Cyanobacteria Associated with Coral Black Band Disease in Caribbean and Indo-Pacific Reefs

For 30 years it has been assumed that a single species of cyanobacteria, Phormidium corallyticum, is the volumetrically dominant component of all cases of black band disease (BBD) in coral. Cyanobacterium-specific 16S rRNA gene primers and terminal restriction fragment length polymorphism analyses were used to determine the phylogenetic diversity of these BBD cyanobacteria on coral reefs in the Caribbean and Indo-Pacific Seas. These analyses indicate that the cyanobacteria that inhabit BBD bacterial mats collected from the Caribbean and Indo-Pacific Seas belong to at least three different taxa, despite the fact that the corals in each case exhibit similar signs and patterns of BBD mat development.     

Black Band Disease Microbial Community Variation on Corals in Three Regions of the Wider Caribbean

Black band disease (BBD) is a pathogenic consortium of microorganisms that primarily affects massive framework-building scleractinian corals on reefs worldwide. There has been considerable debate concerning the microbial community composition of BBD. The aim of this study was to utilize microbial profiling to assess overall patterns of variation in the BBD bacterial community with respect to geographic location, host coral species, time, and nutrient regime. Length heterogeneity polymerase chain reaction (LH-PCR) was employed to differentiate BBD communities based on the natural variation in the sequence lengths within hypervariable domains of the 16S rRNA gene. Analysis of LH-PCR profiles of 97 BBD samples using multivariate ordination methods and analysis of similarity revealed significant clustering with respect to geographic region when comparing BBD sampled from reefs near Lee Stocking Island in the Bahamas’ Exuma Chain, the Northern Florida Keys (NFK), and St. John in the US Virgin Islands. There was much variability in BBD community composition on a regional basis, between sites in the NFK, and in terms of coral host species. The observed differences among BBD microbial community profiles were driven primarily by variation in relative abundance of 313–316-bp amplicons, which correspond to cyanobacteria and α-proteobacteria. The results obtained in this study support previous reports of intrinsic variability and complexity of the BBD microbial community but also suggest that this variability has biogeographic patterns.     

Soundscapes from a Tropical Eastern Pacific reef and a Caribbean Sea reef

Underwater soundscapes vary due to the abiotic and biological components of the habitat. We quantitatively characterized the acoustic environments of two coral reef habitats, one in the Tropical Eastern Pacific (Panama) and one in the Caribbean (Florida Keys), over 2-day recording durations in July 2011. We examined the frequency distribution, temporal variability, and biological patterns of sound production and found clear differences. The Pacific reef exhibited clear biological patterns and high temporal variability, such as the onset of snapping shrimp noise at night, as well as a 400-Hz daytime band likely produced by damselfish. In contrast, the Caribbean reef had high sound levels in the lowest frequencies, but lacked clear temporal patterns. We suggest that acoustic measures are an important element to include in reef monitoring programs, as the acoustic environment plays an important role in the ecology of reef organisms at multiple life-history stages.     

Herbivore abundance and grazing intensity on a Caribbean coral reef

Herbivory is a primary factor in determining the structure of coral reef communities. Spatial variation among reef habitats in the intensity of herbivory has been documented, but underlying variation in species composition and abundance within the herbivore guild has received little attention. The distribution and relative abundances of herbivorous fishes and sea urchins across several habitats were studied on the Belizean barrier reef off the Caribbean coast of Central America. Marked variation in total herbivore density as well as major changes in the composition of the herbivore guild were found across reef habitats. Acanthurids (surgeonfishes) predominated in shallow areas (< 5 m) while scarids (parrotfishes) were dominant in deeper habitats. Significant differences among habitats in an experimental assay of grazing intensity were strongly correlated with herbivore abundance. The spatial distribution of herbivorous fishes across reef habitats does not appear to be simply explained by differences in reef topography, but may depend on complex interactions among proximity to nearby shelter, predator abundance, density of territorial competitors, and local availability of food resources.     

Algal resistance to herbivory on a Caribbean barrier reef

Field and laboratory research at Carrie Bow Cay, Belize showed that macroalgae, grouped in functional-form units resisted fish and urchin herbivory in the following order (from high to low resistance): Crustose-Group, Jointed Calcareous-Group, Thick Leathery-Group, Coarsely Branched-Group, Filamentous-Group and Sheet-Group; thereby supporting the hypothesis that crustose, calcareous and thick algae have evolved antipredator defenses and should show the greatest resistance to herbivory with a gradation of increasing palatability towards filaments and sheets. Of the 21 species examined, several (e.g.,Dictyota cervicornis on grids,Laurencia obtusa andStypopodium zonale) had exceptionally low losses to fish grazing, probably due to chemical defences. The sea urchin,Diadema antillarum, was more inclined to feed on algae with known toxic secondary metabolites than were herbivorous fishes; hypothetically related to the differences in mobility and concomitant modes of feeding. Tough leathery forms such asSargassum polyceratium andTurbinaria turbinata resisted grazing by bottom feeding parrotfishes (Scaridae) and surgeonfishes (Acanthuridae) but were susceptible when suspended midway in the water column, possibly due to the presence of rudderfishes (Kyphosidae) which readily consume drift Sargassaceae. The overall tendencies support our predicted relationship between grazer-resistance and algal morphology. In conjunction with our previously reported findings concerning primary productivity, toughness and calorimetry for many of the same species, these results lend credence to generalizations relating form with function in marine macroalgae.     

Macroalgae Has No Effect on the Severity and Dynamics of Caribbean Yellow Band Disease

By removing herbivores and promoting increases in macroalgae, overfishing is thought to indirectly cause coral disease and mortality. We performed three field manipulations to test the general hypothesis that overfishing and the subsequent alteration of coral reef trophic dynamics are a cause of coral epizootics. Specifically, we asked whether the presence of macroalgae can influence within- and among-colony spread rates of Caribbean Yellow Band Disease in Montastraea faveolata. Macroalgae were placed next to infected and healthy, adult and small coral colonies to measure effects on disease spread rate, coral growth and coral survival. Surprisingly, the addition of macroalgae did not affect disease severity or coral fitness. Our results indicate that macroalgae have no effect on the severity and dynamics of Caribbean Yellow Band Disease and that fisheries management alone will not mitigate the effects of this important epizootic.     

Habitat complexity and consumer-mediated positive feedbacks on a Caribbean coral reef

Theoretical and empirical evidence suggest that positive feedbacks can increase resilience in ecological communities. On Caribbean coral reefs, there have been striking shifts from physically complex communities with high coral cover to relatively homogenous communities dominated by macroalgae, which have persisted for decades. However, little is known about positive feedbacks that may maintain coral reef community states. Here, I explore a potential consumer-mediated feedback on a Jamaican reef by examining how grazing by a keystone herbivore ( Diadema antillarum ) is enhanced by physical structure, which offer refugia from predation. Surveys revealed that habitat complexity and Diadema density were positively related. Increasing habitat complexity by adding physical structure significantly decreased macroalgal cover and increased the proportion of urchins in algal habitats in field manipulations. Experimental increases in urchin density also decreased macroalgal cover, but did not affect the proportion of urchins in algal habitats. These results suggest that the low habitat complexity of macroalgal-dominated reefs may inhibit an urchin-mediated shift to coral dominance and that positive feedbacks must be considered in reef restoration efforts.     

Reproductive ecology of Caribbean reef corals

The last decade has seen a resurgence of interest in the processes of sexual reproduction by scleractinian reef corals. Earlier investigations had focused fortuitously on brooding (planulating) species, which resulted in the general misconception that brooding was the main form of larval development of reef corals. More recent work on Indo-Pacific species has shown broadcast spawning and short annual reproductive periods to predominate. This report presents the reproductive patterns of eleven Caribbean coral species and attempts to explain the adaptive features and selective pressures that have led to the evolution of the four reproductive patterns described to date: (a) hermaphroditic broadcasters; (b) gonochoric broadcasters; (c) hermaphroditic broadcasters; (b) gonochoric brooders. Both (a) and (b) correlate with large colony size and short annual spawning periods; and (c) and (d) correlate with small colony size, multiple planulating cycles per year, and occupation of unstable habitats. Selection for outcrossing between long-lived individuals is proposed as the reason for gonochorism and for synchronous spawning of hermaphroditic broadcasters, and also for the large amount of sperm produced by hermaphroditic brooders. Selection for high rates of local recruitment is proposed as the force behind the evolution of brooding by species inhabiting unstable habitats and suffering high rates of adult mortality.     

Bacterial Community Associated with Black Band Disease in Corals

Black band disease (BBD) is a virulent polymicrobial disease primarily affecting massive-framework-building species of scleractinian corals. While it has been well established that the BBD bacterial mat is dominated by a cyanobacterium, the quantitative composition of the BBD bacterial mat community has not described previously. Terminal-restriction fragment length polymorphism (T-RFLP) analysis was used to characterize the infectious bacterial community of the bacterial mat causing BBD. These analyses revealed that the bacterial composition of the BBD mat does not vary between different coral species but does vary when different species of cyanobacteria are dominant within the mat. On the basis of the results of a new method developed to identify organisms detected by T-RFLP analysis, our data show that besides the cyanobacterium, five species of the division Firmicutes, two species of the Cytophaga-Flexibacter-Bacteroides (CFB) group, and one species of δ-proteobacteria are also consistently abundant within the infectious mat. Of these dominant taxa, six were consistently detected in healthy corals. However, four of the six were found in much higher numbers in BBD mats than in healthy corals. One species of the CFB group and one species of Firmicutes were not always associated with the bacterial communities present in healthy corals. Of the eight dominant bacteria identified, two species were previously found in clone libraries obtained from BBD samples; however, these were not previously recognized as important. Furthermore, despite having been described as an important component of the pathogenetic mat, a Beggiatoa species was not detected in any of the samples analyzed. These results will permit the dominant BBD bacteria to be targeted for isolation and culturing experiments aimed at deciphering the disease etiology.     

Feeding complementarity versus redundancy among herbivorous fishes on a Caribbean reef

Herbivory is an important driver of community structure on coral reefs. Adequate understanding of herbivory will mandate better knowledge of how specific herbivores impact reef communities and the redundancy versus complementarity of their ecological roles. We used algal communities generated by herbivore manipulations to assess such roles among Caribbean herbivorous fishes. We created large enclosures on a 16- to 18-m-deep reef to create treatments grazed for 10 months by: (1) only Sparisoma aurofrenatum, (2) only Acanthurus bahianus, (3) no large herbivorous fishes, or (4) natural densities of all reef fishes. After 10 months, we removed cages and filmed how free-ranging reef fishes fed among these treatments that differed in algal community structure. In general, Acanthurus spp. and Scarus spp. rapidly grazed exclosure and Sparisoma-only treatments, while Sparisoma spp. preferentially grazed exclosure and Acanthurus-only treatments. These patterns suggest complementarity between Sparisoma spp. and both Acanthurus spp. and Scarus spp. but redundancy between Acanthurus spp. and Scarus spp. Despite these generalities, there was also within-genera variance in response to the different treatments. For example, large Scarus spp., such as Scarus guacamaia, fed more similarly to Sparisoma spp., particularly Sparisoma viride, than to other Scarus spp. Moreover, the three common Sparisoma species differed considerably in the macroalgae to which they exhibited positive or negative relationships. Thus, herbivorous reef fishes vary considerably in their response to different algal communities and exhibit complex patterns of compensatory feeding and functional redundancy that are poorly predicted by taxonomy alone.     

Recruitment hotspots and bottlenecks mediate the distribution of corals on a Caribbean reef

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

Context-dependent corallivory by parrotfishes in a Caribbean reef ecosystem

This study assesses the patterns of corallivory by parrotfishes across reefs of the Florida Keys, USA. These reefs represent a relatively unique combination within the wider Caribbean of low coral cover and high parrotfish abundance suggesting that predation pressure could be intense. Surveys across eight shallow forereefs documented the abundance of corals, corallivorous parrotfishes, and predation scars on corals. The corals Porites porites and Porites astreoides were preyed on most frequently with the rates of predation an order of magnitude greater than has been documented for other areas of the Caribbean. In fact, parrotfish bite density on these preferred corals was up to 34 times greater than reported for corals on other reefs worldwide. On reefs where coral cover was low and corals such as Montastraea faveolata, often preferred prey for parrotfishes, were rare, predation rates on P. porites and P. astreoides, and other less common corals, intensified further. The intensity of parrotfish predation increased significantly as coral cover decreased. However, parrotfish abundance showed only a marginal positive relationship with predation pressure on corals, likely because corallivorous parrotfish were abundant across all reefs. Parrotfishes often have significant positive impacts on coral cover by facilitating coral recruitment, survival, and growth via their grazing of algae. However, abundant corallivorous parrotfishes combined with low coral cover may result in higher predation on corals and intensify the negative impact that parrotfishes have on remaining corals.     

Vigilance and antipredator responses of Caribbean reef squid

Antipredator responses, especially those of open-ocean squid, have been seldom studied in the natural environment. Sepioteuthis sepioidea, observed by snorkellers near the shore in early morning/late afternoon, produced an average of eight moves of over 1?m per hour, apparently mostly antipredator behaviours. Close approaches by herbivorous parrotfish elicited no response in 74% of encounters; otherwise, squid produced agonistic zebra stripes or startle-mantle-dots skin patterns. Predatory bar jack fish caused flight but not zebra displays, and squid usually paled and fled quickly (66%) from snapper. The speed of approach was the best predictor for flight and display responses to snapper, but for bar jack and parrotfish, the relative fish size and distance were the predictors for escalated responses. Paired dorsolateral mantle dots were produced when squid approached the sea bottom or hunted outside the group and in reaction to fish approaches; 56% of these were to the very common parrotfish. Reactive pairs of spots were selected from four possible mantle locations and they were significantly likely to be directional towards fish, presumably as startle/warning, but not directed towards conspecifics as indicators of predator presence. Thus the evasion techniques of the cephalopods and their ability to produce different display patterns on the skin show how an intelligent animal modifies an otherwise simple decision about which pairs of spots to select from four possible mantle locations and whether to flee from potentially dangerous animals.     

Decadal-scale changes in abundance of non-scleractinian invertebrates on a Caribbean coral reef

The benthic community structure of the shallow reefs of St. John, US Virgin Islands, was studied from 1992 to 2007 to test the hypotheses that the abundances of non-scleractinian invertebrates have changed, and further, that the changes are associated with variation in the percentage cover of scleractinian coral and macroalgae. The study utilized photoquadrats (0.25 m²) from fringing reefs (7-9 m depth) characterized by igneous boulders and low (< 5%) coral cover, and results from six sites were pooled to describe these reefs as a single habitat. Photoquadrats were analyzed biennially for the abundance of invertebrates, particularly those belonging to four well represented classes (Anthozoa, Demospongiae, Echinoidea, and Polychaeta), as well as the cover of scleractinians and macroalgae. Overall, the combined (multivariate) abundance of 30 invertebrate taxa changed over time, the combined (multivariate) abundance of the four invertebrate classes changed over time, and the individual (univariate) abundance of anthozoans, sponges, and echinoids changed over time. Throughout the study, coral cover remained < 5%, and while it varied significantly, it did not display a consistent trajectory of change; in contrast, the cover of macroalgae increased throughout the study. While it is unsurprising that the abundances of invertebrates changed over 15 y, notably they varied even though coral cover remained stable, in only a few cases were they related positively to macroalgal cover, and in most cases it was members of the suspension feeding guild that became more abundant. These outcomes suggest that: (1) benthic invertebrates on the shallow reefs of St. John may be more strongly influenced by regional (e.g., larva supply) than local (e.g., coral cover) conditions, (2) Caribbean reefs have changed more since the early 1990s than can be inferred from variation in cover of coral and macroalgae, and (3) suspension feeding invertebrates have become more common on shallow fringing reefs in at least one location.     

Biogeographical homogeneity of Caribbean coral reef benthos

Caribbean reef benthic assemblages have been considered biogeographically homogeneous at regional scales, but this concept was recently challenged by Williams et al. (2015, Journal of Biogeography, 42 , 1327–1335). These authors concluded that benthic assemblages exhibit considerable biogeographical variability at regional and smaller scales, that rugosity and wave exposure play key roles in structuring assemblages, and that homogenization has yet to occur at a regional scale. We reassess their conclusions using recently published benthic and fish surveys that targeted sites either protected from fishing or intensively overfished. For sponges, regional variation in assemblages is mostly attributable to the removal of chemically undefended species by sponge‐eating fishes at sites protected from overfishing. We maintain that Caribbean benthic assemblages are remarkably homogeneous when compared to reefs in other tropical regions, and were likely more homogeneous before the localized effects of intensive fishing resulted in top‐down ecosystem alterations in benthic assemblages.     

Meteorology and hydrography of a shoaling reef flat on the Caribbean coast of Panama

We describe the seasonal patterns and frequency distributions of meteorological and hydrographic conditions on a windward, shoaling reef flat at Punta Galeta, Panama (9° 24 N lat.), between 1974 and 1985. The factors monitored were wind speed, wind direction, air temperature, rainfall, solar radiation, water level, water temperature, and salinity. All conditions showed strong seasonal periodicity; however, the timing, duration, and amplitude of the seasonal fluctuations differed among years. The greatest variation occurred in 1981 and 1982, leading into an El Niño event. Emergence of the reef flat and extremes of water temperature were the most apparent physiological stresses; both were dependent on mean water levels. The seasonal pattern of emergence time was inversely related to mean water level. Extreme water temperatures only occurred during low water tevels, ranging between 22° to >37°C in depths <15 cm, but staying between 25°C and 30.9°C in depths <35 cm. Water temperatures averaged 1.5°C higher than air temperatures. Although rainfall was 200 to 400 cm year^-1, salinity remained between 24 and 36 ppt, with more than 98% of the records 30 ppt. The relationship of water temperature to depth is consistent with the hypothesis that the physical environment becomes more stressful when a reef reaches sea level and forms a shoaling platform. Mailing address: Smithsonian Tropical Research Institute, APO Miami, Florida 34002-0011, USA     

The spawning seasons of Caribbean reef fishes

Observations are recorded on the time of spawning of 83 species of Caribbean reef fishes. Details of seasonal variations in the percentage of sexually active fishes are given for 35 species. The majority of species spawn mostly when water temperatures are minimal, in February, March and April. The expected biomass of reef fish eggs in the plankton is about twelve times greater in the above mentioned period than in the period from June to December. Normal recruitment to insular fisheries may be very greatly dependent upon the success of parent stocks spawning elsewhere in the Caribbean and the abundance of recruits to a particular area may be dependent upon the speed and direction of the prevailing currents which carry larvae and juveniles to the insular shelves.     

Island biogeography of Caribbean coral reef fish

Aim The goal of our study was to test fundamental predictions of biogeographical theories in tropical reef fish assemblages, in particular relationships between fish species richness and island area, isolation and oceanographic variables (temperature and productivity) in the insular Caribbean. These analyses complement an analogous and more voluminous body of work from the tropical Indo‐Pacific. The Caribbean is more limited in area with smaller inter‐island distances than the Indo‐Pacific, providing a unique context to consider fundamental processes likely to affect richness patterns of reef fish. Location Caribbean Sea. Methods We compiled a set of data describing reef‐associated fish assemblages from 24 island nations across the Caribbean Sea, representing a wide range of isolation and varying in land area from 53 to 110,860 km². Regression‐based analyses compared the univariate and combined effects of island‐specific physical predictors on fish species richness. Results We found that diversity of reef‐associated fishes increases strongly with increasing island area and with decreasing isolation. Richness also increases with increasing nearshore productivity. Analyses of various subsets of the entire data set reveal the robustness of the richness data and biogeographical patterns. Main conclusions Within the relatively small and densely packed Caribbean basin, fish species richness fits the classical species–area relationship. Richness also was related negatively to isolation, suggesting direct effects of dispersal limitation in community assembly. Because oceanic productivity was correlated with isolation, however, the related effects of system‐wide productivity on richness cannot be disentangled. These results highlight fundamental mechanisms that underlie spatial patterns of biodiversity among Caribbean coral reefs, and which are probably also are functioning in the more widespread and heterogeneous reefs of the Indo‐Pacific.     

Species density and associations in Caribbean reef corals

The density of scleractinian corals on portions of two reefs in the Grenadine Islands, W. I. has been investigated by direct observation by SCUBA divers. Grid, transect, and random quadrat methods were compared with total samples to determine the precision and accuracy of these techniques. Transect methods were generally satisfactory provided at least 15% of a 400 m² total grid area was included in the sample. The data strongly indicated clumped distributions of all species, although numerical analysis does not indicate the distinct zones reported by other workers. Species associations based on Jaccard's coefficient and cluster analysis showed possible similarities in physical requirements, although few strong associations were found. Data based on 4 m² quadrats generally provided a more reliable estimate of species associations than did data based on 1 m² quadrats. It is suggested that surveys of these reef types may be better based on a number of parallel transects rather than a single transect, and that well-defined zones are more likely to be the exception than the rule.