Cap 17
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Cap 17


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feeding on flagellates and other ciliates; 
in brackish and freshwater habitats; four genera. 
 \u2013 Actinobolina Strand, 1928 
 \u2013 Belonophrya André, 1914 
 \u2013 Dactylochlamys Lauterborn, 1901 
 \u2013 Legendrea Fauré-Fremiet, 1908 
 Family APERTOSPATHULIDAE Foissner, Xu, & 
Kreutz, 2005 
 Size, small to medium; shape, elongate ovoid; 
free-swimming; somatic ciliation, holotrichous; 
brosse kineties, three; extrusomes as somatic 
mucocysts and oral toxicysts; oral region, api-
cal to subapical, forming a lasso-shaped bulge 
surrounded by an unclosed ring of circumoral 
dikinetids that extends further posteriorly on 
the right side of the oral region than on the left 
side ; macronucleus, ellipsoid to elongate ellipsoid; 
micronucleus, present; contractile vacuole, present; 
cytoproct (?); feeding on flagellates, amoebae, and 
other ciliates; in freshwater and terrestrial habitats; 
three genera. 
 \u2013 Apertospathula Foissner, Agatha, & Berger, 
2002*
 \u2013 Longispatha Foissner, Xu, & Kreutz, 2005 *
 \u2013 Rhinothrix Foissner, Xu, & Kreutz, 2005 *
 Family DIDINIIDAE Poche, 1913 
 (for Cyclodinidae ; syns. Cyclotrichiidae , Didiniina, 
Liliimorphidae ) 
 Size, small to medium; shape, ovoid to ellipsoid, 
often with flattened anterior end, and some taxa with 
368 17. The Ciliate Taxa Including Families and Genera
an anterior protuberance; free- swimming; somatic
cilia as series of apparently short kinetofrag-
ments in one or more girdles around body, but 
in non-ciliated regions, non-ciliated kinetosomes 
are arranged in meridional kineties; brosse, 
typically a field of clavate cilia or \u201csensory bris-
tles\u201d usually clearly detectable in 3\u20135 kineties ; 
extrusomes as somatic mucocysts and multiple 
kinds of oral toxicysts; oral region, apical, often 
in a conical form with eversible cytopharynx in 
some species; macronucleus, ellipsoid to extremely 
elongate and ribbon-like; micronucleus, present; 
contractile vacuole, present; cytoproct (?); feeding 
on flagellates, microalgae, and other ciliates; in 
marine and freshwater habitats, typically plank-
tonic; eight genera. 
 \u2013 Choanostoma Wang, 1931 
 \u2013 Cyclotrichium Meunier, 1910 
 \u2013 Dicyclotrichium Xu, Song, & Hu, 2005 *
 \u2013 Didinium Stein, 1859 
 \u2013 Liliimorpha Gajewskaja, 1928 
 \u2013 Monodinium Fabre-Domergue, 1888 
 \u2013 Pelagovasicola Jankowski, 1980 
 \u2013 Zonotrichium Meunier, 1910 
 Family ENCHELYIDAE Ehrenberg, 1838 
 (syns. Enchelidae , Enchelyina p.p ., Enchelynidae , 
 Enchelyodontidae p.p .) 
 Size, small to medium; shape, ovoid to flask-
like; free-swimming; somatic ciliation, holotri-
chous, with its kineties abutting on oral region 
without arching; brosse kineties, 2\u20134; dikinetids 
or oralized somatic monokinetids surround-
ing cytostome, and all as anterior extremi-
ties of somatic kineties ; extrusomes as somatic 
mucocysts and oral toxicyts; oral region, apical, 
typically flat with cytostome in number of 
species located at distal end of flexible neck ; 
macronucleus, single or multiple, ellipsoid to 
elongate; micronucleus, present; contractile vacu-
ole, present; cytoproct (?); feeding on flagellates 
and other ciliates; typically in freshwater habitats, 
often somewhat anaerobic; 20 genera and two 
incertae sedis . 
NOTE : Some of the genera included in this fam-
ily have oralized somatic monokinetids, while oth-
ers have the typical circumoral dikinetids. It may be 
that oralized somatic monokinetids should be used 
as a synapomorphy for a Family Enchelyodontidae
Foissner, Agatha, and Berger, 2002, but we await 
confirmation of this by gene sequences. 
 \u2013 Apoenchelys Foissner, Agatha, & Berger, 2002 *
 \u2013 Balantidion Eberhard, 1862 
 \u2013 Crobylura André, 1914 
 \u2013 Enchelydium Kahl, 1930 
 \u2013 Enchelys O. F. Müller, 1773 
 \u2013 Haematophagus Woodcock & Lodge, 1921 
 \u2013 Ileonema Stokes, 1884 
 \u2013 Lagynurus Mansfeld, 1923 
 \u2013 Microregma Kahl, 1930 
 \u2013 Nannophrya Kahl, 1933 
 \u2013 Obliquostoma Foissner, Agatha, & Berger, 
2002* [junior homonym of bryozoan] 
 \u2013 Papillorhabdos Foissner, 1984 
 \u2013 Pithothorax Kahl, 1926 
 \u2013 Quasillagilis Busch, 1920 
 \u2013 Rhopalophrya Kahl, 1926 
 \u2013 Schewiakoffia Corliss, 1960 [ nomen dubium ] 
 \u2013 Spasmostoma Kahl, 1927 
 \u2013 Sphaerobactrum Schmidt, 1920 
 \u2013 Thalassiomastix Busch, 1923 
 \u2013 Urochaenia Savi, 1913 
Incertae sedis in Family Enchelyidae 
 \u2013 Microcardiosoma Vuxanovici, 1963 
 \u2013 Microchoanostoma Vuxanovici, 1963 
 Family HELICOPRORODONTIDAE Small & 
Lynn, 1985 
 Size, large, may be >1,000 µm; shape, elongate, 
vermiform, contractile; free-swimming; somatic 
ciliation, holotrichous; brosse kineties, 2\u20135; extru-
somes as somatic mucocysts and up to several 
types of oral toxicysts; toxicysts, distributed along 
perioral ridge that makes from one turn in
Trachelotractus to a spiral of perioral kineties 
in Helicoprorodon ; oral region, apical, with oral 
dikinetids at ends of perioral kineties, and \u201coralized\u201d 
somatic monokinetids also giving rise to nematodes-
mata; macronucleus, many, as isolated nodules or in 
moniliform grouping; micronucleus, not observed; 
contractile vacuole, present; cytoproct (?); feeding 
on flagellates and smaller protists; in marine sands; 
two genera. 
 \u2013 Helicoprorodon Fauré-Fremiet, 1950 
 \u2013 Trachelotractus Foissner, 1997 
17.3 The Ciliate Taxa to Genus 369
 Family HOMALOZOONIDAE Jankowski, 1980 
(syn. Homalozo[on]idea) 
 Size, medium to large; shape, worm-like, laterally 
compressed, contractile ; free-swimming, typically 
gliding on the substrate; somatic ciliation, holotri-
chous, with kineties packed much more densely on 
the right (?) side ; brosse kineties, three; extrusomes 
as somatic mucocyts and oral toxicysts; oral region, 
apical, with temporary cytostome; macronucleus, a 
single elongate ribbon to multiple ellipsoid nodules, 
which may be in a moniliform grouping; micronu-
cleus, present; contractile vacuole, present; cytoproct 
(?); feeding on flagellates and smaller protists; in 
freshwater habitats, especially ponds; one genus. 
 \u2013 Homalozoon Stokes, 1890 
 Family LACRYMARIIDAE de Fromentel, 1876 
 (syn. Lacrymariina p.p .) 
 Size, small to medium; shape, elongate, often 
flask-shaped, with some species having an extremely 
extensible neck-like anterior region; free-swimming, 
rarely sedentary; somatic ciliation, holotrichous; 
brosse kineties, two or more; extrusomes as somatic 
mucocysts and oral toxicysts; anterior region of the 
body (= head), bulb-like, covered by short oblique 
kineties with densely packed kinetids that abut 
the circumoral dikinetids ; oral region, apical, with 
oral dikinetids and cytostome at the anterior end 
of the bulb-like swelling; macronucleus, single or 
multiple, typically ellipsoid; micronucleus, present; 
contractile vacuole, present; cytoproct (?); feeding 
on flagellates and ciliates; in marine, freshwater, and 
terrestrial habitats, with some species planktonic; 
four genera. 
 \u2013 Lacrymaria Bory de St. Vincent, 1824 
 \u2013 Pelagolacrymaria Foissner, Berger, & 
Schaumberg, 1999 
 \u2013 Phialina Bory de St. Vincent, 1824 
 \u2013 Phialinides Foissner, 1988 
 Family PLEUROPLITIDAE Foissner, 1996 
 Size, small; shape, elongate; free-swimming; 
somatic ciliation, holotrichous; brosse kineties, 
several; extrusomes as somatic toxicysts in an 
extracytostomal bundle on the ventral side ; oral 
region, apical; macronucleus, ellipsoid; micronu-
cleus, present; contractile vacuole, present; cyto-
proct (?); feeding on flagellates and small ciliates; 
in terrestrial habitats; two genera. 
 \u2013 Pleuroplites Foissner, 1988 
 \u2013 Pleuroplitoides Foissner, 1996 
 Family PSEUDOHOLOPHRYIDAE Berger, 
Foissner, & Adam, 1984 
 Size, small; shape, ovoid to elongate; free-swim-
ming; somatic ciliation, holotrichous; somatic 
kineties having a slight right spiral; brosse kine-
ties of many rows in which clavate dikinetids 
alternate with typical somatic monokinetids ; 
extrusomes as oral toxicysts; oral region, apical,