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a b <br /> 0.30 -2003 20 <br /> •Positive trajectory .• <br /> -2007 15 •Negative trajectory • <br /> 0.25 -2011 0 10 . + <br /> • <br /> w m 2014 m • •' '• <br /> 0 0.20 5 <br /> - - - - - <br /> 0 0 -5 <br /> a Range in o-10 ....- <br /> 0.05 mean cover U 15 <br /> -20 f <br /> 0 10 20 30. .40 50 60 70 80 (O O10 OA 00 O°j .�O .�'� .�`1' .�`5 Na <br /> Coral cover(%) 'pO ry0 'p 'p ry0 ry0 ry0 ry0 ry0 ry0 ry0 ry0 <br /> C Year <br /> o • • • • • • N M • • N <br /> U <br /> U1 <br /> F -0.3 -0.2 -0.1 0 0.1 0.2 0.3 <br /> Squared canonical correlation(83.2%) <br /> d <br /> 400 20.0 <br /> 250 <br /> 100 10.0 <br /> 80 2.0 <br /> 60 1.5 <br /> 40 m 15 • 1.0 <br /> 20 0 0.5 t 0 a • 1� <br /> 2i 0 0 • • <br /> 0� <br /> 20 0.5 <br /> >° <br /> ° <br /> 0 <br /> 40 Z 1.0 <br /> 60 1.5 <br /> 10.0 <br /> 80 <br /> ■�■ am" <br /> _ <br /> r r r o tir �r r F �A <br /> \0c5 1 z1 <br /> o0 <br /> 0 �aQe oly G�av >L`0c Q0y \tea o'(`\ �O `ey�� .QO� \act` �,�� �ap� �ap� <br /> ,o`° �&� 0 0� Oa ee+ ec Oc Q° ear �oQ `o �a`c `a`c <br /> oe a��O�ace t1 1 Lei <br /> O<` <br /> 1 c 0 ec r� Oa <br /> S� <br /> �a <br /> 0 5r z <br /> Fig.21 Reeftrajectories predisturbance and associated local land-sea increasingly more distinct set ofconditions than expected by chance alone). <br /> human impacts and environmental factors.a,Coral cover distributions cl,Mean difference(dots)in drop-onejackknife values with upper and lower <br /> among surveyed reefs between 2003 and 2014(n=23).b,Coral cover bars representing the respective maximum and minimum differences in local <br /> trajectories ofindividual reefs.A reelfwas considered on a positive trajectory human impacts and environmental factors between positive and negative <br /> (blue;n=10)or negative trajectory(red;n=8)ifcoral cover between 2003 and trajectory reefs(n=same as in b).Blue and red shaded regions indicate factors <br /> 2014 changed by more than 3%.This cut-offwas based on mean coral cover that were greater on reefs that had positive and negative trajectories, <br /> range among all 23 reefs for the l2-yearpredisturbance period(range 2.8%;min respectively.Zero line represents equal values.See Extended Data Fig.3for <br /> 34.1%;max 36.9%).Reefs with no coral cover change(within±3%)are not shown. the percentage difference in local conditions between positive and negative <br /> c,Difference in local conditions between positive versus negative trajectory trajectory reefs.We included all local human impacts and environmental <br /> reefs(PERMANOVA,pseudo-F1,17=3.38,P=0.001)visualized along a single factors indto provide a general comparison of local conditions between reefs <br /> multivariate axis(capturing the multidimensional and correlated nature ofthe with divergent trajectories.See Fig.lb for reef locations and Supplementary <br /> data,Supplementary Fig.2)using a canonical analysis ofprincipal coordinates Fig.3 for predictor variable distributions.See Methods,Extended Data Table 1 <br /> (n=same as in b).Allocation success equalled 90 and 87.5%for positive and Supplementary Information for detailed information on local land-sea <br /> and negative trajectory reefs,respectively(more than 50%indicates an human impacts and environmental factors. <br /> inclusive of this time frame(Fig.2c).For example,the average biomass a 15 km radius).This finding supports the notion that human popula- <br /> of all fishes,all herbivorous fishes and groups of herbivorous fishes that tion density is a poor indicator of human-driven land-sea impacts at <br /> fill important ecological roles such as scrapers,grazers and browsers30 local scales".We observed minimal differences between positive and <br /> were 24-113%(29-214 kg ha-')greater on reefs with positive trajectories negative trajectory reefs relative to fishing gear restrictions,depth, <br /> compared to those with negative trajectories(Fig.2d and Extended Data sediment input,ocean temperatures,phytoplankton biomass and <br /> Fig.3).These patterns probably reflect positive feedbacks,whereby rainfall.Wave exposure was slightly higher(8.6 kW II on reefs with <br /> increasing coral cover promotes habitat suitability for reef fishes,with positive trajectories,but the difference is minor because the entire <br /> herbivorous fishes then facilitating coral growth by reducing com- study region is generally protected from large wave events34. <br /> petitive exclusion by Fleshyalgae32.By contrast,wastewater pollution, <br /> nutrient loading and urban runoff were 46-80%greater on reefs with <br /> negative trajectories compared to those with positive trajectories. Coral response to the marine heatWave <br /> Despitethese land-based human stressors being comparatively higher In 2015,the Hawaiian Islands experienced the strongest marine Keat- <br /> on reefs with negative trajectories,reefswith positive trajectories had wave on record over the past120 years(Extended Data Fig.2).Ocean <br /> 63%greater human population density(the number of people within temperatures across our study region were 2.2°C above normal and <br /> Nature I www.nature.com 1 3 <br />