Extreme spatial heterogeneity in carbonate accretion potential on a Caribbean fringing reef linked to local human disturbance gradients

The capacity of coral reefs to maintain their structurally complex frameworks and to retain the potential for vertical accretion is vitally important to the persistence of their ecological functioning and the ecosystem services they sustain. However, datasets to support detailed along‐coast assessments of framework production rates and accretion potential do not presently exist. Here, we estimate, based on gross bioaccretion and bioerosion measures, the carbonate budgets and resultant estimated accretion rates (EAR) of the shallow reef zone of leeward Bonaire – between 5 and 12 m depth – at unique fine spatial resolution along this coast (115 sites). Whilst the fringing reef of Bonaire is often reported to be in a better ecological condition than most sites throughout the wider Caribbean region, our data show that the carbonate budgets of the reefs and derived EAR varied considerably across this ~58 km long fringing reef complex. Some areas, in particular the marine reserves, were indeed still dominated by structurally complex coral communities with high net carbonate production (>10 kg CaCO₃ m^?2 year^?1), high live coral cover and complex structural topography. The majority of the studied sites, however, were defined by relatively low budget states (<2 kg CaCO₃ m^?2 year^?1) or were in a state of net erosion. These data highlight the marked spatial heterogeneity that can occur in budget states, and thus in reef accretion potential, even between quite closely spaced areas of individual reef complexes. This heterogeneity is linked strongly to the degree of localized land‐based impacts along the coast, and resultant differences in the abundance of reef framework building coral species. The major impact of this variability is that those sections of reef defined by low‐accretion rates will have limited capacity to maintain their structural integrity and to keep pace with current projections of climate change induced sea‐level rise (SLR), thus posing a threat to reef functioning and biodiversity, potentially leading to trophic cascades. Since many Caribbean reefs are more severely degraded than those found around Bonaire, it is to be expected that the findings presented here are rather the rule than the exception, but the study also highlights the need for similar high spatial resolution (along‐coast) assessments of budget states and accretion rates to meaningfully explore increasing coastal risk at the country level. The findings also more generally underline the significance of reducing local anthropogenic disturbance and restoring framework building coral assemblages. Appropriately focussed local preservation efforts may aid in averting future large‐scale above reef water depth increases on Caribbean coral reefs and will limit the social and economic implications associated with the loss of reef goods and services.     

Holocene coral reef accretion in Hawaii: a function of wave exposure and sea level history

 In the high Hawaiian Islands, significant accretion due to coral reef growth is limited by wave exposure and sea level. Holocene coral growth and reef accretion was measured at four stations off Oahu, Hawaii, chosen along a gradient in wave energy from minimum to maximum exposures. The results show that coral growth of living colonies (linear extension) at optimal depths is comparable at all stations (7.7–10.1 mm/y), but significant reef accretion occurs only at wave sheltered stations. At wave sheltered stations in Hanauma Bay and Kaneohe Bay, rates of long term reef accretion are about 2.0 mm/y. At wave exposed stations, off Mamala Bay and Sunset Beach, reef accretion rates are virtually zero in both shallow (1 m) and deeper (optimal) depths (12 m). At wave sheltered stations, such as Kaneohe Bay and Hanauma Bay, Holocene reef accretion is on the order of 10–15 m thick. At wave exposed stations, Holocene accretion is represented by only a thin veneer of living corals resting on antecedent Pleistocene limestone foundations. Modern coral communities in wave exposed environments undergo constant turnover associated with mortality and recruitment or re-growth of fragmented colonies and are rarely thicker than a single living colony. Breakage, scour, and abrasion of living corals during high wave events appears to be the major source of mortality and ultimately limits accretion to wave sheltered environments. Depth is particularly important as a modulator of wave energy. The lack of coral reef accretion along shallow open ocean coastlines may explain the absence of mature barrier reefs in the high Hawaiian Islands. Accepted: 14 May 1998     

Ecosystem‐based management of coral reefs under climate change

Coral reefs provide food and livelihoods for hundreds of millions of people as well as harbour some of the highest regions of biodiversity in the ocean. However, overexploitation, land‐use change and other local anthropogenic threats to coral reefs have left many degraded. Additionally, coral reefs are faced with the dual emerging threats of ocean warming and acidification due to rising CO₂ emissions, with dire predictions that they will not survive the century. This review evaluates the impacts of climate change on coral reef organisms, communities and ecosystems, focusing on the interactions between climate change factors and local anthropogenic stressors. It then explores the shortcomings of existing management and the move towards ecosystem‐based management and resilience thinking, before highlighting the need for climate change‐ready marine protected areas (MPAs), reduction in local anthropogenic stressors, novel approaches such as human‐assisted evolution and the importance of sustainable socialecological systems. It concludes that designation of climate change‐ready MPAs, integrated with other management strategies involving stakeholders and participation at multiple scales such as marine spatial planning, will be required to maximise coral reef resilience under climate change. However, efforts to reduce carbon emissions are critical if the long‐term efficacy of local management actions is to be maintained and coral reefs are to survive.     

Global change and coral reefs: impacts on reefs, economies and human cultures

Coral reefs have reconstituted themselves after previous large sea-level variations, and climate changes. For the past 6000 years of unusually stable sea-level, reefs have grown without serious interruptions. During recent decades, however, new stresses threaten localized devastation of many reefs. A new period of global climate change is occurring, stimulated by anthropogenic increases in greenhouse gases. Coral reefs will cope well with predicted sea-level rises of 4.5 cm per decade, but reef islands will not. Higher sea levels will provide corals with greater room for growth across reef flats, but there are no foreseeable mechanisms for reef island growth to keep pace with sea-level rise, therefore many low islands may ultimately become uninhabitable. Climate change will introduce localized variations in weather patterns, but changes to individual reefs cannot be predicted. Reefs on average should cope well with regional climate change, as they have coped with similar previous fluctuations. Air temperature increases of 0.2–0.3 °C/decade will induce slower increases in sea-surface temperatures, which may cause localized, or regional increases in coral bleaching. Changes in rainfall will impact on reefs near land masses. Likewise, increased storms and variations in El Nino Southern Oscillation (ENSO) may stress some reefs, but not others. The greatest impact of climate change will be a synergistic enhancement of direct anthropogenic stresses (excessive sediment and pollution from the land; over-fishing, especially via destructive methods; mining of coral rock and sand; and engineering modifications), which currently cause most damage to coral reefs. Many of the world's reefs have been degraded and more will be damaged as anthropogenic impacts increase under the ‘demophoric’ increases in population (demos) and economic (phoric) activity. This biotic and habitat loss will result in severe economic and social losses. Reefs, however, have considerable recovery powers and losses can be minimized by effective management of direct human impacts and reducing indirect threats of global climate change.     

Large‐scale coral reef rehabilitation after blast fishing in Indonesia

The severely degraded condition of many coral reefs worldwide calls for active interventions to rehabilitate their physical and biological structure and function, in addition to effective management of fisheries and no‐take reserves. Rehabilitation efforts to stabilize reef substratum sufficiently to support coral growth have been limited in size. We documented a large coral reef rehabilitation in Indonesia aiming to restore ecosystem functions by increasing live coral cover on a reef severely damaged by blast fishing and coral mining. The project deployed small, modular, open structures to stabilize rubble and to support transplanted coral fragments. Between 2013 to 2015, approximately 11,000 structures covering 7,000 m² were deployed over 2 ha of a reef at a cost of US$174,000. Live coral cover on the structures increased from less than 10% initially to greater than 60% depending on depth, deployment date and location, and disturbances. The mean live coral cover in the rehabilitation area in October 2017 was higher than reported for reefs in many other areas in the Coral Triangle, including marine protected areas, but lower than in the no‐take reference reef. At least 42 coral species were observed growing on the structures. Surprisingly, during the massive coral bleaching in other regions during the 2014–2016 El Niño–Southern Oscillation event, bleaching in the rehabilitation area was less than 5% cover despite warm water (≥30°C). This project demonstrates that coral rehabilitation is achievable over large scales where coral reefs have been severely damaged and are under continuous anthropogenic disturbances in warming waters.     

Millennial-scale change in the structure of a Caribbean reef ecosystem and the role of human and natural disturbance

Caribbean coral reefs have transformed into algal-dominated habitats over the past half-century, but the role of specific anthropogenic drivers is unresolved due to the lack of ecosystem-level data predating human disturbance. To better understand the extent and causes of long-term Caribbean reef declines, we produced a continuous 3000-yr record of the ecosystem state of three reefs in Bocas del Toro, Caribbean Panama. From fossils and sediments obtained from reef matrix cores, we tracked changes in reef accretion rates and the taxonomic and functional group composition of fish, coral, urchin, bivalve and benthic foraminifera. This dataset provided a comprehensive picture of reef community and environmental change. At all sites, reefs shifted from systems with greater relative abundance of herbivorous fish, epifaunal suspension feeding bivalves and Diadema urchins to systems with greater relative abundance of micropredator fish, infaunal bivalves and Echinometra urchins. These transitions were initiated a millennium ago at two less-degraded reefs fringing offshore islands and ~250 yr ago at a degraded patch reef near the continental coast. Ecosystem shifts were accompanied by a decline in reef accretion rates, and at the patch reef, a decline in water quality since the 18th century. Within all cores, synchronous increases in infaunal bivalves and declines in herbivorous fish regardless of water quality suggest a loss of hard substrate and increasingly hypoxic sediment conditions related to herbivore loss. While the early timing of ecosystem transitions at the fringing reefs implicates large-scale hydrological change, the more recent timing of change and loss of water quality at the patch reef implicates terrigenous runoff from land-clearing. Our whole-ecosystem reconstruction reveals that reef ecosystem deterioration appears to follow a predictable trajectory whether driven by natural or anthropogenic disturbances and that historical local human activities have quickly unraveled reefs at a scale similar to longer-term natural environmental change.     

Regional‐scale dominance of non‐framework building corals on Caribbean reefs affects carbonate production and future reef growth

Coral cover on Caribbean reefs has declined rapidly since the early 1980's. Diseases have been a major driver, decimating communities of framework building Acropora and Orbicella coral species, and reportedly leading to the emergence of novel coral assemblages often dominated by domed and plating species of the genera Agaricia, Porites and Siderastrea. These corals were not historically important Caribbean framework builders, and typically have much smaller stature and lower calcification rates, fuelling concerns over reef carbonate production and growth potential. Using data from 75 reefs from across the Caribbean we quantify: (i) the magnitude of non‐framework building coral dominance throughout the region and (ii) the contribution of these corals to contemporary carbonate production. Our data show that live coral cover averages 18.2% across our sites and coral carbonate production 4.1 kg CaCO₃ m^?2 yr^?1. However, non‐framework building coral species dominate and are major carbonate producers at a high proportion of sites; they are more abundant than Acropora and Orbicella at 73% of sites; contribute an average 68% of the carbonate produced; and produce more than half the carbonate at 79% of sites. Coral cover and carbonate production rate are strongly correlated but, as relative abundance of non‐framework building corals increases, average carbonate production rates decline. Consequently, the use of coral cover as a predictor of carbonate budget status, without species level production rate data, needs to be treated with caution. Our findings provide compelling evidence for the Caribbean‐wide dominance of non‐framework building coral taxa, and that these species are now major regional carbonate producers. However, because these species typically have lower calcification rates, continued transitions to states dominated by non‐framework building coral species will further reduce carbonate production rates below ‘predecline’ levels, resulting in shifts towards negative carbonate budget states and reducing reef growth potential.     

Holocene sea level changes and reef development in the southwestern Indian Ocean

 The sedimentological and chronological study of Holocene reef sequences recovered in drill cores through modern reefs of Mauritius, Réunion Island and Mayotte allows the reconstruction of sea level changes and reef growth patterns during the Holocene. The branching-coral facies systematically predominates over coral head facies throughout the Holocene reef sequences, and Acropora is the main frame builder among the branching forms. The reconstructed sea level curves, based both on identification of coral assemblages and on radiometric U/Th ages, are characterized by a rapid rise between 10 and 7.5 ky BP, followed by a clear inflection between 7.5 and 7 ky BP. The stabilization of sea level at its present level occurred between 2000 and 3000 years ago, probably without a higher sea level stand. Rates of vertical reef accretion range between 0.9 and 7 mm. y^-1. In Mauritius, and also probably in Réunion Island, the reef first tracked, then caught-up to sea level to reach an equilibrium position (“catch-up” growth), while the barrier reef margin off Mayotte has been able to keep pace with rising sea level (“keep-up” growth). Accepted: 1 March 1997     

Evidence for and against the existence of alternate attractors on coral reefs

Synthesis Coral reefs are widely thought to exhibit multiple attractors which have profound implications for people that depend on them. If reefs become ‘stuck’ within a self‐reinforcing state dominated by seaweed, it becomes disproportionately difficult and expensive for managers to shift the system back towards its natural, productive, coral state. The existence of multiple attractors is controversial. We assess various forms of evidence and conclude that there remains no incontrovertible proof of multiple attractors on reefs. However, the most compelling evidence, which combines ecological models and field data, is far more consistent with multiple attractors than the competing hypothesis of only a single, coral attractor. Managers should exercise caution and assume that degraded reefs can become stuck there. Testing for the existence of alternate attractors in ecosystems that possess slow dynamics and frequent pulse perturbation is exceptionally challenging. Coral reefs typify such conditions and the existence of alternate attractors is controversial. We analyse different forms of evidence and assess whether they support or challenge the existence of multiple attractors on Caribbean reefs, many of which have shown profound phase shifts in community structure from coral to algal dominance. Field studies alone provide no insight into multiple attractors because the non‐equilibrial nature of reef dynamics prevents equilibria from being observed. Statistical models risk failing to sample the parameter space in which multiple attractors occur, and have failed to account for the confounding effects of heterogeneous environments, anthropogenic drivers (e.g. fishing), and major disturbances (e.g. hurricanes). Simple and complex models all find multiple attractors over some – though not all – regions of a system driver (fishing). Tests of model predictions with field data closely match theory of alternate attractors but a forward‐leaning monotonic curve with only a single attractor can also be fitted to these data. Deeper consideration of the assumptions of this monotonic relationship reveal significant ecological problems which disappear under a model of multiple attractors. To date, there is no evidence against the existence of multiple attractors on Caribbean reefs and while there remains no definitive proof, the balance of evidence and ecological reasoning favours their existence. Theory predicts that Caribbean reefs do not exhibit alternate attractors in their natural state but that disease‐induced loss of two key functional groups has generated bistability. Whether alternate attractors becomes a persistent element of reef dynamics or a brief moment in their geological history will depend, in part, on the ability of functional groups to recover and the impacts of climate change and ocean acidification on coral growth and mortality.     

Region‐wide temporal and spatial variation in Caribbean reef architecture: is coral cover the whole story?

The architectural complexity of coral reefs is largely generated by reef‐building corals, yet the effects of current regional‐scale declines in coral cover on reef complexity are poorly understood. In particular, both the extent to which declines in coral cover lead to declines in complexity and the length of time it takes for reefs to collapse following coral mortality are unknown. Here we assess the extent of temporal and spatial covariation between coral cover and reef architectural complexity using a Caribbean‐wide dataset of temporally replicated estimates spanning four decades. Both coral cover and architectural complexity have declined rapidly over time, with little evidence of a time‐lag. However, annual rates of change in coral cover and complexity do not covary, and levels of complexity vary greatly among reefs with similar coral cover. These findings suggest that the stressors influencing Caribbean reefs are sufficiently severe and widespread to produce similar regional‐scale declines in coral cover and reef complexity, even though reef architectural complexity is not a direct function of coral cover at local scales. Given that architectural complexity is not a simple function of coral cover, it is important that conservation monitoring and restoration give due consideration to both architecture and coral cover. This will help ensure that the ecosystem services supported by architectural complexity, such as nutrient recycling, dissipation of wave energy, fish production and diversity, are maintained and enhanced.     

Is the coral‐algae symbiosis really ‘mutually beneficial’ for the partners?

The consideration of ‘mutual benefits’ and partner cooperation have long been the accepted standpoint from which to draw inference about the onset, maintenance and breakdown of the coral‐algae endosymbiosis. In this paper, I review recent research into the climate‐induced breakdown of this important symbiosis (namely ‘coral bleaching’) that challenges the validity of this long‐standing belief. Indeed, I introduce a more parsimonious explanation, in which the coral host exerts a ‘controlled parasitism’ over its algal symbionts that is akin to an enforced domestication arrangement. Far from being pathogenic, a range of well‐established cellular processes are reviewed that support the role of the coral host as an active ‘farmer’ of the energy‐rich photoassimilates from its captive symbionts. Importantly, this new paradigm reposes the deleterious bleaching response in terms of an envelope of environmental conditions in which the exploitative and captive measures of the coral host are severely restricted. The ramification of this new paradigm for developing management strategies that may assist the evolution of bleaching resistance in corals is discussed.     

Escaping the heat: range shifts of reef coral taxa in coastal Western Australia

One of the most critical challenges facing ecologists today is to understand the changing geographic distribution of species in response to current and predicted global warming. Coastal Western Australia is a natural laboratory in which to assess the effect of climate change on reef coral communities over a temporal scale unavailable to studies conducted solely on modern communities. Reef corals composing Late Pleistocene reef assemblages exposed at five distinct localities along the west Australian coast were censused and the results compared with coral occurrence data published for the modern reefs offshore of each locality. The resulting comparative data set comprises modern and Late Pleistocene reef coral communities occurring over approximately 12° of latitude. For the modern reefs this gradient includes the zone of overlap between the Dampierian and Flindersian Provinces. Modern reef coral communities show a pronounced gradient in coral composition over the latitudinal range encompassed by the study, while the gradient in community composition is not as strong for Pleistocene communities. Tropical‐adapted taxa contracted their ranges north since Late Pleistocene time, emplacing two biogeographic provinces in a region in which a single province had existed previously. Beta diversity values for adjacent communities also reflect this change. Modern reefs show a distinct peak in beta diversity in the middle of the region; the peak is not matched by Pleistocene reefs. Beta diversity is correlated with distance only for comparisons between modern reefs in the north and the fossil assemblages, further supporting change in distribution of the biogeographic provinces in the study area. Coral taxa present in modern communities clearly expanded and contracted their geographic ranges in response to climate change. Those taxa that distinguish Pleistocene from modern reefs are predicted to migrate south in response to future climate change, and potentially persist in ‘temperature refugia’ as tropical reef communities farther north decline.     

Holocene reef evolution in a macrotidal setting: Buccaneer Archipelago,Kimberley Bioregion,Northwest Australia

This study uses information derived from cores to describe the Holocene accretion history of coral reefs in the macrotidal (up to 11 m tidal range) Buccaneer Archipelago of the southern Kimberley coast, Western Australia. The internal architecture of all cored reefs is broadly similar, constituting well-preserved detrital coral fragments, predominantly branching Acropora, in a poorly sorted sandy mud matrix. However, once the reefs reach sea level, they diverge into two types: low intertidal reefs that maintain their detrital character and develop relatively narrow, horizontal or gently sloping reef flats at approximately mean low water spring, and high intertidal reefs that develop broad coralline algal-dominated reef flats at elevations between mean low water neap and mean high water neap. The high intertidal reefs develop where strong, ebb-dominated, tidal asymmetry retains seawater over the low tide and allows continued accretion. Both reef types are ultimately constrained by sea level but differ in elevation by 3–4 m.     

Coral reef growth in an era of rapidly rising sea level: predictions and suggestions for long-term research

Coral reef growth is intimately linked to sea level. It has been postulated that over the next century, sea level will rise at a probable average rate of 15 mm/year, in response to fossil fuel emissions, heating, and melting of the Antarctic ice cap. This predicted rate of sea level rise is five times the present modal rate of vertical accretion on coral reef flats and 50% greater than the maximum vertical accretion rates apparently attained by coral reefs. We use these predictions and observations to offer the following hypothesis for reef growth over the next century. The vertical accretion rates of protected reef flats will accelerate from the present modal rate up to the maximum rate, in response to the more rapidly rising sea level. This more rapid vertical accretion rate will be insufficient to keep up with sea level rise, if present predictions prove to be correct. Less protected reef flats will slow their rate of growth as they become inundated and subjected to erosion by progressively larger waves. This projected sea level rise and postulated reef response will provide an opportunity for long-term studies of the response of coral reef systems to a predictable and measurable forcing function. If the scientific benefits from this uncontrolled global experiment are to be maximized, it will be necessary to establish a permanent international coordinating body to assist with the identification and preservation of long-term study sites and to provide guidelines for baseline data surverys, methods selection and comparison, and other procedures and decisions.     

Life histories predict coral community disassembly under multiple stressors

Climate change is reshaping biological communities against a background of existing human pressure. Evaluating the impacts of multiple stressors on community dynamics can be particularly challenging in species‐rich ecosystems, such as coral reefs. Here, we investigate whether life‐history strategies and cotolerance to different stressors can predict community responses to fishing and temperature‐driven bleaching using a 20‐year time series of coral assemblages in Kenya. We found that the initial life‐history composition of coral taxa largely determined the impacts of bleaching and coral loss. Prior to the 1998 bleaching event, coral assemblages within no‐take marine reserves were composed of three distinct life histories – competitive, stress‐tolerant and weedy– and exhibited strong declines following bleaching with limited subsequent recovery. In contrast, fished reefs had lower coral cover, fewer genera and were composed of stress‐tolerant and weedy corals that were less affected by bleaching over the long term. Despite these general patterns, we found limited evidence for cotolerance as coral genera and life histories were variable in their sensitivities to fishing and bleaching. Overall, fishing and bleaching have reduced coral diversity and led to altered coral communities of ‘survivor’ species with stress‐tolerant and weedy life histories. Our findings are consistent with expectations that climate change interacting with existing human pressure will result in the loss of coral diversity and critical reef habitat.     

Chromogenic and fluorescent ‘turn‐on’ chemodosimeter for fluoride based on a F–‐triggered cascade reaction

We developed a new chromogenic and fluorescent ‘turn‐on’ chemodosimeter 1 based on a F^–‐triggered cascade reaction. This system displayed significant changes in UV/vis absorption and fluorescence emission intensities selectively for F^– over other anions in a mixture of CH₃CN/H₂O (95 : 5, v/v) and in acetonitrile. Copyright © 2014 John Wiley & Sons, Ltd.     

Coral reef conservation in Bali in light of international best practice,a literature review

Bali, Indonesia sits within the coral triangle and is internationally recognised for its high coral reef diversity. The health of Bali’s marine ecosystems has declined in recent decades, and this is thought to be due to threats from climate change, destructive fishing practices, pollution, outbreaks coral eating invertebrates, coral disease and unsustainable tourism. As a response, multiple conservation strategies have been introduced by the island’s communities, non-government organisations and governments, with the aim of preventing further decline, as well as restoring already degraded coral reefs. This literature review provides an in-depth analysis of the tools used to conserve Bali’s coral reefs, and compares them to those used in other countries. In light of international ‘best practice’ in coral reef conservation, this review makes suggestions on how Bali could better conserve its coral reef ecosystems. These include (1) increasing its designation of official Marine Protected Areas (MPAS) and strengthening management of existing ones, (2) creating an MPA network, (3) substantially reducing marine plastic pollution, (4) continuing artificial reef construction in degraded habitats, (5) continuing to develop Bali as an ecotourism destination, (6) increasing engagement in global science to inform marine conservation decision-making, and (7) developing more marine monitoring programmes.     

Caribbean mesophotic coral ecosystems are unlikely climate change refugia

Deeper coral reefs experience reduced temperatures and light and are often shielded from localized anthropogenic stressors such as pollution and fishing. The deep reef refugia hypothesis posits that light‐dependent stony coral species at deeper depths are buffered from thermal stress and will avoid bleaching‐related mass mortalities caused by increasing sea surface temperatures under climate change. This hypothesis has not been tested because data collection on deeper coral reefs is difficult. Here we show that deeper (mesophotic) reefs, 30–75 m depth, in the Caribbean are not refugia because they have lower bleaching threshold temperatures than shallow reefs. Over two thermal stress events, mesophotic reef bleaching was driven by a bleaching threshold that declines 0.26 °C every +10 m depth. Thus, the main premise of the deep reef refugia hypothesis that cooler environments are protective is incorrect; any increase in temperatures above the local mean warmest conditions can lead to thermal stress and bleaching. Thus, relatively cooler temperatures can no longer be considered a de facto refugium for corals and it is likely that many deeper coral reefs are as vulnerable to climate change as shallow water reefs.