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There are major concerns about the ecological impact of extreme weather events. In the oceans, marine heatwaves (MHWs) are an increasing threat causing, for example, recent devastation to coral reefs around the world. We show that these impacts extend to adjacent terrestrial systems and could negatively affect the breeding of endangered species. We demonstrate that during an MHW that resulted in major coral bleaching and mortality in a large, remote marine protected area, anomalously warm temperatures also occurred on sea turtle nesting beaches. Granger causality testing showed that variations in sea surface temperature strongly influenced sand temperatures on beaches. We estimate that the warm conditions on both coral reefs and sandy beaches during the MHW were unprecedented in the last 70 years. Model predictions suggest that the most extreme female-biased hatchling sex ratio and the lowest hatchling survival in nests in the last 70 years both occurred during the heatwave. Our work shows that predicted increases in the frequency and intensity of MHWs will likely have growing impacts on sea turtle nesting beaches as well as other terrestrial coastal environments.

The global decline of reef corals has been driven largely by several marine heatwaves. This has greatly reduced coral cover but has reduced coral diversity also. While there is a lack of data in most locations to detect coral species losses, reefs of the Chagos Archipelago, central Indian Ocean, have long term monitoring data extending back to the late 1970s. Severe declines in cover have occurred since the 1970s, with regional extinctions of some species and key species assemblages. There is a severe decline in coral settlement, along with a substantial loss of habitat quality which has reduced the habitat available for settlement. This is a clear precursor to positive feedback. Regional species extinctions here occur mainly when total coral cover is <10% of pre-warming levels. Climate models predict more frequent and more severe marine heatwaves, and even if this ecosystem recovers it will contain fewer species., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Crown Copyright © 2020. Published by Elsevier Ltd. All rights reserved.)

As in the tropical Atlantic, Acropora populations in the southern Persian/Arabian Gulf plummeted within two decades after having been ecosystem engineers on most wave-exposed reefs since the Pleistocene. Since 1996/1998 live coral cover in the Gulf declined by over 90% in many areas, primarily due to bleaching and diseases caused by rising temperatures. In the formerly dominant table-coral species A. downingi, population dynamics corresponding to disturbance regimes was quantified in three transition matrices (lower disturbance pre-1996; moderate disturbance from 1998 to 2010 and 2013 to 2017, disturbed in 1996/1998, 2010/11/12, 2017). Increased disturbance frequency and severity caused progressive reduction in coral size, cover, and population fecundity. Small size-classes were bolstered more by partial colony mortality than sexual recruitment. Some large corals had a size refuge and resisted die-back but were also lost with increasing disturbance. Matrix and biophysical larval flow models suggested one metapopulation. Southern, Arabian, populations could be connected to northern, Iranian, populations but this connectivity was lost under assumptions of pelagic larval duration at rising temperatures shortened to a third. Then, the metapopulation disintegrated into isolated populations. Connectivity required to avoid extinctions increased exponentially with disturbance frequency and correlation of disturbances across the metapopulation. Populations became unsustainable at eight disturbances in 15 years, when even highest theoretical recruitment no longer compensated mortality. This lethal disturbance frequency was 3-fold that of the moderately disturbed monitoring period and 4-fold of the preceding low-disturbance period-suggesting ongoing shortening of the disturbance-free period. Observed population collapse and environmental changes in the Gulf suggest that A. downingi is heading toward at least functional extinction mainly due to increasingly frequent temperature-induced mortality events, clearly linked to climate change., (© 2018 John Wiley & Sons Ltd.)

This article discusses two key issues: firstly, the demise of reefs in the Gulf which is happening probably more rapidly than elsewhere; and secondly, the reasons why this remains such an intractable problem. Most reasons for this decline are scientifically well understood, though clearly not by the region's managers. Several factors may cause people to ignore the problem, even though habitat loss is vastly costly to the region. About 70% of the Gulf's reefs have essentially disappeared in a few decades, and although scientific indicators confirm that this is happening, it is commonly discounted as even being a possibility. Management of human interactions with the Gulf's marine systems remains very inadequate, to the detriment of the Gulf's marine systems and its people. It is clear that this not a scientific issue any longer but rather it is a political problem and failure., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Most reviews concerning the impact of climate change on coral reefs discuss independent effects of warming or ocean acidification. However, the interactions between these, and between these and direct local stressors are less well addressed. This review underlines that coral bleaching, acidification, and diseases are expected to interact synergistically, and will negatively influence survival, growth, reproduction, larval development, settlement, and post-settlement development of corals. Interactions with local stress factors such as pollution, sedimentation, and overfishing are further expected to compound effects of climate change. Reduced coral cover and species composition following coral bleaching events affect coral reef fish community structure, with variable outcomes depending on their habitat dependence and trophic specialisation. Ocean acidification itself impacts fish mainly indirectly through disruption of predation- and habitat-associated behavior changes. Zooxanthellate octocorals on reefs are often overlooked but are substantial occupiers of space; these also are highly susceptible to bleaching but because they tend to be more heterotrophic, climate change impacts mainly manifest in terms of changes in species composition and population structure. Non-calcifying macroalgae are expected to respond positively to ocean acidification and promote microbe-induced coral mortality via the release of dissolved compounds, thus intensifying phase-shifts from coral to macroalgal domination. Adaptation of corals to these consequences of CO2 rise through increased tolerance of corals and successful mutualistic associations between corals and zooxanthellae is likely to be insufficient to match the rate and frequency of the projected changes. Impacts are interactive and magnified, and because there is a limited capacity for corals to adapt to climate change, global targets of carbon emission reductions are insufficient for coral reefs, so lower targets should be pursued. Alleviation of most local stress factors such as nutrient discharges, sedimentation, and overfishing is also imperative if sufficient overall resilience of reefs to climate change is to be achieved., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Stylophora pistillata is a widely used coral "lab-rat" species with highly variable morphology and a broad biogeographic range (Red Sea to western central Pacific). Here we show, by analysing Cytochorme Oxidase I sequences, from 241 samples across this range, that this taxon in fact comprises four deeply divergent clades corresponding to the Pacific-Western Australia, Chagos-Madagascar-South Africa, Gulf of Aden-Zanzibar-Madagascar, and Red Sea-Persian/Arabian Gulf-Kenya. On the basis of the fossil record of Stylophora, these four clades diverged from one another 51.5-29.6 Mya, i.e., long before the closure of the Tethyan connection between the tropical Indo-West Pacific and Atlantic in the early Miocene (16-24 Mya) and should be recognised as four distinct species. These findings have implications for comparative ecological and/or physiological studies carried out using Stylophora pistillata as a model species, and highlight the fact that phenotypic plasticity, thought to be common in scleractinian corals, can mask significant genetic variation.

The Chagos Archipelago designated as a no-take marine protected area in 2010, lying about 500 km south of the Maldives in the Indian Ocean, has a high conservation priority, particularly because of its fast recovery from the ocean-wide massive coral mortality following the 1998 coral bleaching event. The aims of this study were to examine Symbiodinium diversity and distribution associated with scleractinian corals in five atolls of the Chagos Archipelago, spread over 10,000 km(2). Symbiodinium clade diversity in 262 samples of seven common coral species, Acropora muricata, Isopora palifera, Pocillopora damicornis, P. verrucosa, P. eydouxi, Seriatopora hystrix, and Stylophora pistillata were determined using PCR-SSCP of the ribosomal internal transcribed spacer 1 (ITS1), PCR-DDGE of ITS2, and phylogenetic analyses. The results indicated that Symbiodinium in clade C were the dominant symbiont group in the seven coral species. Our analysis revealed types of Symbiodinium clade C specific to coral species. Types C1 and C3 (with C3z and C3i variants) were dominant in Acroporidae and C1 and C1c were the dominant types in Pocilloporidae. We also found 2 novel ITS2 types in S. hystrix and 1 novel ITS2 type of Symbiodinium in A. muricata. Some colonies of A. muricata and I. palifera were also associated with Symbiodinium A1. These results suggest that corals in the Chagos Archipelago host different assemblages of Symbiodinium types then their conspecifics from other locations in the Indian Ocean; and that future research will show whether these patterns in Symbiodinium genotypes may be due to local adaptation to specific conditions in the Chagos.

We explore impacts on pristine atolls subjected to anthropogenic near-field (human habitation) and far-field (climate and environmental change) pressure. Using literature data of human impacts on reefs, we parameterize forecast models to evaluate trajectories in coral cover under impact scenarios that primarily act via recruitment and increased mortality of larger corals. From surveys across the Chagos, we investigate the regeneration dynamics of coral populations distant from human habitation after natural disturbances. Using a size-based mathematical model based on a time-series of coral community and population data from 1999-2006, we provide hind- and forecast data for coral population dynamics within lagoons and on ocean-facing reefs verified against monitoring from 1979-2009. Environmental data (currents, temperatures) were used for calibration. The coral community was simplified into growth typologies: branching and encrusting, arboresent and massive corals. Community patterns observed in the field were influenced by bleaching-related mortality, most notably in 1998. Survival had been highest in deep lagoonal settings, which suggests a refuge. Recruitment levels were higher in lagoons than on ocean-facing reefs. When adding stress by direct human pressure, climate and environmental change as increased disturbance frequency and modified recruitment and mortality levels (due to eutrophication, overfishing, pollution, heat, acidification, etc), models suggest steep declines in coral populations and loss of community diversification among habitats. We found it likely that degradation of lagoonal coral populations would impact regeneration potential of all coral populations, also on ocean-facing reefs, thus decreasing reef resilience on the entire atoll.

Modern hard corals (Class Hexacorallia; Order Scleractinia) are widely studied because of their fundamental role in reef building and their superb fossil record extending back to the Triassic. Nevertheless, interpretations of their evolutionary relationships have been in flux for over a decade. Recent analyses undermine the legitimacy of traditional suborders, families and genera, and suggest that a non-skeletal sister clade (Order Corallimorpharia) might be imbedded within the stony corals. However, these studies either sampled a relatively limited array of taxa or assembled trees from heterogeneous data sets. Here we provide a more comprehensive analysis of Scleractinia (127 species, 75 genera, 17 families) and various outgroups, based on two mitochondrial genes (cytochrome oxidase I, cytochrome b), with analyses of nuclear genes (ss-tubulin, ribosomal DNA) of a subset of taxa to test unexpected relationships. Eleven of 16 families were found to be polyphyletic. Strikingly, over one third of all families as conventionally defined contain representatives from the highly divergent "robust" and "complex" clades. However, the recent suggestion that corallimorpharians are true corals that have lost their skeletons was not upheld. Relationships were supported not only by mitochondrial and nuclear genes, but also often by morphological characters which had been ignored or never noted previously. The concordance of molecular characters and more carefully examined morphological characters suggests a future of greater taxonomic stability, as well as the potential to trace the evolutionary history of this ecologically important group using fossils.

Coral reefs have emerged as one of the ecosystems most vulnerable to climate variation and change. While the contribution of a warming climate to the loss of live coral cover has been well documented across large spatial and temporal scales, the associated effects on fish have not. Here, we respond to recent and repeated calls to assess the importance of local management in conserving coral reefs in the context of global climate change. Such information is important, as coral reef fish assemblages are the most species dense vertebrate communities on earth, contributing critical ecosystem functions and providing crucial ecosystem services to human societies in tropical countries. Our assessment of the impacts of the 1998 mass bleaching event on coral cover, reef structural complexity, and reef associated fishes spans 7 countries, 66 sites and 26 degrees of latitude in the Indian Ocean. Using Bayesian meta-analysis we show that changes in the size structure, diversity and trophic composition of the reef fish community have followed coral declines. Although the ocean scale integrity of these coral reef ecosystems has been lost, it is positive to see the effects are spatially variable at multiple scales, with impacts and vulnerability affected by geography but not management regime. Existing no-take marine protected areas still support high biomass of fish, however they had no positive affect on the ecosystem response to large-scale disturbance. This suggests a need for future conservation and management efforts to identify and protect regional refugia, which should be integrated into existing management frameworks and combined with policies to improve system-wide resilience to climate variation and change.

The conservation status of 845 zooxanthellate reef-building coral species was assessed by using International Union for Conservation of Nature Red List Criteria. Of the 704 species that could be assigned conservation status, 32.8% are in categories with elevated risk of extinction. Declines in abundance are associated with bleaching and diseases driven by elevated sea surface temperatures, with extinction risk further exacerbated by local-scale anthropogenic disturbances. The proportion of corals threatened with extinction has increased dramatically in recent decades and exceeds that of most terrestrial groups. The Caribbean has the largest proportion of corals in high extinction risk categories, whereas the Coral Triangle (western Pacific) has the highest proportion of species in all categories of elevated extinction risk. Our results emphasize the widespread plight of coral reefs and the urgent need to enact conservation measures.

In 1998, more than 90% of shallow corals were killed on most Indian Ocean reefs. High sea surface temperature (SST) was a primary cause, acting directly or by interacting with other factors. Mean SSTs have been forecast to rise above the 1998 values in a few decades; however, forecast SSTs rarely flow seamlessly from historical data, or may show erroneous seasonal oscillations, precluding an accurate prediction of when lethal SSTs will recur. Differential acclimation by corals in different places complicates this further. Here I scale forecast SSTs at 33 Indian Ocean sites where most shallow corals died in 1998 (ref. 1) to identify geographical patterns in the timing of probable repeat occurrences. Reefs located 10-15 degrees south will be affected every 5 years by 2010-2025. North and south from this, dates recede in a pattern not directly related to present SSTs; paradoxically, some of the warmest sites may be affected last. Temperatures lethal to corals vary in this region by 6 degrees C, and acclimation of a modest 2 degrees C by corals could prolong their survival by nearly 100 years.

The HadISST1 sea surface temperature data set is examined for two contrasting areas: the Chagos Archipelago, central Indian Ocean which has a small (approximately 3 degrees C) annual temperature fluctuation, and Abu Dhabi in the southern Arabian Gulf whose annual air temperature fluctuation of approximately 24 degrees C is the largest known for coral reef habitats. The HadISST1 data are shown to match air temperature records closely, both in terms of annual moving averages and residual analysis. Temperatures in 1998 caused massive mortality of corals in the Indian Ocean: sea surface temperature (SST) values causing this were 33.8 degrees C in the Arabian Gulf at a time when average daily air temperature was over 40 degrees C, while in Chagos the SST lethal to corals was 29.8-29.9 degrees C, when air temperatures peaked at about 31 degrees C. The HadISST1 record was searched back to 1870 for previous abnormal peaks: one of 29.7 degrees C was found for Chagos SST in 1972, though this did not cause coral mortality. Analysis of 12-month running means of the residuals from the annual cycle show that, between 1870 and 1999, the largest SST deviations occurred between October 1997 and May 1998 in Chagos and between August 1998 and July 1999 near Abu Dhabi. The event of 1998-1999 was the largest in these regions for at least 130 years. SSTs have risen over the last three decades at rates of about 0.22 degrees or 0.23 degrees per decade in both locations.