Reef-Building Corals and Reefs of Vietnam: 1. The Gulf of Thailand

This paper deals with the history and investigations of the reefs and coral communities of the Gulf of Thailand based on published and unpublished materials, including the author's. The state of the art in the study of reef-building scleractinian corals and reefs of this region is presented. Characterized by remarkable distinctive features, the coral fauna and reefs of the Gulf of Thailand exhibit high similarity in coral species composition to other regions of Vietnam and form a single complex of species of the equatorial Indo-Pacific.     

Reef-Building Corals of Vietnam as a Part of the Indo-Pacific Reef Ecosystem

The paper analyzes both published and unpublished results of the investigations of Vietnamese reef-building corals and reefs performed in the last decades of the twentieth century. The state of the art in the study of reef-building scleractinian corals and reefs is presented. The scleractinian fauna of Vietnam is shown to match in species diversity (366 species of 70 genera) the tropical coral fauna of the Indonesian-Malacca fertile center, from which Indo-Pacific reef-building corals originated. The whole Vietnam coast from the Gulf of Tonkin to the Gulf of Siam is a biogeographically single whole and is a part of the Indo-Polynesian Province of the Indo-Pacific Area.     

The Evolution of Calcification in Reef-Building Corals

Corals build the structural foundation of coral reefs, one of the most diverse and productive ecosystems on our planet. Although the process of coral calcification that allows corals to build these immense structures has been extensively investigated, we still know little about the evolutionary processes that allowed the soft-bodied ancestor of corals to become the ecosystem builders they are today. Using a combination of phylogenomics, proteomics, and immunohistochemistry, we show that scleractinian corals likely acquired the ability to calcify sometime between ∼308 and ∼265 Ma through a combination of lineage-specific gene duplications and the co-option of existing genes to the calcification process. Our results suggest that coral calcification did not require extensive evolutionary changes, but rather few coral-specific gene duplications and a series of small, gradual optimizations of ancestral proteins and their co-option to the calcification process.     

The Diversity and Biogeography of Western Indian Ocean Reef-Building Corals

This study assesses the biogeographic classification of the Western Indian Ocean (WIO) on the basis of the species diversity and distribution of reef-building corals. Twenty one locations were sampled between 2002 and 2011. Presence/absence of scleractinian corals was noted on SCUBA, with the aid of underwater digital photographs and reference publications for species identification. Sampling effort varied from 7 to 37 samples per location, with 15 to 45 minutes per dive allocated to species observations, depending on the logistics on each trip. Species presence/absence was analyzed using the Bray-Curtis similarity coefficient, followed by cluster analysis and multi-dimensional scaling. Total (asymptotic) species number per location was estimated using the Michaelis-Menten equation. Three hundred and sixty nine coral species were named with stable identifications and used for analysis. At the location level, estimated maximum species richness ranged from 297 (Nacala, Mozambique) to 174 (Farquhar, Seychelles). Locations in the northern Mozambique Channel had the highest diversity and similarity, forming a core region defined by its unique oceanography of variable meso-scale eddies that confer high connectivity within this region. A distinction between mainland and island fauna was not found; instead, diversity decreased radially from the northern Mozambique Channel. The Chagos archipelago was closely related to the northern Mozambique Channel region, and analysis of hard coral data in the IUCN Red List found Chagos to be more closely related to the WIO than to the Maldives, India and Sri Lanka. Diversity patterns were consistent with primary oceanographic drivers in the WIO, reflecting inflow of the South Equatorial Current, maintenance of high diversity in the northern Mozambique Channel, and export from this central region to the north and south, and to the Seychelles and Mascarene islands.     

Reef-Building Corals—Symbiotic Autotrophic Organisms: 2. Pathways and Mechanisms of Adaptation to Light

Ways, mechanisms, and responses of ontogenetic adaptation of reef-building corals to light are discussed on the basis of original and literature data. The possible ways of photoacclimation of corals within the range of tolerance to light are shown: (1) adaptation to bright light (>70% incident photosynthetic active radiation, PAR_s), (2) adaptation to moderate shade (50–10% PAR_s), and (3) adaptation to extremely low light (<5% PAR_s). In each of the ways, general and specific mechanisms and reactions are involved in photoacclimation of corals. Adaptive changes take place in plant (zooxanthellae) and animal (polyps) components of the symbiotic organism. They involve morphology, physiology, and biochemistry of colonial polyps and their zooxanthellae. During changes of the light regime, some adaptive reactions last several months and others occur within a few days. Some physiological and biochemical alterations occur as early as the next day after the light regime changes. The wide range of light tolerance of corals and a great number of mechanisms and reactions of photoacclimation allowing corals to adapt to bright and low light with minimum losses in their metabolic activity give grounds to classify them as a single ecological group of light- and shade-tolerant organisms.     

Adaptive Evolution of Multicolored Fluorescent Proteins in Reef-Building Corals

Here we investigate the evolutionary scenarios that led to the appearance of fluorescent color diversity in reef-building corals. We show that the mutations that have been responsible for the generation of new cyan and red phenotypes from the ancestral green were fixed with the help of positive natural selection. This fact strongly suggests that the color diversity is a product of adaptive evolution. An unexpected finding was a set of residues arranged as an intermolecular binding interface, which was also identified as a target of positive selection but is nevertheless not related to color diversification. We hypothesize that multicolored fluorescent proteins evolved as part of a mechanism regulating the relationships between the coral and its algal endosymbionts (zooxanthellae). We envision that the effect of the proteins’ fluorescence on algal physiology may be achieved not only through photosynthesis modulation, but also through regulatory photosensors analogous to phytochromes and cryptochromes of higher plants. Such a regulation would require relatively subtle, but spectrally precise, modifications of the light field. Evolution of such a mechanism would explain both the adaptive diversification of colors and the coevolutionary chase at the putative algae-protein binding interface in coral fluorescent proteins. Electronic Supplementary Material Electronic Supplementary material is available for this article at and accessible for authorised users. [Reviewing Editor: Dr. Rasmus Neilsen]     

Application of 1H-NMR Metabolomic Profiling for Reef-Building Corals

In light of global reef decline new methods to accurately, cheaply, and quickly evaluate coral metabolic states are needed to assess reef health. Metabolomic profiling can describe the response of individuals to disturbance (i.e., shifts in environmental conditions) across biological models and is a powerful approach for characterizing and comparing coral metabolism. For the first time, we assess the utility of a proton-nuclear magnetic resonance spectroscopy (¹H-NMR)-based metabolomics approach in characterizing coral metabolite profiles by 1) investigating technical, intra-, and inter-sample variation, 2) evaluating the ability to recover targeted metabolite spikes, and 3) assessing the potential for this method to differentiate among coral species. Our results indicate ¹H-NMR profiling of Porites compressa corals is highly reproducible and exhibits low levels of variability within and among colonies. The spiking experiments validate the sensitivity of our methods and showcase the capacity of orthogonal partial least squares discriminate analysis (OPLS-DA) to distinguish between profiles spiked with varying metabolite concentrations (0 mM, 0.1 mM, and 10 mM). Finally, ¹H-NMR metabolomics coupled with OPLS-DA, revealed species-specific patterns in metabolite profiles among four reef-building corals (Pocillopora damicornis, Porites lobata, Montipora aequituberculata, and Seriatopora hystrix). Collectively, these data indicate that ¹H-NMR metabolomic techniques can profile reef-building coral metabolomes and have the potential to provide an integrated picture of the coral phenotype in response to environmental change.     

Partial Mortality in Porites Corals: Variation among Philippine Reefs

Partial mortality or tissue necrosis was quantified in the massive scleractinian coral Porites at three sites in The Philippines (Bolinao, NW Luzon; Puerto Galera, Mindoro; and El Nido, N Palawan). Overall, 15 ± 1 (mean ± 1 standard error, 642 replicates) percent of colony area was dead, mean colony area was 1135 plusmn; 127 cm², and lesion density was 1.7 ± 0.1 dm^—2. Total live coral cover varied between 20 and 63% in belt transects, and Porites and Acropora cover were inversely correlated. ANOVA models incorporating effects of site, colony size, sedimentation rates, wave exposure and depth were highly significant but explained only a small proportion of the variation observed in lesion density and percent dead area (respectively 8 and 2%). Lesion density was found to vary significantly with site (contributed 29% to this explained variance), decrease with increasing colony area (33%), and increase with increasing sedimentation (23%) and wave exposure (14%). Colony size was significantly explained by the factor site (contributing 61% to the total 29% explained variance) and depth (34%), with the smallest colonies being observed in Bolinao and the largest in El Nido. Densities of lesions were highest in Bolinao, intermediate in Puerto Galera, and lowest in El Nido. This pattern is parallel to intensity of human reef exploitation and opposite to that in colony size, live coral cover and Acropora cover. Since only a small part of the observed variance in partial mortality estimators was explained by the ANOVAs, other factors not quantified here must have been more important (e.g. disease incidence, predation, human exploitation).     

Species Composition of Corals in Silty Shallows of the Bai Thu Long Archipelago (Gulf of Tonkin,South China Sea)

Coral communities were investigated on 16 reefs in open and closed bays, in straits, and around capes of different islands of the Bai Thu Long Archipelago (Gulf of Tonkin, South China Sea). Variations observed in the communities of Acropora aspera + Goniopora columna, Goniopora stokesi + Galaxea fascicularis, Acropora aspera, Goniopora columna, and Pavona decussata + Pavona cactus are caused by different ecological conditions and may be linked with sedimentation flux within the water column and near the bottom, as well as with the rate of integral water exchange and resuspension above the coral settlements.     

Marine Reserves Enhance the Recovery of Corals on Caribbean Reefs

The fisheries and biodiversity benefits of marine reserves are widely recognised but there is mounting interest in exploiting the importance of herbivorous fishes as a tool to help ecosystems recover from climate change impacts. This approach might be particularly suitable for coral reefs, which are acutely threatened by climate change, yet the trophic cascades generated by reserves are strong enough that they might theoretically enhance the rate of coral recovery after disturbance. However, evidence for reserves facilitating coral recovery has been lacking. Here we investigate whether reductions in macroalgal cover, caused by recovery of herbivorous parrotfishes within a reserve, have resulted in a faster rate of coral recovery than in areas subject to fishing. Surveys of ten sites inside and outside a Bahamian marine reserve over a 2.5-year period demonstrated that increases in coral cover, including adjustments for the initial size-distribution of corals, were significantly higher at reserve sites than those in non-reserve sites. Furthermore, macroalgal cover was significantly negatively correlated with the change in total coral cover over time. Recovery rates of individual species were generally consistent with small-scale manipulations on coral-macroalgal interactions, but also revealed differences that demonstrate the difficulties of translating experiments across spatial scales. Size-frequency data indicated that species which were particularly affected by high abundances of macroalgae outside the reserve had a population bottleneck restricting the supply of smaller corals to larger size classes. Importantly, because coral cover increased from a heavily degraded state, and recovery from such states has not previously been described, similar or better outcomes should be expected for many reefs in the region. Reducing herbivore exploitation as part of an ecosystem-based management strategy for coral reefs appears to be justified.     

The Sino-Vietnamese Agreement on Maritime Boundary Delimitation in the Gulf of Tonkin

This article addresses the recently ratified Sino-Vietnamese Boundary Delimitation Agreement in the Gulf of Tonkin and its implications for bilateral cooperation and development of friendly and neighboring relations between China and Vietnam. As the first maritime boundary line for China, the delimitation in the Gulf of Tonkin is indicative of China's positive attitude towards the resolution of other maritime issues with Vietnam and other neighboring countries.     

Maritime Delimitation and Fishery Cooperation in the Tonkin Gulf

In June 2004 China and Vietnam ratified both a maritime boundary agreement for the Tonkin Gulf (Beibu Gulf) and a fisheries cooperation agreement for the Gulf. These agreements end years of negotiation and debate regarding the rights of the respective states to the ocean areas and resources in the Gulf.     

Communities of Coral Reefs of Central Vietnam

Communities of macrobenthos on fringing reefs of central Vietnam were studied by the hydrobiological method using scuba diving. The species composition (more than 600 species), population densities of common species of the macrobenthos, and the degree of coverage of the substrate by corals and macrophytes were examined. Bottom communities classified on the basis of these data are indicative of the similarity of the investigated reefs with those of southern and northern Vietnam.     

Deep-Sequencing Method for Quantifying Background Abundances of Symbiodinium Types: Exploring the Rare Symbiodinium Biosphere in Reef-Building Corals

The capacity of reef-building corals to associate with environmentally-appropriate types of endosymbionts from the dinoflagellate genus Symbiodinium contributes significantly to their success at local scales. Additionally, some corals are able to acclimatize to environmental perturbations by shuffling the relative proportions of different Symbiodinium types hosted. Understanding the dynamics of these symbioses requires a sensitive and quantitative method of Symbiodinium genotyping. Electrophoresis methods, still widely utilized for this purpose, are predominantly qualitative and cannot guarantee detection of a background type below 10% of the total Symbiodinium population. Here, the relative abundances of four Symbiodinium types (A13, C1, C3, and D1) in mixed samples of known composition were quantified using deep sequencing of the internal transcribed spacer of the ribosomal RNA gene (ITS-2) by means of Next Generation Sequencing (NGS) using Roche 454. In samples dominated by each of the four Symbiodinium types tested, background levels of the other three types were detected when present at 5%, 1%, and 0.1% levels, and their relative abundances were quantified with high (A13, C1, D1) to variable (C3) accuracy. The potential of this deep sequencing method for resolving fine-scale genetic diversity within a symbiont type was further demonstrated in a natural symbiosis using ITS-1, and uncovered reef-specific differences in the composition of Symbiodinium microadriaticum in two species of acroporid corals (Acropora digitifera and A. hyacinthus) from Palau. The ability of deep sequencing of the ITS locus (1 and 2) to detect and quantify low-abundant Symbiodinium types, as well as finer-scale diversity below the type level, will enable more robust quantification of local genetic diversity in Symbiodinium populations. This method will help to elucidate the role that background types have in maximizing coral fitness across diverse environments and in response to environmental change.     

Present-Day State of Coral Reefs of Nha Trang Bay (Southern Vietnam) and Possible Reasons for the Disturbance of Habitats of Scleractinian Corals

Our investigations were conducted from 1990 to 2002. Sampling of bottom sediments and biological objects, as well as photo and video shooting, were performed during scuba diving. The state of the environment and coral reef communities was assessed using the chemical–analytical, fluorometric, and luminometric methods, as well as the Ames test and the transect technique. The research results suggest that the spectrum and distribution pattern of persistent congeners of PCDD/Fs (dioxins) in bottom sediments are similar to those of the defoliant Agent Orange and that the bottom sediments are toxic and display photo inhibition and a mutagenic effect. The bottom of the bay is heavily silted throughout its depth. Many large dead colonies of corals without mechanical damage were observed everywhere. The total coverage by live corals in all sites investigated does not exceed 30%. Although, without a doubt, many factors contributed much to the disturbance of the bay ecosystems, the actual trigger for the degradation of the coral reefs seems to be the input of dioxin-containing chemicals used as defoliants during the American–Vietnamese war (Vietnam War).     

Reef-Building Corals—Symbiotic Autotrophic Organisms: 1. General Structure,Feeding Pattern,and Light-Dependent Distribution in the Shelf

General features of structure, feeding patterns, mutualistic relationships between animal and plant, as well as light-dependent distribution on the shelf, are described on the basis of the original and literature data for reef-building hermatypic corals (mainly scleractinians).     

Assessment of Epinephelus coioides (Hamilton, 1822) aquaculture systems in the Gulf of Tonkin,Vietnam, by using fish parasites

The parasite fauna of Epinephelus coioides can be used as an indicator to assess different aquaculture systems. A total of 133 E. coioides were collected from fishermen (wild fish) and marine fish farms (net cages, pond farms) in the Gulf of Tonkin (Ha Long, Cat Ba and Nghia Hung), Vietnam in 2013 and 2014, and examined for the presence of protozoan and metazoan parasites. Different patterns of parasite infections were found. The monogenean Pseudorhabdosynochus spp. was predominant in the net cage system; and the protozoans Trichodina spp. and Cryptocaryon irritans were found exclusively in pond and net cage cultivation sites in Nghia Hung and Cat Ba. These three species were assessed in addition to seven further indicator species and two dominance indices, to visualize the potential of assessing different aquaculture systems in Vietnam. This suggests the use of former shrimp ponds for grouper cultivation, reducing the pressure to further extend net cage cultivation in already highly influenced mariculture regions in Vietnam.     

Occurrence and Distribution of Stony Corals in the Gulf of Cariaco,Venezuela

Occurrence and distribution of stony corals (Milleporina and Scleractinia) in the Gulf of Cariaco were investigated quantitatively using skin-diving methods. Unusual temperature conditions due to cold upwelling limit coral growth to a depth of 15 m. A clear vertical zonation of species was observed, with five zones, in order of increasing depth, dominated by Porites porites, Millepora complanata, Siderastrea siderea, Dichocoenia stokesi, and Solenastrea hyades. Twenty-one species of scleractinians and three species of hydrocorals have so far been found in the gulf. Of the total area covered by corals, over 50% is inhabited by Siderastrea siderea, Millepora spp., and Porites porites. A horizontal zonation of the littoral was established based on species composition and density of the coral cover.     

Coral communities in the northern part of the Gulf of Tonkin, south China Sea

The reefs and benthic communities of 15 islands in Ha Long Bay, listed by UNESCO as a world heritage site, have been studied using scuba. Two reef types have been distinguished: structured and unstructured reefs. The changes in the composition and structure of reef-building corals takes place in these reefs under conditions of heavy silting in the surrounding waters. Poritids that primarily form the reef carcass become dominant scleractinians. More than 250 species of macrobenthos are found; the greatest species diversity was recorded in the reefs of Bo Hung and Kong Do islands, which we recommend be organized into a national nature reserve.