Corals are critical to marine biodiversity. Reproduction and dispersal are key to their resilience, but rarely quantified in nature. Exploiting a unique system-a fully censused, longitudinally characterized, semi-isolated population inhabiting mangroves-we used 2bRAD sequencing to demonstrate that rampant asexual reproduction most likely via parthenogenesis and limited dispersal enable the persistence of a natural population of thin-finger coral (Porites divaricata). Unlike previous studies on coral dispersal, knowledge of colony age and location enabled us to identify plausible parent-offspring relationships within multiple clonal lineages and develop tightly constrained estimates of larval dispersal; the best-fitting model indicates dispersal is largely limited to a few metres from parent colonies. Our results explain why this species is adept at colonizing mangroves but suggest limited genetic diversity in mangrove populations and limited connectivity between mangroves and nearby reefs. As P. divaricata is gonochoristic, and parthenogenesis would be restricted to females (whereas fragmentation, which is presumably common in reef and seagrass habitats, is not), mangrove populations likely exhibit skewed sex ratios. These findings suggest that coral reproductive diversity can lead to distinctly different demographic outcomes in different habitats. Thus, coral conservation will require the protection of the entire coral habitat mosaic, and not just reefs.

IntroductionCoral species are negatively impacted by anthropogenic stressors worldwide. Nearshore coastal ecosystems provide an excellent study system for the investigation of the impacts of local land-based stressors such as nutrients on coral assemblages due to their proximity to land based sources of pollution. The nearshore environs of the Florida Keys are an intensively developed and intensively managed system; Florida Keys waters have been characterized by chronic eutrophication since 1999. Benthic communities are known to change or “shift” under chronic eutrophication.ObjectivesThis research examines the patterns of occurrence in stony corals correlating to detailed nearshore water quality measurements. The questions addressed are a) What are the coral assemblages within 500 meters of the shoreline, and which species are resilient to these environs? b) How does water quality impact coral occurrence?MethodsThis study examined the immediate nearshore (0-500m) areas of 13 sites using a randomized blocked experimental design for both water quality and benthic sampling. Water quality parameters measured included chlorophyll-a, nitrates + nitrites (NOx), total nitrogen, total Kjeldahl nitrogen, total phosphorus, Salinity, pH, temperature, and dissolved oxygen. Benthic surveys documented stony coral occurrence using belt transects with point intercept measurements.ResultsStony coral species occurrence was limited at most of the study sites. Siderastrea radians and Porites divaricata were the two common species that appear to have a higher tolerance to eutrophic conditions under these shallow water conditions. Of the 13 study sites, 2 sites had no coral species occurring; this analysis was based on the comparison between sites with and without stony coral species. Water column Chlorophyll-a, pH, dissolved oxygen saturation, temperature, NOx, N:P ratio, total nitrogen and total phosphorus were found to be significantly different between the sites with and without corals. Further multivariate analysis showed a clear clustering of the two groups.ConclusionThe low overall diversity and the higher occurrences of stress tolerant species provide evidence of the effects of coastal eutrophication on the coastal Florida Keys ecosystem. Our data suggest that more conservative water quality thresholds should be posed to improve nearshore biodiversity and ecological functioning of coastal systems.

Coral reefs are among the most diverse and complex ecosystems in the world that provide important ecological and economical services. Increases in sea surface temperature linked to global climate change threatens these ecosystems by inducing coral bleaching. However, it is not fully known if natural intra- or inter-annual physiological variability is linked to bleaching resilience or recovery capacity of corals. Here, we monitored the coral physiology of three common Caribbean species (Porites divaricata, Porites astreoides, Orbicella faveolata) at six time points over 2 years by measuring the following traits: calcification, biomass, lipids, proteins, carbohydrates, chlorophyll a, algal endosymbiont density, stable carbon isotopes of the host and endosymbiotic algae, and the stable carbon and oxygen isotopes of the skeleton. The overall physiological profile of all three species varied over time and that of P. divaricata was consistently different from the two other coral species. Porites divaricata had higher energy reserves coupled with higher contributions of heterotrophically derived carbon to host tissues than both P. astreoides and O. faveolata. Consistently higher overall energy reserves and heterotrophic contributions to tissues appear to buffer against environmental stress, including bleaching events. Thus, natural physiological variability among coral species appears to be a stronger predictor of coral bleaching resilience than intra- or inter-annual physiological variability within a coral species.

Abstract As coral reefs experience dramatic declines in coral cover throughout the tropics, there is an urgent need to understand the role that non-reef habitats, such as mangroves, play in the ecological niche of corals. Mangrove habitats present a challenge to reef-dwelling corals because they can differ dramatically from adjacent reef habitats with respect to key environmental parameters, such as light. Because variation in light within reef habitats is known to drive intraspecific differences in coral phenotype, we hypothesized that coral species that can exploit both reef and mangrove habitats will exhibit predictable differences in phenotypes between habitats. To investigate how intraspecific variation, driven by either local adaptation or phenotypic plasticity, might enable particular coral species to exploit these two qualitatively different habitat types, we compared the phenotypes of two widespread Caribbean corals, Porites divaricata and Porites astreoides, in mangrove versus lagoon habitats on Turneffe Atoll, Belize. We document significant differences in colony size, color, structural complexity, and corallite morphology between habitats. In every instance, the phenotypic differences between mangrove prop root and lagoon corals exhibited consistent trends in both P. divaricata and P. astreoides. We believe this study is the first to document intraspecific phenotypic diversity in corals occupying mangrove prop root versus lagoonal patch reef habitats. A difference in the capacity to adopt an alternative phenotype that is well suited to the mangrove habitat may explain why some reef coral species can exploit mangroves, while others cannot.

Corals with high levels of total lipids are known to have increased resilience potential to bleaching, and lipid class management may shed further light on why some species are more resilient to, or are able to acclimatize to, annual bleaching stress. Here, we measured the lipid class composition of three species of Caribbean corals (Porites astreoides, Porites divaricata, and Orbicella faveolata) collected in July 2009 near Puerto Morelos, Mexico (20° 50′ N, 86° 52′ W) that were experimentally bleached 2 years in a row. Our results show that single bleaching can significantly alter lipid class composition in all species, while repeated bleaching can result in stable (i.e., acclimatized) or even more altered (i.e., not acclimatized) lipid class composition depending on the species. Specifically, P. divaricata and O. faveolata both had altered lipid class composition with losses in storage lipids following single bleaching, but maintained lipid class composition following repeated bleaching stress. However, both single and repeated bleaching altered the lipid class composition in P. astreoides, with changes persisting for the 6 weeks after repeated bleaching stress. This study provides evidence that lipid class management is part of the suite of variables associated with coral resilience, that P. divaricata and O. faveolata acclimatize their lipid class management in response to repeated bleaching stress, but that P. astreoides does not. Corals like P. divaricata and O. faveolata may, therefore, be more suitable for coral restoration efforts since they are more likely to persist under chronic repeat bleaching scenarios predicted for later this century.

Long-term recovery of Caribbean corals from bleaching

During a recent reef coral survey at the submarine Saba Bank (Eastern Caribbean), an uncommon and diverse assemblage of unattached scleractinian corals (coralliths) was encountered, which has not been reported from the Atlantic before. Four different types of these free-living (unattached) corals were distinguished. They were observed on a relatively flat seafloor (15–20 m deep) with poor coral cover and full exposure to oceanic swell. Much of the substratum was not consolidated and consisted mainly of sand and fragments of branching coralline algae. One of the four types is the (1) anthocyathus stage in the life history of the free-living species Manicina areolata and Meandrina danae. The other three are coralliths formed as ecophenotypic varieties: (2) spheroidal–amoeboidal (= globular and (sub)massive) in Porites astreoides, Siderastrea radians, S. siderea, and Stephanocoenia intersepta; (3) tumbleweed-like (= globular and ramose) in Porites divaricata and P. furcata; and (4) discoidal (flat and circular with short branches) in Madracis decactis and possibly in M. cf. auretenra. This assemblage of free-living corals is likely related to a combination of abiotic factors consisting of wave exposure (swell), depths that waves can reach, a horizontal sea floor with little relief, an unconsolidated substratum, and low coral cover.

Annual coral bleaching events due to increasing sea surface temperatures are predicted to occur globally by the mid-century and as early as 2025 in the Caribbean, and severely impact coral reefs. We hypothesize that heterotrophic carbon (C) in the form of zooplankton and dissolved organic carbon (DOC) is a significant source of C to bleached corals. Thus, the ability to utilize multiple pools of fixed carbon and/or increase the amount of fixed carbon acquired from one or more pools of fixed carbon (defined here as heterotrophic plasticity) could underlie coral acclimatization and persistence under future ocean-warming scenarios. Here, three species of Caribbean coral—Porites divaricata, P. astreoides, and Orbicella faveolata—were experimentally bleached for 2.5 weeks in two successive years and allowed to recover in the field. Zooplankton feeding was assessed after single and repeat bleaching, while DOC fluxes and the contribution of DOC to the total C budget were determined after single bleaching, 11 months on the reef, and repeat bleaching. Zooplankton was a large C source for P. astreoides, but only following single bleaching. DOC was a source of C for single-bleached corals and accounted for 11–36 % of daily metabolic demand (CHARDOC), but represented a net loss of C in repeat-bleached corals. In repeat-bleached corals, DOC loss exacerbated the negative C budgets in all three species. Thus, the capacity for heterotrophic plasticity in corals is compromised under annual bleaching, and heterotrophic uptake of DOC and zooplankton does not mitigate C budget deficits in annually bleached corals. Overall, these findings suggest that some Caribbean corals may be more susceptible to repeat bleaching than to single bleaching due to a lack of heterotrophic plasticity, and coral persistence under increasing bleaching frequency may ultimately depend on other factors such as energy reserves and symbiont shuffling.

Mass bleaching events are predicted to occur annually later this century. Nevertheless, it remains unknown whether corals will be able to recover between annual bleaching events. Using a combined tank and field experiment, we simulated annual bleaching by exposing three Caribbean coral species (Porites divaricata, Porites astreoides and Orbicella faveolata) to elevated temperatures for 2.5 weeks in 2 consecutive years. The impact of annual bleaching stress on chlorophyll a, energy reserves, calcification, and tissue C and N isotopes was assessed immediately after the second bleaching and after both short- and long-term recovery on the reef (1.5 and 11 months, respectively). While P. divaricata and O. faveolata were able to recover from repeat bleaching within 1 year, P. astreoides experienced cumulative damage that prevented full recovery within this time frame, suggesting that repeat bleaching had diminished its recovery capacity. Specifically, P. astreoides was not able to recover protein and carbohydrate concentrations. As energy reserves promote bleaching resistance, failure to recover from annual bleaching within 1 year will likely result in the future demise of heat-sensitive coral species.

There has been ongoing flattening of Caribbean coral reefs with the loss of habitat having severe implications for these systems. Complexity and its structural components are important to fish species richness and community composition, but little is known about its role for other taxa or species-specific responses. This study reveals the importance of reef habitat complexity and structural components to different taxa of macrofauna, total species richness, and individual coral and fish species in the Caribbean. Species presence and richness of different taxa were visually quantified in one hundred 25-m(2) plots in three marine reserves in the Caribbean. Sampling was evenly distributed across five levels of visually estimated reef complexity, with five structural components also recorded: the number of corals, number of large corals, slope angle, maximum sponge and maximum octocoral height. Taking advantage of natural heterogeneity in structural complexity within a particular coral reef habitat (Orbicella reefs) and discrete environmental envelope, thus minimizing other sources of variability, the relative importance of reef complexity and structural components was quantified for different taxa and individual fish and coral species on Caribbean coral reefs using boosted regression trees (BRTs). Boosted regression tree models performed very well when explaining variability in total (82·3%), coral (80·6%) and fish species richness (77·3%), for which the greatest declines in richness occurred below intermediate reef complexity levels. Complexity accounted for very little of the variability in octocorals, sponges, arthropods, annelids or anemones. BRTs revealed species-specific variability and importance for reef complexity and structural components. Coral and fish species occupancy generally declined at low complexity levels, with the exception of two coral species (Pseudodiploria strigosa and Porites divaricata) and four fish species (Halichoeres bivittatus, H. maculipinna, Malacoctenus triangulatus and Stegastes partitus) more common at lower reef complexity levels. A significant interaction between country and reef complexity revealed a non-additive decline in species richness in areas of low complexity and the reserve in Puerto Rico. Flattening of Caribbean coral reefs will result in substantial species losses, with few winners. Individual structural components have considerable value to different species, and their loss may have profound impacts on population responses of coral and fish due to identity effects of key species, which underpin population richness and resilience and may affect essential ecosystem processes and services.

Mass coral bleaching events caused by elevated seawater temperatures result in extensive coral loss throughout the tropics, and are projected to increase in frequency and severity. If bleaching becomes an annual event later in this century, more than 90% of coral reefs worldwide may be at risk of long-term degradation. While corals can recover from single isolated bleaching and can acclimate to recurring bleaching events that are separated by multiple years, it is currently unknown if and how they will survive and possibly acclimatize to annual coral bleaching. Here, we demonstrate for the first time that annual coral bleaching can dramatically alter thermal tolerance in Caribbean corals. We found that high coral energy reserves and changes in the dominant algal endosymbiont type (Symbiodinium spp.) facilitated rapid acclimation in Porites divaricata, whereas low energy reserves and a lack of algal phenotypic plasticity significantly increased susceptibility in Porites astreoides to bleaching the following year. Phenotypic plasticity in the dominant endosymbiont type of Orbicella faveolata did not prevent repeat bleaching, but may have facilitated rapid recovery. Thus, coral holobiont response to an isolated single bleaching event is not an accurate predictor of its response to bleaching the following year. Rather, the cumulative impact of annual coral bleaching can turn some coral species 'winners' into 'losers', and can also facilitate acclimation and turn some coral species 'losers' into 'winners'. Overall, these findings indicate that cumulative impact of annual coral bleaching could result in some species becoming increasingly susceptible to bleaching and face a long-term decline, while phenotypically plastic coral species will acclimatize and persist. Thus, annual coral bleaching and recovery could contribute to the selective loss of coral diversity as well as the overall decline of coral reefs in the Caribbean.

Scleractinian corals were exposed to 6 combinations of temperature and solar radiation to evaluate effects on coral bleaching, survival, and tissue surface area changes during and after exposure. A recirculating coral exposure system was coupled to a solar simulator to allow laboratory testing of 6 species of Caribbean corals (Diploria clivosa, Montastraea faveolata, Porites divaricata, Stephanocoenia intersepta, Siderastrea radians, and Siderastrea siderea). Significant bleaching occurred in all of the corals exposed to high irradiance except S. siderea. Elevated light levels resulted in a decrease in photochemical efficiency for all species during the exposure period, with S. siderea showing the smallest decrease. The most prominent reductions in photochemical efficiency occurred in M. faveolata and S. intersepta, and these species exhibited extensive tissue loss and the highest mortality. In contrast to high irradiance, high temperatures significantly decreased photochemical efficiency for only D. clivosa and did not lead to severe tissue loss for this species. These results demonstrate species-specific responses to solar radiation and temperatures, with M. faveolata and S. intersepta being the most susceptible to bleaching due to high irradiance.

Rising atmospheric CO2 and its equilibration with surface ocean seawater is lowering both the pH and carbonate saturation state (Ω) of the oceans. Numerous calcifying organisms, including reef-building corals, may be severely impacted by declining aragonite and calcite saturation, but the fate of coral reef ecosystems in response to ocean acidification remains largely unexplored. Naturally low saturation (Ω ~ 0.5) low pH (6.70–7.30) groundwater has been discharging for millennia at localized submarine springs (called “ojos”) at Puerto Morelos, México near the Mesoamerican Reef. This ecosystem provides insights into potential long term responses of coral ecosystems to low saturation conditions. In-situ chemical and biological data indicate that both coral species richness and coral colony size decline with increasing proximity to low-saturation, low-pH waters at the ojo centers. Only three scleractinian coral species (Porites astreoides, Porites divaricata, and Siderastrea radians) occur in undersaturated waters at all ojos examined. Because these three species are rarely major contributors to Caribbean reef framework, these data may indicate that today’s more complex frame-building species may be replaced by smaller, possibly patchy, colonies of only a few species along the Mesoamerican Barrier Reef. The growth of these scleractinian coral species at undersaturated conditions illustrates that the response to ocean acidification is likely to vary across species and environments; thus, our data emphasize the need to better understand the mechanisms of calcification to more accurately predict future impacts of ocean acidification.

The present study examines coral bleaching along the Florida Keys reef tract (USA) dur- ing a major thermal-stress event in the summer of 2005, and during the summers of 2006 and 2007, which were mild thermal stress years. The primary objectives were to (1) examine the spatial pattern of coral bleaching and its relationship to coral community composition and habitat and (2) determine the relationship between environmental parameters and coral-bleaching prevalence (i.e. the propor- tion of colonies that bleached within each population). Over 50% of all coral species bleached along much of the reef tract in 2005. The Lower Keys supported more colonies per unit area than elsewhere and showed the highest number of bleached coral colonies; Biscayne and the Middle Keys showed the highest coral-bleaching prevalence. The most thermally sensitive corals were Porites furcata, Millepora complanata, Siderastrea radians, Porites divaricata, Agaricia agaricites, Millepora alcicor- nis, and Porites porites. The most extensive bleaching was recorded for large colonies, ≥30 cm, except in 2005 when small branching Porites colonies (<30 cm) showed higher bleaching prevalence. Shallow-water coral colonies and corals at localities with high productivity, in the form of chlorophyll a and dissolved inorganic nitrogen concentration, showed higher bleaching prevalence than both deep coral colonies and corals at localities with low productivity. By locally regulating waste-water discharge from the land, and thereby reducing local primary productivity, the severity of coral bleaching may be reduced when subjected to high regional water temperatures.

The synergistic effect of ultraviolet radiation (UVR) and a polycyclic aromatic hydrocarbon (PAH) was tested on the coral Porites divaricata. Small branches were incubated in different concentrations of fluoranthene followed by exposure to ecologically relevant levels of natural solar radiation, with and without UVR. Exposure to the highest concentration of fluoranthene (60 μ g L− 1) resulted in decreased photochemical efficiency of corals exposed concomitantly to UVR but not in corals exposed to 60 μ g L− 1 fluoranthene without UVR. After 6 days, most coral samples that were exposed to fluoranthene and UVR bleached or died (78%) on the upper side of the branches. At 60 μ g L− 1 fluoranthene with UVR, 11 out of 12 samples died and the remaining sample bleached, much greater than the number of samples that bleached in manipulation or solvent controls. On the under side of the same samples, where the coral polyps are naturally protected from UVR, 11 out of 12 samples remained healthy and intact. The high UVR doses that shallow water dwelling corals receive combined with the results of this study show that photoinduced toxicity of PAHs is a stress factor that needs to be studied in more detail in coral reef ecosystems.

Well-preserved, Holocene coral reefs provide the opportunity to discriminate between models of intrinsically driven succession and extrinsically driven species replacement, especially when paleontological patterns can be combined with ecological observations of the underlying mechanisms. Rhomboid shoals in the central shelf lagoon of the Belizean Barrier Reef experienced a recent and dramatic change in community composition. Agaricia tenuifolia replaced Acropora cervicornis as the dominant coral species at 3–15 m depth along the flanks of the reefs. We tested the hypothesis that shallowing upward caused this shift in dominance. A core extracted from 0.5 m water depth on one of the shoals, Channel Cay, revealed a shallowing-upward shift in dominance from Acropora to Porites divaricata. This successional sequence was quite different from the Acropora-to-Agaricia transition observed in four cores from 6–11 m water depth. Ecological observations showed that Agaricia became the dominant at ≥3 m depth after Acropora populations were decimated by a regional outbreak of white-band disease. The Acropora-to-Agaricia transition was clearly a case of extrinsically driven species replacement rather than an intrinsically driven, successional, shallowing-upward sequence.

Animals rely on their diet for amino acids that they are incapable either of synthesizing or of synthesizing in sufficient quantities to meet metabolic needs. These are the so-called 'essential amino acids'. This set of amino acids is similar among the vertebrates and many of the invertebrates. Previously, no information was available for amino acid synthesis by the most primitive invertebrates, the Cnidaria. The purpose of this study was to examine amino acid synthesis by representative cnidarians within the Order Scleractinia. Three species of zooxanthellate reef coral, Montastraea faveolata, Acropora cervicornis and Porites divaricata, and two species of non-zooxanthellate coral, Tubastrea coccinea and Astrangia poculata, were incubated with 14C-labelled glucose or with the 14C-labelled amino acids glutamic acid, lysine or valine. Radiolabel tracer was followed into protein amino acids. A total of 17 amino acids, including hydroxyproline, were distinguishable by the techniques used. Of these, only threonine was not found radiolabelled in any of the samples. We could not detect tryptophan or cysteine, nor distinguish between the amino acid pairs glutamic acid and glutamine, or aspartic acid and asparagine. Eight amino acids normally considered essential for animals were made by the five corals tested, although some of them were made only in small quantities. These eight amino acids are valine, isoleucine, leucine, tyrosine, phenylalanine histidine, methionine and lysine. The ability of cnidarians to synthesize these amino acids could be yet another indicator of a separate evolutionary history of the cnidarians from the rest of the Metazoa.