Chapter17 The Ciliate Taxa Including Families and Genera

339

T his chapter, in large measure, is independent of the others. Here, we diagnose or characterize in a succinct manner, all suprageneric taxa assign-able to the Phylum Ciliophora. In each charac-terization, we have highlighted text in bold that we believe refers to synapomorphies or shared derived traits for these taxa. These characteriza-tions are revisions of those provided in Corliss (1979), Lynn and Small (2002), de Puytorac and collaborators (1994a), and other references subse-quent to these publications. Kahl’s (1930–1935) great series has invaluable species descriptions, but omitted entirely certain major higher groups, and there is no comparison between the size of the total assemblage then and now. All genera are listed, supplying author and date. Synonyms are occasionally indicated where genera are con-sidered problematic, and homonyms are noted. We have also indicated those genera designated as n omen nudum by Aescht (2001) as these highlight generic concepts for which additional research may confirm the conclusions of the original describer. Nevertheless, our coverage does not go to the nomenclatural and taxonomic which can be found in the second volume of the ist works: the chapters of the Traité volume have been separately cited in the previous chapters

devoted to treatment of the classes. We have included newly described genera published since Aescht (2001) and Berger (1999, 2001), and have indicated these genera by marking them with an asterisk (*).

D etails, discussions of controversial matters,

revelation of the rationale behind the new classi-fication, evolutionary interrelationships, definition of terminology employed, and rich citation of per-tinent literature sources are still to be found only in the preceding chapters.

17.1 Style and Format T he classes are arranged, in supposed phylogenetic order, based on the data provided by molecular phylogenetic analyses and ultrastructural stud-ies ( s ee Chapter 16 ). However, other taxa are presented alphabetically, each name followed by its author(s) and date of first valid description. Throughout, we have generally followed Aescht (2001) as the nomenclatural authority on genera. Remarks and comments are kept to a minimum but C haracterizations of taxa are as complete as possible at the levels of phylum, subphy-ence is made to size, the following criteria were used: small means < 80 μm in b ody length ; medium means 80–200 μm in b ody length ; and

C hapter 17

T

he Ciliate Taxa Including Families and Genera

13 March 2000. Nor do we provide diagnoses and or shared-derived characters for each taxon are characterizations of genera or subgenera, many of with a minimum characterization. Diagnostic of the literature in relation to genera published up to should be consulted for a comprehensive treatment depth of Aescht (2001), whose scholarly work are included when considered indispensable. Traité edited by de Puytorac and collaborators lum, class, and family. Other taxa are provided (1994a), Jankowski (2007), and in other special-highlighted in bold typeface. Whenever refer-

340 17. The Ciliate Taxa Including Families and Genera

large means > 200 μm in b ody length . Taxa are considered “ f ree-swimming ” if they are typically not attached to a substrate by a s talk or l orica or some other a ttachment organelle . In relation to life history, we make the broad distinction between “ f ree-living ” (i.e., NOT associated with other organisms) and s ymbiotic . We use the fol-lowing classification of symbioses: c ommensal-ism – the symbiont benefits but the host does not; m utualism – both symbiont and host benefit; and p arasitism – the symbiont benefits and the host is obviously harmed. Ciliates have often been called p arasitic , but they are most often c ommensalistic . In the characterizations of fami-lies, it is common that a general feature might be followed by “(?)” (e.g., feeding (?) ). This is meant to indicate that we have not been able to discover information about that character from the literature. We would be grateful to receive correspondence that would remove these areas of ignorance, either through personal observations by the correspondent or direction to the literature on the group.

W e have reserved the use of the category i ncertae sedis for families and genera that seem to demand restudy to determine their appropriate assignment in our overall scheme.

17.2 Nomenclatural Notes, Abbreviations, and Figure References

N ames of all authors of a taxonomic name are written out in full, even when the authorship is multiple, with a single exception. Whenever “de Puytorac et al., 1974” appears after a name, it is to be understood to stand for the following complete list of co-authors: de Puytorac, Batisse, Bohatier, Corliss, Deroux, Didier, Dragesco, Fryd-Versavel, Grain, Grolière, Hovasse, Iftode, Laval, Roque, Savoie, and Tuffrau.

A bbreviations ( n ot italicized if directly follow-ing a g eneric name) include: hom. = homonym; n. n.= n omen novum;n on= not; p.p. = p ro parte; s.l. = s ensu lato;s.s. = s ensu stricto; subj. syn. = subjective synonym; syn(s). = synonym(s). In the case of subjective synonyms, we have noted all those that were listed by Aescht (2001) and for which we currently have reserved judgement on their validity. When comments are used, they are preceded by “ N OTE” or are in brackets.

S ynonymies have been established either by reference to the original literature or, when that was unavailable, by using the informa-tion provided in the Z oological Record.For many of the genera established, for example, by Jankowski (1978, 1979, 1980, 1981, and some-times later) and others, and for which authors have not provided assignment to family, we have assumed that the genus remains within the family to which its t ype species was assigned at the time the author proposed the new genus. For example, P elagotrichidium Jankowski, 1978 has H ypotrichidium faurei as the type spe-cies, but Jankowski (1978) did not assign the new genus to a family. Therefore, we have assumed that the new genus P elagotrichidium remains in the Family SPIROFILIDAE to which H ypotrichidium was assigned.

S ince Corliss (1979), numerous monotypic fam-ilies have been established. In the vast majority of cases, we have not affirmed these families, but rather placed them in synonymy with the family from which the type genus of the new monotypic family originated. Future research, especially using molecular genetic approaches, may dem-onstrate the genetic distinctiveness of these pro-posed monotypic families. This would provide the additional support necessary to remove them from genus S chizocaryum as i ncertae sedis in the latter family. Examination of S chizocaryum by electron microscopy (Lynn & Frombach, 1987) and gene sequencing (Lynn & Strüder-Kypke, 2002) dem-onstrated in the former case that its morphology was very different from the p lagiopylids , justify-ing the establishment of a monotypic family, and demonstrated in the latter case that the family should be placed within the Order P hilasterida synonymy. For example, Corliss (1979) synonymized the monotypic Family SCHIZOCARYIDAE with the Family PLAGIOPYLIDAE , and placed its type (see p. 414, 417).

17.3 The Ciliate Taxa to Genus 341

17.3 The Ciliate Taxa to Genus P hylum CILIOPHORA Doflein, 1901 (syns. C iliae ,C iliozoa ,C ytoidea ,E ozoa ,H etero c aryota ,H eterokaryota ,I nfusoria ,also C iliata [ C iliatea ,C iliaside ,E uciliara ]+ S uctoria [ S uctorea ], G ymnostomea + C iliostomea + T entaculifera ,+ P ostciliodesmatophora ,R habdophora + C yrtophora ,E piplasmata + M em b ranellophora ) E ukaryotic, unicellular, protists; size, 10–4,500 μm; free-swimming or sessile; pellicular alveoli; w ith simple or compound cilia in at least one stage of the life cycle ; complex cortical infra-ciliature, divided into somatic and oral regions; c or-tical micro t ubular or microfibrillar structures associated with the kinetosome including a later-ally- or anteriorly-directed kinetodesmal fibril at kineto s omal triplets 5–7, a tangential or radial transverse microtubular ribbon at triplets 3–5, and a postciliary microtubular ribbon at triplet somatic cilium; extrusomes, common, with somatic extrusomes as mucocysts and oral extrusomes as of micronucleus ; contractile vacuole, typically trophic, feeding modes ranging from osmotrophy to phagotrophy, and some mixotrophy; broadly dis-ecto- or endosymbionts common; two subphyla.

Subphylum POSTCILIODESMATOPHORA Gerassimova & Seravin, 1976 (syns.Ciliostomatophora p.p .,Heterotricha +Kinetofragminophora p.p .,Homotricha p.p .,P olyhymenophora p .p ., T ubulicorticata p .p .) S ize, generally large; shape, typically elongate,

highly contractile; sessile and free-swimming; cor-tical alveoli, poorly developed;

s omatic ciliation with dikinetids that have postciliodesmata, a special arrangement of overlapping postciliary

microtubular ribbons ;

parasomal sacs, absent; somatic extrusomes as mucocysts, rhabdocysts, and/or pigmentocysts; oral structures, highly vari-able, from prostomatous to a highly differentiated adoral zone of polykinetids; during stomatogen-esis, the oral apparatus of the proter usually under-going regression and redevelopment prior to and/or conjugation, temporary and isogamontic; diverse feeding habits; species widely distributed; rarely symbiotic; two classes. C lass K ARYORELICTEA Corliss, 1974 (syns.Antostomatina p.p .,Epitrichina ,Infero-trichina ,K aryorelictida ,K aryorelictina ,L oxodina ,P rimociliatida -K aryorelictina ,R hynchophorina )fragile, flattened, often contractile; free-swimming and highly thigmotactic; many genera having a more-or-less conspicuous non-ciliated or sparsely nuclei, non-dividing, two to many, containing approximately, sometimes slightly more than, the diploid amount of DNA; macronuclei form only by division of micronuclei, then differenti-ate ;

conjugation, temporary; contractile vacuole, often apparently absent; in marine and freshwater sands; three orders.

N OTE : Foissner (1995a), and see also Foissner and Dragesco (1996b), placed the Family K ENTROPHORIDAE in the Order L oxodida , whose members he defined as having a specialized dorsolateral ‘bristle’ kinety and an epipellicular

mucus and/or scale layer on the left body surface. However, members of the Family L OXODIDAE do not have the scale layer, while some members of the Family T RACHELOCERCIDAE show a ‘bristle-like’ kinety. Molecular genetic data should

K inetodesmatop h ora 9 ; parasomal sac often adjacent to the base of the

during cytokinesis; fission almost always isotomic; S ize, generally, large; shape, long, vermiform, toxicysts; oral region, generally monostomic, but some and all taxa having a bristle-kinety surrounding

ciliated surface region (= glabrous zone or stripe), ciliodesmata whose major microtubular ribbons

are typically separated by one microtubule ;

sto-the glabrous zone; s

omatic kinetids with post-matogenesis, parakinetal or buccokinetal;

m acro-present; cytoproct, often present; typically hetero-presumed diploid micronuclei and one to several multiple; nuclear dualism with one or more perkinetal, isotomic or anisotomic, and occasionally or telokinetal; fission homothetogenic and often conjugation, which may be temporary or total, apokinetal, parakinetal, mixokinetal, buccokinetal habitats, typically interstitial forms of marine groups mouthless or polystomic; stomatogenesis ampliploid (rarely diploid or oligoploid) macro-with gametic nuclei formed by meiotic division nuclei , with acentric mitosis; sexual process by tributed in diverse aquatic and terrestrial habitats;

+ P olyhymenophorea ,T ubuli c orticata + F ilicorticata +Kinetofragminophora + Oligohymenophorea

342 17. The Ciliate Taxa Including Families and Genera

demonstrate the phylogenetic affinity of the k en-trophorids and help to demonstrate which morpho-logical features are ancestral and which derived.

O rder P rotostomatida Small & Lynn, 1985 (syns. K entrophorida p.p., S ymbiophagina p.p., T hysanophorida p.p., T rachelocercida ,T rachelo-cercina )

S hape, long to very long, highly contractile; v entral (= right?) somatic surface densely cili-ated, dorsal (= left?) somatic surface glabrous (= non-ciliated) and of varying width; border between somatic surfaces ringed by a ‘bristle’ or ‘bristle-like’ kinety that may extend from the left margin around the posterior end and anteriorly along the right margin, possibly with circumoral kinetids inserted in the anterior right region; oral region, apical or ventral; in marine or estuarine interstitial habitats; two families.

F amily K ENTROPHORIDAE Jankowski, 1980

S ize, large; shape, flattened, vermiform, often C-shaped in cross-section; free-swimming; ventral (= right?) surface densely ciliated, covered by kine-ties of ciliated dikinetids; d orsal (= left?) surface only bordered by ‘bristle’ kinety and covered by mucous material colonized by symbiotic sul-phur bacteria; extrusomes as somatic secretory ampullae (= mucocysts?); oral region, apical, and oral ciliature, if present, highly reduced to rem-nants of kinetids; cytostome, not permanent; nuclei in clusters, typically more than 10, commonly with four macro n uclei surrounding one micronucleus; contractile vacuole (?); cytoproct (?); feeding pri-marily by ingesting ‘its’ symbiotic bacteria through habitats; one genus.

–K entrophoros Sauerbre

F amily T RACHELOCERCIDAE Kent, 1881 (syn. P rototrachelocercidae ,S ultanophryidae )

S ize, large; shape, elongate, contractile, often with distinct ‘head and neck’; free-swimming; somatic ciliature as files of dikinetids, which may cover the body completely or leave a gla-brous longitudinal zone of varying width on the dorsal (= left?) side; glabrous zone, if present, bordered by a ‘bristle-like’ kinety; extrusomes as small somatic mucocysts; o ral area apical, surrounded by circumoral ciliature of diki-netids and accompanied by a brosse of either short ciliated rows or an unstructured ciliated tuft; cytostome, inconspicuous, not perma-nent; nuclei in one or more clusters, with several macronuclei surrounding one or two micro-nuclei; contractile vacuole, absent; cytoproct (?); feeding on bacteria and smaller protists; in marine interstitial habitats; six genera.

–K ovalevaia Foissner, 1997

–P rototrachelocerca Foissner, 1996

–S ultanophrys Foissner & Al-Rasheid, 1999

–T rachelocerca Ehrenberg, 1840

–T rachelolophos Foissner & Dragesco, 1996

–T racheloraphis Dragesco, 1960

O rder L oxodida Jankowski, 1980

(syns. C ryptopharyngina ,L oxodina )

S ize, medium to large; s hape, laterally flat-tened, non-contractile;free-swimming; somatic cilia as files of dikinetids mainly on the right surface with the left surface barren except for sin-gle marginal (= ‘bristle’?) kinety; extrusomes as somatic cnidocyst-like organelles in some genera; o ral region, subapical on the narrow ventral surface; oral kinetids as dikinetids surrounding the oral area as two perioral kineties and one intraoral (= intrabuccal) kinety;stomatogenesis, monoparakinetal or buccokinetal; nuclei in clus-ters, typically of two macronuclei and one micro-nucleus; in marine and freshwater (only L oxodes) habitats, typically in anoxic sediments and anoxic water columns; two families.

Family CRYPTOPHARYNGIDAE Jankowski, 1980

S ize, small to medium; shape, ovoid, flat, margin often serrate; e pipellicular ornamen-tal scales embedded in mucous layer on left body surface; free-swimming; somatic ciliature as files of dikinetids on the right (= ventral?) surface and a single left lateral kinety border-ing the left (= dorsal?) surface; extrusomes, not reported; o ral region, circular to ovoid, ringed by inner and outer dikinetid files with

the naked dorsal surface; in marine interstitial

y, 1928

17.3 The Ciliate Taxa to Genus 343

an intrabuccal kinety, which may be short or long; nuclei in clusters, typically two macronu-clei and one micronucleus; contractile vacuole, absent; cytoproct (?); feeding on bacteria and smaller protists; in marine interstitial habitats; two genera.

–A pocryptopharynx Foissner, 1996

–C ryptopharynx Kahl, 1928

F amily L OXODIDAE Bütschli, 1889

(syns. C iliofaureidae ,D repanostom(at)idea )

S ize, medium to very large; s hape, long, flat, with beak-like anterior rostrum, which inter-rupts perioral kineties at the anterior end;free-swimming; somatic ciliature as files of dikinetids on the right (= ventral?) surface and a single left lateral kinety bordering the left (= dorsal?) surface; extrusomes as somatic cnidocyst-like organelles in R emanella;Müllerian vesicles in the endo-plasm, containing strontium ( R emanella)and barium ( L oxodes)salts;oral area in long ven-tral groove behind rostrum, with inner and outer dikinetid files and a long, rectilinear intrabuccal kinety; nuclei in clusters with one or two macronuclei and a single micronucleus; con-tractile vacuole, in freshwater species; cytoproct (?); feeding on bacteria and other protists, such as microalgae; in freshwater ( L oxodes)and marine ( R emanella) habitats, typically in sediments but ranging into the water column when oxygen levels decline; two genera.

–L oxodes Ehrenberg, 1830

–R emanella Foissner, 1996

O rder P rotoheterotrichida Nouzarède, 1977

S ize, large; shape, elongate, highly contractile; often pigmented, appearing black in transmitted microscopic light; free-swimming; somatic cili-ation, holotrichous; o ral region, ventral, with an elaborate oral ciliature including a series of transverse rows of monokinetids, conspicu-ous either on the left or the right side of the oral region, depending upon the genus;sto-matogenesis, not described; nuclei in clusters, typically two macronuclei and one micronu-cleus; contractile vacuole, absent; cytoproct (?); feeding on microalgae and other protists; in marine or brackish coastal sands and gravels; one family.

F amily

G ELEIIDAE Kahl, 1933

(syn. A veliidae )

W ith characteristics of the order; four genera.

–A velia Nouzarède, 1977

–G eleia Kahl in Foissner, 1998

–G ellertia Dragesco, 1999

–P arduczia Dragesco, 1999

I ncertae sedis in Class K ARYORELICTEA

–C iliofaurea Dragesco, 1960

–C orlissia Dragesco, 1954

C lass H ETEROTRICHEA Stein, 1859

p.p .)

S ize, medium to large; shape, variable, from compressed to conical, often elongate and con-tractile; free-swimming, but some species are temporarily or permanently sessile; some species are pigmented, often brightly, with pigment in ribbons are typically separated by 1 + 2 micro-xtrusomes, sometimes as mucocysts, but polykinetids conspi c uous, typically paramem-branelles, encircling the anterior end clockwise before plunging into the oral cavity; one or more “parorals” on the right side of the oral cav-ity; stomatogenesis, parakinetal; m acronucleus,

ple; conjugation, temporary; contractile vacuole, often conspicuous, with long collecting canals; cytoproct, present; distributed widely in marine, freshwater, and terrestrial habitats; one order.

O rder H eterotrichida Stein, 1859

(syns. B lepharismina p.p., C oliphorida p.p.,Colipho-rina p.p.,Condylostomatina p.p.,Peritromida p.p.,Stentorina p.p.)

W ith characteristics of the class; nine families. highly polyploid, divided by extra-macro-nuclear microtubules; micronuclei, typically multi-specialized pigmentocysts; somatic ciliation, holo-postciliodesmata whose major microtubular tubules; e

trichous; s omatic kinetids as dikinetids with pigmentocysts can also be secreted; l eft serial oral (syns.Heterotricha,Heterotrichorina ,M embra-nellata p.p.,Membranellophora p.p.,Spirotricha p.p, Spirotrichophora

344 17. The Ciliate Taxa Including Families and Genera

F amily B LEPHARISMIDAE Jankowski in Small & Lynn, 1985

S ize, medium to large; shape, pyriform or ellipsoid, somewhat narrowed anteriorly, laterally compressed; free-swimming; pigmentocysts, common, filled with polykinetids extending along the left margin of the oral region and circling counter-clockwise into a shallow posterior oral cavity; p aroral dikinetids anterior of cytostome (precytostomal);macro-nucleus, ovoid to sometimes nodular; micronucleus, may be multiple; contractile vacuole, present; cyto-proct, present; feeding on bacteria, microalgae, and becoming cannibalistic; in marine, freshwater, and terrestrial habitats; four genera.

–A nigsteinia Isquith, 1968 –B lepharisma Perty, 1849 –P arablepharisma Kahl, 1932 [nomen nudum] –P seudoblepharisma Kahl, 1927 F amily C HATTONIDIIDAE Villeneuve-Brachon, 1940

S ize, large; shape, somewhat rotund, contractile, with gently pointed posterior end; free-swimming; somatic ciliation, holotrichous; a unique somatic “posteroaxial cavity,” containing 6–7 ciliary organelles, opening at posterior pole ; oral cavity opening apically and bearing full circle of strong oral polykinetids; macronucleus, very long, with loops; micronucleus, present; contractile vacuole (?); cytoproct (?); feeding on bacteria, microalgae, and other protists, including ciliates; in hypereu-trophic brackish waters; one genus.

–C hattonidium Villeneuve, 1937

F amily C LIMACOSTOMIDAE Repak, 1972

(syn. P ediostomatidae )

S ize, medium to large; shape, ovoid, often anteri-

o ral region anterior, with very prominent, ciliated peristomial field occupying much of the anterior part of the body, bordered at least on the posterior and right margins by the serial adoral polyki-netids ; paroral ciliature, inconspicuous; macronu-cleus, usually as a thick ribbon, often coiled and lengthy; micronucleus, present; contractile vacuole, at least in freshwater forms; cytoproct (?); symbiotic zoochlorellae in some species; feeding on bacteria, microalgae, and other protists, including ciliates; in marine and freshwater habitats; three genera. –C limacostomum Stein, 1859 –F abrea Henneguy, 1890 –P ediostomum Kahl, 1932

F amily C ONDYLOSTOMATIDAE Kahl in Doflein S ize, medium to large; shape, very elongate in some forms, nearly ellipsoidal in others; free-swimming; somatic ciliation, holotrichous; extrusomes as saccu-late somatic mucocysts in some species; o ral region expansive; oral ciliature, including adoral polyki-istomial field absent ; macronucleus, typically long

and moniliform; micronucleus, typically multiple; contractile vacuole often with long collecting canal;

cytoproct, present; feeding on bacteria, microalgae,

and other protists; in marine, freshwater, and terrestrial habitats; seven genera and one genus i ncertae sedis .–C ondylostoma Bory de St. Vincent, 1824 –C ondylostomides Da Silva Neto, 1994 –C opemetopus Villeneuve-Brachon, 1940 –E lectostoma Jankowski, 1979

–L inostomella Aescht in Foissner, Berger, & Schaumberg, 1999

–P redurostyla Jankowski, 1978 –P rocondylostoma Jankowski, 1979 I ncertae sedis in Family C ondylostomatidae –D ellochus Corliss, 1960 F amily F OLLICULINIDAE Dons, 1914

(syn. C oliphorida ,C oliphorina ) S ize, medium to large; shape, typically elongate, contractile; often pigmented; m ature forms, sessile and sedentary, always residing in a lorica; body, especially in neck region in some species, with conspicuous pair of “peristomial wings” bearing the prominent oral ciliature ;a t division, complex stage that “recapitulates” a typical heterotrich

netids and a prominent paroral membrane ;per-the pigment blepharismin; somatic ciliation, holo-& Reichenow, 1929 (syns.Condylostomidae,Condylostentoridae)trichous; extrusomes as sacculate mucocysts; oral other protists, including ciliates, with some species orly pointed; free-swimming; somatic ciliation, holo-morphogenesis with vermiform migratory larval trichous; extrusomes as sacculate somatic mucocysts;

17.3 The Ciliate Taxa to Genus 345

with reduced spiralling of the adoral zone;macro-nucleus, variable, as single ellipsoid to multiple and moniliform; micronucleus, single to multiple; contrac-tile vacuole, present at least in freshwater forms; cyto-proct (?); feeding on bacteria, microalgae, and other protists; widely distributed in marine habitats, but a few species in freshwater, attached to algae, higher aquatic plants, or integument or shells of invertebrates (e.g., molluscs, various crustaceans, bryozoa, coelen-terates); 30 genera including some fossil forms.

–A mpullofolliculina Hadzi, 1951

–A scobius Henneguy, 1884

–A triofolliculina Hadzi, 1951 [nomen nudum]

–A ulofolliculina Hadzi, 1951

–B otticula Dioni, 1972

–C laustrofolliculina Hadzi, 1951

–D iafolliculina Hadzi, 1951 [nomen nudum]

–E chinofolliculina Dons, 1934

–E pifolliculina Hadzi, 1951

–E ufolliculina Hadzi, 1951

–F olliculina Lamarck, 1816

–F olliculinopsis Fauré-Fremiet, 1936 [nomen nudum]

–H alofolliculina Hadzi, 1951 [nomen nudum]

–L agotia Wright, 1857

–L atifolliculina Hadzi, 1951

–M agnifolliculina Uhlig, 1964 [nomen nudum] –M etafolliculina Dons, 1924

–M irofolliculina Dons, 1928

–P achyfolliculina Hadzi, 1951

–P arafolliculina Dons, 1914

–P ebrilla Giard, 1888

–P erifolliculina Hadzi, 1951

–P lanifolliculina Hadzi, 1951

–P latyfolliculina Hadzi, 1938

–P riscofolliculina Deflandre & Deunff, 1957 (fossil) –P seudofolliculina Dons, 1914

–P seudoparafolliculina Andrews & Nelson, 1942 –S plitofolliculina Hadzi, 1951 [nomen nudum]

–S tentofolliculina Hadzi, 1938

–V alletofolliculina Andrews, 1953

Family MARISTENTORIDAE Miao, Simpson, Fu, & Lobban, 2005

S ize, large, majestic when fully extended; s hape, trumpet-shaped with apical area expanded into a bilobed “cap” divided by a ventral indentation, very contractile; free-swimming, but typically tem-porarily attached to substrate; pigmented, due both to blood-red pigmentocysts and to the presence of symbiotic zooxanthellae; somatic ciliation, holotri-chous; o ral ciliature of over 300 oral polykinetids, spirals around flared-out anterior end, encircling an anterior peristomial field with scattered cili-ate dikinetids, not arranged in kineties;paroral, very reduced; macronucleus, ellipsoid; micronuclei, multiple; contractile vacuole, absent; cytoproct, not reported; feeding on bacteria, microalgae, and other protists; in marine habitats, at least associated with coral reefs; one genus.

–M aristentor Lobban, Schefter, Simpson, Pochon, Pawlowski, & Foissner, 2002 *

F amily P ERITROMIDAE Stein, 1867

S ize, medium; shape, ellipsoidal, dorsoventrally flattened, contractile; free-swimming; somatic cili-ation primarily on ventral (= right?) surface; s ingle somatic kinety on slightly convex dorsal (= left?) surface; spine-like cilia of dorsal kinetids emerg-ing from wart-like papillae; extrusomes, not reported; o ral region expansive with left serial oral polykinetids extending anteriorly from equatorial oral cavity along anterior border of cell and paroral extending parallel to serial oral polykinetids; macronucleus, typically in two lobes; micronucleus, present; contractile vacuole, present; cytoproct, present; feeding on bacteria, microalgae, and other protists; generally in marine habitats, including salt marshes; one genus.

–P eritromus Stein, 1863

F amily S PIROSTOMIDAE Stein, 1867

(syn. S pirostomatidae )

S ize, medium to large; shape, often elongate, cylin-drical and very contractile; some pigmented forms; free-swimming; somatic ciliation, holotrichous; extru-somes, not reported; oral region in anterior half; p eristomial field, long, narrow, not ciliated; oral ciliature sometimes relatively inconspicuous, but still with many serial oral polykinetids, extend-ing 1/3–1/2 body length; paroral may extend almost entire length of oral region, paralleling the adoral zone; macronucleus, moniliform; micronucleus, present; contractile vacuole posterior, frequently large, and may have lengthy collecting canal; cytoproct, present;

346 17. The Ciliate Taxa Including Families and Genera

feeding on bacteria, microalgae, and other protists; predominantly in freshwater habitats; two genera and two genera i ncertae sedis.

–G ruberia Kahl, 1932

–S pirostomum Ehrenberg, 1834

I ncertae sedis in Family S pirostomidae

–D iplogmus Mansfeld, 1923

–P ropygocirrus Mansfeld, 1923

F amily S TENTORIDAE Carus, 1863

S ize, medium to large, becoming majestic in size and movement; s hape, trumpet-shaped with apical area not bilobed, very contractile;often pigmented due to pigmentocysts with the pig-ment stentorin and/or with symbiotic zoochlo-rellae; free-swimming, but typically temporarily attached to the substrate, with a few species residing in mucilaginous loricae; somatic ciliation, holotri-chous, with posterior end having thigmotactic cilia permitting temporary attachment to substrate; o ral ciliature spirals nearly 360° around flared-out anterior end, encircling an anterior peristomial field that is covered by ciliated kinetids arranged in ordered kineties; paroral accompanying entire length of adoral zone of polykinetids;macro-nucleus, ellipsoid to ribbon-like and moniliform; micronucleus, one to many; contractile vacuole, may be multiple; cytoproct, present; feeding on bac-teria, microalgae, and other protists, including other ciliates; typically in freshwater habitats only; one genus and three genera i ncertae sedis.

–S tentor Oken, 1815

I ncertae sedis in Family S tentoridae

–H eterostentor Song & Wilbert, 2002 *

–P arastentor Vuxanovici, 1961

–S tentoropsis Dogiel & Bychowsky, 1934

S ubphylum I NTRAMACRONUCLEA T A Lynn, 1996 (C iliostomatophora p.p., H omoiotricha p.p., H omo-tricha p.p., K inetodesmatophora p.p., P ostcilio-nematophora p.p., T ubulicorticata + F ilicorticata + E piplasmata + M embranellophora p.p., T rans-S ize, small to large; shape, variable, from globular to ellipsoid to elongate; free-swimming or sessile; cortical alveolar system typically well-developed; somatic ciliation, holotrichous, but forms with girdles and strips, and even non-ciliated taxa are known; parasomal sacs, present; extrusomes as somatic mucocysts and trichocysts and oral and somatic toxicysts; oral structures, variable, minimally with oral dikinetids, either encircling the cytostome or on the right side as a paroral, but some forms also with several to many oral polykinetids in an adoral zone and other forms astomatous or with oralized somatic kinetids; stomatogenesis, variable from telokinetal to buccokinetal; fission, typically isotomic, rarely anisotomic and multiple; m acronuclear genome typically differentiated by fragmentation of micro-nuclear chromosomes during anlage development;

nuclear microtubules; micronucleus, present; con-major classes and subclasses as obligate symbionts,

freshwater, and terrestrial habitats; nine classes.

N OTE: Lynn (1996a) suggested that the rapid radiation within this subphylum arose from a fundamentally different property of ciliate cellular the micronuclear chromosomes and processing of the genomic DNA during macronuclear develop-ment (see also Riley & Katz, 2001).

C lass S PIROTRICHEA Bütschli, 1889 (syns. H alteriia p.p., H alteriida p.p., M embranellata p.p., M embranellophora P olytrichidea + O ligotrichidea ,P ostcilio d esmatophora p.p., S pirotricha S ize, small to large; shape, variable, from spheroid to cone-shape to dorsoventrally flattened; free-swim-ming or sessile, with some loricate forms that may be attached to substrates and/or sedentary attached within lorica; somatic ciliation, holotrichous in some groups, but nearly devoid of cilia in others; somatic dikinetids usually with the anterior or both kineto-somes ciliated or somatic polykinetids, called cirri; extrusomes as mucocysts or trichocyst-like trichites; o ral ciliature conspicuous, with adoral zone of oral polykinetids, typically as paramembranelles, especially prominent and often encircling oral region clockwise before entering the oral cavity; one or more “parorals” on the right, and if two, as paroral (= outer) and endoral (= inner) membranes; stomatogenesis, typically parakinetal or apokinetal, but mixokinetal in P rotocruzia; fission almost always ampliploid macronucleus dividing by intramacro-sometimes parasitic; widely distributed in marine, jugation, typically temporary and isogamontic, but some forms showing complete conjugation with an-isogamonty; feeding habits, diverse, including several organization, perhaps related to fragmentation of

p.p.,

versonematophora p.p.,Ventrata p.p.)

p.p., Spirotrichia p.p.)

17.3 The Ciliate Taxa to Genus 347

isotomic; m acronuclear DNA replication by repli-cation bands (except in Subclasses P rotocruziidia a nd probably P hacodiniidia(but latter needs con-firmation) ); conjugation, typically temporary and isogamontic, but at least one case of total conjugation; feeding habits, diverse, ranging from microphagous bacterivores to predators on other ciliates and even small metazoans; widely distributed in marine, fresh-water, and terrestrial habitats; seven subclasses.

that the Subclasses P rotocruziidia ,P hacodiniidia , L icnophoria ,H ypotrichia ,C horeotrichia ,O ligotrichia , and S tichotrichia belong to a monophyletic group here called the Class S PIROTRICHEA (see Lynn & Strüder-Kypke, 2002). Macronuclear replication bands have not been demonstrated in p rotocruziids and p hacodini-ids .P rotocruziids demonstrate a form of macronuclear division unique within the phylum (Ammermann, 1968; Ruthmann & Hauser, 1974) and may deserve separate status as a monotypic class. P hacodiniids need careful restudy to determine whether or not they have replication bands or something akin to them. Subclass Protocruziidia de Puytorac, Grain, & Mignot, 1987

S ize, small; shape, ovoid; free-swimming; alveoli not well-developed; body with monokinetid or diki-netid field on right side and dorsal surface; somatid dikinetids with short kinetodesmal fibrils and over-lapping postciliary microtubular ribbon, which do not form postciliodesmata; extrusomes, trichocyst-like; adoral zone of 6 (5–8) oral polykinetids on left of oral region; paroral to posterior and right of oral region, composed of dikinetids; stomatogenesis, mixokinetal; n uclear apparatus a cluster of similar-sized nuclei with paradiploid macronuclei surrounding one or more micronuclei; macronuclear division appar-ently as separation of two composite (?) chromo-somes per macronucleus; conjugation, not reported; contractile vacuole (?); cytoproct (?); microphagous, on bacteria, microalgae, and smaller protists; in marine and brackish water habitats; one order.

O rder P rotocruziida Jankowski, 1980

W ith characters of subclass; one family.

F amily P ROTOCRUZIIDAE Jankowski, 1980

W ith characters of order; one genus.

–P rotocruzia de Faria, da Cunha, & Pinto, 1922 S ubclass P hacodiniidia Small & Lynn, 1985

ally; free-swimming; cortex with rigid and ribbed pellicle (= cuirass) and well-developed alveoli; s omatic ciliature, in widely spaced rows, of linear polykinetids of 6–8 kinetosomes with delicate cilia; only a few cirrus-like somatic polykinetids, composed of two rows of kineto-somes; o ral region, long, with an adoral zone

at the cytostome, very near posterior pole of

short files of kinetosomes;macronucleus horse-shoe-shaped; micronuclei, multiple; contractile vacuole, present; cytoproct, present; feeding on bacteria, microalgae, and smaller protists; mainly in freshwater and terrestrial habitats (e.g., moss on trees); one order.

O rder P hacodiniida Small & Lynn, 1985

(syn. P rotohypotrichina p.p.)

W ith characters of the subclass; one family.

F AMILY P HACODINIIDAE Corliss, 1979

W ith characters of the order; one genus.

–P hacodinium Pro

S ubclass L icnophoria Corliss, 1957

S ize, medium; s hape, in form of hour-glass, with prominent oral disc apically and conspicu-free-swimming, but typically attached to substrate; s omatic cilia, essentially absent, except for pos-terior ciliary rings encircling attachment disc; adoral zone of oral polykinetids encircling oral region; paroral as single file of kinetosomes; sto-matogenesis, apokinetal; macronucleus, typically moniliform; micronucleus, one to several; con-jugation, temporary; contractile vacuole, absent; cytoproct, absent; microphagous on bacteria, microalgae, and perhaps organic detrital particles derived from host’s feeding activities; in marine habitats as “ectocommensals” on organisms ranging from an alga (substratum for one species) to a variety of invertebrates (e.g., tunicates, coelenterates, annelids, molluscs, and the respiratory trees of sea cucumbers); one order.

NOTE: Molecular evolution studies suggest of conspicuous oral polykinetids, terminating

organism; paroral, series of obliquely-oriented,

S ize, medium; shape, ovoid, compressed later-

wazek, 1900

ous aboral attachment disc at posterior pole;

(syn.Scaiotricha p.p.)

348 17. The Ciliate Taxa Including Families and Genera

O rder L icnophorida Corliss, 1957

W ith characteristics of subclass; one family. F amily L ICNOPHORIDAE Bütschli, 1887

W ith characteristics of order; two genera. –L icnophora Claparède, 1867 –P rolicnophora Jankowksi, 1978

S ubclass H ypotrichia Stein, 1859

(syns.Euplotia ,Hypotricha p.p .,Hypotrichea

p .p .,Hypotrichida p.p .,Hypotrichina p.p .,H ypotrichorida p .p ., P seudohypotrichina ) S ize, small to medium; shape, dorsoventrally

flattened, typically rigid, oval to rectangular; free-swimming; a lveoli well-developed and, at least

in euplotids, filled with a protein, called platein ;somatic ciliature commonly represented by rows or localized groups of polykinetids, called cirri, conspicuous on the ventral surface; d orsally, files

of widely spaced dikinetids with short cilia

(“sensory bristles”) and retention of a laterally-directed kinetodesmal fibril ; files of marginal cirri, incomplete or absent; s omatic infraciliature typically retained during encystment ;prominent adoral zone of generally numerous oral polyki-netids, as paramembranelles, on left-anterior por-tion of the ventral surface, bordering a broad, non-ciliated peristomial field and sometimes con-tinuing over apical end of body onto the dorsal surface; paroral as paroral and/or endoral in diplo- or polystichomonad condition; stomatogenesis, generally apokinetal, beginning in a cortical pocket in some forms, but sometimes parakinetal; macro-nucleus, ellipsoid to band-shaped or in fragments, with replication bands moving from ends to mid-dle when the nucleus is elongated; micronucleus, one to several; conjugation, temporary; contractile one inquilinic in an echinoid; two orders. rapid sequence evolution. This may explain why the representative genera that have been sequenced do not form a strong monophyletic grouping (Chen & Song, 2001). We conservatively maintain

this subclass until sequence data from additional genes provide evidence that the class should be subdivided.

O rder K iitrichida Nozawa, 1941

(syn. P rotohypotrichina p .p .) S ize, medium; shape, small, rounded-elliptical in outline; f rontoventral cirri, relatively small polykinetids, uniform in size, in 7–10 curving files

along right side of ventral surface ; oral region,

broad area, on right; adoral zone of polykinetids

bordering left margin of body and extending from

near posterior end to near anterior end; paroral, bor-dering almost entire length of right margin of oral

region; macronucleus, ovoid; micronucleus, present;

contractile vacuole, absent; cytoproct (?); feeding

on smaller protists; in marine habitats; one family.

F amily K IITRICHIDAE Nozawa, 1941

W ith characteristics of order; three genera. –C aryotricha Kahl, 1932 –K iitricha Nozawa, 1941 –M usajevella Alekperov, 1984 O rder E uplotida Small & Lynn, 1985 S ize, small to medium; shape, ovoid to rectangu-lar; free-swimming; frontoventral cirri, sporadically

scattered over ventral surface, but never forming more than one conspicuous file on ventral surface, except in G astrocirrhidae ; oral structures, as for subclass; d uring cell division, only the ventral tion of the dorsal ciliature typically occurs within an equatorial band and within the parental kine-ties (i.e., intrakinetally) ; caudal cirri, when present, derived from dorsal kinety anlagen; two suborders. made distinct from main body by more or less obvi-thigmotactic; right marginal cirri usually present, but not on D iscocephalus ;f ile of left marginal cirri typi-cally divided into anterior and posterior- l ateral vacuole, at least present in freshwater forms;S uborder D iscocephalina Wicklow, 1982 microphagous and macrophagous; in marine, fresh- S ize, small to medium; s hape, elongate ovoid NOTE : Sequences of small subunit rRNA genes with anterior “head-like” part bearing oral region free-living forms, but a few species as ecto-ous neck-like constriction ; free-swimming, but quite commensals on various invertebrates and one of species assigned to this subclass generally show water, and terrestrial habitats, widely distributed as

somatic infraciliature is replaced while replica-

17.3 The Ciliate Taxa to Genus 349

parts; transverse cirri, conspicuous with well-devel-oped microtubular rootlets; caudal cirri, present; oral structures, as for subclass; macronucleus, often in many fragments; micronucleus, present; contractile vacuole, absent; cytoproct (?); feeding on microalgae and smaller protists; in marine habitats, especially sands; one family.

N OTE: The Family E rionellidae was placed in this suborder by Lynn and Small (2002). However, its sole genus E rionella Jankowski, 1978 is likely a synonym of H olosticha(Aescht, 2001).

F amily D ISCOCEPHALIDAE Jankowski, 1979 (syns. D iscocephalinae ,D iscocephaloidea ,M argi-notrichinae )

W ith characteristics of the suborder; three genera. N OTE:P sammocephalus Wicklow, 1982 has been included in this family. Lin, Song, and Warren (2004) view it as a junior synonym of P rodiscocephalus Jankowski, 1979.

–D iscocephalus Ehrenberg in Hemprich & Ehrenberg, 1831

–M arginotricha Jankowski, 1978

–P rodiscocephalus Jankowski, 1979

S uborder E uplotina Jankowski, 1979

(syn. E uplotia ,E uplotiidea ,G astrocirrhida p.p., U ronychiida p.p.)

S ize, small to medium; shape, ovoid to ovorec-tangular; free-swimming; r ight marginal cirri, absent;l eft marginal cirri, when present, not as two distinct groups; oral structures, as for subclass; c ontractile vacuole, typically in right posterior of body; in marine, freshwater, and terrestrial habitats; five families.

F amily A SPIDISCIDAE Ehrenberg, 1830

(syn. A spidiscina ,A spidiscoidea ,E uplotaspinae , P araeuplotidae )

S ize, small; shape, flattened and disc-like; free-swimming, highly thigmotactic; dorsal surface may be ridged; n o left marginal cirri;transverse cirri, conspicuous; c audal cirri, absent; reduced number of oral membranelles, located centrally and inconspicuously on ventral surface;paroral, reduced or absent; macronucleus usually C-shaped; micronucleus, present; contractile vacuole, present; cytoproct (?); feeding on bacteria, microalgae, and smaller protists; in marine and freshwater habitats, widely distributed, often benthic but including com-mensals in the echinoid gut and ascidian branchial cavity; two genera and one genus i ncertae sedis.

–A spidisca Ehrenberg, 1830

–E uplotaspis Chatton & Séguéla, 1936

I ncertae sedis in Family A spidiscidae

–P araeuplotes Wichterman, 1942

F amily C ERTESIIDAE Borror & Hill, 1995 (syn. C ertesiina ,C ertesiinae )

S ize, s mall; s hape, b roadly o void; f ree-swimming; u nique condylopallium in anterior end of cell; left marginal cirri more than three;transverse cirri, large, well-developed; caudal cirri, absent; adoral zone of polykinetids, well-developed and continuous; paroral polykinetid conspicuous, exten-ding along two thirds length of the oral cavity; macronucleus, ellipsoid, two to four in number; micronuclei, several; contractile vacuole, present; cytoproct (?); feeding on diatoms and other smaller protists; in marine habitats; one genus.

–C ertesia Fabre-Domergue, 1885

F amily E UPLOTIDAE Ehrenberg, 1838

(syn.Euplotiidea,Euplotidiidae,Euplotidiinae, E uplotina ,E uplotinae ,P loesconiidae )

S ize, small to medium; shape, ovoid, ventrally-flattened; body, rigid; free-swimming; extrusomes as small vesicles (i.e., ampules) associated with dorsal bristle dikinetids; frontoventral and transverse cirri, dispersed in conspicuous groups; l eft marginal cirri, reduced typically to fewer than three; caudal cirri, ventral; v entrally-oriented oral cavity with distinct, contiguous, adoral zone of oral polykinetids forming a “collar” and “lapel”; p aroral as polykinetid on right of oral area accompanied by single endoral file of kineto-somes; macronucleus, more or less C-shaped; micronucleus, present; contractile vacuole, present; cytoproct (?); feeding on bacteria, microalgae, and smaller protists; in marine, freshwater, and terres-trial habitats, widely distributed but predominantly marine, with one E uplotes species found in the intestines of sea urchins; four genera.

350 17. The Ciliate Taxa Including Families and Genera

–E uplotes Ehrenberg in Hemprich & Ehrenberg, 1831

–E uplotoides Borror & Hill, 1995

–E uplotopsis Borror & Hill, 1995

–M oneuplotes Jankowski, 1978

F amily

G ASTROCIRRHIDAE Fauré-Fremiet, 1961 (syns.Cytharoidinae,Gastrocirrhida,Gastro-cirrhina ,G astrocirrhinae ,G astrocirrhoidea )

S ize, medium; shape, conoid, nearly round in cross-section; free-swimming; f rontoventral cirri apparently in two files; left marginal cirri, inconspicuous or absent;t ransverse cirri, many, conspicuous, in U-shape; expansive, anteriorly-opened oral cavity with anterior end of body remarkably truncate; macronucleus, ellipsoid, typically in two fragments; micronucleus, present; contractile vacuole, absent; cytoproct (?); feed-ing on bacteria, microalgae, and smaller protists; in marine habitats, either planktonic or psam-mophilic; three genera.

N OTE:P araeuplotidium Lei, Choi, and Xu, 2002 is considered a junior synonym of E uplotidium since the single left marginal cirrus proposed by Lei, Choi, and Xu (2002) as a character of generic distinctiveness must be corroborated first by gene sequence data to confirm its significance.

–C ytharoides Tuffrau, 1975

–E uplotidium Noland, 1937

–G astrocirrhus Lepsi, 1928

F amily U RONYCHIIDAE Jankowski, 1975 (syns. D iophryidae ,S wedmarkiidae ,U ronychiida ) S ize, small to medium; shape, blunt ovoid, nearly circular in cross-section; free-swimming; cirri, gen-erally conspicuous with frontoventral cirri reduced to groupings on right side; transverse cirri, well-devel-oped; r ight caudal cirri, dorsal, well-developed; left marginal cirri, may be conspicuous; o ral region, expansive, with 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.

–D iophryopsis Hill & Borror, 1992

–D iophrys Dujardin, 1841

–P aradiophrys Jankowski, 1978

–S wedmarkia Dragesco, 1954

–U ronychia Stein, 1859

I ncertae sedis in Subclass H ypotrichia

F amily R EICHENOWELLIDAE Kahl, 1932 (syn. T ransitellidae )

S ize, medium; shape, ellipsoidal; free- s wimming; somatic ciliation, holotrichous, with kineties occa-sionally slightly spiraled; v entral somatic kinetids as groupings of 2–6 dikinetids forming deli-cate “cirri” and dorsal somatic dikinetids as bristles; extrusomes, not reported; o ral 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;–B alantidioides Penard in Kahl, 1930

–R eichenowella Kahl, 1932

S ubclass C horeotrichia Small & Lynn, 1985

S ize, small to large; shape, typically conical or when loricate, as in Order T intinnida ); 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. S trombidinopsis) to extremely reduced (e.g. L ohmanniella); a doral 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 P arastrombidinopsis ; inner ends of some of these outer polykinetids may extend two genera.

Strobilia p.p.)

bell-shaped, sometimes tailed; free-swimming (even (syns.Oligotricha p.p., Oligotrichorida p.p.,

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- s urface 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.

O rder T intinnida Kofoid & Campbell, 1929 (syns. A rchaetintinnoinea ,E utintinnina p .p ., T intin-nina ,Tintinnoida ,Tintinnoidea ,Tintinniona ,T intinnoinea ) S ize, small to large; shape, cylindrical or cone-posterior end; a ttached 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 T intinnidiidae , 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 N OTE ).

N OTE : 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. W e do not accept the all-fossil Families C alpio-

i ncertae sedis among other families (see earlier review not be easily placed within the families that include diversity of the tintinnids would provide a useful per-spective on the “historical” diversity of this group. F amily A SCAMPBELLIELLIDAE Corliss, 1960 (for C raterellidae ) S ize, small; lorica, cup-shaped, not elongate, with smooth to denticulate oral rim and trilaminar wall; l orica 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 i ncertae sedis .–A canthostomella J?rgensen, 1927 –A scampbelliella Corliss, 1960 I ncertae sedis in Family A scampbelliellidae –L uxiella Lecal, 1953

–N iemarshallia Corliss, 1960 F amily C ODONELLIDAE Kent, 1881

(syns.Calpionellidae p.p .,C rassicollariidae ,R emaniellidae ) S ize, 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; l orica wall,

unilaminar, commonly reticulate and agglom-merated ; predominantly in marine habitats, neritic

and eupelagic forms, but a few species (e.g., of

C odonella and T intinnopsis ) ab of freshwater lakes, rivers, and ponds; numerous

fossil as well as widespread contemporary forms;

28 genera and 16 fossil genera i ncertae sedis .–A mphorellina Colom, 1948 (fossil) –B ignotella Willems, 1975 (fossil) –C hitinoidella Doben, 1963 (fossil) –C laretinella Keij, 1974 (fossil) –C odonaria Kofoid & Campbell, 1939 C odonella Haeckel, 1873 (some fossils)

undant in the plankton

shaped, highly contractile, often with elongate Chitinoidellidae ,

–Grün and Blau, 1997, Crassicolariidae Makarieva, nellidae Bonet, 1956, C alpionellopsidae Makar ieva, 1982, Colomiellidae Bonet, 1956, Chitinoidellidae tinoidellidae Nowak, 1978, distributing their genera as data by a taxonomist familiar with the contemporary contemporary taxa. A thorough review of these fossil by Loeblich & Tappan, 1968). Several recently estab-of this section with their included genera, which can-lished families for fossil genera are listed at the end 1982, Remaniellidae Makarieva,1982, and S emichi-

352 17. The Ciliate Taxa Including Families and Genera

–C odonopsis Kofoid & Campbell, 1939

–C oxliellina Colom, 1948 (fossil)

–C rassicollaria Remane, 1962 (fossil)

–D icloeopella Eicher, 1965 (fossil)

–D urandella Dragastan, 1972 (fossil)

–L orenziella Knauer & Nagy, 1964 (fossil)

–P arachitinoidella Trejo, 1972 (fossil)

–P oroecus Cleve, 1902

–P raetintinnopsella Borza, 1969 (fossil)

–P seudarcella Spandel, 1909 (fossil)

–R emanellina Tappan & Loeblich, 1968 (fossil) –R emaniella Catalano, 1965 (fossil)

–S alpingellina Colom, 1948 (fossil)

–S avroniella Belokrys, 1995 (fossil)

–S pinarcella Keij, 1969 (fossil)

–S pinophenia Szczechura, 1969 (fossil)

–T intinnopsella Colom, 1948 (fossil)

–T intinnopsis Stein, 1867 (some fossils)

–T ytthocorys Tappan & Loeblich, 1968 (fossil)

–Urnulella Szczechura, 1969 [not listed in Aescht] (fossil)

–V autrinella Cuvillier & Sacal, 1963 (fossil)

–Y vonniellina Tappan & Loeblich, 1968 (fossil) I ncertae sedis in Family Codonellidae

–B acculinella Konenkova, 2000 (fossil)

–B iconvexellina Konenkova, 1999 (fossil)

–B icornella Bugrova, 2003 *(fossil) [junior hom-onym of ostracoda]

–B orzaiella Makarieva, 1982 (fossil)

–B orziella Pop, 1987 (fossil)

–C arpathella Pop, 1998 (fossil)

–C ubanites Aescht, 2001 (fossil)

–C ylindriconella Aescht, 2001 (fossil)

–D aciella Pop, 1998 (fossil)

–D obeniella Pop, 1997 (fossil)

–F oliacella Makarieva, 1979 (fossil)

–L ongicollaria Pop, 1997 (fossil)

–P opiella Rehakova, 2002 *(fossil)

–R ossielella Aescht, 2001 (fossil)

–S calpratella Makarieva, 1979 (fossil)

–T ianella Bugrova, 2003 *(fossil)

F amily C ODONELLOPSIDAE Kofoid & Campbell, 1929

(syns.Calpionellidae p.p.,Calpionellopsidae, C olomiellidae ,S emichitinoidellidae )

S ize, small to large; lorica, top-shaped with abo-ral end rounded to apiculate; l orica collar, hyaline, cylindrical, delicate but sometimes ridged;lorica wall, thick, but wall of bowl thicker than collar and agglomerated with mineral particles; in marine habitats, neritic and eupelagic forms; many fossil as well as extant forms; 12 genera and 12 fossil genera i ncertae sedis.

–C alpionella Lorenz, 1902 (fossil)

–C alpionellites Colom, 1948 (fossil)

–C alpionellopsella Trejo, 1975 (fossil)

–C alpionellopsis Colom, 1948 (fossil) (subj. syn.

R emaniella)

–C odonellopsis J?rgensen, 1924

–C olomiella Bonet, 1956 (fossil)

–D eflandronella Trejo, 1975 (fossil)

–L aackmanniella Kofoid & Campbell, 1929

–L uminella Kofoid & Campbell, 1939

–P raecalpionellopsis Borza, 1971 (fossil)

–S tenosemella J?rgensen, 1924

–S tenosemellopsis Colom, 1948 (fossil)

I ncertae sedis in Family C odonellopsidae

–B aranella Nagy, 1989 (fossil)

–B orzaites Aescht, 2001 (fossil)

–C alpionelloides Colom, 1948 (fossil)

–C alpionellopsites Nagy, 1986 (fossil)

–C rassicalpionella Nagy, 1989 (fossil)

–F urssenkoiella Makarieva, 1979 (fossil)

–L orenziellites Nagy, 1986 (fossil)

–L orenziellopsis Nagy, 1989 (fossil)

–P raecalpionellites Pop, 1986 (fossil)

–S emichitinoidella Nowak, 1978 (fossil)

–S opianella Nagy, 1989 (fossil)

–S turiella Borza, 1981 (fossil)

Family CYTTAROCYLIDIDAE Kofoid& Campbell, 1939

(syns. C yttarocylidae )

S ize, small to medium; lorica, bell- or kettle-shaped, sometimes elongate; l orica wall with very conspicuous meshwork between wall layers and conspicuous wall material in broad bars sepa-rating polygons; lorica collar slightly flared, with inner suboral shelf; in marine habitats, predomi-nantly eupelagic; no fossil species; one genus.

–C yttarocylis Fol, 1881

F amily D ICTYOCYSTIDAE Haeckel, 1873

S ize, small to medium; lorica, ovoid to coni-cal with dense bowl; l orica collar with one or

17.3 The Ciliate Taxa to Genus 353

two rows of open-arched frames, windows or fenestrae (latter with or without panes);lorica wall of bowl reticulate and, in some species, agglomerated with coccoliths or mineral particles; in marine habitats, eupelagic; no fossil species; two genera.

–D ictyocysta Ehrenberg, 1854 –W angiella Nie, 1934

F amily E PIPLOCYLIDIDAE Kofoid & Campbell, 1939

(syns. E piplocyclididae ,E piplocylidae ) S ize, small to medium; l orica, short, acorn-shaped with aboral end blunt acuminate or with horn ; lorica oral rim, smooth, with collarette or suboral shelf; l orica wall, thin, hyaline, with its surface partially or entirely ornamented with polygons bounded by raised edges ; in marine habitats, eupe-lagic; no fossil species; three genera. –E picancella Kofoid & Campbell, 1929 –E piplocylis J?rgensen, 1924 –E piplocyloides Hada, 1938

F amily M ETACYLIDIDAE Kofoid & Campbell,

1929

(syns.Calpionellidae p.p .,Coxliellidae p.p .,M etacycoidinae ,M etacylin[e]ae ) S ize, medium to large; l orica, tubular or g oblet-shaped, d elicate a nd w ith r ings o r s pirals a s w ound lamina, at least in oral half ; lorica aboral end sometimes with horn; l orica wall, three-layered, usually with distinct and delicate alveoli, some-times with agglomerated coccoliths ;paralorica,coxlielliform; sometimes an epilorica; in marine habitats, eupelagic and neritic, though occasion-ally one found in fresh water; a number of fossil species; eight genera and two genera i ncertae sedis . N OTE : The genus C oxliella via its type-spe-cies, at least, may actually have no reality. Laval-Peuto (1977, 1994a) has demonstrated that there is a “coxlielliform” stage in the life cycle of many t intinnines , including species from a number of families. Thus, Corliss (1979) placed the generic replaced the former familial name ( C oxliellidae ) with the next name available for the group, v iz .,

M etacylididae (originally the second included sub-family). –C limacocylis J?rgensen, 1924

–F avelloides Thalmann, 1942 (fossil) –H elicostomella J?rgensen, 1924 –M etacylis J?rgensen, 1924 –P seudometacylis Balech, 1968

–R habdonelloides Colom, 1939 (fossil) –

S piroxystonellites Knauer, 1969 (fossil)

N omen inquirendum :Coxliella Brandt, 1906 (fossil)

–R hizodomus Strelk –S tylicauda Balech, 1951 Small, & Russo, 1991 S ize, small; l orica, hyaline, bell-shaped, with hinged, oral flaps that close lorica opening when habitats, particularly estuarine and coastal plankton; –N olaclusilis Snyder & Brownlee, 1991 F amily P ETALOTRICHIDAE Kofoid & Campbell,

1929 S ize, medium; lorica, bell-shaped with flared lip on top of vertical cone; l orica lip and bowl with –P arapetalotricha Hada, 1970 –P etalotricha Kent, 1881

F amily P TYCHOCYLIDIDAE Kofoid & Campbell, 1929

P tychocyclidae ,P tychacyclididae ,P tycho-cylidae ) S ize, small to large; l orica conical or chalice-shaped with several annular bulges and pointed or blunt pedicel ; lorica oral rim, often denticulate, with no collar; l orica wall, apparently trilaminate, alveolar in midsection ; paralorica, coxlielliform; epilorica, if present, possibly spiralled; in marine genera.

name in a “questionable” status (see below ) and habitats, mostly eupelagic; no fossil species; five ov & Wirketis, 1950 F amily N OLACLUSILIIDAE Sniezek, Capriulo, small, suboral fenestrae;in marine habitats, eu-ciliate retracts ; macronucleus, bilobed; in marine pelagic; no fossil species; two genera. (syns. Incertae sedis in Family Metacylididae no fossil species; one genus.

354 17. The Ciliate Taxa Including Families and Genera

–C ymatocylis Laackmann, 1910

–F avella J?rgensen, 1924

–P rotocymatocylis Kofoid & Campbell, 1929

–P tychocylis Brandt, 1896

–W ailesia Kofoid & Campbell, 1939

F amily R HABDONELLIDAE Kofoid & Campbell, 1929

S ize, medium to large; lorica, acorn to chalice-shaped, often very long with aboral horn in some species; lorica oral rim, smooth; l orica wall, tril-aminate, hyaline with longitudinal, low ridges that may be simple, branched or anastomosing, sometimes with pores; in marine habitats, mostly eupelagic; no fossil species; four genera.

–E pirhabdonella Kofoid & Campbell, 1939

–P rotorhabdonella J?rgensen, 1924

–R habdonella Brandt, 1906

–R habdonellopsis Kofoid & Campbell, 1929

F amily T INTINNIDAE Claparède & Lachmann, 1858

(syns.Salpingellinae p.p.,Stelidiellinae p.p., T intinninae p.p.)

S ize, medium to large; lorica elongate, typically tubular, and in some species, with b oth ends of lorica open; lorica oral end often flared, smooth or denticulate; l orica wall, thin, hyaline, probably unilaminar, appearing homogeneous with clear inner and outer layers; lorica surface ornamentation frequently as ridges and crests, which may be spiralled; prima-rily in marine habitats, eupelagic, but a few found in brackish habitats; no fossil species; 24 genera.

–A lbatrossiella Kofoid & Campbell, 1929

–A mphorellopsis Kofoid & Campbell, 1929

–A mphorides Strand, 1928

–B randtiella Kofoid & Campbell, 1929

–B ursaopsis Kofoid & Campbell, 1929

–B uschiella Corliss, 1960

–C anthariella Kofoid & Campbell, 1929

–C levea Balech, 1948

–D adayiella Kofoid & Campbell, 1929

–D aturella Kofoid & Campbell, 1929 [nomen dubium]

–E picranella Kofoid & Campbell, 1929

–E utintinnus Kofoid & Campbell, 1939

–F unnela Li & Zhang, 2006 *(fossil) –O dontophorella Kofoid & Campbell, 1929

–O rmosella Kofoid & Campbell, 1929

–P roamphorella Kofoid & Campbell, 1939

–P rostelidiella Kofoid & Campbell, 1939

–R habdosella Kofoid & Campbell, 1929

–S alpingacantha Kofoid & Campbell, 1929

–S alpingella J?rgensen, 1924

–S alpingelloides Campbell, 1942

–S teenstrupiella Kofoid & Campbell, 1929

–S telidiella Kofoid & Campbell, 1929

–T intinnus Schrank, 1803

F amily T INTINNIDIIDAE Kofoid & Campbell, 1929

(syn. T intinnididae )

S ize, small to large; lorica, tubular or flaring, with aboral end open or closed, may be attached to the substrate; lorica collar, rarely visible; l orica wall soft, gelatinous, with agglomerated par-ticles; in marine, brackish, and (occasionally) genera.

–L eprotintinnus J?rgensen, 1900

–M embranicola Foissner, Berger, & Schaumburg, 1999

–T intinnidium Kent, 1881

F amily U NDELLIDAE Kofoid & Campbell, 1929 S ize, small to medium; loricae goblet- or urn-shaped, occasionally elongate; lorica oral rim sometimes with suboral ledge and perhaps an inner collar; l orica wall, hyaline, conspicuously thick, obviously trilaminate, with obvious inner and outer layers; lorica surface, smooth, and some-times with annuli; in marine habitats, eupelagic; no fossil species; seven genera and one genus i ncertae sedis.

–A mplectella Kofoid & Campbell, 1929

–A mplectellopsis Kofoid & Campbell, 1929

–C ricundella Kofoid & Campbell, 1929

–M icrundella Loeblich & Tappan, 1968

–P roplectella Kofoid & Campbell, 1929

–U ndella Daday, 1887

–U ndellopsis Kofoid & Campbell, 1929

I ncertae sedis in Family U ndellidae

–R otundocylis Kufferath, 1952 [ n omen nudum] freshwater habitats; no fossil species; three

17.3 The Ciliate Taxa to Genus 355

F amily X YSTONELLIDAE Kofoid & Campbell, 1929

(syn. X istonellidae )

S ize, medium to large; lorica elongate, chalice-shaped with aboral end long and narrow, sometimes with pedicel; lorica oral rim, usually denticulate; l orica wall, hyaline, trilaminate, reticulate, show-ing an irregular and conspicuous meshwork between lorica wall layers, with inconspicuous, thin wall material separating polygons; in marine habitats, mainly eupelagic; a few fossil forms known; five genera and one genus i ncertae sedis.

–P arafavella Kofoid & Campbell, 1929 (fossil) –P arundella J?rgensen, 1924

–S piroxystonella Kofoid & Campbell, 1929

–X ystonella Brandt, 1906

–X ystonellopsis J?rgensen, 1924

I ncertae sedis in Family X ystonellidae

–P arafavelloides Deflandre & Deflandre, 1949

A DDENDUM: The following families and included genera of fossil forms are listed here as belonging to the Order T intinnida . Assignment to families that include contemporary genera has not been possible at this time.

I ncertae sedis in Order T intinnida : unas-signed genera – A ubertianella Szczechura, 1969, D aturellina Radoi?i?, 1959, P raecolomiella Borza, 1979, S pirocystomellites Colom, 1988; S yringella Paulmier, 1997 [not listed in Aescht; junior homo-nym of anthozoan/poriferan], T intinnoidella Elicki, 1994; unassigned Family B erounkellidae Koshevoj, 1987 with included genera B atiola Koshevoj, 1987, B erounkella Koshevoj, 1987, C hervurt s kella Koshevoj, 1987, K ejvia Koshevoj, 1987, O lgella Koshevoj, 1987, O llella Aescht, 2001, T undrella Koshevoj, 1987, and V elavella Koshevoj, 1987; unassigned Family C adosinidae Wanner, 1940 with included genera C adosina Wanner, 1940, C adosinopsis Scheiber, 1967, and C rustocadosina Rehanek, 1985; u nassigned F amily C alcisphaerulidae Bonet, 1956 with included genera B onetcardiella Dufour, 1968 and C alcisphaerula Bonet, 1956; unassigned Family C ausiidae K o shevoj, 1987 with included genus C ausella Aescht, 2001; and unas-signed Family S tomiosphaeridae Wanner, 1940 with included genera C arpistomiosphaera Nowak, 1968, C olomisphaera Nowak, 1968, C ommittosphaera Rehanek, 1985, I nocardion Masters & Scott, 1978, and S tomiosphaera Wanner, 1940. O rder C horeotrichida Small & Lynn, 1985

S ize, small to large; shape, globular to ellip-soid; free-swimming or not sessile, but may be temporarily attached to substratum; n o lorica; in marine and freshwater habitats, widespread, planktonic with some benthic and symbiotic spe-cies; four suborders.

S uborder L eegaardiellina Laval-Peuto, Grain, & Deroux, 1994

S ize, small to medium; shape, spheroid; free-swimming; somatic kineties composed of dikinetids, restricted to aboral half; s erial oral polykinetids of the outer circle divided into an inner and outer part, with inner oral polykinetids, apparently clearly separated from outer oral polykinetids; oral cavity with a set of internal oral polykinetids on the outer wall and paroral along the margin of micronucleus, not observed; contractile vacuole, absent; cytoproct (?); feeding on microalgae and smaller protists; in marine habitats, planktonic; one family.

Family LEEGAARDIELLIDAE Lynn& Montagnes, 1988

W ith characters of the suborder; one genus.

–L eegaardiella Lynn & Montagnes, 1988

S uborder L ohmanniellina Laval-Peuto, Grain, & Deroux, 1994

S ize, small; shape, spheroid; s omatic kineties, short, posterior, possibly composed of monoki-netids; serial oral polykinetids of outer circle undivided; inner oral polykinetids, apparently as extensions of several outer oral polykinetids, but with a break between the outer and inner parts; observed; contractile vacuole, absent; cytoproct (?); feeding on microalgae and smaller protists; marine plankton; one family.

Family LOHMANNIELLIDAE Montagnes& Lynn, 1991

W ith characters of the suborder; one genus.

–L ohmanniella Leegaard, 1915

the inner wall; macronuclei, two ellipsoid nodules; macronucleus, curved, ellipsoid; micronucleus, not

356 17. The Ciliate Taxa Including Families and Genera

S uborder S trobilidiina Small & Lynn, 1985

S ize, small to medium; shape, spheroid to conoid; free-swimming, but some forms attached to substrates by mucous thread; s omatic kine-ties, one to several, usually not as long as the body, composed of monokinetids with a corti-cal flap covering the bases of cilia and direct-ing them to the cell’s left and underlain (when protargol-stained) by densely-stained mate-rial ; somatic cilia, relatively short, typically <5 μm long; outer and inner oral polykinetids typical of the subclass; macronucleus, typically single, variably shaped from ellipsoid to an elon-gate band; micronucleus, if observed, typically single; contractile vacuole, at least present in freshwater forms; cytoproct (?); feeding on bac-teria, microalgae, and smaller protists; in marine mensals of echinoids; one family.

F amily S TROBILIDIIDAE Kahl in Doflein & Reichenow, 1929 (syn. T orquatellidae ) W ith characters of the suborder; three genera and two i ncertae sedis .–P elagostrobilidium Petz, Song, & Wilbert, 1995 –R imostrombidium Jankowski, 1978

–S trobilidium Schewiakoff, 1893

I ncertae sedis in Family S trobilidiidae –C iliospina Leegaard, 1915 [ n omen dubium ]–P atronella Corliss, 1979 (subj. syn.

S trombidium ) S uborder S trombidinopsina Small & Lynn, 1985

S ize, small to medium; shape, spheroid to conoid; free-swimming; s omatic kineties of ciliated diki-netids, equally distributed around body, extend-ing as simple files from the outer ring of serial oral polykinetids towards cell’s posterior ;outer and inner oral polykinetids typical of the subclass, although P arastrombidinopsis may not have com-pletely closed outer circle when alive; macronu-clei, ellipsoid, typically two; micronucleus, when observed, typically single; contractile vacuole, at least present in freshwater forms; cytoproct (?); feeding on bacteria and microalgae; typically in marine habitats, planktonic; one family.

F amily S TROMBIDINOPSIDAE Small & Lynn, 1985

W ith characters of the suborder; two genera. –P arastrombidinopsis Kim, Jeong, Strüder-Kypke, Lynn, Kim, Kim, & Lee, 2005

*

–S trombidinopsis Kent, 1881

S ubclass S tichotrichia Small & Lynn, 1985

(syns.Euhypotrichina ,Hypotricha p.p .,Hypo-trichea p .p ., H ypotrichida p .p ., H ypotrichina p .p ., H ypotrichorida p .p .)

S ize, small to large; shape, often elongate, sometimes very drawn out posteriorly, in cross-section round to dorsoventrally compressed; free-swimming with a few loricate forms; perilemma in some groups; p ellicular alveoli weakly devel-oped; somatic ventral ciliature as ventral cirri ranging from small and quite inconspicuous, occasionally as few as 2–3 cilia per cirrus, arranged in longitudinal, sometimes spiraled,

files to a few, larger cirri in scattered groups, with in the latter case marginal files of cirri dif-ferentiated ; transverse cirri, may or may not be present; caudal cirri, may or may not be present; dorsal somatic ciliature as one to many kineties

– typically three – of dikinetids without kinetodes-mal fibril, but with short, bristle cilium on anterior kinetosome; adoral zone of oral polykinetids as paramembranelles in “collar” and “lapel”, each

typically of four rows of kinetosomes, with the

first two rows equally long and the fourth row quite short; right oral cilia variable, but usually as

a paroral and endoral; stomatogenesis, parakinetal in those with conspicuous kineties to apokinetal in those with scattered cirri; d ivision morphogenesis may involve replacement of all somatic cili-ature of both proter and opisthe ;macronucleus,typically two nodules, but often multiple, each component typically with one replication band; micronucleus, one to many; conjugation, typi-cally temporary, but sometimes total; contractile vacuole, at least present in freshwater and ter-restrial forms, and typically on the middle left of the body; cytoproct, very likely present; feeding strategies ranging from bacterivorous to cannibal-istic;

e ncysted forms typically dedifferentiate all kinetosomes ;

in marine, freshwater, and terrestrial habitats, free-living with some symbiotic forms as

endo- and ectocommensals; three orders. and freshwater habitats, with some species com-

17.3 The Ciliate Taxa to Genus 357

N OTE: The taxonomy of stichotrichians is one of the most confused in the phylum. The revision below relies heavily on the morphology of the dif-ferentiated individual. There is, however, a trend in recent years to rely more heavily on the similarities in the pattern of division morphogenesis (e.g., Berger, 1999, 2006b; Eigner & Foissner, 1994). Stability may only be achieved when complete division morphogenetic patterns and molecular genetic information for several genes are available on the majority of genera (see Foissner et al., 2004).

O rder S tichotrichida Fauré-Fremiet, 1961 (syns.Chaetospirina p.p.,Oxytrichida p.p., O xytri-china p.p., P lagiotomida p.p., P lagiotomoidea p.p.) S ize, small to large; shape, often elongate, some-times very drawn out posteriorly; free-swimming with a few loricate forms; v entral cirri as one or more longitudinal files of varied lengths, linear (not zig-zag as in U rostylida); dorsal ciliature, typically regularly distributed in longitudinal files; oral structures as for subclass; stomatogenesis, parakinetal or apokinetal; six families.

F amily A MPHISIELLIDAE Jankowski, 1979 (syns.Gastrostylidae p.p.,Gastrostylina p.p., O rthoamphisiellidae )

S ize, small to large; shape, elongate ovoid; free-swimming; v entral cirral file, single with anterior segment of this file formed by cirri from right-most ventral anlage and posterior segment from the second ventral anlage from right;marginal files of cirri, typically extending from anterior to posterior on left and right sides; transverse cirri, may or may not be present; caudal cirri, may or may not be present; dorsal kineties, typically fewer than ten, composed of dikinetids; oral structures as for order with paroral and endoral; macronucleus, from two to many globular to ellipsoid nodules; micro-nuclei, one often accompanying each macronuclear nodule; contractile vacuole, present; cytoproct, likely present; feeding on bacteria and smaller protists; in marine, freshwater, and terrestrial habitats;

11 genera and two genera i ncertae sedis.

–A froamphisiella Foissner, Agatha, & Berger, 2002 *

–A mphisiella Gourret & Roeser, 1888

–A mphisiellides Foissner, 1988

–H emiamphisiella Foissner, 1988 –N udiamphisiella Foissner, Agatha, & Berger, 2002 *

–L amtostyla Buitkamp, 1977

–O rthoamphisiella Eigner & Foissner, 1991

–P aramphisiella Foissner, 1988

–P escozoon Jankowski, 1978

–U roleptoides Wenzel, 1953

–U rospinula Corliss, 1960 (subj. syn. P silotricha) I ncertae sedis in Family A mphisiellidae

–B alladyna Kowalewski, 1882 (subj. syn.

C yrtohymena)

–C ircinella Foissner, 1994

F amily K AHLIELLIDAE Tuffrau, 1979

(syns. B anyulsellidae ,C ladotrichidae ,L acazeidae , P arakahliellidae )

S ize, medium to large; shape, elongate ovoid; free-swimming; somatic ventral ciliature with at least two, typically more than two, ventral cirral files, often not distinctly different from right and left marginal cirral files; v entral cirral files may be preserved through a variable number of cell divisions (= cell generations) before being resorbed and replaced through additional new (= neokinetal) anlagen;transverse cirri,typi-cally absent; caudal cirri, typically absent; dorsal ciliature as several files of dikinetids; oral ciliature as for order with paroral and endoral; macronu-cleus, two to many nodules; micronuclei, several to many; contractile vacuole, present; cytoproct, likely present; feeding on bacteria, microalgae, and smaller protists; in marine, freshwater, and terres-trial habitats; ten genera and four genera i ncertae sedis.

–C ladotricha Gajewskaja, 1926

–D eviata Eigner, 1995

–E ngelmanniella Foissner, 1982

–K ahliella Corliss, 1960

–N eogeneia Eigner, 1995

–P arakahliella Berger, Foissner, & Adam, 1985 –P lesiotricha Dragesco, 1970 (subj. syn. K ahliella)–P seudokahliella Berger, Foissner, & Adam, 1985 –T rachelochaeta?rámek-Husek, 1954

–W allackia Foissner, 1976

I ncertae sedis in Family K ahliellidae

–B anyulsella Dragesco, 1954

–F ragmocirrus Foissner, 2000

358 17. The Ciliate Taxa Including Families and Genera

–L acazea Dragesco, 1960

–P seudouroleptus Hemberger, 1985

F amily K ERONIDAE Dujardin, 1840

(syns. K eronina ,K eronopsidae ,K eronopsina )

S ize, medium; shape, from broad to elongate, even tailed in some species; free-swimming; somatic ventral ciliature as frontoventral cirri gen-erally in several oblique rows across the ventral surface, coursing between right and left marginal cirral files; f irst ventral row, so-called “frontal cirri”, as curved row along ventral anterior border of left serial oral polykinetids and differ-entiating from one to several anlagen;transverse cirri, present; caudal cirri, present; dorsal somatic ciliature as several files of bristle dikinetids; oral ciliature as for order with paroral and endoral; cell division in cyst, except for K erona;macronu-cleus, two to many globular to ellipsoid nodules; micronucleus, present; contractile vacuole, present; cytoproct (?); feeding on bacteria, microalgae, and smaller protists; in marine and freshwater habi-tats, including sand, and one species well-known ectocommensal on H ydra; four genera.

–K erona Müller, 1786

–K eronopsis Penard, 1922

–P araholosticha Wenzel, 1953

–P arakeronopsis Shi, Song, & Shi, 1999 [nomen nudum]

F amily P LAGIOTOMIDAE Bütschli, 1887

S ize, medium; shape, laterally flattened, elongate-ovoid, with right side slightly concave; free-s wimming; s omatic ciliation, holotrichous, dense, of small polykinetids or cirri with no differentia-tion into cirral groups on either body surface; oral ciliature of extensive adoral zone on its left side, coursing from apical end to subequatorial posi-tion and then entering a deep oral cavity and with paroral and endoral along right wall of oral cavity; stomatogenesis, parakinetal; macronucleus, as an irregular bunch of nodules; micronuclei, several, relatively large; contractile vacuole, present; cyto-proct (?); feeding on bacteria and organic matter in host’s digestive system (?); i n terrestrial habitats as an endocommensals solely in certain species of lumbricid oligochaete annelids; one genus.

–P lagiotoma Dujardin, 1841 F amily P SILOTRICHIDAE Bütschli, 1889 (syn. P silotrichinae )

S ize, small to medium; shape, oval to elliptical in outline, with posterior spiny extensions in some species, and sometimes with zoochlorellae in cyto-plasm; free-swimming; s omatic ventral ciliature as long and sparse cirri in seven slightly curved cirral files with the postoral oblique cirral file developing from the anlage file IV; frontal cirri; marginal cirri, strongly reduced; transverse cirri, present; caudal cirri, present; dorsal somatic cili-ature as several files of monokinetids (?); peristo-mial areal limited to anterior third of organism with oral ciliature typical of order, including paroral and endoral; macronucleus, two globular to ellipsoid nodules; micronucleus, present; contractile vacu-ole, present; cytoproct (?); feeding (?); in freshwa-–P silotricha Stein, 1859

–H emiholosticha von Gelei, 1954

F amily S PIROFILIDAE von Gelei, 1929 (syns. A tractidae ,C haetospiridae ,C haetospirina , C haetospirinae ,C haetospiroidea ,H ypotrichidiidae , Microspirettidae,Spiretellidae,Spirofilopsidae, Stichotrichinae,Strongylidae,Strongylidiidae, S trongylidiinae ,S trongylidioidea )

S ize, small to medium; shape, varied, with some forms tailed and others more elongate at the ante-rior end; free-swimming, but lorica produced by some species; s omatic ventral ciliature as incon-spicuous ventral cirri in files curved or spiralling obliquely around body, some ending on dorsal surface; transverse cirri, present or absent; caudal cirri, present or absent; s omatic dorsal ciliature as several dorsal files of bristle dikinetids in “dor-sal strip” helically winding around the body; adoral zone not highly prominent with oral cili-ature t ypical of order and with paroral and endoral; macronucleus, globular to ellipsoid, single to sev-eral nodules; micronucleus, present; contractile vacuole, present; cytoproct, likely present; feeding on bacteria, algae, and smaller protists; in marine and freshwater habitats, both benthic and plank-tonic; 12 genera and one genus i ncertae sedis.

–A tractos V?r?sváry, 1950

–C haetospira Lachmann, 1856

–H ypotrichidium Ilowaisky, 1921

ter and terrestrial habitats; two genera.

17.3 The Ciliate Taxa to Genus 359

–M icrospiretta Jankowski, 1975

–M ucotrichidium Foissner, Oleksiv, & Müller, 1990 –P arastrongylidium Fleury & Fryd-V ersavel, 1985 –P elagotrichidium Jankowski, 1978

–P lanitrichidium Jankowski, 1979 [nomen nudum] –S pirofilopsis Corliss, 1960

–S tichotricha Perty, 1849

–S trongylidium Sterki, 1878

–U rostrongylum Kahl, 1932

I ncertae sedis in Family S pirofilidae

–K ahliela Tucolesco, 1962

I ncertae sedis in Order S tichotrichida

–B alladinopsis Ghosh, 1921 [nomen dubium]

–K lonostricha Jankowski, 1979

–P silotrix Gourret & Roeser, 1888

–S tylonethes Sterki, 1878 [nomen dubium]

O rder S poradotrichida Fauré-Fremiet, 1961 (syn. H alteriina p.p., O xytrichina p.p., P leurotrichina p.p., S poradotrichorina )

S ize, small to large; shape, sometimes elongate, even tailed, but often oval to elliptical in outline; free-swimming; s omatic ventral ciliature as fron-toventral cirri, typically heavy and conspicuous, arranged in specific, localized frontal and ven-tral groups, except in a few taxa (e.g., Family Halteriidae, L aurentiella ,O nychodromus ,and Styxophrya); marginal cirri, typically present; transverse cirri, may or may not be present; caudal cirri, may or may not be present; dorsal somatic ciliature, typically as files of dikinetids with a single bristle cilium; oral ciliature as for subclass; s tomatogenesis, apokinetal, usually with five or six anlagen streaks in two groups for differentia-tion of ventral somatic ciliature; in marine, fresh-water, and terrestrial habitats, widely distributed, primarily benthic with some forms planktonic, others interstitial, and a few species symbiotic, either as ectocommensals on the integument or in the branchial cavity of several invertebrates or as intestinal inquilines of echinoids; three families.

N OTE: Gene sequences of both actins and small subunit rRNA unambiguously place several genera of the Family H alteriidae (e.g., H alteria,M eseres) close to the o xytrichid clade (Croft et al., 2003; Foissner et al., 2004; Hewitt et al., 2003). We have therefore transferred this family to this order from the Subclass O ligotrichia . We consider h alteriids as planktonic descendants of this primarily benthic lineage.

F amily H ALTERIIDAE Claparède & Lachmann, 1858

(syn. M eseridae [for L ieberkuehnidae ])

S ize, small; shape, spheroid to subovoid and conical; free-swimming, often darting through the water; s omatic ciliature as somatic kinetids with long cilia (i.e., >10 μm long), often as cirrus-like “bristles” or stiff cilia; oral cili-ature, with the “collar” as an “open” circle of apical oral polykinetids, the “lapel” on the left side of the oral cavity, and only with the paroral (or endoral?) on the right side of the oral cavity; m acronucleus, single, ellipsoid; micronucleus, present; contractile vacuole, present; cytoproct (?); feeding on bacteria and microalgae; predominantly in freshwater habitats, typically planktonic but some in terrestrial habi-–C ephalotrichium Meunier, 1910 (subj. syn.

S trobilidium)

–H alteria Dujardin, 1841

–H alterioforma Horváth, 1956 [ n omen oblitum]

–M eseres Schewiakoff, 1893

–P elagohalteria Foissner, Skogstad, & Pratt, 1988 I ncertae sedis in Family H alteriidae

–J eannellia Tucolesco, 1962 (subj. syn. H alter-ioforma)

–O ctocirrus Madhava Rao, 1929 [ n omen dubium] F amily O XYTRICHIDAE Ehrenberg, 1830 (syns.Ancystropodiinae,Gastrostylidae p.p., Gastrostylina p.p.,Onychodromusidae,Oxytri-chinae ,O xytrichoidea ,P attersoniellidae ,P leuro-trichidae ,R igidotrichidae (?), S tylonychinae )

S ize, small to large; shape, relatively elongate; free-swimming; s omatic ventral ciliature as fron-toventral and transverse cirri, typically 18 in number, large and distinctive, scattered over mid-area of ventral surface, between right and left marginal cirral files, usually with three fron-toventral cirri posterior to posterior vertex of the oral region, but several genera with conspicuous files of cirri (e.g., L aurentiella,O nychodromus); tats; five genera and two i ncertae sedis.

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