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Reef-building corals are found across > 30° of latitude from tropical to temperate regions, where they occupy habitats greatly differing in seawater temperature and light regimes. It remains largely unknown, however, how the demography of corals differs across this gradient of environmental conditions. Variation in coral growth is especially important to coral populations, because aspects of coral demography are dependent on colony size, with both fecundity and survivorship increasing with larger colonies. Here we tested for latitudinal variation in annual growth rate and survival of juvenile corals, using 11 study locations extending from 17° S to 33° N in the West and South Pacific. Regression analyses revealed a significant decline in annual growth rates with increasing latitude, whereas no significant latitudinal pattern was detected in annual survival. Seawater temperature showed a significant and positive association with annual growth rates. Growth rates varied among the four common genera, allowing them to be ranked Acropora > Pocillopora > Porites > Dipsastraea. Acropora and Pocillopora showed more variation in growth rates across latitudes than Porites and Dipsastraea. Although the present data have limitations with regard to difference in depths, survey periods, and replication among locations, they provide evidence that a higher capacity for growth of individual colonies may facilitate population growth, and hence population recovery following disturbances, at lower latitudes. These trends are likely to be best developed in Acropora and Pocillopora, which have high rates of colony growth.

Within the past decade, a functional group of encrusting red macroalgae defined as peyssonnelid algal crusts (PAC) has been rapidly spreading on shallow Caribbean reefs, frequently encrusting corals and sponges. This study focused on PAC growing on the reefs of St. John, US Virgin Islands, to (1) expand the monitoring of PAC abundance in this location, and (2) test two mechanisms, rapid linear growth and competitive dominance, by which PAC is hypothesized to enhance its capacity to spread. In August 2019, we found that the percentage coverage of PAC reached 46.6% at Tektite (3 m depth), and 63.8% at Cabritte Horn (3 m depth). Overall, the cover of PAC at these sites in 2019 was identical to, or up to 2.4-fold higher than in August 2017, and 2.1–3.7-fold higher than in November 2017, just after two category 5 storms impacted St. John. PAC was 1.8–29.1-fold more abundant at 3 m versus 9 m depth, and 1.4–22.0-fold more abundant at sites close to the tips of headlands than sites sheltered within bays. From August 2019 to January 2020, the rate of linear extension of PAC at each site and depth ranged from 26 to 43 µm d − 1, and of 80 PAC-coral interactions that were tagged in August 2019, none involved corals that had become completely overgrown by PAC by January 2020. The slow linear growth and absence of cases in which PAC completely overgrew coral colonies in 6 months are inconsistent with the high rate at which PAC has occupied space on Caribbean reefs over the past decade. One means to reconcile these observations is the possibility that recruitment of peyssonnelid spores to benthic surfaces facilitates the rapid spread of PAC.

Increasing abundance of arborescent octocorals (often referred to as gorgonians) on Caribbean reefs raises the question of whether habitat structure provided by octocorals can mediate a transition between coral- and algal- dominated states by increasing fish abundance and herbivory. This study tested the hypotheses that feeding rates and densities of demersal reef fishes are affected by the habitat structure provided by dense octocoral communities. Surveys of fishes on coral reefs in St John, US Virgin Islands, found 1.7-fold higher densities, and 2.4-fold higher feeding rates within versus outside of dense octocoral canopies. This difference, however, was only seen at sites with octocoral densities > 8 colonies m−2. Furthemore, the proximity of octocoral colonies to fish had an effect on the grazing rate of key herbivores (surgeonfishes and parrotfishes), with a 53% higher feeding rate (1.90 ± 0.11 bites min−1 m−2) near octocorals (

Coral reefs have long attracted attention because of their biological and economic importance, but this interest now has turned to examining the possibility of functional extirpation. Widespread declines in coral abundances have fueled the shift in motivation for studying reefs and catalyzed the proliferation of monitoring to record the changes underway. Despite appreciation of monitoring as a scientific endeavor, its primary use has continued to be the quantification of cover of coral, macroalgae, and a few other space holders. The limitations of coral cover in evaluating the consequences of changing coral abundance were highlighted decades ago. Yet neglect of the tools most appropriate for this task (demographic approaches) and continuing emphasis on a tool (coral cover) that is not ideal, indicates that these limitations are not widely appreciated. Reef monitoring therefore continues to underperform with respect to its potential, thus depriving scientists of the approaches necessary to project the fate of coral reefs and test hypotheses focused on the proximal causes of declining coral cover. We make the case that the coral reef crisis creates a need for coral demography that is more acute now than 4 decades ago. Modern demographic approaches are well suited to meet this need, but to realize their potential, consideration will need to be given to the possibility of expanding ecological monitoring of coral reefs to provide the data necessary for demographic analyses of their foundation taxon, the Scleractinia.

Recruitment hotspots are locations where organisms are added to populations at high rates. On tropical reefs where coral abundance has declined, recruitment hotspots are important because they have the potential to promote population recovery. Around St. John, US Virgin Islands, coral recruitment at five sites revealed a hotspot that has persistent for 14 years. Recruitment created a hotspot in density of juvenile corals that was 600 m southeast of the recruitment hotspot. Neither hotspot led to increased coral cover, thus revealing the stringency of the demographic bottleneck impeding progression of recruits to adult sizes and preventing population growth. Recruitment hotspots in low-density coral populations are valuable targets for conservation and sources of corals for restoration.

Ocean acidification (OA) is negatively affecting calcification in a wide variety of marine organisms. These effects are acute for many tropical scleractinian corals under short-term experimental conditions, but it is unclear how these effects interact with ecological processes, such as competition for space, to impact coral communities over multiple years. This study sought to test the use of individual-based models (IBMs) as a tool to scale up the effects of OA recorded in short-term studies to community-scale impacts, combining data from field surveys and mesocosm experiments to parameterize an IBM of coral community recovery on the fore reef of Moorea, French Polynesia. Focusing on the dominant coral genera from the fore reef, Pocillopora, Acropora, Montipora and Porites, model efficacy first was evaluated through the comparison of simulated and empirical dynamics from 2010–2016, when the reef was recovering from sequential acute disturbances (a crown-of-thorns seastar outbreak followed by a cyclone) that reduced coral cover to ~0% by 2010. The model then was used to evaluate how the effects of OA (1,100–1,200 µatm pCO2) on coral growth and competition among corals affected recovery rates (as assessed by changes in % cover y−1) of each coral population between 2010–2016. The model indicated that recovery rates for the fore reef community was halved by OA over 7 years, with cover increasing at 11% y−1 under ambient conditions and 4.8% y−1 under OA conditions. However, when OA was implemented to affect coral growth and not competition among corals, coral community recovery increased to 7.2% y−1, highlighting mechanisms other than growth suppression (i.e., competition), through which OA can impact recovery. Our study reveals the potential for IBMs to assess the impacts of OA on coral communities at temporal and spatial scales beyond the capabilities of experimental studies, but this potential will not be realized unless empirical analyses address a wider variety of response variables representing ecological, physiological and functional domains.

The biological world is rapidly changing following decades of anthropogenic disturbances. Under these conditions, species with stable or increasing abundances have been described as winners with the potential for future success, but this assertion is unreliable without knowledge of the selective basis of winning. The incentive to find winners is acute for reef corals, for which large declines in abundance have motivated restoration efforts targeting winning corals. On Caribbean reefs, Porites astreoides has emerged as a potential winner, but the demographic basis of this categorization is poorly known. Here we test for demographic benchmarks of winning in this species by quantifying abundances and sizes of colonies over 28 yr on the south coast of St. John, US Virgin Islands. From 1992 to 2001, the density of colonies of P. astreoides showed little variation while colonies increased in size by 58%, but from 2002 to 2019, population density increased 2.7‐fold, and colony size declined by 41%; accompanying these trends, the mean absolute cover of Porites spp. declined by 46% from 2010 to 2019. Low recruitment and rising abundances of colonies ≤4 cm diameter suggest that partial mortality and fission depressed colony sizes. The reversal over three decades of a positive demographic trend for a ubiquitous coral underscores the challenges of identifying winners from short‐term population performance. Without a mechanistic understanding of fitness, the search for winners based on demographic trends may be futile.

In recent decades, many Caribbean reefs have experienced large declines in abundance of scleractinian corals, and blooms of fleshy macroalgae have often accompanied these trends. In 2010 a new macroalgal threat emerged in Lac Bay, Bonaire, where peyssonnelid algal crusts (PAC) rapidly spread in shallow water and overgrew corals and sponges. Similar growths have been reported in Puerto Rico and the Virgin Islands, and here, we describe the spread of PAC on the shallow reefs of St. John, US Virgin Islands, ~ 760 km northeast of Bonaire. In 2015, PAC covered 0.2–20.6% of hard benthic surfaces at ≤ 7 m depth. By August 2017, at the same sites, PAC had increased to 3.2–61.0% cover, at 5 m depth at 10 other sites along 5 km of shore, it covered 1.4–61.6% of reef surfaces, and at 9 m depth at five sites, it covered 0.8–41.0% of reef surfaces. At 5 m depth in August 2017, scleractinians and octocorals were frequently contacting PAC (42–47% of colonies), and more scleractinians (74%) than octocorals (39%) were overgrown by PAC. Following surveys in August 2017, St. John was hit by two Category 5 hurricanes, yet the shore-wide mean cover of PAC at 5 m depth was only reduced from 26 to 23%. Our results suggest PAC is poised to cause significant ecological change on the reefs of St. John and potentially will promote a community shift favoring octocorals over scleractinians. PAC constitutes an emerging regional threat on shallow Caribbean reefs to which researchers and resource managers will need to quickly respond.

Variation among functionally similar species in their response to environmental stress buffers ecosystems from changing states. Functionally similar species may often be cryptic species representing evolutionarily distinct genetic lineages that are morphologically indistinguishable. However, the extent to which cryptic species differ in their response to stress, and could therefore provide a source of response diversity, remains unclear because they are often not identified or are assumed to be ecologically equivalent. Here, we uncover differences in the bleaching response between sympatric cryptic species of the common Indo‐Pacific coral, Pocillopora. In April 2019, prolonged ocean heating occurred at Moorea, French Polynesia. 72% of pocilloporid colonies bleached after 22 d of severe heating (>8oC‐days) at 10 m depth on the north shore fore reef. Colony mortality ranged from 11% to 42% around the island four months after heating subsided. The majority (86%) of pocilloporids that died from bleaching belonged to a single haplotype, despite twelve haplotypes, representing at least five species, being sampled. Mitochondrial (open reading frame) sequence variation was greater between the haplotypes that experienced mortality versus haplotypes that all survived than it was between nominal species that all survived. Colonies > 30 cm in diameter were identified as the haplotype experiencing the most mortality, and in 1125 colonies that were not genetically identified, bleaching and mortality increased with colony size. Mortality did not increase with colony size within the haplotype suffering the highest mortality, suggesting that size‐dependent bleaching and mortality at the genus level was caused instead by differences among cryptic species. The relative abundance of haplotypes shifted between February and August, driven by declines in the same common haplotype for which mortality was estimated directly, at sites where heat accumulation was greatest, and where larger colony sizes occurred. The identification of morphologically indistinguishable species that differ in their response to thermal stress, but share a similar ecological function in terms of maintaining a coral‐dominated state, has important consequences for uncovering response diversity that drives resilience, especially in systems with low or declining functional diversity.

Since about the turn of the millennium, octocorals have been increasing in abundance on Caribbean reefs. The mechanisms underlying this trend have not been resolved, but the emergent species assemblage appears to be more resilient than the scleractinians they are replacing. The sea fan Gorgonia ventalina is an iconic species in the contemporary octocoral fauna, and here its population dynamics are described from St. John, US Virgin Islands, from 2013 to 2019. Mean densities of G. ventalina at Yawzi Point (9-m depth) varied from 1.4–1.5 colonies m−2, and their mean heights from 24–30 cm; nearby at Tektite (14-m depth), they varied from 0.6–0.8 colonies m−2 and from 25–33 cm. These reefs were impacted by two Category 5 hurricanes in 2017, but neither the density of G. ventalina, the density of their recruits (< 5-cm tall), nor the height of colonies, differed among years, although growth was depressed after the hurricanes. Nevertheless, at Tektite, colony height trended upwards over time, in part because colonies 10.1–20 cm tall were reduced in abundance after the hurricanes. These trends were sustained without density-associated effects mediating recruitment or self-thinning of adults. The dynamics of G. ventalina over seven years reveals the high resilience of this species that will contribute to the persistence of octocorals as a dominant state on Caribbean reefs.

This study tested the hypothesis that sponge assemblages on the reefs of St. John, US Virgin Islands (18.315°N, 64.716°W), changed from 1992 to 2017. Sponges were identified to species or genus in photoquadrats and were quantified at 2–3 y intervals by density, with linear dimensions used to estimate volume as a proxy for biomass. From 1992 to July 2017, overall sponge density (pooled among taxa) increased, although trends varied among the most common species for which density increased (four species), decreased (one species), or showed no linear relationship with time (2 species). Hurricanes Irma and Maria caused a 51% decline in overall sponge density from July to November 2017, with density in November 2017 being 31% lower than in 1992. For the common sponges, densities of 6 declined by 82–100% between July and November 2017, but densities in November 2017 were 13–736% higher than in 1992 for four species, and 47–100% lower than in 1992 for 3 species. Sponge volume did not change linearly between 1992 and November 2017, but it was reduced by 54% following the hurricanes. Multivariate variation over time in benthic community structure was not strongly associated with sponge density alone, although a significant association with rainfall highlighted the potential roles of this environmental factor in directly, or indirectly, mediating changes in community structure. Overall, the mean density of sponges, as well as most of the common sponge species, trended upwards prior to the hurricanes, but these trends reversed following the storms.

One response to the coral reef crisis has been human intervention to enhance selection on the fittest corals through cultivation. This requires genotypes to be identified for intervention, with a primary basis for this choice being growth: corals that quickly grow on contemporary reefs might be future winners. To test for temporal stability of growth as a predictor of future performance, genotypes of the coral Porites spp. were grown in common gardens in Mo'orea, French Polynesia. Growth was measured every two to four months throughout 2018, and each period was used as a predictor of growth over the subsequent period. Area-normalized growth explained less than 29% of the variance in subsequent growth, but for biomass-normalized growth this increased to 45–60%, and was highest when summer growth was used to predict autumn growth. The capacity of initial growth to predict future performance is dependent on the units of measurement and the time of year in which it is measured. The final choice of traits to quantify performance must be informed through consideration of the species and the normalization that best capture the information inherent in the biological processes mediating variation in traits values.

Reef-building corals, like many long-lived organisms, experience environmental change as a combination of separate but concurrent processes, some of which are gradual yet long-lasting, while others are more acute but short-lived. For corals, some chronic environmental stressors, such as rising temperature and ocean acidification, are thought to induce gradual changes in colonies' vital rates. Meanwhile, other environmental changes, such as the intensification of tropical cyclones, change the disturbance regime that corals experience. Here, we use a physiologically structured population model to explore how chronic environmental stressors that impact the vital rates of individual coral colonies interact with the intensity and magnitude of disturbance to affect coral population dynamics and cover. We find that when disturbances are relatively benign, intraspecific density dependence driven by space competition partially buffers coral populations against gradual changes in vital rates. However, the impact of chronic stressors is amplified in more highly disturbed environments, because disturbance weakens the buffering effect of space competition. We also show that coral cover is more sensitive to changes in colony growth and mortality than to external recruitment, at least in open populations, and that space competition and size structure mediate the extent and pace of coral-population recovery following a large-scale mortality event. Understanding the complex interplay among chronic environmental stressors, mass-mortality events, and population size structure sharpens our ability to manage and to restore coral-reef ecosystems in an increasingly disturbed future.

Several species of crustose coralline algae (CCA) and their associated microbial biofilms play important roles in determining the settlement location of scleractinian corals on tropical reefs. In recent decades, peyssonnelid algal crusts (PAC) have become spatial dominants across large areas of shallow Caribbean reefs, where they appear to deter the recruitment of scleractinians. Our genetic investigations of PAC in St. John, US Virgin Islands, amplifying the large-subunit ribosomal RNA and psbA protein D1 marker genes, revealed them to be identical to Ramicrusta textilis previously reported overgrowing corals in Jamaica. Specimens of PAC sampled from the Honduras were likewise identical, confirming that this crustose alga inhabits the easternmost and westernmost regions of the Caribbean. We also analysed 16S rDNA tag amplicon libraries of the biofilms associated with PAC and sympatric CCA, which is favoured for coral settlement. Our results show that the microbial communities on PAC (vs. CCA) are characterized by significantly lower numbers of the epibiotic bacterial genus Pseudoalteromonas, which facilitates the recruitment and settlement of marine invertebrates. From these data, we infer that PAC are therefore unlikely to be attractive as settlement sites for coral larvae. Given the significant ecological change anticipated on these reefs due to increasing cover of PAC, there is an urgent need to further investigate competitive interactions between PAC and scleractinian corals, and elucidate the role of PAC and their associated microbiomes in accentuating phase shifts from coral to algae on tropical reefs.

Reef-building corals, like many long-lived organisms, experience environmental change as a combination of separate but concurrent processes, some of which are gradual yet long-lasting, while others are more acute but short-lived. For corals, some chronic environmental stressors, such as rising temperature and ocean acidification, are thought to induce gradual changes in colonies’ vital rates. Meanwhile, other environmental changes, such as the intensification of tropical cyclones, change the disturbance regime that corals experience. Here, we use a physiologically structured population model to explore how chronic environmental stressors that impact the vital rates of individual coral colonies interact with the intensity and magnitude of disturbance to affect coral population dynamics and cover. We find that when disturbances are relatively benign, intraspecific density dependence driven by space competition partially buffers coral populations against gradual changes in vital rates. However, the impact of chronic stressors is amplified in more highly disturbed environments, because disturbance weakens the buffering effect of space competition. We also show that coral cover is more sensitive to changes in colony growth and mortality than to external recruitment, at least in non-self-seeding populations, and that space competition and size structure mediate the extent and pace of coral-population recovery following a large-scale mortality event. Understanding the complex interplay among chronic environmental stressors, mass-mortality events, and population size structure sharpens our ability to project coral dynamics in an increasingly disturbed future.

Coral reefs throughout the tropics have experienced large declines in the abundance of scleractinian corals over the last few decades, and some reefs are becoming functionally dominated by animal taxa other than scleractinians. This phenomenon is striking on many shallow reefs in the tropical western Atlantic, where arborescent octocorals now are numerically and functionally dominant. Octocorals are one of several taxa that have been overlooked for decades in analyses of coral reef community dynamics, and our understanding of why octocorals are favoured (whereas scleractinians are not) on some modern reefs, and how they will affect the function of future reef communities, is not commensurate with the task of scientifically responding to the coral reef crisis. We summarize the biological and ecological features predisposing octocorals for success under contemporary conditions, and focus on those features that could have generated resistance and resilience of octocoral populations to environmental change on modern reefs. There is a rich set of opportunities for rapid advancement in understanding the factors driving the success of octocorals on modern reefs, but we underscore three lines of inquiry: (1) the functional implications of strongly mixotrophic, polytrophic, and plastic nutrition, (2) the capacity to recruit at high densities and maintain rapid initial rates of vertical growth, and (3) the emergent properties associated with dense animal forests at high colony densities.

Experiments with coral fragments (i.e., nubbins) have shown that net calcification is depressed by elevated PCO2. Evaluating the implications of this finding requires scaling of results from nubbins to colonies, yet the experiments to codify this process have not been completed. Building from our previous research demonstrating that net calcification of Pocillopora verrucosa (2–13 cm diameter) was unaffected by PCO2 (400 µtam and 1,000 µatm) and temperature (26.5°C and 29.7°C) we sought generality to this outcome by testing how colony size modulates PCO2 and temperature sensitivity in a branching acroporid. Together these taxa represent two of the dominant lineages of branching corals on Indo-Pacific coral reefs. Two trials conducted over two years tested the hypothesis that the seasonal range in seawater temperature (26.5°C and 29.2°C) and a future PCO2 (1,062 µatm versus an ambient of 461 µatm) affect net calcification of an ecologically relevant size range (5–20 cm diameter) of colonies of Acropora hyacinthus. Like P. verrucosa, the effects of temperature and PCO2 on net calcification (mg d−1) of A. verrucosa were not statistically detectable. These results support the generality of a null outcome on net calcification of exposing intact colonies of branching corals to environmental conditions contrasting seasonal variation in temperature and predicted future variation in PCO2. While there is a need to expand beyond an experimental culture relying on coral nubbins as tractable replicates, rigorously responding to this need poses substantial ethical and logistical challenges.

Invertebrate ectosymbionts within the coralla of scleractinians enhance host fitness through protection from corallivores and nutrient addition. Here, we explore the ectosymbiotic relationship between the coral Pocillopora verrucosa and the crab Trapezia serenei and the shrimp Alpheus spp., to test for effects on coral calcification under contrasts of seawater temperature (27.7 °C and 29.9 °C) and pH (ambient, 8.0 and reduced, 7.7). Regardless of temperature, ectosymbionts depressed calcification by 55% (vs without ectosymbionts) at ambient pH; however, ectosymbionts only depressed calcification under ambient pH but not at reduced pH. These results suggest that P. verrucosa grows fastest at ambient pH without ectosymbionts, but when ectosymbionts are present, colonies are protected from further declines in calcification at reduced pH. This implies that there may be a change from a currently parasitic ectosymbiont–coral relationship to a commensal relationship that could increase fitness advantages for corals hosting crustacean ectosymbionts under ocean acidification conditions.

An important goal of coral reef science is to understand the roles played by environmental conditions in determining benthic community structure. Pursuit of this goal typically involves testing for associations between community structure and environmental conditions, and in recent years, attention has focused on temperature and seawater pH. Such analyses are helpful in projecting coral reef community structure into the future, but their efficacy depends on the extent to which changes in community structure are correctly associated with causative processes, and whether future conditions are a subset of those experienced in the past. Here, we describe variation from 2014 to 2017 in one environmental factor, photosynthetically active radiation (PAR), at 19-m depth on the reefs of St. John, US Virgin Islands, and evaluate the extent to which this factor could cause changes in benthic community structure. The transmission of downwelling surface PAR to 19-m depth (TPAR-19) varied from 2.0 to 40.0% among days, and from 4.8 to 15.8% among months within a year, and was lower and more variable in the autumn versus other seasons. Monthly TPAR-19 was inversely associated with monthly rainfall, suggesting that runoff affected seawater clarity. The monthly diffuse attenuation coefficient (Kd-PAR) varied 1.7-fold from 0.097 to 0.164 m−1 and was within the range expected for coral reefs. Given the high temporal variation in underwater light intensity, and its importance to foundational reef taxa, understanding of the response of coral reefs to temperature and pH (and other factors) will remain incomplete without closer attention to this physical environmental condition.

In marine benthic habitats, structural complexity is positively associated with diversity, and the area of hard surfaces modulates community structure. These principles are well developed on tropical coral reefs, where stony corals create complex surfaces that are favored for settlement. This study explores the relationship between substratum orientation and the distribution of arborescent octocorals on shallow reefs (

Ocean acidification is expected to affect coral reefs in multiple ways, in part, by depressing the calcification of scleractinian corals. To evaluate how coral communities will respond to ocean acidification, research into the effects on ecological processes determining community structure is now needed. The present study focused on corals utilizing soft tissues (i.e., mesenterial filaments) as agonistic mechanism, and evaluated their ability to compete for space under ocean acidification. Using aquarium-reared specimens in Monaco, single polyps of Galaxea fascicularis were paired with branch tips of Acropora hyacinthus to stimulate competitive interactions, which were evaluated through the production and use of mesenterial filaments in causing tissue damage under ambient (~ 600 μatm) and elevated pCO (~ 1200 μatm). At 1200 μatm pCO, and when paired with A. hyacinthus, the extrusion of mesenterial filaments from G. fascicularis occurred 2 days earlier than under ambient pCO, although ultimately the mesenterial filaments caused the same amount of tissue necrosis on A. hyacinthus under both pCO regimes after 7 days. This outcome supports the hypothesis that some kinds of competitive mechanisms utilized by scleractinian corals (i.e., mesenterial filaments) will be unaffected by short exposure to pCO as high as 1200 μatm.

Metabolic scaling is the relationship between organismal metabolic rate and body mass. Understanding the patterns and causes of metabolic scaling provides a powerful foundation for predicting biological processes at the level of individuals, populations, communities, and ecosystems. Despite intense interest in, and debate on, the mechanistic basis of metabolic scaling, relatively little attention has been paid to metabolic scaling in clonal animals with modular construction, such as colonial cnidarians, bryozoans, and colonial ascidians. Unlike unitary animals, modular animals are structural individuals subdivided into repeated morphological units, or modules, each able to acquire, process, and share resources. A modular design allows flexibility in organism size and shape with consequences for metabolic scaling. Furthermore, with careful consideration of the biology of modular animals, the size and shape of individual colonies can be experimentally manipulated to test competing theories pertaining to metabolic scaling. Here, we review metabolic scaling in modular animals and find that a wide range of scaling exponents, rather than a single value, has been reported for a variety of modular animals. We identify factors influencing variation in intraspecific scaling in this group that relate to the general observation that not all modules within a colony are identical. We highlight current gaps in our understanding of metabolic scaling in modular animals, and suggest future research directions, such as manipulating metabolic states and comparisons among species that differ in extent of module integration.

Peyssonnelid algal crusts (PAC) have rapidly spread on coral reefs throughout the Caribbean since 2010 and have become dominant space holders in multiple locations. In 2019, PAC covered 31–86% of the shallow reefs ( Porites spp., Siderastrea radians, and S. siderea. As Diadema halos remained in the same location for at least 6 months, they may function as refuges for coral recruitment in locations where PAC is becoming spatially dominant.

Ocean acidification (OA), attributed to the sequestration of atmospheric carbon dioxide (CO 2 ) into the surface ocean, and coastal eutrophication, attributed in part to land-use change and terrestrial runoff of fertilizers, have received recent attention in an experimental framework examining the effects of each on coral reef net ecosystem calcification (G net ). However, OA and eutrophication in conjunction have yet to receive attention from the perspective of coral reef sediment dissolution. To address this omission, CO 2 and nitrate (NO 3 − ) addition experiments were performed in Mo'orea, French Polynesia. Incubation chambers were used to measure sediment G net during the day and night under three different [NO 3 − ] (0, 9.8, and 19.7 μM) that were nested within four separate constructed coral reef communities maintained at different P CO2 levels (417, 721, 1030, and 1333 μatm, respectively). P CO2 negatively affected sediment G net during the day and night, resulting in a shift to diel net dissolution at a P CO2 of 1030 μatm. Elevated NO 3 − alone, and the combination of NO 3 − and P CO2 , both negatively affected sediment G net at night. However, the response of G net to NO 3 − was less clear during the day, where diurnal sediment G net was enhanced under the combined treatment of elevated NO 3 − and P CO2 , resulting in no net effect of NO 3 − on sediment G net on diel timescales. Overall, these results show that ocean acidification represents a greater threat to the balance of calcification and dissolution in Mo'orea's back reef sediment communities than the potential impact of NO 3 − enrichment on relatively short timescales.

The threat represented by ocean acidification (OA) for coral reefs has received considerable attention because of the sensitivity of calcifiers to changing seawater carbonate chemistry. However, most studies have focused on the organismic response of calcification to OA, and only a few have addressed community-level effects, or investigated parameters other than calcification, such as photosynthesis. Light (photosynthetically active radiation, PAR) is a driver of biological processes on coral reefs, and the possibility that these processes might be perturbed by OA has important implications for community function. Here we investigate how CO2 enrichment affects the relationships between PAR and community net O2 production (Pnet), and between PAR and community net calcification (Gnet), using experiments on three coral communities constructed to match (i) the back reef of Mo'orea, French Polynesia, (ii) the fore reef of Mo'orea, and (iii) the back reef of O'ahu, Hawaii. The results were used to test the hypothesis that OA affects the relationship between Pnet and Gnet. For the three communities tested, pCO2 did not affect the Pnet–PAR relationship, but it affected the intercept of the hyperbolic tangent curve fitting the Gnet–PAR relationship for both reef communities in Mo'orea (but not in O'ahu). For the three communities, the slopes of the linear relationships between Pnet and Gnet were not affected by OA, although the intercepts were depressed by the inhibitory effect of high pCO2 on Gnet. Our result indicates that OA can modify the balance between net calcification and net photosynthesis of reef communities by depressing community calcification, but without affecting community photosynthesis.

This study tested the bleaching response of the Pacific coral Seriatopora caliendrum to short-term exposure to high temperature and elevated partial pressure of carbon dioxide (pCO2). Juvenile colonies collected from Nanwan Bay, Taiwan, were used in a factorial experimental design in which 2 temperatures (∼27.6 °C and ∼30.4 °C) and 2 pCO2 values (∼47.2 Pa and ∼90.7 Pa) were crossed to evaluate, over 12 days, the effects on the densities and physiology of the symbiotic dinoflagellates (Symbiodinium) in the corals. Thermal bleaching, as defined by a reduction of Symbiodinium densities at high temperature, was unaffected by high pCO2. The division, or mitotic index (MI), of Symbiodinium remaining in thermally bleached corals was about 35% lower than in control colonies, but they contained about 53% more chlorophyll. Bleaching was highly variable among colonies, but the differences were unrelated to MI or pigment content of Symbiodinium remaining in the coral host. At the end of the study, all of the ...

This study tested the hypothesis that intraspecific morphological plasticity within a scleractinian coral elicits differential responses to elevated PCO2 and temperature. In Mo'orea, French Polynesia, two short-term laboratory experiments (21 and 14 days) were conducted to test the effects of PCO2 (400 vs. 700 μatm), and PCO2 (400 vs 1000 μatm) combined with temperature (27.0 vs. 29.8 °C), on branches and plates of Porites rus. Experiments employed two irradiances (~ 1000 vs 200 μmol photons m− 2 s− 1), which characterized the microenvironments on the shallow fringing reefs where branching and plating morphologies are common, respectively. Calcification of both morphologies was insensitive to PCO2, as well as the combined effects of elevated PCO2 and temperature. Mean calcification rates were faster in high light than in low light for both morphologies, and biomass was greater in plates than branches in all treatments. Together, our results suggest P. rus is robust to increased PCO2 and high temperature within the constraints of the treatments applied. Morphological plasticity in this species does not mediate physiological resistance to low pH and high temperature.

Scleractinian corals on tropical reefs are exposed to many natural and anthropogenic disturbances, and while much is known about their responses to such conditions, it is unclear whether the responses will remain the same in a future affected by climate change and ocean acidification. To evaluate how one aspect of these effects — wound repair — might be influenced by ocean acidification, small colonies of massive Porites spp. from the back reef of Moorea, French Polynesia, were damaged to simulate the effects of single bites by corallivorous fishes, and healing was measured under contrasting P CO2 regimes. Using experiments lasting 19–20 d and employing superficial (2013) or deep (2014) lesions, the effects of damage were evaluated at ~ 400 μatm (ambient) and ~ 1000 μatm P CO2 (both at ~ 28.5 °C) using calcification and healing as dependent variables. Damage reduced calcification of massive Porites spp. in 2013, but not in 2014, and P CO2 had no effect on area-normalized calcification or healing in both years, although biomass-normalized calcification was reduced by high P CO2 in 2014. Overall these results reveal the physiological resilience of this functional group of corals to the extent of ocean acidification expected by the end of this century, and suggest that over this period small colonies will remain capable of recovery from minor damage arising from fish corallivory.

On most tropical coral reefs, decades of disturbances have ratcheted down coral cover to create low abundance communities. In such a state, the reefs of St. John, US Virgin Islands, were hit by two Category 5 hurricanes in September 2017, yet the effects on two sites dominated by Orbicella annularis were minor in terms of coral cover. To explore the implications of this outcome, the fates of O. annularis colonies were determined from photoquadrats and used to prepare size‐based matrix models for the year preceding the storms and the four months bracketing the storms. The populations displayed contrasting dynamics from 1988 to July 2017, with coral cover declining from 43% to 5% at Yawzi Point but remaining at ~30% at Tektite. Over this period, colony sizes declined, with ≥82% having planar areas ≤50 cm2 (i.e., the smallest size class) by July 2017, and while densities declined from 47 to 8 colonies/m2 at Yawzi Point, they increased from 36 to 51 colonies/m2 at Tektite. Hurricanes Irma and Maria depressed coral cover by 1–4%, transitioned colonies into the smallest size class (>87% by November), killed 27% and 5% of the colonies in the smallest size class at Yawzi Point and Tektite, respectively, and depressed the 5‐yr intrinsic rate of population growth (λ) to 0.53–0.87. Twenty‐year projections suggested these demographic effects will not have ecologically meaningful impacts on population size, at least compared to projections initiated assuming Hurricanes Irma and Maria had not occurred. With low cover of O. annularis distributed among many small colonies, future disturbances may play more important roles in winnowing the few remaining host genotypes rather than further depressing coral cover.

Coral reefs are threatened by ocean acidification (OA), which depresses net calcification of corals, calcified algae, and coral reef communities. These effects have been quantified for many organisms, but most experiments last weeks-to-months, and do not test for effects on community structure. Here, the effects of OA on back reef communities from Mo’orea, French Polynesia (17.492 S, 149.826 W), were tested from 12 November 2015 to 16 November 2016 in outdoor flumes maintained at mean pCO2 levels of 364 µatm, 564 µatm, 761 µatm, and 1067 µatm. The communities consisted of four corals and two calcified algae, with change in mass (Gnet, a combination of gross accretion and dissolution) and percent cover recorded monthly. For massive Porites and Montipora spp., Gnet differed among treatments, and at 1067 µatm (relative to ambient) was reduced and still positive; for Porolithon onkodes, all of which died, Gnet was negative at high pCO2, revealing dissolution (sample sizes were too small for analysis of Gnet for other taxa). Growth rates (% cover month−1) were unaffected by pCO2 for Montipora spp., P. rus, Pocillopora verrucosa, and Lithophyllum kotschyanum, but were depressed for massive Porites at 564 µatm. Multivariate community structure changed among seasons, and the variation under all elevated pCO2 treatments differed from that recorded at 364 µatm, and was greatest under 564 µatm and 761 µatm pCO2. Temporal variation in multivariate community structure could not be attributed solely to the effects of OA on the chemical and physical properties of seawater. Together, these results suggest that coral reef community structure may be more resilient to OA than suggested by the negative effects of high pCO2 on Gnet of their component organisms.

Some of the best-known disturbances affecting coral reefs are storms, yet their impacts on light are poorly known. Here, we describe the underwater light on a reef off St. John, US Virgin Islands (18°18′37.04N, 63°43′23.17W), during two hurricanes and multiple tropical waves that occurred between 17 August 2017 and 30 November 2017. Photosynthetically active radiation (PAR) was recorded at 19-m depth and at the surface, and rainfall was measured as a cause of turbidity affecting underwater light. Hurricanes Irma and Maria reduced maximum daily underwater photosynthetic photon flux density (PPFD) to

There was an error published in J. Exp. Biol. (2016) 219 , [3896-3906][1] ([doi:10.1242/jeb.146381][2]). In Table S1, some of the values were incorrectly assigned to tank treatments and were averaged over an incorrect period that did not correspond to the number of days from the beginning to the

We quantified benthic community structure on shallow (10 m isobath) reefs separated by 3–130 km on four islands in the south Pacific, and evaluated the roles of disturbances vs. coral recruitment as causes of spatial heterogeneity. Reefs were surveyed in 2013 on Moorea, Tahiti, Tetiaroa, and Maiao, with community structure sampled at two sites on each island using photoquadrats. The effects of coral recruitment on population structure were evaluated through genetic analyses of Pocillopora on three of the islands. Benthic community structure with functional group resolution differed among islands and generally among sites, but coral community structure (generic resolution) differed among islands, but generally not among sites. Genetic analyses of Pocillopora using the open reading frame of host mtDNA revealed varying relative abundances of Pocillopora meandrina/Pocillopora eydouxi, Pocillopora verrucosa, Pocillopora effuses, and two unnamed haplotypes on each island. These results suggest that corals on each island represent unique samplings of genetically discrete larval assemblages rather than random samplings of a single larval assemblage. Together, our findings emphasize the extent to which coral community structure varies over a scale of

The effects of ocean acidification (OA) on coral reefs have been studied thoroughly with a focus on the response of calcification of corals and calcified algae. However, there are still large gaps in our knowledge of the effects of OA on photosynthesis and respiration of these organisms. Comparisons among species and determination of the functional relationships between pCO2 and either photosynthesis or respiration are difficult using previously published data, because experimental conditions typically vary widely between studies. Here, we tested the response of net photosynthesis, gross photosynthesis, dark respiration, and light-enhanced dark respiration (LEDR) of eight coral taxa and seven calcified alga taxa to six different pCO2 levels (from 280 to 2000 µatm). Organisms were maintained during 7–10 days incubations in identical conditions of light, temperature, and pCO2 to facilitate comparisons among species. Net photosynthesis was not affected by pCO2 in seven of eight corals or any of the algae; gross photosynthesis did not respond to pCO2 in six coral taxa and six algal taxa; dark respiration also was unaffected by pCO2 in six coral and six algae; and LEDR did not respond to pCO2 in any of the tested species. Overall, our results show that pCO2 levels up to 2000 µatm likely will not fertilize photosynthesis or modify respiration rates of most of the main calcifiers on the back reef of Moorea, French Polynesia.

We tested the hypothesis that ocean acidification (OA) affects spatial competition among scleractinian corals. Competitive ability was evaluated indirectly by linear extension of Porites lutea and Montipora aequituberculata placed in intraspecific, interspecific, and control pairings (paired with dead coral skeleton) and exposed to ambient (400 μatm) and elevated (1000 μatm) pCO in experiments conducted in Moorea, French Polynesia, and Okinawa, Japan. High pCO had no effect on linear extension of M. aequituberculata in Moorea, but in Okinawa, it reduced linear extension 37%; high pCO had no significant effect on linear extension of P. lutea in Okinawa. Much of the negative effect of high pCO on linear extension for M. aequituberculata in Okinawa was due to reduced extension in control pairings, with corals engaged in intra- and interspecific competition unaffected by OA. Linear extension of M. aequituberculata and P. lutea in interspecific pairings decreased relative to control pairings at ambient pCO by 39 and 71%, respectively, indicating a strong effect of competition on extension rates. These differences however disappeared at elevated pCO when the linear extension of controls was depressed. Together, our results show that OA can negatively affect the linear extension of corals not engaged in competition, as shown in the control pairings, and suggest that OA does not directly affect the ability of corals to compete with one another for space.

To evaluate the possibility that density-associated effects modulate octocoral abundance on a Caribbean coral reef, we tested the hypothesis that the density of octocoral recruits (colonies ≤ 4 cm tall) and adult colonies are positively associated on shallow reefs (≤ 14 m depth) in St. John, US Virgin Islands. Both life stages were censused for density at 8–10 sites along 7 km of shore in 2013 and 2014, and a correlative approach was used to evaluate the extent to which the densities were associated using sites as replicates. For 8 sites censused in both years, mean densities of adults (pooled among taxa) varied from 2.95 ± 1.16 colonies m− 2 to 20.60 ± 2.62 colonies m− 2 in 2013, and from 3.20 ± 0.75 colonies m− 2 to 13.00 ± 1.04 colonies m− 2 in 2014; for recruits, mean densities varied from 1.05 ± 0.34 colonies m− 2 to 4.25 ± 0.81 colonies m− 2 in 2013, and from 0.60 ± 0.31 colonies m− 2 to 1.44 ± 0.40 colonies m− 2 in 2014 (all ± SE). The most common taxa in both years among all sites were Antillogorgia spp., Gorgonia spp., and plexaurids. Density of recruits was significantly and positively correlated with population density of adult octocorals (pooled among taxa) and plexaurids in both years, and for Gorgonia spp. in 2013 (with a similar trend in 2014). Densities of recruits and adults of Antillogorgia spp. were not associated in either year. Together, these data suggest that densities of adult octocorals positively influence the density of co-occurring octocoral recruits, thereby potentially promoting population growth.

Coral reefs experience biologically-driven pCO2 oscillations that are predicted to become more extreme in magnitude and duration under ocean acidification (OA) regimes. Understanding the plasticity of responses in common reef-building corals to oscillations in pCO2 will allow for better predictions of their function in future seawater conditions. This study explored the effects of variation in seawater pCO2 on coral calcification using experiments conducted over one month between 9 April 2018 and 18 May 2018. Branches (~4-cm long) of Acropora retusa were sampled from colonies at 10-m depth on the fore reef of Mo'orea, French Polynesia (17° 28′ 53.9004" S, 149° 49′ 50.5992" W). We tested the hypothesis that depressed calcification caused by elevated pCO2 (~1000 μatm) is relaxed (i.e., calcification increases) upon return to ambient pCO2 (~400 μatm). Corals first were incubated in ambient or elevated pCO2 for 19 days, with the result that calcification integrated over this period was reduced by 31% under elevated pCO2. The same corals were then incubated at ambient pCO2 for 11 days, during which calcification was independent of the experimental pCO2 exposure history. Our results suggest that a quick relaxation of pCO2-depressed calcification in A. retusa following cessation of high pCO2 indicates that corals are capable of a reversible plastic response of calcification when confronted by pCO2 oscillations.

Density dependence (DD) controls community recovery following widespread mortality, yet this principle rarely has been applied to coral assemblages. The reefs of Mo'orea, French Polynesia, provide the opportunity to study DD of coral population growth, because coral assemblages in this location responded to declines in abundance with high recruitment and an increase in cover during which recruitment of pocilloporid corals was inversely associated with density. This study tests for DD in this system, first, by describing the context within which it operates: coral cover changed from 46% in 2005, to

Ocean acidification (OA) poses a severe threat to tropical coral reefs, yet much of what is know about these effects comes from individual corals and algae incubated in isolation under high pCO2. Studies of similar effects on coral reef communities are scarce. To investigate the response of coral reef communities to OA, we used large outdoor flumes in which communities composed of calcified algae, corals, and sediment were combined to match the percentage cover of benthic communities in the shallow back reef of Moorea, French Polynesia. Reef communities in the flumes were exposed to ambient (~ 400 μatm) and high pCO2 (~ 1300 μatm) for 8 weeks, and calcification rates measured for the constructed communities including the sediments. Community calcification was reduced by 59% under high pCO2, with sediment dissolution explaining ~ 50% of this decrease; net calcification of corals and calcified algae remained positive but was reduced by 29% under elevated pCO2. These results show that, despite the capacity of coral reef calcifiers to maintain positive net accretion of calcium carbonate under OA conditions, reef communities might transition to net dissolution as pCO2 increases, particularly at night, due to enhanced sediment dissolution.

Populations of marine organisms on coral reef islands (CRI) are connected in space and time by seawater that transports propagules of plants, animals, and algae. Yet, despite this reality, it is often assumed that routine replenishment of populations of marine organisms on CRI is supported by locally-sourced propagules (hereafter, larvae). Following large disturbances, however, distantly-sourced larvae from less disturbed CRI within a regional meta-population are likely to be important for local population recovery, but evaluating the roles of locally- vs. distantly-sourced larvae remains difficult. While larval sources are relatively well-known for some fishes, they remain virtually unknown for most taxa, particularly those associated with the benthos, as exemplified by scleractinian corals. This review focuses on reef recovery and larval connectivity. Using corals as examples, we argue that CRI can serve as natural laboratories in which studies of these issues can enhance understanding of coral reef community dynamics under future disturbance regimes. Rather than focusing on synthesizing empirical data, we focus on the capacity for CRI to realize their potential in this research area, concluding that progress is impeded by the limited breadth, detail, and spatio-temporal concordance of existing research. Using long-term observational programs of coral reefs in Mo'orea (French Polynesia), Okinawa (Japan), and St. John (US Virgin Islands) as examples of the data currently available, we make the case that new modes of multidisciplinary and collaborative research will be required to exploit the value of CRI in understanding the role of connectivity in mediating ecosystem resilience in a future affected by anthropogenic disturbances.

Coral abundance continues to decline on tropical reefs around the world, and this trend suggests that coral reefs may not persist beyond the current century. In contrast, this study describes the near-complete mortality of corals on the outer reef (10 m and 17 m depth) of the north shore of Mo’orea, French Polynesia, from 2005 to 2010, followed by unprecedented recovery from 2011 to 2017. Intense corallivory and a cyclone drove coral cover from 33–48% to −1) to 25–74% cover by 2017. The thirteen-year, U-shape trajectory of coral cover over time created by the loss and replacement of millions of corals through sexual reproduction underscores the potential for beneficial genetic responses to environmental conditions for at least one genus, Pocillopora. The high ecological resilience of this coral community appears to have been enhanced by variation among genera in the susceptibility to declining cover, and the capacity for population growth (i.e., response diversity). These results suggest that the outer coral communities of Mo’orea may be poised for genetic changes that could affect their capacity to persistence.

Much of the research on the effects of ocean acidification on tropical coral reefs has focused on the calcification rates of individual coral colonies, and less attention has been given to carbonate production and dissolution at the community scale. Using flumes (5.0 × 0.3 × 0.3 m) located outdoors in Moorea, French Polynesia, we assembled local back reef communities with ~ 25% coral cover, and tested their response to pCO2 levels of 344, 633, 870 and 1146 μatm. Incubations began in late Austral spring (November 2015), and net community calcification (Gnet) and net community primary production (Pnet) were measured prior to treatments, 24 h after treatment began, and biweekly or monthly thereafter until early Austral autumn (March 2016). Gnet was depressed under elevated pCO2 over 4 months, although the magnitude of the response varied over time. The proportional decline in Gnet as a function of saturation state of aragonite (Ωar) depended on the initial Ωar, but was 24% for a decline in Ωar from 4.0 to 3.0, which is nearly twice as sensitive to variation in Ωar than the previously published values for the net calcification of ex situ coral colonies. However, community Gnet was less sensitive to Ωar than coral reefs that have been analyzed in situ. Pnet was unaffected by pCO2, but Pnet and Gnet expressed on a hourly time base were positively associated, thus revealing the tight coupling between these metabolic processes. The high sensitivity of Gnet to pCO2 for the back reef of Moorea, versus lower sensitivity of individual coral colony calcification to pCO2, underscores the challenges of scaling-up experimental results on the effects of pCO2 from coral reef organisms to coral reef communities.

Octocorals have increased in abundance on a number of Caribbean reefs, but this trend has largely been reported with functional group or genus resolution. A species-level analysis of octocoral communities in St. John, US Virgin Islands was conducted to better understand how this taxon will respond to changing conditions based on their synecology at two sites that are 1.5 km apart and differ in physical conditions. East Cabritte is characterized by moderate wave energy, low sedimentation, and clear water, while contrasting conditions characterize Europa Bay. Surveys conducted in 2014 and 2015 showed that the abundance and size of adult octocorals differed between sites, with taller and denser communities at East Cabritte than Europa Bay (mean height of 32 versus 20 cm; 18 versus 8 colonies m−2). Octocoral diversity and evenness were similar between sites, although multivariate octocoral community structure differed between sites regardless of whether octocorals were resolved to genera or species. Genus-resolution masked differences between sites for speciose genera like Eunicea. The broad overlap in species representation at both sites suggests that diversity is less responsive than community structure to differing environmental conditions, perhaps because the ecological niches of these species are broad. With 35 octocoral species, and diversity and abundances comparable to those studied > 40 years ago on shallow Caribbean reefs, the dense octocoral communities appearing on some present-day reefs reflect expanded benthic occupancy by a well-known (rather than a novel) community type.