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calculated following ref.81.Sediment load was assumed to scale with 2000-2014.Fishing gear restrictions were from marine managed <br /> discharge according to a approximate ratings curve following ref.82 area designation at the onset of reef surveys and the depth came from <br /> (Supplementary Figs.23 and 24). in-water diver-assessed values. <br /> The difference in local human impacts and environmental factors <br /> Fishing gear restrictions.We created a categorical value for local fish- between positiveand negative trajectory reefswere then calculated as <br /> inggear restrictions usingregulation information and marine managed the difference in the mean drop-one jackknife values for each impact <br /> area boundary designations updated from ref.80.All regulations were or factor88.Upper and lower bars in Fig.2d represent the respective <br /> evaluated for prohibition of gear categories in relation to fishing for maximum and minimum differences in drop-one jackknife values <br /> reef finfish species over time:line fishing,lay nets,spear fishing and between positive and negative trajectory reefs.Before calculating <br /> aquarium collection.Ranked fishing gear categories are as follows: the drop-one jackknife values,we identified and removed outliers <br /> (1)full no-take,(2)no lay net,spear or aquarium,(3)no lay net or aquarium, that fell outside a threshold of+2standard deviations of the median. <br /> (4)no lay net,(5)no aquarium and(6)open to all gear types(Supple- We formally tested for a difference in the local conditions of positive <br /> mentaryTable 1 and Supplementary Fig.25). versus negative trajectory reefs using a multivariate permutational <br /> analysis of variance(PERMANOVA)89 based on a Euclidean distance <br /> Sea surface temperature and heat stress.The mean and variability similarity matrix,type III(partial)sums-of-squares and unrestricted <br /> (that is,standard deviation)in summertime sea surface temperature permutationsof the normalized data.Wevisualized the results in Fig.2c <br /> (SST)was calculated over a90-day window centred on the maximum using a constrained analysis of principal coordinates90 and calculated <br /> value of a 7-day moving window average for each SST pixel(Supple- the cross-validation allocation success(a measure of group distinct- <br /> mentary Fig.26).Mean regional temperature(Fig.3a)wascalculated by ness)from the leave-one-out procedure of the constrained analysis of <br /> takingthe 7-day runningmean of daily values and then averagingacross principal coordinates analysis. <br /> all coastal pixels within our study region.Heat stress on reefs during <br /> the 2015 marine heatwave was assessed using DHW35,a widely used Coral response to the 2015 marine heatwave.Our goal was to as- <br /> metric by coral reef scientists across the world.All data were NOANs sess the local land-sea human impacts and environmental factors <br /> Coral Reef Watch v.3.1,available daily at5 km resolution35. that best explained changes in coral cover as a consequence of the <br /> 2015 marine heatwave.Any potential to observe change,however, <br /> Phytoplankton biomass and irradiance.We used satellite derived could be influenced by variations in starting condition.Reefs with <br /> chlorophyll-a(mg m-3;a proxyfor phytoplankton biomass)and irradi- higher initial cover(such as those on positive coral cover trajectories <br /> ance(E m-Z d-')from two sources.The long-term mean(2002-2013)in predisturbance,Fig.2b)had greater scope for loss and vice versa91 <br /> 8-day,4 km data were obtained from ref.80 and shown in Fig.2d and (Extended Data Fig.S).To account for this and ensure comparability <br /> Extended Data Fig.3.All subsequent analysis used thevisible-infrared across reefs(Supplementary Fig.4)we calculated coral cover change <br /> imaging/radiometer suite,which has high spatial(750 m)and tempo- following ref.92 as: <br /> ral(daily)resolution data starting in 2014(provided by NOANs Coral <br /> Reef Watch).All data were quality controlled and masked to account %differenceA=[(Az,,;-Ab,;)/AbJ j X 100 <br /> for cloud cover(Supplementary Information)and optically shallow <br /> waters following ref.83(Supplementary Fig.27). whereAb and A.are the mean coral cover values at each reef in 2014 or <br /> 2015,and 2016,respectively. <br /> Wave exposure.Wave power(kW m-')combines wave height and Wethen calculated the following predictors based on current litera- <br /> period and provides a more representative metric of wave exposure ture and our hypotheses of the principal factors that drive changes in <br /> than wave heightalone'.A series of nestedgrids(fromglobal to 50 m) coral cover owing to severe heat stress(Extended Data Table 1).Fish <br /> using WAVEWATCH III"'and Simulating Waves Nearshore116were used biomass metrics included the mean of fish data that were coupled <br /> to quantify wave transformation over the reef environment at SO m, with benthic surveys:2014(n=40)or 2015(n=40)and 2016(n=80); <br /> at hourly intervals across our study region from ref.87 and updated human population,wastewater pollution,nutrient loading,urban run- <br /> for this study.Annual data were then generated for each 50 m grid off,annual rainfall,peak rainfall and wave exposure were taken from <br /> cell by taking the mean of the top 97.5%in daily maximum wave power the mean of all data from 2012 to 2016,sediment was measured from <br /> (Supplementary Fig.28). the mean of the top three events from 2006 to 2016;SST mean and <br /> SST variability were taken from the mean from 2000 to 2014;DH W <br /> Depth.Depth of the reef floor(m)was measured using diver depth was the maximum value for 2015;phytoplankton biomass and irra- <br /> gauges during the in-water reef surveys. diance was the mean from June to November 2015,representing the <br /> time inclusive of the marine heatwave;fishing gear restrictions was <br /> Statistical analyses the marine managed area designation before the marine heatwave <br /> Coral reef trajectories predisturbance.We quantified the change (2014 or 2015,depending on the reef surveyed)and depth came from <br /> in coral cover at 23 reefs from 2003 to 2014.A reef was considered to in-water diver-assessed values. <br /> have a positive trajectory or negative trajectory if coral cover from We then tested for correlations between coral loss and our suite of <br /> the 2003 survey to the 2014 survey increased or decreased bygreater predictor variables using a generalized additive mixed-effects mod- <br /> than 3%,respectively(Fig.2b).This cut-off was based on the range in elling(GAMM)framework24 with the gamm4(ref.93)package for R <br /> mean coral cover among all 23 reefs across the 12-year period(range (www.r-project.org)v.4.0.2.Before model fitting,we identified the <br /> 2.8%;minimum 34.1%;maximum of 36.9%).We then quantified local presence of outliers in our predictor variables as any point that fell <br /> human impacts and environmental factors at each reef as follows: outside a threshold of±2 standard deviations of the median.We then <br /> fish biomass metrics were from the mean of all annual surveys for appliedan additional stepto retain any point above this threshold that <br /> each year from 2003 to 2014;human population,wastewater pol- was within 25%of the maximum predictorvalue below thethreshold. <br /> lution,nutrient loading,urban runoff,annual rainfall,peak rainfall, This ensured that no data points were unnecessarily discarded from <br /> SST mean and SST variability from the mean of all data from 2000 to our formal model-fitting process because of applying an arbitrary <br /> 2014.Phytoplankton biomass and irradiance were from the maximum threshold cut-off for data inclusion.The following predictors were <br /> monthly climatology from 2002 to 2013.Sedimentand waveexposure square-root transformed to down-weightthe influenceof values at the <br /> came from the mean of the top five events from each year spanning extreme ends of their distributions:all fish biomass metrics,wastewater <br />