Bleaching Response Research Articles

Metabolomic shifts associated with heat stress in coral holobionts.

Understanding the response of the coral holobiont to environmental change is crucial to inform conservation efforts. The most pressing problem is "coral bleaching," usually precipitated by prolonged thermal stress. We used untargeted, polar metabolite profiling to investigate the physiological response of the coral species Montipora capitata and Pocillopora acuta to heat stress. Our goal was to identify diagnostic markers present early in the bleaching response. From the untargeted UHPLC-MS data, a variety of co-regulated dipeptides were found that have the highest differential accumulation in both species. The structures of four dipeptides were determined and showed differential accumulation in symbiotic and aposymbiotic (alga-free) populations of the sea anemone Aiptasia (Exaiptasia pallida), suggesting the deep evolutionary origins of these dipeptides and their involvement in symbiosis. These and other metabolites may be used as diagnostic markers for thermal stress in wild coral.

Quartz bleaching by phosphoric acid: An investigation on the relationship between Fe rejection rate and colour response

Advances in high-tech industries have led to increasing the demand for highly pure quartz. Research on the beneficiation of quartz has been conducted to meet market requirements by different processes including hydrometallurgical ones. Even though phosphoric acid (PA) has a capacity to complex iron, and therefore has the potential to bleach quartz, this reagent has not attracted sufficient attention. This study was conducted to investigate the relationship between Fe remaining in concentrate by PA bleaching and colour responses of quartz ore sample employing Box-Behnken Design, a common designs of Response Surface Methodology. Effects of bleaching parameters (PA concentration 0–2 M, temperature 65–95 °C, bleaching time 45–180 min.) were investigated in agitated pulp condition at 400 rpm. The efficiency of the process was determined by colorimetric analysis and chemical analysis (ICP-MS). Experimental results revealed that the linear effects of bleaching parameters on Fe removal by PA were statistically significant. The lowest Fe remaining in concentrate, 79 ppm, was observed in two different conditions tested at the highest initial PA concentration (2 M) at 95 °C, over 90 min, and at 85 °C, over 180 min of bleaching. The increase in acidity resulted in the higher Fe rejection rate and the lower yellowness. Significant improvements in whiteness and yellowness were obtained by PA bleaching, whereas redness could not be improved considerably. This finding has been attributed to the presence of free dark coloured particles, and the resistance of redness source minerals like hematite to react with PA ions.

Host genotype and stable differences in algal symbiont communities explain patterns of thermal stress response of Montipora capitata following thermal pre-exposure and across multiple bleaching events

As sea surface temperatures increase worldwide due to climate change, coral bleaching events are becoming more frequent and severe, resulting in reef degradation. Leveraging the inherent ability of reef-building corals to acclimatize to thermal stress via pre-exposure to protective temperature treatments may become an important tool in improving the resilience of coral reefs to rapid environmental change. We investigated whether historical bleaching phenotype, coral host genotype, and exposure to protective temperature treatments would affect the response of the Hawaiian coral Montipora capitata to natural thermal stress. Fragments were collected from colonies that demonstrated different bleaching responses during the 2014–2015 event in Kāne‘ohe Bay (O‘ahu, Hawai‘i) and exposed to four different artificial temperature pre-treatments (and a control at ambient temperature). After recovery, fragments experienced a natural thermal stress event either in laboratory conditions or their native reef environment. Response to thermal stress was quantified by measuring changes in the algal symbionts’ photochemical efficiency, community composition, and relative density. Historical bleaching phenotype was reflected in stable differences in symbiont community composition, with historically bleached corals containing only Cladocopium symbionts and historically non-bleached corals having mixed symbiont communities dominated by Durusdinium. Mixed-community corals lost more Cladocopium than Cladocopium-only corals during the natural thermal stress event and preferentially recovered with Durusdinium. Laboratory pre-treatments exposed corals to more thermal stress than anticipated, causing photochemical damage that varied significantly by genotype. While none of the treatments had a protective effect, temperature variation during treatments had a significant detrimental effect on photochemical efficiency during the thermal stress event. These results show that acclimatization potential is affected by fine-scale differences in temperature regime, host genotype, and relatively stable differences in symbiont community composition that underpin historical bleaching phenotypes in M. capitata.

Coral husbandry for ocean futures: leveraging abiotic factors to increase survivorship, growth, and resilience in juvenile Montipora capitata

Reef restoration via direct outplanting of sexually propagated juvenile corals is a key strategy in preserving coral reef ecosystem function in the face of global and local stressors (e.g. ocean warming). To advance our capacity to scale and maximize the efficiency of restoration initiatives, we examined how abiotic conditions (i.e. larval rearing temperature, substrate condition, light intensity, and flow rate) interact to enhance post-settlement survival and growth of sexually propagated juvenileMontipora capitata. Larvae were reared at 3 temperatures (high: 28.9°C, ambient: 27.2°C, low: 24.5°C) for 72 h during larval development, and were subsequently settled on aragonite plugs conditioned in seawater (1 or 10 wk) and raised in different light and flow regimes. These juvenile corals underwent a natural bleaching event in Kāne‘ohe Bay, O‘ahu, Hawai‘i (USA), in summer 2019, allowing us to opportunistically measure bleaching response in addition to survivorship and growth. This study demonstrates how leveraging light and flow can increase the survivorship and growth of juvenileM. capitata. In contrast, larval preconditioning and substrate conditioning had little overall effect on survivorship, growth, or bleaching response. Importantly, there was no optimal combination of abiotic conditions that maximized survival and growth in addition to bleaching tolerances. This study highlights the ability to tailor sexual reproduction for specific restoration goals by addressing knowledge gaps and incorporating practices that could improve resilience in propagated stocks.

The projected degradation of subtropical coral assemblages by recurrent thermal stress.

Subtropical coral assemblages are threatened by similar extreme thermal stress events to their tropical counterparts. Yet, the mid- and long-term thermal stress responses of corals in subtropical environments remain largely unquantified, limiting our capacity to predict their future viability. The annual survival, growth and recruitment of 311 individual corals within the Solitary Islands Marine Park (Australia) was recorded over a 3-year period (2016-2018), including the 2015/2016 thermal stress event. These data were used to parameterise integral projection models quantifying the effect of thermal stress within a subtropical coral assemblage. Stochastic simulations were also applied to evaluate the implications of recurrent thermal stress scenarios predicted by four different Representative Concentration Pathways. We report differential shifts in population growth rates (λ) among coral populations during both stress and non-stress periods, confirming contrasting bleaching responses among taxa. However, even during non-stress periods, the observed dynamics for all taxa were unable to maintain current community composition, highlighting the need for external recruitment sources to support the community structure. Across all coral taxa, projected stochastic growth rates (λs ) were found to be lowest under higher emissions scenarios. Correspondingly, predicted increases in recurrent thermal stress regimes may accelerate the loss of coral coverage, species diversity and structural complexity within subtropical regions. We suggest that these trends are primarily due to the susceptibility of subtropical specialists and endemic species, such as Pocillopora aliciae, to thermal stress. Similarly, the viability of many tropical coral populations at higher latitudes is highly dependent on the persistence of up-current tropical systems. As such, the inherent dynamics of subtropical coral populations appear unable to support their future persistence under unprecedented thermal disturbance scenarios.

Low coral bleaching prevalence at the Bolinao-Anda Reef Complex, northwestern Philippines during the 2016 thermal stress event

Here, we examined the coral bleaching responses during the 2016 thermal stress event and post-bleaching changes in coral communities in the heavily disturbed reefs of the Bolinao-Anda Reef Complex (BARC), northwestern Philippines. Less than 25% of colonies bleached, with 77% attributed to five genera (Dipsastrea, Porites, Fungia, Seriatopora, and Montipora). Coral bleaching prevalence was associated with site location, coral composition, and coral abundance, suggesting that small-scale variation (<20 km) in coral communities (taxa and density) influences spatial variation in coral bleaching prevalence. There was no noticeable change in coral composition and cover two years after the bleaching event as exposure to chronic disturbance likely selected for the dominance of stress tolerant coral taxa and communities. Results show that the 2016 thermal stress event caused coral bleaching but with low prevalence at the BARC, which suggests that disturbed reefs may provide spatial refuge to coral communities from thermal stress.

Experimental investigation of tidal and freshwater influence on Symbiodiniaceae abundance in Anthopleura elegantissima.

The San Francisco Bay outflow creates a tidally influenced low-salinity plume that affects adjacent coastal sites. In the study region, Anthopleura elegantissima (Cnidaria; Anthozoa) hosts a single symbiont, the dinoflagellate Breviolum muscatinei. Salinity, temperature, and aerial stress induce a bleaching response similar to corals where symbionts are expelled, causing further energetic stress. Using field observations of environmental conditions and symbiont abundance at sites on a gradient of exposure to estuarine outflow, along with a fully crossed multifactorial lab experiment, we tested for changes in symbiont abundance in response to various combinations of three stressors. Lab experiments were designed to mimic short term outflow events with low salinity, high temperature, and aerial exposure treatments. The lab aerial exposure treatment was a statistically significant factor in suppressing symbiont repopulation (ANOVA, p = .017). In the field, symbiont density decreased with increasing tidal height at the site closest to freshwater outflow (ANOVA, p = .007), suggesting that aerial exposure may affect symbiont density more than sea surface temperature and salinity. Unanticipated documentation of survival in 9 months of sand burial and subsequent repopulation of symbionts is reported as a six-month extension to past observations, exemplifying strong tolerance to environmental insult in this Cnidarian mutualism. The study of this symbiosis is useful in examining predicted changes in ocean conditions in tidepool communities and considering relative sources of stress.

Highly variable taxa-specific coral bleaching responses to thermal stresses

Complex histories of chronic and acute sea surface temperature (SST) stresses are expected to trigger taxon- and location-specific responses that will ultimately lead to novel coral communities. The 2016 El Niño-Southern Oscillation provided an opportunity to examine large-scale and recent environmental histories on emerging patterns in 226 coral communities distributed across 12 countries from East Africa to Fiji. Six main coral communities were identified that largely varied across a gradient of Acropora to massive Porites dominance. Bleaching intensity was taxon-specific and was associated with complex interactions among the 20 environmental variables that we examined. Coral community structure was better aligned with the historical temperature patterns between 1985 and 2015 than the 2016 extreme temperature event. Additionally, bleaching responses observed during 2016 differed from historical reports during past warm years. Consequently, coral communities present in 2016 are likely to have been reorganized by both long-term community change and acclimation mechanisms. For example, less disturbed sites with cooler baseline temperatures, higher mean historical SST background variability, and infrequent extreme warm temperature stresses were associated with Acropora-dominated communities, while more disturbed sites with lower historical SST background variability and frequent acute warm stress were dominated by stress-resistant massive Porites corals. Overall, the combination of taxon-specific responses, community-level reorganization over time, geographic variation, and multiple environmental stressors suggest complex responses and a diversity of future coral communities that can help contextualize management priorities and activities.

All-in-one, reversible yellow-green color switching tungsten oxide electrochromic devices

Tungsten oxide (WO3) sol doped with carbon nanodots (CDs) was utilized for constructing an electrolyte-free, all-in-one electrochromic device (ECD). Different from the traditional WO3-based ECDs, showing the color transition between colorless and blue, the demonstrated device could realize reversible color changes between yellow and green due to the intrinsic color of CDs. The optical contrast of the optimized ECD was up to 53.6% at 650 nm and the coloration response time was 27 s. It could also be automatically bleached under 0 V with the bleaching response time of 80 s. In addition, attributing to the unique photoluminescence (PL) characteristics of CDs, the ECD could also present tunable PL under various excitation wavelengths, which potentially broadens the applications of ECDs in multiple anti-counterfeiting. These results provide simple and effective guidelines for constructing high-performance ECDs with simple architecture and extended color palette.

Unravelling the different causes of nitrate and ammonium effects on coral bleaching

Mass coral bleaching represents one of the greatest threats to coral reefs and has mainly been attributed to seawater warming. However, reduced water quality can also interact with warming to increase coral bleaching, but this interaction depends on nutrient ratios and forms. In particular, nitrate (NO3−) enrichment reduces thermal tolerance while ammonium (NH4+) enrichment tends to benefit coral health. The biochemical mechanisms underpinning the different bleaching responses of corals exposed to DIN enrichment still need to be investigated. Here, we demonstrated that the coral Stylophora pistillata underwent a severe oxidative stress condition and reduced aerobic scope when exposed to NO3− enrichment combined with thermal stress. Such condition resulted in increased bleaching intensity compared to a low-nitrogen condition. On the contrary, NH4+ enrichment was able to amend the deleterious effects of thermal stress by favoring the oxidative status and energy metabolism of the coral holobiont. Overall, our results demonstrate that the opposite effects of nitrate and ammonium enrichment on coral bleaching are related to the effects on corals’ energy/redox status. As nitrate loading in coastal waters is predicted to significantly increase in the future due to agriculture and land-based pollution, there is the need for urgent management actions to prevent increases in nitrate levels in seawater. In addition, the maintenance of important fish stocks, which provide corals with recycled nitrogen such as ammonium, should be favoured.

Genome‐wide SNP analysis reveals an increase in adaptive genetic variation through selective breeding of coral

Marine heat waves are increasing in magnitude, duration, and frequency as a result of climate change and are the principal global driver of mortality in reef-building corals. Resilience-based genetic management may increase coral heat tolerance, but it is unclear how temperature responses are regulated at the genome level and thus how corals may adapt to warming naturally or through selective breeding. Here we combine phenotypic, pedigree, and genomic marker data from colonies sourced from a warm reef on the Great Barrier Reef reproductively crossed with conspecific colonies from a cooler reef to produce combinations of warm purebreds and warm-cool hybrid larvae and juveniles. Interpopulation breeding created significantly greater genetic diversity across the coral genome compared to breeding between populations and maintained diversity in key regions associated with heat tolerance and fitness. High-density genome-wide scans of single nucleotide polymorphisms (SNPs) identified alleles significantly associated with larval families reared at 27.5°C (87-2,224 loci), including loci putatively associated with proteins involved in responses to heat stress (cell membrane formation, metabolism, and immune responses). Underlying genetics of these families explained 43% of PCoA multilocus variation in survival, growth, and bleaching responses at 27.5°C and 31°C at the juvenile stage. Genetic marker contribution to total variation in fitness traits (narrow-sense heritability) was high for survival but not for growth and bleaching in juveniles, with heritability of these traits being higher at 31°C relative to 27.5°C. While based on only a limited number of crosses, the mechanistic understanding presented here demonstrates that allele frequencies are affected by one generation of selective breeding, key information for the assessments of genetic intervention feasibility and modelling of reef futures.

Uncovering the role of Symbiodiniaceae assemblage composition and abundance in coral bleaching response by minimizing sampling and evolutionary biases

BackgroundBiodiversity and productivity of coral-reef ecosystems depend upon reef-building corals and their associations with endosymbiotic Symbiodiniaceae, which offer diverse functional capabilities to their hosts. The number of unique symbiotic partners (richness) and relative abundances (evenness) have been hypothesized to affect host response to climate change induced thermal stress. Symbiodiniaceae assemblages with many unique phylotypes may provide greater physiological flexibility or form less stable symbioses; assemblages with low abundance phylotypes may allow corals to retain thermotolerant symbionts or represent associations with less-suitable symbionts.ResultsHere we demonstrate that true richness of Symbiodiniaceae phylotype assemblages is generally not discoverable from direct enumeration of unique phylotypes in association records and that cross host-species comparisons are biased by sampling and evolutionary patterns among species. These biases can be minimized through rarefaction of richness (rarefied-richness) and evenness (Probability of Interspecific Encounter, PIE), and analyses that account for phylogenetic patterns. These standardized metrics were calculated for individual Symbiodiniaceae assemblages composed of 377 unique ITS2 phylotypes associated with 123 coral species. Rarefied-richness minimized correlations with sampling effort, while maintaining important underlying characteristics across host bathymetry and geography. Phylogenetic comparative methods reveal significant increases in coral bleaching and mortality associated with increasing Symbiodiniaceae assemblage richness and evenness at the level of host species.ConclusionsThese results indicate that the potential flexibility afforded by assemblages characterized by many phylotypes present at similar relative abundances does not result in decreased bleaching risk and point to the need to characterize the overall functional and genetic diversity of Symbiodiniaceae assemblages to quantify their effect on host fitness under climate change.

Electrochromic smart windows using 2D-MoS2 nanostructures protected silver nanowire based flexible transparent electrodes

In recent years, the harvesting and management of energy has become a topic of intense interest because of the growing energy demands of society, the depletion of fossil fuel sources, and the research on eco-friendly development of efficient energy management technologies. In this connection, we proposed to utilize the energy-saving properties of electrochromic technology-based MoS2/silver nanowire (AgNW) electrodes for flexible smart windows. MoS2 is used to overcome the difficulties of stability, corrosion, and oxidation of AgNWs in the electrolyte. Transition metal chalcogenide (2D-MoS2) structures provide a unique platform to stabilize the surface of the AgNWs at the electrolyte interface. The prepared WO3/MoS2/AgNW based electrochromic device attained an optical contract of 52% at the low voltage of −1.2 V, which yielded a coloration efficiency (η) of 31.92 cm2/C. The MoS2/AgNW electrode exhibited high coloration and bleaching response, Tc = 11.5 s and Tb = 4.5 s, respectively, which is a faster response than that of the AgNW based electrochromic devices. Most of the existing electrochromic devices have advantages for next-generation smart windows and flexible and wearable optoelectronic applications.

Gene regulation underpinning increased thermal tolerance in a laboratory-evolved coral photosymbiont.

Small increases in ocean temperature can disrupt the obligate symbiosis between corals and dinoflagellate microalgae, resulting in coral bleaching. Little is known about the genes that drive the physiological and bleaching response of algal symbionts to elevated temperature. Moreover, many studies to-date have compared highly divergent strains, making it challenging to accredit specific genes to contrasting traits. Here, we compare transcriptional responses at ambient (27°C) and bleaching-relevant (31°C) temperatures in a monoclonal, wild-type (WT) strain of Symbiodiniaceae to those of a selected-strain (SS), derived from the same monoclonal culture and experimentally evolved to elevated temperature over 80 generations (2.5years). Thousands of genes were differentially expressed at a log fold-change of >8 between the WT and SS over a 35days temperature treatment period. At 31°C, WT cells exhibited a temporally unstable transcriptomic response upregulating genes involved in the universal stress response such as molecular chaperoning, protein repair, protein degradation and DNA repair. Comparatively, SS cells exhibited a temporally stable transcriptomic response and downregulated many stress response genes that were upregulated by the WT. Among the most highly upregulated genes in the SS at 31°C were algal transcription factors and a gene probably of bacterial origin that encodes a type II secretion system protein, suggesting interactions with bacteria may contribute to the increased thermal tolerance of the SS. Genes and functional pathways conferring thermal tolerance in the SS could be targeted in future genetic engineering experiments designed to develop thermally resilient algal symbionts for use in coral restoration and conservation.

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.

Contrasting patterns of mortality in Polynesian coral reefs following the third global coral bleaching event in 2016

In 2016, many tropical corals worldwide were exposed to anomalously high temperatures due to one of the strongest El Nino events ever recorded. Bleaching impacts were reported on 23 islands within three archipelagos of French Polynesia (Tuamotu, Society and Marquesas archipelagos). A detailed study on the effects of elevated temperatures on corals was performed on five islands (Mo’orea, Makemo, Hikueru, Marutea and Katiu) and revealed contrasting patterns of coral bleaching responses between Mo’orea (Society Archipelago) and the four islands of the Tuamotu Archipelago. While some reefs from the Tuamotu lost more than half of their coral cover, in Mo’orea, less than 1% mortality was recorded 6 months after bleaching. During the 2016 bleaching event, certain reefs at 12 m depth in the outer reef habitats were not exposed to sufficiently long high-temperature periods (heat stress not exceeding 1.1 °C weeks in Mo’orea) to cause large-scale bleaching-related coral mortality. In contrast, other reefs in the Tuamotu Archipelago had DHW reaching up to 9.2 °C weeks and experienced severe mortality (up to 71%). Our study showed how differential heat stress exposure across reefs of French Polynesia led to different impacts on corals. Until now, Mo’orea reefs have been spared abnormally high temperatures leading to mortality and should be considered an important source of larvae to help maintain reefs on the surrounding islands.

Successive marine heatwaves cause disproportionate coral bleaching during a fast phase transition from El Niño to La Niña.

The frequency and intensity of marine heatwaves that result in coral bleaching events have increased over recent decades and led to catastrophic losses of reef-building corals in many regions. The high-latitude coral assemblages at Lord Howe Island, which is a UNESCO listed site is the world southernmost coral community, were exposed to successive thermal anomalies following a fast phase-transition of the record-breaking 2009 to 2010 warm pool El Niño in the Central Pacific to a strong La Niña event in late 2010. The coral community experienced severe and unprecedented consecutive bleaching in both 2010 and 2011. Coral health surveys completed between March 2010 and September 2012 quantified the response and recovery of approximately 43,700 coral colonies to these successive marine heatwaves. In March 2010, coral bleaching ranged from severe, with 99% of colonies bleached at some shallow lagoon sites, to mild at deeper reef slope sites, with only 17% of individuals affected. Significant immediate mortality from thermal stress was evident during the peak of the bleaching event. Overall, species in the genera Pocillopora, Stylophora, Seriatopora and Porites were the most affected, while minimal bleaching and mortality was recorded among members of other coral families (e.g. Acroporidae, Dendrophyllidae & Merulinidae). Surviving corals underwent a subsequent, but much less intense, thermal anomaly in 2011 that led to a disproportionate bleaching response among susceptible taxa. While this observation indicates that the capacity of thermally susceptible high-latitude corals to acclimatize to future ocean warming may be limited, particularly if bleaching events occur annually, our long-term survey data shows that coral cover at most sites recovered to pre-bleaching levels within three years in the absence of further thermal anomalies.

Diverse symbiont bleaching responses are evident from 2-degree heating week bleaching conditions as thermal stress intensifies in coral

Coral bleaching is the dysfunction of the coral–algal endosymbiosis and is characterised as a loss of Symbiodiniaceae cells from host tissues or the loss of photosynthetic pigments. This breakdown of symbiosis occurs as a result of elevated temperature beyond the organism’s thermal threshold. The thermal tipping points within the symbiosis have not yet been well resolved, and the mechanisms underlying the various cellular processes of the corals bleaching response remain unknown. This study characterised the cellular responses of the symbiont Cladocopium sp. (syn. clade C3) within the host coral Acropora aspera during exposure to thermal stress. Exposure to temperatures between 2 and 3°C below the bleaching threshold, equating to 2-degree heating weeks (DHWs), results in changes to the symbiont cell morphology and cell division rates. Once corals were exposed to 4 DHWs, over 90% of the symbiont cells showed signs of degradation. Although sub-bleaching thermal stress is not sufficient to trigger bleaching alerts at an ecological scale, this stressor substantially affects the coral symbiosis. It is therefore vital that we begin to quantify how sub-bleaching thermal stress affects the fitness of Symbiodiniacea populations, their coral hosts and subsequently reefs worldwide.

Refugia under threat: Mass bleaching of coral assemblages in high-latitude eastern Australia.

Environmental anomalies that trigger adverse physiological responses and mortality are occurring with increasing frequency due to climate change. At species' range peripheries, environmental anomalies are particularly concerning because species often exist at their environmental tolerance limits and may not be able to migrate to escape unfavourable conditions. Here, we investigated the bleaching response and mortality of 14 coral genera across high-latitude eastern Australia during a global heat stress event in 2016. We evaluated whether the severity of assemblage-scale and genus-level bleaching responses was associated with cumulative heat stress and/or local environmental history, including long-term mean temperatures during the hottest month of each year (SSTLTMAX ), and annual fluctuations in water temperature (SSTVAR ) and solar irradiance (PARZVAR ). The most severely-bleached genera included species that were either endemic to the region (Pocillopora aliciae) or rare in the tropics (e.g. Porites heronensis). Pocillopora spp., in particular, showed high rates of immediate mortality. Bleaching severity of Pocillopora was high where SSTLTMAX was low or PARZVAR was high, whereas bleaching severity of Porites was directly associated with cumulative heat stress. While many tropical Acropora species are extremely vulnerable to bleaching, the Acropora species common at high latitudes, such as A. glauca and A. solitaryensis, showed little incidence of bleaching and immediate mortality. Two other regionally-abundant genera, Goniastrea and Turbinaria, were also largely unaffected by the thermal anomaly. The severity of assemblage-scale bleaching responses was poorly explained by the environmental parameters we examined. Instead, the severity of assemblage-scale bleaching was associated with local differences in species abundance and taxon-specific bleaching responses. The marked taxonomic disparity in bleaching severity, coupled with high mortality of high-latitude endemics, point to climate-driven simplification of assemblage structures and progressive homogenisation of reef functions at these high-latitude locations.

What drives phenotypic divergence among coral clonemates of Acropora palmata?

Evolutionary rescue of populations depends on their ability to produce phenotypic variation that is heritable and adaptive. DNA mutations are the best understood mechanisms to create phenotypic variation, but other, less well‐studied mechanisms exist. Marine benthic foundation species provide opportunities to study these mechanisms because many are dominated by isogenic stands produced through asexual reproduction. For example, Caribbean acroporid corals are long lived and reproduce asexually via breakage of branches. Fragmentation is often the dominant mode of local population maintenance. Thus, large genets with many ramets (colonies) are common. Here, we observed phenotypic variation in stress responses within genets following the coral bleaching events in 2014 and 2015 caused by high water temperatures. This was not due to genetic variation in their symbiotic dinoflagellates (Symbiodinium “fitti”) because each genet of this coral species typically harbours a single strain of S. “fitti”. Characterization of the microbiome via 16S tag sequencing correlated the abundance of only two microbiome members (Tepidiphilus, Endozoicomonas) with a bleaching response. Epigenetic changes were significantly correlated with the host's genetic background, the location of the sampled polyps within the colonies (e.g., branch vs. base of colony), and differences in the colonies’ condition during the bleaching event. We conclude that long‐term microenvironmental differences led to changes in the way the ramets methylated their genomes, contributing to the differential bleaching response. However, most of the variation in differential bleaching response among clonemates of Acropora palmata remains unexplained. This research provides novel data and hypotheses to help understand intragenet variability in stress phenotypes of sessile marine species.