Trace metal pollution gradients in a tropical seagrass ecosystem.

Trace metals are one of the most serious pollutants in tropical seagrass meadows given their persistence and toxicity. Whereas quantity is frequently measured, there is no information on the spatial extent of metal pollution in these systems. Here, we use an island in Indonesia (Barang Lompo) as a model system to study the impact radius of two major and eight trace metals in sediment and seagrass leaves. We provide evidence for exponential decay in both the metal pollution index and concentrations of most metals with increasing distance from the island (k = -0.01 to -0.08 m-1). Consequently, there is an impact radius of approximately 100 m around the island. The comparative analysis of both seagrass species further revealed interspecific differences in metal loads. This study highlights the importance of assessing the spatial extent of metal pollution in addition to its quantity.

Sustainable reference points for multispecies coral reef fisheries.

Sustainably managing fisheries requires regular and reliable evaluation of stock status. However, most multispecies reef fisheries around the globe tend to lack research and monitoring capacity, preventing the estimation of sustainable reference points against which stocks can be assessed. Here, combining fish biomass data for >2000 coral reefs, we estimate site-specific sustainable reference points for coral reef fisheries and use these and available catch estimates to assess the status of global coral reef fish stocks. We reveal that >50% of sites and jurisdictions with available information have stocks of conservation concern, having failed at least one fisheries sustainability benchmark. We quantify the trade-offs between biodiversity, fish length, and ecosystem functions relative to key benchmarks and highlight the ecological benefits of increasing sustainability. Our approach yields multispecies sustainable reference points for coral reef fisheries using environmental conditions, a promising means for enhancing the sustainability of the world's coral reef fisheries.

Temperature, species identity and morphological traits predict carbonate excretion and mineralogy in tropical reef fishes.

Anthropogenic pressures are restructuring coral reefs globally. Sound predictions of the expected changes in key reef functions require adequate knowledge of their drivers. Here we investigate the determinants of a poorly-studied yet relevant biogeochemical function sustained by marine bony fishes: the excretion of intestinal carbonates. Compiling carbonate excretion rates and mineralogical composition from 382 individual coral reef fishes (85 species and 35 families), we identify the environmental factors and fish traits that predict them. We find that body mass and relative intestinal length (RIL) are the strongest predictors of carbonate excretion. Larger fishes and those with longer intestines excrete disproportionately less carbonate per unit mass than smaller fishes and those with shorter intestines. The mineralogical composition of excreted carbonates is highly conserved within families, but also controlled by RIL and temperature. These results fundamentally advance our understanding of the role of fishes in inorganic carbon cycling and how this contribution will change as community composition shifts under increasing anthropogenic pressures.

First insights into coral recruit and juvenile abundances at remote Aldabra Atoll, Seychelles.

Coral recruitment and successive growth are essential for post-disturbance reef recovery. As coral recruit and juvenile abundances vary across locations and under different environmental regimes, their assessment at remote, undisturbed reefs improves our understanding of early life stage dynamics of corals. Here, we first explored changes in coral juvenile abundance across three locations (lagoon, seaward west and east) at remote Aldabra Atoll (Seychelles) between 2015 and 2019, which spanned the 2015/16 global coral bleaching event. Secondly, we measured variation in coral recruit abundance on settlement tiles from two sites (lagoon, seaward reef) during August 2018-August 2019. Juvenile abundance decreased from 14.1 ± 1.2 to 7.4 ± 0.5 colonies m-2 (mean ± SE) during 2015-2016 and increased to 22.4 ± 1.2 colonies m-2 during 2016-2019. Whilst juvenile abundance increased two- to three-fold at the lagoonal and seaward western sites during 2016-2018 (from 7.7-8.3 to 17.3-24.7 colonies m-2), increases at the seaward eastern sites occurred later (2018-2019; from 5.8-6.9 to 16.6-24.1 colonies m-2). The composition of coral recruits on settlement tiles was dominated by Pocilloporidae (64-92% of all recruits), and recruit abundance was 7- to 47-fold higher inside than outside the lagoon. Recruit abundance was highest in October-December 2018 (2164 ± 453 recruits m-2) and lowest in June-August 2019 (240 ± 98 recruits m-2). As Acroporid recruit abundance corresponded to this trend, the results suggest that broadcast spawning occurred during October-December, when water temperature increased from 26 to 29°C. This study provides the first published record on coral recruit abundance in the Seychelles Outer Islands, indicates a rapid (2-3 years) increase of juvenile corals following a bleaching event, and provides crucial baseline data for future research on reef resilience and connectivity within the region.

A survey of current trends and suggested future directions in coral transplantation for reef restoration.

Coral transplantation has been used in reef restoration for several decades, but information on the type of projects, their scope, scale, and success is mostly limited to published scientific studies and technical reports. Many practitioners do not have the capacity to share their progress in peer-reviewed literature, yet likely have a wealth of information to share on how to improve the efficiency of transplantation efforts. In order to incorporate non-published data on coral transplantation projects and gain an overview of the general features of these projects, we conducted an initial systematic online survey of projects run by various practitioners. Surveyed projects (n = 50) covered most of the tropical belt and ranged in size from a few hundred transplanted corals to >5000 transplants. The most frequent source of coral fragments were corals already broken from some previous impact ("corals of opportunity"; 58% of projects), followed by fragments stored in different types of aquaculture systems (42% of projects). The use of sexual reproduction was very limited. Fast-growing, branching corals were used in 96% of projects, being by far the most common transplanted growth form. About half of the projects mentioned undertaking maintenance of the transplantation plots. The majority of projects undertook subsequent monitoring (80%), yet the available data indicates that duration of monitoring efforts was not adequate to evaluate long-term success. The findings underline that while some general principles for successful coral restoration projects are reasonably well established, others need to be mainstreamed better in order to improve the effectiveness of coral transplantation for reef restoration. This relates in particular to sustainable funding, adequate site assessment, and long-term monitoring using established protocols. Additional information is needed to better understand and address potential challenges with regards to the sourcing of transplants and use of slow-growing species. A better integration of practitioners is necessary to improve the understanding of coral transplantation effectiveness. The results underline a need to develop and use monitoring protocols that allow gauging and comparing the effectiveness of coral transplantation among various projects, as well as for accessible platform(s) to allow the exchange of experiences made in different projects. Regular surveys of restoration projects are recommended to collate and share information among practitioners. We provide a number of recommendations for items to include in future surveys.

Meeting fisheries, ecosystem function, and biodiversity goals in a human-dominated world.

The worldwide decline of coral reefs necessitates targeting management solutions that can sustain reefs and the livelihoods of the people who depend on them. However, little is known about the context in which different reef management tools can help to achieve multiple social and ecological goals. Because of nonlinearities in the likelihood of achieving combined fisheries, ecological function, and biodiversity goals along a gradient of human pressure, relatively small changes in the context in which management is implemented could have substantial impacts on whether these goals are likely to be met. Critically, management can provide substantial conservation benefits to most reefs for fisheries and ecological function, but not biodiversity goals, given their degraded state and the levels of human pressure they face.

Coral restoration - A systematic review of current methods, successes, failures and future directions.

Coral reef ecosystems have suffered an unprecedented loss of habitat-forming hard corals in recent decades. While marine conservation has historically focused on passive habitat protection, demand for and interest in active restoration has been growing in recent decades. However, a disconnect between coral restoration practitioners, coral reef managers and scientists has resulted in a disjointed field where it is difficult to gain an overview of existing knowledge. To address this, we aimed to synthesise the available knowledge in a comprehensive global review of coral restoration methods, incorporating data from the peer-reviewed scientific literature, complemented with grey literature and through a survey of coral restoration practitioners. We found that coral restoration case studies are dominated by short-term projects, with 60% of all projects reporting less than 18 months of monitoring of the restored sites. Similarly, most projects are relatively small in spatial scale, with a median size of restored area of 100 m2. A diverse range of species are represented in the dataset, with 229 different species from 72 coral genera. Overall, coral restoration projects focused primarily on fast-growing branching corals (59% of studies), and report survival between 60 and 70%. To date, the relatively young field of coral restoration has been plagued by similar 'growing pains' as ecological restoration in other ecosystems. These include 1) a lack of clear and achievable objectives, 2) a lack of appropriate and standardised monitoring and reporting and, 3) poorly designed projects in relation to stated objectives. Mitigating these will be crucial to successfully scale up projects, and to retain public trust in restoration as a tool for resilience based management. Finally, while it is clear that practitioners have developed effective methods to successfully grow corals at small scales, it is critical not to view restoration as a replacement for meaningful action on climate change.

Herbivorous fish rise as a destructive fishing practice falls in an Indonesian marine national park.

Securing ecosystem functions is challenging, yet common priority in conservation efforts. While marine parks aim to meet this challenge by regulating fishing through zoning plans, their effectiveness hinges on compliance levels and may respond to changes in fishing practices. Here we use a speciose assemblage of nominally herbivorous reef fish in Karimunjawa National Park (zoned since 1989) to investigate whether areas subject to a restrictive management regime sustained higher biomass over seven years compared to areas where moderate and permissive regulations apply. Using a trait-based approach we characterize the functional space of the entire species pool and ask whether changes in biomass translate into changes in functional structure. We track changes in predator biomass, benthic community structure, and fishing practices that could influence herbivore trajectories. Overall herbivore biomass doubled in 2012 compared to 2006-2009 and remained high in 2013 across all management regimes. We found no evidence that this biomass build-up resulted from predator depletion or increased food availability but suggest it emerged in response to a park-wide cessation of fishing with large drive nets known as muroami. The biomass increase was accompanied by a modest increase in taxonomic richness and a slight decrease in community-scale rarity that did not alter functional redundancy levels. Subtle changes in both functional specialization and identity of assemblages emerged as generalist species with low intrinsic vulnerability to fishing recovered sooner than more vulnerable specialists. While this implies a recovery of mechanisms responsible for the grazing of algal turfs and detritus, restoring other facets of herbivory (e.g., macroalgal consumption) may require more time. An increase in the cost-benefit ratio per journey of muroami fishing facilitated a ban on muroami nets that met minimal resistance. Similar windows of opportunity may emerge elsewhere in which gear-based regulations can supplement zoning plans, especially when compliance is low. This does not advocate for implementing such regulations once a fishery has become unprofitable. Rather, it underlines their importance for breaking the cycle of resource depletion and low compliance to zoning, thus alleviating the resulting threats to food security and ecosystem integrity.

Positive association between epiphytes and competitiveness of the brown algal genus Lobophora against corals.

Observations of coral-algal competition can provide valuable information about the state of coral reef ecosystems. Here, we report contact rates and apparent competition states for six shallow lagoonal reefs in Fiji. A total of 81.4% of examined coral perimeters were found to be in contact with algae, with turf algae (54.7%) and macroalgae of the genus Lobophora (16.8%) representing the most frequently observed contacts. Turf algae competitiveness was low, with 21.8% of coral-turf contacts being won by the algae (i.e. overgrowth or bleaching of coral tissue). In contrast, Lobophora competitiveness against corals was high, with 62.5% of contacts being won by the alga. The presence of epiphytic algae on Lobophora was associated with significantly greater algal competitiveness against corals, with 75.8% and 21.1% of interactions recorded as algal wins in the presence and absence of epiphytes, respectively. Sedimentation rate, herbivorous fish biomass, and coral colony size did not have a significant effect on Lobophora-coral interactions. This research indicates a novel and important role of epiphytes in driving the outcome of coral-algal contacts.

Citizen science in data and resource-limited areas: A tool to detect long-term ecosystem changes.

Coral reefs are threatened by numerous global and local stressors. In the face of predicted large-scale coral degradation over the coming decades, the importance of long-term monitoring of stress-induced ecosystem changes has been widely recognised. In areas where sustained funding is unavailable, citizen science monitoring has the potential to be a powerful alternative to conventional monitoring programmes. In this study we used data collected by volunteers in Southeast Sulawesi (Indonesia), to demonstrate the potential of marine citizen science programmes to provide scientifically sound information necessary for detecting ecosystem changes in areas where no alternative data are available. Data were collected annually between 2002 and 2012 and consisted of percent benthic biotic and abiotic cover and fish counts. Analyses revealed long-term coral reef ecosystem change. We observed a continuous decline of hard coral, which in turn had a significant effect on the associated fishes, at community, family and species levels. We provide evidence of the importance of marine citizen science programmes in detecting long-term ecosystem change as an effective way of delivering conservation data to local government and national agencies. This is particularly true for areas where funding for monitoring is unavailable, resulting in an absence of ecological data. For citizen science data to contribute to ecological monitoring and local decision-making, the data collection protocols need to adhere to sound scientific standards, and protocols for data evaluation need to be available to local stakeholders. Here, we describe the monitoring design, data treatment and statistical analyses to be used as potential guidelines in future marine citizen science projects.

Community-wide scan identifies fish species associated with coral reef services across the Indo-Pacific.

Determining whether many functionally complementary species or only a subset of key species are necessary to maintain ecosystem functioning and services is a critical question in community ecology and biodiversity conservation. Identifying such key species remains challenging, especially in the tropics where many species co-occur and can potentially support the same or different processes. Here, we developed a new community-wide scan (CWS) approach, analogous to the genome-wide scan, to identify fish species that significantly contribute, beyond the socio-environmental and species richness effects, to the biomass and coral cover on Indo-Pacific reefs. We found that only a limited set of species (51 out of approx. 400, approx. 13%), belonging to various functional groups and evolutionary lineages, are strongly and positively associated with fish biomass and live coral cover. Many of these species have not previously been identified as functionally important, and thus may be involved in unknown, yet important, biological mechanisms that help sustain healthy and productive coral reefs. CWS has the potential to reveal species that are key to ecosystem functioning and services and to guide management strategies as well as new experiments to decipher underlying causal ecological processes.

Gravity of human impacts mediates coral reef conservation gains.

Coral reefs provide ecosystem goods and services for millions of people in the tropics, but reef conditions are declining worldwide. Effective solutions to the crisis facing coral reefs depend in part on understanding the context under which different types of conservation benefits can be maximized. Our global analysis of nearly 1,800 tropical reefs reveals how the intensity of human impacts in the surrounding seascape, measured as a function of human population size and accessibility to reefs ("gravity"), diminishes the effectiveness of marine reserves at sustaining reef fish biomass and the presence of top predators, even where compliance with reserve rules is high. Critically, fish biomass in high-compliance marine reserves located where human impacts were intensive tended to be less than a quarter that of reserves where human impacts were low. Similarly, the probability of encountering top predators on reefs with high human impacts was close to zero, even in high-compliance marine reserves. However, we find that the relative difference between openly fished sites and reserves (what we refer to as conservation gains) are highest for fish biomass (excluding predators) where human impacts are moderate and for top predators where human impacts are low. Our results illustrate critical ecological trade-offs in meeting key conservation objectives: reserves placed where there are moderate-to-high human impacts can provide substantial conservation gains for fish biomass, yet they are unlikely to support key ecosystem functions like higher-order predation, which is more prevalent in reserve locations with low human impacts.

Effects of sandfish (Holothuria scabra) removal on shallow-water sediments in Fiji.

Sea cucumbers play an important role in the recycling and remineralization of organic matter (OM) in reef sands through feeding, excretion, and bioturbation processes. Growing demand from Asian markets has driven the overexploitation of these animals globally. The implications of sea cucumber fisheries for shallow coastal ecosystems and their management remain poorly understood. To address this knowledge gap, the current study manipulated densities of Holothuria scabra within enclosures on a reef flat in Fiji, between August 2015 and February 2016, to study the effects of sea cucumber removal on sedimentary function as a biocatalytic filter system. Three treatments were investigated: (i) high density (350 g m-2 wet weight; ca. 15 individuals); (ii) natural density (60 g m-2; ca. 3 individuals); and (iii) exclusion (0 g m-2). Quantity of sediment reworked through ingestion by H. scabra, grain size distribution, O2 penetration depth, and sedimentary oxygen consumption (SOC) were quantified within each treatment. Findings revealed that the natural population of H. scabra at the study site can rework ca. 10,590 kg dry sediment 1,000 m-2 year-1; more than twice the turnover rate recorded for H. atra and Stichopus chloronotus. There was a shift towards finer fraction grains in the high treatment. In the exclusion treatment, the O2 penetration depth decreased by 63% following a 6 °C increase in water temperature over the course of two months, while in the high treatment no such change was observed. SOC rates increased ca. two-fold in the exclusion treatment within the first month, and were consistently higher than in the high treatment. These results suggest that the removal of sea cucumbers can reduce the capacity of sediments to buffer OM pulses, impeding the function and productivity of shallow coastal ecosystems.

Mutualistic damselfish induce higher photosynthetic rates in their host coral.

Coral reefs are amongst the most diverse ecosystems on Earth where complex inter-specific interactions are ubiquitous. An example of such interactions is the mutualistic relationship between damselfishes and branching corals in the Northern Red Sea, where the fish use corals as shelter and provide them with nutrients, enhance the flow between their branches, and protect them from predators. By enhancing the flow between the coral branches, the fish ventilate the coral's inner zone, mitigating hypoxic conditions that otherwise develop within that zone during the night. Here, we tested, for the first time, the effects of the damselfish Dascyllus marginatus on photosynthesis and respiration in its host coral Stylophora pistillata Laboratory experiments using an intermittent-flow respirometer showed that the presence of fish between the coral branches under light conditions augmented the coral's photosynthetic rate. No effect on the coral's respiration was found under dark conditions. When a fish was allowed to enter the inner zone of a dead coral skeleton, its respiration was higher than when it was in a live coral. Field observations indicated that damselfish were present between coral branches 18-34% of the time during daylight hours and at all times during the night. Considering the changes induced by the fish together with the proportion of time they were found between coral branches in the field, the effect of the fish amounted to an augmentation of 3-6% of the coral's daily photosynthesis. Our findings reveal a previously unknown positive contribution of coral-dwelling fish to their host's photosynthesis.

Abundance and physiology of dominant soft corals linked to water quality in Jakarta Bay, Indonesia.

Declining water quality is one of the main reasons of coral reef degradation in the Thousand Islands off the megacity Jakarta, Indonesia. Shifts in benthic community composition to higher soft coral abundances have been reported for many degraded reefs throughout the Indo-Pacific. However, it is not clear to what extent soft coral abundance and physiology are influenced by water quality. In this study, live benthic cover and water quality (i.e. dissolved inorganic nutrients (DIN), turbidity (NTU), and sedimentation) were assessed at three sites (< 20 km north of Jakarta) in Jakarta Bay (JB) and five sites along the outer Thousand Islands (20-60 km north of Jakarta). This was supplemented by measurements of photosynthetic yield and, for the first time, respiratory electron transport system (ETS) activity of two dominant soft coral genera, Sarcophyton spp. and Nephthea spp. Findings revealed highly eutrophic water conditions in JB compared to the outer Thousand Islands, with 44% higher DIN load (7.65 µM/L), 67% higher NTU (1.49 NTU) and 47% higher sedimentation rate (30.4 g m-2 d-1). Soft corals were the dominant type of coral cover within the bay (2.4% hard and 12.8% soft coral cover) compared to the outer Thousand Islands (28.3% hard and 6.9% soft coral cover). Soft coral abundances, photosynthetic yield, and ETS activity were highly correlated with key water quality parameters, particularly DIN and sedimentation rates. The findings suggest water quality controls the relative abundance and physiology of dominant soft corals in JB and may thus contribute to phase shifts from hard to soft coral dominance, highlighting the need to better manage water quality in order to prevent or reverse phase shifts.

Under pressure: Investigating marine resource-based livelihoods in Jakarta Bay and the Thousand Islands.

Jakarta Bay, next to the Jakarta Metropolitan Area with around 30 million inhabitants, is facing extreme pollution. Although local coral reefs are degraded and marine resources heavily exploited, they provide livelihoods for millions of people. This study investigates anthropogenic pressures on local fisheries resources and associated livelihoods. Questionnaire surveys were conducted in 15 coastal communities (10 coastal neighborhoods in Jakarta Bay on the mainland and 5 of the offshore Thousand Islands). The most economically valuable species were Caesio cuning (Redbelly yellowtail fusilier) on the islands and Rastrelliger kanagurta (Indian mackerel) on the mainland. Over 80% of all interviewed fishermen regarded the current state of marine resources as declining, mainly due to pollution and overexploitation. While perceptions of declining resources were equally high on the islands and the mainland, pollution was listed as the principal cause of degradation significantly more on the mainland. Findings are discussed in the context of coastal livelihood vulnerability.

Bright spots among the world's coral reefs.

Ongoing declines in the structure and function of the world's coral reefs require novel approaches to sustain these ecosystems and the millions of people who depend on them3. A presently unexplored approach that draws on theory and practice in human health and rural development is to systematically identify and learn from the 'outliers'­places where ecosystems are substantially better ('bright spots') or worse ('dark spots') than expected, given the environmental conditions and socioeconomic drivers they are exposed to. Here we compile data from more than 2,500 reefs worldwide and develop a Bayesian hierarchical model to generate expectations of how standing stocks of reef fish biomass are related to 18 socioeconomic drivers and environmental conditions. We identify 15 bright spots and 35 dark spots among our global survey of coral reefs, defined as sites that have biomass levels more than two standard deviations from expectations. Importantly, bright spots are not simply comprised of remote areas with low fishing pressure; they include localities where human populations and use of ecosystem resources is high, potentially providing insights into how communities have successfully confronted strong drivers of change. Conversely, dark spots are not necessarily the sites with the lowest absolute biomass and even include some remote, uninhabited locations often considered near pristine6. We surveyed local experts about social, institutional, and environmental conditions at these sites to reveal that bright spots are characterized by strong sociocultural institutions such as customary taboos and marine tenure, high levels of local engagement in management, high dependence on marine resources, and beneficial environmental conditions such as deep-water refuges. Alternatively, dark spots are characterized by intensive capture and storage technology and a recent history of environmental shocks. Our results suggest that investments in strengthening fisheries governance, particularly aspects such as participation and property rights, could facilitate innovative conservation actions that help communities defy expectations of global reef degradation.

Non-Random Variability in Functional Composition of Coral Reef Fish Communities along an Environmental Gradient.

Changes in the coral reef complex can affect predator-prey relationships, resource availability and niche utilisation in the associated fish community, which may be reflected in decreased stability of the functional traits present in a community. This is because particular traits may be favoured by a changing environment, or by habitat degradation. Furthermore, other traits can be selected against because degradation can relax the association between fishes and benthic habitat. We characterised six important ecological traits for fish species occurring at seven sites across a disturbed coral reef archipelago in Indonesia, where reefs have been exposed to eutrophication and destructive fishing practices for decades. Functional diversity was assessed using two complementary indices (FRic and RaoQ) and correlated to important environmental factors (live coral cover and rugosity, representing local reef health, and distance from shore, representing a cross-shelf environmental gradient). Indices were examined for both a change in their mean, as well as temporal (short-term; hours) and spatial (cross-shelf) variability, to assess whether fish-habitat association became relaxed along with habitat degradation. Furthermore, variability in individual traits was examined to identify the traits that are most affected by habitat change. Increases in the general reef health indicators, live coral cover and rugosity (correlated with distance from the mainland), were associated with decreases in the variability of functional diversity and with community-level changes in the abundance of several traits (notably home range size, maximum length, microalgae, detritus and small invertebrate feeding and reproductive turnover). A decrease in coral cover increased variability of RaoQ while rugosity and distance both inversely affected variability of FRic; however, averages for these indices did not reveal patterns associated with the environment. These results suggest that increased degradation of coral reefs is associated with increased variability in fish community functional composition resulting from selective impacts on specific traits, thereby affecting the functional response of these communities to increasing perturbations.

Local and Regional Impacts of Pollution on Coral Reefs along the Thousand Islands North of the Megacity Jakarta, Indonesia.

Worldwide, coral reefs are challenged by multiple stressors due to growing urbanization, industrialization and coastal development. Coral reefs along the Thousand Islands off Jakarta, one of the largest megacities worldwide, have degraded dramatically over recent decades. The shift and decline in coral cover and composition has been extensively studied with a focus on large-scale gradients (i.e. regional drivers), however special focus on local drivers in shaping spatial community composition is still lacking. Here, the spatial impact of anthropogenic stressors on local and regional scales on coral reefs north of Jakarta was investigated. Results indicate that the direct impact of Jakarta is mainly restricted to inshore reefs, separating reefs in Jakarta Bay from reefs along the Thousand Islands further north. A spatial patchwork of differentially degraded reefs is present along the islands as a result of localized anthropogenic effects rather than regional gradients. Pollution is the main anthropogenic stressor, with over 80% of variation in benthic community composition driven by sedimentation rate, NO2, PO4 and Chlorophyll a. Thus, the spatial structure of reefs is directly related to intense anthropogenic pressure from local as well as regional sources. Therefore, improved spatial management that accounts for both local and regional stressors is needed for effective marine conservation.