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In 1983 to 1984, a mass mortality event caused a Caribbean-wide, >95% population reduction of the echinoid grazer, Diadema antillarum . This led to blooms of algae contributing to the devastation of scleractinian coral populations. Since then, D. antillarum exhibited only limited and patchy population recovery in shallow water, and in 2022 was struck by a second mass mortality reported over many reef localities in the Caribbean. Half-a-century time-series analyses of populations of this sea urchin from St. John, US Virgin Islands, reveal that the 2022 event has reduced population densities by 98.00% compared to 2021, and by 99.96% compared to 1983. In 2021, coral cover throughout the Caribbean was approaching the lowest values recorded in modern times. However, prior to 2022, locations with small aggregations of D. antillarum produced grazing halos in which weedy corals were able to successfully recruit and become the dominant coral taxa. The 2022 mortality has eliminated these algal-free halos on St. John and perhaps many other regions, thereby increasing the risk that these reefs will further transition into coral-free communities.

For nearly 50 years, analyses of coral physiology have used small coral fragments (nubbins) to make inferences about larger colonies. However, scaling in corals shows that linear extrapolations from nubbins to whole colonies can be misleading, because polyps in nubbins are divorced of their morphologically complex and physiologically integrated corallum. We tested for the effects of integration among branches in determining size-dependent calcification of the coral Pocillopora spp. under elevated P CO 2 . Area-normalized net calcification was compared between branches (nubbins), aggregates of nubbins (complex morphologies without integration) and whole colonies (physiologically integrated) at 400 versus approximately 1000 µatm P CO 2 . Net calcification was unaffected by P CO 2 , but differed among colony types. Single nubbins grew faster than whole colonies, but when aggregated, nubbins changed calcification to match whole colonies even though they lacked integration among branches. Corallum morphology causes the phenotype of branching corals to differ from the summation of their branches.

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.

Sessile organisms exploit a life-history strategy in which adults are immobile and their growth position is determined at settlement. The morphological strategy exploited by these organisms has strong selective value, because it can allow beneficial matching of morphology to environmental and biological conditions. In benthic marine environments, a ‘sheet-tree’ morphology is a classic mechanism exploited by select sessile organisms, and milleporine hydrocorals provide one of the best examples of this strategy. Using 30-year analysis of Millepora sp. on the reefs of St. John, US Virgin Islands, I tested for the benefits of a sheet-tree morphology in mediating the ecological success of an important functional group of benthic space holders. The abundance of Millepora sp. chaotically changed from 1992 to 2021 in concert with hurricanes, bleaching and macroalgal crowding. Millepora sp. responded to these disturbances by exploiting their morphological strategy to increase the use of trees when their sheets were compromised by bleaching and spatial competition with macroalgae, and the use of sheets when their trees were broken by storms. Together, these results reveal the selective value of a plastic sheet-tree morphology, which can be exploited by sessile organisms to respond to decadal-scale variation in environmental conditions.

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.

The severity of marine heatwaves (MHWs) that are increasingly impacting ocean ecosystems, including vulnerable coral reefs, has primarily been assessed using remotely sensed sea-surface temperatures (SSTs), without information relevant to heating across ecosystem depths. Here, using a rare combination of SST, high-resolution in-situ temperatures, and sea level anomalies observed over 15 years near Moorea, French Polynesia, we document subsurface MHWs that have been paradoxical in comparison to SST metrics and associated with unexpected coral bleaching across depths. Variations in the depth range and severity of MHWs was driven by mesoscale (10s to 100s of km) eddies that altered sea levels and thermocline depths and decreased (2007, 2017 and 2019) or increased (2012, 2015, 2016) internal-wave cooling. Pronounced eddy-induced reductions in internal waves during early 2019 contributed to a prolonged subsurface MHW and unexpectedly severe coral bleaching, with subsequent mortality offsetting almost a decade of coral recovery. Variability in mesoscale eddy fields, and thus thermocline depths, is expected to increase with climate change, which, along with strengthening and deepening stratification, could increase the occurrence of subsurface MHWs over ecosystems historically insulated from surface ocean heating by the cooling effects of internal waves.

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 (

After centuries of human-mediated disturbances, Caribbean reef communities are vastly different from those described in the 1950s. Many are functionally dominated by macroalgae, but this community state represents only one of several possibilities into which present-day coral reefs can transition. Octocorals have always been abundant on Caribbean reefs, but increases in their abundance over the last few decades suggest that arborescent octocorals have the potential to expand their populations on reefs that hitherto had been dominated by scleractinians. Here we show that octocoral-dominated communities at three sites on the fringing reefs of St. John, US Virgin Islands, were resilient to the effects of two Category 5 hurricanes in 2017. We describe the dynamics of octocoral communities over five years at three sites on shallow reefs (~9-m depth), and test for the effects of Hurricanes Irma and Maria. The hurricanes depressed the densities of juvenile and adult octocoral colonies as much as 47%. However, there were only weak effects on species richness and the relative abundances of the octocoral species. The hurricanes did not alter patterns of spatial variability in octocoral community structure that existed among sites prior to the storms. The density of octocoral recruits (individuals ≤ 5 cm high) was reduced in the year following the hurricanes, mainly due to a decline in abundance of recruits

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.

In this study, fore reef coral communities were exposed to high pCO2 for a year to explore the relationship between net accretion (Gnet) and community structure (planar area growth). Coral reef communities simulating the fore reef at 17-m depth on Mo’orea, French Polynesia, were assembled in three outdoor flumes (each 500 l) that were maintained at ambient (396 µatm), 782 µatm, and 1434 µatm pCO2, supplied with seawater at 300 l h−1, and exposed to light simulating 17-m depth. The communities were constructed using corals from the fore reef, and the responses of massive Porites spp., Acropora spp., and Pocillopora verrucosa were assessed through monthly measurements of Gnet and planar area. High pCO2 depressed Gnet but did not affect colony area by taxon, although the areas of Acropora spp. and P. verrucosa summed to cause multivariate community structure to differ among treatments. These results suggest that skeletal plasticity modulates the effects of reduced Gnet at high pCO2 on planar growth, at least over a year. The low sensitivity of the planar growth of fore reef corals to the effects of ocean acidification (OA) on net calcification supports the counterintuitive conclusion that coral community structure may not be strongly affected by OA.

Identifying relatively intact areas within ecosystems and determining the conditions favoring their existence is necessary for effective management in the context of widespread environmental degradation. In this study, we used 3766 surveys of randomly selected sites in the United States and U.S. Territories to identify the correlates of sites categorized as "oases" (defined as sites with relatively high total coral cover). We used occupancy models to evaluate the influence of 10 environmental predictors on the probability that an area (21.2-km

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.

Ecosystems are changing in complex and unpredictable ways, and analysis of these changes is facilitated by coordinated, long‐term research. Meeting diverse societal needs requires an understanding of what populations and communities will be dominant in 20, 50, and 100 yr. This paper is a product of a synthesis effort of the U.S. National Science Foundation funded Long‐Term Ecological Research (LTER) network addressing the LTER core research area of populations and communities. This analysis revealed that each LTER site had at least one compelling story about what their site would look like in 50 or 100 yr. As the stories were prepared, themes emerged, and the stories were grouped into papers along five themes for this special issue: state change, connectivity, resilience, time lags, and cascading effects. This paper addresses the resilience theme and includes stories from the Baltimore (urban), Hubbard Brook (northern hardwood forest), Andrews (temperate rain forest), Moorea (coral reef), Cedar Creek (grassland), and North Temperate Lakes (lakes) sites. The concept of resilience (the capacity of a system to maintain structure and processes in the face of disturbance) is an old topic that has seen a resurgence of interest as the nature and extent of global environmental change have intensified. The stories we present here show the power of long‐term manipulation experiments (Cedar Creek), the value of long‐term monitoring of forests in both natural (Andrews, Hubbard Brook) and urban settings (Baltimore), and insights that can be gained from modeling and/or experimental approaches paired with long‐term observations (North Temperate Lakes, Moorea). Three main conclusions emerge from the analysis: (1) Resilience research has matured over the past 40 yr of the LTER program; (2) there are many examples of high resilience among the ecosystems in the LTER network; (3) there are also many warning signs of declining resilience of the ecosystems we study. These stories highlight the need for long‐term studies to address this complex topic and show how the diversity of sites within the LTER network facilitates the emergence of overarching concepts about this important driver of ecosystem structure, function, services, and futures.

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.

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.

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.

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.