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feeding on flagellates and other ciliates; 
in brackish and freshwater habitats; four genera. 
 – Actinobolina Strand, 1928 
 – Belonophrya André, 1914 
 – Dactylochlamys Lauterborn, 1901 
 – 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. 
 – Apertospathula Foissner, Agatha, & Berger, 
2002*
 – Longispatha Foissner, Xu, & Kreutz, 2005 *
 – 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 “sensory bris-
tles” usually clearly detectable in 3–5 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. 
 – Choanostoma Wang, 1931 
 – Cyclotrichium Meunier, 1910 
 – Dicyclotrichium Xu, Song, & Hu, 2005 *
 – Didinium Stein, 1859 
 – Liliimorpha Gajewskaja, 1928 
 – Monodinium Fabre-Domergue, 1888 
 – Pelagovasicola Jankowski, 1980 
 – 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, 2–4; 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. 
 – Apoenchelys Foissner, Agatha, & Berger, 2002 *
 – Balantidion Eberhard, 1862 
 – Crobylura André, 1914 
 – Enchelydium Kahl, 1930 
 – Enchelys O. F. Müller, 1773 
 – Haematophagus Woodcock & Lodge, 1921 
 – Ileonema Stokes, 1884 
 – Lagynurus Mansfeld, 1923 
 – Microregma Kahl, 1930 
 – Nannophrya Kahl, 1933 
 – Obliquostoma Foissner, Agatha, & Berger, 
2002* [junior homonym of bryozoan] 
 – Papillorhabdos Foissner, 1984 
 – Pithothorax Kahl, 1926 
 – Quasillagilis Busch, 1920 
 – Rhopalophrya Kahl, 1926 
 – Schewiakoffia Corliss, 1960 [ nomen dubium ] 
 – Spasmostoma Kahl, 1927 
 – Sphaerobactrum Schmidt, 1920 
 – Thalassiomastix Busch, 1923 
 – Urochaenia Savi, 1913 
Incertae sedis in Family Enchelyidae 
 – Microcardiosoma Vuxanovici, 1963 
 – 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, 2–5; 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 “oralized” 
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. 
 – Helicoprorodon Fauré-Fremiet, 1950 
 – 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. 
 – 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. 
 – Lacrymaria Bory de St. Vincent, 1824 
 – Pelagolacrymaria Foissner, Berger, & 
Schaumberg, 1999 
 – Phialina Bory de St. Vincent, 1824 
 – 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. 
 – Pleuroplites Foissner, 1988 
 – 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,