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Biome degradation characterizes the Anthropocene Epoch, and modern ecology is deeply involved with describing the changes underway. Most research has focused on the role of acute disturbances in causing conspicuous changes in ecosystem structure, which leads to an underappreciation of the chronic effects causing large changes through the cumulative effects of small perturbations over decades. Coral reefs epitomize this trend, because the changes in community structure are profound, yet the data to quantify these effects are usually insufficient to evaluate the relative roles of different disturbance types. Here, four decades of surveys from two coral reefs (9 and 14 m depth) off St. John, US Virgin Islands, are used to quantify the associations of acute and chronic events with the changes in benthic community structure. These reefs profoundly changed over 36 years, with coral death altering species assemblages to depress abundances of the ecologically important coral Orbicella spp. and elevating the coverage of macroalgae and crustose coralline algae/turf/bare space (CTB). Linear mixed models revealed the prominent role of chronic variation in temperature in accounting for changes in coverage of corals, macroalgae, and CTB, with rising temperature associated with increases in coral cover on the deep reef, and declines on the shallow reef. Hurricanes were also associated with declines in coral cover on the shallow reef, and increases on the deep reef. Multivariate analyses revealed strong associations between community structure and temperature, but weaker associations with hurricanes, bleaching, and diseases. These results highlight the overwhelming importance of chronically increasing temperature in altering the benthic community structure of Caribbean reefs. [ABSTRACT FROM AUTHOR]

Monitoring coral cover can describe the ecology of reef degradation, but rarely can it reveal the proximal mechanisms of change, or achieve its full potential in informing conservation actions. Describing temporal variation in Symbiodiniaceae within corals can help address these limitations, but this is rarely a research priority. Here, we augmented an ecological time series of the coral reefs of St. John, US Virgin Islands, by describing the genetic complement of symbiotic algae in common corals. Seventy-five corals from nine species were marked and sampled in 2017. Of these colonies, 41% were sampled in 2018, and 72% in 2019; 28% could not be found and were assumed to have died. Symbiodiniaceae ITS2 sequencing identified 525 distinct sequences (comprising 42 ITS2 type profiles), and symbiont diversity differed among host species and individuals, but was in most cases preserved within hosts over 3 yrs that were marked by physical disturbances from major hurricanes (2017) and the regional onset of stony coral tissue loss disease (2019). While changes in symbiont communities were slight and stochastic over time within colonies, variation in the dominant symbionts among colonies was observed for all host species. Together, these results indicate that declining host abundances could lead to the loss of rare algal lineages that are found in a low proportion of few coral colonies left on many reefs, especially if coral declines are symbiont-specific. These findings highlight the importance of identifying Symbiodiniaceae as part of a time series of coral communities to support holistic conservation planning. Repeated sampling of tagged corals is unlikely to be viable for this purpose, because many Caribbean corals are dying before they can be sampled multiple times. Instead, random sampling of large numbers of corals may be more effective in capturing the diversity and temporal dynamics of Symbiodiniaceae metacommunities in reef corals. [ABSTRACT FROM AUTHOR]

As the major form of coral reef regime shift, stony coral to macroalgal transitions have received considerable attention. In the Caribbean, however, regime shifts in which scleractinian corals are replaced by octocoral assemblages hold potential for maintaining reef associated communities. Accordingly, forecasting the resilience of octocoral assemblages to future disturbance regimes is necessary to understand these assemblages' capacity to maintain reef biodiversity. We parameterised integral projection models quantifying the survival, growth, and recruitment of the octocorals, Antillogorgia americana, Gorgonia ventalina, and Eunicea flexuosa, in St John, US Virgin Islands, before, during, and after severe hurricane disturbance. Using these models, we forecast the density of populations of each species under varying future hurricane regimes. We demonstrate that although hurricanes reduce population growth, A. americana, G. ventalina, and E. flexuosa each display a capacity for quick recovery following storm disturbance. Despite this recovery potential, we illustrate how the population dynamics of each species correspond with a longer-term decline in their population densities. Despite their resilience to periodic physical disturbance events, ongoing global change jeopardises the future viability of octocoral assemblages. [ABSTRACT FROM AUTHOR]

Understanding population dynamics is a long-standing objective of ecology, but the need for progress in this area has become urgent. For coral reefs, achieving this objective is impeded by a lack of information on settlement versus post-settlement events in determining recruitment and population size. Declines in coral abundance are often inferred to be associated with reduced densities of recruits, which could arise from mechanisms occurring at larval settlement, or throughout post-settlement stages. This study uses annual measurements from 2008 to 2021 of coral cover, the density of coral settlers (S), the density of small corals (SC), and environmental conditions, to evaluate the roles of settlement versus post-settlement events in determining rates of coral recruitment and changes in coral cover at Moorea, French Polynesia. Coral cover, S, SC, and the SC:S ratio (a proxy for post-settlement success), and environmental conditions, were used in generalized additive models (GAMs) to show that: (a) coral cover was more strongly related to SC and SC:S than S, and (b) SC:S was highest when preceded by cool seawater, low concentrations of Chlorophyll a, and low flow speeds, and S showed evidence of declining with elevated temperature. Together, these results suggest that changes in coral cover in Moorea are more strongly influenced by post-settlement events than settlement. The key to understanding coral community resilience may lie in elucidating the factors attenuating the bottleneck between settlers and small corals. [ABSTRACT FROM AUTHOR]

The implications of ocean acidification are acute for calcifying organisms, notably tropical reef corals, for which accretion generally is depressed and dissolution enhanced at reduced seawater pH. We describe year‐long experiments in which back reef and fore reef (17‐m depth) communities from Moorea, French Polynesia, were incubated outdoors under pCO2 regimes reflecting endpoints of representative concentration pathways (RCPs) expected by the end the century. Incubations were completed in three to four flumes (5.0 × 0.3 m, 500 L) in which seawater was refreshed and circulated at 0.1 m s−1, and the response of the communities was evaluated monthly by measurements of net community calcification (NCC) and net community productivity (NCP). For both communities, NCC (but not NCP) was affected by treatments and time, with NCC declining with increasing pCO2, and for the fore reef, becoming negative (i.e., dissolution was occurring) at the highest pCO2 (1067–1433 μatm, RCP8.5). There was scant evidence of community adjustment to reduce the negative effects of ocean acidification, and inhibition of NCC intensified in the back reef as the abundance of massive Porites spp. declined. These results highlight the risks of dissolution under ocean acidification for coral reefs and suggest these effects will be most acute in fore reef habitats. Without signs of amelioration of the negative effects of ocean acidification during year‐long experiments, it is reasonable to expect that the future of coral reefs in acidic seas can be predicted from their current known susceptibility to ocean acidification. [ABSTRACT FROM AUTHOR]

Coral recruitment describes the addition of new individuals to populations, and it is one of the most fundamental demographic processes contributing to population size. As many coral reefs around the world have experienced large declines in coral cover and abundance, there has been great interest in understanding the factors causing coral recruitment to vary and the conditions under which it can support community resilience. While progress in these areas is being facilitated by technological and scientific advances, one of the best tools to quantify recruitment remains the humble settlement tile, variants of which have been in use for over a century. Here I review the biology and ecology of coral recruits and the recruitment process, largely as resolved through the use of settlement tiles, by: (i) defining how the terms 'recruit' and 'recruitment' have been used, and explaining why loose terminology has impeded scientific advancement; (ii) describing how coral recruitment is measured and why settlement tiles have value for this purpose; (iii) summarizing previous efforts to review quantitative analyses of coral recruitment; (iv) describing advances from hypothesis‐driven studies in determining how refuges, seawater flow, and grazers can modulate coral recruitment; (v) reviewing the biology of small corals (i.e. recruits) to understand better how they respond to environmental conditions; and (vi) updating a quantitative compilation of coral recruitment studies extending from 1974 to present, thus revealing long‐term global declines in density of recruits, juxtaposed with apparent resilience to coral bleaching. Finally, I review future directions in the study of coral recruitment, and highlight the need to expand studies to deliver taxonomic resolution, and explain why time series of settlement tile deployments are likely to remain pivotal in quantifying coral recruitment. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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.Hidden marine heatwaves, associated with ocean eddies that modulate undersea internal waves, threaten coastal ecosystems by driving unexpected sub-surface heating and severe coral bleaching and mortality across depths. [ABSTRACT FROM AUTHOR]

Marine fouling communities have long provided model systems for studying the ecology of community development, and settlement plates are the tool of choice for this purpose. Decades of plate deployments provide a baseline against which present-day trends can be interpreted, with one classic trend being the ultimate dominance of plates by colonial and encrusting taxa. Here we report the results of annual deployments of settlement plates from 2010 to 2021 in the shallow sub-tidal of southern California, where the recruitment of invertebrates and algae was recorded photographically, and resolved to functional group (solitary, encrusting, and arborescent) and the lowest taxon possible. The communities on these plates differed among years, with trends in abundances varying by functional group and taxon; solitary taxa consistently were abundant, but encrusting taxa declined in abundance. Seawater temperature and the subsurface concentration of chlorophyll a differed among years, and there was a weak inverse association between temperature and the abundances of encrusting taxa. Long-term increases in seawater temperature therefore could serve as a mechanism causing fouling communities to change. Because of the prominence of encrusting taxa in fouling communities, the shifts in abundance of this functional group reported here may portend ecologically significant changes in fouling communities exposed to warmer seawater because of an alleviation of competition for a classically limiting resource (i.e., space). [ABSTRACT FROM AUTHOR]

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‐km2 cell) would harbor coral oases defined at four spatial extents: cross‐basin, basin, region, and subregion. Across all four spatial extents, oases were more likely to occur in habitats with high light attenuation. The influence of the other environmental predictors on the probability of oasis occurrence were less consistent and varied with the scale of observation. Oases were most likely in areas of low human population density, but this effect was evident only at the cross‐basin and subregional extents. At the regional and subregional extents oases were more likely where sea‐surface temperature was more variable, whereas at the larger spatial extents the opposite was true. By identifying the correlates of oasis occurrence, the model can inform the prioritization of reef areas for management. Areas with biophysical conditions that confer corals with physiological resilience, as well as limited human impacts, likely support coral reef oases across spatial extents. Our approach is widely applicable to the development of conservation strategies to protect biodiversity and ecosystems in an era of magnified human disturbance. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Most tropical coral reefs are experiencing declining coral cover, yet interpretation of this generality is tempered by spatial variation in coral cover among reefs separated over 20–200 km. This study addresses such landscape scale variation in coral reefs at 12 sites (7–10 m depth) around St. John (18° 18´ 37.04 N, 63° 43´ 23.17 W) and St. Thomas (18° 20´ 43.57 N, 65° 55´ 13.88 W), US Virgin Islands. Surveys completed from 2011 to 2021 were used to test for spatial variation in community dynamics among islands, shores, and sites. Community synchrony (φ) was used to evaluate portfolio effects in mediating changes in coral communities. From 2011 to 2021, changes in benthic communities differed among sites and times (i.e., there were site × year interactions), and, while coral cover declined at 11 sites, the decline was 2.6-fold faster around St. Thomas than St. John. The loss of coral cover was driven by multiple taxa that differed among sites, thus revealing asynchronous responses to prevailing conditions. Asynchrony suggests that coral communities at some of the sites have the capacity to exploit portfolio effects to modulate stability, yet these effects did not prevent declines in coral cover that reflect island-scale phenomena. In the US Virgin Islands, coral death is overwhelming the capacity for resilience of coral communities that historically may have benefitted from portfolio effects. Until coral assemblages are depleted through taxonomic extirpation, maintenance of the assemblage composition retains the possibility that increased resilience might emerge if environmental challenges can be alleviated. [ABSTRACT FROM AUTHOR]

This article discusses the fate of dead coral colonies on tropical reefs, specifically focusing on the reefs of Moorea, French Polynesia. The authors describe an unusual mode of decay where dead Pocillopora spp corals rapidly dissolve, leaving behind "ghosts" with hollow branches. These hollowed corals are hidden beneath a calcified façade colonized by algae, but their internal voids indicate imminent skeletal crumbling. The article raises questions about how coral reef recovery is affected by the structural legacies of dead corals and whether ghost colonies will become the new normal for reefs facing coral mortality due to ocean acidification and climate change. [Extracted from the article]

Measurements of coral recruitment can improve understanding of coral population dynamics, but in short duration studies high variance impedes interpretation of this vital rate. Here, coral recruitment in Moorea, French Polynesia, was measured over 13 yr and tested for associations with environmental conditions. Recruitment of spawning pocilloporid corals was recorded using settlement tiles immersed for ~ 6 months at 10 m and 17 m depth, biannually, and the environment was quantified through seawater clarity (Kd490), surface and bottom flow speeds, coral cover, and temperature. Using GAMs, most of the variation (63–74%) in recruitment was explained using depth and three predictors. Recruitment was high in January/February following a year of low seawater clarity (high Kd490), and in August/September following a year of high clarity. Recruitment in January/February was elevated by high coral cover, and by high surface flow during larval dispersal; recruitment was low in August/September following high temperature during gametogenesis. These effects reveal the extent to which early life stages of corals are exposed to annually variable environmental conditions that have differential consequences depending on the phenology of reproduction and recruitment. The temporal mosaic defining interactions between reproduction, recruitment, and environmental conditions create opportunities for acclimatization to affect where and when coral recruitment is successful. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Coral bleaching in 2019 affected ~72% of I Pocillopora i spp. corals, killing 11-42% of Moorea colonies (French Polynesia). GLO:69VH/01jul21:bes21885-fig-0005.jpg PHOTO (COLOR): . gl Photo 5: Researcher collecting standardized images of corals, using a photoquadrat frame, to measure the size and survival of individual colonies. GLO:69VH/01jul21:bes21885-fig-0002.jpg PHOTO (COLOR): . gl Photo 2: Coral bleaching in 2019 at Moorea appeared to affect the larger colonies more than the smaller colonies of coral in the genus Pocillopora. [Extracted from the article]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The supply of propagules mediates recruitment and population dynamics, thereby driving community resilience following disturbances. These relationships are of interest on tropical reefs, where coral populations have drastically declined in abundance and sexual recruitment is the only means by which they will recover. To better understand the causes and implications of variation in this vital rate (i.e., recruitment), coral recruitment was measured in Mo'orea, French Polynesia, and St. John, U.S. Virgin Islands, using settlement tiles deployed from 2005 to 2019. The results were used to test two hypotheses: (1) annual variation in recruitment is a weak predictor of long‐term variation in recruitment, but (2) it is associated with seawater temperature. Coral recruitment varied over space and time, so that differences in recruitment between consecutive years were uninformative of long‐term trends. Recruitment varied among years in an apparently chaotic manner, but the variation reflected linear and quadratic associations with mean annual temperature and the daily variation in temperature. These associations are consistent with theory addressing the mechanisms by which temperature affects coral larvae and recruitment. Comprehension of these mechanisms is required to accurately interpret evidence of coral recruitment collapse, and to elucidate the conditions favoring recovery of coral communities through recruitment. [ABSTRACT FROM AUTHOR]

With coral cover declining on most tropical reefs, efforts are intensifying to understand the mechanisms by which environmental stressors are driving populations into demographic deficit. Here, the population dynamics of sexual cohorts of small corals (≤ 4 cm diameter) in St. John, US Virgin Islands were used to test the hypothesis that the association of vital rates with environmental conditions has changed over 22 yr. The annual fate of 3999 corals at six sites and 7--9 m depth were used to populate Leslie Matrices and calculate assemblage growth rates (λ, without recruitment), mortality of the smallest corals, and graduation from the largest size class. Vital rates were tested for associations with temperature, the number of days with seawater warmer than 29.4°C (i.e., hot days), and rainfall. Transition probabilities varied among years, and usually more corals remained in their original size class than moved among size classes. Corals in the smallest size class had the highest mortality rate (42% yr-1), λ had a grand mean of 0.688, and there were trends for λ to vary with the number of hot days and rainfall through quadratic relationships. The linear dependence of λ on rainfall over consecutive 3-yr periods declined over two decades, suggesting that this colony size-defined coral assemblage has undergone phenotypic change in their responsiveness to varying environmental conditions. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

In this study, fore reef coral communities were exposed to high pCO2 for a year to explore the relationship between net accretion (G net) 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 G net and planar area. High pCO2 depressed G net 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 G net 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. [ABSTRACT FROM AUTHOR]

Given the severe implications of climate change and ocean acidification (OA) for marine ecosystems, there is an urgent need to quantify ecosystem function in present‐day conditions to determine the impacts of future changes in environmental conditions. For tropical coral reefs that are acutely threatened by these effects, the metabolism of benthic communities provides several metrics suitable for this purpose, but the application of infrastructure to manipulate conditions and measure community responses is not fully realized. To date, most studies of the effects of OA on coral reefs have been conducted ex situ, and while greater ecological relevance can be achieved through free ocean carbon enrichment (FOCE) experiments on undisturbed areas of reef, such approaches have been deterred by technical challenges (e.g., spatial scale and duration, stable maintenance of conditions). In this study, we describe novel experimental infrastructure called shallow coral reef (SCoRe) FOCE to overcome these challenges and present data from a proof of concept application in Mo'orea, French Polynesia. Our objectives were to (1) implement an autonomous system that could be deployed kilometers from shore, (2) regulate the chemical (pCO2) and physical properties of seawater over undisturbed, shallow (∼2–5‐m depth) coral reef over multiple weeks, and (3) measure the metabolic response of the coral community to the treatment conditions. We describe the design, function, and application of the SCoRe FOCE, and present data demonstrating its efficacy. This infrastructure has great potential for advancing ecologically relevant studies of the effects of changing environmental conditions on coral reefs. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Human activities have led to widespread ecological decline; however, the severity of degradation is spatially heterogeneous due to some locations resisting, escaping, or rebounding from disturbances.We developed a framework for identifying oases within coral reef regions using long‐term monitoring data. We calculated standardised estimates of coral cover (z‐scores) to distinguish sites that deviated positively from regional means. We also used the coefficient of variation (CV) of coral cover to quantify how oases varied temporally, and to distinguish among types of oases. We estimated "coral calcification capacity" (CCC), a measure of the coral community's ability to produce calcium carbonate structures and tested for an association between this metric and z‐scores of coral cover.We illustrated our z‐score approach within a modelling framework by extracting z‐scores and CVs from simulated data based on four generalized trajectories of coral cover. We then applied the approach to time‐series data from long‐term reef monitoring programmes in four focal regions in the Pacific (the main Hawaiian Islands and Mo'orea, French Polynesia) and western Atlantic (the Florida Keys and St. John, US Virgin Islands). Among the 123 sites analysed, 38 had positive z‐scores for median coral cover and were categorised as oases.Synthesis and applications. Our framework provides ecosystem managers with a valuable tool for conservation by identifying "oases" within degraded areas. By evaluating trajectories of change in state (e.g., coral cover) among oases, our approach may help in identifying the mechanisms responsible for spatial variability in ecosystem condition. Increased mechanistic understanding can guide whether management of a particular location should emphasise protection, mitigation or restoration. Analysis of the empirical data suggest that the majority of our coral reef oases originated by either escaping or resisting disturbances, although some sites showed a high capacity for recovery, while others were candidates for restoration. Finally, our measure of reef condition (i.e., median z‐scores of coral cover) correlated positively with coral calcification capacity suggesting that our approach identified oases that are also exceptional for one critical component of ecological function. Our framework provides ecosystem managers with a valuable tool for conservation by identifying "oases" within degraded areas. By evaluating trajectories of change in state (e.g., coral cover) among oases, our approach may help in identifying the mechanisms responsible for spatial variability in ecosystem condition. Increased mechanistic understanding can guide whether management of a particular location should emphasise protection, mitigation or restoration. Analysis of the empirical data suggest that the majority of our coral reef oases originated by either escaping or resisting disturbances, although some sites showed a high capacity for recovery, while others were candidates for restoration. Finally, our measure of reef condition (i.e., median z‐scores of coral cover) correlated positively with coral calcification capacity suggesting that our approach identified oases that are also exceptional for one critical component of ecological function. [ABSTRACT FROM AUTHOR]

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 <1% in 2010 following an outbreak of a corallivorous sea star and a cyclone, and then increased to 74% by 2017, in large part through inverse density‐associated pocilloporid recruitment. Second, a test for DD of recruitment was conducted by decreasing Pocillopora spp. cover from 33% to 19%: one year later, the density of Pocillopora spp. recruits was 1.65‐fold higher in the low vs. high cover treatment. Finally, the effects of DD were investigated by comparing simulated and empirical distributions of pocilloporid colonies: as predicted by DD, small colonies were randomly distributed, while large colonies were uniformly distributed. Together these results demonstrate DD of population regulation for Pocillopora spp. corals, thus revealing the potential importance of this ecological principle in determining the resilience of coral assemblages. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The coral reef crisis has resulted in many reefs stabilizing in a low coral cover state, and this condition is exemplified by the fringing reefs of St. John, US Virgin Islands, where coral cover has remained low for decades. To evaluate the demographic features maintaining this condition, a decade of coral abundance was examined in three domains: recruits, small colonies (≤ 40-mm diameter), and cover. While cover was stable from 2007 to 2016, other aspects of coral community structure changed in taxonomically unique ways differing among domains. Most changes occurred gradually, although the warm year of 2010 was associated with perturbations in growth, mortality, and density that differed among domains and taxa, and summed to a weak signal distinguishing communities before and after ~ 2011/2012. As a result, coral community structure was taxonomically shuffled within each domain, while low cover was maintained. These low coral cover communities may be characterized by functional redundancy among taxa and a weakening of interactions among them that traditionally have supported emergent properties like community calcification. With persistent failure of recruitment to augment coral cover, some corals may already occur at such low abundances that they cannot perform the ecological services with which they were once associated. [ABSTRACT FROM AUTHOR]

Abstract: On Caribbean reefs, colonies of the hydrocoral Millepora alcicornis are capable of pursuing and overgrowing arborescent octocorals (Wahle ). Nearly four decades since these interactions were described, we quantified arborescent octocorals encrusted by Millepora spp. ( Linnaeus 1758) on shallow reefs in St. John, U.S. Virgin Islands, and evaluated encrusted colonies for evidence of an origin through pursuit. In 2014, 8% of octocorals (n=1684 colonies) were encrusted by Millepora spp., and in 2015, 12% were encrusted (n=847 colonies). Millepora spp. encrusted colonies of 10 octocoral genera, and in 2014, the most frequently encrusted were Eunicea spp. (40% of encrusted octocorals); in 2015, the most frequently encrusted were Gorgonia spp. (46% of encrusted colonies). In both years, ≥67% of encrusted octocorals were >21 cm from other colonies of Millepora spp.; 7% of octocorals were >1.3 m from colonies of Millepora spp. in 2014; and 16% of octocorals were >2.0 m from colonies of Millepora spp. in 2015. Relative to the ~5% of octocorals encrusted by Millepora spp. in Jamaica in the late 1970s, a high percentage (~9%) of octocorals were encrusted by Millepora spp. in St. John in 2014 and 2015. The large distances from most encrusted octocorals to the nearest colonies of Millepora spp. in St. John are inconsistent with a hypothesized origin through pursuit, contact, and overgrowth (sensu Wahle ). [ABSTRACT FROM AUTHOR]

For poikilothermic organisms in aquatic environment, temperature strongly affects metabolism and modulates the extent to which it is mass transfer limited. The effects of temperature on metabolism are translated into organism performance depending on the organism size, morphology, and the flow regime to which they are exposed. Environmental forcing of metabolism is particularly germane to tropical reef corals, because they increasingly are exposed to upward temperature perturbations to which they display variable responses that remain incompletely understood. In this study, we tested the effects of colony size and seawater turbulence on the response of the common Indo-Pacific branching coral, Pocillopora verrucosa, to different seawater temperatures. Using whole-colony calcification as a response variable, 12 tanks (each 150 l) were used in two trials lasting 14 days to contrast the effects of seawater turbulence (two levels) and temperature (25.5 vs 29.5 °C) on colonies varying in size from ~ 4 to 13-cm diameter. Turbulence in the tanks was measured as the root mean square (rms) turbulent flow speed ( q ), which quantified the variation in speed of complex and non-linear flow, regardless of direction. Treatments contrasted low ( q = 0.63 cm s) and high ( q = 2.07 cm s) turbulence that were ecologically relevant for shallow coral reefs. The effects of turbulent flow speed and temperature on calcification (mg day) differed among colony sizes: at 25.5 °C, calcification increased with tissue surface area at a slower rate under high, compared to low turbulent flow speeds; at low turbulent flow speeds, calcification increased with tissue surface area more slowly at 29.5 versus 25.5 °C. These results show that the response of branching corals to temperatures depends on colony size and turbulent flow speed. This outcome suggests that spatial heterogeneity in turbulent flow regimes across a coral reef may provide opportunities for branching corals to exploit colony size to modulate susceptibility to rising temperature. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The negative implications of the thermal sensitivity of reef corals became clear with coral bleaching throughout the Caribbean in the 1980’s, and later globally, with the severe El Niño of 1998 and extensive seawater warming in 2005. These events have substantially contributed to declines in coral cover, and therefore the El Niño of 2016 raised concerns over the implications for coral reefs; on the Great Barrier Reef these concerns have been realized. A different outcome developed in Mo’orea, French Polynesia, where in situ seawater temperature from 15 March 2016 to 15 April 2016 was an average of 0.4°C above the upper 95% CI of the decadal mean temperature, and the NOAA Degree Heating Weeks (DHW) metric supported a Level 1 bleaching alert (DHW ≥ 4.0). Starting 1 September 2016 and for the rest of the year (122 d), in situ seawater temperature was an average of 0.4°C above the 95% CI of long-term values, although DHW remained at zero. Minor coral bleaching (0.2–2.6% of the coral) occurred on the outer reef (10-m and 17-m depth) in April 2016, by May 2016 it had intensified to affect 1.3–16.8% of the coral, but by August 2016, only 1.4–3.0% of the coral was bleached. Relative to the previous decade, recruitment of scleractinians to settlement tiles on the outer- (10 m) and back- (2 m) reef over 2016/17 was high, both from January 2016 to August 2016, and from August 2016 to January 2017, with increased relative abundances of pocilloporids on the outer reef, and acroporids in the back reef. The 2016 El Niño created a distinctive signature in seawater temperature for Mo’orea, but it did not cause widespread coral bleaching or mortality, rather, it was associated with high coral recruitment. While the 2016 El Niño has negatively affected other coral reefs in the Indo-Pacific, the coral communities of Mo’orea continue to show signs of resilience, thus cautioning against general statements regarding the effects of the 2015/16 El Niño on coral reefs in the region. [ABSTRACT FROM AUTHOR]

Genetic variation underlying differences in organism performance is subject to natural selection, and organisms with high values of genetically determined phenotypic measures of fitness should perform better than those that do not. Using small scleractinian corals (i.e., ≤40-mm diameter), this principle was tested with 20 yr of census data from St. John, US Virgin Islands. Using growth rate (change in diameter) as a measure of fitness, growth in one year was tested for association with growth and survivorship in the following two years, and this process was repeated over 20 yr using a 3-yr sliding window. Virtually all variation in growth was independent of colony size, and growth among pairs of years was highly variable, with corals that grew fast in one year rarely growing fast in the next 2 yr. While growth in some pairs of years was positively correlated, ≤4% of the growth variance was explained by growth in the preceding 2 yr. Survivorship was related positively to growth in the preceding year, but the association was weak, it did not extend over 3 yr, and was inconsistent over the study. These results demonstrate the importance of the environment in translating phenotypic measures of fitness into future performance, and for small Caribbean corals, they suggest that environmental conditions may preempt genotype in determining short-term success. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 corals contained clade C Symbiodinium (either C1d or C15), and the genetic variation of symbionts did not account for among-colony bleaching differences. These results showed that high temperature causes coral bleaching independent of pCO2, and underscores the potential role of the coral host in driving intraspecific variation in coral bleaching. [ABSTRACT FROM AUTHOR]

In densely populated communities, such as coral reefs, organisms can modify the physical and chemical environment for neighbouring individuals. We tested the hypothesis that colony density (12 colonies each placed -0.5 cm apart versus -8 cm apart) can modulate the physiological response (measured through rates of calcification, photosynthesis and respiration in the light and dark) of the coral Pocillopora verrucosa to partial pressure of CO2 (PCO2) treatments (~400 µatm and -1200 µatm) by altering the seawater flow regimes experienced by colonies placed in aggregations within a flume at a single flow speed. While light calcification decreased 20% under elevated versus ambient PCO2 for colonies in low-density aggregations, light calcification of high-density aggregations increased 23% at elevated versus ambient PCO2. As a result, densely aggregated corals maintained calcification rates over 24 h that were comparable to those maintained under ambient PCO2, despite a 45% decrease in dark calcification at elevated versus ambient PCO2. Additionally, densely aggregated corals experienced reduced flow speeds and higher seawater retention times between colonies owing to the formation of eddies. These results support recent indications that neighbouring organisms, such as the conspecific coral colonies in the present example, can create small-scale refugia from the negative effects of ocean acidification. [ABSTRACT FROM AUTHOR]

The threat represented by ocean acidification (OA) for coral reef has received considerable attention because of the sensitivity of calcifiers to changing water 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 Moorea, French Polynesia, (ii) the fore reef of Moorea, and (iii) the reef flat of Oahu, 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 Moorea (but not in Oahu). 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. [ABSTRACT FROM AUTHOR]

Body size has large effects on organism physiology, but these effects remain poorly understood in modular animals with complex morphologies. Using two trials of a ~24 day experiment conducted in 2014 and 2015, we tested the hypothesis that colony size of the coral Pocillopora verrucosa affects the response of calcification, aerobic respiration and gross photosynthesis to temperature (~26.5 and ~29.7°C) and PCO2 (~40 and ~1000 μatm). Large corals calcified more than small corals, but at a slower size-specific rate; areanormalized calcification declined with size. Whole-colony and areanormalized calcification were unaffected by temperature, PCO2, or the interaction between the two. Whole-colony respiration increased with colony size, but the slopes of these relationships differed between treatments. Area-normalized gross photosynthesis declined with colony size, but whole-colony photosynthesis was unaffected by PCO2, and showed a weak response to temperature. When scaled up to predict the response of large corals, area-normalized metrics of physiological performance measured using small corals provide inaccurate estimates of the physiological performance of large colonies. Together, these results demonstrate the importance of colony size in modulating the response of branching corals to elevated temperature and high PCO2. [ABSTRACT FROM AUTHOR]

We tested for depth-dependent vertical zonation in coral reef community structure with surveys conducted in 2013 at 10 m and 27 m depth off the islands of Moorea, Tahiti, Maiao, and Tetiaroa, which are distributed over ~3500 km2 of the tropical south Pacific. Benthic communities were censused using photoquadrats to obtain percentage cover, first by functional groups (scleractinians, macroalgae, and crustose coralline algae, algal turf, and bare space combined [CTB]), and second by genus for scleractinians and Millepora. Virtually every aspect of community structure differed between the two depths, but the effects were dependent on spatial scale of investigation. Interactive effects of islands and depths determined the abundance of functional groups, as well as seven of eight common coral genera. On the two islands at which two sites were censused, the abundance of functional groups also differed between sites, and was affected by depth × site interactions. The spatially variable effects of depth on community structure underscores the importance of context-specific synergy between biotic and abiotic factors in driving these patterns. Spatial scale-dependent vertical zonation also shows the potential for these effects to be modulated by the changes that have affected coral community structure on contemporary reefs over the last few decades. [ABSTRACT FROM AUTHOR]

There are concerns about the future of coral reefs in the face of ocean acidification and warming, and although studies of these phenomena have advanced quickly, efforts have focused on pieces of the puzzle rather than integrating them to evaluate ecosystem-level effects. The field is now poised to begin this task, but there are information gaps that first must be overcome before progress can be made. Many of these gaps focus on calcification at the levels of cells, organisms, populations, communities, and ecosystem, and their closure will be made difficult by the complexity of the interdependent processes by which coral reefs respond to ocean acidification, with effects scaling from cells to ecosystems and from microns to kilometers. Existing ecological theories provide an important and largely untapped resource for overcoming these difficulties, and they offer great potential for integrating the effects of ocean acidification across scales on coral reefs. [ABSTRACT FROM AUTHOR]

K-12 science programs in the United States are implementing inquiry-based lessons to expose students to the scientific process, and in some states these changes are mandated by regulations. At the same time, university faculty in science, technology, engineering, and math (STEM) disciplines are being encouraged by agencies funding their research to develop outreach activities that typically involve schools. Outreach programs promoting interactions between schools and universities are one means to address this need, and here we describe such a program between a Los Angeles independent school and California State University, Northridge. Our program exploits experiential learning focused on marine biology to engage students in ecological research with application to contemporary environmental issues such as climate change. Moreover, it addresses multiple aspects of the Next Generation Science Standards within a flexible framework that can be adapted to multiple curricular needs. Coral reefs are used as a model system in this program, but the concepts can easily be extended to other coastal marine environments. The program has evolved through three phases: (1) fostering interactions among educational partners, and developing curricula through research that exploits problem-solving skills; (2) completion of scientific activities in the classroom and field; and (3) engaging students in the scientific process through professional conferences and publication. These efforts have led to a self-sustaining program of interactions among schoolchildren, undergraduates, graduate students, K-12 educators, and university faculty that is promoting STEM careers and generating peer-reviewed publications. [ABSTRACT FROM AUTHOR]

Many organisms exhibit depressed metabolism when resources are limited, a change that makes it possible to balance an energy budget. For symbiotic reef corals, daily cycles of light and periods of intense cloud cover can be chronic causes of food limitation through reduced photosynthesis. Furthermore, coral bleaching is common in present-day reefs, creating a context in which metabolic depression could have beneficial value to corals. In the present study, corals (massive Porites spp.) were exposed to an extreme case of resource limitation by starving them of food and light for 20 days. When resources were limited, the corals depressed area-normalized respiration to 37% of initial rates, and coral biomass declined to 64% of initial amounts, yet the corals continued to produce skeletal mass. However, the declines in biomass cannot account for the declines in area-normalized respiration, as mass-specific respiration declined to 30% of the first recorded time point. Thus, these corals appear to be capable of metabolic depression. It is possible that some coral species are better able to depress metabolic rates than others; such variation could explain differential survival during conditions that limit resources (e.g. shading). Furthermore, we found that maintenance of existing biomass, in part, supports the production of skeletal mass. This association could be explained if maintenance supplies needed energy (e.g. ATP) or inorganic carbon (i.e. CO2) that otherwise limits the production of skeletal mass. Finally, the observed metabolic depression can be explained as a change in pool sizes, and does not require a change in metabolic rules. [ABSTRACT FROM AUTHOR]

To date, studies of ocean acidification ( OA) on coral reefs have focused on organisms rather than communities, and the few community effects that have been addressed have focused on shallow back reef habitats. The effects of OA on outer barrier reefs, which are the most striking of coral reef habitats and are functionally and physically different from back reefs, are unknown. Using 5-m long outdoor flumes to create treatment conditions, we constructed coral reef communities comprised of calcified algae, corals, and reef pavement that were assembled to match the community structure at 17 m depth on the outer barrier reef of Moorea, French Polynesia. Communities were maintained under ambient and 1200 μatm pCO2 for 7 weeks, and net calcification rates were measured at different flow speeds. Community net calcification was significantly affected by OA, especially at night when net calcification was depressed ~78% compared to ambient pCO2. Flow speed (2-14 cm s−1) enhanced net calcification only at night under elevated pCO2. Reef pavement also was affected by OA, with dissolution ~86% higher under elevated pCO2 compared to ambient pCO2. These results suggest that net accretion of outer barrier reef communities will decline under OA conditions predicted within the next 100 years, largely because of increased dissolution of reef pavement. Such extensive dissolution poses a threat to the carbonate foundation of barrier reef communities. [ABSTRACT FROM AUTHOR]

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 <200 km, and suggests that recruitment from spatially discrete pools of coral larvae plays an important role in creating spatial variation in community structure, even where reefs are connected by prevailing currents. [ABSTRACT FROM AUTHOR]