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Insights into assemblages that can persist in extreme environments are still emerging. Ocean warming and acidification select against species with low physiological tolerance (trait-based 'filtering'). However, intraspecific trait variation can promote species adaptation and persistence, with potentially large effects on assemblage structure. By sampling nine coral traits (four morphological, four tissue and one skeletal) along an offshore-inshore gradient in temperature and pH, we show that distantly related coral species undergo consistent intraspecific changes as they cross into warm, acidic environments. Intraspecific variation and species turnover each favoured colonies with greater tissue biomass, higher symbiont densities and reduced skeletal investments, indicating strong filtering on colony physiology within and across species. Physiological tissue traits were highly variable within species and were independent of morphology, enabling morphologically diverse species to cross into sites of elevated temperature and acidity. Widespread intraspecific change can therefore counter the loss of biodiversity and morphological structure across a steep environmental gradient.

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

Increasing incidence of major disturbances is contributing to extensive and widespread coral loss, thereby undermining the biodiversity, structure and function of reef ecosystems. The composition of coral assemblages is already changing due to selective effects of recurrent disturbances, combined with marked differences in the underlying life-history dynamics of corals, which affects their recovery. This study quantifies the effects of varying disturbance regimes on two groups of corals with divergent life histories: short-lived species with rapid growth (bushy and tabular Acropora) and long-lived species with slow growth (massive and columnar Porites). Inter-decadal shifts in the coral assemblages across four locations suggest that a high frequency of moderate disturbances favours Porites, whereas infrequent, but severe disturbances favour rapidly replenishing Acropora. Using empirical modelling, we expand these observations to show that Acropora continues to dominate so long as the interval between major disturbances is > 2 years. The only disturbance regime we considered that favoured Porites was high frequency (2-year recurrence) of moderate disturbance, whereas high frequency of severe disturbances led to local extirpation of both Acropora and Porites. Our results show that increasing incidence of major disturbances will not necessarily lead to selective loss of species that are most susceptible to disturbance, as long as these species can continue to colonise vacant space and grow quickly in the aftermath of such disturbances. This study highlights the need to consider the sensitivity of taxa to changes in both disturbance frequency and severity when forecasting changes in the composition of coral assemblages under new disturbance regimes.

Humans have long sought to restore species, but little attention has been directed at how to best select a subset of foundation species for maintaining rich assemblages that support ecosystems, like coral reefs and rainforests that are increasingly threatened by environmental change.We propose a two-part hedging approach that selects optimized sets of species for restoration. The first part acknowledges that biodiversity supports ecosystem functions and services, and so it takes precaution against loss by ensuring an even spread of phenotypic traits. The second part maximizes species and ecosystem persistence by weighting species based on characteristics that are known to improve ecological persistence—e.g., abundance, species range and tolerance to environmental change.Using existing phenotypic trait and ecological characteristic data for reef building corals, we identified sets of ecologically persistent species by examining marginal returns in occupancy of phenotypic trait space. We compared optimal sets of species with those from the world’s southern-most coral reef which naturally harbors low coral diversity to show these occupy much of the trait space. Comparison with an existing coral restoration program indicated that current corals used for restoration only cover part of the desired trait space and may be improved by including species with different traits.Synthesis and applications. While there are many possible criteria for selecting species for restoration, the approach proposed here addresses the need to insure against unpredictable losses of ecosystem services by focusing on a wide range of phenotypic traits and ecological characteristics. Furthermore, the flexibility of the approach enables the functional goals of restoration to vary depending on environmental context, stakeholder values, and the spatial and temporal scales at which meaningful impacts can be achieved.

Structure-from-motion (SfM) photogrammetry is a technique used to generate three-dimensional (3D) reconstructions from a sequence of two-dimensional (2D) images. SfM methods are becoming increasingly popular as a noninvasive way to monitor many systems, including anthropogenic and natural landscapes, geologic structures, and both terrestrial and aquatic ecosystems. Here, a detailed protocol is provided for collecting SfM imagery to generate 3D models of benthic habitats. Additionally, the cost, time efficiency, and output quality of employing a Digital Single Lens Reflex (DSLR) camera versus a less expensive action camera have been compared. A tradeoff between computational time and resolution was observed, with the DSLR camera producing models with more than twice the resolution, but taking approximately 1.4-times longer to produce than the action camera. This primer aims to provide a thorough description of the steps necessary to collect SfM data in benthic habitats for those who are unfamiliar with the technique as well as for those already using similar methods.

Changes in the size structure of coral populations have major consequences for population dynamics and community function, yet many coral reef monitoring projects do not record this critical feature. Consequently, our understanding of current and future trajectories in coral size structure, and the demographic processes underlying these changes, is still emerging. Here, we provide a conceptual summary of the benefits to be gained from more comprehensive attention to the size of coral colonies in reef monitoring projects, and we support our argument through the use of case-history examples and a simplified ecological model. We neither seek to review the available empirical data, or to rigorously explore causes and implications of changes in coral size, we seek to reveal the advantages to modifying ongoing programs to embrace the information inherent in changing coral colony size. Within this framework, we evaluate and forecast the mechanics and implications of changes in the population structure of corals that are transitioning from high to low abundance, and from large to small colonies, sometimes without striking effects on planar coral cover. Using two coral reef locations that have been sampled for coral size, we use demographic data to underscore the limitations of coral cover in understanding the causes and consequences of long-term declining coral size, and abundance. A stage-structured matrix model is used to evaluate the demographic causes of declining coral colony size and abundance, particularly with respect to the risks of extinction. The model revealed differential effects of mortality, growth and fecundity on coral size distributions. It also suggested that colony rarity and declining colony size in association with partial tissue mortality and chronic declines in fecundity, can lead to a demographic bottleneck with the potential to prolong the existence of coral populations when they are characterized by mostly very small colonies. Such bottlenecks could have ecological importance if they can delay extinction and provide time for human intervention to alleviate the environmental degradation driving reductions in coral abundance.

The disturbance regimes of ecosystems are changing, and prospects for continued recovery remain unclear. New assemblages with altered species composition may be deficient in key functional traits. Alternatively, important traits may be sustained by species that replace those in decline (response diversity). Here, we quantify the recovery and response diversity of coral assemblages using case studies of disturbance in three locations. Despite return trajectories of coral cover, the original assemblages with diverse functional attributes failed to recover at each location. Response diversity and the reassembly of trait space was limited, and varied according to biogeographic differences in the attributes of dominant, rapidly recovering species. The deficits in recovering assemblages identified here suggest that the return of coral cover cannot assure the reassembly of reef trait diversity, and that shortening intervals between disturbances can limit recovery among functionally important species.

Changes in the size structure of coral populations have major consequences for population dynamics and community function, yet many coral reef monitoring projects do not record this critical feature. Consequently, our understanding of current and future trajectories in coral size structure, and the demographic processes underlying these changes, is still emerging. Here, we provide a conceptual summary of the benefits to be gained from more comprehensive attention to the size of coral colonies in reef monitoring projects, and we support our argument through the use of case-history examples and a simplified ecological model. We neither seek to review the available empirical data, or to rigorously explore causes and implications of changes in coral size, we seek to reveal the advantages to modifying ongoing programs to embrace the information inherent in changing coral colony size. Within this framework, we evaluate and forecast the mechanics and implications of changes in the population structure of corals that are transitioning from high to low abundance, and from large to small colonies, sometimes without striking effects on planar coral cover. Using two coral reef locations that have been sampled for coral size, we use demographic data to underscore the limitations of coral cover in understanding the causes and consequences of long-term declining coral size, and abundance. A stage-structured matrix model is used to evaluate the demographic causes of declining coral colony size and abundance, particularly with respect to the risks of extinction. The model revealed differential effects of mortality, growth and fecundity on coral size distributions. It also suggested that colony rarity and declining colony size in association with partial tissue mortality and chronic declines in fecundity, can lead to a demographic bottleneck with the potential to prolong the existence of coral populations when they are characterized by mostly very small colonies. Such bottlenecks could have ecological importance if they can delay extinction and provide time for human intervention to alleviate the environmental degradation driving reductions in coral abundance.

Corals are major contributors to a range of key ecosystem functions on tropical reefs, including calcification, photosynthesis, nutrient cycling, and the provision of habitat structure. The abundance of corals is declining at multiple scales, and the species composition of assemblages is responding to escalating human pressures, including anthropogenic global warming. An urgent challenge is to understand the functional consequences of these shifts in abundance and composition in different biogeographical contexts. While global patterns of coral species richness are well known, the biogeography of coral functions in provinces and domains with high and low redundancy is poorly understood. Here, we quantify the functional traits of all currently recognized zooxanthellate coral species (n = 821) in both the Indo-Pacific and Atlantic domains to examine the relationships between species richness and the diversity and redundancy of functional trait space. We find that trait diversity is remarkably conserved (>75% of the global total) along latitudinal and longitudinal gradients in species richness, falling away only in species-poor provinces (n < 200), such as the Persian Gulf (52% of the global total), Hawaii (37%), the Caribbean (26%), and the East-Pacific (20%), where redundancy is also diminished. In the more species-poor provinces, large and ecologically important areas of trait space are empty, or occupied by just a few, highly distinctive species. These striking biogeographical differences in redundancy could affect the resilience of critical reef functions and highlight the vulnerability of relatively depauperate, peripheral locations, which are often a low priority for targeted conservation efforts.

Scleractinian corals often exhibit high levels of morphological plasticity, which is potentially important in enabling individual species to occupy benthic spaces across a wide range of environmental gradients. This study tested for differences in the three-dimensional (3D) geometry of three branching corals, Acropora nasuta, Pocillopora spp. and Stylophora pistillata among inner-, mid- and outer-shelf reefs in the central Great Barrier Reef, Australia. Important attributes of coral morphology (e.g., surface area to volume ratio) were expected to vary linearly across the shelf in accordance with marked gradients in environmental conditions, but instead, we detected non-linear trends in the colony structure of A. nasuta and Pocillopora spp. The surface area to volume ratio of both A. nasuta and Pocillopora spp. was highest at mid-shelf locations, (reflecting higher colony complexity) and was significantly lower at both inner-shelf and outer-shelf reefs. The branching structure of these corals was also far more tightly packed at inner-shelf and outer-shelf reefs, compared to mid-shelf reefs. Apparent declines in complexity and inter-branch spacing at inner and outer-shelf reefs (compared to conspecifics from mid-shelf reefs) may reflect changes driven by gradients of sedimentation and hydrodynamics. The generality and explanations of observed patterns warrant further investigation, which is very feasible using the 3D-photogrammetry techniques used in this study.

Mutualisms play a critical role in ecological communities; however, the importance and prevalence of mutualistic associations can be modified by external stressors. On coral reefs, elevated sediment deposition can be a major stressor reducing the health of corals and reef resilience. Here, we investigated the influence of severe sedimentation on the mutualistic relationship between small damselfishes ( Pomacentrus moluccensis and Dascyllus aruanus ) and their coral host ( Pocillopora damicornis ). In an aquarium experiment, corals were exposed to sedimentation rates of approximately 100 mg cm −2 d −1 , with and without fishes present, to test whether: (i) fishes influence the accumulation of sediments on coral hosts, and (ii) fishes moderate partial colony mortality and/or coral tissue condition. Colonies with fishes accumulated much less sediment compared with colonies without fishes, and this effect was strongest for colonies with D. aruanus (fivefold less sediment than controls) as opposed to P. moluccensis (twofold less sediment than controls). Colonies with symbiont fishes also had up to 10-fold less sediment-induced partial mortality, as well as higher chlorophyll and protein concentrations. These results demonstrate that fish mutualisms vary in the strength of their benefits, and indicate that some mutualistic or facilitative interactions might become more important for species health and resilience at high-stress levels.

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.

Sustaining ecological functions as biodiversity changes will be a major challenge in the 21st century [1]. However, our understanding of the relationship between biodiversity and ecosystem function is still emerging on tropical coral reefs [2], where reef-building corals form highly productive assemblages [3, 4] and species respond in different ways to their neighbors [5] and their environment (e.g., water flow) [6]. Experimental coral communities were assembled to quantify the performance of coral colonies with and without neighbors and in the presence of conspecifics versus heterospecifics. Under higher flow, we identified a positive effect of coral species richness on primary productivity (gross and net photosynthesis) indicated by a 53% increase in productivity in multispecies assemblages (2-4 species) relative to monocultures. Productivity in monocultures was predicted by surface areas associated with different species morphologies. In contrast, multispecies assemblages maintained high levels of productivity even in the absence of the most productive species, reflecting non-additive effects of species richness on community functioning. Assemblage performances were regulated by positive and negative interactions between colonies, with many colonies performing better among heterospecific neighbors than in isolation (facilitation). Facilitation occurred primarily among flow-sensitive taxa with simple morphologies and did not occur under lower flow, suggesting that modifications to flow microclimates by corals generated beneficial, interspecific interactions. Our results show that competition and facilitation among neighbors may be important mechanisms regulating coral assemblage productivity in variable environments. Furthermore, shifts in the diversity and identity of neighbors can impair these interactions, with potentially widespread consequences for coral community functioning.

During 2015-2016, record temperatures triggered a pan-tropical episode of coral bleaching, the third global-scale event since mass bleaching was first documented in the 1980s. Here we examine how and why the severity of recurrent major bleaching events has varied at multiple scales, using aerial and underwater surveys of Australian reefs combined with satellite-derived sea surface temperatures. The distinctive geographic footprints of recurrent bleaching on the Great Barrier Reef in 1998, 2002 and 2016 were determined by the spatial pattern of sea temperatures in each year. Water quality and fishing pressure had minimal effect on the unprecedented bleaching in 2016, suggesting that local protection of reefs affords little or no resistance to extreme heat. Similarly, past exposure to bleaching in 1998 and 2002 did not lessen the severity of bleaching in 2016. Consequently, immediate global action to curb future warming is essential to secure a future for coral reefs.