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.     

Morphology and Infraciliature of Trochiliopsis australis N. Sp., Pelagohalteria viridis (Fromentel, 1876) N. G., N. Comb., and Strobilidium lacustris N. Sp. (Protozoa,Ciliophora)1

Three ciliate species from Australia and Norway were examined by silver impregnation, biometry, and scanning electron microscopy. Trochiliopsis australis n. sp. (order Nassulida, family Microthoracidae) differs from the single other known species of the genus, T. opaca, by its prominent ridges on the left side, the distinctly longer anterior segment of somatic kinety 2, and by the doubled total number of kinetosomes. Pelagohalteria n. g. (order Oligotrichida, family Halteriidae) differs from Halteria by the structure of the bristle complexes, which are composed of a vertically and a horizontally oriented row of kinetosomes. Two Halteria species belong to the new genus: P. viridis (Fromentel, 1876) n. comb., which is redescribed here, and P. cirrifera (Kahl, 1935) n. comb. Strobilidium lacustris n. sp. (order Oligotrichida, family Strobilidiidae) differs from its nearest relative, S. velox, by the smooth pellicle, the absence of symbiotic green algae, and by its larger size.     

Marine Heterotrophic Amoebae, Flagellates and Heliozoa From Belize (Central America) and Tenerife (Canary Islands), With Descriptions of New Species, Luffisphaera Bulbochaete N. Sp., L. Longihastis N. Sp., L. Turriformis N. Sp. and Paulinella Intermedia N. Sp.

ABSTRACT. Thirty four taxa of heterotrophic protists (amoebae, flagellates and heliozoa) were encountered in cultures established from marine samples from Belize (Central America) and Tenerife (Canary Islands). Most species are flagellates drawn from the choanoflagellates, the cryptophyceans, the euglenids, the kinetoplastids, the bicosoecids, the chromulinids, the pedinellids and a variety of laxa of uncertain affinities (Protista incertae sedis). the identity of the thecate choanoflagellates Salpingoeca ringens Kent, 1880, and S. tuba Kent, 1880, is discussed, and four new species of heterotrophic protists are described: one new species of the amoeba genus Paulinella (Paulinella intermedia n. sp.) and three new species of the incertae sedis genus Luffisphaera Belcher & Swale, 1975 ( Luffisphaera bulbochaete n. sp.; L. longihastis n. sp.; L. turriformis n. sp.).     

Molecular phylogeny of Cercomonadidae and kinetid patterns of Cercomonas and Eocercomonas gen. nov. (Cercomonadida, Cercozoa)

Cercomonads are among the most abundant and widespread zooflagellates in soil and freshwater. We cultured 22 strains and report their complete 18S rRNA sequences and light microscopic morphology. Phylogenetic analysis of 51 Cercomonas rRNA genes shows in each previously identified major clade (A, B) two very robust, highly divergent, multi-species subclades (A1, A2; B1, B2). We studied kinetid ultrastructure of five clade A representatives by serial sections. All have two closely associated left ventral posterior microtubular roots, an anterior dorsal root, a microtubule-nucleating left anterior root, and a cone of microtubules passing to the nucleus. Anterior centrioles (=basal bodies, kinetosomes) of A1 have cartwheels; the posterior centriole does not, suggesting it is older, and implying flagellar transformation similar to other bikonts. Strain C-80 (subclade A2) differs greatly, having a dorsal posterior microtubule band, but lacking the A1-specific fibrillar striated root, nuclear extension to the centrioles, centriolar diaphragm, extrusomes; both mature centrioles lack cartwheels. For clade A2 we establish Eocercomonas gen. n., with type Eocercomonas ramosa sp. n., and for clade B1 Paracercomonas gen. n. (type Paracercomonas marina sp. n.). We establish Paracercomonas ekelundi sp. n. for culture SCCAP C1 and propose a Cercomonas longicauda neotype and Cercomonas (=Neocercomonas) jutlandica comb. n. and Paracercomonas (=Cercomonas) metabolica comb. n.     

Description of Two New, Mycophagous Soil Ciliates (Ciliophora, Colpodea): Fungiphrya strobli n. g., n. sp. and Grossglockneria ovata n. sp.

ABSTRACT The morphology and infraciliature of two new, mycophagous soil ciliates are described. Specimens were isolated from dried, rewetted soil samples with the non-flooded Petri dish method and investigated in vivo and with various silver impregnation techniques. Fungiphrya strobli n. g., n. sp. was discovered in the mud of rock-pools on the summit of Table Mountain, Republic of South Africa. It is a holotrichously ciliated, about 50 × 40 μm-sized grossglockneriid with the oral apparatus on the right side of the cell. The adoral ciliature, minute in all other members of the group, is well developed and has a mean of eight kineties forming a conspicuous left, oral polykinetid, highly reminiscent of that found in small species of the genus Colpoda . The ejected extrusomes have a unique, inflated distal end. Grossglockneria ovata n. sp. was discovered in leaf litter from the Lackawanna State Forest in Pennsylvania, USA. It differs from the other members of the genus by the ovate shape, smooth cortex, and the sparse, irregularly-shaped mucocysts. Taxonomic characters and ranking of grossglockneriids are discussed. Because of the complex, unique feeding tube associated with a unique feeding strategy, mycophagy, it is argued that grossglockneriid ciliates should be classified in a separate order, in spite of their close genetic relatedness to members of the order Colpodida.     

A New Marine Interstitial Ciliate from the Northwest Atlantic Coast Tracheloraphis primitarum Sp. N. (Ciliophora, Kinetofragminophora, Karyorelictida, Trachelocercidae)

ABSTRACT A new species of marine interstitial ciliate, Tracheloraphis primitarum sp. n., is described from intertidal sands (160-200-μm medium grain size) along the Northwest Atlantic coast. Living specimens are not transparent under incident light, 0.4-0.9 mm in length, and are characterized by an elongated body, bulbous anterior and rounded posterior regions. Cytostome is simple without a cleft on the lip. Protargol silver impregnation shows 18-29 longitudinal kineties. The glabrous zone is very narrow, 6-7 μm wide, and corresponds to the area occupied by one kinety and two interkinetal spaces. Ten to fifteen kineties terminate against the glabrous zone. Interkinetal extrusomes are present. The nuclear apparatus is a single centrally-placed nuclear group that consists of three large irregular macronuclei and 2-3 micronuclei; these are either clustered or encapsulated.     

New and little-known species of Rhagidiidae from talus ecosystems, in the Czech Republic and Austria

Two new species of mites, Foveacheles (Proxistella) simulata n.sp. and Rhagidia longiseta n.sp., are described, and the little-known Foveacheles (Proxistella) terricola (C. L. Koch) is redescribed. All species were found inhabiting underground spaces in talus formations in the Czech Republic and Austria.     

Corticocolpoda kaneshiroae N. G., N. Sp., a New Colpodid Ciliate (Protozoa, Ciliophora) from the Bark of Ohia Trees in Hawaii

Corticocolpoda kaneshiroae n. g., n. sp. was discovered in the bark of Ohia trees (Metrosideros polymorpha) from the Hawaiian Archipelago. Its morphology and infraciliature were studied in live cells, with the scanning electron microscope, as well as in specimens impregnated with protargol, silver nitrate, and silver carbonate. The new genus, Corticocolpoda, belongs to the family Colpodidae and is unique in having a left oral ciliary field (polykinetid) composed of 13–20 equidistantly spaced, paired rows of monokinetids. The new species, C. kaneshiroae, measures 100–300 × 60–250 μm and has a huge vestibulum used to capture large food items, mainly other ciliates. The oral ciliary fields are within the vestibulum and of equal length; the right field is a paroral membrane composed of a single row of dikinetids. Several micronuclei are attached to the ellipsoid macronucleus. The mucocysts are inconspicuous and arranged around the somatic dikinetids. The resting cyst is globular and has a 1–2 μm thin, firm, yellowish inner wall and a 20-μm-thick, fragile, hyaline outer wall. Convergent evolutionary paths and a possible polyphyly of the Colpodidae are discussed. An improved key to the genera of the family Colpodidae is provided. Twenty-three species of ciliates and one flagel***iate (Hemimastix amphikineta), which occurred together with C. kaneshiroae, are new for the fauna of Hawaii and listed in the ecology section.     

Nadelspora canceri N. G., N. Sp., an Unusual Microsporidian Parasite of the Dungeness Crab, Cancer Magister

ABSTRACT. The microsporidium Nadelspora canceri n. g., n. sp., is described from the striated musculature of the Dungeness crab ( Cancer magister ) in Oregon, USA. The needle-shaped spores were rounded anteriorly, tapered to a posterior point and measured 7.1–11.8 × 0.2–0.3 μm in fixed preparations. The extremely narrow spore diameter prevented observation of morphological details at the light microscopic level and ultrastructural details of mature spores were difficult to resolve. Meronts were not observed and the monokaryotic merozoites and sporonts were not contained within either parasitophorous or sporophorous vesicles. Sporonts were disporoblastic and gave rise to monokaryotic sporoblasts that became narrow and elongate as they developed into immature spores with a developing polar filament. The nucleus was not clearly resolved in mature spores and may have been surrounded by the lamellar polaroplast. The polar filament was of nearly uniform diameter throughout most of its length and ended abruptly about three-fourths of the distance from the anterior end of the spore. Unusual spherical non-membrane bound granules surrounded the polar filament in a spiral arrangement. The new microsporidium resembles members of the family Mrazekiidiae, but differs in lacking a diplokaryon at any stage. It is probably most closely related to Baculea daphniae from which it differs primarily by spore shape and size. The familial relationships of the genus Baculea have not been determined and it is proposed to include it with Nadelspora in the new family Nadelsporidae.     

Morphological and taxonomical studies on two soil haptorid ciliates (Ciliophora, Litostomatea): Clavoplites haranti sp. n. and Enchelys terrenum (Foissner, 1984) comb. n., and taxonomy of the family Enchelyidae Ehrenberg, 1838

The morphology and infraciliature of two soil haptorid ciliates, Clavoplites haranti sp. n. and Enchelys terrenum (Foissner, 1984) comb. n., collected from Malé Karpaty Mts. and Biele Karpaty Mts. (Slovakia), were investigated using live observation and protargol impregnation. Clavoplites haranti is distinguished from its congeners by a combination of characters including: theronts spatulate to slenderly fusiform, trophonts bursiform, size about 60 x 25 microm in vivo; massive oral bulge with broadly clavate to lemon-shaped, 1.5-2 microm long extrusomes arranged in a ring in oral bulge; ellipsoidal macronucleus and one globular micronucleus; on average 13 ciliary rows, 3 anteriorly differentiated to an inconspicuous dorsal brush. Enchelys terrenum is about 85 x 30 microm in size and differs from its congeners by a combination of characters including spatulate shape; massive oral bulge with 4-6 microm long, fusiform extrusomes arranged in a ring in oral bulge; ellipsoidal to reniform macronucleus and one globular micronucleus; usually 19 ciliary rows, 3 anteriorly differentiated to an ordinary dorsal brush. The diagnoses of the family Enchelyidae and the genus Enchelys are improved. The new genus Armatoenchelys differs from the genera Enchelys and Apoenchelys in having both body and oral bulge extrusomes. Enchelys geleii, E. longitricha, and E. vermiformis are transferred to the new genus.     

Fine Structure of the Feeding Stage of A Sorogenic Ciliate, Sorogena Stoianovitchae Gen. N., Sp. N.

SYNOPSIS. A new species of kinetophragminophoran ciliate, collected from dried vegetation and capable of forming an aerial sorocarp, is described and named Sorogena stoianovitchae gen. n., sp. n. This ciliate is a voracious predator that feeds on species of Colpoda , and, when the latter is depleted in numbers, aggregates to forms sorogens. Each sorogen rises into the air from the surface of the water, forming a secreted stalk with a sorus of cysts at its apex. the feeding stage of the ciliate resembles an Enchelys in that it has an apical, slit-like mouth surrounded by a lip, a somewhat dorso-ventrally flattened body, and meridional kineties. Its length ranges from 40–75 μm and width from 23–55 μm. It has a typical rhabdos type of cytopharynx, but no specialized oral ciliature. the somatic kineties are formed of rows of paired kinetosomes with associated microfibrils, the arrangement of which differs a little from that of other ciliates of this subclass. Sorogena has tentatively been placed in the order Haptorida although it lacks toxicysts, recognizable mucocysts, and clavate cilia. Its unique life cycle and some of the details of its fine structure indicate differences between Sorogena and other haptorids so profound that a new family, SOROGENIDAE, is created for it. the type species (PNG76-73) was collected on dry figs at the Wau Ecology Institute, Papua New Guinea.     

A Comparative Analysis of the Morphology of Holosticha corlissi N. Sp. (Ciliophora,Hypotrichida) and Related Species1

ABSTRACT The morphology of the hypotrichous ciliate, Holosticha corlissi n. sp., found in the moss Calliergonella cuspidata taken from the beech wood of Montejo de la Sierra (Madrid, Spain) is described. Some characteristics (organization of the mid-ventral, frontal and fronto-terminal cirri, presence of a buccal cirrus and the number of transverse cirri) are sufficiently different from the closely related species H. intermedia, H. sigmoidea and H. xanthichroma to suggest that it is a separate species, although its body shape, nucleus and buccal apparatus are very similar.     

Morphological and Molecular Characterization of a New Protist Family, Sandmanniellidae n. fam. (Ciliophora, Colpodea), with Description of Sandmanniella terricola n. g., n. sp. from the Chobe Floodplain in Botswana

ABSTRACT. Sandmanniella terricola n. g., n. sp. was discovered in soil from the Chobe floodplain, Botswana, southern Africa. Its morphology and 18S rDNA gene sequence were studied with standard methods. Sandmanniella terricola is very likely an adversity strategist because it reaches peak abundances 6–12 h after rewetting the soil and maintains trophic food vacuoles with undigested bacteria in the resting cyst, a highly specific feature suggested as an indicator for an adversity life strategy. Possibly, the energy of the stored food vacuoles is used for reproduction and support of the cyst wall. Morphologically, Sandmanniella terricola is inconspicuous, having a size of only 50 × 40 μm and a simple, ellipsoidal shape. The main characteristics of the genus are a colpodid silverline pattern; a perioral cilia condensation; a flat, dish-shaped oral cavity, in the centre of which originates a long, conical oral basket resembling that of certain nassulid ciliates; and a vertically oriented left oral polykinetid composed of brick-shaped adoral organelles. This unique mixture of features and the gene sequence trees, where Sandmanniella shows an isolated position, suggest establishing a new family, the Sandmanniellidae n. fam., possibly related to the families Colpodidae or Bryophryidae. The curious oral basket provides some support for the hypothesis of a common ancestor of colpodid and nassulid ciliates.     

Epulopiscium fishelsoni N. G., N. Sp., a Protist of Uncertain Taxonomic Affinities from the Gut of an Herbivorous Reef Fish1

Epulopiscium fishelsoni n. g., n. sp., is described from the gut of Acanthurus nigrofuscus, an herbivorous surgeonfish (Acanthuridae: Teleostei) from the Red Sea, Israel. The symbiont is placed in the kingdom Protoctista (or Protista) but more specific taxonomic assignment is impossible at this time. The organism is of an elongate oval shape, round in cross-section, and highly mobile, with the ability to reverse direction rapidly. It lacks a mouth or other external structures, with the exception of fine surface filaments of an undetermined nature. Internally, there is a complex of folded membranes or tubules and a nucleus. No other organelles are evident. Reproduction is characterized by doubling of nuclei within the parental cell and emergence of daughter cells from a perforation in the parental cortex. Symbionts appear to exhibit circadian cycles of growth, reproductive activity, and movement within the gut. Although the organism greatly reduces local gut pH when present in large numbers, we know nothing of interactions between host and symbiont.     

Morphology of Three Species of Amphileptus (Protozoa,Ciliophora, Pleurostomatida) from the South China Sea,with Note on Phylogeny of A. dragescoi sp. n.

Two new and one problematic species of pleurostomatids, Amphileptus dragescoi sp. n., A. wilberti sp. n., and A. marinus from coastal areas of the South China Sea, are described based on observations of live and protargol‐impregnated specimens. Amphileptus dragescoi is different from its congeners by the presence of an apical group of extrusomes and the possession of 12–15 right and five left somatic kineties, two macronuclear nodules, and a single terminally positioned contractile vacuole. Amphileptus wilberti is diagnosed by oval or pyriform body, 15–19 right and seven or eight left somatic kineties, extrusomes arranged only in anterior portion of oral slit, usually three ventrally located contractile vacuoles, and two macronuclear nodules. Amphileptus marinus (Kahl, 1931) Song et al., 2004 is redescribed and its diagnosis is improved. One isolate which was misidentified as A. marinus by Song et al. (2004) is believed to represent an unknown form, named here as Amphileptus songi sp. n. Phylogenetic analyses of the SSU rDNA sequences indicate that the genus Amphileptus is paraphyletic, but its monophyly is not rejected by statistical tree topology tests.     

On Keelungia pulex nov. gen. et nov. sp., a heterotrophic euglenoid flagellate that lacks pellicular plates (Euglenophyceae,Euglenida)

Keelungia pulex nov. gen. et nov. sp. is described from coastal waters of NE Taiwan. The new species is heterotrophic and feeds on bacteria. Cells are oblong-ovoid, biflagellate and glide along the sides of the flask. Each cell is approximately 8–11 μm long, and one of the smallest euglenoid flagellates presently known. Keelungia lacks pellicular plates and in this respect resembles diplonemids and Symbiontida, which are thought to be among the basal groups of Euglenozoa. SEM showed the presence of 10 evenly spaced longitudinal striae in the cell surface, but the striae are difficult to see in the light microscope. TEM showed each stria to comprise a double set of very low longitudinal ridges separated by a shallow furrow, and supported by ca 5 microtubules beneath the plasmalemma, unlike the situation in diplonemids and Symbiontida. The cell surface was further subtended by an extensive system of rough cisternae of endoplasmic reticulum. Keelungia pulex is phylogenetically related to species of Ploeotia and to Lentomonas applanata, but differs in details of the feeding apparatus and in the absence of pellicular plates. Sequencing of SSU rDNA indicates that Ploeotia, Keelungia and Entosiphon form a clade near the base of the euglenoid phylogenetic tree.     

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.     

Kiitricha minuta n. sp., a peculiar hypotrichous ciliate (Ciliophora, Spirotrichea) from the Yellow Sea

The morphology and infraciliature of a new ciliate, Kiitricha minuta n. sp., isolated from the Yellow Sea, were investigated using live observation and protargol impregnation. Kiitricha minuta represents a third member of the rarely known order Kiitrichida. It is unique in the subclass Hypotrichia in having many rows of small uniform cirri along the right side of the body and the dorsal kineties composed of dikinetids, most of which bear two cilia. Kiitricha minuta n. sp. is ovoid and measures about 60 x 45 microm in vivo. It has a huge buccal cavity occupying about 80% of the body length, numerous body extrusomes, one macronucleus and two micronuclei, 27-27 adoral membranelles, 9-12 frontoventral cirral rows, a submarginal row of 7-9 cirri, 6 or 7 transverse cirri, and roughly 7-9 dorsal kineties. This new species differs distinctly from its only congener Kiitricha marina by its smaller size (60 microm vs. 80-150 microm), the presence of body extrusomes (vs. absent), the different macronuclear pattern (one vs. two nodules), and the lower number of frontoventral cirral rows (9-12 vs. 21-26), which terminate at the anterior two-thirds of body (vs. extend to the posterior). The new term "submarginal cirral row" is introduced to distinguish from the marginal cirral row in typical hypotrichs sensu lato. Based on our new observations and the literature, an improved diagnosis for the genus Kiitricha is provided and its phylogenetic importance is discussed.     

Three new species of dicyemid mesozoans (phylum Dicyemida) from Amphioctopus fangsiao (Mollusca: Cephalopoda), with comments on the occurrence patterns of dicyemids

Three new species of dicyemid mesozoans are described from the renal appendages of Amphioctopus fangsiao, collected off Akashi, in Harima Nada, and from Osaka Bay. Dicyema akashiense n. sp. is a small species that reaches about 900 microm in length. The vermiform stages are characterized as having 15-17 peripheral cells, a conical calotte, and an axial cell that extends to the base of the metapolar cells. Infusoriform embryos consist of 37 cells; two nuclei are present in each urn cell, and the refringent bodies are solid. Dicyema helocephalum n. sp. is a small species that reaches about 800 microm in length. The vermiform stages are characterized as having 22 peripheral cells, a disc-shaped calotte, and an axial cell that extends to the base of the propolar cells. Infusoriform embryos consist of 37 cells; a single nucleus is present in each urn cell, and the refringent bodies are solid. Dicyema awajiense n. sp. is a small species that reaches about 300 microm in length. The vermiform stages are characterized as having 22 peripheral cells, a conical calotte, and an axial cell that extends to the middle of the propolar cells. Infusoriform embryos consist of 37 cells; a single nucleus is present in each urn cell, and the refringent bodies are solid. In A. fangsiao various occurrence patterns of dicyemid species were observed, including instances where different dicyemid species were found in the renal appendage on each side. This suggests that dicyemids infect each renal appendage independently. The prevalence, reproductive traits, calotte shapes, and co-occurrence patterns of dicyemids are briefly discussed.     

Morphology and Phylogeny of Two New Pleurostomatid Ciliates,Epiphyllum shenzhenense n. sp. and Loxophyllum spirellum n. sp. (Protozoa,Ciliophora) from A Mangrove Wetland,South China

ABSTRACT. The morphology, infraciliature, and small subunit (SSU) rRNA gene sequences of two new pleurostomatid ciliates, Epiphyllum shenzhenense n. sp. and Loxophyllum spirellum n. sp., isolated from a mangrove wetland near Shenzhen, South China, were investigated. Epiphyllum shenzhenense n. sp. is morphologically characterized by leaf‐shaped cell about 150 × 35 μm in vivo, usually with four contractile vacuoles, 20–29 right kineties and 10–26 left kineties, ca. four macronuclear nodules, and two types of extrusomes (i.e. short spindle‐shaped and long bar‐shaped). As a new species, L. spirellum n. sp. is distinguished from its congeners by its posterior dorsal margin twisted onto the left side, the distribution of extrusomes (evenly arranged along the oral slit, the posterior end, and clustered to 7–13 warts on dorsal margin), the subterminally positioned contractile vacuole, the number of kineties (8–10 on right side, 4–5 on left side), and its genetic distance from congeners. Phylogenetic trees based on the SSU rRNA gene sequence for both organisms were constructed, which indicate that Epiphyllum is a distinct genus and occupies a basal position in the Pleurostomatida clade; L. spirellum n. sp. falls well into the Loxophyllum clade, which has a close relationship with Litonotus and Spiroloxophyllum.