Taxonomy,morphology and molecular systematics of a new oligotrich ciliate,Williophrya maedai gen. nov., sp. nov., with redescriptions of Strombidium basimorphum and Pseudotontonia simplicidens (Protozoa,Ciliophora, Oligotrichia)

The morphology and small subunit rRNA (SSrRNA) gene sequence of a marine oligotrich ciliate, Williophrya maedai gen. nov., sp. nov., are reported. The new genus Williophrya is characterized by the adoral zone with no differentiation of membranelles, and the reduced somatic ciliature which comprises a bipartite girdle kinety only. In addition, the in vivo morphologies of two other oligotrichs, namely Strombidium basimorphum Martin & Montagnes, 1993 and Pseudotontonia simplicidens (Lynn & Gilron, 1993 Lynn, D. H. and Gilron, G. L. 1993. Strombidiid ciliates from coastal waters near Kingston Harbour, Jamaica (Ciliophora, Oligotrichia, Strombidiidae). Journal of the Marine Biological Association of the United Kingdom, 73: 47–65. [Crossref] , [Google Scholar]) Agatha, 2004, are reported for the first time based on Chinese populations. Improved diagnoses of both species are supplied. The phylogenetic position of Williophrya maedai is investigated based on SSrRNA gene sequence data. These show that: (1) Williophrya maedai is most closely related to Strombidium purpureum and S. apolatum; (2) Williophrya is assigned to the family Strombidiidae although it has some unique morphological features regarding its oral and somatic ciliatures.     

Taxonomy and molecular systematics of three oligotrich (s.l.) ciliates including descriptions of two new species,Strombidium guangdongense sp. nov. and Strombidinopsis sinicum sp. nov. (Protozoa,Ciliophora)

In this study we investigated the morphology of three oligotrich (s.l.) ciliates, Strombidium guangdongense sp. nov., Cyrtostrombidium paralongisomum Tsai et al., 2015 and Strombidinopsis sinicum sp. nov. Strombidium guangdongense sp. nov. is characterized by its elongate obconical to obovoidal body shape and widely spaced dikinetids in the girdle and ventral kineties. Another new species, Strombidinopsis sinicum sp. nov. is diagnosed by its small size and semi-globular body shape without mineral envelopes. Some additional morphological data of the recently described species Cyrtostrombidium paralongisomum Tsai et al., 2015, such as the endoral membrane, are supplied based on our population. We also analysed the molecular phylogeny of each species based on small subunit rRNA (SSU rRNA) gene sequence data. The monophyly of Cyrtostrombidium is supported by our phylogenetic analyses, but the monophyly of Strombidinopsis and of the family Strombidinopsidae are both rejected by AU tests. In addition, Strombidium species have a tail branch separately from one another in phylogenetic trees, whereas strombidiids with a pigment spot group together, suggesting the latter character is a synapomorphy for this group of strombidiids.

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Taxonomy and phylogeny of two poorly studied genera of marine oligotrich ciliates including descriptions of two new species: Cyrtostrombidium paraboreale sp. n. and Apostrombidium orientale sp. n. (Ciliophora: Spirotrichea)

Although it is widely recognized that oligotrich ciliates are the dominant constituent of microzooplankton communities and perform key functions in energy flow and material cycling in marine microbial food webs, knowledge of their diversity is scant. In the present study, we investigate the oligotrich genera, Cyrtostrombidium and Apostrombidium, with emphasis on their morphology and evolutionary relationships. Three isolates were collected from coastal waters of northern and southern China including two new species, viz., Cyrtostrombidium paraboreale sp. n., Apostrombidium orientale sp. n., and Apostrombidium pseudokielum Xu et al., 2009. Cyrtostrombidium paraboreale sp. n. is characterized by possessing 64–98 cytopharyngeal rods and two macronuclear nodules. Apostrombidium orientale sp. n. is characterized by its somatic kinety consisting of five fragments including a horizontally orientated subterminal fragment and possessing conspicuously long dorsal cilia. Apostrombidium pseudokielum is redescribed based on the new population and a re-examination of the type material. Phylogenetic analyses were performed for the subclass Oligotrichia, incorporating SSU rRNA gene sequences of the three species investigated here. The results indicate that the genus Cyrtostrombidium is monophyletic with C. paraboreale sp. n. occupying the basal position. The genus Apostrombidium is not monophyletic as Varistrombidium kielum is nested within it.     

The Morphology of Three Loxophyllum Species (Ciliophora,Pleurostomatida) from Southern China,L. lembum sp. n., L. vesiculosum sp. n. and L. perihoplophorum Buddenbrock, 1920, with Notes on the Molecular Phylogeny of Loxophyllum

Two new pleurostomatid ciliates, Loxophyllum lembum sp. n., L. vesiculosum sp. n., and the poorly known L. perihoplophorum Buddenbrock, 1920, isolated from brackish waters in coastal regions of southern China, are described following observations of live cells and protargol‐impregnated specimens. Loxophyllum lembum sp. n. is distinguished by a combination of characters including two macronuclear nodules, 6–9 contractile vacuoles along the ventral margin, 11–14 right and 6–8 left kineties and the presence of cortical granules. Loxophyllum vesiculosum sp. n. differs from its congeners mainly by the unique distribution of contractile vacuoles, several of which lie along the dorsal margin and one on the ventral margin, and 15–21 right and 6–8 left kineties. Loxophyllum perihoplophorum is characterized by its large cell size (350–450 μm long in vivo), 3–5 contractile vacuoles along the dorsal margin in the posterior region of the body, and 19–23 right and 7–9 left kineties. An improved diagnosis of L. perihoplophorum is provided. The SSU rDNA sequence of L. perihoplophorum is reported for the first time and its molecular phylogeny is analyzed. Maximum likelihood and Bayesian inference analyses of SSU rDNA sequence data recover the monophyly both of the order Pleurostomatida and of the genus Loxophyllum.     

Morphological Characterizations of Four Species of Parallelostrombidium (Ciliophora,Oligotrichia), with a Note on the Phylogeny of the Genus

The morphology and phylogeny of four oligotrichid ciliates, Parallelostrombidium paraellipticum sp. n., P. dragescoi sp. n., P. jankowskii (Xu et al. 2009) comb. n., and P. kahli (Xu et al. 2009) comb. n., are described or redescribed based on live observation, protargol stained material, and SSU rRNA gene sequences. The new species P. paraellipticum sp. n. is characterized by its obovoidal cell shape, adoral zone composed of 17–21 collar, 9–11 buccal, and two thigmotactic membranelles, and extrusomes attached in one row along the girdle kinety. The new species P. dragescoi sp. n. is distinguished from its congeners by its obovoidal cell shape and a lack of thigmotactic membranelles. Based on ciliary patterns recognizable in the original slides, Omegastrombidium jankowskii Xu et al. 2009 and O. kahli Xu et al. 2009 should be transferred to the genus Parallelostrombidium Agatha 2004. Phylogenetic analyses based on SSU rRNA gene sequence data demonstrate that all four new sequences cluster with previously described congeners. The genus Parallelostrombidium is separated into two clusters, suggesting its non‐monophyly and probably corresponding to the two subgenera proposed by Agatha and Strüder‐Kypke (2014), as well as their morphological difference (cell dorsoventrally flattened vs. unflattened).     

Morphology and molecular phylogeny of two new species of Spirostrombidium (Ciliophora,Oligotrichia), with a key to the species of Spirostrombidium

Microscopic methods were used to investigate the morphological characterization of two novel oligotrich ciliates, Spirostrombidium paraurceolare sp. nov. and Spirostrombidium faurefremieti sp. nov., isolated from a mangrove wetland in Zhanjiang and an intertidal sandy beach in Qingdao, respectively. Spirostrombidium paraurceolare sp. nov. is characterized by three thigmotactic and 8–10 buccal membranelles, the girdle kinety spiralling around cell with one and a half whorls, and located at right anterior third of dorsal side anteriorly. Spirostrombidium faurefremieti sp. nov. can be recognized by a prominently deep and broad buccal cavity, two thigmotactic and 15–19 buccal membranelles, and the girdle kinety spiralling around cell with two whorls. The small subunit ribosomal RNA genes of these two species were sequenced and compared with those of their congeners to reveal nucleotide differences. Phylogenetic analyses revealed that the genus Spirostrombidium is non-monophyletic. Spirostrombidium faurefremieti sp. nov. falls into a clade comprising most congeners, but Spirostrombidium paraurceolare sp. nov. branches off and groups with Varistrombidium kielum with moderate support. A key to the identification of Spirostrombidium species is also provided.www.zoobank.org/urn:lsid:zoobank.org:pub:AB96BEE6-BE3A-4B95-B75A-3469B1C53ABB2     

Redefinition of the hypotrichous ciliate Uncinata,with descriptions of the morphology and phylogeny of three urostylids (Protista,Ciliophora)

We investigated the morphology, morphogenesis and small subunit rRNA gene-based phylogeny of three marine urostylids, Uncinata gigantea Bullington, 1940 Bullington, W. E. (1940). Some ciliates from Tortugas. Papers from the Tortugas Laboratory, 32, 179–221. [Google Scholar], Holosticha heterofoissneri Hu & Song, 2001 Hu, X., & Song, W. (2001). Morphology and morphogenesis of Holosticha heterofoissneri n. sp. from the Yellow Sea, China (Ciliophora, Hypotrichida). Hydrobiologia, 448, 171–179. doi:10.1023/A:1017553406031.[Crossref], [Web of Science ®] , [Google Scholar], and Holosticha cf. heterofoissneri. The dorsal morphogenesis of Uncinata gigantea shows de novo formation of two groups of anlagen near the marginal rows. Holosticha cf. heterofoissneri demonstrates fragmentation of the first dorsal kinety anlage as in Holosticha heterofoissneri. Our population of H. heterofoissneri corresponds well with previously described populations in terms of its general morphology and ciliary pattern. Uncinata gigantea can be recognized by its large and highly contractile body, yellowish to brownish cell colour, two types of cortical granules, and 20–30 transversely oriented and densely arranged cirri in the left marginal row, which often overlie the buccal vertex. Based on the new data, especially infraciliature, the genus Uncinata is here redefined. Both the morphology and phylogenetic analyses suggest that the genus Uncinata should be classified within the family Urostylidae. In addition, both morphological and morphogenetic data suggest that Holosticha bradburyae Gong et al., 2001 Gong, J., Song, W., Hu, X., Ma, H., & Zhu, M. (2001). Morphology and infraciliature of Holosticha bradburyae n. sp. (Ciliophora, Hypotrichida) from the Yellow Sea, China. Hydrobiologia, 464, 63–69. doi:10.1023/A:1013901621439.[Crossref], [Web of Science ®] , [Google Scholar] should be transferred to Uncinata as U. bradburyae (Gong et al., 2001 Gong, J., Song, W., Hu, X., Ma, H., & Zhu, M. (2001). Morphology and infraciliature of Holosticha bradburyae n. sp. (Ciliophora, Hypotrichida) from the Yellow Sea, China. Hydrobiologia, 464, 63–69. doi:10.1023/A:1013901621439.[Crossref], [Web of Science ®] , [Google Scholar]) comb. nov., due to its possession of a characteristically prominent beak-like, leftwards curved projection and the developmental mode of the dorsal kineties. This assignment is supported by the phylogenetic analyses, which placed Uncinata gigantea in a clade with U. bradburyae (Gong et al., 2001 Gong, J., Song, W., Hu, X., Ma, H., & Zhu, M. (2001). Morphology and infraciliature of Holosticha bradburyae n. sp. (Ciliophora, Hypotrichida) from the Yellow Sea, China. Hydrobiologia, 464, 63–69. doi:10.1023/A:1013901621439.[Crossref], [Web of Science ®] , [Google Scholar]) comb. nov., and revealed only 1.13% (19 bp) difference in their SSU-rDNA gene sequence.     

Morphology and small subunit rDNA phylogeny of a new soil ciliate, Bistichella variabilis n. sp. (Ciliophora, Stichotrichia)

We isolated a new ciliate, Bistichella variabilis n. sp., from slightly saline soil of the Yellow River Delta in China and studied it using live observation, protargol staining, and small subunit (SSU) rRNA gene sequencing. Bistichella variabilis is elongated ellipsoidal and about 220 × 70 μm in vivo, with a variable number and arrangement of cirri- ventral row VI mostly very short, but a long row observed also in one specimen, and an extra row occasionally exists besides the left or the right marginal row. Bistichella variabilis differs from all congeners except Bistichella buitkampi (Foissner, 1982) in the macronuclear pattern (four vs. two or many nodules); and from B. buitkampi by the usual oblique arrangement of transverse cirri (vs. perpendicular); and the ventral row VI mostly restricted in the posterior quarter of the cell (vs. long and stretching to the anterior quarter). Phylogenetic analyses of SSU rDNA sequences indicated that B. variabilis grouped with the clade of Orthoamphisiella breviseries and Uroleptoides magnigranulosus with high support values. Despite their distinct differences in morphology, these three species have sequence similarities of 99.5-99.7%. Nonetheless, it is still premature to assign Bistichella together with Uroleptoides to the family Orthoamphisiellidae because of the taxonomic uncertainties and undersampling.     

Candida dajiaensis sp. nov., Candida yuanshanicus sp. nov., Candida jianshihensis sp. nov., and Candida sanyiensis sp. nov., four anamorphic, ascomycetous yeast species isolated from soil in Taiwan

Nine anamorphic, ascomycetous yeast strains belonging to the Pichia anomala clade were recovered from forest soil in 2006 in Taiwan. The nine yeast strains represent four novel yeast species based on the sequences of their D1/D2 domain of the large subunit (LSU) rRNA gene and their physiological characteristics. The scientific names of Candida dajiaensis sp. nov., Candida yuanshanicus sp. nov., Candida jianshihensis sp. nov., and Candida sanyiensis sp. nov. are proposed for these novel yeast species. The type strains are C. dajiaensis SM11S03(T) (=CBS 10590(T)=BCRC 23099(T)), C. yuanshanicus SY3S02(T) (=CBS 10589(T)=BCRC 23100(T)), C. jianshihensis SM8S04(T) (=CBS 10591(T)=BCRC 23096(T)), and C. sanyiensis SA1S06(T) (=CBS 10592(T)=BCRC 23094(T)). Sequence analysis of the D1/D2 of the LSU rRNA gene revealed that the three species, C. dajiaensis, C. yuanshanicus and Pichia onychis, shared a separate branch in the phylogenetic tree, C. jianshihensis is phylogenetically related to Candida ulmi and Pichia alni, and the phylogenetically closest relative of C. sanyiensis is Pichia populi.     

New SSU-rDNA sequences for eleven colpodeans (Ciliophora,Colpodea) and description of Apocyrtolophosis nov. gen

Using 11 new SSU-rDNA sequences, we analyze relationships within the class Colpodea, especially of some uncommon taxa, such as Kalometopia duplicata, Cyrtolophosis minor, and Jaroschia sumptuosa. The sequences do not change the basic structure of the molecular Colpodea tree, i.e., all belong to one of the four molecular clades recognized by Foissner et al. (2011): Colpodida, Cyrtolophosidida, Bursariomorphida, and Platyophryida. The addition of three Colpoda sequences strengthens the observation that species of this genus are distributed over the whole molecular Colpodea tree. Very likely, this is caused by a fast radiation of Colpoda, several species of which then evolved independently, forming new genera and families. Cyrtolophosis minor, which belongs to the molecular Pseudocyrtolophosis clade, is referred to a new genus, Apocyrtolophosis nov. gen., characterized by a comparably large, deltoid oral opening, an unciliated posterior region, and the absence of an oblique kinety in the left oral polykinetid. Bryometopus triquetrus does not erase the paraphyly of its genus. Platyophrya vorax, P. spumacola, and P. bromelicola form a highly supported clade in the order Platyophryida. Platyophryides and Ottowphrya are close genetically but differ in the silverline pattern (colpodid vs. platyophryid).     

Novistrombidium sinicum n. sp. and Novistrombidium orientale n. sp. (Protozoa: Ciliophora): Two New Oligotrich Ciliates from a Mangrove Wetland, South China

ABSTRACT. The morphology and infraciliature of two new marine oligotrich ciliates, Novistrombidium sinicum n. sp. and Novistrombidium orientale n. sp., isolated from a mangrove wetland near Guangzhou, southern China, were studied from live and protargol-stained specimens. Novistrombidium sinicum is different from its congeners by the combination of the following characters: three posteriorly directed thigmotactic membranelles, one ellipsoidal macronucleus, the extrusomes equidistantly arranged, and the ventral kinety commencing below the right end of the girdle kinety. The small-sized N. orientale can be separated from its congeners by two posteriorly directed thigmotactic membranelles, one ellipsoidal macronucleus, the extrusomes equidistantly arranged, and the ventral kinety commencing below the right end of the girdle kinety.     

Morphology,ontogenesis and molecular phylogeny of Platynematum salinarum nov. spec., a new scuticociliate (Ciliophora,Scuticociliatia) from a solar saltern

Platynematum salinarum nov. spec. was discovered in a hypersaline (∼120‰) solar saltern in Portugal. Its morphology, ontogenesis, and 18S rRNA were studied with routine methods. Platynematum salinarum has a size of about 35 μm × 18 μm and differs from other platynematids (= Platynematum and Pseudoplatynematum) in having an only slightly flattened body without any spines or notches. Both, the oral and somatic infraciliature resemble other platynematids and the tetrahymenid pattern in general. The ontogenesis is scuticobuccokinetal, being unique in generating protomembranelle 1 from kinetids produced by the paroral membrane of the proter and of the scutica. This composite divides transversely: the right half becomes the paroral membrane of the opisthe, the left half transforms to opisthe's adoral membranelle 1. The scutica and the molecular sequence classify P. salinarum into the order Scuticociliatida, family Cinetochilidae. The 18S rRNA sequence shows 92.7% similarity to the closest relative deposited in public databases (the scuticociliate Sathrophilus holtae), and our study provides the first sequence for the genus Platynematum. Experiments at different salinities show growth between 120‰ and 300‰, survival at 100‰, and cell death around 60‰ salinity, characterizing P. salinarum as a true halophile.     

Ontogenesis and molecular phylogeny of a new marine urostylid ciliate, Anteholosticha petzi n. sp. (Ciliophora, Urostylida)

A new urostylid ciliate, Anteholosticha petzi n. sp., isolated from the northern China seas, was studied using live observation and protargol impregnation. It differs from all the congeners mainly in the highly flexible and variable body shape, its red-brown to brick-red colour, and the possession of three types of cortical granules. Analysis of morphometric and molecular data confirm the validity of A. petzi as a distinct species. Anteholosticha petzi n. sp. is characterized as follows: by having 3 frontal, 1 buccal, 2 frontoterminal, 2 pretransverse, and 8-11 transverse cirri; a midventral row that comprises 10-16 cirral pairs and extends ca. 60% of cell length; and 3 bipolar dorsal kineties. Ontogenesis is similar to that in Anteholosticha warreni except for the formation of the oral primordium and is characterized by: (1) the parental adoral membranelles are completely renewed and the oral primordium is formed de novo; and (2) the anlagen for the frontal-ventral-transverse cirri are formed as primary primordia. The small subunit rRNA gene sequence of A. petzi n. sp. was also compared with that of its congeners with the sequence differences ranging from 0.39% to 8.62%. Phylogenetic trees based on the SSU rRNA gene sequence were constructed, indicating the nonmonophyly of the genus Anteholosticha, as supported by the approximately unbiased test.     

Morphology and molecular phylogeny of three new oligotrich ciliates (Protozoa,Ciliophora) from the South China Sea

The present paper documents the morphology and systematic positions of three new oligotrich ciliates, P arallelostrombidium obesum sp. nov. , P arallelostrombidium ellipticum sp. nov. , and S trombidium tropicum sp. nov. , which were sampled from habitats with different salinities in southern China. P arallelostrombidium obesum sp. nov. is characterized by a fat body and the posterior portions of the girdle and ventral kineties extending transversely on the dorsal side. P arallelostrombidium ellipticum sp. nov. is recognizable by the anterior ends of the girdle and ventral kineties being close to each other and the posterior ends of the girdle and ventral kineties intersecting on the dorsal side. S trombidium tropicum sp. nov. is distinguished by a ventrally opened girdle kinety that is slightly spiralled with the right end shifted posteriad. Small subunit rRNA gene trees show that P . obesum sp. nov. and P . ellipticum sp. nov. fall into a mixed group composed of Parallelostrombidium and some Novistrombidium species, and that S . tropicum sp. nov. branches at the base of the clade containing non‐Strombidium species. The relationships of Parallelostrombidium species and that of Strombidium species are both not resolved considering their low support values in our phylogenetic analysis. © 2015 The Linnean Society of London     

Morphology,morphogenesis and molecular phylogeny of a new marine ciliate,Trichototaxis marina n. sp. (Ciliophora,Urostylida)

The live morphology, infraciliature and morphogenesis of a new urostylid ciliate, Trichototaxis marina n. sp., collected from coastal water in Qingdao, China, were studied based on the observations of live and silver stained specimens. The new species is characterised as follows: body very flexible and contractile, slight to brick-reddish in colour due to irregularly-shaped, brick-red pigments; ca. 70 adoral membranelles; about 17 frontal cirri arranged in a bicorona; average 67 midventral pairs, the right base of each pair being conspicuously larger than the left base; five to seven transverse cirri; constantly two frontoterminal, one buccal and two pretransverse ventral cirri; two or three left marginal rows; right and innermost left marginal rows with 56–92 and 66–106 cirri, respectively; six bipolar dorsal kineties; more than 100 macronuclear nodules. The characteristic morphogenetic feature in T. marina is the development of the left marginal rows, that is, only one left marginal row is newly built the other one or two being retained from the parental cell. Phylogenetic analyses based on small subunit ribosomal gene sequence data reveal a close relationship of T. marina with members of family Pseudokeronopsidae.     

Morphology and taxonomy of Pseudokeronopsis rubra (Ehrenberg, 1836) and Pseudokeronopsis parasongi sp. nov. (Ciliophora,Hypotrichia, Urostylida) from the Yellow Sea

Pseudokeronopsis rubra (Ehrenberg, 1836), type species of Pseudokeronopsis, is a widely reported but taxonomically confused species with most available sequences either misidentified or dubious. Based on the Yellow Sea population, of which the molecular data have been analyzed in a previous work, we redescribe P. rubra and try to clarify the identifications of the populations designated as P. rubra or those allocated to this species. Nonetheless, there is still a need of integrated investigation of the species combing both morphological and molecular data based on specimens from its type locality. Recent molecular analyses indicate a Yellow Sea population of Pseudokeronopsis (marked as P. cf. songi) likely represents a new species. Here we describe it as P. parasongi sp. nov. based on the specimens from the same population. The new species possesses a yellow-brownish body with both pigmented cortical granules and colorless blood cell-shaped structures underneath the body surface, invariably one contractile vacuole at 66% of body length and occasionally a second one at 25%, and the midventral complex terminating 2–23 μm ahead of 2–4 transverse cirri. It differs from closely related species by its yellow-brownish body color and separates from them in the ITS1-5.8S-ITS2 and COI genetic distances.     

Morphology and Molecular Phylogeny of Two New Marine Euplotids,Pseudodiophrys nigricans n. g., n. sp., and Paradiophrys zhangi n. sp. (Ciliophora: Euplotida)

ABSTRACT. The morphology, infraciliature, and molecular phylogeny of Pseudodiophrys nigricans n. g., n. sp., and Paradiophrys zhangi n. sp., isolated from a sandy beach near Qingdao, China, were investigated. Pseudodiophrys is characterized by the Diophrys‐like ciliature pattern, but having only a single, reduced undulating membrane. Pseudodiophrys nigricans, the type species by monotypy, is described from live and silver‐impregnated specimens. Paradiophrys zhangi is similar to the type species Paradiophrys irmgard but can be recognized by its border body and the number (7 vs. 8–10) and arrangement (in rows vs. sparsely distributed) of the frontoventral cirri. Small subunit (SSU) rRNA gene sequence data support the validity of both species. Phylogenetic analyses based on the SSU rRNA gene sequence data currently available for uronychiids and other related taxa indicate that P. nigricans is most closely related to Diophrys scutum and Diophrys apoligothrix, while P. zhangi clusters most closely with Apodiophrys ovalis within a clade that also includes two other Paradiophrys species.