Climate change impacts on mesophotic regions of the Great Barrier Reef.

Climate change projections for coral reefs are founded exclusively on sea surface temperatures (SST). While SST projections are relevant for the shallowest reefs, neglecting ocean stratification overlooks the striking differences in temperature experienced by deeper reefs for all or part of the year. Density stratification creates a buoyancy barrier partitioning the upper and lower parts of the water column. Here, we mechanistically downscale climate models and quantify patterns of thermal stratification above mesophotic corals (depth 30 to 50 m) of the Great Barrier Reef (GBR). Stratification insulates many offshore regions of the GBR from heatwaves at the surface. However, this protection is lost once global average temperatures exceed ~3 °C above preindustrial, after which mesophotic temperatures surpass a recognized threshold of 30 °C for coral mortality. Bottom temperatures on the GBR (30 to 50 m) from 2050 to 2060 are estimated to increase by ~0.5 to 1 °C under lower climate emissions (SSP1-1.9) and ~1.2 to 1.7 °C under higher climate emissions (SSP5-8.5). In short, mesophotic coral reefs are also threatened by climate change and research might prioritize the sensitivity of such corals to stress.

A diverse portfolio of marine protected areas can better advance global conservation and equity.

Marine protected areas (MPAs) are widely used for ocean conservation, yet the relative impacts of various types of MPAs are poorly understood. We estimated impacts on fish biomass from no-take and multiple-use (fished) MPAs, employing a rigorous matched counterfactual design with a global dataset of >14,000 surveys in and around 216 MPAs. Both no-take and multiple-use MPAs generated positive conservation outcomes relative to no protection (58.2% and 12.6% fish biomass increases, respectively), with smaller estimated differences between the two MPA types when controlling for additional confounding factors (8.3% increase). Relative performance depended on context and management: no-take MPAs performed better in areas of high human pressure but similar to multiple-use in remote locations. Multiple-use MPA performance was low in high-pressure areas but improved significantly with better management, producing similar outcomes to no-take MPAs when adequately staffed and appropriate use regulations were applied. For priority conservation areas where no-take restrictions are not possible or ethical, our findings show that a portfolio of well-designed and well-managed multiple-use MPAs represents a viable and potentially equitable pathway to advance local and global conservation.

Mapping the susceptibility of reefs to rubble accumulation across the Great Barrier Reef.

Disturbance-induced rubble accumulations are described as "killing fields" on coral reefs as coral recruits suffer high post-settlement mortality, creating a bottleneck for reef recovery. The increasing frequency of coral bleaching events, that can generate rubble once coral dies, has heightened concerns that rubble beds will become more widespread and persistent. But we currently lack the tools to predict where rubble is most likely to accumulate. Here, we developed a modelling framework to identify areas that are likely to accumulate rubble on forereef slopes across the Great Barrier Reef. The algorithm uses new high-resolution bathymetric and geomorphic datasets from satellite remote sensing. We found that 47 km of reef slope (3% of the entire reef surveyed), primarily in the southern region, could potentially reach 50% rubble cover. Despite being statistically significant (p < 0.001), the effects of depth and aspect on rubble cover were minimal, with a 0.2% difference in rubble cover between deeper and shallower regions, as well as a maximum difference of 0.8% among slopes facing various directions. Therefore, we conclude that the effects of depth and aspect were insufficient to influence ecological processes such as larval recruitment and recovery in different coral communities. Maps of potential rubble accumulation can be used to prioritise surveys and potential restoration, particularly after major disturbances have occurred.

Emergent increase in coral thermal tolerance reduces mass bleaching under climate change.

Recurrent mass bleaching events threaten the future of coral reefs. To persist under climate change, corals will need to endure progressively more intense and frequent marine heatwaves, yet it remains unknown whether their thermal tolerance can keep pace with warming. Here, we reveal an emergent increase in the thermal tolerance of coral assemblages at a rate of 0.1 °C/decade for a remote Pacific coral reef system. This led to less severe bleaching impacts than would have been predicted otherwise, indicating adaptation, acclimatisation or shifts in community structure. Using future climate projections, we show that if thermal tolerance continues to rise over the coming century at the most-likely historic rate, substantial reductions in bleaching trajectories are possible. High-frequency bleaching can be fully mitigated at some reefs under low-to-middle emissions scenarios, yet can only be delayed under high emissions scenarios. Collectively, our results indicate a potential ecological resilience to climate change, but still highlight the need for reducing carbon emissions in line with Paris Agreement commitments to preserve coral reefs.

Moving beyond heritability in the search for coral adaptive potential.

Global environmental change is happening at unprecedented rates. Coral reefs are among the ecosystems most threatened by global change. For wild populations to persist, they must adapt. Knowledge shortfalls about corals' complex ecological and evolutionary dynamics, however, stymie predictions about potential adaptation to future conditions. Here, we review adaptation through the lens of quantitative genetics. We argue that coral adaptation studies can benefit greatly from "wild" quantitative genetic methods, where traits are studied in wild populations undergoing natural selection, genomic relationship matrices can replace breeding experiments, and analyses can be extended to examine genetic constraints among traits. In addition, individuals with advantageous genotypes for anticipated future conditions can be identified. Finally, genomic genotyping supports simultaneous consideration of how genetic diversity is arrayed across geographic and environmental distances, providing greater context for predictions of phenotypic evolution at a metapopulation scale.

Research priorities for the sustainability of coral-rich western Pacific seascapes.

Nearly a billion people depend on tropical seascapes. The need to ensure sustainable use of these vital areas is recognised, as one of 17 policy commitments made by world leaders, in Sustainable Development Goal (SDG) 14 ('Life below Water') of the United Nations. SDG 14 seeks to secure marine sustainability by 2030. In a time of increasing social-ecological unpredictability and risk, scientists and policymakers working towards SDG 14 in the Asia-Pacific region need to know: (1) How are seascapes changing? (2) What can global society do about these changes? and (3) How can science and society together achieve sustainable seascape futures? Through a horizon scan, we identified nine emerging research priorities that clarify potential research contributions to marine sustainability in locations with high coral reef abundance. They include research on seascape geological and biological evolution and adaptation; elucidating drivers and mechanisms of change; understanding how seascape functions and services are produced, and how people depend on them; costs, benefits, and trade-offs to people in changing seascapes; improving seascape technologies and practices; learning to govern and manage seascapes for all; sustainable use, justice, and human well-being; bridging communities and epistemologies for innovative, equitable, and scale-crossing solutions; and informing resilient seascape futures through modelling and synthesis. Researchers can contribute to the sustainability of tropical seascapes by co-developing transdisciplinary understandings of people and ecosystems, emphasising the importance of equity and justice, and improving knowledge of key cross-scale and cross-level processes, feedbacks, and thresholds.

Demographic resilience may sustain significant coral populations in a 2°C-warmer world.

Projections of coral reefs under climate change have important policy implications, but most analyses have focused on the intensification of climate-related physical stress rather than explicitly modelling how coral populations respond to stressors. Here, we analyse the future of the Great Barrier Reef (GBR) under multiple, spatially realistic drivers which allows less impacted sites to facilitate recovery. Under a Representative Concentration Pathway (RCP) 2.6 CMIP5 climate ensemble, where warming is capped at ~2°C, GBR mean coral cover declined mid-century but approached present-day levels towards 2100. This is considerably more optimistic than most analyses. However, under RCP4.5, mean coral cover declined by >80% by late-century, and reached near zero under RCP ≥6.0. While these models do not allow for adaptation, they significantly extend past studies by revealing demographic resilience of coral populations to low levels of additional warming, though more pessimistic outcomes might be expected under CMIP6. Substantive coral populations under RCP2.6 would facilitate long-term genetic adaptation, adding value to ambitious greenhouse emissions mitigation.

No apparent trade-offs associated with heat tolerance in a reef-building coral.

As marine species adapt to climate change, their heat tolerance will likely be under strong selection. Yet trade-offs between heat tolerance and other life history traits could compromise natural adaptation or assisted evolution. This is particularly important for ecosystem engineers, such as reef-building corals, which support biodiversity yet are vulnerable to heatwave-induced mass bleaching and mortality. Here, we exposed 70 colonies of the reef-building coral Acropora digitifera to a long-term marine heatwave emulation experiment. We tested for trade-offs between heat tolerance and three traits measured from the colonies in situ - colony growth, fecundity, and symbiont community composition. Despite observing remarkable within-population variability in heat tolerance, all colonies were dominated by Cladocopium C40 symbionts. We found no evidence for trade-offs between heat tolerance and fecundity or growth. Contrary to expectations, positive associations emerged with growth, such that faster-growing colonies tended to bleach and die at higher levels of heat stress. Collectively, our results suggest that these corals exist on an energetic continuum where some high-performing individuals excel across multiple traits. Within populations, trade-offs between heat tolerance and growth or fecundity may not be major barriers to natural adaptation or the success of assisted evolution interventions.

Emigration patterns of motile cryptofauna and their implications for trophic functioning in coral reefs.

Patterns of movement of marine species can reflect strategies of reproduction and dispersal, species' interactions, trophodynamics, and susceptibility to change, and thus critically inform how we manage populations and ecosystems. On coral reefs, the density and diversity of metazoan taxa are greatest in dead coral and rubble, which are suggested to fuel food webs from the bottom up. Yet, biomass and secondary productivity in rubble is predominantly available in some of the smallest individuals, limiting how accessible this energy is to higher trophic levels. We address the bioavailability of motile coral reef cryptofauna based on small-scale patterns of emigration in rubble. We deployed modified RUbble Biodiversity Samplers (RUBS) and emergence traps in a shallow rubble patch at Heron Island, Great Barrier Reef, to detect community-level differences in the directional influx of motile cryptofauna under five habitat accessibility regimes. The mean density (0.13-4.5 ind cm-3) and biomass (0.14-5.2 mg cm-3) of cryptofauna were high and varied depending on microhabitat accessibility. Emergent zooplankton represented a distinct community (dominated by the Appendicularia and Calanoida) with the lowest density and biomass, indicating constraints on nocturnal resource availability. Mean cryptofauna density and biomass were greatest when interstitial access within rubble was blocked, driven by the rapid proliferation of small harpacticoid copepods from the rubble surface, leading to trophic simplification. Individuals with high biomass (e.g., decapods, gobies, and echinoderms) were greatest when interstitial access within rubble was unrestricted. Treatments with a closed rubble surface did not differ from those completely open, suggesting that top-down predation does not diminish rubble-derived resources. Our results show that conspecific cues and species' interactions (e.g., competition and predation) within rubble are most critical in shaping ecological outcomes within the cryptobiome. These findings have implications for prey accessibility through trophic and community size structuring in rubble, which may become increasingly relevant as benthic reef complexity shifts in the Anthropocene.

Setting sustainable limits on anchoring to improve the resilience of coral reefs.

Boat anchoring is common at coral reefs that have high economic or social value, but anchoring has received relatively little attention in reef resilience studies. We developed an individual-based model of coral populations and simulated the effects of anchor damage over time. The model allowed us to estimate the carrying capacity of anchoring for four different coral assemblages and different starting levels of coral cover. The carrying capacity of small to medium-sized recreational vessels across these four assemblages was between 0 and 3.1 anchor strikes ha-1 day-1. In a case study of two Great Barrier Reef archipelagos, we modelled the benefits of anchoring mitigation under bleaching regimes expected for four climate scenarios. The partial mitigation of even a very mild anchoring incidence (1.17 strikes ha-1 day-1) resulted in median coral gains of 2.6-7.7 % absolute cover under RCP2.6, though benefits varied temporally and depended on the Atmosphere-Ocean General Circulation Model used.

Drivers of global mangrove loss and gain in social-ecological systems.

Mangrove forests store high amounts of carbon, protect communities from storms, and support fisheries. Mangroves exist in complex social-ecological systems, hence identifying socioeconomic conditions associated with decreasing losses and increasing gains remains challenging albeit important. The impact of national governance and conservation policies on mangrove conservation at the landscape-scale has not been assessed to date, nor have the interactions with local economic pressures and biophysical drivers. Here, we assess the relationship between socioeconomic and biophysical variables and mangrove change across coastal geomorphic units worldwide from 1996 to 2016. Globally, we find that drivers of loss can also be drivers of gain, and that drivers have changed over 20 years. The association with economic growth appears to have reversed, shifting from negatively impacting mangroves in the first decade to enabling mangrove expansion in the second decade. Importantly, we find that community forestry is promoting mangrove expansion, whereas conversion to agriculture and aquaculture, often occurring in protected areas, results in high loss. Sustainable development, community forestry, and co-management of protected areas are promising strategies to reverse mangrove losses, increasing the capacity of mangroves to support human-livelihoods and combat climate change.

Functional and phylogenetic responses of motile cryptofauna to habitat degradation.

Biodiversity of terrestrial and marine ecosystems, including coral reefs, is dominated by small, often cryptic, invertebrate taxa that play important roles in ecosystem structure and functioning. While cryptofauna community structure is determined by strong small-scale microhabitat associations, the extent to which ecological and environmental factors shape these communities are largely unknown, as is the relative importance of particular microhabitats in supporting reef trophodynamics from the bottom up. The goal of this study was to address these knowledge gaps, provided coral reefs are increasingly exposed to multiple disturbances and environmental gradients that influence habitat complexity, condition and ecosystem functioning. We compared the density, biomass, size range, phylogenetic diversity and functional roles of motile cryptofauna in Palau, Western Micronesia, among four coral-derived microhabitats representing various states of degradation (live coral [Acropora and Pocillopora], dead coral and coral rubble) from reefs along a gradient of effluent exposure. In total, 122 families across ten phyla were identified, dominated by the Arthropoda (Crustacea) and Mollusca. Cryptofauna biomass was greatest in live Pocillopora, while coral rubble contained the greatest density and diversity. Size ranges were broader in live corals than both dead coral and rubble. From a bottom-up perspective, effluent exposure had mixed effects on cryptic communities including a decline in total biomass in rubble. From a top-down perspective, cryptofauna were generally unaffected by predator biomass. Our data show that, as coral reef ecosystems continue to decline in response to more frequent and severe disturbances, habitats other than live coral may become increasingly important in supporting coral reef biodiversity and food webs.

Climate refugia on the Great Barrier Reef fail when global warming exceeds 3°C.

Increases in the magnitude, frequency, and duration of warm seawater temperatures are causing mass coral mortality events across the globe. Although, even during the most extensive bleaching events, some reefs escape exposure to severe stress, constituting potential refugia. Here, we identify present-day climate refugia on the Great Barrier Reef (GBR) and project their persistence into the future. To do this, we apply semi-dynamic downscaling to an ensemble of climate projections released for the IPCC's recent sixth Assessment Report. We find that GBR locations experiencing the least thermal stress over the past 20 years have done so because of their oceanographic circumstance, which implies that longer-term persistence of climate refugia is feasible. Specifically, tidal and wind mixing of warm water away from the sea surface appears to provide relief from warming. However, on average this relative advantage only persists until global warming exceeds ~3°C.

A roadmap to integrating resilience into the practice of coral reef restoration.

Recent warm temperatures driven by climate change have caused mass coral bleaching and mortality across the world, prompting managers, policymakers, and conservation practitioners to embrace restoration as a strategy to sustain coral reefs. Despite a proliferation of new coral reef restoration efforts globally and increasing scientific recognition and research on interventions aimed at supporting reef resilience to climate impacts, few restoration programs are currently incorporating climate change and resilience in project design. As climate change will continue to degrade coral reefs for decades to come, guidance is needed to support managers and restoration practitioners to conduct restoration that promotes resilience through enhanced coral reef recovery, resistance, and adaptation. Here, we address this critical implementation gap by providing recommendations that integrate resilience principles into restoration design and practice, including for project planning and design, coral selection, site selection, and broader ecosystem context. We also discuss future opportunities to improve restoration methods to support enhanced outcomes for coral reefs in response to climate change. As coral reefs are one of the most vulnerable ecosystems to climate change, interventions that enhance reef resilience will help to ensure restoration efforts have a greater chance of success in a warming world. They are also more likely to provide essential contributions to global targets to protect natural biodiversity and the human communities that rely on reefs.

The importance of 1.5°C warming for the Great Barrier Reef.

Tropical coral reefs are among the most sensitive ecosystems to climate change and will benefit from the more ambitious aims of the United Nations Framework Convention on Climate Change's Paris Agreement, which proposed to limit global warming to 1.5° rather than 2°C above pre-industrial levels. Only in the latest Intergovernmental Panel on Climate Change focussed assessment, the Coupled Model Intercomparison Project phase 6 (CMIP6), have climate models been used to investigate the 1.5° warming scenario directly. Here, we combine the most recent model updates from CMIP6 with a semi-dynamic downscaling to evaluate the difference between the 1.5 and 2°C global warming targets on coral thermal stress metrics for the Great Barrier Reef (GBR). By ~2080, severe bleaching events are expected to occur annually under intensifying emissions (shared socioeconomic pathway SSP5-8.5). Adherence to 2° warming (SSP1-2.6) halves this frequency but the main benefit of confining warming to 1.5° (SSP1-1.9) is that bleaching events are reduced further to 3 events per decade. Attaining low emissions of 1.5° is also paramount to prevent the mean magnitude of thermal stress from stabilizing close to a critical thermal threshold (8 Degree Heating Weeks). Thermal stress under the more pessimistic pathways SSP3-7.0 and SSP5-8.5 is three to fourfold higher than the present day, with grave implications for future reef ecosystem health. As global warming continues, our projections also indicate more regional warming in the central and southern GBR than the far north and northern GBR.

Projecting coral responses to intensifying marine heatwaves under ocean acidification.

Over this century, coral reefs will run the gauntlet of climate change, as marine heatwaves (MHWs) become more intense and frequent, and ocean acidification (OA) progresses. However, we still lack a quantitative assessment of how, and to what degree, OA will moderate the responses of corals to MHWs as they intensify throughout this century. Here, we first projected future MHW intensities for tropical regions under three future greenhouse gas emissions scenario (representative concentration pathways, RCP2.6, RCP4.5 and RCP8.5) for the near-term (2021-2040), mid-century (2041-2060) and late-century (2081-2100). We then combined these MHW intensity projections with a global data set of 1,788 experiments to assess coral attribute performance and survival under the three emissions scenarios for the near-term, mid-century and late-century in the presence and absence of OA. Although warming and OA had predominately additive impacts on the coral responses, the contribution of OA in affecting most coral attributes was minor relative to the dominant role of intensifying MHWs. However, the addition of OA led to greater decreases in photosynthesis and survival under intermediate and unrestricted emissions scenario for the mid- and late-century than if intensifying MHWs were considered as the only driver. These results show that role of OA in modulating coral responses to intensifying MHWs depended on the focal coral attribute and extremity of the scenario examined. Specifically, intensifying MHWs and OA will cause increasing instances of coral bleaching and substantial declines in coral productivity, calcification and survival within the next two decades under the low and intermediate emissions scenario. These projections suggest that corals must rapidly adapt or acclimatize to projected ocean conditions to persist, which is far more likely under a low emissions scenario and with increasing efforts to manage reefs to enhance resilience.

Cumulative bleaching undermines systemic resilience of the Great Barrier Reef.

Climate change and ENSO have triggered five mass coral bleaching events on Australia's Great Barrier Reef (GBR), three of which occurred in the last 5 years.1-5 Here, we explore the cumulative nature of recent impacts and how they fragment the reef's connectivity. The coverage and intensity of thermal stress have increased steadily over time. Cumulative bleaching in 2016, 2017, and 2020 is predicted to have reduced systemic larval supply by 26%, 50%, and 71%, respectively. Larval disruption is patchy and can guide interventions. The majority of severely bleached reefs (75%) are predicted to have experienced an 80%-100% loss of larval supply. Yet restoration would not be cost-effective in the 2% of such reefs (∼30) that still experience high larval supply. Managing such climate change impacts will benefit from emerging theory on the facilitation of genetic adaptation,6,7 which requires the existence of regions with predictably high or low thermal stress. We find that a third of reefs constitute warm spots that have consistently experienced bleaching stress. Moreover, 13% of the GBR are potential refugia that avoid significant warming more than expected by chance, with a modest proportion (14%) within highly protected areas. Coral connectivity is likely to become increasingly disrupted given the predicted escalation of climate-driven disturbances,8 but the existence of thermal refugia, potentially capable of delivering larvae to 58% of the GBR, may provide pockets of systemic resilience in the near-term. Theories of conservation planning for climate change will need to consider a shifting portfolio of thermal environments over time.