The world's tropical reef ecosystems, and the people who depend on them, are increasingly 60 impacted by climate change [1][2][3][4][5][6][7] Reef, as well as the potential influence of water quality and fishing pressure on the severity of 71 bleaching. 72The geographic footprints of mass bleaching of corals on the Great Barrier Reef have varied 73 strikingly during three major events in 1998 , 2002 and 2016). In 1998, bleaching was 74 primarily coastal and most severe in the central and southern regions. In 2002, bleaching was 75 more widespread, and affected offshore reefs in the central region that had escaped in 1998 8 . 76In 2016, bleaching was even more extensive and much more severe, especially in the 77 northern, and to a lesser extent the central regions, where many coastal, mid-shelf and 78 offshore reefs were affected (Fig. 1a, b). In 2016, the proportion of reefs experiencing 79 extreme bleaching (>60% of corals bleached) was over four times higher compared to 1998 80 or 2002 (Fig. 1f) The severity and distinctive geographic footprints of bleaching in each of the three 88 years can be explained by differences in the magnitude and spatial distribution of sea-surface 89 temperature anomalies (Fig. 1a, b 102The geographic pattern of bleaching also demonstrates how marine heatwaves can be (Fig. 2a) (Fig. 1g). largely escaped bleaching in the two earlier events (Fig. 1a). Thirty-five percent of the reefs (Fig. 1b, e). We conclude that the overlap of disparate geographic bleaching at the scale of both individual reefs and the entire Great Barrier Reef (Fig. 1a, b). 134We found a similar strong relationship between the amount of bleaching measured 135 underwater, and the satellite-based estimates of heat exposure on individual reefs (Fig. 3). 136Low levels of bleaching was observed at some locations when DHW values were only 2-3 137 o C-weeks. Typically, 30-40% of corals bleached on reefs exposed to 4 o C-weeks, whereas an 138 average of 70-90% of corals bleached on reefs that experience 8 o C-weeks or more (Fig. 3). 139Resistance and adaptation to bleaching 140 Once we account for the amount of heat stress experienced on each reef, adding 141 chlorophyll-a, a proxy for water quality, to our statistical model yielded no support for the 142 hypothesis that good water quality confers resistance to bleaching 13 . Rather, the estimated 143 effect of chlorophyll-a was to significantly reduce the DHW threshold for bleaching 144 (Extended Data Table 1). However, despite the statistical significance, the effect in real terms 145 beyond heat stress alone is very small (Extended Data Fig. 1). Similarly, we found no effect 146 of the level of protection (in fished or protected zones) on bleaching (P > 0.1: Extended Data 147 Table 1). These results are consistent with the broad-scale pattern of severe bleaching in the 148 northern Great Barrier Reef, which affected hundreds of reefs across inshore-offshore 149 gradients in water quality, and regardless of their zoning (protection) status (Fig. 1a, b). 150Simila...

Tropical reef systems are transitioning to a new era in which the interval between recurrent bouts of coral bleaching is too short for a full recovery of mature assemblages. We analyzed bleaching records at 100 globally distributed reef locations from 1980 to 2016. The median return time between pairs of severe bleaching events has diminished steadily since 1980 and is now only 6 years. As global warming has progressed, tropical sea surface temperatures are warmer now during current La Niña conditions than they were during El Niño events three decades ago. Consequently, as we transition to the Anthropocene, coral bleaching is occurring more frequently in all El Niño-Southern Oscillation phases, increasing the likelihood of annual bleaching in the coming decades.

Many coral reefs worldwide have undergone phase shifts to alternate, degraded assemblages because of the combined effects of over-fishing, declining water quality, and the direct and indirect impacts of climate change. Here, we experimentally manipulated the density of large herbivorous fishes to test their influence on the resilience of coral assemblages in the aftermath of regional-scale bleaching in 1998, the largest coral mortality event recorded to date. The experiment was undertaken on the Great Barrier Reef, within a no-fishing reserve where coral abundances and diversity had been sharply reduced by bleaching. In control areas, where fishes were abundant, algal abundance remained low, whereas coral cover almost doubled (to 20%) over a 3 year period, primarily because of recruitment of species that had been locally extirpated by bleaching. In contrast, exclusion of large herbivorous fishes caused a dramatic explosion of macroalgae, which suppressed the fecundity, recruitment, and survival of corals. Consequently, management of fish stocks is a key component in preventing phase shifts and managing reef resilience. Importantly, local stewardship of fishing effort is a tractable goal for conservation of reefs, and this local action can also provide some insurance against larger-scale disturbances such as mass bleaching, which are impractical to manage directly.

Global warming is rapidly emerging as a universal threat to ecological integrity and function, highlighting the urgent need for a better understanding of the impact of heat exposure on the resilience of ecosystems and the people who depend on them . Here we show that in the aftermath of the record-breaking marine heatwave on the Great Barrier Reef in 2016 , corals began to die immediately on reefs where the accumulated heat exposure exceeded a critical threshold of degree heating weeks, which was 3-4 °C-weeks. After eight months, an exposure of 6 °C-weeks or more drove an unprecedented, regional-scale shift in the composition of coral assemblages, reflecting markedly divergent responses to heat stress by different taxa. Fast-growing staghorn and tabular corals suffered a catastrophic die-off, transforming the three-dimensionality and ecological functioning of 29% of the 3,863 reefs comprising the world's largest coral reef system. Our study bridges the gap between the theory and practice of assessing the risk of ecosystem collapse, under the emerging framework for the International Union for Conservation of Nature (IUCN) Red List of Ecosystems , by rigorously defining both the initial and collapsed states, identifying the major driver of change, and establishing quantitative collapse thresholds. The increasing prevalence of post-bleaching mass mortality of corals represents a radical shift in the disturbance regimes of tropical reefs, both adding to and far exceeding the influence of recurrent cyclones and other local pulse events, presenting a fundamental challenge to the long-term future of these iconic ecosystems.

Increased frequency of disturbances and anthropogenic activities are predicted to have a devastating impact on coral reefs that will ultimately change the composition of reef associated fish communities. We reviewed and analysed studies that document the effects of disturbance-mediated coral loss on coral reef fishes. Meta-analysis of 17 independent studies revealed that 62% of fish species declined in abundance within 3 years of disturbances that resulted in 410% decline in coral cover. Abundances of species reliant on live coral for food and shelter consistently declined during this time frame, while abundance of some species that feed on invertebrates, algae and/or detritus increased. The response of species, particularly those expected to benefit from the immediate loss of coral, is, however, variable and is attributed to erratic replenishment of stocks, ecological versatility of species and sublethal responses, such as changes in growth, body condition and feeding rates. The diversity of fish communities was found to be negatively and linearly correlated to disturbance-mediated coral loss. Coral loss 420% typically resulted in a decline in species richness of fish communities, although diversity may initially increase following small declines in coral cover from high coverage. Disturbances that result in an immediate loss of habitat complexity (e.g. severe tropical storms), have a greater impact on fishes from all trophic levels, compared with disturbances that kill corals, but leave the reef framework intact (e.g. coral bleaching and outbreaks of Acanthaster planci). This is most evident among small bodied species and suggests the long-term consequences of coral loss through coral bleaching and crown-ofthorn starfish outbreaks may be much more substantial than the short-term effects currently documented.

The persistence of most coastal marine species depends on larvae finding suitable adult habitat at the end of an offshore dispersive stage that can last weeks or months. We tested the effects that ocean acidification from elevated levels of atmospheric carbon dioxide (CO2) could have on the ability of larvae to detect olfactory cues from adult habitats. Larval clownfish reared in control seawater (pH 8.15) discriminated between a range of cues that could help them locate reef habitat and suitable settlement sites. This discriminatory ability was disrupted when larvae were reared in conditions simulating CO2-induced ocean acidification. Larvae became strongly attracted to olfactory stimuli they normally avoided when reared at levels of ocean pH that could occur ca. 2100 (pH 7.8) and they no longer responded to any olfactory cues when reared at pH levels (pH 7.6) that might be attained later next century on a business-as-usual carbon-dioxide emissions trajectory. If acidification continues unabated, the impairment of sensory ability will reduce population sustainability of many marine species, with potentially profound consequences for marine diversity.climate change ͉ larval sensory mechanisms ͉ population connectivity ͉ population replenishment O cean acidification caused by the uptake of additional carbon dioxide (CO 2 ) at the ocean surface is now recognized as a serious threat to marine ecosystems (1-4). At least 30% of the anthropogenic CO 2 released into the atmosphere in the past 200 years has been absorbed by the oceans, causing ocean pH to decline at a rate Ϸ100 times faster than at any time in the past 650,000 years (1, 4). Global ocean pH is estimated to have dropped by 0.1 units since preindustrial times and is projected to fall another 0.3-0.4 units by 2100 because of existing and future CO 2 emissions (1, 5-6). Considerable research effort has focused on predicting the impact that reduced carbonate-ion saturation states that accompany ocean acidification will have on calcifying marine organisms, particularly corals and other invertebrates that precipitate aragonite skeletons (2-3, 6). However, the effects that ocean acidification will have on other marine organisms, including fishes, remain almost completely unknown, especially for conditions of atmospheric carbon dioxide and seawater pH that could occur in the near future (4, 7-9).The persistence of most coastal marine species depends on the ability of larvae to locate suitable settlement habitat at the end of a pelagic stage that can last weeks or months. Accumulating evidence for reef fishes suggests that both reef sounds (10) and olfactory cues (11-13) are used by larvae to locate reefs. The olfactory organs of many reef fishes are well-developed by the end of the larval phase (14-15), and it has recently been shown that larvae of some species can discriminate the smell of water from their natal reef compared with water from other reefs (13), which provides a mechanism to explain high levels of selfrecruitment in some reef fish populations (16)(...

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Coral reef recovery from major disturbance is hypothesized to depend on the arrival of propagules from nearby undisturbed reefs. Therefore, reefs isolated by distance or current patterns are thought to be highly vulnerable to catastrophic disturbance. We found that on an isolated reef system in north Western Australia, coral cover increased from 9% to 44% within 12 years of a coral bleaching event, despite a 94% reduction in larval supply for 6 years after the bleaching. The initial increase in coral cover was the result of high rates of growth and survival of remnant colonies, followed by a rapid increase in juvenile recruitment as colonies matured. We show that isolated reefs can recover from major disturbance, and that the benefits of their isolation from chronic anthropogenic pressures can outweigh the costs of limited connectivity.