Characterising the distribution of mangroves along the southern coast of Vietnam using multi-spectral indices and a deep learning model.

Mangroves are an ecologically and economically valuable ecosystem that provides a range of ecological services, including habitat for a diverse range of plant and animal species, protection of coastlines from erosion and storms, carbon sequestration, and improvement of water quality. Despite their significant ecological role, in many areas, including in Vietnam, large scale losses have occurred, although restoration efforts have been underway. Understanding the scale of the loss and the efficacy of restoration requires high resolution temporal monitoring of mangrove cover on large scales. We have produced a time series of 10-m-resolution mangrove cover maps using the Multispectral Instrument on the Sentinel 2 satellites and with this tool measured the changes in mangrove distribution on the Vietnamese Southern Coast (VSC). We extracted the annual mangrove cover ranging from 2016 to 2023 using a deep learning model with a U-Net architecture based on 17 spectral indices. Additionally, a comparison of misclassification by the model with global products was conducted, indicating that the U-Net architecture demonstrated superior performance when compared to experiments including multispectral bands of Sentinel-2 and time-series of Sentinel-1 data, as shown by the highest performing spectral indices. The generated performance metrics, including overall accuracy, precision, recall, and F1-score were above 90 % for entire years. Water indices were investigated as the most important variables for mangrove extraction. Our study revealed some misclassifications by global products such as World Cover and Global Mangrove Watch and highlighted the significance of our study for local analysis. While we did observe a loss of 34,778 ha (42.2 %) of mangrove area in the region, 47,688 ha (57.8 %) of new mangrove area appeared, resulting in a net gain of 12,910 ha (15.65 %) over the eight-year period of the study. The majority of new mangrove areas were concentrated in Ca Mau peninsulas and within estuaries undergoing recovery programs and natural recovery processes. Mangrove loss occurred in regions where industrial development, wind farm projects, reclaimed land, and shrimp pond expansion is occurring. Our study provides a theoretical framework as well as up-to-date data for mapping and monitoring mangrove cover change that can be readily applied at other sites.

Planning hydrological restoration of coastal wetlands: Key model considerations and solutions.

The hydrological restoration of coastal wetlands is an emerging approach for mitigating and adapting to climate change and enhancing ecosystem services such as improved water quality and biodiversity. This paper synthesises current knowledge on selecting appropriate modelling approaches for hydrological restoration projects. The selection of a modelling approach is based on project-specific factors, such as costs, risks, and uncertainties, and aligns with the overall project objectives. We provide guidance on model selection, emphasising the use of simpler and less expensive modelling approaches when appropriate, and identifying situations when models may not be required for project managers to make informed decisions. This paper recognises and supports the widespread use of hydrological restoration in coastal wetlands by bridging the gap between hydrological science and restoration practices. It underscores the significance of project objectives, budget, and available data and offers decision-making frameworks, such as decision trees, to aid in matching modelling methods with specific project outcomes.

Mangroves provide blue carbon ecological value at a low freshwater cost.

"Blue carbon" wetland vegetation has a limited freshwater requirement. One type, mangroves, utilizes less freshwater during transpiration than adjacent terrestrial ecoregions, equating to only 43% (average) to 57% (potential) of evapotranspiration ([Formula: see text]). Here, we demonstrate that comparative consumptive water use by mangrove vegetation is as much as 2905 kL H2O ha-1 year-1 less than adjacent ecoregions with [Formula: see text]-to-[Formula: see text] ratios of 47-70%. Lower porewater salinity would, however, increase mangrove [Formula: see text]-to-[Formula: see text] ratios by affecting leaf-, tree-, and stand-level eco-physiological controls on transpiration. Restricted water use is also additive to other ecosystem services provided by mangroves, such as high carbon sequestration, coastal protection and support of biodiversity within estuarine and marine environments. Low freshwater demand enables mangroves to sustain ecological values of connected estuarine ecosystems with future reductions in freshwater while not competing with the freshwater needs of humans. Conservative water use may also be a characteristic of other emergent blue carbon wetlands.

Characterizing the spatial distribution of coral reefs in the South-Central Coast region of Viet Nam using Planetscope imagery.

This study aims to understand the spatial distribution of coral reefs in the central region of Viet Nam. We classified live coral cover in Son Tra Peninsula (ST) and Cu Lao Cham Island (CLC) in the South-Central Coast Region of Viet Nam using the Maximum Likelihood Classifier on 3 m Planetscope imagery. Confusion matrices and the accuracy of the classifier were assessed using field data (1,543 and 1,560 photographs in ST and CLC, respectively). The results showed that the reef's width ranged from 30 to 300 m across the study site, and we were able to detect live coral cover across a depth gradient of 2 to 6 m below the sea surface. The overall accuracies of the classifier (the Kappa coefficient) were 76.78% (0.76) and 78.08% (0.78) for ST and CLC, respectively. We found that 60.25% of coral reefs in ST were unhealthy and the live coral cover was less than 50%, while 25.75% and 11.46% of those in CLC were in good and excellent conditions, respectively. This study demonstrates the feasibility of utilizing Planetscope imagery to monitor shallow coral reefs of small islands at a high spatial resolution of 3 m. The results of this study provide valuable information for coral reef protection and conservation.

Author Correction: Future response of global coastal wetlands to sea-level rise.

Change history: In Fig. 2b of this Letter, 'Relative wetland change (km2)' should have read 'Relative wetland change (%)' and equations (2) and (3) have been changed from 'RSLRcrit = (m × TRe) × Sed + i' and 'Sedcrit = (RSLR - i)/(m × TRe)', respectively. The definition of the variables in equation (2) has been updated. These errors have been corrected online.

Future response of global coastal wetlands to sea-level rise.

The response of coastal wetlands to sea-level rise during the twenty-first century remains uncertain. Global-scale projections suggest that between 20 and 90 per cent (for low and high sea-level rise scenarios, respectively) of the present-day coastal wetland area will be lost, which will in turn result in the loss of biodiversity and highly valued ecosystem services1-3. These projections do not necessarily take into account all essential geomorphological4-7 and socio-economic system feedbacks8. Here we present an integrated global modelling approach that considers both the ability of coastal wetlands to build up vertically by sediment accretion, and the accommodation space, namely, the vertical and lateral space available for fine sediments to accumulate and be colonized by wetland vegetation. We use this approach to assess global-scale changes in coastal wetland area in response to global sea-level rise and anthropogenic coastal occupation during the twenty-first century. On the basis of our simulations, we find that, globally, rather than losses, wetland gains of up to 60 per cent of the current area are possible, if more than 37 per cent (our upper estimate for current accommodation space) of coastal wetlands have sufficient accommodation space, and sediment supply remains at present levels. In contrast to previous studies1-3, we project that until 2100, the loss of global coastal wetland area will range between 0 and 30 per cent, assuming no further accommodation space in addition to current levels. Our simulations suggest that the resilience of global wetlands is primarily driven by the availability of accommodation space, which is strongly influenced by the building of anthropogenic infrastructure in the coastal zone and such infrastructure is expected to change over the twenty-first century. Rather than being an inevitable consequence of global sea-level rise, our findings indicate that large-scale loss of coastal wetlands might be avoidable, if sufficient additional accommodation space can be created through careful nature-based adaptation solutions to coastal management.

Mangrove dieback during fluctuating sea levels.

Recent evidence indicates that climate change and intensification of the El Niño Southern Oscillation (ENSO) has increased variation in sea level. Although widespread impacts on intertidal ecosystems are anticipated to arise from the sea level seesaw associated with climate change, none have yet been demonstrated. Intertidal ecosystems, including mangrove forests are among those ecosystems that are highly vulnerable to sea level rise, but they may also be vulnerable to sea level variability and extreme low sea level events. During 16 years of monitoring of a mangrove forest in Mangrove Bay in north Western Australia, we documented two forest dieback events, the most recent one being coincident with the large-scale dieback of mangroves in the Gulf of Carpentaria in northern Australia. Diebacks in Mangrove Bay were coincident with periods of very low sea level, which were associated with increased soil salinization of 20-30% above pre-event levels, leading to canopy loss, reduced Normalized Difference Vegetation Index (NDVI) and reduced recruitment. Our study indicates that an intensification of ENSO will have negative effects on some mangrove forests in parts of the Indo-Pacific that will exacerbate other pressures.

The effects of elevated CO2 and eutrophication on surface elevation gain in a European salt marsh.

Salt marshes can play a vital role in mitigating the effects of global environmental change by dissipating incident storm wave energy and, through accretion, tracking increasing water depths consequent upon sea level rise. Atmospheric CO2 concentrations and nutrient availability are two key variables that can affect the biological processes that contribute to marsh surface elevation gain. We measured the effects of CO2 concentrations and nutrient availability on surface elevation change in intact mixed-species blocks of UK salt marsh using six open-top chambers receiving CO2 -enriched (800 ppm) or ambient (400 ppm) air. We found more rapid surface elevation gain in elevated CO2 conditions: an average increase of 3.4 mm over the growing season relative to ambient CO2 . Boosted regression analysis to determine the relative influence of different parameters on elevation change identified that a 10% reduction in microbial activity in elevated CO2 -grown blocks had a positive influence on elevation. The biomass of Puccinellia maritima also had a positive influence on elevation, while other salt marsh species (e.g. Suaeda maritima) had no influence or a negative impact on elevation. Reduced rates of water use by the vegetation in the high CO2 treatment could be contributing to elevation gain, either directly through reduced soil shrinkage or indirectly by decreasing microbial respiration rates due to lower redox levels in the soil. Eutrophication did not influence elevation change in either CO2 treatment despite doubling aboveground biomass. The role of belowground processes (transpiration, root growth and decomposition) in the vertical adjustment of European salt marshes, which are primarily minerogenic in composition, could increase as atmospheric CO2 concentrations rise and should be considered in future wetland models for the region. Elevated CO2 conditions could enhance resilience in vulnerable systems such as those with low mineral sediment supply or where migration upwards within the tidal frame is constrained.

The effects of CO2 and nutrient fertilisation on the growth and temperature response of the mangrove Avicennia germinans.

In order to understand plant responses to both the widespread phenomenon of increased nutrient inputs to coastal zones and the concurrent rise in atmospheric CO2 concentrations, CO2-nutrient interactions need to be considered. In addition to its potential stimulating effect on photosynthesis and growth, elevated CO2 affects the temperature response of photosynthesis. The scarcity of experiments testing how elevated CO2 affects the temperature response of tropical trees hinders our ability to model future primary productivity. In a glasshouse study, we examined the effects of elevated CO2 (800 ppm) and nutrient availability on seedlings of the widespread mangrove Avicennia germinans. We assessed photosynthetic performance, the temperature response of photosynthesis, seedling growth and biomass allocation. We found large synergistic gains in both growth (42 %) and photosynthesis (115 %) when seedlings grown under elevated CO2 were supplied with elevated nutrient concentrations relative to their ambient growing conditions. Growth was significantly enhanced under elevated CO2 only under high-nutrient conditions, mainly in above-ground tissues. Under low-nutrient conditions and elevated CO2, root volume was more than double that of seedlings grown under ambient CO2 levels. Elevated CO2 significantly increased the temperature optimum for photosynthesis by ca. 4 °C. Rising CO2 concentrations are likely to have a significant positive effect on the growth rate of A. germinans over the next century, especially in areas where nutrient availability is high.

The vulnerability of Indo-Pacific mangrove forests to sea-level rise.

Sea-level rise can threaten the long-term sustainability of coastal communities and valuable ecosystems such as coral reefs, salt marshes and mangroves. Mangrove forests have the capacity to keep pace with sea-level rise and to avoid inundation through vertical accretion of sediments, which allows them to maintain wetland soil elevations suitable for plant growth. The Indo-Pacific region holds most of the world's mangrove forests, but sediment delivery in this region is declining, owing to anthropogenic activities such as damming of rivers. This decline is of particular concern because the Indo-Pacific region is expected to have variable, but high, rates of future sea-level rise. Here we analyse recent trends in mangrove surface elevation changes across the Indo-Pacific region using data from a network of surface elevation table instruments. We find that sediment availability can enable mangrove forests to maintain rates of soil-surface elevation gain that match or exceed that of sea-level rise, but for 69 per cent of our study sites the current rate of sea-level rise exceeded the soil surface elevation gain. We also present a model based on our field data, which suggests that mangrove forests at sites with low tidal range and low sediment supply could be submerged as early as 2070.

The effect of atmospheric carbon dioxide concentrations on the performance of the mangrove Avicennia germinans over a range of salinities.

By increasing water use efficiency and carbon assimilation, increasing atmospheric CO2 concentrations could potentially improve plant productivity and growth at high salinities. To assess the effect of elevated CO2 on the salinity response of a woody halophyte, we grew seedlings of the mangrove Avicennia germinans under a combination of five salinity treatments [from 5 to 65 parts per thousand (ppt)] and three CO2 concentrations (280, 400 and 800 ppm). We measured survivorship, growth rate, photosynthetic gas exchange, root architecture and foliar nutrient and ion concentrations. The salinity optima for growth shifted higher with increasing concentrations of CO2 , from 0 ppt at 280 ppm to 35 ppt at 800 ppm. At optimal salinity conditions, carbon assimilation rates were significantly higher under elevated CO2 concentrations. However, at salinities above the salinity optima, salinity had an expected negative effect on mangrove growth and carbon assimilation, which was not alleviated by elevated CO2 , despite a significant improvement in photosynthetic water use efficiency. This is likely due to non-stomatal limitations to growth at high salinities, as indicated by our measurements of foliar ion concentrations that show a displacement of K(+) by Na(+) at elevated salinities that is not affected by CO2 . The observed shift in the optimal salinity for growth with increasing CO2 concentrations changes the fundamental niche of this species and could have significant effects on future mangrove distribution patterns and interspecific interactions.

Regulation of water balance in mangroves.

BACKGROUND: Mangroves are a group of highly salt-tolerant woody plants. The high water use efficiency of mangroves under saline conditions suggests that regulation of water transport is a crucial component of their salinity tolerance. SCOPE: This review focuses on the processes that contribute to the ability of mangroves to maintain water uptake and limit water loss to the soil and the atmosphere under saline conditions, from micro to macro scales. These processes include: (1) efficient filtering of the incoming water to exclude salt; (2) maintenance of internal osmotic potentials lower than that of the rhizosphere; (3) water-saving properties; and (4) efficient exploitation of less-saline water sources when these become available. CONCLUSIONS: Mangroves are inherently plastic and can change their structure at the root, leaf and stand levels in response to salinity in order to exclude salt from the xylem stream, maintain leaf hydraulic conductance, avoid cavitation and regulate water loss (e.g. suberization of roots and alterations of leaf size, succulence and angle, hydraulic anatomy and biomass partitioning). However, much is still unknown about the regulation of water uptake in mangroves, such as how they sense and respond to heterogeneity in root zone salinity, the extent to which they utilize non-stomatally derived CO2 as a water-saving measure and whether they can exploit atmospheric water sources.

How mangrove forests adjust to rising sea level.

Mangroves are among the most well described and widely studied wetland communities in the world. The greatest threats to mangrove persistence are deforestation and other anthropogenic disturbances that can compromise habitat stability and resilience to sea-level rise. To persist, mangrove ecosystems must adjust to rising sea level by building vertically or become submerged. Mangroves may directly or indirectly influence soil accretion processes through the production and accumulation of organic matter, as well as the trapping and retention of mineral sediment. In this review, we provide a general overview of research on mangrove elevation dynamics, emphasizing the role of the vegetation in maintaining soil surface elevations (i.e. position of the soil surface in the vertical plane). We summarize the primary ways in which mangroves may influence sediment accretion and vertical land development, for example, through root contributions to soil volume and upward expansion of the soil surface. We also examine how hydrological, geomorphological and climatic processes may interact with plant processes to influence mangrove capacity to keep pace with rising sea level. We draw on a variety of studies to describe the important, and often under-appreciated, role that plants play in shaping the trajectory of an ecosystem undergoing change.

The effect of nutrient enrichment on the growth, nucleic acid concentrations, and elemental stoichiometry of coral reef macroalgae.

The growth rate hypothesis (GRH) links growth rates with organism elemental stoichiometry. Support for the GRH was found for many animal species, but less so for plants. This is the first study to test the GRH in macroalgae. Tropical coral reef macroalgae from three lineages, Caulerpa serrulata (Chlorophyta), Laurencia intricata (Rhodophyta), and Sargassum polyphyllum (Phaeophyceae) were grown enriched with nitrogen or phosphorous and under control conditions at Heron Island on the Great Barrier Reef, Australia. Growth rate, photosynthesis, nucleic acid composition, and elemental stoichiometry were measured. Nutrient enrichment had positive effects on photosynthetic rates and on investment in RNA. However, growth rate was not correlated with either photosynthetic rates or RNA content; thus, we did not find support for the GRH in tropical macroalgae. Macroalgae, especially L. intricata, accumulated P to very high levels (>0.6% of dry weight). The growth rate response to tissue P concentrations was unimodal. Above 0.21%, P accumulation had negative effects on growth. Nitrogen was not stored, but evidence of futile cycling was observed. The capacity to store large amounts of P is probably an adaptation to the low and patchy nutrient environment of the tropical oceans.

Differential responses of the mangrove Avicennia marina to salinity and abscisic acid.

Salinisation of the soil can cause plant water deficits, ion and nutrient imbalances and toxic reactions. The halophyte, Avicennia marina (Forssk.) Vierh., is a mangrove that tolerates a wide range of soil salinities. In order to understand how salinity affects plant growth and functioning and how salinity responses are influenced by the water deficit signalling hormone abscisic acid (ABA) we grew A. marina seedlings under two non-growth limiting salinities: 60% seawater and 90% seawater and with and without exogenously supplied ABA. We measured growth, photosynthesis, sap flow, aquaporin gene expression, hydraulic anatomy and nutrient status as well as sap ABA concentrations. ABA addition resulted in a drought phenotype (reduced sap flow, transpiration rates and photosynthesis and increased water use efficiency and aquaporin expression). In contrast, growth in high salinity did not lead to responses that are typical for water deficits, but rather, could be characterised as drought avoidance strategies (no reduction in sap flow, transpiration rates and photosynthesis and reduced aquaporin expression). Tissue nutrient concentrations were higher in seedlings grown at high salinities. We did not find evidence for a role for ABA in the mangrove salinity response, suggesting ABA is not produced directly in response to high concentrations of NaCl ions.

IMPORTANCE OF MACRO- VERSUS MICROSTRUCTURE IN MODULATING LIGHT LEVELS INSIDE CORAL COLONIES(1).

Adjusting the light exposure and capture of their symbiotic photosynthetic dinoflagellates (genus Symbiodinium Freud.) is central to the success of reef-building corals (order Scleractinia) across high spatio-temporal variation in the light environment of coral reefs. We tested the hypothesis that optical properties of tissues in some coral species can provide light management at the tissue scale comparable to light modulation by colony architecture in other species. We compared within-tissue scalar irradiance in two coral species from the same light habitat but with contrasting colony growth forms: branching Stylophora pistillata and massive Lobophyllia corymbosa. Scalar irradiance at the level of the symbionts (2 mm into the coral tissues) were <10% of ambient irradiance and nearly identical for the two species, despite substantially different light environments at the tissue surface. In S. pistillata, light attenuation (90% relative to ambient) was observed predominantly at the colony level as a result of branch-to-branch self-shading, while in L. corymbosa, near-complete light attenuation (97% relative to ambient) was occurring due to tissue optical properties. The latter could be explained partly by differences in photosynthetic pigment content in the symbiont cells and pigmentation in the coral host tissue. Our results demonstrate that different strategies of light modulation at colony, polyp, and cellular levels by contrasting morphologies are equally effective in achieving favorable irradiances at the level of coral photosymbionts.

Intense storms and the delivery of materials that relieve nutrient limitations in mangroves of an arid zone estuary.

Tropical cyclones can be devastating to ecosystems, but they can also result in pulses of fresh water and sediments delivered in floodwaters to the coastal zone. In the arid zone the pulses provided by cyclones may be particularly important for the maintenance of productivity. We examined the impacts of Cyclone Pancho on growth and nutrient limitations to growth in mangroves on the arid coast of Western Australia. We found that growth of trees was enhanced after the cyclone, more than doubling their rates of stem extension. Fertilisation studies showed that before the cyclone tree growth was nutrient limited. After the cyclone fertilisation treatments had no significant effect on growth, indicating nutrients had been delivered during the storm. Additionally, before the cyclone the efficiency of resorption of phosphorus and nitrogen from senescent leaves was higher than after the cyclone, suggesting that nutrient availability was enhanced. Analysis of stable isotopes of leaf tissue indicated that the cyclone was associated with small changes in water use efficiency, consistent with decreased soil salinity associated with the cyclone. There was, however, significant reductions in δ15N indicating enhanced N supply potentially from a new source. We conclude that in the arid zone, floodwater associated with cyclones is important for the delivery of nutrient subsidies that stimulate mangrove growth and that predicted future reductions in the frequency of cyclones will have negative impacts on the productivity of these ecosystems.

Nutrition of mangroves.

Mangrove forests dominate the world's tropical and subtropical coastlines. Similar to other plant communities, nutrient availability is one of the major factors influencing mangrove forest structure and productivity. Many mangrove soils have extremely low nutrient availability, although nutrient availability can vary greatly among and within mangrove forests. Nutrient-conserving processes in mangroves are well developed and include evergreeness, resorption of nutrients prior to leaf fall, the immobilization of nutrients in leaf litter during decomposition, high root/shoot ratios and the repeated use of old root channels. Both nitrogen-use efficiency and nutrient resorption efficiencies in mangroves are amongst the highest recorded for angiosperms. A complex range of interacting abiotic and biotic factors controls the availability of nutrients to mangrove trees, and mangroves are characteristically plastic in their ability to opportunistically utilize nutrients when these become available. Nitrogen and phosphorus have been implicated as the nutrients most likely to limit growth in mangroves. Ammonium is the primary form of nitrogen in mangrove soils, in part as a result of anoxic soil conditions, and tree growth is supported mainly by ammonium uptake. Nutrient enrichment is a major threat to marine ecosystems. Although mangroves have been proposed to protect the marine environment from land-derived nutrient pollution, nutrient enrichment can have negative consequences for mangrove forests and their capacity for retention of nutrients may be limited.

Coral skeletons defend against ultraviolet radiation.

BACKGROUND: Many coral reef organisms are photosynthetic or have evolved in tight symbiosis with photosynthetic symbionts. As such, the tissues of reef organisms are often exposed to intense solar radiation in clear tropical waters and have adapted to trap and harness photosynthetically active radiation (PAR). High levels of ultraviolet radiation (UVR) associated with sunlight, however, represent a potential problem in terms of tissue damage. METHODOLOGY/PRINCIPAL FINDINGS: By measuring UVR and PAR reflectance from intact and ground bare coral skeletons we show that the property of calcium carbonate skeletons to absorb downwelling UVR to a significant extent, while reflecting PAR back to the overlying tissue, has biological advantages. We placed cnidarians on top of bare skeletons and a UVR reflective substrate and showed that under ambient UVR levels, UVR transmitted through the tissues of cnidarians placed on top of bare skeletons were four times lower compared to their counterparts placed on a UVR reflective white substrate. In accordance with the lower levels of UVR measured in cnidarians on top of coral skeletons, a similar drop in UVR damage to their DNA was detected. The skeletons emitted absorbed UVR as yellow fluorescence, which allows for safe dissipation of the otherwise harmful radiation. CONCLUSIONS/SIGNIFICANCE: Our study presents a novel defensive role for coral skeletons and reveals that the strong UVR absorbance by the skeleton can contribute to the ability of corals, and potentially other calcifiers, to thrive under UVR levels that are detrimental to most marine life.

Photoreactivation is the main repair pathway for UV-induced DNA damage in coral planulae.

The larvae of most coral species spend some time in the plankton, floating just below the surface and hence exposed to high levels of ultraviolet radiation (UVR). The high levels of UVR are potentially stressful and damaging to DNA and other cellular components, such as proteins, reducing survivorship. Consequently, mechanisms to either shade (prevent) or repair damage potentially play an important role. In this study, the role of photoreactivation in the survival of coral planulae was examined. Photoreactivation is a light-stimulated response to UV-damaged DNA in which photolyase proteins repair damaged DNA. Photoreactivation rates, as well as the localization of photolyase, were explored in planulae under conditions where photoreactivation was or was not inhibited. The results indicate that photoreactivation is the main DNA repair pathway in coral planulae, repairing UV-induced DNA damage swiftly (K=1.75 h(-1) and a half-life of repair of 23 min), with no evidence of any light-independent DNA repair mechanisms, such as nucleotide excision repair (NER), at work. Photolyase mRNA was localized to both the ectoderm and endoderm of the larvae. The amount of cell death in the coral planulae increased significantly when photoreactivation was inhibited, by blocking photoreactivating light. We found that photoreactivation, along with additional UV shielding in the form of five mycosporine-like amino acids, are sufficient for survival in surface tropical waters and that planulae do not accumulate DNA damage despite being exposed to high UVR.