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stage as migratory telotroch ; 
oral region and oral ciliature as for subclass; fis-
sion, isotomic or anisotomic, followed in many 
species by development into arboroid colonies; 
resting cysts; feeding on bacteria, microalgae, and 
sometimes detritus; free-living or ectosymbionts, 
with a few species as endosymbiotic forms; 14 
families. 
NOTE : Recent molecular genetic research has 
demonstrated that the morphology of peritrichs 
may be misleading us and suggests we are a long 
way from achieving a natural classification of 
this order (Clamp & Williams, 2006; Gong et al., 
2006).
 Family ASTYLOZOIDAE Kahl, 1935 
 (syns. Astylozooidae , Astylozoonidae , Hast-
atellidae ) 
 Size, small to medium; shape, bell-shaped; tro-
phont, free-swimming, a secondarily(?) stalkless 
form, which swims with oral end forward using 
oral ciliature; telotrochs, lacking somatic cili-
ature; trophonts, with one or two rigid caudal 
bristles or short caudal cilia arising from scop-
ula ; oral ciliature as for subclass; macronucleus, 
band-shaped; micronucleus, present; contractile 
vacuole, present; cytoproct, not observed; cysts, 
in some species; in freshwater habitats, especially 
the plankton of temporary or small ponds; two 
genera.
 – Astylozoon Engelmann, 1862 
 – Hastatella Erlanger, 1890 
 Family ELLOBIOPHRYIDAE Chatton & Lwoff, 
1929
 Size, medium; shape, ovoid-cylindrical, with 
a pair of elongate, cylindrical, and contrac-
tile aboral projections, encircling filamentous 
body parts of the host and cemented together 
by derivatives of the scopula at their tips to 
form a closed circle in firm attachment ; oral 
region as for subclass, but with epistomial disk 
slightly depressed in relation to prominent collar-
ette; fission, anisotomic; macronucleus, ellipsoid; 
micronucleus, present; contractile vacuole, present; 
cytoproct, not observed; in marine habitats as 
ectocommensals on gills of lamellibranch molluscs 
430 17. The Ciliate Taxa Including Families and Genera
and fishes , scale spines of fishes , and oral tentacles 
of bryozoans ; two genera. 
 – Caliperia Laird, 1953 
 – Ellobiophrya Chatton & Lwoff, 1923 
 Family EPISTYLIDIDAE Kahl, 1933 
 (syns. Apiosomatidae , Epistylidae , Ichthyophy-
llinae , Nedulidae , Nidulidae , Nidulinae ) 
 Size, from small to large with some species 
of Campanella and Epistylis having zooids up to 
600 µm in length; shape, ovoid, cylindrical-conical 
or campanulate; trophonts, contractile, on non-
contractile stalk, which may be extremely dif-
ficult to resolve with light microscopy (e.g., 
Apiosoma ) ; if stalkless, in lorica; solitary or colo-
nial; oral region as for subclass, but with peris-
tomial lip and with epistomial disk, only slightly 
projecting; oral ciliature, making from one to as 
many as five complete turns around the peristome; 
macronucleus, elongate, ellipsoid or band-shaped; 
micronucleus, present; contractile vacuole, present; 
cytoproct, not observed; in freshwater and occa-
sionally marine habitats as free-living forms or as 
symphorionts associated with diverse hosts – from 
other peritrichs to molluscs , crustaceans , aquatic 
insects, and some vertebrates, such as freshwater 
 fishes on whose integument Apiosoma is especially 
widely found; 11 genera, of which one is a fossil 
genus from the Lower Triassic. 
 – Apiosoma Blanchard, 1885 
 – Campanella Goldfuss, 1820 
 – Epistylis Ehrenberg, 1830 
 – Foissnerella Jankowski, 1986 
 – Heteropolaria Foissner & Schubert, 1977 
 – Ichthyophyllum Jankowski, 1976 [nomen nudum] 
 – Nuchterleinella Matthes, 1990 
 – Opisthostyla Stokes, 1886 (subj. syn. Rhabdostyla ) 
 – Rhabdostyla Kent, 1881 
 – Triadopercularia Weitschat & Guhl, 1994 
(fossil) 
 – Uvelinus Jankowski, 1985 
 Family LAGENOPHRYIDAE Bütschli, 1889 
 (syns. Lagenophryiidae , Lagenophryinae , Stylo-
hedrinae ) 
 Size, small to medium; shape, ovoid, flattened, 
bilaterally symmetrical; trophont, completely 
enclosed in an ellipsoid, ovoid or hemispheroid 
lorica to which they are attached only at part 
of the interior margin of the aperture; lorica, 
closed by opposing lip-like folds, oriented paral-
lel to the transverse axis of the lorica or with 
a flap-like operculum ; stalk, absent; scopula, 
greatly enlarged in area, apparently acting only 
during attachment of the telotroch; oral region 
as for subclass, but without peristomial lip and 
with an epistomial disk extensible by means of an 
elongate contractile base; fission, anisotomic or 
asymmetric, with telotroch forming to left side 
of parent (when viewed from above) ; macro-
nucleus, elongate, often band-shaped or compact 
ellipsoid; micronucleus, present; contractile vacu-
ole, present; cytoproct, present; in marine, brack-
ish, and freshwater habitats as symphorionts, most 
commonly on gills of freshwater amphipods and 
 decapods , especially crayfish , but also shrimps and 
 crabs , but also on aquatic plants; six genera and 
one genus incertae sedis . 
 – Clistolagenophrys Clamp, 1991 
 – Lagenophrys Stein, 1852 
 – Operculigera Kane, 1969 
 – Paralagenophrys Clamp, 1987 
 – Setonophrys Jankowski, 1986 
 – Stylohedra Kellicott, 1884 (subj. syn. Lage-
nophrys ) 
Incertae sedis in Family Lagenophryidae 
 – Eilymophrys Corliss, 1979 
 Family OPERCULARIIDAE Fauré-Fremiet in 
Corliss, 1979 
 (syns. Bezedniellidae , Entziellidae, Entziellinae, 
 Operculariinae ) 
 Size, small to medium; shape, typically ovoid 
to elongate cylindrical; zooid, contractile, but 
peristomial lip does not fold outward on eversion; 
stalk, typically non-contractile; solitary or colonial, 
with highly developed theca in many species; oral
region as for subclass, but without distinct peris-
tomial lip and with a prominent, extensible and 
contractile epistomial disk that is on a stalk with 
a furrow separating and elevating this disc from 
the margin of the peristome, but epistomial disk 
secondarily reduced in some endocommensal 
species ; oral cavity with an infundibulum, often 
dilated dorsally, into which contractile vacuole 
and cytoproct open; macronucleus, band-shaped or 
17.3 The Ciliate Taxa to Genus 431
ellipsoid; micronuclues, present; contractile vacu-
ole, present; cytoproct, present; in freshwater habi-
tats, very commonly as epibionts on insects and 
other arthropods, but Operculariella is endocom-
mensal in the esophagus of a beetle and Orsomia is 
associated with an oligochaete annelid ; 13 genera 
and one genus incertae sedis . 
 – Ballodora Dogiel & Furssenko, 1921 
 – Bezedniella Stloukal & Matis, 1994 
 – Cyathopercularia Jankowski, 1980 
 – Discophryson Jankowski, 1985 
 – Discotheca Jankowski, 1967 
 – Entziella Stiller, 1951 
 – Opercularia Goldfuss, 1820 
 – Operculariella Stammer, 1948 
 – Orbopercularia Lust in Guhl, 1979 
 – Orbopyxidiella Lust in Guhl, 1979 
 – Propyxidium Corliss, 1979 
 – Scyphidiella Guhl, 1979 
 – Spirocochlearia Jankowski, 1976 [nomen nudum] 
 – Syncyathella Jankowski, 1976 [nomen nudum] 
Incertae sedis in Family Operculariidae 
 – Orsomia Baer, 1952 
 Family OPHRYDIIDAE Ehrenberg, 1838 
 Size, medium to large; shape, bottle-, vase-, 
or spindle-shaped, often with long and highly 
contractile oral end, neck-like ; zooids, solitary 
(Gerda ) or forming gelatinous colonies, often up 
to 15 cm in diameter; green-colored from endo-
symbiotic zoochlorellae in the individual zooids; 
stalk, atypical, with scopula producing long 
peduncular fibers ; oral region as for subclass; 
macronucleus, elongate; micronucleus (?); con-
tractile vacuole, aborally located, with a long 
canal connecting it to the infundibulum ; cyto-
proct (?); cysts; exclusively in freshwater habitats; 
two genera. 
 – Gerda Claparède & Lachmann, 1858 
 – Ophrydium Bory de St. Vincent, 1824 
 Family OPISTHONECTIDAE Foissner, 1976 
 (syns. Telotrochididae , Telotrochidiidae , Telot-
rochiidae ) 
 Size, medium;

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