Environmental Concentrations of Herbicide Prometryn Render Stress-Tolerant Corals Susceptible to Ocean Warming.

Global warming has caused the degradation of coral reefs around the world. While stress-tolerant corals have demonstrated the ability to acclimatize to ocean warming, it remains unclear whether they can sustain their thermal resilience when superimposed with other coastal environmental stressors. We report the combined impacts of a photosystem II (PSII) herbicide, prometryn, and ocean warming on the stress-tolerant coral Galaxea fascicularis through physiological and omics analyses. The results demonstrate that the heat-stress-induced inhibition of photosynthetic efficiency in G. fascicularis is exacerbated in the presence of prometryn. Transcriptomics and metabolomics analyses indicate that the prometryn exposure may overwhelm the photosystem repair mechanism in stress-tolerant corals, thereby compromising their capacity for thermal acclimation. Moreover, prometryn might amplify the adverse effects of heat stress on key energy and nutrient metabolism pathways and induce a stronger response to oxidative stress in stress-tolerant corals. The findings indicate that the presence of prometryn at environmentally relevant concentrations would render corals more susceptible to heat stress and exacerbate the breakdown of coral Symbiodiniaceae symbiosis. The present study provides valuable insights into the necessity of prioritizing PSII herbicide pollution reduction in coral reef protection efforts while mitigating the effects of climate change.

Spatio-temporal coupling analysis and tipping points detection of China's coastal integrated land-human activity-ocean system.

The coastal zone is typically highly developed and its ocean environment is vastly exposed to the onshore activities. Land-based pollution, as the "metabolite" of terrestrial human activities, significantly impacts the ocean environment. Although numerous studies have investigated these effects, few have quantified the interactions among land-human activity-ocean across both spatial and temporal scales. In this study, we have developed a land-human activity-ocean systemic framework integrating the coupling coordination degree model and tipping point to quantify the spatiotemporal dynamic interaction mechanism among the land-based pollution, human activities, and ocean environment in China from 2001 to 2020. Our findings revealed that the overall coupling coordination degree of the China's coastal zone increased by 36.9 % over last two decades. Furthermore, the effect of human activities on China's coastal environment remained within acceptable thresholds, as no universal tipping points for coastal pollution or ocean environment has been found over the 20-year period. Notably, the lag time for algal blooms, the key indicator of ocean environment health, was found to be 0-3 years in response to the land economic development and 0-4 years in response to land-based pollution. Based on the differences in spatiotemporal interactions among land-human activity-ocean system, we employed cluster analysis to categorize China's coastal provinces into four types and to develop appropriate management measures. Quantifying the interaction mechanism within the land-human activity-ocean system could aid decision-makers in creating sustainable coastal development strategies. This enables efficient use of land and ocean resources, supports coastal conservation and restoration efforts, and fosters effective management recommendations to enhance coastal sustainability and resilience.

Barriers and corridors of gene flow in an urbanized tropical reef system.

Information about the distribution of alleles among marine populations is critical for determining patterns of genetic connectivity that are essential in modern conservation planning. To estimate population connectivity in Singapore's urbanized equatorial reef system, we analysed single nucleotide polymorphisms (SNPs) from two species of reef-building corals with distinct life histories. For Porites sp., a broadcast-spawning coral, we found cryptic lineages that were differentially distributed at inshore and central-offshore sites that could be attributed to contemporary surface current regimes. Near panmixia was observed for Pocillopora acuta with differentiation of colonies at the farthest site from mainland Singapore, a possible consequence of the brooding nature and relatively long pelagic larval duration of the species. Furthermore, analysis of recent gene flow showed that 60-80% of colonies in each population were nonmigrants, underscoring self-recruitment as an important demographic process in this reef system. Apart from helping to enhance the management of Singapore's coral reef ecosystems, findings here pave the way for better understanding of the evolution of marine populations in South-East Asia.

Coral community composition and carbonate production in an urbanized seascape.

Coastal urbanization causes environmental modifications that directly and indirectly influence the distribution and functioning of coral reefs. However, the capacity of urban infrastructure to support corals and vertically accrete is less understood. Here, we investigated if coral communities on reefs and seawalls in Singapore are distinct, and examined the environmental variables influencing coral carbonate production. Surveys at 22 sites yielded 134 coral species, with richness significantly higher on reefs. Coral cover and Shannon index did not differ between habitat types. Community composition was distinct between habitat types, with seawalls supporting a higher proportion of massive and thick-plating species. 'Distance from mainland' was the single most important variable influencing normalized carbonate production rates (a function of species-specific linear extension rate and skeletal bulk density and site coral cover), which were higher further from the mainland where human activity and development pressures were greater. Our results indicate that environmental filtering strongly shapes coral communities and may influence ecosystem functioning in Singapore's urbanized reef system. The findings will guide the management of reefs on increasingly urbanized coastlines.

Gene Expression and Photophysiological Changes in Pocillopora acuta Coral Holobiont Following Heat Stress and Recovery.

The ability of corals to withstand changes in their surroundings is a critical survival mechanism for coping with environmental stress. While many studies have examined responses of the coral holobiont to stressful conditions, its capacity to reverse responses and recover when the stressor is removed is not well-understood. In this study, we investigated among-colony responses of Pocillopora acuta from two sites with differing distance to the mainland (Kusu (closer to the mainland) and Raffles Lighthouse (further from the mainland)) to heat stress through differential expression analysis of target genes and quantification of photophysiological metrics. We then examined how these attributes were regulated after the stressor was removed to assess the recovery potential of P. acuta. The fragments that were subjected to heat stress (2 °C above ambient levels) generally exhibited significant reduction in their endosymbiont densities, but the extent of recovery following stress removal varied depending on natal site and colony. There were minimal changes in chl a concentration and maximum quantum yield (Fv/Fm, the proportion of variable fluorescence (Fv) to maximum fluorescence (Fm)) in heat-stressed corals, suggesting that the algal endosymbionts' Photosystem II was not severely compromised. Significant changes in gene expression levels of selected genes of interest (GOI) were observed following heat exposure and stress removal among sites and colonies, including Actin, calcium/calmodulin-dependent protein kinase type IV (Camk4), kinesin-like protein (KIF9), and small heat shock protein 16.1 (Hsp16.1). The most responsive GOIs were Actin, a major component of the cytoskeleton, and the adaptive immune-related Camk4 which both showed significant reduction following heat exposure and subsequent upregulation during the recovery phase. Our findings clearly demonstrate specific responses of P. acuta in both photophysiological attributes and gene expression levels, suggesting differential capacity of P. acuta corals to tolerate heat stress depending on the colony, so that certain colonies may be more resilient than others.

Responses of urban reef corals during the 2016 mass bleaching event.

Predicting the bleaching responses of corals is crucial in light of frequent heat stress events to manage further losses of biodiversity and ecosystem functioning, especially for reefs impacted by urbanisation. We examined if the coral cover and community at various Singapore sites changed during the 2016 global coral bleaching event. Bleaching prevalence varied widely among sites in June 2016, and was best explained by site and coral species. While some sites were minimally impacted, others registered significant decreases in coral cover and community changes persisting till March 2017, when normal colouration was mostly regained by corals. Bleaching susceptibility was associated with larger corallites in hermaphrodites and smaller corallites in gonochores (probably due to the cost of maintaining dual sexual functions in hermaphrodites), and with increasing proximity between polyps (likely because thermal damage would be less contained among polyps with greater physiological integration). However, bleaching resilience-the capacity to regain baseline pigmentation-was poorly explained by the traits studied. Our findings suggest that the interplay between local conditions and species composition strongly affects bleaching outcomes on urbanised reefs, and underscore the utility of coral traits for predicting bleaching responses to help in formulating appropriate management strategies.

From marine park to future genomic observatory? Enhancing marine biodiversity assessments using a biocode approach.

Few tropical marine sites have been thoroughly characterised for their animal species, even though they constitute the largest proportion of multicellular diversity. A number of focused biodiversity sampling programmes have amassed immense collections to address this shortfall, but obstacles remain due to the lack of identification tools and large proportion of undescribed species globally. These problems can be partially addressed with DNA barcodes ("biocodes"), which have the potential to facilitate the estimation of species diversity and identify animals to named species via barcode databases. Here, we present the first results of what is intended to be a sustained, systematic study of the marine fauna of Singapore's first marine park, reporting more than 365 animal species, determined based on DNA barcodes and/or morphology represented by 931 specimens (367 zooplankton, 564 macrofauna including 36 fish). Due to the lack of morphological and molecular identification tools, only a small proportion could be identified to species solely based on either morphology (24.5%) or barcodes (24.6%). Estimation of species numbers for some taxa was difficult because of the lack of sufficiently clear barcoding gaps. The specimens were imaged and added to "Biodiversity of Singapore" (http://singapore.biodiversity.online), which now contains images for > 13,000 species occurring in the country.

Bleaching and necrosis of staghorn coral (Acropora formosa) in laboratory assays: Immediate impact of LDPE microplastics.

The impact of low-density polyethylene (LDPE) microplastics (<100 µm; P100-A P100-B, P100-C, 100-200 µm; P200, 200-500 µm; P500) on Acropora formosa was investigated. This study investigated the bleaching and necrosis extent of A. formosa caused by LDPE contamination via laboratory assay. The staghorn coral ingested the microplastics, resulting in bleaching and necrosis that concomitantly occurred with the release of zooxanthellae. P100-A experimentation was the worst case, showing bleaching by day 2 (10.8 ±â€¯2.2%) and continued bleaching to 93.6% ±â€¯2.0 by day 14 followed by 5.9 ±â€¯2.5% necrosis. The overall results confirmed that the LDPE concentration impacts coral health. We highlighted that microplastics have been ingested and partially egested. Their presence showed either a direct or indirect impact on coral polyps via direct interaction or through photosynthesis perturbation due to microplastics that cover the coral surface.

Seasonal variation of Sargassum ilicifolium (Phaeophyceae) growth on equatorial coral reefs.

Temporal and spatial variations in Sargassum ilicifolium thallus density and length were investigated on equatorial coral reefs in Singapore from November 2011 to October 2012. Thalli density varied little throughout the year, however, we found strong seasonal patterns in thallus length and identified temperature as the significant driver. Sargassum ilicifolium reached maximum length in December (110.39 ± 2.37 cm) during periods of cooler water temperatures, and minimum length in May (9.88 ± 0.48 cm) during periods of warmer water temperatures. Significant spatial variation was also observed for both thallus density and length of S. ilicifolium among reefs. Within reefs, densities of S. ilicifolium were higher on reef flats (20.40 ± 0.40 individuals · 0.25 m-2 ) compared to upper reef slopes (5.66 ± 0.23 individuals · 0.25 m-2 ). Our findings highlight that marked seasonality in the growth of canopy-forming macroalgae can occur within equatorial reef systems where temperature ranges are restricted (<3°C).

Urban coral reefs: Degradation and resilience of hard coral assemblages in coastal cities of East and Southeast Asia.

Given predicted increases in urbanization in tropical and subtropical regions, understanding the processes shaping urban coral reefs may be essential for anticipating future conservation challenges. We used a case study approach to identify unifying patterns of urban coral reefs and clarify the effects of urbanization on hard coral assemblages. Data were compiled from 11 cities throughout East and Southeast Asia, with particular focus on Singapore, Jakarta, Hong Kong, and Naha (Okinawa). Our review highlights several key characteristics of urban coral reefs, including "reef compression" (a decline in bathymetric range with increasing turbidity and decreasing water clarity over time and relative to shore), dominance by domed coral growth forms and low reef complexity, variable city-specific inshore-offshore gradients, early declines in coral cover with recent fluctuating periods of acute impacts and rapid recovery, and colonization of urban infrastructure by hard corals. We present hypotheses for urban reef community dynamics and discuss potential of ecological engineering for corals in urban areas.

The origin and evolution of coral species richness in a marine biodiversity hotspot.

The Coral Triangle (CT) region of the Indo-Pacific realm harbors an extraordinary number of species, with richness decreasing away from this biodiversity hotspot. Despite multiple competing hypotheses, the dynamics underlying this regional diversity pattern remain poorly understood. Here, we use a time-calibrated evolutionary tree of living reef coral species, their current geographic ranges, and model-based estimates of regional rates of speciation, extinction, and geographic range shifts to show that origination rates within the CT are lower than in surrounding regions, a result inconsistent with the long-standing center of origin hypothesis. Furthermore, endemism of coral species in the CT is low, and the CT endemics are older than relatives found outside this region. Overall, our model results suggest that the high diversity of reef corals in the CT is largely due to range expansions into this region of species that evolved elsewhere. These findings strongly support the notion that geographic range shifts play a critical role in generating species diversity gradients. They also show that preserving the processes that gave rise to the striking diversity of corals in the CT requires protecting not just reefs within the hotspot, but also those in the surrounding areas.

Differential Response of Coral Assemblages to Thermal Stress Underscores the Complexity in Predicting Bleaching Susceptibility.

Coral bleaching events have been predicted to occur more frequently in the coming decades with global warming. The susceptibility of corals to bleaching during thermal stress episodes is dependent on many factors and an understanding of these underlying drivers is crucial for conservation management. In 2013, a mild bleaching episode ensued in response to elevated sea temperature on the sediment-burdened reefs in Singapore. Surveys of seven sites highlighted variable bleaching susceptibility among coral genera-Pachyseris and Podabacia were the most impacted (31% of colonies of both genera bleached). The most susceptible genera such as Acropora and Pocillopora, which were expected to bleach, did not. Susceptibility varied between less than 6% and more than 11% of the corals bleached, at four and three sites respectively. Analysis of four of the most bleached genera revealed that a statistical model that included a combination of the factors (genus, colony size and site) provided a better explanation of the observed bleaching patterns than any single factor alone. This underscored the complexity in predicting the coral susceptibility to future thermal stress events and the importance of monitoring coral bleaching episodes to facilitate more effective management of coral reefs under climate change.

Augmenting the post-transplantation growth and survivorship of juvenile scleractinian corals via nutritional enhancement.

Size-dependent mortality influences the recolonization success of juvenile corals transplanted for reef restoration and assisting juvenile corals attain a refuge size would thus improve post-transplantation survivorship. To explore colony size augmentation strategies, recruits of the scleractinian coral Pocillopora damicornis were fed with live Artemia salina nauplii twice a week for 24 weeks in an ex situ coral nursery. Fed recruits grew significantly faster than unfed ones, with corals in the 3600, 1800, 600 and 0 (control) nauplii/L groups exhibiting volumetric growth rates of 10.65 ± 1.46, 4.69 ± 0.9, 3.64 ± 0.55 and 1.18 ± 0.37 mm3/week, respectively. Corals supplied with the highest density of nauplii increased their ecological volume by more than 74 times their initial size, achieving a mean final volume of 248.38 ± 33.44 mm3. The benefits of feeding were apparent even after transplantation to the reef. The corals in the 3600, 1800, 600 and 0 nauplii/L groups grew to final sizes of 4875 ± 260 mm3, 2036 ± 627 mm3, 1066 ± 70 mm3 and 512 ± 116 mm3, respectively. The fed corals had significantly higher survival rates than the unfed ones after transplantation (63%, 59%, 56% and 38% for the 3600, 1800, 600 and 0 nauplii/L treatments respectively). Additionally, cost-effectiveness analysis revealed that the costs per unit volumetric growth were drastically reduced with increasing feed densities. Corals fed with the highest density of nauplii were the most cost-effective (US$0.02/mm3), and were more than 12 times cheaper than the controls. This study demonstrated that nutrition enhancement can augment coral growth and post-transplantation survival, and is a biologically and economically viable option that can be used to supplement existing coral mariculture procedures and enhance reef restoration outcomes.

Contrasting patterns of coral bleaching susceptibility in 2010 suggest an adaptive response to thermal stress.

BACKGROUND: Coral bleaching events vary in severity, however, to date, the hierarchy of susceptibility to bleaching among coral taxa has been consistent over a broad geographic range and among bleaching episodes. Here we examine the extent of spatial and temporal variation in thermal tolerance among scleractinian coral taxa and between locations during the 2010 thermally induced, large-scale bleaching event in South East Asia. METHODOLOGY/PRINCIPAL FINDINGS: Surveys to estimate the bleaching and mortality indices of coral genera were carried out at three locations with contrasting thermal and bleaching histories. Despite the magnitude of thermal stress being similar among locations in 2010, there was a remarkable contrast in the patterns of bleaching susceptibility. Comparisons of bleaching susceptibility within coral taxa and among locations revealed no significant differences between locations with similar thermal histories, but significant differences between locations with contrasting thermal histories (Friedman = 34.97; p<0.001). Bleaching was much less severe at locations that bleached during 1998, that had greater historical temperature variability and lower rates of warming. Remarkably, Acropora and Pocillopora, taxa that are typically highly susceptible, although among the most susceptible in Pulau Weh (Sumatra, Indonesia) where respectively, 94% and 87% of colonies died, were among the least susceptible in Singapore, where only 5% and 12% of colonies died. CONCLUSIONS/SIGNIFICANCE: The pattern of susceptibility among coral genera documented here is unprecedented. A parsimonious explanation for these results is that coral populations that bleached during the last major warming event in 1998 have adapted and/or acclimatised to thermal stress. These data also lend support to the hypothesis that corals in regions subject to more variable temperature regimes are more resistant to thermal stress than those in less variable environments.

Periophthalmus variabilis (Teleostei: Gobiidae: Oxudercinae), a valid species of mudskipper, and a re-diagnosis of Periophthalmus novemradiatus.

The current concept of Periophthalmus novemradiatus (Hamilton, 1822) includes two species. Procurement of fresh material and examination of extant type specimens revealed that P. novemradiatus differs from P. variabilis Eggert, 1935, primarily in the number of anal fin rays (I, 12 or 13 vs. I, 10-12); the lateral scale series (62-67 vs. 48-60); the extent of fusion of the pelvic fins (56.0-90.0% vs. 32.0-48.8%); and the patterns of the first and second dorsal fins.

More evidence for pervasive paraphyly in scleractinian corals: systematic study of Southeast Asian Faviidae (Cnidaria; Scleractinia) based on molecular and morphological data.

Coral taxonomy and systematics continue to be plagued by a host of problems. Due to high phenotypic variability within species, morphological approaches have often failed to recognize natural taxa, and molecular techniques have yet to be applied to many groups. Here, we summarize the levels of paraphyly found for scleractinian corals and test, based on new data, whether paraphyly is also a significant problem in Faviidae, the second-most speciose hermatypic scleractinian family. Using both DNA sequence and morphological data we find that, regardless of analysis technique (maximum parsimony, maximum likelihood and Bayesian likelihood), many conventional taxonomic groups are not monophyletic. Based on two mitochondrial markers (COI and a noncoding region) that we amplified for 81 samples representing 41 faviid species and 13 genera, five genera that are represented by more than one species are paraphyletic, as is the family Faviidae. The morphological characters currently used to identify these corals similarly fail to recover many genera. Furthermore, trees based on both data types are incongruent, and total evidence analysis does little to salvage conventional taxonomic groupings. Morphological convergence, phenotypic variability in response to the environment, and recent speciation are likely causes for these conflicts, which suggest that the present classification of corals is in need of a major overhaul. We propose more detailed studies of problematic faviid taxa using standardized morphological, mitochondrial, and nuclear genetic markers to facilitate combining of data.

Slow mitochondrial COI sequence evolution at the base of the metazoan tree and its implications for DNA barcoding.

The evolution rates of mtDNA in early metazoans hold important implications for DNA barcoding. Here, we present a comprehensive analysis of intra- and interspecific COI variabilities in Porifera and Cnidaria (separately as Anthozoa, Hydrozoa, and Scyphozoa) using a data set of 619 sequences from 224 species. We found variation within and between species to be much lower in Porifera and Anthozoa compared to Medusozoa (Hydrozoa and Scyphozoa), which has divergences similar to typical metazoans. Given that recent evidence has shown that fungi also exhibit limited COI divergence, slow-evolving mtDNA is likely to be plesiomorphic for the Metazoa. Higher rates of evolution could have originated independently in Medusozoa and Bilateria or been acquired in the Cnidaria + Bilateria clade and lost in the Anthozoa. Low identification success and substantial overlap between intra- and interspecific COI distances render the Anthozoa unsuitable for DNA barcoding. Caution is also advised for Porifera and Hydrozoa because of relatively low identification success rates as even threshold divergence that maximizes the "barcoding gap" does not improve identification success.

Taxonomical and morphological notes on two species of mudskippers, Periophthalmus walailakae and Periophthalmodon schlosseri (Teleostei: Gobiidae) from Singapore.

The mudskipper Periophthalmus walailakae is recorded from Singapore, where it was previously misidentified as Periophthalmodon schlosseri, with which it is syntopic. Periophthalmus walailakae is distinguished from its congeners by the following combination of characters: pelvic fins completely united and shaped like a disk, and first dorsal fin dark brown or black, with a rounded posterior edge and a white distal margin. This species most closely resembles Pn. schlosseri but has one row of teeth on the upper jaw, scales on the isthmus, and a different upper lip and jaw morphology. Contrary to an earlier report, scales are present on the snout, interorbital, and isthmus of Ps. walailakae. The two species can also be distinguished by size, external morphology, and body color patterns.

Proportion and cluster analyses of the skull in various species of the tree shrews.

The skull adaptation was functional-morphologically examined in 14 species of the tree shrews. From the data of the proportion indices, the similarities were confirmed between T. minor and T. gracilis, T. tana and T. dorsalis, and T. longipes and T. glis. We demonstrated that the splanchnocranium was elongated in terrestrial T. tana and T. dorsalis and shortened in arboreal T. minor and T. gracilis from the proportion data. In both dendrogram from the matrix of the Q-mode correlation coefficients and scattergram from the canonical discriminant analysis, the morphological similarities in the skull shape suggested the terrestrial-insectivorous adaptation of T. tana and T. dorsalis, and the arboreal adaptation of T. minor and T. gracilis. Since the osteometrical skull similarities were indicated among the three species of Tupaia by cluster and canonical discriminant analyses, the arbo-terrestrial behavior and its functional-morphological adaptation may be commonly established in T. montana, T. longipes and T. glis.