Diversification of refugia types needed to secure the future of coral reefs subject to climate change.

Identifying locations of refugia from the thermal stresses of climate change for coral reefs and better managing them is one of the key recommendations for climate change adaptation. We review and summarize approximately 30 years of applied research focused on identifying climate refugia to prioritize the conservation actions for coral reefs under rapid climate change. We found that currently proposed climate refugia and the locations predicted to avoid future coral losses are highly reliant on  excess heat metrics, such as degree heating weeks. However, many existing alternative environmental, ecological, and life-history variables could be used to identify other types of refugia that lead to the desired diversified portfolio for coral reef conservation. To improve conservation priorities for coral reefs, there is a need to evaluate and validate the predictions of climate refugia with long-term field data on coral abundance, diversity, and functioning. There is also the need to identify and safeguard locations displaying resistance toprolonged exposure to heat waves and the ability to recover quickly after thermal exposure. We recommend using more metrics to identify a portfolio of potential refugia sites for coral reefs that can avoid, resist, and recover from exposure to high ocean temperatures and the consequences of climate change, thereby shifting past efforts focused on avoidance to a diversified risk-spreading portfolio that can be used to improve strategic coral reef conservation in a rapidly warming climate.

Diversificación de los tipos de refugio necesarios para asegurar el futuro de los arrecifes de coral sujetos al cambio climático Resumen Una de las principales recomendaciones para la adaptación al cambio climático es identificar los refugios de los arrecifes de coral frente al estrés térmico del cambio climático y mejorar su gestión. Revisamos y resumimos ∼30 años de investigación aplicada centrada en la identificación de refugios climáticos para priorizar las acciones de conservación de los arrecifes de coral bajo un rápido cambio climático. Descubrimos que los refugios climáticos propuestos actualmente y las ubicaciones que pueden evitarlos dependen en gran medida de métricas de exceso de calor, como las semanas de calentamiento en grados (SCG). Sin embargo, existen muchas variables alternativas de historia vital, ambientales y ecológicas que podrían utilizarse para identificar otros tipos de refugios que resulten en el acervo diversificado que se desea para la conservación de los arrecifes de coral. Para mejorar las prioridades de conservación de los arrecifes de coral, es necesario evaluar y validar las predicciones sobre refugios climáticos con datos de campo a largo plazo sobre abundancia, diversidad y funcionamiento de los corales. También es necesario identificar y salvaguardar lugares que muestren resistencia a la exposición climática prolongada a olas de calor y la capacidad de recuperarse rápidamente tras la exposición térmica. Recomendamos utilizar más métricas para identificar un acervo de posibles lugares de refugio para los arrecifes de coral que puedan evitar, resistir y recuperarse de la exposición a las altas temperaturas oceánicas y las consecuencias del cambio climático, para así desplazar los esfuerzos pasados centrados en la evitación hacia un acervo diversificado de riesgos que pueda utilizarse para mejorar la conservación estratégica de los arrecifes de coral en un clima que se calienta rápidamente.

Mid-Holocene expansion of the Indian Ocean warm pool documented in coral Sr/Ca records from Kenya.

Proxy reconstructions suggest that mid-Holocene East African temperatures were warmer than today between 8 and 5 ka BP, but climate models cannot replicate this warming. Precessional forcing caused a shift of maximum insolation from boreal spring to fall in the mid-Holocene, which may have favored intense warming at the start of the warm season. Here, we use three Porites corals from Kenya that represent time windows from 6.55 to 5.87 ka BP to reconstruct past sea surface temperature (SST) seasonality from coral Sr/Ca ratios in the western Indian Ocean during the mid-Holocene. Although the Indian monsoon was reportedly stronger in the mid-Holocene, which should have amplified the seasonal cycle of SST in the western Indian Ocean, the corals suggest reduced seasonality (mean 3.2 °C) compared to the modern record (mean 4.3 °C). Warming in austral spring is followed by a prolonged period of warm SSTs, suggesting that an upper limit of tropical SSTs under mid-Holocene conditions was reached at the start of the warm season, and SSTs then remained stable. Similar changes are seen at the Seychelles. Bootstrap estimates suggest a reduction in SST seasonality of 1.3 ± 0.22 °C at Kenya and 1.7 ± 0.32 °C at the Seychelles. SST seasonality at Kenya corresponds to present-day SST seasonality at 55° E-60° E, while SST seasonality at the Seychelles corresponds to present day SST seasonality at ~ 65° E. This implies a significant westward expansion of the Indian Ocean warm pool. Furthermore, the coral data suggests that SST seasonality deviates from seasonal changes in orbital insolation due to ocean-atmosphere interactions.

Corals reveal ENSO-driven synchrony of climate impacts on both terrestrial and marine ecosystems in northern Borneo.

Extreme climate events, such as the El Niños in 1997/1998 and 2015/16, have led to considerable forest loss in the Southeast Asian region following unprecedented drought and wildfires. In Borneo, the effects of extreme climate events have been exacerbated by rapid urbanization, accelerated deforestation and soil erosion since the 1980s. However, studies quantifying the impact of interannual and long-term (>3 decades) climatic and anthropogenic change affecting Borneo's coastal and coral reef environments are lacking. Here, we used coral cores collected in Miri-Sibuti Coral Reefs National Park, Sarawak (Malaysia) to reconstruct the spatio-temporal dynamics of sea surface temperature and oxygen isotopic composition of seawater from 1982 to 2016, based on paired oxygen isotope and Sr/Ca measurements. The results revealed rising sea surface temperatures of 0.26 ± 0.04 °C per decade since 1982. Reconstructed δ18Osw displayed positive excursion during major El Niño events of 1983, 1997/98 and 2015/16, indicating drought conditions with less river runoff, rainfall and higher ocean salinities. La Niñas were generally associated with lower δ18Osw. We observed a long-term shift from more saline conditions between 1982 and 1995 towards less saline conditions after 1995, which are in agreement with the regional freshening trend, punctuated by saline excursion during El Niños. The decadal shifts were found to be driven by the Pacific Decadal Oscillation (PDO). This study provides the first long-term data on El Niño Southern Oscillation (ENSO)-driven synchrony of climate impacts on both terrestrial and marine ecosystems in northern Borneo. Our results suggest that coral records from northern Borneo are invaluable archives to detect regional ENSO and PDO impacts, and their interaction with the Asian-Australian monsoon, on the hydrological balance in the southern South China Sea beyond the past three decades.

Borneo coral reefs subject to high sediment loads show evidence of resilience to various environmental stressors.

For reefs in South East Asia the synergistic effects of rapid land development, insufficient environmental policies and a lack of enforcement has led to poor water quality and compromised coral health from increased sediment and pollution. Those inshore turbid coral reefs, subject to significant sediment inputs, may also inherit some resilience to the effects of thermal stress and coral bleaching. We studied the inshore turbid reefs near Miri, in northwest Borneo through a comprehensive assessment of coral cover and health in addition to quantifying sediment-related parameters. Although Miri's Reefs had comparatively low coral species diversity, dominated by massive and encrusting forms of Diploastrea, Porites, Montipora, Favites, Dipsastrea and Pachyseris, they were characterized by a healthy cover ranging from 22 to 39%. We found a strong inshore to offshore gradient in hard coral cover, diversity and community composition as a direct result of spatial differences in sediment at distances <10 km. As well as distance to shore, we included other environmental variables like reef depth and sediment trap accumulation and particle size that explained 62.5% of variation in benthic composition among sites. Miri's reefs showed little evidence of coral disease and relatively low prevalence of compromised health signs including bleaching (6.7%), bioerosion (6.6%), pigmentation response (2.2%), scars (1.1%) and excessive mucus production (0.5%). Tagged colonies of Diploastrea and Pachyseris suffering partial bleaching in 2016 had fully (90-100%) recovered the following year. There were, however, seasonal differences in bioerosion rates, which increased five-fold after the 2017 wet season. Differences in measures of coral physiology, like that of symbiont density and chlorophyll a for Montipora, Pachyseris and Acropora, were not detected among sites. We conclude that Miri's reefs may be in a temporally stable state given minimal recently dead coral and a limited decline in coral cover over the last two decades. This study provides further evidence that turbid coral reefs exposed to seasonally elevated sediment loads can exhibit relatively high coral cover and be resilient to disease and elevated sea surface temperatures.

Opposite polarities of ENSO drive distinct patterns of coral bleaching potentials in the southeast Indian Ocean.

Episodic anomalously warm sea surface temperature (SST) extremes, or marine heatwaves (MHWs), amplify ocean warming effects and may lead to severe impacts on marine ecosystems. MHW-induced coral bleaching events have been observed frequently in recent decades in the southeast Indian Ocean (SEIO), a region traditionally regarded to have resilience to global warming. In this study, we assess the contribution of El Niño-Southern Oscillation (ENSO) to MHWs across the mostly understudied reefs in the SEIO. We find that in extended summer months, the MHWs at tropical and subtropical reefs (divided at ~20°S) are driven by opposite ENSO polarities: MHWs are more likely to occur at the tropical reefs during eastern Pacific El Niño, driven by enhanced solar radiation and weaker Australian Monsoon, some likely alleviated by positive Indian Ocean Dipole events, and at the subtropical reefs during central Pacific La Niña, mainly caused by increased horizontal heat transport, and in some cases reinforced by local air-sea interactions. Madden-Julian Oscillations (MJO) also modulate the MHW occurrences. Projected future increases in ENSO and MJO intensity with greenhouse warming will enhance thermal stress across the SEIO. Implementing forecasting systems of MHWs can be used to anticipate future coral bleaching patterns and prepare management responses.

Coral Sr/Ca-based sea surface temperature and air temperature variability from the inshore and offshore corals in the Seribu Islands, Indonesia.

The ability of massive Porites corals to faithfully record temperature is assessed. Porites corals from Kepulauan Seribu were sampled from one inshore and one offshore site and analyzed for their Sr/Ca variation. The results show that Sr/Ca of the offshore coral tracked SST, while Sr/Ca variation of the inshore coral tracked ambient air temperature. In particular, the inshore SST variation is related to air temperature anomalies of the urban center of Jakarta. The latter we relate to air-sea interactions modifying inshore SST associated with the land-sea breeze mechanism and/or monsoonal circulation. The correlation pattern of monthly coral Sr/Ca with the Niño3.4 index and SEIO-SST reveals that corals in the Seribu islands region respond differently to remote forcing. An opposite response is observed for inshore and offshore corals in response to El Niño onset, yet similar to El Niño mature phase (December to February). SEIO SSTs co-vary strongly with SST and air temperature variability across the Seribu island reef complex. The results of this study clearly indicate that locations of coral proxy record in Indonesia need to be chosen carefully in order to identify the seasonal climate response to local and remote climate and anthropogenic forcing.

Evidence for climate-driven synchrony of marine and terrestrial ecosystems in northwest Australia.

The effects of climate change are difficult to predict for many marine species because little is known of their response to climate variations in the past. However, long-term chronologies of growth, a variable that integrates multiple physical and biological factors, are now available for several marine taxa. These allow us to search for climate-driven synchrony in growth across multiple taxa and ecosystems, identifying the key processes driving biological responses at very large spatial scales. We hypothesized that in northwest (NW) Australia, a region that is predicted to be strongly influenced by climate change, the El Niño Southern Oscillation (ENSO) phenomenon would be an important factor influencing the growth patterns of organisms in both marine and terrestrial environments. To test this idea, we analyzed existing growth chronologies of the marine fish Lutjanus argentimaculatus, the coral Porites spp. and the tree Callitris columellaris and developed a new chronology for another marine fish, Lethrinus nebulosus. Principal components analysis and linear model selection showed evidence of ENSO-driven synchrony in growth among all four taxa at interannual time scales, the first such result for the Southern Hemisphere. Rainfall, sea surface temperatures, and sea surface salinities, which are linked to the ENSO system, influenced the annual growth of fishes, trees, and corals. All four taxa had negative relationships with the Niño-4 index (a measure of ENSO status), with positive growth patterns occurring during strong La Niña years. This finding implies that future changes in the strength and frequency of ENSO events are likely to have major consequences for both marine and terrestrial taxa. Strong similarities in the growth patterns of fish and trees offer the possibility of using tree-ring chronologies, which span longer time periods than those of fish, to aid understanding of both historical and future responses of fish populations to climate variation.

Contrasting environmental drivers of adult and juvenile growth in a marine fish: implications for the effects of climate change.

Many marine fishes have life history strategies that involve ontogenetic changes in the use of coastal habitats. Such ontogenetic shifts may place these species at particular risk from climate change, because the successive environments they inhabit can differ in the type, frequency and severity of changes related to global warming. We used a dendrochronology approach to examine the physical and biological drivers of growth of adult and juvenile mangrove jack (Lutjanus argentimaculatus) from tropical north-western Australia. Juveniles of this species inhabit estuarine environments and adults reside on coastal reefs. The Niño-4 index, a measure of the status of the El Niño-Southern Oscillation (ENSO) had the highest correlation with adult growth chronologies, with La Niña years (characterised by warmer temperatures and lower salinities) having positive impacts on growth. Atmospheric and oceanographic phenomena operating at ocean-basin scales seem to be important correlates of the processes driving growth in local coastal habitats. Conversely, terrestrial factors influencing precipitation and river runoff were positively correlated with the growth of juveniles in estuaries. Our results show that the impacts of climate change on these two life history stages are likely to be different, with implications for resilience and management of populations.

Coral luminescence identifies the Pacific Decadal Oscillation as a primary driver of river runoff variability impacting the southern Great Barrier Reef.

The Pacific Decadal Oscillation (PDO) is a large-scale climatic phenomenon modulating ocean-atmosphere variability on decadal time scales. While precipitation and river flow variability in the Great Barrier Reef (GBR) catchments are sensitive to PDO phases, the extent to which the PDO influences coral reefs is poorly understood. Here, six Porites coral cores were used to produce a composite record of coral luminescence variability (runoff proxy) and identify drivers of terrestrial influence on the Keppel reefs, southern GBR. We found that coral skeletal luminescence effectively captured seasonal, inter-annual and decadal variability of river discharge and rainfall from the Fitzroy River catchment. Most importantly, although the influence of El Niño-Southern Oscillation (ENSO) events was evident in the luminescence records, the variability in the coral luminescence composite record was significantly explained by the PDO. Negative luminescence anomalies (reduced runoff) were associated with El Niño years during positive PDO phases while positive luminescence anomalies (increased runoff) coincided with strong/moderate La Niña years during negative PDO phases. This study provides clear evidence that not only ENSO but also the PDO have significantly affected runoff regimes at the Keppel reefs for at least a century, and suggests that upcoming hydrological disturbances and ecological responses in the southern GBR region will be mediated by the future evolution of these sources of climate variability.

Human deforestation outweighs future climate change impacts of sedimentation on coral reefs.

Near-shore coral reef systems are experiencing increased sediment supply due to conversion of forests to other land uses. Counteracting increased sediment loads requires an understanding of the relationship between forest cover and sediment supply, and how this relationship might change in the future. Here we study this relationship by simulating river flow and sediment supply in four watersheds that are adjacent to Madagascar's major coral reef ecosystems for a range of future climate change projections and land-use change scenarios. We show that by 2090, all four watersheds are predicted to experience temperature increases and/or precipitation declines that, when combined, result in decreases in river flow and sediment load. However, these climate change-driven declines are outweighed by the impact of deforestation. Consequently, our analyses suggest that regional land-use management is more important than mediating climate change for influencing sedimentation of Malagasy coral reefs.

Linking coral river runoff proxies with climate variability, hydrology and land-use in Madagascar catchments.

Understanding the linkages between coastal watersheds and adjacent coral reefs is expected to lead to better coral reef conservation strategies. Our study aims to examine the main predictors of environmental proxies recorded in near shore corals and therefore how linked near shore reefs are to the catchment physical processes. To achieve these, we developed models to simulate hydrology of two watersheds in Madagascar. We examined relationships between environmental proxies derived from massive Porites spp. coral cores (spectral luminescence and barium/calcium ratios), and corresponding time-series (1950-2006) data of hydrology, climate, land use and human population growth. Results suggest regional differences in the main environmental drivers of reef sedimentation: on annual time-scales, precipitation, river flow and sediment load explained the variability in coral proxies of river discharge for the northeast region, while El Niño-Southern Oscillation (ENSO) and temperature (air and sea surface) were the best predictors in the southwest region.