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Scleractinian corals are colonial animals with a range of life history strategies, making up diverse species assemblages that define coral reefs. We tagged and tracked approximately 30 colonies from each of 11 species during seven trips spanning six years (2009-2015) in order to measure their vital rates and competitive interactions on the reef crest at Trimodal Reef, Lizard Island, Australia. Pairs of species were chosen from five growth forms where one species of the pair was locally rare (R) and the other common (C). The sampled growth forms were massive [Goniastrea pectinata (R) and G. retiformis (C)], digitate [Acropora humilis (R) and A. cf. digitifera (C)], corymbose [A. millepora (R) and A. nasuta (C)], tabular [A. cytherea (R) and A. hyacinthus (C)] and arborescent [A. robusta (R) and A. intermedia (C)]. An extra corymbose species with intermediate abundance, A. spathulata was included when it became apparent that A. millepora was too rare on the reef crest, making the 11 species in total. The tagged colonies were visited each year in the weeks prior to spawning. During visits, two or more observers each took 2-3 photographs of each tagged colony from directly above and on the horizontal plane with a scale plate to track planar area. Dead or missing colonies were recorded and new colonies tagged in order to maintain approximately 30 colonies per species throughout the six years of the study. In addition to tracking tagged corals, 30 fragments were collected from neighboring untagged colonies of each species for counting numbers of eggs per polyp (fecundity); and fragments of untagged colonies were brought into the laboratory where spawned eggs were collected for biomass and energy measurements. We also conducted surveys at the study site to generate size structure data for each species in several of the years. Each tagged colony photograph was digitized by at least two people. Therefore, we could examine sources of error in planar area for both photographers and outliners. Competitive interactions were recorded for a subset of species by measuring the margins of tagged colony outlines interacting with neighboring corals. The study was abruptly ended by Tropical Cyclone Nathan (Category 4) that killed all but nine of the over 300 tagged colonies in early 2015. Nonetheless, these data will be of use to other researchers interested in coral demography and coexistence, functional ecology, and parametrizing population, community and ecosystem models. The data set is not copyright restricted, and users should cite this paper when using the data. Publisher PDF

Global warming is killing corals; however, the effects of warming on population connectivity, a process fundamental to reef recovery, are largely unexplored. Using a high-resolution (as high as 200 m), empirically calibrated biophysical model of coral larval dispersal for the southern Great Barrier Reef, we show that the increased larval mortality and reduced competency duration under a 2 °C warming alter dispersal patterns, whereas projected changes in large-scale currents have limited effects. Overall, there was on average a 7% decrease in the distance larvae disperse (among-reef interquartile range (IQR), −10% to −4%), an 8% decrease in the number of connections into each reef (IQR, −11% to −3%) and a 20% increase in local retention (IQR, 0% to +49%). Collectively, these shifts imply that 2 °C of warming will reduce inter-reef connectivity, hampering recovery after disturbances and reducing the spread of warm-adapted genes. Such changes make protections more effective locally, but may require reducing spacing between protected areas.

Ecosystems have always been shaped by disturbances, but many of these events are becoming larger, more severe and more frequent. The recovery capacity of depleted populations depends on the frequency of disturbances, the spatial distribution of mortality and the scale of dispersal. Here, we show that four mass coral bleaching events on the Great Barrier Reef (in 1998, 2002, 2016 and 2017) each had markedly larger disturbance footprints and were less patchy than a severe category 5 tropical cyclone (Cyclone Yasi, 2011). Severely bleached reefs in 2016 and 2017 were isolated from the nearest lightly affected reefs by up to 146 and 200 km, respectively. In contrast, reefs damaged by Cyclone Yasi were on average 20 km away from relatively undisturbed reefs, well within the estimated range of larval dispersal for most corals. Based on these results, we present a model of coral reef disturbance and recovery to examine (1) how the spatial clustering of disturbances modifies large-scale recovery rates; and (2) how recovery rates are shaped by species' dispersal abilities. Our findings illustrate that the spatial footprint of the recent mass bleaching events poses an unprecedented threat to the resilience of coral species in human history, a threat that is even larger than the amount of mortality suggests.

Sustainably managing fisheries requires regular and reliable evaluation of stock status. However, most multispecies reef fisheries around the globe tend to be data-poor and lack research and monitoring capacity (e.g., long-term fishery data), preventing the estimation of sustainable reference points against which stocks can be assessed. Here, combining fish biomass data for more than 2000 coral reefs with catch estimates from 99 jurisdictions, we estimate site-specific sustainable reference points for coral reef fisheries and use these to assess the status of coral reef fish stocks. We reveal that more than half of jurisdictions with available information have stocks of conservation concern, having failed at least one fisheries sustainability benchmark. We quantify the trade-offs between biodiversity, mean 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.TeaserA global assessment of the sustainability of multispecies reef fisheries indicates that more than half of jurisdictions have failed at least one of two key sustainability benchmarks.

JSM and MM were supported by the National Science Foundation (NSF)1948946. MD is supported by the Warman Foundation, the Leverhulme Centre for Anthropocene Biodiversity (RC-2018-021) and NSF-NERC grant NE/V009338/1. MM is supported by a Leverhulme Trust Early Career Fellowship (ECF-2021-512). Life-history traits are promising tools to predict species commonness and rarity because they influence a population's fitness in a given environment. Yet, species with similar traits can have vastly different abundances, challenging the prospect of robust trait-based predictions. Using long-term demographic monitoring, we show that coral populations with similar morphological and life-history traits show persistent (decade-long) differences in abundance. Morphological groups predicted species positions along two, well-known life-history axes (the fast-slow continuum and size-specific fecundity). However, integral projection models revealed that density-independent population growth (λ) was more variable within morphological groups, and was consistently higher in dominant species relative to rare species. Within-group λ differences projected large abundance differences among similar species in short timeframes, and were generated by small but compounding variation in growth, survival, and reproduction. Our study shows that easily-measured morphological traits predict demographic strategies, yet small life-history differences can accumulate into large differences in λ and abundance among similar species. Quantifying the net effects of multiple traits on population dynamics is therefore essential to anticipate species commonness and rarity. Publisher PDF

Environmental fluctuations are becoming increasingly volatile in many ecosystems, highlighting the need to better understand how stochastic and deterministic processes shape patterns of commonness and rarity, particularly in high-diversity systems like coral reefs. Here, we analyze reef fish time-series across the Great Barrier Reef to show that approximately 75% of the variance in relative species abundance is attributable to deterministic, intrinsic species differences. Nevertheless, the relative importance of stochastic factors is markedly higher on reefs that have experienced stronger coral cover volatility. By contrast, α-diversity and species composition are independent of coral cover volatility but depend on environmental gradients. Our findings imply that increased environmental volatility on coral reefs erodes assemblage’s niche structure, an erosion that is not detectable from static measures of biodiversity.One-Sentence SummaryCoral cover volatility modulates how stochastic and deterministic processes shape commonness and rarity in coral reef fishes.

Coral reef fisheries support the livelihoods of millions of people in tropical countries, despite large-scale depletion of fish biomass. While human adaptability can help to explain the resistance of fisheries to biomass depletion, compensatory ecological mechanisms may also be involved. If this is the case, high productivity should coexist with low biomass under relatively high exploitation. Here we integrate large spatial scale empirical data analysis and a theory-driven modelling approach to unveil the effects of human exploitation on reef fish productivity-biomass relationships. We show that differences in how productivity and biomass respond to overexploitation can decouple their relationship. As size-selective exploitation depletes fish biomass, it triggers increased production per unit biomass, averting immediate productivity collapse in both the modelling and the empirical systems. This 'buffering productivity' exposes the danger of assuming resource production-biomass equivalence, but may help to explain why some biomass-depleted fish assemblages still provide ecosystem goods under continued global fishing exploitation.

Changes in disturbance regimes due to climate change are increasingly challenging the capacity of ecosystems to absorb recurrent shocks and reassemble afterwards, escalating the risk of widespread ecological collapse of current ecosystems and the emergence of novel assemblages1–3. In marine systems, the production of larvae and recruitment of functionally important species are fundamental processes for rebuilding depleted adult populations, maintaining resilience and avoiding regime shifts in the face of rising environmental pressures4,5. Here we document a regional-scale shift in stock–recruitment relationships of corals along the Great Barrier Reef—the world’s largest coral reef system—following unprecedented back-to-back mass bleaching events caused by global warming. As a consequence of mass mortality of adult brood stock in 2016 and 2017 owing to heat stress6, the amount of larval recruitment declined in 2018 by 89% compared to historical levels. For the first time, brooding pocilloporids replaced spawning acroporids as the dominant taxon in the depleted recruitment pool. The collapse in stock–recruitment relationships indicates that the low resistance of adult brood stocks to repeated episodes of coral bleaching is inexorably tied to an impaired capacity for recovery, which highlights the multifaceted processes that underlie the global decline of coral reefs. The extent to which the Great Barrier Reef will be able to recover from the collapse in stock–recruitment relationships remains uncertain, given the projected increased frequency of extreme climate events over the next two decades7. A regional-scale shift in the relationships between adult stock and recruitment of corals occurred along the Great Barrier Reef, following mass bleaching events in 2016 and 2017 caused by global warming.

Anthropogenic climate change is a rapidly intensifying selection pressure on biodiversity across the globe and, particularly, on the world's coral reefs. The rate of adaptation to climate change is proportional to the amount of phenotypic variation that can be inherited by subsequent generations (i.e., narrow-sense heritability, h2 ). Thus, traits that have higher heritability (e.g., h2 > 0.5) are likely to adapt to future conditions faster than traits with lower heritability (e.g., h2

The discovery of multi-species synchronous spawning of scleractinian corals on the Great Barrier Reef in the 1980s stimulated an extraordinary effort to document spawning times in other parts of the globe. Unfortunately, most of these data remain unpublished which limits our understanding of regional and global reproductive patterns. The Coral Spawning Database (CSD) collates much of these disparate data into a single place. The CSD includes 6178 observations (3085 of which were unpublished) of the time or day of spawning for over 300 scleractinian species in 61 genera from 101 sites in the Indo-Pacific. The goal of the CSD is to provide open access to coral spawning data to accelerate our understanding of coral reproductive biology and to provide a baseline against which to evaluate any future changes in reproductive phenology., info:eu-repo/semantics/published

Marine multicellular organisms host a diverse collection of bacteria, archaea, microbial eukaryotes, and viruses that form their microbiome. Such host-associated microbes can significantly influence the host’s physiological capacities; however, the identity and functional role(s) of key members of the microbiome (“core microbiome”) in most marine hosts coexisting in natural settings remain obscure. Also unclear is how dynamic interactions between hosts and the immense standing pool of microbial genetic variation will affect marine ecosystems’ capacity to adjust to environmental changes. Here, we argue that significantly advancing our understanding of how host-associated microbes shape marine hosts’ plastic and adaptive responses to environmental change requires (i) recognizing that individual host–microbe systems do not exist in an ecological or evolutionary vacuum and (ii) expanding the field toward long-term, multidisciplinary research on entire communities of hosts and microbes. Natural experiments, such as time-calibrated geological events associated with well-characterized environmental gradients, provide unique ecological and evolutionary contexts to address this challenge. We focus here particularly on mutualistic interactions between hosts and microbes, but note that many of the same lessons and approaches would apply to other types of interactions., This Essay argues that in order to truly understand how marine hosts benefit from the immense diversity of microbes, we need to expand towards long-term, multi-disciplinary research focussing on few areas of the world’s ocean that we refer to as “natural experiments,” where processes can be studied at scales that far exceed those captured in laboratory experiments.

The age or size structure of a population has a marked influence on its demography and reproductive capacity. While declines in coral cover are well documented, concomitant shifts in the size-frequency distribution of coral colonies are rarely measured at large spatial scales. Here, we document major shifts in the colony size structure of coral populations along the 2300 km length of the Great Barrier Reef relative to historical baselines (1995/1996). Coral colony abundances on reef crests and slopes have declined sharply across all colony size classes and in all coral taxa compared to historical baselines. Declines were particularly pronounced in the northern and central regions of the Great Barrier Reef, following mass coral bleaching in 2016 and 2017. The relative abundances of large colonies remained relatively stable, but this apparent stability masks steep declines in absolute abundance. The potential for recovery of older fecund corals is uncertain given the increasing frequency and intensity of disturbance events. The systematic decline in smaller colonies across regions, habitats and taxa, suggests that a decline in recruitment has further eroded the recovery potential and resilience of coral populations.

Reef-building coral assemblages are typically species-rich, yet the processes maintaining coral biodiversity remain poorly understood. Disturbance has long been believed to promote coral species coexistence by reducing the strength of competition. However, such disturbance-induced effects have since been shown to be insufficient on their own to prevent competitive exclusion. Nevertheless, Modern Coexistence Theory has revealed other mechanisms by which disturbance and, more generally, environmental variation can favour coexistence. Here, we formulate, calibrate, and analyze a size-structured, stochastic coral competition model using field data from two common colony morphologies. These two coral morphologies, tabular and digitate, differ in their size-dependent vulnerability to dislodgement caused by wave action. We confirm that fluctuations in wave action can promote coral species coexistence. However, using a recently proposed partitioning framework, we show that, contrast to previous expectations, temporal variability in strength of competition did not promote coexistence. Instead, coexistence was enabled by differential fluctuations in size-dependent mortality among competitors. Frequent and intense disturbances resulted in monocultures of digitate corals, which are more robust to wave action than tabular corals. In contrast, infrequent or weak disturbances resulted in monocultures of tabular corals. Coexistence was only possible under intermediate levels of disturbance frequency and intensity. Given the sensitivity of coexistence to disturbance frequency and intensity, anthropogenic changes in disturbance regimes are likely to affect biodiversity in coral assemblages in ways that are not predictable from single population models.

Knowledge of a species’ abundance is critically important for assessing its risk of extinction, but for the vast majority of wild animal and plant species such data are scarce at biogeographic scales. Here, we estimate the total number of reef-building corals and the population sizes of more than 300 individual species on reefs spanning the Pacific Ocean biodiversity gradient, from Indonesia to French Polynesia. Our analysis suggests that approximately half a trillion corals (0.3 × 1012–0.8 × 1012) inhabit these coral reefs, similar to the number of trees in the Amazon. Two-thirds of the examined species have population sizes exceeding 100 million colonies, and one-fifth of the species even have population sizes greater than 1 billion colonies. Our findings suggest that, while local depletions pose imminent threats that can have ecologically devastating impacts to coral reefs, the global extinction risk of most coral species is lower than previously estimated. Approximately half a trillion reef-building corals live across the Pacific Ocean, based on estimates of the population sizes of more than 300 species.

Climate change is radically altering the frequency, intensity and spatial scale of severe weather events, such as heatwaves, droughts, floods and fires1. As the time interval shrinks between recurrent shocks2–5, the responses of ecosystems to each new disturbance are increasingly likely to be contingent on the history of other recent extreme events. Ecological memory—defined as the ability of the past to influence the present trajectory of ecosystems6,7—is also critically important for understanding how species assemblages are responding to rapid changes in disturbance regimes due to anthropogenic climate change2,3,6–8. Here, we show the emergence of ecological memory during unprecedented back-to-back mass bleaching of corals along the 2,300 km length of the Great Barrier Reef in 2016, and again in 2017, whereby the impacts of the second severe heatwave, and its geographic footprint, were contingent on the first. Our results underscore the need to understand the strengthening interactions among sequences of climate-driven events, and highlight the accelerating and cumulative impacts of novel disturbance regimes on vulnerable ecosystems. The increasing frequency of marine heatwaves suggests that the impacts of successive events may be influenced by previous events. The extent of the 2016 and 2017 bleaching events on the Great Barrier Reef shows that ecological memory played a role in the impacts of the second heatwave.

Funding was provided by the Australian Research Council Centre of Excellence for Coral Reef Studies (CE140100020), a Future Fellowship (A.H.B.), an Australian Professorial Fellowship (S.R.C.) and the Templeton Foundation (grant no. 60501, ‘Putting the Extended Evolutionary Synthesis to the Test’). M.D. is grateful to the Scottish Funding Council (MASTS, grant reference HR09011) and the European Research Council (grant BioTIME). Disturbances, such as cyclones, have a major effect on the structure and dynamics of coral reef assemblages. However, the effect of cyclones on demographic traits, such as fecundity, has rarely been quantified, and direct estimates of mortality at the species level are rare. Here, we document the effect of Severe Tropical Cyclone Nathan on the demography of corals on the reef crest at Trimodal Reef in the northern Great Barrier Reef. Mortality rates based on tagged colonies were very high, ranging from 85.2% in Goniastrea retiformis to 100% in 6 Acropora species, 3 to 40 times higher than averages rates in the 5 yr preceding Cyclone Nathan. Fecundity was lower in 3 out of the 4 species examined following the cyclone, and egg carbon content was reduced by 58-63% in the 2 species examined. These results suggest that energy normally invested in reproduction was diverted to other processes such as injury repair and demonstrate that cyclones have important sub-lethal effects in addition to high rates of whole colony mortality. Coral cover was reduced from 34.9 ± 3.9% (mean ± SE) to 3.4 ± 1.5%, with reductions in the cover of all taxa except those with predominantly massive morphologies such as the Poritidae. A projected increase in the frequency of tropical storms as a result of global warming, combined with an increase in the frequency and scale of coral bleaching, suggest a fundamental shift in mortality regimes on reefs which has the potential to threaten their long-term persistence. Postprint

Funding was provided by the Australian Council Centre of Excellence for Coral Reef Studies (COE140100020) and the John Templeton Foundation (M.D., J.S.M. grant #60501 'Putting the Extended Evolutionary Synthesis to the Test’). Coral reefs are being subjected to an increase in the frequency and intensity of disturbance, such as bleaching and cyclones, and it is important to document the effects of such disturbance on reef coral assemblages. Between March 2014 and May 2017, the reefs of Lizard Island in the northern section of the Great Barrier Reef were affected by 4 consecutive disturbances: severe tropical cyclones Ita and Nathan in 2014 and 2015, and mass bleaching events in 2016 and 2017. Loss of coral cover following the cyclones was patchy and dependent on the direction of the waves generated. In contrast, loss of cover following bleaching was much more uniform. Overall, coral cover declined 5-fold from 36% pre-cyclone Ita to 7% post-bleaching in 2017, while mean species richness dropped from 10 to 4 species per transect. The spatial scale and magnitude of the loss of coral cover in the region suggests that it will be many years before these reefs recover. Postprint

Up to 90% of global coral reefs are predicted to be severely degraded by 2050 under “business-as-usual” scenarios. To meet the scale and scope of this challenge, we propose designing and demonstrating a multi-modal system that can incorporate data from remote sensing (satellites, aircraft, and aerial drones), acoustics, genetics, sensor arrays, and low-cost imaging systems. The latter will be collected by low-cost smart sensing and autonomous underwater vehicles (AUVs) guided by adaptive sampling modeling software and rapidly analyzed using automated machine learning systems. Development and deployment will be linked to extensive and diversity-enhancing training programs. The Coral Sentinel System will be globally deployed to enable rapid-response adaptive management and to build public engagement in conservation interventions to save coral reefs.Phase 1 (Year 1) will involve testing assumptions, coalition building, fundraising, and initial system development. Phase 2 (Years 2‐4) will focus on engineering and development with a pilot deployment in the Caribbean. Phase 3 (Years 5‐6) will involve system expansion and iteration along the Tropical Eastern Pacific corridor. Phase 4 (Years 7‐10) will involve global deployment to over 50 reef sites. This will lead during the following decade (Phase 5) to provisioning of low-cost Sentinel systems to coastal communities globally.

The frequency, intensity, and spatial scale of climate extremes are changing rapidly due to anthropogenic global warming.1,2 A growing research challenge is to understand how multiple climate-driven disturbances interact with each other over multi-decadal time frames, generating combined effects that cannot be predicted from single events alone.3, 4, 5 Here we examine the emergent dynamics of five coral bleaching events along the 2,300 km length of the Great Barrier Reef that affected >98% of the Reef between 1998 and 2020. We show that the bleaching responses of corals to a given level of heat exposure differed in each event and were strongly influenced by contingency and the spatial overlap and strength of interactions between events. Naive regions that escaped bleaching for a decade or longer were the most susceptible to bouts of heat exposure. Conversely, when pairs of successive bleaching episodes were close together (1–3 years apart), the thermal threshold for severe bleaching increased because the earlier event hardened regions of the Great Barrier Reef to further impacts. In the near future, the biological responses to recurrent bleaching events may become stronger as the cumulative geographic footprint expands further, potentially impairing the stock-recruitment relationships among lightly and severely bleached reefs with diverse recent histories. Understanding the emergent properties and collective dynamics of recurrent disturbances will be critical for predicting spatial refuges and cumulative ecological responses, and for managing the longer-term impacts of anthropogenic climate change on ecosystems.

Not enough time for recovery Coral bleaching occurs when stressful conditions result in the expulsion of the algal partner from the coral. Before anthropogenic climate warming, such events were relatively rare, allowing for recovery of the reef between events. Hughes et al. looked at 100 reefs globally and found that the average interval between bleaching events is now less than half what it was before. Such narrow recovery windows do not allow for full recovery. Furthermore, warming events such as El Niño are warmer than previously, as are general ocean conditions. Such changes are likely to make it more and more difficult for reefs to recover between stressful events. Science , this issue p. 80

Macroecology has traditionally relied on descriptive characterization of large-scale ecological patterns to offer narrative explanations for the origin and maintenance of those patterns. Only recently have macroecologists begun to employ models termed ‘process-based’ and ‘mechanistic’, in contrast to other areas of ecology, where such models have a longer history. Here, we define and differentiate between process-based and mechanistic features of models, and we identify and discuss important advantages of working with models possessing such features. We describe some of the risks associated with process-based and mechanistic model-centered research programs, and we propose ways to mitigate these risks. Giving process-based and mechanistic models a more central role in research programs can reinvigorate macroecology by strengthening the link between theory and data.

Contemporary science educators must equip their students with the knowledge and practical know-how to connect multiple disciplines like mathematics, computing and the natural sciences to gain a richer and deeper understanding of a scientific problem. However, many biology and earth science students are prejudiced against mathematics due to negative emotions like high mathematical anxiety and low mathematical confidence. Here, we present a theoretical framework that investigates linkages between student engagement, mathematical anxiety, mathematical confidence, student achievement and subject mastery. We implement this framework in a large, first-year interdisciplinary science subject and monitor its impact over several years from 2010 to 2015. The implementation of the framework coincided with an easing of anxiety and enhanced confidence, as well as higher student satisfaction, retention and achievement. The framework offers interdisciplinary science educators greater flexibility and confidence in their approach to designing and delivering subjects that rely on mathematical concepts and practices.

Abundance patterns in ecological communities have important implications for biodiversity maintenance and ecosystem functioning. However, ecological theory has been largely unsuccessful at capturing multiple macroecological abundance patterns simultaneously. Here, we propose a parsimonious model that unifies widespread ecological relationships involving local aggregation, species-abundance distributions, and species associations, and we test this model against the metacommunity structure of reef-building corals and coral reef fishes across the western and central Pacific. For both corals and fishes, the unified model simultaneously captures extremely well local species-abundance distributions, interspecific variation in the strength of spatial aggregation, patterns of community similarity, species accumulation, and regional species richness, performing far better than alternative models also examined here and in previous work on coral reefs. Our approach contributes to the development of synthetic theory for large-scale patterns of community structure in nature, and to addressing ongoing challenges in biodiversity conservation at macroecological scales.

Aim: To test hypothesized biogeographic partitions of the tropical Indo-Pacific Ocean with phylogeographic data from 56 taxa, and to evaluate the strength and nature of barriers emerging from this test. Location: The Indo-Pacific Ocean. Time period: Pliocene through the Holocene. Major taxa studied: Fifty-six marine species. Methods: We tested eight biogeographic hypotheses for partitioning of the Indo-Pacific using a novel modification to analysis of molecular variance. Putative barriers to gene flow emerging from this analysis were evaluated for pairwise ΦST, and these ΦST distributions were compared to distributions from randomized datasets and simple coalescent simulations of vicariance arising from the Last Glacial Maximum. We then weighed the relative contribution of distance versus environmental or geographic barriers to pairwise ΦST with a distance-based redundancy analysis (dbRDA). Results: We observed a diversity of outcomes, although the majority of species fit a few broad biogeographic regions. Repeated coalescent simulation of a simple vicariance model yielded a wide distribution of pairwise ΦST that was very similar to empirical distributions observed across five putative barriers to gene flow. Three of these barriers had median ΦST that were significantly larger than random expectation. Only 21 of 52 species analysed with dbRDA rejected the null model. Among these, 15 had overwater distance as a significant predictor of pairwise ΦST, while 11 were significant for geographic or environmental barriers other than distance. Main conclusions: Although there is support for three previously described barriers, phylogeographic discordance in the Indo-Pacific Ocean indicates incongruity between processes shaping the distributions of diversity at the species and population levels. Among the many possible causes of this incongruity, genetic drift provides the most compelling explanation: given massive effective population sizes of Indo-Pacific species, even hard vicariance for tens of thousands of years can yield ΦSTvalues that range from 0 to nearly 0.5.

Marine reserve networks are increasingly implemented to conserve biodiversity and enhance the persistence and resilience of exploited species and ecosystems. However, the efficacy of marine reserve networks in frequently disturbed systems, such as coral reefs, has rarely been evaluated. Here we analyze a well-mixed larval pool model and a spatially explicit model based on a well-documented coral trout (Plectropomus spp.) metapopulation in the Great Barrier Reef Marine Park, Australia, to determine the effects of marine reserve coverage and placement (in relation to larval connectivity and disturbance heterogeneity) on the temporal stability of fisheries yields and population biomass in environmentally disturbed systems. We show that marine reserves can contribute to stabilizing fishery yield while increasing metapopulation persistence, irrespective of whether reserves enhance or diminish average fishery yields. However, reserve placement and the level of larval connectivity among subpopulations were important factors affecting the stability and sustainability of fisheries and fish metapopulations. Protecting a mix of disturbed and non-disturbed reefs, rather than focusing on the least-disturbed habitats, was the most consistently beneficial approach across a range of dispersal and reserve coverage scenarios. Placing reserves only in non-disturbed areas was the most beneficial for biomass enhancement, but had variable results for fisheries and could potentially destabilize yields in systems with well-mixed larval or those that are moderately fished. We also found that focusing protection on highly disturbed areas could actually increase variability in yields and biomass, especially when degraded reef reserves were distant and poorly connected to the meta-population. Our findings have implications for the design and implementation of reserve networks in the presence of stochastic, patchy environmental disturbances.

We thank the University of St Andrews MHD Cluster and the Bioinformatics Unit (Wellcome Trust ISSF grant 105621/Z/14/Z). L.H.A. was supported by Fundacao para a Ciencia e Tecnologia, Portugal (POPH/FSE SFRH/BD/90469/2012). A.E.M. acknowledges the ERC (BioTIME 250189). M.D. acknowledges funding from the Marine Alliance for Science and Technology Scotland (MASTS), funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions.

Removal of predators is often hypothesized to alter community structure through trophic cascades. However, despite recent advances in our understanding of trophic cascades, evidence is often circumstantial on coral reefs because fishing pressure frequently co-varies with other anthropogenic effects, such as fishing for herbivorous fishes and changes in water quality due to pollution. Australia's outer Great Barrier Reef (GBR) has experienced fishing-induced declines of apex predators and mesopredators, but pollution and targeting of herbivorous fishes are minimal. Here, we quantify fish and benthic assemblages across a fishing-induced predator density gradient on the outer GBR, including apex predators and mesopredators to herbivores and benthic assemblages, to test for evidence of trophic cascades and alternative hypotheses to trophic cascade theory. Using structural equation models, we found no cascading effects from apex predators to lower trophic levels: a loss of apex predators did not lead to higher levels of mesopredators, and this did not suppress mobile herbivores and drive algal proliferation. Likewise, we found no effects of mesopredators on lower trophic levels: a decline of mesopredators was not associated with higher abundances of algae-farming damselfishes and algae-dominated reefs. These findings indicate that top-down forces on coral reefs are weak, at least on the outer GBR. We conclude that predator-mediated trophic cascades are probably the exception rather than the rule in complex ecosystems such as the outer GBR.

Predicted increases in sea-surface temperatures due to climate change are likely to alter the physiology of marine organisms and ultimately influence the distribution and abundance of their populations. The consequences of increased temperatures for marine species, including decreased survival and altered rates of development, growth and settlement, are well known and often attributed to imbalances between energy supply and demand. To test this hypothesis, we calibrated the effect of temperature on rates of survival and lipid depletion for larvae of the common stony coral Acropora tenuis over a 7 °C temperature range. Temperature had a pronounced, linearly increasing effect on larval mortality, with a sixfold decrease in median survival time. Contrary to expectation, however, temperature had a quasi-parabolic effect on lipid use; rates declined as temperatures either increased above or decreased below the ambient temperature at the time of spawning. This contrasts with previous work suggesting that increased energy depletion is the cause of larval mortality at higher temperatures. Our results highlight the sensitivity of coral larvae to temperature and have implications for dispersal potential because fewer larvae will survive to disperse. Such projected declines in connectivity among coral populations are likely to undermine reef resilience.

Ecologists have often used indirect proxies to represent variables that are difficult or impossible to measure directly. In phytoplankton, the internal concentration of the most limiting nutrient in a cell determines its growth rate. However, directly measuring the concentration of nutrients within cells is inaccurate, expensive, destructive, and time-consuming, substantially impairing our ability to model growth rates in nutrient-limited phytoplankton populations. The red chlorophyll autofluorescence (hereafter "red fluorescence") signal emitted by a cell is highly correlated with nitrogen quota in nitrogen-limited phytoplankton species. The aim of this study was to evaluate the reliability of including flow cytometric red fluorescence as a proxy for internal nitrogen status to model phytoplankton growth rates. To this end, we used the classic Quota model and designed three approaches to calibrate its model parameters to data: where empirical observations on cell internal nitrogen quota were used to fit the model ("Nitrogen-Quota approach"), where quota dynamics were inferred only from changes in medium nutrient depletion and population density ("Virtual-Quota approach"), or where red fluorescence emission of a cell was used as an indirect proxy for its internal nitrogen quota ("Fluorescence-Quota approach"). Two separate analyses were carried out. In the first analysis, stochastic model simulations were parameterized from published empirical relationships and used to generate dynamics of phytoplankton communities reared under nitrogen-limited conditions. Quota models were fitted to the dynamics of each simulated species with the three different approaches and the performance of each model was compared. In the second analysis, we fit Quota models to laboratory time-series and we calculate the ability of each calibration approach to describe the observed trajectories of internal nitrogen quota in the culture. Results from both analyses concluded that the Fluorescence-Quota approach including per-cell red fluorescence as a proxy of internal nitrogen substantially improved the ability of Quota models to describe phytoplankton dynamics, while still accounting for the biologically important process of cell nitrogen storage. More broadly, many population models in ecology implicitly recognize the importance of accounting for storage mechanisms to describe the dynamics of individual organisms. Hence, the approach documented here with phytoplankton dynamics may also be useful for evaluating the potential of indirect proxies in other ecological systems.

Coral reefs are biologically diverse and ecologically complex ecosystems constructed by stony corals. Despite decades of research, basic coral population biology and community ecology questions remain. Quantifying trait variation among species can help resolve these questions, but progress has been hampered by a paucity of trait data for the many, often rare, species and by a reliance on nonquantitative approaches. Therefore, we propose filling data gaps by prioritizing traits that are easy to measure, estimating key traits for species with missing data, and identifying 'supertraits' that capture a large amount of variation for a range of biological and ecological processes. Such an approach can accelerate our understanding of coral ecology and our ability to protect critically threatened global ecosystems.

Marine no-take reserves, where fishing and other extractive activities are prohibited, have well-established conservation benefits [1], yet their impacts on fisheries remains contentious [2-4]. For fishery species, reserves are often implemented alongside more conventional harvest strategies, including catch and size limits [2, 5]. However, catch and fish abundances observed post-intervention are often attributed to reserves, without explicitly estimating the potential contribution of concurrent management interventions [2, 3, 6-9]. Here we test a metapopulation model against observed fishery [10] and population [11] data for an important coral reef fishery (coral trout; Plectropomus spp.) in Australia's Great Barrier Reef Marine Park (GBRMP) to evaluate how the combined increase in reserve area [12] and reduction in fishing effort [13, 14] in 2004 influenced changes in fish stocks and the commercial fishery. We found that declines in catch, increases in catch rates, and increases in biomass since 2004 were substantially attributable to the integration of direct effort controls with the rezoning, rather than the rezoning alone. The combined management approach was estimated to have been more productive for fish and fisheries than if the rezoning had occurred alone and comparable to what would have been obtained with effort controls alone. Sensitivity analyses indicate that the direct effort controls prevented initial decreases in catch per unit effort that would have otherwise occurred with the rezoning. Our findings demonstrate that by concurrently restructuring the fishery, the conservation benefits of reserves were enhanced and the fishery cost of rezoning the reserve network was socialized, mitigating negative impacts on individual fishers.

We are grateful to the North Queensland Algal Identification and Culturing Facility (NQAIF), in particular Stan Hudson and Florian Berner. We are grateful to the Ecological Modelling Research Group at James Cook University, especially Dr Loic Thibaut for comments on modelling techniques and R codes. Also, thanks to the High Performance Computing team at James Cook University. We thank Dr Lyndon Llewellyn, Dr Christian Lonborg and Dr Catia Carreira for helpful advice. Finally, we thank A/Prof A. Martiny and an anonymous reviewer for detailed and constructive comments on the manuscript. This research was supported by AIMS@JCU (aims.jcu.edu.au), the Australian Institute of Marine Science (www.aims.gov.au), the Advanced Manufacturing Cooperative Research Centre (Project 2.3.4) and James Cook University (www.jcu.edu.au). The authors declare no conflict of interest.

Marine reserves are often established in areas that support fisheries. Larval export from reserves is argued to help compensate for the loss of fishable habitat; however, previous modeling studies have focused on long-term equilibrium outcomes. We examined the transient consequences of reserve establishment for fished metapopulations, considering both a well-mixed larval pool and a spatially explicit model based on a coral trout (Plectropomus spp.) metapopulation. When fishing pressure was reallocated relative to the area protected, yields decreased initially, then recovered, and ultimately exceeded pre-reserve levels. However, recovery time was on the order of several years to decades. If fishing pressure intensified to maintain pre-reserve yields, reserves were sometimes unable to support the increased mortality and the metapopulation collapsed. This was more likely when reserves were small, or located peripherally within the metapopulation. Overall, reserves can achieve positive conservation and fishery benefits, but fisheries management complementary to reserve implementation is essential.

Models of dispersal potential are required to predict connectivity between populations of sessile organisms. However, to date, such models do not allow for time-varying rates of acquisition and loss of competence to settle and metamorphose, and permit only a limited range of possible survivorship curves. We collect high-resolution observations of coral larval survival and metamorphosis, and apply a piecewise modeling approach that incorporates a broad range of temporally varying rates of mortality and loss of competence. Our analysis identified marked changes in competence loss and mortality rates, the timing of which implicates developmental failure and depletion of energy reserves. Asymmetric demographic rates suggest more intermediate-range dispersal, less local retention, and less long-distance dispersal than predicted by previously employed non-piecewise models. Because vital rates are likely temporally asymmetric, at least for nonfeeding broadcast-spawned larvae, piecewise analysis of demographic rates will likely yield more reliable predictions of dispersal potential.

The extent to which populations in nature are regulated by density-dependent processes is unresolved. While experiments increasingly find evidence of strong density dependence, unmanipulated population time series yield much more ambiguous evidence of regulation, especially when accounting for effects of observation error. Here, we reexamine the evidence for density dependence in time series of population sizes in nature, by conducting an aggregate analysis of the populations in the Global Population Dynamics Database (GPDD). First, following the conventional approach, we fit a density-dependent and a density-independent variant of the Gompertz state-space model to each time series. Then, we conduct an aggregate analysis of the entire database by considering two random-effects density-dependent models that leverage information across data sets. When individual time series are tested independently, we find very little evidence for density dependence. However, in the aggregate, we find very strong evidence for density dependence, even though, paradoxically, estimated strengths of density dependence for individual time series tend to be weaker than when each individual time series is analyzed independently. Furthermore, a hierarchical model that accounts for taxonomic variation in the strength of density dependence reveals that density dependence is consistently stronger in insects and fish than in birds and mammals. Our findings resolve apparent inconsistencies between observational and experimental studies of density dependence by revealing that the observational record does indeed contain strong support for the hypothesis that density dependence is widespread in nature.

Changes in disturbance regimes due to climate change are increasingly challenging the capacity of ecosystems to absorb recurrent shocks and reassemble afterwards, escalating the risk of widespread ecological collapse of current ecosystems and the emergence of novel assemblages

Nitrate and ammonium are the two most important ionic forms of inorganic nitrogen driving biomass production in marine and freshwater aquatic systems. The performance of plants and algae often changes when reared with either of these two forms of nitrogen individually, as well as when they are both present, or when cells have experienced previous periods of nitrogen starvation. Current functional responses quantifying how ambient nitrogen drives changes in population density are unable to capture interacting and transient effects of nitrate and ammonium. Hence, in this paper we formulate, calibrate, and test a new nitrate–ammonium quota model that accounts for nitrate and ammonium uptake, as well as the effects of nitrogen starvation length and ammonium-induced nitrate uptake inhibition. We fit the model with several time-series from the green alga Chlorella sp. reared in laboratory batch cultures under multiple initial conditions. We show that a single set of calibrated model parameters can capture time-series collected from experiments inoculated at 12 different initial concentrations of nitrate, ammonium, and biomass. The model also performed well when validated against time-series from two novel initial conditions withheld from model calibration. Our model therefore provides a framework for evaluating the potential broader ecological and environmental consequences of ambient nitrate and ammonium regimes for phytoplankton communities in nature and aquaculture.

We are grateful to the North Queensland Algal Identification and Culturing Facility (NQAIF), in particular Stan Hudson and Florian Berner. We also thank A/Prof Bruce Bowden and Prof James Burnell for the assistance in laboratory protocols. Finally, we thank Dr Lyndon Llewellyn, Dr Christian Lonborg, Dr Murray Logan, and Dr Catia Carreira for the helpful advice. This research was supported by AIMS@JCU (aims.jcu.edu.au), the Australian Institute of Marine Science (www.aims.gov.au), the Advanced Manufacturing Cooperative Research Centre (Project 2.3.4), and James Cook University (www.jcu.edu.au). We also thank the reviewers, whose comments and suggestions helped improve the manuscript.

Saura () claims that studies using the Probability of Connectivity metric (PC) had already demonstrated the importance of including node self-connections in network metrics. As originally defined and used, PC cannot test the importance of self-connections. However, with key terms redefined, PC could be a useful tool in future work.

Trait-based approaches advance ecological and evolutionary research because traits provide a strong link to an organism's function and fitness. Trait-based research might lead to a deeper understanding of the functions of, and services provided by, ecosystems, thereby improving management, which is vital in the current era of rapid environmental change. Coral reef scientists have long collected trait data for corals; however, these are difficult to access and often under-utilized in addressing large-scale questions. We present the Coral Trait Database initiative that aims to bring together physiological, morphological, ecological, phylogenetic and biogeographic trait information into a single repository. The database houses species- and individual-level data from published field and experimental studies alongside contextual data that provide important framing for analyses. In this data descriptor, we release data for 56 traits for 1547 species, and present a collaborative platform on which other trait data are being actively federated. Our overall goal is for the Coral Trait Database to become an open-source, community-led data clearinghouse that accelerates coral reef research.

MD was supported by the ERC (BioTIME 250189) and the Scottish Funding Council (MASTS –HR09011). JM, AB and SC were supported by fellowships from the Australian Research Council (FT110100609, FT0990652 and DP0880544, respectively). Competition is an important determinant of assemblage structure and population regulation, often resulting in decreased growth, fecundity or survival. In corals, most studies testing for an effect of competition on demographic traits, such as growth, have been experimental and often impose very high levels of competition upon colonies. To more realistically assess the role of competition on coral traits, multispecies studies in the wild are required. Here, we use 5 yr of data that includes 11 coral species on the reef crest at Lizard Island on the Great Barrier Reef to quantify the effect of competition on growth. Additionally, we test whether species differ in their susceptibility to direct‐contact (overgrowth and digestion) and overtopping competition, and whether species from some morphological groups are more likely to compete with one another than with species from other morphological groups. We also investigate the relationships between competitive ability and three key traits: growth rate, mechanical stability and fecundity. In contrast to most previous work using field manipulations of competition, we found a negligible effect of competition on growth. Acropora species consistently won overtopping encounters but lost in direct‐contact encounters, and these results were consistent among the four Acropora morphological groups. In contrast, the massive Goniastrea spp. were poor at overtopping but generally won direct‐contact encounters. Only tabular colonies were disproportionally more likely to compete against one another than with other morphologies. This propensity increases intraspecific relative to interspecific competition, a phenomenon that can promote coexistence when it is present among dominant competitors. Good competitors grew more quickly and had higher fecundity but were less mechanically stable, implying a tradeoff between performance during disturbance vs. performance in the absence of disturbance. We conclude that competition among adults is less likely to influence community dynamics than previously thought. If competition does have an effect, it is more likely to occur at life‐stages other than adults. Publisher PDF

Global warming is rapidly emerging as a universal threat to ecological integrity and function, highlighting the urgent need for a better understanding of the impact of heat exposure on the resilience of ecosystems and the people who depend on them 1 . Here we show that in the aftermath of the record-breaking marine heatwave on the Great Barrier Reef in 2016 2 , corals began to die immediately on reefs where the accumulated heat exposure exceeded a critical threshold of degree heating weeks, which was 3–4 °C-weeks. After eight months, an exposure of 6 °C-weeks or more drove an unprecedented, regional-scale shift in the composition of coral assemblages, reflecting markedly divergent responses to heat stress by different taxa. Fast-growing staghorn and tabular corals suffered a catastrophic die-off, transforming the three-dimensionality and ecological functioning of 29% of the 3,863 reefs comprising the world’s largest coral reef system. Our study bridges the gap between the theory and practice of assessing the risk of ecosystem collapse, under the emerging framework for the International Union for Conservation of Nature (IUCN) Red List of Ecosystems 3 , by rigorously defining both the initial and collapsed states, identifying the major driver of change, and establishing quantitative collapse thresholds. The increasing prevalence of post-bleaching mass mortality of corals represents a radical shift in the disturbance regimes of tropical reefs, both adding to and far exceeding the influence of recurrent cyclones and other local pulse events, presenting a fundamental challenge to the long-term future of these iconic ecosystems.