Cap 17
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Cap 17


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oral polykinetids of \u201clapel\u201d and 
\u201ccollar\u201d separated, the latter anteriodorsal; 
paroral, prominent, as polystichomonad, encir-
cling right border of the oral region from its right 
rear to its anterior left ; macronucleus, ellipsoid, 
as two or more separated nodules or moniliform; 
micronucleus, present; contractile vacuole, may be 
present; cytoproct (?); feeding on bacteria, micro-
algae, and smaller protists; in marine habitats, free-
living and in the mantle cavity of molluscs; five 
genera. 
 \u2013 Diophryopsis Hill & Borror, 1992 
 \u2013 Diophrys Dujardin, 1841 
 \u2013 Paradiophrys Jankowski, 1978 
 \u2013 Swedmarkia Dragesco, 1954 
 \u2013 Uronychia Stein, 1859 
Incertae sedis in Subclass Hypotrichia 
 Family REICHENOWELLIDAE Kahl, 1932 
 (syn. Transitellidae ) 
 Size, medium; shape, ellipsoidal; free- swimming; 
somatic ciliation, holotrichous, with kineties occa-
sionally slightly spiraled; ventral somatic kinetids 
as groupings of 2\u20136 dikinetids forming deli-
cate \u201ccirri\u201d and dorsal somatic dikinetids as 
bristles ; extrusomes, not reported; oral region, 
a narrowed peristomial field with an oral cav-
ity supported by a basket of nematodesmata, 
originating from kinetosomes of the oral polyki-
netids and paroral ; adoral zone of polykinetids 
on left side of oral region; paroral and/or endoral 
on right side of oral region; macronucleus, elon-
gate; micronucleus, present; contractile vacuole, 
present; cytoproct (?); feeding on microalgae and 
smaller protists; in freshwater or terrestrial habitats;
 three genera. 
 \u2013 Balantidioides Penard in Kahl, 1930 
 \u2013 Reichenowella Kahl, 1932 
 Subclass Choreotrichia Small & Lynn, 1985 
 (syns. Oligotricha , Oligotrichorida p.p ., Strobilia 
p.p .) 
 Size, small to large; shape, typically conical or 
bell-shaped, sometimes tailed; free-swimming (even 
when loricate, as in Order Tintinnida ); a perilemma, 
often present external to the cell (plasma) membrane; 
extrusomes, restricted to the oral region as capsules 
torqueés, at least in tintinnids; somatic ciliature, as 
dikinetids or monokinetids, poorly developed, rang-
ing from weakly holotrichous (e.g. Strombidinopsis ) 
to extremely reduced (e.g. Lohmanniella ); adoral 
zone of oral polykinetids, used in locomotion 
and feeding, forming closed, outer circle around 
broader anterior end, but slightly open at least 
in the living forms of Parastrombidinopsis ; inner 
ends of some of these outer polykinetids may extend 
17.3 The Ciliate Taxa to Genus 351
into the oral cavity where they may accompany a 
smaller number of inner oral polykinetids restricted 
to the inner oral cavity; paroral, typically composed 
of single file of kinetosomes (monostichomad); 
stomatogenesis, apokinetal, ultimately developing 
in a below- surface pouch; macronucleus, typically 
as two ellipsoid fragments, but various other shapes 
possible; micronucleus, present; contractile vacuole, 
present at least in freshwater forms; cytoproct, pos-
sibly absent; feeding on bacteria, microalgae, and 
other protists; mainly in marine habitats, but some 
freshwater forms, typically planktonic; two orders. 
 Order Tintinnida Kofoid & Campbell, 1929 
 (syns. Archaetintinnoinea , Eutintinnina p.p ., Tintin-
nina , Tintinnoida , Tintinnoidea , Tintinniona , 
 Tintinnoinea ) 
 Size, small to large; shape, cylindrical or cone-
haped, highly contractile, often with elongate 
posterior end; attached to inside of lorica, and sed-
entary within lorica ; loricae, 100\u2013200 (24\u20131,000) 
µm in length, but up to 3,000 µm if certain aberrant 
questionable fossil material is included; loricae, typ-
ically rigid, but gelatinous in Family Tintinnidiidae , 
hyaline or agglomerated with mineral or organic 
particles; free-swimming or sessile; tentaculoids 
containing extrusomes (\u201ccapsules torquées\u201d) inter-
spersed between oral polykinetids in some taxa; 
macronuclei, typically two; micronucleus, present; 
in marine and freshwater habitats, typically marine, 
widespread in pelagic and neritic plankton (with fos-
sil evidence for past aeons); 15 families, excluding 
several families based on fossil taxa (see NOTE ). 
NOTE : If phylogenies derived from small sub-
unit rRNA gene sequences represent the true phy-
logenetic relationships, the taxonomy of this order, 
based on lorica morphology, is very probably 
incorrect (Strüder-Kypke & Lynn, 2003). Agatha 
and Riedel-Lorjé (2006) noted that fewer than 20 
species have the kinetome described in sufficient 
detail from silver impregnation to permit a rigorous 
comparative morphological analysis. Until more 
complete morphological and gene sequence data 
are available, we have conservatively retained the 
classification based on loricae. 
 We do not accept the all-fossil Families Calpio-
nellidae Bonet, 1956, Calpionellopsidae Maka rieva, 
1982, Colomiellidae Bonet, 1956, Chitinoidellidae 
Grün and Blau, 1997, Crassicolariidae Makarieva, 
1982, Remaniellidae Makarieva, 1982, Semichitino-
idellidae Nowak, 1978, distributing their genera as 
incertae sedis among other families (see earlier review 
by Loeblich & Tappan, 1968). Several recently estab-
lished families for fossil genera are listed at the end 
of this section with their included genera, which can-
not be easily placed within the families that include 
contemporary taxa. A thorough review of these fossil 
data by a taxonomist familiar with the contemporary 
diversity of the tintinnids would provide a useful per-
spective on the \u201chistorical\u201d diversity of this group. 
 Family ASCAMPBELLIELLIDAE Corliss, 1960 
 (for Craterellidae ) 
 Size, small; lorica, cup-shaped, not elongate, 
with smooth to denticulate oral rim and trilaminar 
wall; lorica rim of collar as inner collar and 
outer flared rim, with gutter or trough between ; 
lorica of some species with agglutinated coccol-
iths; in marine habitats, mainly eupelagic; no fossil 
species; two genera and two genera incertae sedis . 
 \u2013 Acanthostomella Jörgensen, 1927 
 \u2013 Ascampbelliella Corliss, 1960 
Incertae sedis in Family Ascampbelliellidae 
 \u2013 Luxiella Lecal, 1953 
 \u2013 Niemarshallia Corliss, 1960 
 Family CODONELLIDAE Kent, 1881 
 (syns. Calpionellidae p.p ., Chitinoidellidae , 
 Crassicollariidae , Remaniellidae ) 
 Size, small to medium; lorica flask-, bowl- or chalice-
shaped, with aboral end sometimes pointed and 
closed; lorica collar not clear, but if present may 
or may not have a nuchal constriction; lorica wall, 
unilaminar, commonly reticulate and agglom-
merated ; predominantly in marine habitats, neritic 
and eupelagic forms, but a few species (e.g., of 
Codonella and Tintinnopsis ) abundant in the plankton 
of freshwater lakes, rivers, and ponds; numerous
fossil as well as widespread contemporary forms; 
28 genera and 16 fossil genera incertae sedis . 
 \u2013 Amphorellina Colom, 1948 (fossil) 
 \u2013 Bignotella Willems, 1975 (fossil) 
 \u2013 Chitinoidella Doben, 1963 (fossil) 
 \u2013 Claretinella Keij, 1974 (fossil) 
 \u2013 Codonaria Kofoid & Campbell, 1939 
 \u2013 Codonella Haeckel, 1873 (some fossils) 
352 17. The Ciliate Taxa Including Families and Genera
 \u2013 Codonopsis Kofoid & Campbell, 1939 
 \u2013 Coxliellina Colom, 1948 (fossil) 
 \u2013 Crassicollaria Remane, 1962 (fossil) 
 \u2013 Dicloeopella Eicher, 1965 (fossil) 
 \u2013 Durandella Dragastan, 1972 (fossil) 
 \u2013 Lorenziella Knauer & Nagy, 1964 (fossil) 
 \u2013 Parachitinoidella Trejo, 1972 (fossil) 
 \u2013 Poroecus Cleve, 1902 
 \u2013 Praetintinnopsella Borza, 1969 (fossil) 
 \u2013 Pseudarcella Spandel, 1909 (fossil) 
 \u2013 Remanellina Tappan & Loeblich, 1968 (fossil) 
 \u2013 Remaniella Catalano, 1965 (fossil) 
 \u2013 Salpingellina Colom, 1948 (fossil) 
 \u2013 Savroniella Belokrys, 1995 (fossil) 
 \u2013 Spinarcella Keij, 1969 (fossil) 
 \u2013 Spinophenia Szczechura, 1969 (fossil) 
 \u2013 Tintinnopsella Colom, 1948 (fossil) 
 \u2013 Tintinnopsis Stein, 1867 (some fossils) 
 \u2013 Tytthocorys Tappan & Loeblich, 1968 (fossil) 
 \u2013 Urnulella Szczechura, 1969 [not listed in 
Aescht]