \u2013 Holosticha Wrzesniowski, 1877 
 \u2013 Holostichides Foissner, 1987 
 \u2013 Metabakuella Alekperov, 1989 
 \u2013 Metaurostylopsis Song, Petz, & Warren, 2001 *
 \u2013 Notocephalus Petz, Song, & Wilbert, 1995 
 \u2013 Parabirojimia Hu, Song, & Warren, 2002 *
 \u2013 Paragastrostyla Hemberger, 1985 
 \u2013 Paramitrella Jankowski, 1978 
 \u2013 Paruroleptus Wenzel, 1953 (subj. syn. Uroleptus ) 
 \u2013 Periholosticha Hemberger, 1985 
 \u2013 Perisincirra Jankowski, 1978 
 \u2013 Psammomitra Borror, 1972 (subj. syn. 
Uroleptus ) 
 \u2013 Pseudoamphisiella Song, 1996 
 \u2013 Pseudobakuella Alekperov, 1992 
 \u2013 Tunicothrix Xu, Lei & Choi, 2006 *
 \u2013 Uroleptus Ehrenberg, 1831 
 \u2013 Urostyla Ehrenberg, 1830 
Incertae sedis in Subclass Stichotrichia 
 \u2013 Erniella Foissner, 1987 
 \u2013 Gastrosticha Kahl, 1932 [nomen dubium] 
 \u2013 Saudithrix Berger, Al-Rasheid, & Foissner, 2006 *
 \u2013 Stenotricha Jankowski, 1978 
 \u2013 Uncinata Bullington, 1940 
 Subclass Oligotrichia Bütschli, 1887/1889 
 (syns. Oligotricha , Oligotrichorida ) 
 Size, small to large, shape, typically rounded 
or gently pointed posteriorly, sometimes tailed; 
free-swimming; a perilemma present in many spe-
cies; internal polysaccharide plates in some spe-
cies; somatic kineties, reduced in number and 
variable in pattern, forming girdles and spirals, 
typically derived from an \u201cequatorial\u201d girdle 
kinety and a \u201cventral\u201d kinety of dikinetids that 
originates near the posterior pole ; extrusomes 
often prominent rod-like \u201ctrichites\u201d; oral region 
on anterior half with oral polykinetids extensive 
and conspicuous in two sections \u2013 a \u201ccollar\u201d out 
on the body surface encircling anterior pole of 
organism and a \u201clapel\u201d inside the oral cavity 
proper ; paroral, a single file of kinetosomes (i.e., 
monostichomonad); stomatogenesis, apokinetal, 
often in a below-surface pouch; division morpho-
17.3 The Ciliate Taxa to Genus 363
genesis, enantiotropic-like; macronucleus, globular 
to ellipsoid to band-like, often multiple; micronu-
cleus, present; contractile vacuole, at least present 
in freshwater forms; cytoproct, possibly absent; 
feeding on bacteria, microalgae, and smaller pro-
tists, but mixotrophic species common; in marine 
and freshwater habitats, free-living as plankton but 
several species endocommensals in echinoids; one 
order. 
NOTE : Our classification reflects the molecu-
lar genetic analyses that suggest the halteriids , 
classically considered oligotrichs , arose within
the stichotrichs while the strombidiids are a sepa-
rate lineage probably related to the choreotrichs 
(Snoeyenbos-West, Salcedo, McManus, & Katz, 
2002; Strüder-Kypke & Lynn, 2003). 
 Order Strombidiida Petz & Foissner, 1992 
 (syn. Strombidiina ) 
 With characterististics of the class; two families. 
 Family STROMBIDIIDAE Fauré-Fremiet, 1970 
 (syns. Cyrtostrombidiidae , Pelagostrombidiidae ) 
 With characteristics of the order; without an 
elongate and conspicuous contractile tail, but tail 
is often lost in fixed specimens (see Tontoniidae 
below); 17 genera. 
NOTE : We have listed all the names of strom-
bidiid genera considered valid by Aescht (2001) 
and Agatha (2004). However, it is highly doubtful 
given recent research on the molecular evolution of 
 strombidiids that all these genera represent mono-
phyletic clades (e.g., see Agatha, Strüder-Kypke, 
Beran, & Lynn, 2005; Snoeyenbos-West et al., 
2002; Strüder-Kypke & Lynn, 2003). 
 \u2013 Buehringa Busch, 1921 (subj. syn. Strombidium ) 
 \u2013 Cyrtostrombidium Lynn & Gilron, 1993 
 \u2013 Echinostrombidium Jankowski, 1978 
 \u2013 Laboea Lohmann, 1908 
 \u2013 Limnostrombidium Krainer, 1995 
 \u2013 Lissostrombidium Jankowski, 1978 
 \u2013 Metastrombidium Fauré-Fremiet, 1924 
 \u2013 Novistrombidium Song & Bradbury, 1998 
 \u2013 Omegastrombidium Agatha, 2004 *
 \u2013 Parallelostrombidium Agatha, 2004 *
 \u2013 Pelagostrombidium Krainer, 1991 
 \u2013 Peristrombidium Jankowski, 1978 
 \u2013 Pseudostrombidium Horváth, 1933 
 \u2013 Seravinella Alekperov & Mamajeva, 1992 
 \u2013 Spirostrombidium Jankowski, 1978 
 \u2013 Strombidium Claparède & Lachmann, 1859 
 \u2013 Thigmostrombidium Jankowski, 1978 
 Family TONTONIIDAE Agatha, 2004 
 With characteristics of the order; with elongate 
and conspicuous contractile tail, but tail is often 
lost in fixed specimens ; four genera. 
 \u2013 Paratontonia Jankowski, 1978 (subj. syn. 
Tontonia ) 
 \u2013 Pseudotontonia Agatha, 2004 *
 \u2013 Spirotontonia Agatha, 2004 *
 \u2013 Tontonia Fauré-Fremiet, 1914 
 Class ARMOPHOREA Lynn, 2004 
 Size, small to large; shape, varied from top-
shaped with spines to laterally flattened and 
leaf-like; alveoli, conspicuous to absent; somatic 
dikinetids with both kinetosomes bearing cilia, 
typically distributed in kineties covering entire 
body, but in smaller forms ciliature reduced to 
tufts or cirri; hydrogenosomes, with remnant 
genome retained in some forms, replacing 
mitochondria; oral polykinetids on left side 
of oral cavity, few and inconspicuous to many, 
forming an adoral zone; stomatogenesis, typi-
cally pleurokinetal; macronucleus, single, large, 
and ellipsoid to elongate, but sometimes multi-
ple; conjugation, typically temporary, but may 
be total in some armophorids; contractile vacu-
ole, present, sometimes with collecting canals; 
cytoproct, may be present; bacterivorous, but 
also with endosymbiotic methanogens; in marine, 
freshwater, and rarely terrestrial anaerobic habi-
tats (i.e. polysaprobic), typically in sediments 
and the intestinal