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oral polykinetids of “lapel” and 
“collar” 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. 
 – Diophryopsis Hill & Borror, 1992 
 – Diophrys Dujardin, 1841 
 – Paradiophrys Jankowski, 1978 
 – Swedmarkia Dragesco, 1954 
 – 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 2–6 dikinetids forming deli-
cate “cirri” 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. 
 – Balantidioides Penard in Kahl, 1930 
 – 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, 100–200 (24–1,000) 
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 (“capsules torquées”) 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 “historical” 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 . 
 – Acanthostomella Jörgensen, 1927 
 – Ascampbelliella Corliss, 1960 
Incertae sedis in Family Ascampbelliellidae 
 – Luxiella Lecal, 1953 
 – 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 . 
 – Amphorellina Colom, 1948 (fossil) 
 – Bignotella Willems, 1975 (fossil) 
 – Chitinoidella Doben, 1963 (fossil) 
 – Claretinella Keij, 1974 (fossil) 
 – Codonaria Kofoid & Campbell, 1939 
 – Codonella Haeckel, 1873 (some fossils) 
352 17. The Ciliate Taxa Including Families and Genera
 – Codonopsis Kofoid & Campbell, 1939 
 – Coxliellina Colom, 1948 (fossil) 
 – Crassicollaria Remane, 1962 (fossil) 
 – Dicloeopella Eicher, 1965 (fossil) 
 – Durandella Dragastan, 1972 (fossil) 
 – Lorenziella Knauer & Nagy, 1964 (fossil) 
 – Parachitinoidella Trejo, 1972 (fossil) 
 – Poroecus Cleve, 1902 
 – Praetintinnopsella Borza, 1969 (fossil) 
 – Pseudarcella Spandel, 1909 (fossil) 
 – Remanellina Tappan & Loeblich, 1968 (fossil) 
 – Remaniella Catalano, 1965 (fossil) 
 – Salpingellina Colom, 1948 (fossil) 
 – Savroniella Belokrys, 1995 (fossil) 
 – Spinarcella Keij, 1969 (fossil) 
 – Spinophenia Szczechura, 1969 (fossil) 
 – Tintinnopsella Colom, 1948 (fossil) 
 – Tintinnopsis Stein, 1867 (some fossils) 
 – Tytthocorys Tappan & Loeblich, 1968 (fossil) 
 – Urnulella Szczechura, 1969 [not listed in 
Aescht]