Evidence for managing herbivores for reef resilience.

Herbivore management is an important tool for resilience-based approaches to coral reef conservation, and evidence-based science is needed to enact successful management. We synthesized data from multiple monitoring programs in Hawai'i to measure herbivore biomass and benthic condition over a 10-year period preceding any major coral bleaching. We analysed data from 20 242 transects alongside data on 27 biophysical and human drivers and found herbivore biomass was highly variable throughout Hawai'i, with high values in remote locations and the lowest values near population centres. Both human and biophysical drivers explained variation in herbivore biomass, and among the human drivers both fishing and land-based pollution had negative effects on biomass. We also found evidence that herbivore functional group biomass is strongly linked to benthic condition, and that benthic condition is sensitive to changes in herbivore biomass associated with fishing. We show that when herbivore biomass is below 80% of potential biomass, benthic condition is predicted to decline. We also show that a range of management actions, including area-specific fisheries regulations and gear restrictions, can increase parrotfish biomass. Together, these results provide lines of evidence to support managing herbivores as an effective strategy for maintaining or bolstering reef resilience in a changing climate.

Local human impacts disrupt depth-dependent zonation of tropical reef fish communities.

The influence of depth and associated gradients in light, nutrients and plankton on the ecological organization of tropical reef communities was first described over six decades ago but remains untested across broad geographies. During this time humans have become the dominant driver of planetary change, requiring that we revisit historic ecological paradigms to ensure they capture the dynamics of contemporary ecological systems. Analysing >5,500 in-water reef fish surveys between 0 and 30 m depth on reef slopes of 35 islands across the Pacific, we assess whether a depth gradient consistently predicts variation in reef fish biomass. We reveal predictable ecological organization at unpopulated locations, with increased biomass of planktivores and piscivores and decreased primary consumer biomass with increasing depth. Bathymetric steepness also had a striking influence on biomass patterns, primarily for planktivores, emphasizing potential links between local hydrodynamics and the upslope propagation of pelagic subsidies to the shallows. However, signals of resource-driven change in fish biomass with depth were altered or lost for populated islands, probably due to depleted fish biomass baselines. While principles of depth zonation broadly held, our findings expose limitations of the paradigm for predicting ecological dynamics where human impacts confound connections between ecological communities and their surrounding environment.

Coral reefs benefit from reduced land-sea impacts under ocean warming.

Coral reef ecosystems are being fundamentally restructured by local human impacts and climate-driven marine heatwaves that trigger mass coral bleaching and mortality1. Reducing local impacts can increase reef resistance to and recovery from bleaching2. However, resource managers lack clear advice on targeted actions that best support coral reefs under climate change3 and sector-based governance means most land- and sea-based management efforts remain siloed4. Here we combine surveys of reef change with a unique 20-year time series of land-sea human impacts that encompassed an unprecedented marine heatwave in Hawai'i. Reefs with increased herbivorous fish populations and reduced land-based impacts, such as wastewater pollution and urban runoff, had positive coral cover trajectories predisturbance. These reefs also experienced a modest reduction in coral mortality following severe heat stress compared to reefs with reduced fish populations and enhanced land-based impacts. Scenario modelling indicated that simultaneously reducing land-sea human impacts results in a three- to sixfold greater probability of a reef having high reef-builder cover four years postdisturbance than if either occurred in isolation. International efforts to protect 30% of Earth's land and ocean ecosystems by 2030 are underway5. Our results reveal that integrated land-sea management could help achieve coastal ocean conservation goals and provide coral reefs with the best opportunity to persist in our changing climate.

Surface slicks are pelagic nurseries for diverse ocean fauna.

Most marine animals have a pelagic larval phase that develops in the coastal or open ocean. The fate of larvae has profound effects on replenishment of marine populations that are critical for human and ecosystem health. Larval ecology is expected to be tightly coupled to oceanic features, but for most taxa we know little about the interactions between larvae and the pelagic environment. Here, we provide evidence that surface slicks, a common coastal convergence feature, provide nursery habitat for diverse marine larvae, including > 100 species of commercially and ecologically important fishes. The vast majority of invertebrate and larval fish taxa sampled had mean densities 2-110 times higher in slicks than in ambient water. Combining in-situ surveys with remote sensing, we estimate that slicks contain 39% of neustonic larval fishes, 26% of surface-dwelling zooplankton (prey), and 75% of floating organic debris (shelter) in our 1000 km2 study area in Hawai'i. Results indicate late-larval fishes actively select slick habitats to capitalize on concentrations of diverse prey and shelter. By providing these survival advantages, surface slicks enhance larval supply and replenishment of adult populations from coral reef, epipelagic, and deep-water ecosystems. Our findings suggest that slicks play a critically important role in enhancing productivity in tropical marine ecosystems.

Impacts of pollution, fishing pressure, and reef rugosity on resource fish biomass in West Hawaii.

Human activities and land-use drivers combine in complex ways to affect coral reef health and, in turn, the diversity and abundance of reef fauna. Here we examine the impacts of different marine protected area (MPA) types, and various human and habitat drivers, on resource fish functional groups (i.e., total fish, herbivore, grazer, scraper, and browser biomass) along the 180 km west coast of Hawaii Island. Across survey years from 2008 to 2018, we observed an overall decrease in total fish biomass of 45%, with similar decreases in biomass seen across most fish functional groups. MPAs that prohibited a combination of lay nets, aquarium collection, and spear fishing were most effective in maintaining and/or increasing fish biomass across all functional groups. We also found that pollution, fishing, and habitat drivers all contributed to changes in total fish biomass, where the most negative impact was nitrogen input from land-based sewage disposal. Fish biomass relationships with our study drivers depended on fish functional grouping. For surgeonfish (grazers), changes in biomass linked most strongly to changes in reef rugosity. For parrotfish (scrapers), biomass was better explained by changes in commercial catch where current commercial fishing levels are negatively affecting scraper populations. Our observations suggest that regional management of multiple factors, including habitat, pollution, and fisheries, will benefit resource fish biomass off Hawaii Island.

Prey-size plastics are invading larval fish nurseries.

Life for many of the world's marine fish begins at the ocean surface. Ocean conditions dictate food availability and govern survivorship, yet little is known about the habitat preferences of larval fish during this highly vulnerable life-history stage. Here we show that surface slicks, a ubiquitous coastal ocean convergence feature, are important nurseries for larval fish from many ocean habitats at ecosystem scales. Slicks had higher densities of marine phytoplankton (1.7-fold), zooplankton (larval fish prey; 3.7-fold), and larval fish (8.1-fold) than nearby ambient waters across our study region in Hawai'i. Slicks contained larger, more well-developed individuals with competent swimming abilities compared to ambient waters, suggesting a physiological benefit to increased prey resources. Slicks also disproportionately accumulated prey-size plastics, resulting in a 60-fold higher ratio of plastics to larval fish prey than nearby waters. Dissections of hundreds of larval fish found that 8.6% of individuals in slicks had ingested plastics, a 2.3-fold higher occurrence than larval fish from ambient waters. Plastics were found in 7 of 8 families dissected, including swordfish (Xiphiidae), a commercially targeted species, and flying fish (Exocoetidae), a principal prey item for tuna and seabirds. Scaling up across an ∼1,000 km2 coastal ecosystem in Hawai'i revealed slicks occupied only 8.3% of ocean surface habitat but contained 42.3% of all neustonic larval fish and 91.8% of all floating plastics. The ingestion of plastics by larval fish could reduce survivorship, compounding threats to fisheries productivity posed by overfishing, climate change, and habitat loss.

Parsing human and biophysical drivers of coral reef regimes.

Coral reefs worldwide face unprecedented cumulative anthropogenic effects of interacting local human pressures, global climate change and distal social processes. Reefs are also bound by the natural biophysical environment within which they exist. In this context, a key challenge for effective management is understanding how anthropogenic and biophysical conditions interact to drive distinct coral reef configurations. Here, we use machine learning to conduct explanatory predictions on reef ecosystems defined by both fish and benthic communities. Drawing on the most spatially extensive dataset available across the Hawaiian archipelago-20 anthropogenic and biophysical predictors over 620 survey sites-we model the occurrence of four distinct reef regimes and provide a novel approach to quantify the relative influence of human and environmental variables in shaping reef ecosystems. Our findings highlight the nuances of what underpins different coral reef regimes, the overwhelming importance of biophysical predictors and how a reef's natural setting may either expand or narrow the opportunity space for management interventions. The methods developed through this study can help inform reef practitioners and hold promises for replication across a broad range of ecosystems.

Combining fish and benthic communities into multiple regimes reveals complex reef dynamics.

Coral reefs worldwide face an uncertain future with many reefs reported to transition from being dominated by corals to macroalgae. However, given the complexity and diversity of the ecosystem, research on how regimes vary spatially and temporally is needed. Reef regimes are most often characterised by their benthic components; however, complex dynamics are associated with losses and gains in both fish and benthic assemblages. To capture this complexity, we synthesised 3,345 surveys from Hawai'i to define reef regimes in terms of both fish and benthic assemblages. Model-based clustering revealed five distinct regimes that varied ecologically, and were spatially heterogeneous by island, depth and exposure. We identified a regime characteristic of a degraded state with low coral cover and fish biomass, one that had low coral but high fish biomass, as well as three other regimes that varied significantly in their ecology but were previously considered a single coral dominated regime. Analyses of time series data reflected complex system dynamics, with multiple transitions among regimes that were a function of both local and global stressors. Coupling fish and benthic communities into reef regimes to capture complex dynamics holds promise for monitoring reef change and guiding ecosystem-based management of coral reefs.

Advancing the integration of spatial data to map human and natural drivers on coral reefs.

A major challenge for coral reef conservation and management is understanding how a wide range of interacting human and natural drivers cumulatively impact and shape these ecosystems. Despite the importance of understanding these interactions, a methodological framework to synthesize spatially explicit data of such drivers is lacking. To fill this gap, we established a transferable data synthesis methodology to integrate spatial data on environmental and anthropogenic drivers of coral reefs, and applied this methodology to a case study location-the Main Hawaiian Islands (MHI). Environmental drivers were derived from time series (2002-2013) of climatological ranges and anomalies of remotely sensed sea surface temperature, chlorophyll-a, irradiance, and wave power. Anthropogenic drivers were characterized using empirically derived and modeled datasets of spatial fisheries catch, sedimentation, nutrient input, new development, habitat modification, and invasive species. Within our case study system, resulting driver maps showed high spatial heterogeneity across the MHI, with anthropogenic drivers generally greatest and most widespread on O'ahu, where 70% of the state's population resides, while sedimentation and nutrients were dominant in less populated islands. Together, the spatial integration of environmental and anthropogenic driver data described here provides a first-ever synthetic approach to visualize how the drivers of coral reef state vary in space and demonstrates a methodological framework for implementation of this approach in other regions of the world. By quantifying and synthesizing spatial drivers of change on coral reefs, we provide an avenue for further research to understand how drivers determine reef diversity and resilience, which can ultimately inform policies to protect coral reefs.

Seascape models reveal places to focus coastal fisheries management.

To design effective marine reserves and support fisheries, more information on fishing patterns and impacts for targeted species is needed, as well as better understanding of their key habitats. However, fishing impacts vary geographically and are difficult to disentangle from other factors that influence targeted fish distributions. We developed a set of fishing effort and habitat layers at high resolution and employed machine learning techniques to create regional-scale seascape models and predictive maps of biomass and body length of targeted reef fishes for the main Hawaiian Islands. Spatial patterns of fishing effort were shown to be highly variable and seascape models indicated a low threshold beyond which targeted fish assemblages were severely impacted. Topographic complexity, exposure, depth, and wave power were identified as key habitat variables that influenced targeted fish distributions and defined productive habitats for reef fisheries. High targeted reef fish biomass and body length were found in areas not easily accessed by humans, while model predictions when fishing effort was set to zero showed these high values to be more widely dispersed among suitable habitats. By comparing current targeted fish distributions with those predicted when fishing effort was removed, areas with high recovery potential on each island were revealed, with average biomass recovery of 517% and mean body length increases of 59% on Oahu, the most heavily fished island. Spatial protection of these areas would aid recovery of nearshore coral reef fisheries.

Managing Local Stressors for Coral Reef Condition and Ecosystem Services Delivery Under Climate Scenarios.

Coral reefs provide numerous ecosystem goods and services, but are threatened by multiple environmental and anthropogenic stressors. To identify management scenarios that will reverse or mitigate ecosystem degradation, managers can benefit from tools that can quantify projected changes in ecosystem services due to alternative management options. We used a spatially-explicit biophysical ecosystem model to evaluate socio-ecological trade-offs of land-based vs. marine-based management scenarios, and local-scale vs. global-scale stressors and their cumulative impacts. To increase the relevance of understanding ecological change for the public and decision-makers, we used four ecological production functions to translate the model outputs into the ecosystem services: "State of the Reef," "Trophic Integrity," "Fisheries Production," and "Fisheries Landings." For a case study of Maui Nui, Hawai'i, land-based management attenuated coral cover decline whereas fisheries management promoted higher total fish biomass. Placement of no-take marine protected areas (MPAs) across 30% of coral reef areas led to a reversal of the historical decline in predatory fish biomass, although this outcome depended on the spatial arrangement of MPAs. Coral cover declined less severely under strict sediment mitigation scenarios. However, the benefits of these local management scenarios were largely lost when accounting for climate-related impacts. Climate-related stressors indirectly increased herbivore biomass due to the shift from corals to algae and, hence, greater food availability. The two ecosystem services related to fish biomass increased under climate-related stressors but "Trophic Integrity" of the reef declined, indicating a less resilient reef. "State of the Reef" improved most and "Trophic Integrity" declined least under an optimistic global warming scenario and strict local management. This work provides insight into the relative influence of land-based vs. marine-based management and local vs. global stressors as drivers of changes in ecosystem dynamics while quantifying the tradeoffs between conservation- and extraction-oriented ecosystem services.

Upstream solutions to coral reef conservation: The payoffs of smart and cooperative decision-making.

Land-based source pollutants (LBSP) actively threaten coral reef ecosystems globally. To achieve the greatest conservation outcome at the lowest cost, managers could benefit from appropriate tools that evaluate the benefits (in terms of LBSP reduction) and costs of implementing alternative land management strategies. Here we use a spatially explicit predictive model (InVEST-SDR) that quantifies change in sediment reaching the coast for evaluating the costs and benefits of alternative threat-abatement scenarios. We specifically use the model to examine trade-offs among possible agricultural road repair management actions (water bars to divert runoff and gravel to protect the road surface) across the landscape in West Maui, Hawaii, USA. We investigated changes in sediment delivery to coasts and costs incurred from management decision-making that is (1) cooperative or independent among landowners, and focused on (2) minimizing costs, reducing sediment, or both. The results illuminate which management scenarios most effectively minimize sediment while also minimizing the cost of mitigation efforts. We find targeting specific "hotspots" within all individual parcels is more cost-effective than targeting all road segments. The best outcomes are achieved when landowners cooperate and target cost-effective road repairs, however, a cooperative strategy can be counter-productive in some instances when cost-effectiveness is ignored. Simple models, such as the one developed here, have the potential to help managers make better choices about how to use limited resources.

Identifying Suitable Locations for Mesophotic Hard Corals Offshore of Maui, Hawai'i.

Mesophotic hard corals (MHC) are increasingly threatened by a growing number of anthropogenic stressors, including impacts from fishing, land-based sources of pollution, and ocean acidification. However, little is known about their geographic distributions (particularly around the Pacific islands) because it is logistically challenging and expensive to gather data in the 30 to 150 meter depth range where these organisms typically live. The goal of this study was to begin to fill this knowledge gap by modelling and predicting the spatial distribution of three genera of mesophotic hard corals offshore of Maui in the Main Hawaiian Islands. Maximum Entropy modeling software was used to create separate maps of predicted probability of occurrence and uncertainty for: (1) Leptoseris, (2) Montipora, and (3) Porites. Genera prevalence was derived from the in situ presence/absence data, and used to convert relative habitat suitability to probability of occurrence values. Approximately 1,300 georeferenced records of the occurrence of MHC, and 34 environmental predictors were used to train the model ensembles. Receiver Operating Characteristic (ROC) Area Under the Curve (AUC) values were between 0.89 and 0.97, indicating excellent overall model performance. Mean uncertainty and mean absolute error for the spatial predictions ranged from 0.006% to 0.05% and 3.73% to 17.6%, respectively. Depth, distance from shore, euphotic depth (mean and standard deviation) and sea surface temperature (mean and standard deviation) were identified as the six most influential predictor variables for partitioning habitats among the three genera. MHC were concentrated between Hanaka'o'o and Papawai Points offshore of western Maui most likely because this area hosts warmer, clearer and calmer water conditions almost year round. While these predictions helped to fill some knowledge gaps offshore of Maui, many information gaps remain in the Hawaiian Archipelago and Pacific Islands. This approach may be used to identify other potentially suitable areas for MHCs, helping scientists and resource managers prioritize sites, and focus their limited resources on areas that may be of higher scientific or conservation value.