Khaled A. S. Al-Rasheid

The urostylid family Pseudokeronopsidae Borror and Wicklow, 1983 was considered to be a well-outlined taxon. Nevertheless, recent evidence, including morphological, ontogenetic, and molecular information, has consistently revealed the polyphyly of this family. In the present work, a new population of Thigmokeronopsis stoecki Shao et al., 2008 was found and its binary divisional process was described for the first time. In addition, the morphogenetic features of Thigmokeronopsis species and all the other pseudokeronopsids, for which detailed ontogenetic data are available, were rechecked and compared. This reveals that: (1) the ontogenetic process of T. stoecki corresponds well with its congeners T. jahodai and T. rubra except for the macronuclear behavior; (2) Apokeronopsis and Thigmokeronopsis share a similar ontogenetic mode despite of the differences in the number and origin of their buccal cirri; (3) most pseudokeronopsids share the same pattern in the origins of their oral primordia and fronto-ventral-transverse cirral anlagen, except for Pseudokeronopsis similis, which may not be a valid member of the family Pseudokeronopsidae.
 

The morphology, the infraciliature, some stages of cell division and physiological reorganization, and the SSU rRNA gene sequence of the little-known marine 18-cirri hypotrich Tachysoma rigescens (Kahl, 1932) Borror, 1972 [basionym Oxytricha (Tachysoma) rigescens], isolated from mariculture waters near Qingdao, China, were investigated. This rare species is characterized, inter alia, by narrowly spaced, small, colourless cortical granules and several conspicuous ring-shaped structures in the cytoplasm. The caudal cirri and the simple dorsal kinety pattern (three bipolar kineties) are probably plesiomorphic traits within the Hypotricha, the composition of the adoral zone of the proter from new and parental membranelles, as well as the presence of two ‘extra’ cirri behind the right marginal row strongly suggest a misclassification in Tachysoma. The SSU rRNA gene sequence data indicate that T. rigescens branches off rather basally in the Hypotricha tree, which supports the hypothesis that the 18-cirri pattern occurred very early, probably already in the last common ancestor of the Hypotricha. A detailed survey of the early branching 18-cirri hypotrichs and similar taxa (e.g. Trachelostyla pediculiformis, Hemigastrostyla enigmatica, Protogastrostyla pulchra) reveals that for T. rigescens a new genus (Apogastrostyla gen. nov.) has to be established, because there are important differences, inter alia, in the dorsal infraciliature. Besides the type species, A. rigescens comb. nov., which seems to be confined to the northern hemisphere according to the sparse faunistic data, a second marine species, A. szaboi comb. nov. (basionym Hemigastrostyla szaboi), so far only twice recorded from the Antarctic region, can be included. The Chinese population is fixed as neotype to define the species objectively, because no type material of A. rigescens is present and the original type locality is not known. The species name Tachysoma multinucleate is emended: Tachysoma multinucleatum nom. corr.

Ten new species of myxosporeans found from marine fishes were collected from coastal waters off the Yellow Sea in China: Sphaerospora sebasta sp. n. coelozoic in the gall bladder of Sebastes schlegeli, Ceratomyxa hemitriptera sp. n. coelozoic in the gall bladder of Hemitripterus villosus,Ceratomyxa kareus sp. n. coelozoic in the gall bladders of Kareius bicoloratus and Zebrias zebra,Ceratomyxa lateolabrax sp. n. and Ceratomyxa lomi sp. n. coelozoic in the gall bladder ofLateolabrax japonicus, Ceratomyxa qingdaoensis sp. n. coelozoic in the urinary bladder ofArgyrosomus argentatus, Ceratomyxa saurida sp. n. coelozoic in the gall bladder of Saurida elongata, Ceratomyxa sebastisca sp. n. coelozoic in the gall bladder of Sebastiscus sp., Ceratomyxa simplex sp. n. coelozoic in the gall bladder of Chirolophis japonicus and Ceratomyxa triacantha sp. n. coelozoic in the gall bladder and bile of Triacanthus brevirostris. All those forms were described in a book chapter cited as “known forms” several years ago, but have never been formally established as new taxa which are thus officially reported here. The present contribution only provided the morphology and geographic distribution of these organisms.

The morphology and morphogenesis of two populations of the soil hypotrichous ciliate, Parakahliella macrostoma (Foissner, 1982) Berger et al. 1985, isolated from northwest China, were investigated based on specimens examined in vivo and stained with protargol. Our populations resemble the original one in terms of their live characters and cirral pattern. The main events during binary fission are as follows: (1) the parental adoral zone of membranelles is retained completely by the proter; (2) both in the proter and in the opisthe five frontal-ventral cirral anlagen are recognizable; (3) the marginal rows and dorsal kineties develop intrakinetally. In addition, the SSU rRNA gene was sequenced for the genus Parakahliella for the first time. Molecular phylogenetic analyses suggest that two populations of the genus Parakahliella cluster together and have a close relationship with species of Oxytrichidae. 

The largest known Aspidisca species, A. magna Kahl, 1932, was found from coastal waters near Qingdao (Tsingtao), northern China, and investigated using both the “wet” silver nitrate and protargol staining method. Based on the living observation and impregnated individuals, improved diagnosis and morphometric data are provided. As an additional contribution, the morphogenesis in Aspidisca magna during binary fission was revealed and summarized as follows: 1) the parental adoral zone of membranelles and paroral membranęare retained by the proter; 2) the oral primordium of the opisthe develops hypoapokinetally behind the posterior part of the adoral zone of membranelles; 3) five frontoventral-transverse cirral anlagen are formed de novo, initially as primary primordia, and develop into 3:3:2:2:1 cirri from left to right, respectively in both dividers; 4) the leftmost frontoventral cirrus is generated from an independently formed cirral anlage in both dividers. In the present work, Chinese and Korean populations of A. leptaspis Fresenius, 1865 were investigated respectively to support the importance of living characteristics in identification of Aspidisca species. The isolation of A. magna and A. leptaspis from other congeners are also firmly demonstrated by the SSU rRNA gene sequence alignments.

The morphology and infraciliature of five trachelocercid ciliates: Tracheloraphis huangi spec. nov., T. colubis (Kahl, 1933) comb. nov., T. phoenicopterus (Cohn, 1866) Dragesco, 1960, T. oligostriata (Raikov, 1962) Foissner and Dragesco, 1996 and Trachelocerca incaudata Kahl, 1933, isolated from the intertidal zone of a beach at Qingdao, China, were studied in live and protargol impregnated specimens. Tracheloraphis huangi spec. nov. was distinguished from its congeners mainly by its single nuclear group composed of 25–30 round macronuclei and 29–37 somatic kineties. The poorly known T. colubis is redescribed including for the first time information on its infraciliature. An improved diagnosis is also provided. In light of its infraciliature, T. colubis is transferred to Tracheloraphis from the genus Trachelocerca. Additional data on other three species is supplied based on the Qingdao populations.

This paper investigates the morphology and morphogenesis during binary fission of a Chinese population of Euplotes amieti Dragesco, 1970, a fresh water form which has previously not been well defined. This organism is morphologically very similar to the well-known Euplotes eurystomus but differs from the latter both in the number of dorsal kineties and the molecular data. According to the information obtained, it is characterized by a combination of features including nine frontoventral cirri, ca. 60 membranelles, 12–15 dorsal kineties, a macronucleus in the shape of the number 3, and a ‘double-eurystomus’ type of silverline system. Its morphogenesis proceeds broadly in the same pattern as in its congeners. In this study, the SSU rRNA gene was sequenced for the first time, and phylogenetic analyses indicated that it is closely related to the eurystomus-aediculatus-woodruffi- complex. Considering the extreme similarities in morphology between E. amieti and E.eurystomus, we believe that the four sequences (four isolates) under the name of Euplotes eurystomus (No. FR873716; FR873717; EF193250; AJ310491 deposited in GenBank) are very likely from misidentified material; that is, they represent different populations of Euplotes amieti.

The morphology and infraciliature of four marine scuticociliates, Pleuronema elegans spec. nov., P. setigerum Calkins, 1902, P. grolierei Wang et al., 2008 and Uronema orientalis spec. nov., collected from China seas, were investigated through live observation and protargol staining methods. Pleuronema elegans spec. nov. can be recognized by the combination of the following characters: size in vivo 90–115 × 45–60 µm, slender oval in outline with a distinctly pointed posterior end; about 10 prolonged caudal cilia; consistently two preoral kineties and 18 or 19 somatic kineties; membranelle 2a double-rowed with its posterior end straight; membranelle 3 three-rowed; one macronucleus; marine habitat.Uronema orientalis spec. nov. is distinguished by the following features: in vivo about 40–55 × 20–30 μm with a truncated apical plate; consistently twenty somatic kineties; membranelle 1 single-rowed and divided into two parts which comprise four and three basal bodies respectively; contractile vacuole pore positioned at the end of the second somatic kinety; marine habitat. We also provide improved diagnoses for P. grolierei Wang et al., 2008 and P. setigerum Calkins, 1902 based on current and previous reports. The small subunit rRNA gene of U. orientalis, P. elegans, P. grolierei and P. puytoraci were sequenced. Phylogenetic analyses indicate that Uronema and Pleuronema are not monophyletic.

The morphology, infraciliature, and silverline system of three marine scuticociliates, Uronema marinum Dujardin, 1841, U. heteromarinum nov. spec. and Pleuronema setigerum Calkins, 1902, isolated from coastal waters off Qingdao, China, were investigated using living observation and silver impregnation methods. Due to the great confusion in the species definition of the well-known species U. marinum, we have documented a detailed discussion/comparison and believe that most of the confusion is due to the fact that at least 2 closely-related sibling morphotypes exist which are often not recognized. Based on the data available, U. marinum is strictly defined as follows: marine Uronema ca. 30 × 10 μm in size, with truncated apical frontal plate and smooth pellicle, extrusomes inconspicuous, cytostome located equatorially, 12–14 somatic kineties and one contractile vacuole pore near posterior end of kinety 2. Uronema heteromarinum nov. spec. resembles U. marinum but can be distinguished morphologically by its notched pellicle with conspicuous extrusomes and reticulate ridges, the 15–16 somatic kineties, widely separated membranelle 1 and membranelle 2, as well as the subequatorially positioned cytostome. Based on the Qingdao population, an improved diagnosis for the poorly known Pleuronema setigerum is: marine slender oval-shaped form, in vivo about 40–50 × 15–20 μm; 3–5 preoral kineties and 14–22 somatic kineties; membranelle 1 and 3 three-rowed, and posterior end of M2a ring-like. The small subunit (SSU) rRNA gene for all three organisms were sequenced and analyzed with standard methods.

As a follow-up part of our studies of the tropical ciliate fauna carried out in the last decade, we investigated the morphology and phylogeny of three choreotrich ciliates, viz, Parastrombidinopsis costalis sp. n., P. pelagica (Fauré-Fremiet, 1924) comb. n. and P. minima Tsai et al., 2008. The new species is characterized by its unique asymmetrical cell shape, 19–21 collar membranelles, one buccal membrane, and eight unevenly distributed somatic kineties.An improved diagnosis of P. pelagica(Fauré-Fremiet, 1924) comb. n.is given based on the original and current studies, the species is characterized by a large cell size, an elongate obconical cell shape, 31–36 collar membranelles and 13–15 somatic kineties. The species P. minimais redescribed based on a new population with some new features supplemented.SSU rRNA genes of both the new species and P. minima were sequenced and a phylogenetic review of related taxa obtained has been performed in order to reveal their systematic relationships. The monophyly of the genus Parastrombidinopsis is highly supported in our phylogenetic analyses.

During faunistic studies of ciliates in coastal waters of Daya Bay and Bohai Bay, China, two previously unknown ciliates were discovered and investigated using standard taxonomic methods. Morphological comparative analyses revealed that they represent two novel species in the genusChaenea. Chaenea paucistriata spec. nov. can be distinguished from its congeners by the following traits: body length in vivo about 180–250 µm; eight somatic kineties; dorsal brush rows 1–4 consisting of three, five, seven, and two dikinetids, respectively; rod-like extrusomes, 8 µm long; 63–94 macronuclei; cortical granules minute and colourless. Chaenea sinica spec. nov. differs from its congeners in having: body length in vivo about 140–240 µm; 17–21 somatic kineties; dorsal brush rows 1–4 consisting of 3–7, 10 or 11, 11–13, and 3–6 dikinetids, respectively; rod-like extrusomes about 6–8 µm long; 71–164 macronuclei. A key is presented to assist the identification of allChaenea species.

The morphology and infraciliature of three marine cyrtophorid ciliates, Dysteria lanceolata Claparède and Lachmann, 1859, Lynchella nordica Jankowski, 1968 and Chlamydonyx paucidentatus Deroux, 1976, collected from the coastal waters of Qingdao, China, were investigated using live observations and the protargol impregnation method. D. lanceolata was oval in body outline, about 65 × 45 μm in vivo, with a subcaudally positioned podite, six to seven right kineties, two or three frontoventral kineties, and two ventral contractile vacuoles. The improved diagnosis for L. nordica and a key to all known Lynchella species are supplied. The rediscovery of C. paucidentatus enables us to accept its taxonomic identification (e.g., separation from a morphologically similar species Trochilioides recta). The genus Trochilioides Deroux, nov. gen., which was a nomen nudum according to ICZN (1999), is re-established.

The morphology, the infraciliature, and two stages of physiological reorganization of Hemigastrostyla elongata spec. nov.,isolated from the Yellow Sea near Qingdao (China), are described. The new species differs from the type H. stenocephala, inter alia, by the length of the dorsal bristles and the position of the pretransverse ventral cirri; from H. enigmatica by the number of caudal cirri; and from H. para-enigmatica spec. nov. – established for the H. enigmatica populations from the Yellow Sea – by the arrangement of the postoral ventral cirri and the cortical granulation. A key to the Hemigastrostyla species and some other 18-cirri hypotrichs is provided. Hemigastrostyla szaboi is fixed as type species of Heterooxytricha gen. nov. because the type population lacks the extra cirri which are characteristic for Hemigastrostyla. In addition, Oxytricha geleii is assigned to this new genus, whose species have, like many oxytrichids, 18 frontal-ventraltransverse cirri, but a Gonostomum dorsal kinety pattern. The old, large, and difficult genus Oxytricha is briefly reviewed, mainly on the basis of the dorsal kinety pattern. Very likely, only species with the Oxytricha pattern belong to this genus. Oxytricha marcili and O. pseudofurcata, which have the Urosomoida kinety pattern (i.e. kinety 3 fragmentation lacking), are transferred to Urosomoida which is, inter alia, defined by a more or less distinctly reduced number of ventral and transverse cirri. Some other Oxytricha species with this kinety pattern (O. islandica, O. lanceolata, O. pseudosimilis, O. setigera) are not transferred to Urosomoida, but preliminarily classified as incertae sedis in Oxytricha, because they have the full set of 18 cirri. The available molecular data on O. lanceolata indicate that this type of 18-cirri hypotrichs likely needs a genus of its own because O. lanceolata does not cluster with O. granulifera, type of this genus. The marine Actinotricha saltans, classified for a very long time in Oxytricha, seems to be a non-dorsomarginalian hypotrich according to molecular data, justifying the reactivation of the old genus Actinotricha. Oxytricha shii has a multiple dorsal kinety 3 fragmentation, three dorsomarginal rows, and the undulating membranes arranged in the Cyrtohymena pattern, strongly indicating that it is a member of the subgenus Cyrtohymena (Cyrtohymenides). This brief review is a further step to unravel the complicated systematics of the old, but still little-known genus Oxytricha. The following new combinations are made in this paper: Cyrtohymena (Cyrtohymenides) shii (Shi et al., 1997) comb. nov.; Heterooxytricha szaboi (Wilbert and Song, 2005) comb. nov.; Heterooxytricha geleii (Wilbert, 1986) comb. nov.; Urosomoida marcili (Paiva and Silva-Neto, 2004) comb. nov.; Urosomoida pseudofurcata (Berger, 1999) comb. nov.

The morphology and infraciliature of two marine scuticociliates, Pleuronema puytoraci Grolière and Detcheva, 1974, and Parauronema longum Song, 1995, collected from China, were investigated using live observation and protargol impregnation methods. Based on the data obtained for the China population, new information of the living morphology of Pleuronema puytoraci is documented and details of the complete infraciliature is available for the first time. The stomatogenesis of Parauronema longum is basically similar to that of its congeners and can be summarized as follows: membranelle 1, membranelle 2 and the scutica of the opisthe originate from the parental paroral membrane, whereas membranelle 3 of the opisthe develops from the parental scutica; the paroral membrane originates from the parental paroral membrane.

Living observation and silver impregnation methods were used to investigate the morphology and infraciliature of three Frontonia ciliates (F. guangdongensis spec. nov., F. ocularis Bullington, 1939 and F. schaefferi Bullington, 1939) that were isolated from coastal waters of the South China Sea. Frontonia guangdongensis spec. nov. may be recognized by the combination of the following characteristics: cells about 160 × 35 μm in vivo; elongated body with right margin depressed in anterior third; length to width ratio 4:1 to 5:1; three or four vestibular and four or five postoral kineties; peniculi 1 and 2 each with four rows of kineties, peniculus 3 with two rows; one contractile vacuole in mid-body region right of cell median; brackish water habitat. A key based on morphological data for fourteen marine or brackish water Frontonia species found in China is also provided. In addition, the small subunit (SSU) rDNA gene was sequenced for F. ocularis Bullington, 1939. Our phylogenetic analyses support the contention that the genus Frontonia is not monophyletic.

The morphology, morphogenesis and infraciliature of two marine euplotid ciliates, Euplotes dammamensis n. sp. and Euplotes balteatus (Dujardin, 1841) Kahl, 1932, isolated from a sandy beach of the Arabian Gulf, Saudi Arabia, were investigated using observations in vivo and protargol-impregnation methods. Euplotes dammamensis n. sp. is characterized by a combination of features including its huge body size (100–170 × 80–120 μm), 10 conspicuous dorsal ridges, 10 normal-sized frontoventral and two marginal cirri, and 11 dorsal kineties. Euplotes balteatus is mainly characterized by 10 frontoventral, two caudal, and two left marginal cirri, 7–10 dorsal kineties and 5–7 prominent dorsal ridges as well as double-eurystomus silverline system. The small subunit rRNA (SSU-rRNA) gene sequences were determined for both species and phylogenetic analyses based on these data indicated that E. dammamensis is most closely related to E. parabalteatus Jiang et al., 2010, and E. balteatus clusters with E. plicatum Valbonesi et al., 1997, E. orientalis Jiang et al., 2010, and E. bisulcatus Kahl, 1932.

The present work investigates the living morphology and infraciliature of three marine cyrtophorid ciliates, which were isolated from Qingdao, China. Compared with its congeners, Orthotrochilia sinica spec. nov. can be distinguished by a combination of features: body slender and elliptical in outline, size about 50–60 × 20–25 μm in vivo, 18–21 somatic kineties, the length of the left perioral kinety treble the length of the right one, two ventrally located contractile vacuoles on the right side, and 25–32 nematodesmal rods. Based on current observations and the previous description, the diagnosis of Trochilioides tenuis (Deroux, 1976) Chen et al. 2011 is improved: cell size 30–40 × 20–35 μm in vivo, oval shaped in outline; consistently three right kineties, four left kineties and seven postoral kineties; a single contractile vacuole; marine habitat. A second species of Trochilioides, T. recta (Kahl, 1923) Chen et al. 2011 is re-described based on a Chinese population. Furthermore, a key to the identification of species of the genus Trochilioides whose infraciliature data are available is supplied, and Chlamydonyx trivialis (Fenchel, 1965) comb. nov. [basionym: Trochilioides trivialis Fenchel, 1965] is suggested.

Eight marine scuticociliates, Pseudoplatynematum denticulatum (Kahl, 1933) nov. comb., Protocyclidium sinica nov. spec., Histiobalantium marinum Kahl, 1933, Porpostoma notata Möbius, 1888, Philaster hiatti Thompson, 1969, Parauronema longum Song, 1995, Uronemella parafilificum Gong et al., 2007, and Paranophrys magna Borror, 1972, collected from Chinese coastal waters, were investigated using live observations and silver impregnation methods. Investigations of a Chinese population of Platynematum denticulatum (Kahl, 1933) reveal that it has a highly strengthened pellicle and distinct spines and thus corresponds well with the definition of Pseudoplatynematum Bock, 1952. A new combination, Pseudoplatynematum denticulatum (Kahl, 1933) nov. comb., is therefore proposed and an improved species diagnosis is supplied. Protocyclidium sinica nov. spec. is characterized by: small body size with buccal field approximately 60% of body length; extrusomes present; 13 or 14 somatic kineties; somatic kinety 1 comprising approximately 24 densely arranged kinetids; somatic kinety n shortened posteriorly; single macronucleus. Additional information is documented on the morphology of six other species of scuticociliates based on the China populations.