THE RECLASSIFICATION OF PFIESTERIA SHUMWAYAE (DINOPHYCEAE): PSEUDOPFIESTERIA,GEN. NOV.1

Pfiesteria shumwayae Glasgow et Burkholder is assigned to a new genus Pseudopfiesteria gen. nov. Plate tabulation differences between Pfiesteria and Pseudopfiesteria gen. nov. as well as a maximum likelihood phylogenetic analysis based on rDNA sequence data warrant creation of this new genus. The Kofoidian thecal plate formula for the new genus is Po, cp, X, 4′, 1a, 6′′, 6c, PC, 5+s, 5′′′, 0p, 2′′′′. In addition to having six precingular plates, P. shumwayae comb. nov. also has a distinctive diamond or rectangular‐shaped anterior intercalary plate. Both Pfiesteria and Pseudopfiesteria gen. nov. are reassigned to the order Peridiniales based on an apical pore complex (APC) with a canal (X) plate that contacts a symmetrical 1′, four to five sulcal plates, and the conservative hypothecal tabulation of 5′′′, 0p, and 2′′′′. These morphological characters and the life histories of Pfiesteria and Pseudopfiesteria are consistent with placement of both genera in the Peridiniales. Based on the plate tabulations for P. shumwayae, P. piscicida, and the closely related “cryptoperidiniopsoid” and “lucy” groups, the family Pfiesteriaceae is amended to include species with the following tabulation: 4‐5′, 0‐2a, 5‐6′′, 6c, PC, 5+s, 5′′′, 0p, and 2′′′′ as well as an APC containing a pore plate (Po), a closing plate (cp), and an X plate; the tabulation is expanded to increase the number of sulcal plates and to include a new plate, the peduncle cover (PC) plate. Members of the family have typical dinoflagellate life cycles characterized by a biflagellated free‐living motile stage, a varying number of cyst stages, and the absence of multiple amoeboid stages.     

Genetic identity of free‐living Symbiodinium obtained over a broad latitudinal range in the Japanese coast

The dinoflagellate Symbiodinium is well known to engage symbiosis with various marine animals, including corals. Recent records of environmental Symbiodinium (occurring in the environment and separately from host animals; usually referred to as ‘free‐living’ Symbiodinium) are of special interest, since these environmental populations are essential as symbiont sources for many host animals. In the present study, we carried out a phylogenetic analysis of environmental Symbiodinium isolates (culture strains) from sand, tide pools, or macroalgal surfaces, and environmental DNA clones extracted from the water‐column, at numerous sites around Japan. Our phylogenetic analysis based on the nuclear rRNA gene (internal transcribed spacers ‐1, ‐2, and 5.8S), indicated that most of the environmental isolates form monophyletic subclades within the Clade‐A lineage, and separate from a host‐associated Clade‐A population with high bootstrap values. Results of the partial nuclear 28S rDNA phylogeny and thecal‐plate observations revealed that these environmental isolates were closely related to a previously‐described ‘planktonic species’, Symbiodinium natans Gert Hansen et Daugbjerg, which was isolated from a plankton‐net sample from the Northeast‐Atlantic Ocean. On the other hand, the environmental DNA clones were also noted to be mostly nested within host‐associated Symbiodinium groups scattered in various clades. These results led to the assumption that the environmental Symbiodinium can be divided into two groups. One group, as typified by environmental isolates in the present study and previous reports, may be exclusively free‐living; the other group exists transiently in free‐living forms, possibly having been expelled from animal hosts. The populations within the latter group probably represent environmental sources of viable symbionts, because these are normally host‐associated. However, the Symbiodinium in the former group are not expected to engage in stable symbioses with host cnidarians.     

Brandtodinium gen. nov. and B. nutricula comb. Nov. (Dinophyceae), a dinoflagellate commonly found in symbiosis with polycystine radiolarians

Symbiotic interactions between pelagic hosts and microalgae have received little attention, although they are widespread in the photic layer of the world ocean, where they play a fundamental role in the ecology of the planktonic ecosystem. Polycystine radiolarians (including the orders Spumellaria, Collodaria and Nassellaria) are planktonic heterotrophic protists that are widely distributed and often abundant in the ocean. Many polycystines host symbiotic microalgae within their cytoplasm, mostly thought to be the dinoflagellate Scrippsiella nutricula, a species originally described by Karl Brandt in the late nineteenth century as Zooxanthella nutricula. The free‐living stage of this dinoflagellate has never been characterized in terms of morphology and thecal plate tabulation. We examined morphological characters and sequenced conservative ribosomal markers of clonal cultures of the free‐living stage of symbiotic dinoflagellates isolated from radiolarian hosts from the three polycystine orders. In addition, we sequenced symbiont genes directly from several polycystine‐symbiont holobiont specimens from different oceanic regions. Thecal plate arrangement of the free‐living stage does not match that of Scrippsiella or related genera, and LSU and SSU rDNA‐based molecular phylogenies place these symbionts in a distinct clade within the Peridiniales. Both phylogenetic analyses and the comparison of morphological features of culture strains with those reported for other closely related species support the erection of a new genus that we name Brandtodinium gen. nov. and the recombination of S. nutricula as B. nutricula comb. nov.     

MORPHOLOGY AND MOLECULAR PHYLOGENY OF A NEW MARINE SAND‐DWELLING PROROCENTRUM SPECIES,P. TSAWWASSENENSE (DINOPHYCEAE,PROROCENTRALES), FROM BRITISH COLUMBIA,CANADA1

A new marine benthic, sand‐dwelling Prorocentrum species from the temperate region of the Pacific coast of British Columbia, Canada, is described using LM and EM and molecular phylogenetic analyses. The cells have a broad oval shape, 40.0–55.0 μm long and 30.0–47.5 μm wide, and a wide U‐shaped periflagellar area on the right thecal plate. The left thecal plate consists of a straighter apical outline in the form of a raised ridge. Five to six delicate apical spines in the center of the periflagellar area are present. The nucleus is located in the posterior region of the cell, and a conspicuous pusule is located in the anterior region of the cell. The cells have golden‐brown chloroplasts with a compound, intrachloroplast pyrenoid that lacks a starch sheath. The thecal plates are smooth with round pores of two different sizes. The larger pores are arranged in a specific pattern of radial rows that are evenly spaced around the plate periphery and of irregular rows (or double rows) that form an incomplete “V” at the apical end of the plates. Large pores are absent in the center of the left and right thecal plates. The intercalary band is striated transversely and also has faint horizontal striations. Trichocysts and two types of mucocysts are present. The molecular phylogenetic position of Prorocentrum tsawwassenense sp. nov. was inferred using SSU rDNA sequences. This new species branched with high support in a Prorocentrum clade containing both benthic and planktonic species.     

PHYLOGENETIC ANALYSIS OF A FREE‐LIVING STRAIN OF SYMBIODINIUM ISOLATED FROM JIAOZHOU BAY,P.R. CHINA1

Recently, the isolation of a free‐living strain of symbiotic dinoflagellate belonging to the genus Symbiodinium was reported. Although the specimen procured and characterization from Jiaozhou Bay, P.R. China is a Symbiodinium spp., the manner in which this isolate was classified is inconsistent with the currently used and accepted Symbiodinium cladal nomenclature. To avert unnecessary confusion in the field, I place this important scientific contribution into the proper context and state of Symbiodinium research.     

Laciniporus arabicus gen. et sp. nov. (Dinophyceae,Peridiniales), a new thecate,marine, sand‐dwelling dinoflagellate from the northern Indian Ocean (Arabian Sea)1

A new thecate, photosynthetic, sand‐dwelling marine dinoflagellate, Laciniporus arabicus gen. et sp. nov., is described from the subtidal sediments of the Omani coast in the Arabian Sea, northern Indian Ocean, based on detailed morphological and molecular data. Cells of L. arabicus are small (16.2–30.1 μm long and 13.1–23.2 μm wide), dorsoventrally compressed, with a small apical flap‐shaped projection pointing to the left. The thecal plate pattern is distinguished by minute first precingular plate and sulcus, which extends into the epitheca, with large anterior and right sulcal plates. The Kofoidian thecal tabulation is Po, X, 4′, 2a, 7′′, 6c, 6s, 5′′′, 2′′′′. Morphologically, the revealed plate pattern has an affinity to the Peridiniales, and LSU rDNA based phylogenetic analyses placed L. arabicus within the Thoracosphaeraceae, close to calcareous‐cyst producing scrippsielloids, predatory pfiesteriaceans, and photosynthetic freshwater peridinioids Chimonodinium lomnickii and Apocalathium spp. However, the thecal plate arrangement of L. arabicus differs noticeably from any currently described dinoflagellates, and the species stands out from closely related taxa by extensive differences in physiology and ecology.     

Amoebophrya spp. from the Bloom‐forming Dinoflagellate Cochlodinium polykrikoides: Parasites not Nested in the “Amoebophrya ceratii Complex”

Members of Amoebophrya ceratii complex are known to infect a number of free‐living dinoflagellates including harmful algal bloom species. In August and October 2012, Amoebophrya infections during two bloom events of the dinoflagellate Cochlodinium polykrikoides were observed along southern coastal waters of Korea. Microscopic observations and molecular data revealed that two different Amoebophrya parasites infected the same host species. In addition, while one developed in the host's nucleus, the other in the host's cytoplasm. Phylogenetic analyses showed that the two parasites were not nested in the previously recognized “Amoebophrya ceratii complex clade”, which contained sequences of parasites infecting numerous dinoflagellate species. Instead, they branched as sister taxa to the isolate (possibly Amoebophrya) from radiolarians Hexacontium gigantheum. Our result indicates that the two Amoebophrya parasites infecting C. polykrikoides may be different species from those inside the “complex.”     

Genetics and Morphology Characterize the Dinoflagellate Symbiodinium voratum,n. sp., (Dinophyceae) as the Sole Representative of Symbiodinium Clade E

Dinoflagellates in the genus Symbiodinium are ubiquitous in shallow marine habitats where they commonly exist in symbiosis with cnidarians. Attempts to culture them often retrieve isolates that may not be symbiotic, but instead exist as free‐living species. In particular, cultures of Symbiodinium clade E obtained from temperate environments were recently shown to feed phagotrophically on bacteria and microalgae. Genetic, behavioral, and morphological evidence indicate that strains of clade E obtained from the northwestern, southwestern, and northeastern temperate Pacific Ocean as well as the Mediterranean Sea constitute a single species: Symbiodinium voratum n. sp. Chloroplast ribosomal 23S and mitochondrial cytochrome b nucleotide sequences were the same for all isolates. The D1/D2 domains of nuclear ribosomal DNA were identical among Western Pacific strains, but single nucleotide substitutions differentiated isolates from California (USA) and Spain. Phylogenetic analyses demonstrated that S. voratum is well‐separated evolutionarily from other Symbiodinium spp. The motile, or mastigote, cells from different cultures were morphologically similar when observed using light, scanning, and transmission electron microscopy; and the first complete Kofoidian plate formula for a Symbiodinium sp. was characterized. As the largest of known Symbiodinium spp., the average coccoid cell diameters measured among cultured isolates ranged between 12.2 (± 0.2 SE) and 13.3 (± 0.2 SE) μm. Unique among species in the genus, a high proportion (approximately 10–20%) of cells remain motile in culture during the dark cycle. Although S. voratum occurs on surfaces of various substrates and is potentially common in the plankton of coastal areas, it may be incapable of forming stable mutualistic symbioses.     

COMPARISON OF ENDOSYMBIOTIC AND FREE‐LIVING SYMBIODINIUM (DINOPHYCEAE) DIVERSITY IN A HAWAIIAN REEF ENVIRONMENT1

Many scleractinian corals must acquire their endosymbiotic dinoflagellates (genus Symbiodinium) anew each generation from environmental pools, and exchange between endosymbiotic and environmental pools of Symbiodinium (reef waters and sediments) has been proposed as a mechanism for optimizing coral physiology in the face of environmental change. Our understanding of the diversity of Symbiodinium spp. in environmental pools is poor by comparison to that engaged in endosymbiosis, which reflects the challenges of visualizing the genus against the backdrop of the complex and diverse micro‐eukaryotic communities found free‐living in the environment. Here, the molecular diversity of Symbiodinium living in the waters and sediments of a reef near Coconut Island, O‘ahu, Hawai‘i, sampled at four hourly intervals over a period of 5 d was characterized using a Symbiodinium‐specific hypervariable region of the chloroplast 23S. A comparison of Symbiodinium spp. diversity recovered from environmental samples with the endosymbiotic diversity in coral species that dominate the adjacent reef revealed limited overlap between these communities. These data suggest that the potential for infection, exchange, and/or repopulation of corals with Symbiodinium derived from the environment is limited at this location, a finding that is perhaps consistent with the high proportion of coral species in this geographic region that transmit endosymbionts from generation to generation.     

DESCRIPTION OF A NEW GENUS OF PFIESTERIA‐LIKE DINOFLAGELLATE,LUCIELLA GEN. NOV. (DINOPHYCEAE), INCLUDING TWO NEW SPECIES: LUCIELLA MASANENSIS SP. NOV. AND LUCIELLA ATLANTIS SP. NOV.1

A new genus of Pfiesteria‐like heterotrophic dinoflagellate, Luciella gen. nov., and two new species, Luciella masanensis sp. nov. and Luciella atlantis sp. nov., are described. These species commonly occur with other small (<20 μm) heterotrophic and mixotrophic dinoflagellates in estuaries from Florida to Maryland and the southern coast of Korea, suggesting a possible global distribution. An SEM analysis indicates that members of the genus Luciella have the enhanced Kofoidian plate formula of Po, cp, X, 4′, 2a, 6″, 6c, PC, 5+s, 5?, 0p, and 2″″. The two four‐sided anterior intercalary plates are diamond shaped. The genus Luciella differs from the other genera in the Pfiesteriaceae by a least one plate in the plate tabulation and in the configuration of the two anterior intercalary plates. An SSU rDNA phylogenetic analysis confirmed the genus as monophyletic and distinct from the other genera in the Pfiesteriaceae. The morphology of Luciella masanensis closely resembles Pfiesteria piscicida Steid. et J. M. Burkh. and other Pfiesteria‐like dinoflagellates in size and shape, making it easily misidentified using LM. Luciella atlantis, in contrast, has a more distinctive morphology. It can be distinguished from L. masanensis and other Pfiesteria‐like organisms by a larger cell size, a more conical‐shaped epitheca and hypotheca, larger rhombic‐shaped intercalary plates, and an asymmetrical hypotheca. The genus Luciella is assigned to the order Peridiniales and the family Pfiesteriaceae based on plate tabulation, plate pattern, general morphology, and phylogenetic analysis.     

Cryptic Sex in Symbiodinium (Alveolata,Dinoflagellata) is Supported by an Inventory of Meiotic Genes

Symbiodinium encompasses a diverse clade of dinoflagellates that are ecologically important as symbionts of corals and other marine organisms. Despite decades of study, cytological evidence of sex (karyogamy and meiosis) has not been demonstrated in Symbiodinium, although molecular population genetic patterns support the occurrence of sexual recombination. Here, we provide additional support for sex in Symbiodinium by uncovering six meiosis‐specific and 25 meiosis‐related genes in three published genomes. Cryptic sex may be occurring in Symbiodinium's seldom‐seen free‐living state while being inactive in the symbiotic state.     

Isolation of New <Emphasis Type="Italic">Symbiodinium</Emphasis> Strains from Tridacnid Giant Clam (<Emphasis Type="Italic">Tridacna crocea</Emphasis>) and Sea Slug (<Emphasis Type="Italic">Pteraeolidia ianthina</Emphasis>) Using Culture Medium Containing Giant Clam Tissue Homogenate

Recent molecular biological studies have revealed that some photosymbiotic invertebrates dwelling in coral reefs host several genetically different dinoflagellates, Symbiodinium species, as symbionts. However, little is known about the difference in physiologic characteristics among these symbionts living in a single host, because some Symbiodinium strains are difficult to culture in vitro. To isolate some of these Symbiodinium strains, we have developed an agar culture medium plate containing antibiotics and a giant clam tissue homogenate. Using-this medium we isolated two new Symbiodinium strains from two molluscan hosts, Tridacna crocea and Pteraeolidia ianthina, each of which hosted two different Symbiodinium strains belonging to Symbiodinium C and D, respectively. The tissue homogenate was essential for the growth of Symbiodinium D. Although it was not essential for the growth of Symbiodinium C, it did stimulate the initial growth. For the isolation of some Symbiodinium strains, isolation medium containing host homogenate is effective.     

Ailadinium reticulatum gen. et sp. nov. (Dinophyceae), a New Thecate,Marine, Sand‐Dwelling Dinoflagellate from the Northern Red Sea

A new photosynthetic, sand‐dwelling marine dinoflagellate, Ailadinium reticulatum gen. et sp. nov., is described from the Jordanian coast in the Gulf of Aqaba, northern Red Sea, based on detailed morphological and molecular data. A. reticulatum is a large (53–61 μm long and 38–48 μm wide), dorsoventrally compressed species, with the epitheca smaller than the hypotheca. The theca of this new species is thick and peculiarly ornamented with round to polygonal depressions forming a foveate‐reticulate thecal surface structure. The Kofoidian thecal tabulation is APC (Po, cp), 4′, 2a, 6′′, 6c, 4s, 6′′′, 1p, 1′′′′ or alternatively it can be interpreted as APC, 4′, 2a, 6′′, 6c, 4s, 6′′′, 2′′′′. The plate pattern of A. reticulatum is noticeably different from described dinoflagellate genera. Phylogenetic analyses based on the SSU and LSU rDNA genes did not show any supported affinities with currently known thecate dinoflagellates.     

PROTOPERIDINIUM BELIZEANUM SP. NOV. (DINOPHYCEAE) FROM MANATEE CAY,BELIZE, CENTRAL AMERICA1

A new marine heterotrophic dinoflagellate species, Protoperidinium belizeanum sp. nov., from a coral reef‐mangrove pond was identified from scanning electron micrographs. Recognition of this new species was based on unique features of the thecal morphology, which included cell size and shape, presence of short and wide postcingular plates, sulcal architecture, antapical spines, and intricate thecal plate patterns of ridged hexagonal depressions. The thecal plate formula is as follows: Po, X, 4′, 3a, 7″, 4C (3+t), 6S, 5?, 2″″. Species association of P. be‐lizeanum sp. nov. within the genus Protoperidinium, its habitat, and associated dinoflagellates species are discussed.     

SAME BUT DIFFERENT: TWO NOVEL BICARINATE SPECIES OF EXTANT CALCAREOUS DINOPHYTES (THORACOSPHAERACEAE,PERIDINIALES) FROM THE MEDITERRANEAN SEA1

The diversity of extant calcareous dinophytes (Thoracosphaeraceae, Dinophyceae) is currently not sufficiently recorded. The majority of their coccoid stages are cryptotabulate or entirely atabulate, whereas relatively few forms exhibit at least some degree of tabulation more than the archeopyle. A survey of coastal surface sediment samples from the Mediterranean Sea resulted in the isolation and cultivation of several strains of calcareous dinophytes showing a prominent tabulation. We investigated the morphologies of the thecate and the coccoid cells and conducted phylogenetic analyses using Maximum Likelihood and Bayesian approaches. The coccoid cells showed a distinct reflection of the cingulum (and were thus cingulotabulate), whereas thecal morphology corresponded to the widely distributed and species‐rich Scrippsiella. As inferred from molecular sequence data (including 81 new GenBank entries), the strains belonged to the Scrippsiella sensu lato clade of the Thoracosphaeraceae and represented two distinct species. Morphological details likewise indicated two distinct species with previously unknown coccoid cells that we describe here as new, namely S. bicarinata spec. nov. and S. kirschiae spec. nov. Cingulotabulation results from the fusion of processes representing the pre‐ and postcingular plate series in S. bicarinata, whereas the ridges represent sutures between the cingulum and the pre‐ and postcingular series in S. kirschiae, respectively. Bicarinate cingulotabulation appears homoplasious among calcareous dinophytes, which is further supported by a comparison to similar, but only distantly related fossil forms.     

HOST‐SPECIALIST LINEAGES DOMINATE THE ADAPTIVE RADIATION OF REEF CORAL ENDOSYMBIONTS

Bursts in species diversification are well documented among animals and plants, yet few studies have assessed recent adaptive radiations of eukaryotic microbes. Consequently, we examined the radiation of the most ecologically dominant group of endosymbiotic dinoflagellates found in reef‐building corals, Symbiodinium Clade C, using nuclear ribosomal (ITS2), chloroplast (psbA^ncr), and multilocus microsatellite genotyping. Through a hierarchical analysis of high‐resolution genetic data, we assessed whether ecologically distinct Symbiodinium, differentiated by seemingly equivocal rDNA sequence differences, are independent species lineages. We also considered the role of host specificity in Symbiodinium speciation and the correspondence between endosymbiont diversification and Caribbean paleo‐history. According to phylogenetic, biological, and ecological species concepts, Symbiodinium Clade C comprises many distinct species. Although regional factors contributed to population‐genetic structuring of these lineages, Symbiodinium diversification was mainly driven by host specialization. By combining patterns of the endosymbiont's host specificity, water depth distribution, and phylogeography with paleo‐historical signals of climate change, we inferred that present‐day species diversity on Atlantic coral reefs stemmed mostly from a post‐Miocene adaptive radiation. Host‐generalist progenitors spread, specialized, and diversified during the ensuing epochs of prolonged global cooling and change in reef‐faunal assemblages. Our evolutionary reconstruction thus suggests that Symbiodinium undergoes “boom and bust” phases in diversification and extinction during major climate shifts.     

CONSPECIFICITY AND INDO-PACIFIC DISTRIBUTION OF SYMBIODINIUM GENOTYPES (DINOPHYCEAE) FROM GIANT CLAMS

We previously reported the occurrence of genetically‐diverse symbiotic dinoflagellates (zooxanthellae) within and between 7 giant clam species (Tridacnidae) from the Philippines based on the algal isolates' allozyme and random amplified polymorphic DNA (RAPD) patterns. We also reported that these isolates all belong to clade A of the Symbiodinium phylogeny with identical 18S rDNA sequences. Here we extend the genetic characterization of Symbiodinium isolates from giant clams and propose that they are conspecific. We used the combined DNA sequences of the internal transcribed spacer (ITS)1, 5.8S rDNA, and ITS2 regions (rDNA‐ITS region) because the ITS1 and ITS2 regions evolve faster than 18S rDNA and have been shown to be useful in distinguishing strains of other dinoflagellates. DGGE of the most variable segment of the rDNA‐ITS region, ITS1, from clonal representatives of clades A, B, and C showed minimal intragenomic variation. The rDNA‐ITS region shows similar phylogenetic relationships between Symbiodinium isolates from symbiotic bivalves and some cnidarians as does 18S rDNA, and that there are not many different clade A species or strains among cultured zooxanthellae (CZ) from giant clams. The CZ from giant clams had virtually identical sequences, with only a single nucleotide difference in the ITS2 region separating two groups of isolates. These data suggest that there is one CZ species and perhaps two CZ strains, each CZ strain containing individuals that have diverse allozyme and RAPD genotypes. The CZ isolated from giant clams from different areas in the Philippines (21 isolates, 7 clam species), the Australian Great Barrier Reef (1 isolate, 1 clam species), Palau (8 isolates, 7 clam species), and Okinawa, Japan (1 isolate, 1 clam species) shared the same rDNA‐ITS sequences. Furthermore, analysis of fresh isolates from giant clams collected from these geographical areas shows that these bivalves also host indistinguishable clade C symbionts. These data demonstrate that conspecific Symbiodinium genotypes, particularly clade A symbionts, are distributed in giant clams throughout the Indo‐Pacific.     

Morphology and phylogenetic interpretation of a new Cambrian edrioasteroid (Echinodermata) from Spain

Abstract: The edrioasteroid, Aragocystites belli gen. nov. sp. nov. from the middle Cambrian Murero Formation of Spain, is described based on a small number of very well‐preserved specimens. Important anatomical characteristics include star‐shaped and pseudoclavate theca, rare or absent epispires, well‐developed interradially positioned oral plates and several unorganized cover plates associated with each widely exposed flooring plate. A phylogenetic analysis including several Cambro–Ordovician species shows it is more derived than Stromatocystites and Totiglobus but is a sister group to a clade comprising Cambraster and Edriodiscus. Ontogenetic observations based on juveniles of 5 mm in diameter suggest that this species changed thecal shape markedly during growth. A. belli gen. nov. sp. nov. probably lived in quiet environments where it attached directly to the sea floor on stabilized substrates.     

Free‐living calamyzin chrysopetalids (Annelida) from methane seeps,anoxic basins,and whale falls

Members of Calamyzinae, a clade of free‐living and ectoparasitic chrysopetalids, are mainly associated with deep‐sea chemosynthetic environments. The three currently known free‐living calamyzin species are placed in Vigtorniella. A new free‐living calamyzin species similar to these is described here. Phylogenetic analyses of Calamyzinae using mitochondrial (cytochrome c oxidase subunit I and 16S rDNA) and nuclear (Histone H3 and 18S rDNA) loci showed that Vigtorniella and the new species form a grade with respect to an ectoparasitic clade, requiring two new genera to be erected. All free‐living calamyzins show a similar anterior end and chaetal morphology. Micospina auribohnorum gen. et sp. nov. is described for the small‐bodied new species from deep‐sea whale falls off California and methane seeps off Costa Rica. The maximum‐likelihood and Bayesian analyses show Micospina gen. nov. as sister to the ectoparasitic clade. Boudemos gen. nov. is named for the clade of two larger‐bodied species: Boudemos flokati gen. et comb. nov. and Boudemos ardabilia gen. et comb. nov., which is sister group to all other Calamyzinae. Vigtorniella is retained for the type species, Vigtorniella zaikai (Kiseleva, 1992), with the adults found amongst bacterial mats at the boundary of the hydrogen sulphide zone in the Black Sea. Micospina gen. nov., Boudemos gen. nov., and Vigtorniella form a grade of free‐living taxa that is associated with feeding on organic‐enriched sediments, and the latter two taxa display ontogenetic jaw change. Jaws are absent in Micospina auribohnorum gen. et sp. nov. and most of the calamyzin clade of parasitic forms.     

Thecal tabulation,body scale morphology and phylogeny of Heterocapsa philippinensis sp. nov. (Peridiniales,Dinophyceae) from the Philippines

The thecal tabulation and body scale structure of the marine armoured dinoflagellate Heterocapsa, isolated from Philippines, were examined using LM, SEM and TEM, and its phylogenetic position was inferred from ITS and LSU rDNA sequences. Cells were ovoid and the plate tabulation (Po, cp, X, 5′, 3a, 7′′, 6c, 5s, 5′′′, 2′′′′) was consistent with most Heterocapsa species. The second anterior intercalary plate (2a) had a circular pattern with a thick marginal border free of pores. The nucleus was longitudinally elongated and curved, and located at the dorsal side of the cell. Discoid lobes of brownish chloroplast were peripherally distributed, and a pyrenoid was positioned at the centre. The triradiate body scales, measuring 250–300 nm in diameter, consisted of a roundish basal plate with six radiating ridges, nine peripheral uprights/spines, and three radiating spines. These components were identical to those of H. pseudotriquetra and H. steinii, except for the roundish outline of basal plate. Molecular phylogeny showed that the species clustered with H. pseudotriquetra and H. steinii. This species was differentiated from all other Heterocapsa species in the sausage-shaped nucleus and circular pattern on the 2a plate. This study proposed a novel species Heterocapsa philippinensis sp. nov. for the isolate.