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Showing papers in "Coral Reefs in 2017"


Journal ArticleDOI
TL;DR: It is found that structural complexity and reef zone are the strongest and most consistent predictors of reef fish abundance, biomass, species richness, and trophic structure, and that coral traits, diversity, and life histories provided additional predictive power for models of Reef fish assemblages, and were key drivers of structural complexity.
Abstract: With the ongoing loss of coral cover and the associated flattening of reef architecture, understanding the links between coral habitat and reef fishes is of critical importance. Here, we investigate whether considering coral traits and functional diversity provides new insights into the relationship between structural complexity and reef fish communities, and whether coral traits and community composition can predict structural complexity. Across 157 sites in Seychelles, Maldives, the Chagos Archipelago, and Australia's Great Barrier Reef, we find that structural complexity and reef zone are the strongest and most consistent predictors of reef fish abundance, biomass, species richness, and trophic structure. However, coral traits, diversity, and life histories provided additional predictive power for models of reef fish assemblages, and were key drivers of structural complexity. Our findings highlight that reef complexity relies on living corals-with different traits and life histories-continuing to build carbonate skeletons, and that these nuanced relationships between coral assemblages and habitat complexity can affect the structure of reef fish assemblages. Seascape-level estimates of structural complexity are rapid and cost effective with important implications for the structure and function of fish assemblages, and should be incorporated into monitoring programs.

201 citations


Journal ArticleDOI
TL;DR: In this paper, a small GoPro HERO4 with a modified lens mounted on a DJI Phantom 2 drone was used to perform a 10min flight and collect 306 aerial images with an overlap equal or greater than 90%.
Abstract: We propose a novel technique to measure the small-scale three-dimensional features of a shallow-water coral reef using a small drone equipped with a consumer-grade camera, a handheld GPS and structure from motion (SfM) algorithms. We used a GoPro HERO4 with a modified lens mounted on a DJI Phantom 2 drone (maximum total take-off weight <2 kg) to perform a 10 min flight and collect 306 aerial images with an overlap equal or greater than 90%. We mapped an area of 8380 m2, obtaining as output an ortho-rectified aerial photomosaic and a bathymetric digital elevation model (DEM) with a resolution of 0.78 and 1.56 cm pixel−1, respectively. Through comparison with airborne LiDAR data for the same area, we verified that the location of the ortho-rectified aerial photomosaic is accurate within ~1.4 m. The bathymetric difference between our DEM and the LiDAR dataset is −0.016 ± 0.45 m (1σ). Our results show that it is possible, in conditions of calm waters, low winds and minimal sun glint, to deploy consumer-grade drones as a relatively low-cost and rapid survey technique to produce multispectral and bathymetric data on shallow-water coral reefs. We discuss the utility of such data to monitor temporal changes in topographic complexity of reefs and associated biological processes.

192 citations


Journal ArticleDOI
TL;DR: Study of changes in cover of corals, algal turfs, benthic cyanobacterial mats, macroalgae, sponges and crustose coralline algae at four reef sites of the Caribbean islands of Curaçao and Bonaire over a time span of 40 yr suggests that reefs dominated by algae may be less stable than previously thought.
Abstract: Over the past decades numerous studies have reported declines in stony corals and, in many cases, phase shifts to fleshy macroalgae. However, long-term studies documenting changes in other benthic reef organisms are scarce. Here, we studied changes in cover of corals, algal turfs, benthic cyanobacterial mats, macroalgae, sponges and crustose coralline algae at four reef sites of the Caribbean islands of Curacao and Bonaire over a time span of 40 yr. Permanent 9 m2 quadrats at 10, 20, 30 and 40 m depth were photographed at 3- to 6-yr intervals from 1973 to 2013. The temporal and spatial dynamics in the six dominant benthic groups were assessed based on image point-analysis. Our results show consistent patterns of benthic community change with a decrease in the cover of calcifying organisms across all sites and depths from 32.6 (1973) to 9.2% (2013) for corals and from 6.4 to 1% for crustose coralline algae. Initially, coral cover was replaced by algal turfs increasing from 24.5 (1973) to 38% around the early 1990s. Fleshy macroalgae, still absent in 1973, also proliferated covering 12% of the substratum approximately 20 yr later. However, these new dominants largely declined in abundance from 2002 to 2013 (11 and 2%, respectively), marking the rise of benthic cyanobacterial mats. Cyanobacterial mats became the most dominant benthic component increasing from a mere 7.1 (2002) to 22.2% (2013). The observed increase was paralleled by a small but significant increase in sponge cover (0.5 to 2.3%). Strikingly, this pattern of degradation and phase change occurred over the reef slope down to mesophotic depths of 40 m. These findings suggest that reefs dominated by algae may be less stable than previously thought and that the next phase may be the dominance of slimy cyanobacterial mats with some sponges.

106 citations


Journal ArticleDOI
TL;DR: In this article, the authors validate the effectiveness of degree heating weeks (DHW) as a mass bleaching index by on-site historical observation at eight sites in the northwestern Pacific Ocean.
Abstract: Mass bleaching is the most significant threat to coral reefs. The United States National Oceanic and Atmospheric Administration monitors world sea surface temperature (SST) and releases warnings for bleaching based on degree heating weeks (DHW), which is the accumulation of temperature anomalies exceeding the monthly maximum mean SST for a given region. DHW values >4.0 °C-weeks are thought to induce bleaching, and those >8.0 °C-weeks are thought to result in widespread bleaching and some mortality. This study validates the effectiveness of DHW as a mass bleaching index by on-site historical observation at eight sites in the northwestern Pacific Ocean. The mass bleaching events occurred during different years at different sites. The recorded years of the bleaching events matched well with DHW values >8 °C-weeks, and the logistically projected probability of bleaching against DHW showed a positive relationship. DHW provides a reasonable threshold for bleaching.

100 citations


Journal ArticleDOI
TL;DR: The traditional mesophotic zone is home to three ecological communities in the GoMx, one that is confluent with shallow reefs, a distinct Mesophotic assemblage spanning 60–120 m, and a third that extends onto the outer continental shelf.
Abstract: We analyzed an extensive dataset of over 9000 benthic and suprabenthic species found throughout the Gulf of Mexico (GoMx) to assess whether mesophotic coral ecosystems represent distinct assemblages and evaluate their potential to serve as refugia for shallow reef communities. We assessed community structure of the overall benthic community from 0 to 300 m via non-metric multidimensional scaling (NMDS) of species presence across depth bands. We used the Jaccard index of similarity to calculate the proportion of shared species between adjacent depth bands, measure species turnover with depth, and assess taxonomic overlap between shallow reefs versus progressively deeper depth bands. NMDS ordinations showed that the traditionally defined mesophotic range (30–150 m) as a whole is not a distinct community. In contrast, taxonomically distinct communities, determined by hierarchical clustering, were found at 0–70, 60–120, 110–200, and 190–300 m. Clustering highlighted an important separation in the benthic community at ~60 m, which was especially important for actinopterygian fishes. Species turnover between adjacent depths decreased with depth for all taxa combined and individual taxa, with peaks at ~60, 90–120, and 190–200 m. Fishes showed lower turnover from shallow to upper mesophotic depths (0–50 m) than all taxa combined, a substantial peak at 60 m, followed by a precipitous and continued decline in turnover thereafter. Taxonomic overlap between shallow (0–20 m) and progressively deeper zones declined steadily with depth in all taxa and individual taxa, suggesting that mid- and lower mesophotic habitats have less (but not inconsequential) potential to serve as refugia (60–150 m, 15–25% overlap with shallow habitats) than upper mesophotic zones (30–60 m, 30–45% overlap with shallow habitats) for all taxa combined. We conclude that the traditional mesophotic zone is home to three ecological communities in the GoMx, one that is confluent with shallow reefs, a distinct mesophotic assemblage spanning 60–120 m, and a third that extends onto the outer continental shelf.

82 citations


Journal ArticleDOI
TL;DR: Gas chromatography–mass spectrometry (GC–MS) metabolomics was applied to detect changes in the intracellular free metabolite pools of hospite dinoflagellate symbionts and their coral hosts during early- and late-stage thermal bleaching to further understanding of the metabolic changes that occur to symbiont and host (and its associated microorganisms) during thermalBleaching.
Abstract: Rising seawater temperatures pose a significant threat to the persistence of coral reefs. Despite the importance of these systems, major gaps remain in our understanding of how thermal stress and bleaching affect the metabolic networks that underpin holobiont function. We applied gas chromatography–mass spectrometry (GC–MS) metabolomics to detect changes in the intracellular free metabolite pools (polar and semi-polar compounds) of in hospite dinoflagellate symbionts and their coral hosts (and any associated microorganisms) during early- and late-stage thermal bleaching (a reduction of approximately 50 and 70% in symbiont density, respectively). We detected characteristic changes to the metabolite profiles of each symbiotic partner associated with individual cellular responses to thermal, oxidative and osmotic stress, which progressed with the severity of bleaching. Alterations were also indicative of changes to energy-generating and biosynthesis pathways in both partners, with a shift to the increased catabolism of lipid stores. Specifically, in symbiont intracellular metabolite pools, we observed accumulations of multiple free fatty acids, plus the chloroplast-associated antioxidant alpha-tocopherol. In the host, we detected a decline in the abundance of pools of multiple carbohydrates, amino acids and intermediates, in addition to the antioxidant ascorbate. These findings further our understanding of the metabolic changes that occur to symbiont and host (and its associated microorganisms) during thermal bleaching. These findings also provide further insight into the largely undescribed roles of free metabolite pools in cellular homeostasis, signalling and acclimation to thermal stress in the cnidarian–dinoflagellate symbiosis.

74 citations


Journal ArticleDOI
TL;DR: An extensive dataset collected by restoration practitioners is combined to document early restoration success metrics in Florida and Puerto Rico, USA and provides the basis for a stop-light indicator framework for new or existing restoration programs to evaluate their performance.
Abstract: Coral gardening plays an important role in the recovery of depleted populations of threatened Acropora cervicornis in the Caribbean. Over the past decade, high survival coupled with fast growth of in situ nursery corals have allowed practitioners to create healthy and genotypically diverse nursery stocks. Currently, thousands of corals are propagated and outplanted onto degraded reefs on a yearly basis, representing a substantial increase in the abundance, biomass, and overall footprint of A. cervicornis. Here, we combined an extensive dataset collected by restoration practitioners to document early (1–2 yr) restoration success metrics in Florida and Puerto Rico, USA. By reporting region-specific data on the impacts of fragment collection on donor colonies, survivorship and productivity of nursery corals, and survivorship and productivity of outplanted corals during normal conditions, we provide the basis for a stop-light indicator framework for new or existing restoration programs to evaluate their performance. We show that current restoration methods are very effective, that no excess damage is caused to donor colonies, and that once outplanted, corals behave just as wild colonies. We also provide science-based benchmarks that can be used by programs to evaluate successes and challenges of their efforts, and to make modifications where needed. We propose that up to 10% of the biomass can be collected from healthy, large A. cervicornis donor colonies for nursery propagation. We also propose the following benchmarks for the first year of activities for A. cervicornis restoration: (1) >75% live tissue cover on donor colonies; (2) >80% survivorship of nursery corals; and (3) >70% survivorship of outplanted corals. Finally, we report productivity means of 4.4 cm yr−1 for nursery corals and 4.8 cm yr−1 for outplants as a frame of reference for ranking performance within programs. Such benchmarks, and potential subsequent adaptive actions, are needed to fully assess the long-term success of coral restoration and species recovery programs.

72 citations


Journal ArticleDOI
TL;DR: The spatial patterns of stony corals are non-random and reflect fundamental life-history characteristics of the taxa, suggesting that the reef landscape may, in many cases, have important elements of spatial predictability.
Abstract: For sessile organisms such as reef-building corals, differences in the degree of dispersion of individuals across a landscape may result from important differences in life-history strategies or may reflect patterns of habitat availability. Descriptions of spatial patterns can thus be useful not only for the identification of key biological and physical mechanisms structuring an ecosystem, but also by providing the data necessary to generate and test ecological theory. Here, we used an in situ imaging technique to create large-area photomosaics of 16 plots at Palmyra Atoll, central Pacific, each covering 100 m2 of benthic habitat. We mapped the location of 44,008 coral colonies and identified each to the lowest taxonomic level possible. Using metrics of spatial dispersion, we tested for departures from spatial randomness. We also used targeted model fitting to explore candidate processes leading to differences in spatial patterns among taxa. Most taxa were clustered and the degree of clustering varied by taxon. A small number of taxa did not significantly depart from randomness and none revealed evidence of spatial uniformity. Importantly, taxa that readily fragment or tolerate stress through partial mortality were more clustered. With little exception, clustering patterns were consistent with models of fragmentation and dispersal limitation. In some taxa, dispersion was linearly related to abundance, suggesting density dependence of spatial patterning. The spatial patterns of stony corals are non-random and reflect fundamental life-history characteristics of the taxa, suggesting that the reef landscape may, in many cases, have important elements of spatial predictability.

63 citations


Journal ArticleDOI
TL;DR: Given an increase in fish species descriptions, completeness of taxonomic checklists, and improvement in species-level assignment with custom genetic databases as shown here, it is suggested that the Red Sea region may be ideal for further testing of the eDNA approach.
Abstract: Relatively small volumes of water may contain sufficient environmental DNA (eDNA) to detect target aquatic organisms via genetic sequencing. We therefore assessed the utility of eDNA to document the diversity of coral reef fishes in the central Red Sea. DNA from seawater samples was extracted, amplified using fish-specific 16S mitochondrial DNA primers, and sequenced using a metabarcoding workflow. DNA sequences were assigned to taxa using available genetic repositories or custom genetic databases generated from reference fishes. Our approach revealed a diversity of conspicuous, cryptobenthic, and commercially relevant reef fish at the genus level, with select genera in the family Labridae over-represented. Our approach, however, failed to capture a significant fraction of the fish fauna known to inhabit the Red Sea, which we attribute to limited spatial sampling, amplification stochasticity, and an apparent lack of sequencing depth. Given an increase in fish species descriptions, completeness of taxonomic checklists, and improvement in species-level assignment with custom genetic databases as shown here, we suggest that the Red Sea region may be ideal for further testing of the eDNA approach.

62 citations


Journal ArticleDOI
TL;DR: This study paired population structure from comprehensive genetic sampling and biophysical larval transport modeling to describe how spiny lobster (Panulirus argus) population differentiation is related to biological oceanography, providing a framework for future explorations of wide-scale larval dispersal and marine connectivity.
Abstract: Connectivity, the exchange of individuals among locations, is a fundamental ecological process that explains how otherwise disparate populations interact. For most marine organisms, dispersal occurs primarily during a pelagic larval phase that connects populations. We paired population structure from comprehensive genetic sampling and biophysical larval transport modeling to describe how spiny lobster (Panulirus argus) population differentiation is related to biological oceanography. A total of 581 lobsters were genotyped with 11 microsatellites from ten locations around the greater Caribbean. The overall F ST of 0.0016 (P = 0.005) suggested low yet significant levels of structuring among sites. An isolation by geographic distance model did not explain spatial patterns of genetic differentiation in P. argus (P = 0.19; Mantel r = 0.18), whereas a biophysical connectivity model provided a significant explanation of population differentiation (P = 0.04; Mantel r = 0.47). Thus, even for a widely dispersing species, dispersal occurs over a continuum where basin-wide larval retention creates genetic structure. Our study provides a framework for future explorations of wide-scale larval dispersal and marine connectivity by integrating empirical genetic research and probabilistic modeling.

61 citations


Journal ArticleDOI
TL;DR: This work used natural feeding surfaces, covered with late successional stage reef-grown algal turfs, to examine turf algal removal by the lined bristletooth and brown surgeonfish and found that C. striatus ingests detritus and sediment, leaving mature algalTurfs relatively intact, while A. nigrofuscus directly removes and ingests turf algae.
Abstract: The lined bristletooth, Ctenochaetus striatus, and the brown surgeonfish, Acanthurus nigrofuscus, are among the most abundant surgeonfishes on Indo-Pacific coral reefs. Yet, the functional role of these species has been the focus of an ongoing debate lasting at least six decades. Specifically, to what extent are C. striatus herbivorous like the visually similar A. nigrofuscus? To address this question, we used natural feeding surfaces, covered with late successional stage reef-grown algal turfs, to examine turf algal removal by the two species. Surfaces exposed to C. striatus in laboratory experiments exhibited no significant reductions in turf length or area covered by turfing algae. In marked contrast, A. nigrofuscus reduced turf length by 51% and area covered by turfing algae by 15% in 1 h. The gut contents of specimens from the reef revealed that A. nigrofuscus predominantly ingests algae (the dominant item in 79.6–94.7% of gut content quadrats), while C. striatus ingests detritus and sediments (dominant in 99.6–100% of quadrats). Therefore, C. striatus ingests detritus and sediment, leaving mature algal turfs relatively intact, while A. nigrofuscus directly removes and ingests turf algae. The function of C. striatus differs from cropping herbivorous surgeonfishes such as A. nigrofuscus. On coral reefs, C. striatus brush detrital aggregates from algal turfs, removing microorganisms, organic detritus and inorganic sediment. Confusion over the functional role of C. striatus may stem from an inability to fit it into a single functional category.

Journal ArticleDOI
TL;DR: In this paper, the authors investigated phylogenetic relationships and patterns of population genetic structure and diversity in all four fire coral species occurring in the Brazilian Province to understand patterns of speciation and biogeography in the genus.
Abstract: Fire corals are the only branching corals in the South Atlantic and provide an important ecological role as habitat-builders in the region. With three endemic species (Millepora brazilensis, M. nitida and M. laboreli) and one amphi-Atlantic species (M. alcicornis), fire coral diversity in the Brazilian Province rivals that of the Caribbean Province. Phylogenetic relationships and patterns of population genetic structure and diversity were investigated in all four fire coral species occurring in the Brazilian Province to understand patterns of speciation and biogeography in the genus. A total of 273 colonies from the four species were collected from 17 locations spanning their geographic ranges. Sequences from the 16S ribosomal DNA (rDNA) were used to evaluate phylogenetic relationships. Patterns in genetic diversity and connectivity were inferred by measures of molecular diversity, analyses of molecular variance, pairwise differentiation, and by spatial analyses of molecular variance. Morphometrics of the endemic species M. braziliensis and M. nitida were evaluated by discriminant function analysis; macro-morphological characters were not sufficient to distinguish the two species. Genetic analyses showed that, although they are closely related, each species forms a well-supported clade. Furthermore, the endemic species characterized a distinct biogeographic barrier: M. braziliensis is restricted to the north of the Sao Francisco River, whereas M. nitida occurs only to the south. Millepora laboreli is restricted to a single location and has low genetic diversity. In contrast, the amphi-Atlantic species M. alcicornis shows high genetic connectivity within the Brazilian Province, and within the Caribbean Province (including Bermuda), despite low levels of gene flow between these populations and across the tropical Atlantic. These patterns reflect the importance of the Amazon–Orinoco Plume and the Mid-Atlantic Barrier as biogeographic barriers, and suggest that, while M. alcicornis is capable of long-distance dispersal, the three endemics have restricted ranges and more limited dispersal capabilities.

Journal ArticleDOI
TL;DR: This result highlights the exceptional, atypical thermal tolerance of dominant GoA branching corals and validates the efficacy of protecting GoA reefs from local stressors if they are to serve as a coral refuge in the face of global sea temperature rise.
Abstract: Despite rapidly rising sea surface temperatures and recurrent positive temperature anomalies, corals in the Gulf of Aqaba (GoA) rarely experience thermal bleaching Elsewhere, mass coral bleaching has been observed in corals when the water temperature exceeds 1–2 °C above the local maximum monthly mean (MMM) This threshold value or “bleaching rule” has been used to create predictive models of bleaching from satellite sea surface temperature observations, namely the “degree heating week” index This study aimed to characterize the physiological changes of dominant reef building corals from the GoA in response to a temperature and light stress gradient Coral collection and experiments began after a period of 14 consecutive days above MMM in the field Stylophora pistillata showed negligible changes in symbiont and host physiology parameters after accumulating up to 94 degree heating weeks during peak summer temperatures, for which the index predicts widespread bleaching and some mortality This result demonstrates acute thermal tolerance in S pistillata from the GoA and deviation from the bleaching rule In a second experiment after 4 weeks at 4 °C above peak summer temperatures, S pistillata and Acropora eurystoma in the high-light treatment visibly paled and suffered greater midday and afternoon photoinhibition compared to corals under low-light conditions (35% of high-light treatment) However, light, not temperature (alone or in synergy with light), was the dominant factor in causing paling and the effective quantum yield of corals at 4 °C above ambient was indistinguishable from those in the ambient control This result highlights the exceptional, atypical thermal tolerance of dominant GoA branching corals Concomitantly, it validates the efficacy of protecting GoA reefs from local stressors if they are to serve as a coral refuge in the face of global sea temperature rise

Journal ArticleDOI
TL;DR: Differences in coral community structure over time were found, driven primarily by a fourfold increase of one local species, Plesiastrea versipora, rather than recruitment of warm-affinity species, consistent with a net tropicalisation of temperate reefs.
Abstract: Globally, many temperate marine communities have experienced significant temperature increases over recent decades in the form of gradual warming and heatwaves. As a result, these communities are shifting towards increasingly subtropical and tropical species compositions. Expanding coral populations have been reported from several temperate reef ecosystems along warming coastlines; these changes have been attributed to direct effects of gradual warming over decades. In contrast, increases in coral populations following shorter-term extreme warming events have rarely been documented. In this study, we compared coral populations on 17 temperate reefs in Western Australia before (2005/06) and after (2013) multiple marine heatwaves (2010–2012) affected the entire coastline. We hypothesised that coral communities would expand and change as a consequence of increasing local populations and recruitment of warm-affinity species. We found differences in coral community structure over time, driven primarily by a fourfold increase of one local species, Plesiastrea versipora, rather than recruitment of warm-affinity species. Coral populations became strongly dominated by small size classes, indicative of recent increased recruitment or recruit survival. These changes were likely facilitated by competitive release of corals from dominant temperate seaweeds, which perished during the heatwaves, rather than driven by direct temperature effects. Overall, as corals are inherently warm-water taxa not commonly associated with seaweed-dominated temperate reefs, these findings are consistent with a net tropicalisation. Our study draws attention to processes other than gradual warming that also influence the trajectory of temperate reefs in a changing ocean.

Journal ArticleDOI
TL;DR: Data suggest that in this host species, host–symbiont specificity is a genetically determined trait, and if this level of specificity is widespread among other symbiotic cnidarian–algal symbioses may not be able to respond to rapid, climate change-associated environmental changes by means of between-generation switching of symbionts.
Abstract: Ocean warming and other anthropogenic stresses threaten the symbiosis between tropical reef cnidarians and their dinoflagellate endosymbionts (Symbiodinium). Offspring of many cnidarians acquire their algal symbionts from the environment, and such flexibility could allow corals to respond to environmental changes between generations. To investigate the effect of both habitat and host genotype on symbiont acquisition, we transplanted aposymbiotic offspring of the common Caribbean octocoral Briareum asbestinum to (1) an environmentally different habitat that lacked B. asbestinum and (2) an environmentally similar habitat where local adults harbored Symbiodinium phylotypes that differed from parental colonies. Symbiont acquisition and establishment of symbioses over time was followed using a within-clade DNA marker (23S chloroplast rDNA) and a within-phylotype marker (unique alleles at a single microsatellite locus). Early in the symbiosis, B. asbestinum juveniles harbored multiple symbiont phylotypes, regardless of source (parent or site). However, with time (~4 yr), offspring established symbioses with the symbiont phylotype dominant in the parental colonies, regardless of transplant location. Within-phylotype analyses of the symbionts revealed a similar pattern, with offspring acquiring the allelic variant common in symbionts in the parental population regardless of the environment in which the offspring was reared. These data suggest that in this host species, host–symbiont specificity is a genetically determined trait. If this level of specificity is widespread among other symbiotic cnidarians, many cnidarian–algal symbioses may not be able to respond to rapid, climate change-associated environmental changes by means of between-generation switching of symbionts.

Journal ArticleDOI
TL;DR: The ecological and physical structural changes that occurred across five atoll interior reefs in the southern Maldives using data collected at 6 and 12 months post-bleaching suggest that the physical structure of these reefs will need to decline further before effective recruitment and recovery can begin.
Abstract: Given the severity of the 2016 global bleaching event, there are major questions about how quickly reef communities will recover. Here, we explore the ecological and physical structural changes that occurred across five atoll interior reefs in the southern Maldives using data collected at 6 and 12 months post-bleaching. Following initial severe coral mortality, further minor coral mortality had occurred by 12 months post-bleaching, and coral cover is now low (<6%). In contrast, reef rugosity has continued to decline over time, and our observations suggest transitions to rubble-dominated states will occur in the near future. Juvenile coral densities in shallow fore-reef habitats are also exceptionally low (<6 individuals m−2), well below those measured 9–12 months following the 1998 bleaching event, and below recovery thresholds identified on other Indian Ocean reefs. Our findings suggest that the physical structure of these reefs will need to decline further before effective recruitment and recovery can begin.

Journal ArticleDOI
TL;DR: This is the first study using the newly developed BGS seabed mapping toolbox and an ROV-based microbathymetric grid to explore the environmental variables that control coral growth on cold-water coral reefs.
Abstract: Cold-water corals form substantial biogenic habitats on continental shelves and in deep-sea areas with topographic highs, such as banks and seamounts. In the Atlantic, many reef and mound complexes are engineered by Lophelia pertusa, the dominant framework-forming coral. In this study, a variety of mapping approaches were used at a range of scales to map the distribution of both cold-water coral habitats and individual coral colonies at the Mingulay Reef Complex (west Scotland). The new ArcGIS-based British Geological Survey (BGS) seabed mapping toolbox semi-automatically delineated over 500 Lophelia reef ‘mini-mounds’ from bathymetry data with 2-m resolution. The morphometric and acoustic characteristics of the mini-mounds were also automatically quantified and captured using this toolbox. Coral presence data were derived from high-definition remotely operated vehicle (ROV) records and high-resolution microbathymetry collected by a ROV-mounted multibeam echosounder. With a resolution of 0.35 × 0.35 m, the microbathymetry covers 0.6 km2 in the centre of the study area and allowed identification of individual live coral colonies in acoustic data for the first time. Maximum water depth, maximum rugosity, mean rugosity, bathymetric positioning index and maximum current speed were identified as the environmental variables that contributed most to the prediction of live coral presence. These variables were used to create a predictive map of the likelihood of presence of live cold-water coral colonies in the area of the Mingulay Reef Complex covered by the 2-m resolution data set. Predictive maps of live corals across the reef will be especially valuable for future long-term monitoring surveys, including those needed to understand the impacts of global climate change. This is the first study using the newly developed BGS seabed mapping toolbox and an ROV-based microbathymetric grid to explore the environmental variables that control coral growth on cold-water coral reefs.

Journal ArticleDOI
TL;DR: Strong shifts in the bacterial community of C. aspera are reported and it is hypothesized that these shifts are age related and could be influenced by the observed baseline differences in the microbiome of the 4- and 10-yr-old corals, bacteria–bacteria interactions, and/or host energetics.
Abstract: Shifts in the microbiome of the intertidal coral Coelastrea aspera (formally known as Goniastrea aspera) from Phuket, Thailand, were noted over the course of a 4-d period of spring tides. During this time, corals were naturally exposed to high temperatures, intense solar radiation, sub-aerial exposure and tidally induced water fluxes. Analysis of the 16S microbiome highlighted that the corals harbored both ‘core or stable’ communities and those which appeared to be more ‘transient or sporadic.’ Only relatively few microbial associates were classified as core microbes; the majority were transient or sporadic. Such transient associates were likely to have been governed by tidally induced variations in mucus thickness and water fluxes. Here we report strong shifts in the bacterial community of C. aspera over a short temporal scale. However, we also show significant differences in the timing of shifts between the two age groups of corals studied. More rapid changes (within 2 d of sub-aerial exposure) occurred within the 4-yr-old colonies, but a slightly delayed response was observed in the 10-yr-old colonies, whereby the microbial associates only changed after 4 d. We hypothesize that these shifts are age related and could be influenced by the observed baseline differences in the microbiome of the 4- and 10-yr-old corals, bacteria–bacteria interactions, and/or host energetics.

Journal ArticleDOI
TL;DR: The ctenidia is designated as the site of light-enhanced ammonia absorption with the absorbed ammonia being assimilated to glutamine by the host GS, the first report on light-dependent gene and protein expression of a host enzyme involved in nitrogen metabolism in an alga–invertebrate association.
Abstract: Giant clams harbor symbiotic zooxanthellae and manifest light-enhanced calcification. In light, they also increase the absorption and assimilation of exogenous ammonia, but the roles of the host and symbionts are unclear and the reason for light dependency remains enigmatic. Here, we report the complete coding cDNA sequence of a glutamine synthetase (GS), from the ctenidia (gill) of Tridacna squamosa. The deduced GS amino acid sequence was closely related to other animal GSs, indicating a host origin. The GS/GS transcript level and protein abundance increased significantly in the ctenidia after 12 h of light exposure. These results denote the ctenidia as the site of light-enhanced ammonia absorption with the absorbed ammonia being assimilated to glutamine by the host GS. Immunofluorescence microscopy confirmed that GS was expressed predominantly in the cytoplasm of the epithelial cells separating the tertiary water channels from the hemolymph inside the ctenidia. Hence, unlike symbiotic cnidarians, the host clam probably supplies exogenous nitrogen as glutamine from the ctenidia, through the hemolymph and tubular fluid, to the nitrogen-deficient zooxanthellae which reside extracellularly in the extensible outer mantle. This is the first report on light-dependent gene and protein expression of a host enzyme involved in nitrogen metabolism in an alga–invertebrate association.

Journal ArticleDOI
TL;DR: In this paper, an in situ optical backscatter sediment deposition sensor capable of measuring sedimentation over intervals of a few hours was deployed for 39 d in a highly turbid inshore area in the Great Barrier Reef.
Abstract: Sedimentation is considered the most widespread contemporary, human-induced perturbation on reefs, and yet if the problems associated with its estimation using sediment traps are recognized, there have been few reliable measurements made over time frames relevant to the local organisms. This study describes the design, calibration and testing of an in situ optical backscatter sediment deposition sensor capable of measuring sedimentation over intervals of a few hours. The instrument has been reconfigured from an earlier version to include 15 measurement points instead of one, and to have a more rugose measuring surface with a microtopography similar to a coral. Laboratory tests of the instrument with different sediment types, colours, particle sizes and under different flow regimes gave similar accumulation estimates to SedPods, but lower estimates than sediment traps. At higher flow rates (9–17 cm s−1), the deposition sensor and SedPods gave estimates >10× lower than trap accumulation rates. The instrument was deployed for 39 d in a highly turbid inshore area in the Great Barrier Reef. Sediment deposition varied by several orders of magnitude, occurring in either a relatively uniform (constant) pattern or a pulsed pattern characterized by short-term (4–6 h) periods of ‘enhanced’ deposition, occurring daily or twice daily and modulated by the tidal phase. For the whole deployment, which included several very high wind events and suspended sediment concentrations (SSCs) >100 mg L−1, deposition rates averaged 19 ± 16 mg cm−2 d−1. For the first half of the deployment, where SSCs varied from <1 to 28 mg L−1 which is more typical for the study area, the deposition rate averaged only 8 ± 5 mg cm−2 d−1. The capacity to measure sedimentation rates over a few hours is discussed in terms of examining the risk from sediment deposition associated with catchment run-off, natural wind/wave events and dredging activities.

Journal ArticleDOI
TL;DR: It is demonstrated that branches grown on the substratum had significantly higher skeletal density, measured using computerized tomography, and lower linear extension rates compared to water-column fragments, and suggested that there is genetically based variability in coral calcification performance.
Abstract: Staghorn coral, Acropora cervicornis, is a threatened species and the primary focus of western Atlantic reef restoration efforts to date. We compared linear extension, calcification rate, and skeletal density of nursery-raised A. cervicornis branches reared for 6 months either on blocks attached to substratum or hanging from PVC trees in the water column. We demonstrate that branches grown on the substratum had significantly higher skeletal density, measured using computerized tomography, and lower linear extension rates compared to water-column fragments. Calcification rates determined with buoyant weighing were not statistically different between the two grow-out methods, but did vary among coral genotypes. Whereas skeletal density and extension rates were plastic traits that depended on grow-out method, calcification rate was conserved. Our results show that the two rearing methods generate the same amount of calcium carbonate skeleton but produce colonies with different skeletal characteristics and suggest that there is genetically based variability in coral calcification performance.

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TL;DR: These genetic partitions at the western periphery of the Indo-Pacific reflect similar partitions recently discovered at the eastern periphery of The Hawaiian and the Marquesan Archipelagos, indicating that these disjunctive habitats at the ends of the range may serve as evolutionary incubators for coral reef organisms.
Abstract: The Arabian Sea is a heterogeneous region with high coral cover and warm stable conditions at the western end (Djibouti), in contrast to sparse coral cover, cooler temperatures, and upwelling at the eastern end (southern Oman). We tested for barriers to dispersal across this region (including the Gulf of Aden and Gulf of Oman), using mitochondrial DNA surveys of 11 reef fishes. Study species included seven taxa from six families with broad distributions across the Indo-Pacific and four species restricted to the Arabian Sea (and adjacent areas). Nine species showed no significant genetic partitions, indicating connectivity among contrasting environments spread across 2000 km. One butterflyfish (Chaetodon melannotus) and a snapper (Lutjanus kasmira) showed phylogenetic divergences of d = 0.008 and 0.048, respectively, possibly indicating cryptic species within these broadly distributed taxa. These genetic partitions at the western periphery of the Indo-Pacific reflect similar partitions recently discovered at the eastern periphery of the Indo-Pacific (the Hawaiian and the Marquesan Archipelagos), indicating that these disjunctive habitats at the ends of the range may serve as evolutionary incubators for coral reef organisms.

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TL;DR: The results suggest that a coral’s thermal tolerance is primarily and positively dependent on its symbiont density and is less related to its effective photochemical efficiency.
Abstract: Although it is well established that different coral species have different susceptibilities to thermal stress, the reasons behind this variation are still unclear. In this study, 384 samples across five dominant coral species were collected seasonally between September 2013 and August 2014 at Luhuitou fringing reef in Sanya, Hainan Island, northern South China Sea, and their algal symbiont density and effective photochemical efficiency (Φ PSII) were measured. The results indicated that both the Symbiodinium density and Φ PSII of corals were subject to significant interspecies and seasonal variations. Stress-tolerant coral species, including massive Porites lutea and plating Pavona decussata, had higher symbiont densities but lower Φ PSII compared to the vulnerable branching species of Acropora over the course of all four seasons. Seasonally, coral symbiont densities were the lowest during winter, while during the same period, Φ PSII of corals was at the highest point. Further analysis suggested that dissolved inorganic nutrients and upwelling in the reef area were probably responsible for the observed seasonal variations in symbiont density. The fact that Porites lutea has the lowest Φ PSII during all four seasons is likely related to their symbionts’ lower capacity to provide required photosynthates for calcification. These results suggest that a coral’s thermal tolerance is primarily and positively dependent on its symbiont density and is less related to its effective photochemical efficiency.

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TL;DR: In this paper, the authors measured dissepiment spacing to reconstruct multiple years of monthly skeletal extension rates in two Porites colonies from Palmyra Atoll and in another from Palau that bleached in 1998 under anomalously high sea temperatures.
Abstract: The skeletons of many reef-building corals are accreted with rhythmic structural patterns that serve as valuable sclerochronometers. Annual high- and low-density band couplets, visible in X-radiographs or computed tomography scans, are used to construct age models for paleoclimate reconstructions and to track variability in coral growth over time. In some corals, discrete, anomalously high-density bands, called “stress bands,” preserve information about coral bleaching. However, the mechanisms underlying the formation of coral skeletal density banding remain unclear. Dissepiments—thin, horizontal sheets of calcium carbonate accreted by the coral to support the living polyp—play a key role in the upward growth of the colony. Here, we first conducted a vital staining experiment to test whether dissepiments were accreted with lunar periodicity in Porites coral skeleton, as previously hypothesized. Over 6, 15, and 21 months, dissepiments consistently formed in a 1:1 ratio to the number of full moons elapsed over each study period. We measured dissepiment spacing to reconstruct multiple years of monthly skeletal extension rates in two Porites colonies from Palmyra Atoll and in another from Palau that bleached in 1998 under anomalously high sea temperatures. Spacing between successive dissepiments exhibited strong seasonality in corals containing annual density bands, with narrow (wide) spacing associated with high (low) density, respectively. A high-density “stress band” accreted during the 1998 bleaching event was associated with anomalously low dissepiment spacing and missed dissepiments, implying that thermal stress disrupts skeletal extension. Further, uranium/calcium ratios increased within stress bands, indicating a reduction in the carbonate ion concentration of the coral’s calcifying fluid under stress. Our study verifies the lunar periodicity of dissepiments, provides a mechanistic basis for the formation of annual density bands in Porites, and reveals the underlying cause of high-density stress bands.

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TL;DR: An almost exclusive dominance of zooxanthellate species is revealed at the studied upper MCE reefs, thus indicating an adequate light regime for photosynthesis there and should encourage similar studies on other reefs.
Abstract: Mesophotic coral-reef ecosystems (MCEs), which comprise the light-dependent communities of corals and other organisms found at depths between 30 and ~ 150 m, have received very little study to date. However, current technological advances, such as remotely operated vehicles and closed-circuit rebreather diving, now enable their thorough investigation. Following the reef-building stony corals, octocorals are the second most common benthic component on many shallow reefs and a major component on deep reefs, the Red Sea included. This study is the first to examine octocoral community features on upper MCEs based on species-level identification and to compare them with the shallower reef zones. The study was carried out at Eilat (Gulf of Aqaba, northern Red Sea), comparing octocoral communities at two mesophotic reefs (30–45 m) and two shallow reef zones (reef flat and upper fore-reef) by belt transects. A total of 30 octocoral species were identified, with higher species richness on the upper MCEs compared to the shallower reefs. Although the MCEs were found to host a higher number of species than the shallower reefs, both featured a similar diversity. Each reef zone revealed a unique octocoral species composition and distinct community structure, with only 16% of the species shared by both the MCEs and the shallower reefs. This study has revealed an almost exclusive dominance of zooxanthellate species at the studied upper MCE reefs, thus indicating an adequate light regime for photosynthesis there. The findings should encourage similar studies on other reefs, aimed at understanding the spatiotemporal features and ecological role of octocorals in reef ecosystems down to the deepest limit of the MCEs.

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TL;DR: A more nuanced approach to science will serve managers needs well and help minimise the rise of mediocrity in coral reef ecosystems as discussed by the authors, where strong ecological interactions are discovered, even small changes in abundance can have profound impacts on coral resilience.
Abstract: Climate change will homogenise the environment and generate a preponderance of mediocre reefs. Managing seascapes of mediocrity will be challenging because our science is ill prepared to deal with the ‘shades of grey’ of reef health; we tend to study natural processes in the healthiest reefs available. Yet much can be gained by examining the drivers and implications of even subtle changes in reef state. Where strong ecological interactions are discovered, even small changes in abundance can have profound impacts on coral resilience. Indeed, if we are to develop effective early warnings of critical losses of resilience, then monitoring must place greater emphasis on measuring and interpreting changes in reef recovery rates. In terms of mechanism, a more nuanced approach is needed to explore the generality of what might be considered ‘dogma’. A more nuanced approach to science will serve managers needs well and help minimise the rise of mediocrity in coral reef ecosystems.

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TL;DR: In this paper, the authors quantified the biomass of five benthic foraminifera (LBFs) species in their algal flat habitat in the austral winter (July 2013), spring (October 2013), and summer (February 2014) at One Tree Reef.
Abstract: Large benthic foraminifera (LBFs) are a vital component of coral reef carbonate production, often overlooked due to their small size. These super-abundant calcifiers are crucial to reef calcification by generation of lagoon and beach sands. Reef-scale carbonate production by LBFs is not well understood, and seasonal fluctuations in this important process are largely unquantified. The biomass of five LBF species in their algal flat habitat was quantified in the austral winter (July 2013), spring (October 2013), and summer (February 2014) at One Tree Reef. WorldView-2 satellite images were used to characterize and create LBF habitat maps based on ground-referenced photographs of algal cover. Habitat maps and LBF biomass measurements were combined to estimate carbonate storage across the entire reef flat. Total carbonate storage of LBFs on the reef flat ranged from 270 tonnes (winter) to 380 tonnes (summer). Satellite images indicate that the habitat area used by LBFs ranged from 0.6 (winter) to 0.71 km2 (spring) of a total possible area of 0.96 km2. LBF biomass was highest in the winter when algal habitat area was lowest, but total carbonate storage was the highest in the summer, when algal habitat area was intermediate. Our data suggest that biomass measurements alone do not capture total abundance of LBF populations (carbonate storage), as the area of available habitat is variable. These results suggest LBF carbonate production studies that measure biomass in discrete locations and single time points fail to capture accurate reef-scale production by not incorporating estimates of the associated algal habitat. Reef-scale measurements in this study can be incorporated into carbonate production models to determine the role of LBFs in sedimentary landforms (lagoons, beaches, etc.). Based on previous models of entire reef metabolism, our estimates indicate that LBFs contribute approximately 3.9–5.4% of reef carbonate budgets, a previously underappreciated carbon sink.

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TL;DR: The ontogenetic window of symbiont uptake in a restoration target species is examined and it is found that early exposure to Symbiodinium may benefit laboratory-reared recruits (e.g., enhance growth), but the potential for establishing long-term novel symbiotic associations may be limited.
Abstract: In most coral species, the critical association with a subset of genetically diverse algal endosymbionts, Symbiodinium, is re-established anew each generation in early coral ontogeny. Yet little is known about the window during which these associations are established or the potential for altering symbiont associations through early exposure to non-native, and/or ecologically beneficial (e.g., stress tolerant), symbiont strains. This study examined the ontogenetic window of symbiont uptake in a restoration target species. Orbicella faveolata recruits, maintained aposymbiotic in laboratory tanks for 4 months, showed a significant decrease in symbiont acquisition upon exposure to natural seawater. Recruits initially inoculated with cultured Symbiodinium readily acquired additional strains from environmental symbiont populations upon exposure, but exogenous uptake also decreased in frequency after 4 months of laboratory rearing. Early exposure to Symbiodinium may benefit laboratory-reared recruits (e.g., enhance growth), but the potential for establishing long-term novel symbiotic associations may be limited.

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TL;DR: With at least six reproductive events per year, a short planktonic larval phase, high settlement rates, and a positive response to nutrient enrichment, the broadcast-spawning species D. labyrinthiformis displays a range of reproductive and early life-history traits that are more often associated with brooding coral species, illustrating that classical divisions of coral species by reproductive mode alone do not always reflect the true biology and ecology of their earliest life stages.
Abstract: Despite the fact that most of the severe demographic bottlenecks in coral populations occur during their earliest life stages, information on the reproductive biology and early life history traits of many coral species is limited and often inferred from adult traits only. This study reports on several atypical aspects of the reproductive biology and early life ecology of the grooved brain coral, Diploria labyrinthiformis (Linnaeus, 1758), a conspicuous reef-building species on Caribbean reefs. The timing of gamete release of D. labyrinthiformis was monitored in Curacao over eight consecutive months, and embryogenesis, planulae behavior, and settlement rates were observed and quantified. We further studied growth and symbiont acquisition in juvenile D. labyrinthiformis for 3.5 yr and compared settler survival under ambient and nutrient-enriched conditions in situ. Notably, D. labyrinthiformis reproduced during daylight hours in six consecutive monthly spawning events between May and September 2013, with a peak in June. This is the largest number of reproductive events per year ever observed in a broadcast-spawning Caribbean coral species. In settlement experiments, D. labyrinthiformis planulae swam to the bottom of culture containers 13 h after spawning and rapidly settled when provided with settlement cues (42% within 14 h). After 5 months, the survival and growth rates of settled juveniles were 3.7 and 1.9 times higher, respectively, for settlers that acquired zooxanthellae within 1 month after settlement, compared to those that acquired symbionts later on. Nutrient enrichment increased settler survival fourfold, but only for settlers that had acquired symbionts within 1 month after settlement. With at least six reproductive events per year, a short planktonic larval phase, high settlement rates, and a positive response to nutrient enrichment, the broadcast-spawning species D. labyrinthiformis displays a range of reproductive and early life-history traits that are more often associated with brooding coral species, illustrating that classical divisions of coral species by reproductive mode alone do not always reflect the true biology and ecology of their earliest life stages.

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TL;DR: The importance of crevices as spatial refugia from predators for juvenile corals and the importance of structural complexity for juvenile coral growth and survival that enhances reef recovery are highlighted.
Abstract: Coral recruitment and juvenile growth are essential processes for coral population maintenance and recovery. A growing body of research has evaluated the influence of reef microstructure on coral settlement and post-settlement survival, showing that physical refugia enhance recruitment. These studies have evaluated coral recruit morality from competition with macroalgae and indirect predation by grazing organisms, but the impact of direct predation by corallivorous piscine species on juvenile corals and how this interacts with reef microstructure is relatively unknown. This study examined whether refugia provided by micro-crevices enhance juvenile coral survival from corallivory. Juvenile corals from two different functional groups, the slow-growing massive Porites lobata and fast-growing branching Pocillopora damicornis, with average nubbin sizes of 1.4 cm × 0.3 cm and 0.5 cm × 1.0 cm (diameter × height), respectively, were attached to experimental tiles using small (1.44 cm3) and large (8.0 cm3) crevice sizes and were monitored for 29 d on a forereef in Palau. Full crevices (four sided) enhanced coral survival compared to exposed microhabitats in both coral taxa, but crevice size did not alter survival rates. Corallivores targeted recruits within crevices regardless of crevice size; dominant predators included small triggerfish (Balistidae), butterflyfish (Chaetodon), and wrasse (Cheilinus). Overall, Pocillopora suffered much higher rates of mortality than Porites. All Pocillopora were consumed by day 8 of the experiment, but mortality was significantly delayed in full crevices compared to exposed and partial crevice (three sided) microhabitats. In contrast, Por. lobata located in all microhabitats survived the entire experiment up to 29 d, with high survival in full (>90%) and partial crevices (70%), but only 28% survival in exposed microhabitats. These findings show the importance of crevices as spatial refugia from predators for juvenile corals and highlight the importance of structural complexity for juvenile coral growth and survival that enhances reef recovery.