First report of folliculinid ciliates affecting Caribbean scleractinian corals

This is the first report of a ciliate of the genus Halofolliculina infecting hard coral species of six families (Acroporidae, Agaricidae, Astrocoeniidae, Faviidae, Meandrinidae and Poritidae) and milleporids in the Caribbean. Surveys conducted during 2004–2005 in Venezuela, Panama and México confirmed that this ciliate affects up to 25 scleractinian species. The prevalence of this ciliate at the coral community level was variable across sites, being most commonly found at Los Roques, Venezuela, and at Bocas del Toro, Panama (prevalence 0.2–2.5%), but rarely observed in the Mexican Caribbean. Ciliates were more prevalent within populations of acroporids (Acropora palmata, Acropora cervicornis and Acropora prolifera) in Los Roques. Recent observations also corroborate the presence of these ciliates in Curacao and Puerto Rico. Our observations indicate that ciliates affecting corals have a wider distribution than previously thought, and are no longer exclusively found in the Indo-Pacific and Red Sea.     

Coral mass- and split-spawning at a coastal and an offshore Venezuelan reefs,southern Caribbean

This study aimed to evaluate potential differences in coral spawning behavior between a fringing coastal reef and an offshore reef in the southern Caribbean. For this, scleractinian and gorgonian colonies (N = 324) of 21 species were mapped along eight transects, each 10-m long, at two study sites located in Morrocoy and Los Roques National Parks, Venezuela. Observations were made between 19:30 and 23:00 from August 23 to 30 and from September 26 to 30, 2002. Ninety one colonies belonging to six hard coral and seven octocoral species spawned or planulated during this period. We were unable to observe any signs of reproductive activity in 95 colonies of nine species different from those that reproduced. Despite the differences in environmental conditions between the two sites, we observed high synchrony in the spawning behavior of seven coral species common to both reefs. The most striking result was the ability of colonies of Montastraea faveolata and Eusmilia fastigiata to split spawn up to three times, either in consecutive nights or in different months.     

Evaluating Methods for Transplanting Endangered Elkhorn Corals in the Virgin Islands

Restoration of rare corals is desirable and restoration projects are fairly common, but scientific evaluation of this approach is limited. We tested several techniques for transplant and restabilization of Acropora palmata (the elkhorn coral), an ecologically important Caribbean coral whose populations have suffered severe declines. In rough weather, fragments break‐off colonies of branching corals like A. palmata as a normal form of asexual reproduction. We transplanted naturally produced coral fragments from remnant populations to nearby restoration sites. Untouched control fragments at the donor site died faster and grew slower than fragments attached to the reef, so attaching fragments to the reef is beneficial. Transplanted fragments grew and died at a rate similar to fragments left at the donor site (both groups were attached to the reef), so there were no effects of moving fragments or differences in habitat quality between donor and restoration sites. Growth and survival were similar using four methods of attaching fragments to the reef: cable ties, two types of epoxy resin, and hydrostatic cement. Corals sometimes compete with the macroalgae that dominate degraded reefs, and clearing surrounding algae improved the growth of fragments. After 4 years, transplanted fragments grew to 1,450 cm² in area and so were potentially sexually active. Because the methods tested are simple and cheap, they could be used by volunteer recreational divers to restore locally extirpated A. palmata populations or to enhance reefs where natural recovery is slow.     

High-latitude<Emphasis Type="Italic"> Acropora cervicornis</Emphasis> thickets off Fort Lauderdale,Florida, USA

Baseline studies conducted in 1998 document the presence of robust, non-reef-building Acropora cervicornis thickets in shallow (3–7 m depth), near-shore waters off the coast of Fort Lauderdale, Florida, USA. These thickets thrive in a high-latitude environment, the northernmost in the continental USA, in the midst of potential anthropogenic stressors. Within thickets, the spatial variation in mean percent coral cover, macroalgal cover, scleractinian species richness, density of A. cervicornis juveniles, and the density and size of A. cervicornis colonies and fragments were recorded. Thicket size ranged between ~0.1 and 0.8 ha and mean coral cover varied between ~5 and 28%, with A. cervicornis accounting for ~87–97% of all scleractinians. Mean A. cervicornis colony and fragment densities per thicket were 1.3–3.3 colonies m^–2 and 0.7–2.8 fragments m^–2, respectively. Recruit densities varied between 0 and 1 ind. m^–2. White band disease was detected at all thickets, with a mean A. cervicornis colony surface area affected of 1.8%. For all thickets, densities of the corallivorous polychaete Hermodice carunculata ranged between ~18 and 86 ind. ha^–1, with predation scars affecting <0.2% of the A. cervicornis cover. These flourishing A. cervicornis thickets off Fort Lauderdale provide an interesting counterpoint to the declining and disease-stricken A. cervicornis populations reported in the Florida Keys and wider Caribbean.     

Assessment of Acropora palmata in the Mesoamerican Reef System

The once-dominant shallow reef-building coral Acropora palmata has suffered drastic geographical declines in the wider Caribbean from a disease epidemic that began in the late 1970s. At present there is a lack of quantitative data to determine whether this species is recovering over large spatial scales. Here, we use quantitative surveys conducted in 107 shallow-water reef sites between 2010 and 2012 to investigate the current distribution and abundance of A. palmata along the Mesoamerican Reef System (MRS). Using historical data we also explored how the distribution and abundance of this species has changed in the northern portion of the MRS between 1985 and 2010–2012. A. palmata was recorded in only a fifth of the surveyed reef sites in 2010–2012. In the majority of these reef sites the presence of A. palmata was patchy and rare. Only one site (Limones reef), in the northernmost portion of the MRS, presented considerably high A. palmata cover (mean: 34.7%, SD: 24.5%). At this site, the size-frequency distribution of A. palmata colonies was skewed towards small colony sizes; 84% of the colonies were healthy, however disease prevalence increased with colony size. A comparison with historical data showed that in the northern portion of the MRS, in 1985, A. palmata occurred in 74% of the 31 surveyed sites and had a mean cover of 7.7% (SD = 9.0), whereas in 2010–2012 this species was recorded in 48% of the sites with a mean cover of 2.9% (SD = 7.5). A. palmata populations along the MRS are failing to recover the distribution and abundance they had prior to the 1980s. Investigating the biological (e.g., population genetics) and environmental conditions (e.g., sources of stress) of the few standing reefs with relatively high A. palmata cover is crucial for the development of informed restoration models for this species.     

Bleaching increases likelihood of disease on Acropora palmata (Lamarck) in Hawksnest Bay, St John, US Virgin Islands

Anomalously high water temperatures may enhance the likelihood of coral disease outbreaks by increasing the abundance or virulence of pathogens, or by increasing host susceptibility. This study tested the compromised-host hypothesis, and documented the relationship between disease and temperature, through monthly monitoring of Acropora palmata colonies from May 2004 to December 2006, in Hawksnest Bay, St John, US Virgin Islands (USVI). Disease prevalence and the rate of change in prevalence showed a positive linear relationship with water temperature and rate of change in water temperature, respectively, but only in 2005 during prolonged periods of elevated temperature. Both bleached and unbleached colonies showed a positive relationship between disease prevalence and temperature in 2005, but the average area of disease-associated mortality increased only for bleached corals, indicating host susceptibility, rather than temperature per se, influenced disease severity on A. palmata.     

Bioerosion caused by foraging of the tropical chiton Acanthopleura gemmata at One Tree Reef, southern Great Barrier Reef

The bioerosive potential of the intertidal chiton Acanthopleura gemmata on One Tree Reef was determined by quantification of CaCO₃ in daily faecal pellet production of individuals transplanted into mesocosms after nocturnal-feeding forays. Mean bioerosive potential was estimated at 0.16 kg CaCO₃ chiton⁻¹ yr⁻¹. Bioerosion rates were estimated for populations on two distinct chiton habitats, reef margin (0.013 kg CaCO₃ m⁻² yr⁻¹) and beachrock platform (0.25 kg CaCO₃ m⁻² yr⁻¹). Chiton density on the platform was orders of magnitude greater than on the reef margin. The surface-lowering rate (0.16 mm m⁻² yr) due to bioerosion by the beachrock population is a substantial contribution to the total surface-lowering rate of 2 mm m⁻² yr⁻¹ previously reported for One Tree Reef across all erosive agents. At high densities, the contribution of A. gemmata to coral reef bioerosion budgets may be comparable to other important bioeroders such as echinoids and fish.     

Genetic Susceptibility,Colony Size,and Water Temperature Drive White-Pox Disease on the Coral Acropora palmata

Outbreaks of coral diseases are one of the greatest threats to reef corals in the Caribbean, yet the mechanisms that lead to coral diseases are still largely unknown. Here we examined the spatial-temporal dynamics of white-pox disease on Acropora palmata coral colonies of known genotypes. We took a Bayesian approach, using Integrated Nested Laplace Approximation algorithms, to examine which covariates influenced the presence of white-pox disease over seven years. We showed that colony size, genetic susceptibility of the coral host, and high-water temperatures were the primary tested variables that were positively associated with the presence of white-pox disease on A. palmata colonies. Our study also showed that neither distance from previously diseased individuals, nor colony location, influenced the dynamics of white-pox disease. These results suggest that white-pox disease was most likely a consequence of anomalously high water temperatures that selectively compromised the oldest colonies and the most susceptible coral genotypes.     

Effects of microhabitat characteristics on the settlement and recruitment of a coral reef fish at two spatial scales

Populations of fishes on coral reefs are replenished by the settlement of pelagic larvae to demersal populations. Recruitment varies spatially and temporally and can exert strong effects on the dynamics of reef fish populations. This study examined the effect of microhabitat characteristics on small-scale and large-scale recruitment variation in the three-spot damselfish, Stegastes planifrons (Cuvier). Comparison of 0.25-m² quadrats occupied by three-spots with randomly sampled null quadrats showed that three-spots quadrats contained a higher percent cover of the coral Montastrea annularis than would be expected at random. Manipulative experiments on three types of 1.0-m² patch reefs (living M. annularis, dead Porites Porites and dead Acropora palmata) patch reefs on showed that this non-random distribution was established by microhabitat choice during settlement and not by differential post-settlement survival. The presence of conspecific juveniles did not affect settlement. Recruitment was monitored at nine sites on three islands over 3 years. Recruitment showed no consistent pattern in the relative levels of recruitment among sites. Similarly, no consistent relationship emerged between recruitment levels and microhabitat characteristics at the nine sites. For example, at this large scale, the percent cover of M. annularis explained variation in recruitment in only 1 out of 3 years. These results suggest that small-scale recruitment patterns are influenced by microhabitat choice during settlement, but that these habitat effects do not scale up to influence large-scale variation in recruitment.     

Discovery and quantification of anaerobic nitrogen metabolisms among oxygenated tropical Cuban stony corals

Coral reef health depends on an intricate relationship among the coral animal, photosynthetic algae, and a complex microbial community. The holobiont can impact the nutrient balance of their hosts amid an otherwise oligotrophic environment, including by cycling physiologically important nitrogen compounds. Here we use ¹⁵N-tracer experiments to produce the first simultaneous measurements of ammonium oxidation, nitrate reduction, and nitrous oxide (N₂O) production among five iconic species of reef-building corals (Acropora palmata, Diploria labyrinthiformis, Orbicella faveolata, Porites astreoides, and Porites porites) in the highly protected Jardines de la Reina reefs of Cuba. Nitrate reduction is present in most species, but ammonium oxidation is low potentially due to photoinhibition and assimilatory competition. Coral-associated rates of N₂O production indicate a widespread potential for denitrification, especially among D. labyrinthiformis, at rates of ~1 nmol cm⁻² d⁻¹. In contrast, A. palmata displays minimal active nitrogen metabolism. Enhanced rates of nitrate reduction and N₂O production are observed coincident with dark net respiration periods. Genomes of bacterial cultures isolated from multiple coral species confirm that microorganisms with the ability to respire nitrate anaerobically to either dinitrogen gas or ammonium exist within the holobiont. This confirmation of anaerobic nitrogen metabolisms by coral-associated microorganisms sheds new light on coral and reef productivity.Subject terms: Biogeochemistry, Biogeochemistry     

Attributing mortality among drivers of population decline in <Emphasis Type="Italic">Acropora palmata</Emphasis> in the Florida Keys (USA)

Acropora palmata populations have experienced steep declines over the past 30 years. Although numerous culprits are recognized, their relative contributions to the decline are poorly quantified, making it difficult to prioritize effective conservation measures. In 2004, a demographic monitoring program was implemented in the Florida Keys (USA), aimed at determining the relative importance of various stressors affecting A. palmata. A subset of randomly selected A. palmata colonies within 15 fore-reef plots was tagged and surveyed three to four times per year over 7 years. Colony size, live tissue, prevalence of disease, snail (Coralliophila abbreviata) predation, physical damage and other conditions were assessed at each survey. The estimated effect of each condition causing recent mortality was ranked, and together, these parameters were used to attribute the population-level tissue loss associated with each condition. In addition, all new colonies in the study plots were counted and assessed annually in order to track trends in total colony count and live tissue abundance. Between 2004 and 2010, the study population has shown more than 50% decline in live area from three main conditions: fragmentation, disease and snail predation. Approximately half of this decline occurred during the catastrophic 2005 hurricane season from which recovery has been minimal. Meanwhile, colony abundance has shown gradual decline throughout the study. Snail predation was the most prevalent condition. However, it ranked third in attributed tissue loss, behind breakage that occurred during the 2005 hurricane season, and disease. Thermal bleaching of A. palmata was not observed during this study. Because mortality continues to outpace recruitment and growth, intervention to ameliorate losses to the more manageable threats including predation and breakage could result in substantial conservation of live tissue, buying time for the abatement of less tractable threats to A. palmata recovery such as climate change and disease.     

Severity of the 1998 and 2005 bleaching events in Venezuela, southern Caribbean

This study describes the severity of the 2005 bleaching event at 15 reef sites across Venezuela and compares the 1998 and 2005 bleaching events at one of them. During August and September 2005, bleached corals were first observed on oceanic reefs rather than coastal reefs, affecting 1 to 4% of coral colonies in the community (3 reef sites, n = 736 colonies). At that time, however, no bleached corals were recorded along the eastern coast of Venezuela, an area of seasonal upwelling (3 reefs, n = 181 colonies). On coastal reefs, bleaching started in October but highest levels were reached in November 2005 and January 2006, when 16% of corals were affected among a wide range of taxa (e.g. scleractinians, octocorals, Millepora and zoanthids). In the Acropora habitats of Los Roques (an oceanic reef),no bleached was recorded in 2005 (four sites,n = 643 colonies). At Cayo Sombrero, a coastal reef site, bleaching was less severe in 1998 than in 2005 (9% of the coral colonies involving 2 species vs. 26% involving 23 species, respectively). Our results indicate that bleaching was more severe in 2005 than in 1998 on Venezuelan reefs; however, no mass mortality was observed in either of these two events.     

Present state of diseases and other signs of reef deterioration at seven coral reefs at Los Roques Archipelago National Park, Venezuela

This work was aimed to determine the incidence of coral diseases in six different reef sites at the Parque Nacional Archipiélago de Los Roques, Venezuela: Arrecife de herradura, Arrecife costanero, both at Dos Mosquises Sur Key, Boca de Cote, Carenero, Crasquí and Pelona de Rabusquí. Each reef was surveyed by using ten 10 m2-band transects (10 x 1 m), placed parallel to the long axis of the reef within a depth gradient ranging from 1 to 9 m depth. All healthy and injured corals, along each band transect, were counted and identified to species level. Additionally, all diseases and recent mortality that were still identifiable on each colony were also recorded. The occurrence of diseased colonies and other signs of reef decline between localities were compared by means of a Chi2 test. The absolute, relative and mean incidence was estimated for each disease and other signs of damage observed for all coral species surveyed at each site. The overall incidence of coral diseases was low for all the localities surveyed, only 6.04% of the 3 344 colonies observed, showed signs of diseases. The most important diseases recorded were the Yellow-Blotch Disease (YBD) and Dark Spots Disease (DSD) with 2.1% +/- 1.52 y 2.1% +/- 2.54, respectively. Significant differences were found in the incidence of coral diseases between reef sites (Chi2 p < 0.05). Finally, the occurrence of colonies injured by parrotfish bites and pomacentrids was higher compared with the incidence of coral diseases for all the reefs surveyed. In conclusion, currently the proportion of healthy colonies at Los Roques coral reefs is higher than the percentage of both diseased and injured colonies.     

Adaptive photosynthetic strategies of the Mediterranean maquis species according to their origin

In consideration of their origin the adaptive strategies of the evergreen species of the Mediterranean maquis were analysed. Rosmarinus officinalis L., Erica arborea L., and Erica multiflora L. had the lowest net photosynthetic rate (P_N) in the favourable period [7.8±0.6 mol(CO₂) m^–2s^–1, mean value], the highest P_N decrease (on an average 86 % of the maximum) but the highest recovery capacity (>70 % of the maximum) at the first rainfall in September. Cistus incanus L. and Arbutus unedo L. had the highest P_N during the favourable period [15.5±5.2 mol(CO₂) m^–2s^–1, mean value], 79 % decrease during drought, and a lower recovery capacity (on an average 54 %). Quercus ilex L., Phillyrea latifolia L., and Pistacia lentiscus L. had an intermediate P_N in the favourable period [9.2±1.3 mol(CO2) m^–2s^–1, mean value], a lower reduction during drought (on an average 63 %), and a range from 62 % (Q. ilex and P. latifolia) to 39 % (P. lentiscus) of recovery capacity. The Mediterranean species had higher decrease in P_N and stomatal conductance during drought and a higher recovery capacity than the pre-Mediterranean species. Among the pre-Mediterranean species, P. latifoliahad the best adaptation to long drought periods also by its higher leaf mass per area (LMA) which lowered leaf temperature thus decreasing transpiration rate during drought. Moreover, its leaf longevity determined a more stable leaf biomass during the year. Among the Mediteranean species, R. officinalis was the best adapted species to short drought periods by its ability to rapidly recover. Nevertheless, R. officinalis had the lowest tolerance to high temperatures by its P_N dropping below half its maximum value when leaf temperature was over 33.6°C. R. officinalismay be used as a bioindicator species of global change.This revised version was published online in March 2005 with corrections to the page numbers.     

Nitrogen use efficiency of sugarcane in relation to its BNF potential and population of endophytic diazotrophs at different N levels

The nitrogen fixing bacterial endophytes Gluconacetobacter diazotrophicus and Herbaspirillum spp. have been proposed to benefit sugarcane (Saccaharum spp. hybrids) growth. Variable populations of these endophytes exist depending upon ontogenic and climatic variations as well. This study investigates the effect of variable chemical nitrogen application in soil on the population of endophytic diazotrophs, acetylene reduction ability of excised roots, plant N-nutrient use efficiency and probable interactions among different parameters in eight commercial sugarcane varieties of subtropical India. Recovery efficiency (RE), agronomic efficiency (AE), partial factor productivity (PFP) and physiologic efficiency (PE) indicators were used for accounting N-nutrient use efficiency. The population of G. diazotrophicus was more at N₇₅ compared to N₀ and N₁₅₀, whereas Herbaspirillum population increased from N₀ to N₁₅₀. ARA was positively correlated with Gluconacetobacter population in rhizosphere and root, whereas it had poor correlation with Herbaspirillum population. Positive correlation of RE and AE with ARA of roots, Gluconacetobacter and Herbaspirillum populations in roots and stems indicate their positive contribution in total nitrogen uptake by the plant per kg of N applied. Average PFP was 808.9 at N₇₅ compared to 408.7 at N₁₅₀ indicating that N was utilized efficiently at low N input status in sugarcane. Strong positive correlations of AE₇₅ (agronomic efficiency from 75 kg N ha⁻¹ to 150 kg N ha⁻¹) with N-uptake (r ² = 0.615), cane yield (r ² = 0.758) and PFP (r ² = 0.758) and other parameters compared to AE (agronomic efficiency from 0 kg N ha⁻¹ to 75 kg N ha⁻¹ or 150 kg N ha⁻¹) correlations with N-uptake (r ² = 0.111), cane yield (r ² = 0.368) and PFP (r ² = 0.190) indicated that the AE of sugarcane was strongly directed towards producing more cane yield per unit of N fertilizer once the sugarcane plant has established using initial dose of nitrogen and thus AE₇₅ seems to be a more appropriate indicator for accounting N-nutrient use efficiency in sugarcane.     

Relationship between net photosynthesis and leaf respiration in Mediterranean evergreen species

The relationship between net photosynthetic (P _N) and leaf respiration (R) rates of Quercus ilex, Phillyrea latifolia, Myrtus communis, Arbutus unedo, and Cistus incanus was monitored in the period February 2006 to February 2007. The species investigated had low R and P _N during winter, increasing from March to May, when mean air temperature reached 19.2 °C. During the favourable period, C. incanus and A. unedo had a higher mean P _N (16.4±2.4 μmol m⁻² s⁻¹) than P. latifolia, Q. ilex, and M. communis (10.0±1.3 μmol m⁻² s⁻¹). The highest R (1.89±0.30 μmol m⁻² s⁻¹, mean of the species), associated to a significant P _N decrease (62 % of the maximum, mean value of the species), was measured in July (mean R/P _N ratio 0.447±0.091). Q₁₀, indicating the respiration sensitivity to short-term temperature increase, was in the range 1.49 to 2.21. Global change might modify R/P _N determining differences in dry matter accumulation among the species, and Q. ilex and P. latifolia might be the most favoured species by their ability to maintain sufficiently higher P _N and lower R during stress periods.     

Rapid phase-shift reversal on a Jamaican coral reef

Many Caribbean reefs have experienced a phase-shift in community structure, the principle features being a decline in coral cover and an increase in macroalgal biomass. However, one Jamaican reef—Dairy Bull on the north shore near Discovery Bay—is once again dominated by scleractinian corals and several key species have returned. Living coral cover at 6–8 m depth at Dairy Bull has doubled over the past 9 years and is now ~54%. The absolute cover of Acropora cervicornis was <1% in 1995, but increased to ~11% by January 2004. During this time the cover of macroalgae decreased by 90%, from 45 to 6%. We speculate that long-lived colonies of Montastraea annularis may have facilitated the recovery of this reef by providing structural refugia.     

New insights into the dynamics between reef corals and their associated dinoflagellate endosymbionts from population genetic studies

The mutualistic symbioses between reef‐building corals and micro‐algae form the basis of coral reef ecosystems, yet recent environmental changes threaten their survival. Diversity in host‐symbiont pairings on the sub‐species level could be an unrecognized source of functional variation in response to stress. The Caribbean elkhorn coral, Acropora palmata, associates predominantly with one symbiont species (Symbiodinium ‘fitti’), facilitating investigations of individual‐level (genotype) interactions. Individual genotypes of both host and symbiont were resolved across the entire species’ range. Most colonies of a particular animal genotype were dominated by one symbiont genotype (or strain) that may persist in the host for decades or more. While Symbiodinium are primarily clonal, the occurrence of recombinant genotypes indicates sexual recombination is the source of this genetic variation, and some evidence suggests this happens within the host. When these data are examined at spatial scales spanning the entire distribution of A. palmata, gene flow among animal populations was an order of magnitude greater than among populations of the symbiont. This suggests that independent micro‐evolutionary processes created dissimilar population genetic structures between host and symbiont. The lower effective dispersal exhibited by the dinoflagellate raises questions regarding the extent to which populations of host and symbiont can co‐evolve during times of rapid and substantial climate change. However, these findings also support a growing body of evidence, suggesting that genotype‐by‐genotype interactions may provide significant physiological variation, influencing the adaptive potential of symbiotic reef corals to severe selection.     

Revised paleoenvironmental analysis of the Holocene portion of the Barbados sea-level record: Cobbler’s Reef revisited

Sample elevations corrected for tectonic uplift and assessed relative to local modeled sea levels provide a new perspective on paleoenvironmental history at Cobbler’s Reef, Barbados. Previously, ¹⁴C-dated surface samples of fragmented Acropora palmata plotted above paleo sea level based on their present (uplifted) elevations, suggesting supratidal rubble deposited during a period of extreme storms (4500–3000 cal BP), precipitating reef demise. At several sites, however, A. palmata persisted, existing until ~370 cal BP. Uplift-corrected A. palmata sample elevations lie below the western Atlantic sea-level curve, and ~2 m below ICE-6G-modeled paleo sea level, under slow rates of sea-level rise, negating the possibility that Cobbler’s Reef is a supratidal storm ridge. Most sites show limited age ranges from corals likely damaged/killed on the reef crest, not the mixed ages of rubble ridges, strongly suggesting the reef framework died off in stages over 6500 yr. Reef crest death assemblages invoke multiple paleohistoric causes, from ubiquitous hurricanes to anthropogenic impacts. Comparison of death assemblage ages to dated regional paleotempestological sequences, proxy-based paleotemperatures, recorded hurricanes, tsunamis, European settlement, deforestation, and resulting turbidity, reveals many possible factors inimical to the survival of A. palmata along Cobbler’s Reef.