Cellular mechanisms underlying extraordinary sulfide tolerance in a crustacean holobiont from hydrothermal vents.

The shallow-water hydrothermal vent system of Kueishan Island has been described as one of the world's most acidic and sulfide-rich marine habitats. The only recorded metazoan species living in the direct vicinity of the vents is Xenograpsus testudinatus, a brachyuran crab endemic to marine sulfide-rich vent systems. Despite the toxicity of hydrogen sulfide, X. testudinatus occupies an ecological niche in a sulfide-rich habitat, with the underlying detoxification mechanism remaining unknown. Using laboratory and field-based experiments, we characterized the gills of X. testudinatus that are the major site of sulfide detoxification. Here sulfide is oxidized to thiosulfate or bound to hypotaurine to generate the less toxic thiotaurine. Biochemical and molecular analyses demonstrated that the accumulation of thiosulfate and hypotaurine is mediated by the sodium-independent sulfate anion transporter (SLC26A11) and taurine transporter (Taut), which are expressed in gill epithelia. Histological and metagenomic analyses of gill tissues demonstrated a distinct bacterial signature dominated by Epsilonproteobacteria. Our results suggest that thiotaurine synthesized in gills is used by sulfide-oxidizing endo-symbiotic bacteria, creating an effective sulfide-buffering system. This work identified physiological mechanisms involving host-microbe interactions that support life of a metazoan in one of the most extreme environments on our planet.

A next generation approach to species delimitation reveals the role of hybridization in a cryptic species complex of corals.

BACKGROUND: Our ability to investigate processes shaping the evolutionary diversification of corals (Cnidaria: Anthozoa) is limited by a lack of understanding of species boundaries. Discerning species of corals has been challenging due to a multitude of factors, including homoplasious and plastic morphological characters and the use of molecular markers that are either not informative or have not completely sorted. Hybridization can also blur species boundaries by leading to incongruence between morphology and genetics. We used traditional DNA barcoding and restriction-site associated DNA sequencing combined with coalescence-based and allele-frequency methods to elucidate species boundaries and simultaneously examine the potential role of hybridization in a speciose genus of octocoral, Sinularia. RESULTS: Species delimitations using two widely used DNA barcode markers, mtMutS and 28S rDNA, were incongruent with one another and with the morphospecies identifications. When mtMutS and 28S were concatenated, a 0.3% genetic distance threshold delimited the majority of morphospecies. In contrast, 12 of the 15 examined morphospecies formed well-supported monophyletic clades in both concatenated RAxML phylogenies and SNAPP species trees of > 6000 RADSeq loci. DAPC and Structure analyses also supported morphospecies assignments, but indicated the potential for two additional cryptic species. Three morphologically distinct species pairs could not, however, be distinguished genetically. ABBA-BABA tests demonstrated significant admixture between some of those species, suggesting that hybridization may confound species delimitation in Sinularia. CONCLUSIONS: A genomic approach can help to guide species delimitation while simultaneously elucidating the processes generating coral diversity. Results support the hypothesis that hybridization is an important mechanism in the evolution of Anthozoa, including octocorals, and future research should examine the contribution of this mechanism in generating diversity across the coral tree of life.

Phylogeny and systematics of deep-sea precious corals (Anthozoa: Octocorallia: Coralliidae).

The phylogeny of Coralliidae is being increasingly studied to elucidate their evolutionary history and species delimitation due to global concerns about their conservation. Previous studies on phylogenetic relationships within Coralliidae have pointed out that the two currently recognized genera are not monophyletic and the Coralliidae should be divided into three genera. In order to provide a comprehensive revision of the taxonomy of Coralliidae, we documented 110 specimens using eight mitochondrial and one nuclear loci to reconstruct their phylogeny. The morphological features of 27 type specimens were also examined. Phylogenetic relationships based on both mitochondrial and nuclear markers revealed two reciprocally monophyletic clades of Coralliidae. One of the clades was further split into two subclades with respect to sequence variation and observable morphological features. Based on the results of genealogical analyses and distinctive morphological features, the three genera classification of Coralliidae proposed by Gray (1867) was redefined. In this revised taxonomic system, Corallium, Hemicorallium, and Pleurocorallium consist of 7, 16 and 14 species, respectively. Our results also showed that the cosmopolitan Hemicorallium laauense is a species complex containing a cryptic species.

Identification of potential source regions and long-range transport routes/channels of marine PM2.5 at remote sites in East Asia.

Long-range transport (LRT) of air masses in East Asia and their impacts on marine PM2.5 were explored. Situated in the leeward region of East Asia, Taiwan Island marked by its elevated Central Mountain Range (CMR) separates air masses into two distinct air currents. This study aims to investigate the transport of PM2.5 from the north to the leeward region. Six transport routes (A-F) were identified and further classified them into three main channels (i.e. East, West, and South Channels) based on their transport routes and potential sources. Green Island (Site GR) and Hengchun Peninsula (Site HC) exhibited similarities in their transport routes, with Central China, North China, and Korean Peninsula being the major source regions of PM2.5, particularly during the Asian Northeastern Monsoons (ANMs). Dongsha Island (Site DS) was influenced by both Central China and coastal regions of East China, indicating Asian continental outflow (ACO) as the major source of PM2.5. The positive matrix factorization (PMF) analysis of PM2.5 resolved that soil dust, sea salts, biomass burning, ship emissions, and secondary aerosols were the major sources. Northerly Channels (i.e. East and West Channels) were primarily influenced by ship emissions and secondary aerosols, while South Channel was dominated by oceanic spray and soil dust. The results of W-PSCF and W-CWT analysis indicated that three remote sites experienced significant contributions from Central China in the highest PM2.5 concentration range (75-100%). In contrast, PM2.5 in the 0-25% and 25-50% ranges primarily originated from the open seas, with ship emissions being the prominent source. It suggested that northern regions with heavy industrialization and urbanization have impacts on high PM2.5 concentrations, while open seas are the main sources of low PM2.5 concentrations.

Long-range transport of atmospheric speciated mercury from the eastern waters of Taiwan Island to northern South China Sea.

This study explored the temporospatial distribution, gas-particle partition, and pollution sources of atmospheric speciated mercury (ASM) from the eastern offshore waters of the Taiwan Island (TI) to the northern South China Sea (SCS). Both gaseous and particulate mercury were simultaneously sampled at three remote sites in four seasons. The average concentrations of gaseous elemental mercury (GEM), gaseous oxidized mercury (GOM), and particulate bound mercury (PBM) were 2.05 ± 0.45 ng/m3, 19.17 ± 5.39 pg/m3, and 0.11 ± 0.06 ng/m3, respectively. The concentrations of GEM and PBM in the cold seasons were higher than those in the warm seasons, but those of GOM had an opposite trend. In terms of gas-solid partition, ASM was apportioned as 91.3-97.3% of GEM and 2.7-8.7% of GOM and PBM. The average concentrations of GEM, GOM, and PBM at the Green Island (GI) were 2.21 ± 0.47 ng/m3, 22.31 ± 5.35 pg/m3, and 0.12 ± 0.06 ng/m3; those at the Kenting Peninsula (KT) were 2.11 ± 0.43 ng/m3, 20.57 ± 4.38 pg/m3, and 0.11 ± 0.06 ng/m3; and those at the Dongsha Islands (DS) were 1.84 ± 0.40 ng/m3, 15.19 ± 3.58 pg/m3, and 0.08 ± 0.05 ng/m3, respectively. Overall, the spatial distribution of ASM concentrations showed the order as: GI > KT > DS. Air masses blown mainly from the West Pacific Ocean (WPO) and SCS in summer showed the lowest ASM concentrations. Oppositely, high ASM concentrations were commonly observed in spring and winter when polluted air masses were blown by Asian Northeastern Monsoons (ANMs). The transport routes of polluted air masses were originated mainly from North China, Central China, Northeast China, Korea and Japan, and mostly passed through the urban and industrial regions in the northeastern Asian countries.

Chemical characteristics and spatiotemporal variation of marine fine particles for clustered channels of air masses transporting toward remote background sites in East Asia.

This study investigated the chemical characteristics, spatiotemporal distribution, and source apportionment of marine fine particles (PM2.5) for clustered transport channels/routes of air masses moving toward three remote sites in East Asia. Six transport routes in three channels were clustered based on backward trajectory simulation (BTS) in the order of: West Channel > East Channel > South Channel. Air masses transported toward Dongsha Island (DS) came mainly from the West Channel, while those transported toward Green Island (GR) and Kenting Peninsula (KT) came mostly from the East Channel. High PM2.5 commonly occurred from late fall to early spring during the periods of Asian Northeastern Monsoons (ANMs). Marine PM2.5 was dominated by water-soluble ions (WSIs) which were predominated by secondary inorganic aerosols (SIAs). Although the metallic content of PM2.5 was predominated by crustal elements (Ca, K, Mg, Fe, and Al), enrichment factor clearly showed that trace metals (Ti, Cr, Mn, Ni, Cu, and Zn) came mainly from anthropogenic sources. Organic carbon (OC) was superior to elemental carbon (EC), while OC/EC and SOC/OC ratios in winter and spring were higher than those in other two seasons. Similar trends were observed for levoglucosan and organic acids. The mass ratio of malonic acid and succinic acid (M/S) was commonly higher than unity, showing the influences of biomass burning (BB) and secondary organic aerosols (SOAs) on marine PM2.5. We resolved that the main sources of PM2.5 were sea salts, fugitive dust, boiler combustion, and SIAs. Boiler combustion and fishing boat emissions at the site DS had higher contribution than those at the sites GR and KT. The highest/lowest contribution ratios of cross-boundary transport (CBT) were 84.9/29.6% in winter and summer, respectively.

Inter-comparison of chemical fingerprint and source apportionment of marine fine particles at two islands through the west and east passages of the Taiwan Island.

In this study, the concentrations of marine fine particles (PM2.5) and their chemical fingerprints were inter-compared at two islands located aside from the west and east waters of Taiwan Island and the variability of west and east passages (i.e., Routes A1 and A2) were explored. Marine PM2.5 was simultaneously sampled at the Green and Dongsha Islands and five chemical components (i.e., water-soluble ions, metallic elements, carbonaceous content, anhydrosugars, and organic acids) were further analyzed in PM2.5 to characterize their chemical fingerprints. The highest concentrations of chemical composition and PM2.5 were commonly observed during the Asian Northeastern Monsoons (ANMs) via long-range transport (LRT). Water-soluble ions (WSIs) were dominated by secondary inorganic aerosols (SIAs), and followed by oceanic spray. The major metallic content of PM2.5 was crustal elements, while trace metals originated from anthropogenic sources with an enrichment factor (EF) > 10. In terms of carbonaceous content in PM2.5, organic carbon (OC) was superior to elemental carbon (EC). High levoglucosan concentrations were also observed during the periods of ANMs. Secondary organic aerosols (SOAs) were formed by atmospheric chemical reactions during the LRT procedure. The PM2.5 concentration of Route A1 was 37.51 % higher than that of Route A2, and trace metals (V, Mn, Ni, Pb, Cr, and Cu) increased significantly by 96.16-325.83 %. Positive matrix factorization (PMF) results revealed that the dominant factor of PM2.5 for Route A1 was shipping emissions and vehicular exhausts (41.2 %), while that for Route A2 was oceanic spray (30.2 %). Route A1 was mainly attributed to highly industrialized regions, densely populated urbanized areas, and ship-intensive traffics in East Asia.

From an Old Eroded Carapace: Rediscovery of the Majid Crab Leptomithrax sinensis Rathbun, 1916 (Crustacea, Brachyura, Majidae) from Taiwan and Japan.

Kingsley J. H. Wong, Peter K. L. Ng, and Ming-Shiou Jeng (2018) The majid crab Leptomithrax sinensis Rathbun, 1916 was previously only known from the holotype, a detached and partially eroded carapace collected during the Albatross Philippine Expedition in 1908 from the northern part of the South China Sea. Recent collections of fresh material from precious coral harvest sites off northeastern Taiwan, and Shikoku, Japan made verifying this poorly known species possible. The species is considered to be valid and is here redescribed, illustrated, and compared with its closest East Asian congener, L. bifidus (Ortmann, 1893).

Trophic structure and energy flow in a shallow-water hydrothermal vent: Insights from a stable isotope approach.

Shallow-water hydrothermal vent ecosystems are distinct from the deep-sea counterparts, because they are in receipt of sustenance from both chemosynthetic and photosynthetic production and have a lack of symbiosis. The trophic linkage and energy flow in these ecosystems, however remain elusive, which allows us poor understanding of the whole spectrum of biological components distributed across such environmental gradients. In this study, a thorough isotopic survey was conducted on various biological specimens and suspended particulates collected along four transects across the venting features of a shallow-water hydrothermal field off Kueishan Island, Taiwan. The isotope data combined with a Bayesian-based mixing model indicate that the vent-associated particulate organic matter (vent POM), as primary contribution of chemoautotrophic populations, has a high δ13C value (-18.2 ± 1.1‰) and a low δ15N value (-1.7 ± 0.4‰). Zooplankton and epibenthic crustaceans, as the fundamental consumers, exhibit δ13C and δ15N values ranging from -21.3 to -19.8‰ and +5.1 to +7.5‰, respectively, and can utilize the vent POM for 38-53% of their diets. The vent-obligate crab Xenograpsus testudinatus shows a large variation in δ13C (from -18.8 to -13.9‰) and δ15N values (from 1.1 to 9.8‰), although an omnivorous trophic level (2.5) is identified for it using δ15N values of amino acids, and it can utilize the vent POM for 6-87% of its diet. The consistently low (< 10.0‰) and overlapping δ15N values for most of the analyzed macroinvertebrates suggest extensive ingestion of chemosynthetic production complementing the photosynthetic production, a weak prey-predator relationship and low trophic complexity possibly imposed by the extreme environmental contexts of shallow-water hydrothermal ecosystems.

The Octocorals of Dongsha Atoll (South China Sea): An Iterative Approach to Species Identification Using Classical Taxonomy and Molecular Barcodes.

Yehuda Benayahu, Leendert Pieter van Ofwegen, Chang-feng Dai, Ming-Shiou Jeng, Keryea Soong, Alex Shlagman, Samuel W. Du, Prudence Hong, Nimrah H. Imam, Alice Chung, Tiana Wu, and Catherine S. McFadden (2018) Surveys of octocorals from Dongsha Atoll, Taiwan were conducted during 2011, 2013 and 2015 by SCUBA at a depth range of 6-25 m. The collections yielded ~540 specimens, encompassing the variety of taxa occurring in the explored sites; estimates of their abundances were also recorded. Dongsha features a highly diverse octocoral fauna, and octocorals are the dominant benthic organisms in the surveyed reef sites, often covering the majority of the hard substratum. Specimens were identified to the genus and species levels based on an iterative approach that integrates classical taxonomy with character-based molecular barcodes. A total of 51 nominal species representing 20 genera belonging to seven families were recorded, plus ~30 colonies that could only be assigned to a genus. Members of the family Alcyoniidae were the most abundant and diverse taxa, with 27 nominal species plus at least one potentially new, undescribed species of Sinularia, and 5-7 species each of Cladiella, Lobophytum and Sarcophyton. Problems with the taxonomic identification and phylogenetic relationships of species in these genera are discussed. The peculiarity of the Dongsha octocoral species composition is noted, and the composition is also compared to the other Taiwanese reef systems.

Strong Ion Regulatory Abilities Enable the Crab Xenograpsus testudinatus to Inhabit Highly Acidified Marine Vent Systems.

Hydrothermal vent organisms have evolved physiological adaptations to cope with extreme abiotic conditions including temperature and pH. To date, acid-base regulatory abilities of vent organisms are poorly investigated, although this physiological feature is essential for survival in low pH environments. We report the acid-base regulatory mechanisms of a hydrothermal vent crab, Xenograpsus testudinatus, endemic to highly acidic shallow-water vent habitats with average environment pH-values ranging between 5.4 and 6.6. Within a few hours, X. testudinatus restores extracellular pH (pHe) in response to environmental acidification of pH 6.5 (1.78 kPa pCO2) accompanied by an increase in blood [Formula: see text] levels from 8.8 ± 0.3 to 31 ± 6 mM. Branchial Na(+)/K(+)-ATPase (NKA) and V-type H(+)-ATPase (VHA), the major ion pumps involved in branchial acid-base regulation, showed dynamic increases in response to acidified conditions on the mRNA, protein and activity level. Immunohistochemical analyses demonstrate the presence of NKA in basolateral membranes, whereas the VHA is predominantly localized in cytoplasmic vesicles of branchial epithelial- and pillar-cells. X. testudinatus is closely related to other strong osmo-regulating brachyurans, which is also reflected in the phylogeny of the NKA. Accordingly, our results suggest that the evolution of strong ion regulatory abilities in brachyuran crabs that allowed the occupation of ecological niches in euryhaline, freshwater, and terrestrial habitats are probably also linked to substantial acid-base regulatory abilities. This physiological trait allowed X. testudinatus to successfully inhabit one of the world's most acidic marine environments.

Coralliidae (Anthozoa: Octocorallia) from the INDEMARES 2010 expedition to north and northwest Spain (northeast Atlantic), with delimitation of a new species using both morphological and molecular approaches.

Three species of deep-water bathyal Coralliidae were collected during the INDEMARES 2010 expedition of the Spanish Institute of Oceanography to the Avilés Canyon System and the Galicia Bank (Spain, northeast Atlantic): Corallium occultum n. sp., Corallium cf. bayeri Simpson & Watling, 2011, and Corallium niobe Bayer, 1964. The new species is supported by both morphological and molecular evidence, and its phylogenetic relationship within the Coralliidae is inferred. Corallium cf. bayeri is first recorded from European waters. Corallium johnsoni Gray, 1860 from off Portugal and Madeira, and Corallium tricolor (Johnson, 1898) from Madeira are redescribed from museum material, and their sclerites first depicted by scanning electron microscopy. The sclerome of C. johnsoni is more complex than previously thought, with occurrence of double clubs, and 6-, 7- and 8-radiates. A key is proposed for the identification of all the Atlantic species of the genus Corallium.