The primary problem hindering the study of octocorals is the disordered situation regarding their taxonomy, chiefly caused by insufficient knowledge of valid morphological taxonomic characters. Briareum is an octocoral genus found in the Atlantic and Pacific in shallow tropical and subtropical waters, and occurs in both encrusting and branching colony forms. Their simple morphology and morphological plasticity have hindered taxonomic understanding of this genus. In this study three morphologically distinct types (= type-1, -2, and -3) of Briareum from the Ryukyu Archipelago and their genetic diversity were examined. Colony, anthostele morphology, and sclerite length were examined for each type. Four molecular markers (mitochondrial cytochrome c oxidase subunit 1, mitochondrial mismatch repair gene, nuclear 18S ribosomal DNA, internal transcribed spacer 2 (ITS2)) were used to evaluate molecular phylogenetic status of these variations. Although one morphological type ("deep" small colonies, = type-3) showed small differences in nuclear ITS2 sequences compared to the other two types, the remaining types had identical sequences for all molecular markers examined. The results suggest extremely low genetic diversity despite highly variable morphology of Briareum species in Okinawa. Nevertheless, considering the distribution patterns and discontinuous morphology of type-3 compared to the other two morphotypes, genetic isolation of type-3 is plausible. In Briareum, small variances in nuclear ITS2 sequences of type-3 may have much more importance than in molecular phylogenies of other octocorals. Further phylogenetic investigations and comparison with Briareum specimens from other regions are necessary to conclusively taxonomically identify the three types.
Gregarines are a diverse group of apicomplexan parasites with a conspicuous extracellular feeding stage, called a "trophozoite", that infects the intestines and other body cavities of invertebrate hosts. Although the morphology of trophozoites is very diverse in gregarines as a whole, high degrees of intraspecific variation combined with relatively low degrees of interspecific variation make the delimitation of different species based on trophozoite morphology observed with light microscopy difficult. The coupling of molecular phylogenetic data with comparative morphology has shed considerable light onto the boundaries and interrelationships of different gregarine species. In this study, we isolated a novel marine gregarine from the hepatic region of a Pacific representative of the hemichordate Glossobalanus minutus, and report the first ultrastructural and molecular data from any gregarine infecting this distinctive group of hosts. Molecular phylogenetic analyses of an SSU rDNA sequence derived from two single-cell isolates of this marine gregarine demonstrated a strong and unexpected affiliation with a clade of terrestrial gregarines (e.g. Gregarina). This molecular phylogenetic data combined with a comparison of the morphological features in previous reports of gregarines collected from Atlantic representatives of G. minutus justified the establishment of a new binomial for the new isolate, namely Caliculium glossobalani n. gen. et sp. The molecular phylogenetic analyses demonstrated a clade of terrestrial gregarines associated with a sequence acquired from a marine species, which suggest that different groups of terrestrial/freshwater gregarines evolved independently from marine ancestors.
Despite the Central Indo-Pacific (CIP) and the Indonesian Archipelago being a well-known region of coral reef biodiversity, particularly in the 'Coral Triangle', little published information is available on its zoantharians (Cnidaria: Hexacorallia: Zoantharia). In order to provide a basis for future research on the Indo-Pacific zoantharian fauna and facilitate comparisons between more well-studied regions such as Japan and the Great Barrier Reef, this report deals with CIP zoantharian specimens in the Naturalis collection in Leiden, the Netherlands; 106 specimens were placed into 24 morpho-species and were supplemented with 88 in situ photographic records from Indonesia, the Philippines, and Papua New Guinea. At least nine morpho-species are likely to be undescribed species, indicating that the region needs more research in order to properly understand zoantharian diversity within the CIP. The Naturalis' zoantharian specimens are listed by species, as well as all relevant collection information, and in situ images are provided to aid in future studies on zoantharians in the CIP.
The low evolutionary rate of mitochondrial genes in Anthozoa has challenged their utility for phylogenetic and systematic purposes, especially for DNA barcoding. However, the evolutionary rate of Ceriantharia, one of the most enigmatic "orders" within Anthozoa, has never been specifically examined. In this study, the divergence of mitochondrial DNA of Ceriantharia was compared to members of other Anthozoa and Medusozoa groups. In addition, nuclear markers were used to check the relative phylogenetic position of Ceriantharia in relation to other Cnidaria members. The results demonstrated a pattern of divergence of mitochondrial DNA completely different from those estimated for other anthozoans, and phylogenetic analyses indicate that Ceriantharia is not included within hexacorallians in most performed analyses. Thus, we propose that the Ceriantharia should be addressed as a separate clade.
Stylophora pistillata is a widely used coral "lab-rat" species with highly variable morphology and a broad biogeographic range (Red Sea to western central Pacific). Here we show, by analysing Cytochorme Oxidase I sequences, from 241 samples across this range, that this taxon in fact comprises four deeply divergent clades corresponding to the Pacific-Western Australia, Chagos-Madagascar-South Africa, Gulf of Aden-Zanzibar-Madagascar, and Red Sea-Persian/Arabian Gulf-Kenya. On the basis of the fossil record of Stylophora, these four clades diverged from one another 51.5-29.6?Mya, i.e., long before the closure of the Tethyan connection between the tropical Indo-West Pacific and Atlantic in the early Miocene (16-24?Mya) and should be recognised as four distinct species. These findings have implications for comparative ecological and/or physiological studies carried out using Stylophora pistillata as a model species, and highlight the fact that phenotypic plasticity, thought to be common in scleractinian corals, can mask significant genetic variation.
Little is known about effects of large storm systems on mesophotic reefs. This study reports on how Typhoon 17 (Jelawat) affected Ryugu Reef on Okinawa-jima, Japan in September 2012. Benthic communities were surveyed before and after the typhoon using line intercept transect method. Comparison of the benthic assemblages showed highly significant differences in coral coverage at depths of 25-32 m before and after Typhoon 17. A large deep stand of Pachyseris foliosa was apparently less resistant to the storm than the shallower high diversity area of this reef. Contradictory to common perception, this research shows that large foliose corals at deeper depths are just as susceptible to typhoon damage as shallower branching corals. However, descriptive functional group analyses resulted in only minor changes after the disturbance, suggesting the high likelihood of recovery and the high resilience capacity of this mesophotic reef.
Microfossils of the genus Punctatus include developmental stages such as blastula, gastrula, and hatchlings, and represent the most complete developmental sequence of animals available from the earliest Cambrian. Despite the extremely well-preserved specimens, the evolutionary position of Punctatus has relied only on their conical remains and they have been tentatively assigned to cnidarians. We present a new interpretation of the Punctatus body plan based on the developmental reconstruction aided by recent advances in developmental biology.
The spawning behavior and early embryogenesis of Palythoa tuberculosa (Anthozoa, Zoantharia) were observed in August 2009 off Okinawa Island, Japan. P. tuberculosa released zygotes just after high tide around new moon nights. The mean diameter of zygotes was 365.6 ± s.d.14.8 ?m, and zygotes did not contain any symbiotic algae (zooxanthellae). About 2 h after spawning, the first cleavage furrow appeared on one side of the zygotes, although it was uncertain when eggs were fertilized. After second cleavage, the arrangement of blastomeres was pseudospherical. At 9 h after spawning, the embryo became a concave-convex dish shape, then gastrulation occurred and the blastopore was formed. Seven-day old larvae were ellipsoid and about 700 ?m long, with an open mouth at one end. Two weeks after spawning, the larvae developed a longitudinal band of long cilia (= ventral ciliate band) that is characteristic of zoanthella larvae. In P. tuberculosa, larvae show a non-radial body plan and then metamorphose to almost-radial (in outward appearance) polyps after settlement. These results may support a hypothesis that a common ancestor of Cnidaria had a bilateral body plan that has been secondarily lost in some extant cnidarians.
Zoanthids (Anthozoa, Hexacorallia) are colonial anemones that contain one of the deadliest toxins ever discovered, palytoxin (LD(50) in mice 300 ng/kg), but it is generally believed that highly toxic species are not sold in the home aquarium trade. We previously showed that an unintentionally introduced zoanthid in a home aquarium contained high concentrations of palytoxin and was likely responsible for a severe respiratory reaction when an individual attempted to eliminate the contaminant colonies using boiling water. To assess the availability and potential exposure of palytoxin to marine aquarium hobbyists, we analyzed zoanthid samples collected from local aquarium stores for palytoxin using liquid chromatography and high resolution mass spectrometry and attempted to identify the specimens through genetic analysis of 16S and cytochrome c oxidase 1 (COI) markers. We found four specimens of the same apparent species of zoanthid, that we described previously to be responsible for a severe respiratory reaction in a home aquarium, to be available in three aquarium stores in the Washington D.C. area. We found all of these specimens (n?=?4) to be highly toxic with palytoxin or palytoxin-like compounds (range 0.5-3.5 mg crude toxin/g zoanthid). One of the most potent non-protein compounds ever discovered is present in dangerous quantities in a select species of zoanthid commonly sold in the home aquarium trade.
Zoanthids (Cnidaria, Hexacorallia) of the genus Epizoanthus are often found in association with other marine invertebrates, including gastropods and hermit crabs. However, little information exists on the specificity and nature of these associations due to a lack of investigation into Epizoanthus species diversity, and the taxonomy of Epizoanthus is therefore confused. In this study, analyses of morphological data (tentacle number, polyp size, etc) and molecular data (mitochondrial cytochrome oxidase subunit 1 = COI, 16S ribosomal DNA = 16S rDNA) were used to examine Epizoanthus specimens from Tosa Bay, Kochi, Japan. The Epizoanthus specimens were found on both live gastropods (Gemmula unedo) and hermit crabs (Paguristes palythophilus) inhabiting G. unedo and G. cosmoi shells. While morphological analyses did not show clear differences between examined specimens, both COI and mt 16S rDNA clearly divided the specimens into two groups, one associated only with hermit crabs (= Epizoanthus sp. C), and another associated only with living gastropods (= Epizoanthus sp. S). Unexpectedly, DNA sequences from both groups did not match with two previously reported Epizoanthus species from Japan (E. indicus, E. ramosus), indicating they both may be undescribed species. These results highlight the utility of DNA "barcoding" of unknown zoanthids, and will provide a foundation for re-examinations of Epizoanthus species diversity and specificity, which will be critical in understanding the evolution of these unique marine invertebrates.
Zoanthids comprise an order of benthic, generally colonial cnidarians, which can usually be distinguished from other hexacorallians by embedded sand and detritus in their mesoglea to help strengthen their structure. These animals are becoming increasingly important research subjects in biochemistry and other research fields. Their inclusion of both calcium and silica results in the need for both decalcification and desilification for internal morphological examinations. Since the methodology of hydrofluoric acid (HF) desilification has rarely been documented in zoanthids, histological surveys for zoanthid taxonomy have often been abandoned and their taxonomy is often problematic. Recent investigations utilizing molecular methods have brought a clearer understanding of zoanthid diversity, but standardization of HF treatments are still needed to provide a link between molecular and more traditional techniques, and to properly examine specimens for which molecular methods may not be an option (e.g., formalin-preserved specimens, etc.). Here, we use both "straight" HF and, for the first time with zoanthids, buffered HF (BHF) treatments at different treatment lengths (1-48 h) on polyps from three different species of zoanthids for histological examination. Section conditions were judged based on the presence/absence of embedded detritus, drag marks, and tissue condition. Results show that the BHF treatment resulted in slightly better tissue conditions for all specimens, and suggest that desilification works well regardless of treatment time for species with smaller (polyp diameter <0.5 cm), less heavily encrusted polyps. Desilification of heavily encrusted Palythoa mutuki polyps were still problematic, with at least 24 h treatment needed. To aid future research, we provide guidelines for HF treatments of zoanthid specimens.
The zoanthid genus Neozoanthus was originally described in 1972 from a single species in Madagascar. This monotypic genus was placed within its own family, Neozoanthidae, given its unusual characters of only partial sand encrustation, and an endodermal sphincter muscle combined with a brachycnemic mesenterial arrangement. Recently, undescribed specimens of Neozoanthus were discovered thousands of kilometers away in both Australia and Japan. While the phylogenetic and evolutionary aspects of Neozoanthus spp. are now somewhat well understood, the new specimens remained undescribed. Here we describe the specimens as two new species, Neozoanthus uchinasp. n. from the Middle Ryukyu Islands of southern Japan, and Neozoanthus caleyisp. n. from the waters around Heron Island, on the Great Barrier Reef in Australia. Both species can be distinguished from each other and the type species, Neozoanthus tulearensis, by their distributions, oral disk colors, and average numbers of tentacles. Additionally, each species appears to have subtle differences in their cnidae. The division of Japanese and Australian specimens into two species is strongly supported by recently reported phylogenetic data. The discovery and description of these two species highlights how little is known of zoanthid species diversity in the Indo-Pacific.
The cosmopolitan solitary deep-water scleractinian coral Desmophyllum dianthus (Esper, 1794) was selected as a representative model species of the polyphyletic Caryophylliidae family to (1) examine phylogenetic relationships with respect to the principal Scleractinia taxa, (2) check population structure, (3) test the widespread connectivity hypothesis and (4) assess the utility of different nuclear and mitochondrial markers currently in use. To carry out these goals, DNA sequence data from nuclear (ITS and 28S) and mitochondrial (16S and COI) markers were analyzed for several coral species and for Mediterranean populations of D. dianthus. Three phylogenetic methodologies (ML, MP and BI), based on data from the four molecular markers, all supported D. dianthus as clearly belonging to the "robust" clade, in which the species Lophelia pertusa and D. dianthus not only grouped together, but also shared haplotypes for some DNA markers. Molecular results also showed shared haplotypes among D. dianthus populations distributed in regions separated by several thousands of kilometers and by clear geographic barriers. These results could reflect limited molecular and morphological taxonomic resolution rather than real widespread connectivity. Additional studies are needed in order to find molecular markers and morphological features able to disentangle the complex phylogenetic relationship in the Order Scleractinia and to differentiate isolated populations, thus avoiding the homoplasy found in some morphological characters that are still considered in the literature.
The question of how many marine species exist is important because it provides a metric for how much we do and do not know about life in the oceans. We have compiled the first register of the marine species of the world and used this baseline to estimate how many more species, partitioned among all major eukaryotic groups, may be discovered.
The family Melithaeidae (Octocorallia: Alcyonacea) is distributed in the West Pacific, Indian Ocean and the Red Sea. They are most abundant in warmer waters but can also be found in temperate waters. At present six genera are assigned to this family (Melithaea, Mopsella, Clathraria, Acabaria, Wrightella and Asperaxis), however overlapping characteristics make this groups taxonomic identification difficult and their relationships unclear. There are only a few reports from the Ryukyu Archipelago in southern Japan of melithaeids and most other octocorals, despite the islands being an area of high octocoral diversity. To help resolve the taxonomic confusion in this family, samples from various Ryukyu Archipelago locations were collected and DNA sequences of nuclear 28S ribosomal DNA and mitochondrial cytochrome oxidase I (COI) were obtained. Additionally, SEM micrographs of the sclerites of specimens were taken to further confirm the molecular results. Three strongly supported clades were recovered from the COI and 28S rDNA analyses, corresponding to Melithaea, Acabaria, and Mopsella, and in most cases clades were clearly related with the sclerite shape reported for each genus. These results show clearly that molecular differences are present between the three genera, and also demonstrates the strong need of other molecular markers for resolving intra-generic phylogenies. Our results provide baseline data for future studies of this octocoral family, not only on taxonomy, but also with regards to their distribution in the Ryukyu Islands.
Commensal leucothoid amphipods have been collected from coral rubble samples throughout the Ryukyu Archipelago, Japan. Seven new species are described in two generawith valuable location data. A new locality is presented for Paranamixis misakiensis Thomas, 1997. An identification key to all described Leucothoidae of the Ryukyu Archipelago is provided.
Commensal leucothoid amphipods have been collected from the canals of their sponge hosts throughout the Ryukyu Archipelago, Japan. Eleven new species are described in the genus Leucothoe with valuable location data and host records. An identification key to sponge-dwelling Leucothoidae of the Ryukyu Archipelago is provided.
Commensal leucothoid amphipods have been collected from the branchial chambers of their ascidian hosts throughout the Ryukyu Archipelago, Japan. Seven new species are described in two genera with valuable location data and host records. An identification key to ascidian-dwelling Leucothoidae of the Ryukyu Archipelago is provided.
Research on the soft coral genus Sarcophyton extends over a wide range of fields, including marine natural products and the isolation of a number of cembranoid diterpenes. However, it is still unknown how soft corals produce this diverse array of metabolites, and the relationship between soft coral diversity and cembranoid diterpene production is not clear. In order to understand this relationship, we examined Sarcophyton specimens from Okinawa, Japan, by utilizing three methods: morphological examination of sclerites, chemotype identification, and phylogenetic examination of both Sarcophyton (utilizing mitochondrial protein-coding genes MutS homolog: msh1) and their endosymbiotic Symbiodinium spp. (utilizing nuclear internal transcribed spacer of ribosomal DNA: ITS- rDNA). Chemotypes, molecular phylogenetic clades, and sclerites of Sarcophyton trocheliophorum specimens formed a clear and distinct group, but the relationships between chemotypes, molecular phylogenetic clade types and sclerites of the most common species, Sarcophyton glaucum, was not clear. S. glaucum was divided into four clades. A characteristic chemotype was observed within one phylogenetic clade of S. glaucum. Identities of symbiotic algae Symbiodinium spp. had no apparent relation to chemotypes of Sarcophyton spp. This study demonstrates that the complex results observed for S. glaucum are due to the incomplete and complex taxonomy of this species group. Our novel method of identification should help contribute to classification and taxonomic reassessment of this diverse soft coral genus.
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