Redescription of Licnophora chattoni Villeneuve-Brachon, 1939 (Ciliophora, Spirotrichea), associated with Zyzzyzus warreni Calder, 1988 (Cnidaria, Hydrozoa)

Licnophora chattoni, found in association with Zyzzyzus warreni, a tubulariid hydroid epizoic in sponges from São Sebastião (SP, Brazil), is redescribed and illustrated using light and electron microscopy. The ciliate has a flexible, transparent body formed by an oval anterior region linked to the posterior basal disc via a flexible neck region. Numerous cortical granules are observed scattered throughout the body and densely packed along the neck. The adoral zone is formed by about 81 external and 24 infundibular paramembranelles. The paroral membrane, formed by a row of long cilia arranged in monokinetids, extends through a groove in the body to the adhesive disc. Two dorsal kinetids are present along the right body margin and around the neck. The adhesive disc (18 μm in diameter) lacks cilia in the area above the velum. The velum covers a row of dikinetids bearing long cilia and four dikineties, two or three of which are interrupted on the ventral surface. Nine to twelve macronuclear nodules connected by isthmuses are distributed in the cytoplasm, plus two nodules located in the adhesive disc and between those there is an ovate micronucleus.     

Ultrastructure of Prorodon (Ciliophora, Prostomatida) Ii. Oral Cortex and Phylogenetic Conclusions

ABSTRACT. The ultrastructure of the oral region and the ultrastructural architecture of the basket of Prorodon aklitolophon and Prorodon teres are described. the oral region of Prorodon consists of: 1) A circumoral kinety at the anterior pole of the cell surrounding the typically slit-shaped cytostomial funnel. This kinety is composed of inversely oriented dikinetids in which both kinetosomes are ciliated and are associated with a very short postciliary microtubular ribbon and a few transverse microtubules; 2) Three brush rows aligned in parallel and extended meridionally in the anterior part of the cell. the individual brush rows consist of dikinetids, but in contrast to the dikinetids around the cytopharynx they are not inverted and only the anterior kinetosomes bear specialized short brush cilia and are associated with a divergent-tangential transverse microtubular ribbon. the posterior kinetosome is non-ciliated and bears a prominent convergent postciliary microtubular ribbon. Schematized dikinetid patterns of both oral regions of Prorodon are provided. In addition, a three-dimensional reconstruction of the basket of the genus Prorodon based on serial thin sections is presented. A phylogenetic tree, mainly based on stomatogenic data, is given to show the phylogenetic relationships of some prostomatid genera as well as the hypothesized sistergroup relationship of colpodid and prostomatid ciliates.     

Updating the Trachelocercids (Ciliophora, Karyorelictea). I. A Detailed Description of the Infraciliature of Trachelolophos gigas N. G., N. Sp. and T. filum (Dragesco & Dragesco-Kernéis, 1986) N. Comb

ABSTRACT. Trachelolophos gigas n. g., n. sp. and T. filum (Dragesco & Dragesco-Kernéis, 1986) n. comb. (basionym: Tracheloraphis filum) were discovered in the mesopsammon of the French Atlantic coast at Roscoff. Their morphology and infraciliature were studied in live and protargol impregnated specimens. The new genus, Trachelolophos, belongs to the family Trachelocercidae and is unique in having a conspicuous ciliary tuft, which is very likely a highly modified brosse, in the oral cavity. The two species investigated have a very similar infraciliature, differing only in morphometric characteristics and in the nuclear configuration. The entire somatic and oral infraciliature consists of dikinetids which have both basal bodies ciliated or only the anterior or posterior ones, depending on the region of the cell. The right side is densely and uniformly ciliated. Its kineties extend onto the left side to the glabrous stripe, where an anterior and posterior secant system are formed, reducing the number of kineties in the narrowed neck and tail region. The left side bears a narrow glabrous stripe bordered by slightly irregularly arranged dikinetids having rather stiff cilia (bristles), possibly forming an uninterrupted, prolate ellipsoidal (bristle) kinety as indicated by their ciliation. The bristle kinety commences subapically at the right margin of the glabrous stripe, extends posteriorly, then anteriorly at the left, to end up at the right margin again. The dikinetids of the right posterior portion of the bristle kinety have the posterior basal bodies ciliated, whereas the anterior basal bodies are ciliated in its left and right anterior portion. The ends of the bristle kinety meet distinctly subapically at the right margin of the glabrous stripe, as indicated by the diametrically opposed ciliation of the dikinetids. The anterior region (head) of the cell bears a distinct circumoral kinety composed of very regularly arranged dikinetids, associated with nematodesmata forming an oral basket together with the nematodesmal bundles originating from the oralized somatic dikinetids at the anterior end of the somatic kineties. The systematics of trachelocercid ciliates are briefly reviewed and discussed.     

Morphology and Notes on Morphogenesis during Cell Division of Deviata polycirrata n. sp. and of Deviata bacilliformis (Gelei, 1954) Eigner, 1995 (Ciliophora: Kahliellidae) from Argentina

ABSTRACT. Described herein are the morphology and certain morphogenetic stages of a new freshwater ciliate species, Deviata polycirrata n. sp., and of Deviata bacilliformis recorded in the soil of a dried temporary pond from Argentina. Ciliates were studied alive and after silver impregnation with Protargol. Deviata polycirrata n. sp. measures 130–180 × 45–70 μm in vivo. The species possesses 8–9 long cirral rows on the right and 9–13 on the left of the oral zone, and 3 dorsal rows of dikinetids. The adoral zone is composed of 39–48 membranelles. There are four macronuclear nodules and usually two micronuclei. A single contractile vacuole is located equatorially on the left body margin. This new species mainly differs from its congeners in having a higher number of cirral rows, the three long dorsal rows of dikinetids (vs. usually one to two dorsal rows of dikinetids), and a higher number of adoral membranelles. The other species reported here, D. bacilliformis, is recorded for the first time in Argentina. Unlike previous observations on this species, on the dorsal surface there are cirral rows that are preceded by cilia (combined cirral rows), and stomatogenesis begins with the proliferation of non‐ciliferous basal bodies some distance posterior to the buccal vertex.     

Balanion masanensis n. sp. (ciliophora: prostomatea) from the coastal waters of Korea: morphology and small subunit ribosomal RNA gene sequence

The planktonic ciliate Balanion masanensis n. sp. is described from living cells, from cells prepared by quantitative protargol staining (QPS), scanning electron microscopy (SEM), and transmitted electron microscopy (TEM) preparations, and the sequence of its nuclear small subunit rDNA (SSU rDNA) is reported. This species is almost ovoid with a flattened anterior oral region when the cells are alive and stained. The flattened anterior region of a living cell often forms a dome with the perimeter receded in a groove, and this region is easily inflated or depressed. In SEM photos, a brosse of six to nine monokinetids (or possibly three to five dikinetids) was observed inside the circumoral dikinetids. In TEM photos, circumoral microtubular ribbons were observed below the oral cilia, which along with the oral flaps were 8-16 microm in length. The cytostome is a slight funnel-like central depression on the flattened anterior end. The morphological characteristics of this ciliate are identical to those of the genus Balanion (Order Prorodontida). The ranges (and mean+/-standard deviation) of cell length, cell width, and oral diameter of living cells (n=23-26) were 27-43 microm (35.2+/-4.6), 25-32 microm (28.6+/-2.3), and 25-30 microm (27.6+/-1.3), respectively, while those of the QPS-stained specimens (n=70) were 23-37 microm (30.6+/-3.5), 26-35 microm (30.7+/-2.2), and 26-33 microm (29.5+/-1.5), respectively. Forty-six to 55 somatic kineties (SKs) were equally spaced around the cell body and extended from the oral to near the posterior regions with 24-50 monokinetids per kinety. Each kinetid bore a cilium 2.8-7.2 microm long. A caudal cilium (ca 14 microm long) arose on the posterior end. The single ellipsoid macronucleus is 6.8-13.4 x 6.8-10.5 microm, accompanied by a single micronucleus (2.0-2.8 x 1.5-2.5 microm) visible only in QPS specimens. Because, the cell size, the number of SKs, and the number of kinetosomes per SK of this ciliate were much greater than those of Balanion comatum and Balanion planctonicum, the only two Balanion species so far reported, we have established B. masanensis n. sp. When properly aligned, the sequence of the SSU rDNA of B. masanensis n. sp. (GenBank Accession No. AM412525) was approximately 9% different from that of Coleps hirtus (Colepidae, Prorodontida) and 12% different from that of Prorodon teres (Prorodontidae, Prorodontida).     

Ultrastructural aspects of the somatic and buccal infraciliature ofColeps amphacanthus Ehrenberg 1833

Summary Somatic and buccal infraciliature ofColeps amphacanthus Ehrenberg 1833 were studied by light and electron microscopy. The somatic kineties are composed of monokinetids and 2 dikinetids at the anterior end of each kinety. The monokinetids are associated with postciliary microtubules at triplet 9, a kinetodesmal fiber at triplet 5 and 7 and nearly radially arranged transverse microtubules at triplet 4. The associated fibrillar systems of the posterior kinetosome of the dikinetids are like those of the monokinetids. The anterior kinetosome is associated with transverse microtubules at triplet 4 and one or few postciliary microtubules at triplet 9. The anterior kinetosome bears only a short cilium.The oral ciliature is composed of a kinety of nearly circumorally arranged paroral dikinetids and 3 adoral organelles at the ventral left side of the oral opening. Nematodesmata arising from the oral ciliature form the major component of the cytopharyngeal apparatus which is lined by microtubular ribbons of postciliary origin. The buccal cavity is surrounded by oral papillae which often contain toxicysts.     

Isolation and characterization of cholangiocyte primary cilia

Primary cilia are distinct organelles expressed by many vertebrate cells, including cholangiocytes; however, their functions remain obscure. To begin to explore the physiological role of these organelles in the liver, we described the morphology and structure of cholangiocyte cilia and developed new approaches for their isolation. Primary cilia were present only in bile ducts and were not observed in hepatocytes or in hepatic arterial or portal venous endothelial cells. Each cholangiocyte possesses a single cilium that extends from the apical membrane into the bile duct lumen. In addition, the length of the cilia was proportional to the bile duct diameter. We reproducibly isolated enriched fractions of cilia from normal rat and mouse cholangiocytes by two different approaches as assessed by scanning electron, transmission electron, and confocal microscopy. The purity of isolated ciliary fractions was further analyzed by Western blot analysis using acetylated tubulin as a ciliary marker and P2Y(2) as a nonciliary cell membrane marker. These novel techniques produced enriched ciliary fractions of sufficient purity and quantity for light and electron microscopy and for biochemical analyses. They will permit further assessment of the role of primary cilia in normal and pathological conditions.     

Development of ciliary bands in larvae of the living isocrinid sea lily Metacrinus rotundus

Embryos and larvae of an isocrinid sea lily, Metacrinus rotundus, are described by scanning electron microscopy. Around hatching (35 h after fertilization), the outer surface of the gastrula becomes ubiquitously covered with short cilia. At 40 h, the hatched swimming embryo develops a cilia‐free zone of ectoderm on the ventral side. By 3 days, the very early dipleurula larva develops a cilia‐free zone ventrally, densely ciliated regions laterally, and a sparsely ciliated region dorsally. At this stage, the posterior and anterior ciliary bands first appear: the former runs along a low ridge separating the densely from the sparsely ciliated epidermal regions, while the latter is visible, at first discontinuously, along the boundary between the densely ciliated lateral regions and the cilia‐free ventral zone. In the late dipleurula larva (5 days after fertilization), the anterior and posterior loops of ciliary bands are well defined. The transition from the dipleurula to the semidoliolaria larva occurs at 6 days as the posterior loop becomes rearranged to form incompletely circumferential ciliary bands. The larva becomes competent to settle at this stage. The arrangement of the ciliary bands on the semidoliolaria is maintained during the second week of development, while the larva retains its competence to settle. The larval ciliary patterns described here are compared with those of stalkless crinoids and eleutherozoan echinoderms. The closest morphological similarities are between M. rotundus and the basal eleutherozoan class Asteroidea.     

Fine Structure and Molecular Phylogeny of Parametopidium circumlabens (Ciliophora: Armophorea), Endocommensal of Sea Urchins

Metopid armophoreans are ciliates commonly found in anaerobic environments worldwide; however, very little is known of their fine structure. In this study, the metopid Parametopidium circumlabens (Biggar and Wenrich 1932) Aescht, 1980, a common endocommensal of sea urchins, is investigated for the first time with emphasis on transmission electron microscopy, revealing several previously unknown elements of its morphology. Somatic dikinetids of P. circumlabens have a typical ribbon of transverse microtubules, an isolated microtubule near triplets 4 and 5 of the anterior kinetosome, plus two other microtubules between anterior and posterior kinetosomes, a short kinetodesmal striated fiber and long postciliary microtubules. In the dikinetids of the perizonal stripe, the kinetodesmal fiber is very pronounced, and there is a conspicuous microfibrillar network system associated with the kinetosomes. A new structure, shaped as a dense, roughly cylindrical mass surrounded by microtubules, is found associated with the posterior kinetosome of perizonal dikinetids. The paroral membrane is diplostichomonad and the adoral membranelles are of the “paramembranelle” type. Bayesian inference and maximum‐likelihood analysis of the 18S‐rDNA gene unambiguously placed P. circumlabens as sister group of the cluster formed by ((Atopospira galeata, Atopospira violacea) Metopus laminarius) + Clevelandellida, corroborating its classification within the Metopida.     

Strombidinopsis jeokjo n. sp. (ciliophora: choreotrichida) from the coastal waters off western Korea: morphology and small subunit ribosomal DNA gene sequence

The planktonic ciliate Strombidinopsis jeokjo n. sp. is described from Quantitative Protargol-Stained (QPS) preparations, and the sequence of the small subunit rDNA (SSU rDNA) from cultured cells is reported. This species is ovoid and bluntly tapered towards the posterior. The ranges (and mean +/- standard deviation, n = 31) of cell length, cell width, and oral diameter of the QPS-stained specimens were 100-190 microm (149 +/- 25), 60-105 microm (79 +/- 13), and 55-80 microm (64 +/- 5), respectively. Fifteen to seventeen external oral polykinetids had oral membranelle cilia 20-35 microm long. Twenty-six to twenty-eight somatic kineties were equally spaced around the cell body and extended from the oral to the posterior regions with 23-44 dikinetids per kinety. Both kinetosomes of each kinetid bore cilia 3-7 microm long. Strombidinopsis jeokjo had two ovoid macronuclei of 25-38 microm x 12-15 microm. When properly aligned, the sequence of the SSU rDNA of S. jeokjo (GenBank Accession No. AJ628250) was approximately 2% different from that of an unidentified Strombidinopsis species (GenBank Accession No. AF399132-AF399135), the closest species in the SSU rDNA sequence.     

Ditrypanocystis sp. (Apicomplexa, Gregarinia, Selenidiidae): the mode of survival in the gut of Enchytraeus albidus (Annelida, Oligochaeta, Enchytraeidae) is close to that of the coccidian genus Cryptosporidium

A selenid gregarine Ditrypanocystis sp. (Apicomplexa, Gregarinia, Selenidiidae), harboring the gut lumen of the oligochaete Enchytraeus albidus, was studied by light and electron microscopy. The trophozoite of Ditrypanocystis sp. is attached to the gut wall with its apical end to be anchored eventually between enterocytes in the crypts. Simultaneously, between the surfaces of the parasite and the host cell a peculiar contact is formed made of membranous channels and vesicles of unknown origin, the host cell surface in the contact area lacking cilia. The trophozoite becomes progressively enclosed within a parasitophorous vacuole made of layers of fused ciliar membranes of enterocytes. The fused cilia may be a source of membranes lining channels and vesicles of the contact area. Such a mode of parasitophorous arrangements has never been described before for gregarines, however, it bears a some likeness with that of the coccidian genus Cryptosporidium (similarity and differences being discussed). With regard to some molecular phylogeny constructions, claiming the "sister" relationship between gregarines and the coccidian genus Cryptosporidium (Carreno et al., 1999; Leander et al., 2003), this common feature in host-parasite relationships enabled us to put forward an idea of a possible evolutionary route from extracellularity of gregarines to intracellularity of coccidia, as exemplified by species of Cryptosporidium.     

Microscopic investigation of three species of Diophrys (Ciliophora,Euplotida, Uronychiidae) from Japan,including Diophrys peculiaris nov. spec

Three species of Diophrys, D. peculiaris nov. spec., D. cf. scutum and D. oligothrix, isolated from the New Nagasaki Fishing Port, Nagasaki, Japan, were investigated using live observation and protargol impregnation. Diophrys peculiaris nov. spec. can be recognized by having two characteristic clusters of rod-like structures and two groups of dikinetids located on anterior dorsal portion of cell. Morphogenetic data show that this part of the life cycle basically proceeds as in congeners, except for the formation of dikinetids under the rod-like structures. In the opisthe, the origin of dikinetids under the rod-like structures is still unknown, but the old dikinetids under the rod-like structures may be retained by the proter. The Japanese population of Diophrys cf. scutum resembles other populations of D. scutum well except for moniliform macronuclear segments. Our populations of D. oligothrix correspond well with other populations in terms of general morphology and ciliary pattern, in particular the continuous dorsal kineties with loosely arranged cilia.     

The fine structure of the paralabial organelle in the rumen ciliate Ophryoscolex purkinjei Stein, 1858

The paralabial organelle of the rumen ciliate Ophryoscolex purkinjei, located on the ventral side of the ciliophor, is a highly specialized part of the somatic cortex. It consists of alternating rows of short modified cilia and thin pellicular folds which form a ridge-like structure. The central "top kinety" is composed of monokinetids which bear cilia with 9 + 2 axonemes and 2 microns in length. The top kinety is accompanied by a comb-shaped fold on its distal side and by a broad wedge-shaped fold on its proximal side. To both sides there follow two or three lateral kineties made of dikinetids. The anterior kinetosome of each pair bears a clavate cilium, only 0.5-0.7 micron in length and with a 9 + 0 axoneme while the cilium of the posterior kinetosome is even shorter. Lateral folds with numerous microtubules cover these lateral kineties and rows of barren basal bodies. The fine structure of this supposed sensory organelle show a basic pattern in four other ophryoscolecids, and its increasing complexity parallels the suggested phylogenetic line of evolution of these ciliates.     

Cortical ultrastructure of Coleps bicuspis Noland, 1925 and the phylogeny of the class Prostomatea (Ciliophora)

The ultrastructure of Coleps bicuspis Noland, 1925 is described. The ciliate is a typical prostomate: the somatic kinetid is a monokinetid with a postciliary ribbon at triple 9, a kinetodesmal fibril originating near triplets 5, 6, 7 and an apparently radial transverse ribbon at triplet 4. The oral area is circular and has three brosse kineties associated with it. The brosse kineties are composed of dikinetids whose anterior kinetosome bears a tangential transverse ribbon and whose posterior kinetosome bears the fibrillar associates typical of a somatic monokinetid. The oral dikinetids are oriented parallel to the circumference of the oral cavity, which is surrounded by oral papillae and oral ridges. Pairs of nematodesmata, originating from oral dikinetid kinetosomes, are typically triangular in transection. A phylogeny of rhabdophoran ciliates is presented using the mixed parsimony algorithm and is discussed with reference to the systematic revisions of the phylum Ciliophora.     

The Ultrastructure of Chaenea teres and an Analysis of the Phylogeny of the Haptorid Ciliates

Chaenea teres has typical haptorid ultrastructure. The somatic monokinetid has two transverse microtubular ribbons, an overlapping postciliary microtubular ribbon, and a laterally directed kinetodesmal fiber. The evered cytopharynx forms a dome at the apical end of the cell. The base of the dome is surrounded by oral dikinetids. The left, anterior kinetosome of the oral pair is not ciliated and has a transverse microtubular ribbon, a nematodesmata and a single postciliary microtubule. The right, posterior kinetosome is ciliated and has only postciliary microtubules. The kinetosomes at the anterior ends of the somatic kinetics are close together and their transverse microtubules and nematodesmata contribute to the support of the cytopharynx. The transverse microtubules of these oralized somatic kinetosomes, together with those from the oral dikinetids, line the cytopharynx. Accessory or bulge microtubules arise perpendicular to the transverse microtubules. A dorsal brush of three kineties of clavate cilia is found on the cell surface just posterior to the oral region. Mucocysts and a single type of toxicyst are present. The toxicysts are confined to the oral region. There are multiple ovoid macronuclei that stain weakly. Micronuclei were not observed. Cladistic analysis indicates the Chaenea may be most closely related to Fuscheria and Acropisthium. The cladistic analysis also suggests that existing taxonomies of the subclass Haptoria need to be revised. We propose some modifications to Foissner & Foissner's classification that include transferring Helicoprorodon, Actinobolina, the buetschiliids, and the balantidiids to the order Haptorida and recognizing the close relationship between pleurostomes and spathidiids.     

Ultrastructural Variability in the Locomotor Cortex of the Ciliated Protozoa, Mytilophilus pacificae

ABSTRACT. Mytilophilus pacificae is an endocommensal ciliate found in the mantle cavity of the Pacific Coast mussel Mytilus californianus. In this paper we report our findings on pellicular organization of this species. Transmission and scanning electron microscope examination of the somatic cortex revealed that a number of different types of kinetids, i.e. monokinetids, dikinetids, and polykinetids are found in the locomotor cortex. The type and distribution of the kinetids are described. Surprisingly, the locomotor region was found to be highly variable among individuals with respect to its kinetid distribution; each cell appears to have its own characteristic kinetid pattern. Some cells have mostly monokinetids and dikinetids in their locomotor cortex, while others may have dikinetids and polykinetids but very few monokinetids. In contrast to the locomotor region, the thigmotactic field (a region specialized for attachment) is exclusively composed of dikinetids and shows no heterogeneity. The finding of ultrastructural variability in the locomotor cortex was unexpected since, in the view of the structural conservation hypothesis, the somatic cortex is seen as a "stable" element. These observations raise new questions with regard to cortical pattern formation in this organism.     

Fasciola hepatica: surfaces involved in movement of miracidia and cercariae

Rapid freezing and substitution with fixative prior to scanning electron microscopy was used to demonstrate the pattern of beat and recovery of the cilia of free swimming miracidia of Fasciola hepatica. There were stages of dexioplectic metachronal co-ordination and the power stroke was approximately 15 degrees anticlockwise from the anterior-posterior axis. Around the circumference of the body of the miracidia there were approximately 12 metachronal waves of power and recovery. Free-swimming cercariae were recorded by time-lapse photography and, after conventional fixation, by scanning electron microscopy. Cercarial tail-beats were to the posterior of the body in the lateral plane at a rate of 8 Hz. The tail has paired lateral ridges positioned to act as leading edges. There is an array of 32 sensory papillae on the mid-ventral surface of the tail. The tegument of the most distal part of the tail is described: it is free of sensory endings and the surface shows a spiral pattern.     

The Fine Structure of Saprodinium dentatum Lauterborn, 1908 as a Representative of the Odontostomatida (Ciliophora)

The fine structure of the sapropelic ciliate Saprodinium dentatum is described based on phase-contrast microscopy, silver-staining techniques, cryo-fracture scanning electron microscopy, and thin sections. The study concentrates on a detailed analysis of the somatic cortex and the oral ciliature of this highly asymmetric, laterally compressed ciliate. The cell shape is dominated by a number of site-specific spines and the curving course of 10 somatic kineties (SK 1–10). The SK, composed of dikinetids, show an intrakinety differentiation that seems characteristic for other odontostomes as well. The anterior segment of the SK is mostly ciliated, followed by a non-ciliated segment in which the kinetosomes lack all typical fiber systems. Except for SK 4–6, the posterior segment is ciliated again, forming the spine kinetics associated with particular caudal spines. The anterior segment of SK 3 through SK 7 form the frontal band, which together with the two frontal kineties constitutes the main locomotory organelle for a ciliate that creeps on the substratum. A short kinety with inverse polarity, not seen in earlier light microscopical studies, was observed near the oral spine. We made particular effort to find a logical explanation for the observed association of the SK with the various caudal spines. The oral ciliature consists of nine adoral organelles located in a tripartite oral cavity. The absence of a paroral ciliature together with the position of the cytostome anterior to the adoral organelles may be the result of rotational movement of the oral apparatus during the evolution of these bizarre ciliates. Results are discussed with special reference to the phylogenetic relationship of the Odontostomatida to the Heterotrichida and no conclusive answer was found in this first electron microscopical study of an odontostomatid ciliate.     

Revision of the Genus Askenasia Blochmann, 1895, with Proposal of Two New Species, and Description of Rhabdoaskenasia minima N. G., N. Sp. (Ciliophora, Cyclotrichida)

ABSTRACT. The planktonic ciliate genus Askenasia Blochmann, 1895 is reviewed and the new genus Rhabdoaskenasia n. gen. is established. Askenasia is characterized by three circumferential kinety belts and a circumoral wreath of paired argyrophilic granules without recognizable cilia and nematodesmata. A "brush" is absent. Askenasia apparently lacks the key characters of the Haptorida and is thus transferred to the Cyclotrichida, family Mesodiniidae. Rhabdoaskenasia differs from Askenasia in having single files of basal bodies in all kinety belts and club-shaped extrusomes. It possesses a circumoral kinety composed of dikinetids from which nematodesmata originate, forming a distinct rhabdos. Although very similar to Askenasia in its general appearance, R. minima n. sp. could belong to another order. Based on an extensive review of the literature and on silver impregnated specimens the following Askenasia species are recognized and described in detail: A. volvox (Eichwald, 1852) Kahl, 1930, A. stellaris (Leegaard, 1920) Kahl, 1930, A. acrostomia n. sp., and A. chlorelligera n. sp. Askenasia faurei Kahl, 1930 and A. humilis Gajewskaja, 1928 are transferred to the genus Cyclotrichium: C. faurei (Kahl, 1930) n. comb., C. humilis (Gajewskaja, 1928) n. comb. The systematic position of the genus Askenasia is discussed and keys to the genera of the Mesodiniidae and to the species of Askenasia are provided.     

Changes of the lingual epithelium in Ambystoma mexicanum

Changes in the lingual epithelium during ontogenesis and after induced metamorphosis in Ambystoma mexicanum are described as observed by light microscopy and scanning electron microscopy. The epithelium of the tongue is always multilayered in the larva as well as in the adult. It consists of a stratum germinativum with little differentiated basal cells and a stratum superficiale (superficial layer) with specialized superficial cells and goblet cells. Usually, there are more than two layers because of a stratum intermedium consisting of replacement cells. The apical cell membrane of the superficial cells is perforated by fine pores. Its most typical feature are microridges. Maturing superficial cells possess microvilli. Goblet cells occur in early larvae primarily in the centre of the tongue. They spread throughout the dorsal face of the tongue as their numbers increase during ontogenesis. The small apices of the goblet cells are intercalated in the wedges between the superficial cells. Leydig cells are not found on the larval tongue but on that of adults. Due to metamorphosis, the epithelium of the tongue changes. It is furrowed in its anterior part. The furrows house the openings of the lingual glands. The surface is further modulated by ridges which are densely coated by microvilli and which bear the taste buds. The villi of the tongue which lack extrusion pores show cilia and microvilli but lack microridges. The Leydig cells disappear during metamorphosis. In addition to the two types of goblet cells found in different regions of the glandular tubules, goblet cells occur in the caudal part. They secrete directly into the cavity of the mouth. The posterior part is characterised by a dense coat of cilia.