Holocene growth history of an eastern Pacific fringing reef,Punta Islotes,Costa Rica

Rock and sediment cores reveal that a well-developed fringing reef in Golfo Dulce, Pacific Costa Rica, up to 9 m thick was established on Cretaceous basalt about 5500 y BP. It is presently being smothered with fine sediments and is almost completely dead. This reef is made up of three main facies that are represented by comparable extant reef zones: reef-flat branching coral, fore-reef slope massive coral, and fore-reef talus sediment facies. Reef growth began with the establishment of small patch reefs dominantly formed by the branching coral Pocillopora damicornis. P. damicornis spread across the basalt bench and massive colonies of Porites lobata grew on the outer slopes, eventually blocking the seaward transport of Pocillopora fragments to the fore-reef talus sediments. The reef flourished until 500 years ago. Lower accumulation rates during the past 500 years may be due to deteriorating environmental conditions rather than slower growth after the reef reached sea level. Present-day reef communities are severely degraded with less than 2% living coral cover. The increased turbidity associated with the final stage of degradation of this reef is probably related to human activity on the adjacent shores, including deforestation, mining, and road construction.     

A precise and non-destructive method to calculate the surface area in living scleractinian corals using X-ray computed tomography and 3D modeling

The surface area of corals represents a major reference parameter for the standardization of flux rates, for coral growth investigations, and for investigations of coral metabolism. The methods currently used to determine the surface area of corals are rather approximate approaches lacking accuracy, or are invasive and often destructive methods that are inapplicable for experiments involving living corals. This study introduces a novel precise and non-destructive technique to quantify surface area in living coral colonies by applying computed tomography (CT) and subsequent 3D reconstruction. Living coral colonies of different taxa were scanned by conventional medical CT either in air or in sea water. Resulting data volumes were processed by 3D modeling software providing realistic 3D coral skeleton surface reconstructions, thus enabling surface area measurements. Comparisons of CT datasets obtained from calibration bodies and coral colonies proved the accuracy of the surface area determination. Surface area quantifications derived from two different surface rendering techniques applied for scanning living coral colonies showed congruent results (mean deviation ranging from 1.32 to 2.03%). The validity of surface area measurement was verified by repeated measurements of the same coral colonies by three test persons. No significant differences between all test persons in all coral genera and in both surface rendering techniques were found (independent sample t-test: all n.s.). Data analysis of a single coral colony required approximately 15 to 30 min for a trained user using the isosurface technique regardless of the complexity and growth form of the latter, rendering the method presented in this study as a time-saving and accurate method to quantify surface areas in both living coral colonies and bare coral skeletons. Communicated by Biology Editor Dr Michael Lesser     

The ecology of 'Acroporid white syndrome', a coral disease from the southern Great Barrier Reef

Outbreaks of coral disease have increased worldwide over the last few decades. Despite this, remarkably little is known about the ecology of disease in the Indo-Pacific Region. Here we report the spatiotemporal dynamics of a coral disease termed 'Acroporid white syndrome' observed to affect tabular corals of the genus Acropora on the southern Great Barrier Reef. The syndrome is characterised by rapid tissue loss initiating in the basal margins of colonies, and manifests as a distinct lesion boundary between apparently healthy tissue and exposed white skeleton. Surveys of eight sites around Heron Reef in 2004 revealed a mean prevalence of 8.1±0.9%, affecting the three common species (Acropora cytherea, A. hyacinthus, A. clathrata) and nine other tabular Acropora spp. While all sizes of colonies were affected, white syndrome disproportionately affected larger colonies of tabular Acroporids (>80 cm). The prevalence of white syndrome was strongly related to the abundance of tabular Acroporids within transects, yet the incidence of the syndrome appears unaffected by proximity to other colonies, suggesting that while white syndrome is density dependant, it does not exhibit a strongly aggregated spatial pattern consistent with previous coral disease outbreaks. Acroporid white syndrome was not transmitted by either direct contact in the field or by mucus in aquaria experiments. Monitoring of affected colonies revealed highly variable rates of tissue loss ranging from 0 to 1146 cm(-2) week(-1), amongst the highest documented for a coral disease. Contrary to previous links between temperature and coral disease, rates of tissue loss in affected colonies increased threefold during the winter months. Given the lack of spatial pattern and non-infectious nature of Acroporid white syndrome, further studies are needed to determine causal factors and longer-term implications of disease outbreaks on the Great Barrier Reef.     

Correlations between chaetodontid fishes and coral communities of the Gulf of Aqaba (Red Sea)

Synopsis The relationships existing between the chaetodontid fishes and the surrounding coral communities were investigated in the Gulf of Aqaba. Quantitative data were analysed by a correspondence and a cluster analysis. The results demonstrated a similarity in the spatial distribution of both communities. Significant correlations were found between the density of chaetodontid fishes and the diversity of the coral community as well as the substratum coverage by the coral colonies. The density of exclusive coral browsers was also correlated to the abundance of branching colonies. Among the different genera of branching corals, correlations were significant only for the genusAcropora. These results suggested the existence of strong links between coral and chaetodontid fish assemblages.     

Development and validation of computational fluid dynamics models for prediction of heat transfer and thermal microenvironments of corals

We present Computational Fluid Dynamics (CFD) models of the coupled dynamics of water flow, heat transfer and irradiance in and around corals to predict temperatures experienced by corals. These models were validated against controlled laboratory experiments, under constant and transient irradiance, for hemispherical and branching corals. Our CFD models agree very well with experimental studies. A linear relationship between irradiance and coral surface warming was evident in both the simulation and experimental result agreeing with heat transfer theory. However, CFD models for the steady state simulation produced a better fit to the linear relationship than the experimental data, likely due to experimental error in the empirical measurements. The consistency of our modelling results with experimental observations demonstrates the applicability of CFD simulations, such as the models developed here, to coral bleaching studies. A study of the influence of coral skeletal porosity and skeletal bulk density on surface warming was also undertaken, demonstrating boundary layer behaviour, and interstitial flow magnitude and temperature profiles in coral cross sections. Our models compliment recent studies showing systematic changes in these parameters in some coral colonies and have utility in the prediction of coral bleaching.     

Determination of the physical parameters of coral distributions using line transect data

We present a new method for extracting a comprehensive suite of biologically significant parameters from line transect data of coral communities. In addition to the percentage coral cover (the traditionally extracted parameter), the method extracts the population density of the coral colonies, their mean diameter and associated standard deviation and, for adequate data, their size frequency distribution. The method assumes only that the coral colonies form a system of non-overlapping circles in the plane, that the diameters of the circles are random quantities with an unknown distribution function, and that the transects are placed randomly. We test the method on both theoretical and real data to show that it performs as well as, if not better than, current methods in extracting the traditional parameter as well as being able to extract the additional useful parameters indicated. Because the method makes few restrictive assumptions and seems robust when used with field data, we suggest that it has wide application wherever line transects are used for ecological survey. The method is implemented in a Fortran program available from the senior author.AIMS publication number 222     

Effect of colony size and surrounding substrate on corals experiencing a mild bleaching event on Heron Island reef flat (southern Great Barrier Reef,Australia)

In January–May 2006, Heron Island in the Great Barrier Reef experienced a mild bleaching event. The effect of colony size, morphology and surrounding substrate on the extent of bleaching was explored. In contrast with previous studies, colony size did not influence bleaching sensitivity, suggesting that there may be a threshold of light and temperature stress beyond which size plays a role. Also contrasting with previous studies, massive corals were more affected by bleaching than branching corals. Massive corals surrounded by sand were more affected than the ones surrounded by rubble or dead coral. It is hypothesized that light reflectance from sand increases stress levels experienced by the colonies. This effect is maximized in massive corals as opposed to branching corals that form dense thickets on Heron Island. These results emphasize the importance of the ecological dynamics of coral communities experiencing low, moderate and high levels of bleaching for the understanding of how coral communities may change under the stress of climate change.     

Densitometry from digitized images of X-radiographs: Methodology for measurement of coral skeletal density

Skeletons of massive coral colonies contain annual density bands that are revealed by X-radiography of slices cut along growth axes. These bands allow measurement of skeletal growth parameters such as annual extension rate and annual calcification rate. Such measurements have been important in understanding coral growth, in assessing environmental impacts and in recovering proxy environmental information. Measurements of coral calcification rate from annual density banding require measurements of skeletal density along tracks across skeletal slices and, until now, such density measurements have depended upon specialized and expensive equipment. Here, we describe a straightforward, inexpensive and accurate technique for measuring skeletal density from digitized images of X-radiographs of coral skeletal slices. An aragonitic step-wedge was included in each X-radiograph of a coral slice together with two aluminium bars positioned along the anode-cathode axis. Optical density was measured along tracks across the X-ray images of these different objects. The aragonite step-wedge provided a standard for converting optical density to skeletal density. The aluminium bars were used to correct for the heel effect—a variation in the intensity of the X-ray beam along the anode-cathode axis that would, otherwise, introduce large errors into measurements of skeletal density. Exposure was found to vary from X-radiographs to X-radiograph, necessitating the inclusion of the calibration standards in each X-radiograph of a coral slice. Results obtained using this technique compared well with results obtained by direct gamma densitometry of skeletal slices.     

Coral community structure and risk assessment of high-latitude reefs at Sodwana Bay,South Africa

The importance of studying coral communities at different spatial scales is acknowledged in a growing volume of scientific literature, and principles of landscape ecology were thus used to elucidate the patterns in coral community structure on the high-latitude reefs in South Africa. These reefs are at the southernmost distribution of this fauna in Africa, are surprisingly species rich, and represent a biodiversity peak in this fauna south of the equator, regardless of the marginal nature of the environment. Coral community patterns were identified on and between the reefs at Sodwana Bay, justifying the grouping of reef areas in distinct zones. A number of landscape components were identified, ranging from the entire reef complex (10 km scale), individual reefs (1 km scales) and reef zones, to components that were separated using multivariate statistical analysis of transect data. These components transcended spatial similarities, e.g. the fore-reef on Five-mile Reef was not similar to the fore-reef on Seven-mile Reef, but was rather grouped with the reef flat on Two-mile Reef. This information was “translated” into an index of management intervention, based on risk assessment, and was generated using parameters that measure susceptibility to crown-of-thorns feeding, bleaching, diver-related damage and swell-induced breakage. We also assessed was the time elapsed since the last major disturbance and the proximity to the only boat launch site, a proxy measure of continuous disturbance. The risk assessment suggested that conservation management is most needed in the stable and “climax” coral communities that are usually characterised by a near-equal mix of hard and soft corals at maximal coral species diversity.     

A comparison between coral colonies of the genus Madracis and simulated forms

In addition to experimental studies, computational models provide valuable information about colony development in scleractinian corals. Using our simulation model, we show how environmental factors such as nutrient distribution and light availability affect growth patterns of coral colonies. To compare the simulated coral growth forms with those of real coral colonies, we quantitatively compared our modelling results with coral colonies of the morphologically variable Caribbean coral genus Madracis. Madracis species encompass a relatively large morphological variation in colony morphology and hence represent a suitable genus to compare, for the first time, simulated and real coral growth forms in three dimensions using a quantitative approach. This quantitative analysis of three-dimensional growth forms is based on a number of morphometric parameters (such as branch thickness, branch spacing, etc.). Our results show that simulated coral morphologies share several morphological features with real coral colonies (M. mirabilis, M. decactis, M. formosa and M. carmabi). A significant correlation was found between branch thickness and branch spacing for both real and simulated growth forms. Our present model is able to partly capture the morphological variation in closely related and morphologically variable coral species of the genus Madracis.     

Monitoring the coral disease, plague type II, on coral reefs in St. John, U.S. Virgin Islands

In July 1997, conspicuous white patches of necrotic tissue and bare skeleton began to appear on scleractinian corals in several bays around St. John, US Virgin Islands. Analysis of diseased coral tissue from five different species confirmed the presence of a Sphingomonas-like bacterium, the pathogen for plague type II. To date, 14 species of hard corals have been affected by plague type II around St. John. This disease was monitored at Haulover and Tektite Reefs at depths of 7-12 meters. The study site at Tektite Reef has > 50% cover by scleractinian corals with 90% of hard corals being composed of Montastraea annularis. Monthly surveys at Tektite Reef from December 1997 to May 2001 documented new incidence of disease (bare white patches of skeleton) every month with associated loss of living coral and 90.5% of all disease patches occurred on M. annularis. The frequency of disease within transects ranged from 3 to 58%, and the area of disease patches ranged from 0.25 to 9000 cm2. The average percent cover by the disease within 1 m2 ranged from 0.01% (+/- 0.04 SD) to 1.74% (+/- 9.08 SD). Photo-monitoring of 28 diseased corals of 9 species begun in September 1997 at Haulover Reef revealed no recovery of diseased portions with all necrotic tissue being overgrown rapidly by turf algae, usually within less than one month. Most coral colonies suffered partial mortality. Very limited recruitment (e.g., of Agaricia spp., Favia spp. and sponges) has been noted on the diseased areas. This coral disease has the potential to cause more loss of live coral on St. John reefs than any other stress to date because it targets the dominant reef building species, M. annularis.     

Coral growth on three reefs: development of recovery benchmarks using a space for time approach

This 14-year study (1989–2003) develops recovery benchmarks based on a period of very strong coral recovery in Acropora-dominated assemblages on the Great Barrier Reef (GBR) following major setbacks from the predatory sea-star Acanthaster planci in the early 1980s. A space for time approach was used in developing the benchmarks, made possible by the choice of three study reefs (Green Island, Feather Reef and Rib Reef), spread along 3 degrees of latitude (300 km) of the GBR. The sea-star outbreaks progressed north to south, causing death of corals that reached maximum levels in the years 1980 (Green), 1982 (Feather) and 1984 (Rib). The reefs were initially surveyed in 1989, 1990, 1993 and 1994, which represent recovery years 5–14 in the space for time protocol. Benchmark trajectories for coral abundance, colony sizes, coral cover and diversity were plotted against nominal recovery time (years 5–14) and defined as non-linear functions. A single survey of the same three reefs was conducted in 2003, when the reefs were nominally 1, 3 and 5 years into a second recovery period, following further Acanthaster impacts and coincident coral bleaching events around the turn of the century. The 2003 coral cover was marginally above the benchmark trajectory, but colony density (colonies.m⁻²) was an order of magnitude lower than the benchmark, and size structure was biased toward larger colonies that survived the turn of the century disturbances. The under-representation of small size classes in 2003 suggests that mass recruitment of corals had been suppressed, reflecting low regional coral abundance and depression of coral fecundity by recent bleaching events. The marginally higher cover and large colonies of 2003 were thus indicative of a depleted and aging assemblage not yet rejuvenated by a strong cohort of recruits.     

The development of sampling strategies for population studies of coral reef fishes. A case study

Fish ecologists need to do pilot studies to develop accurate, precise and efficient sampling strategies. This paper presents a case study of the pilot investigations carried out to achieve this for three species of butterflyfish Chaetodon rainfordi, C. plebeius and Chelmon rostratus, at One Tree Reef. The effects of different transect sizes and methods on density estimates and their precision for each species were assessed. Varying transect dimensions (25, 50, 75 and 100 m lengthx1,2 and 3 m width) did not significantly affect the density estimates. However for C. rainfordi the precision of estimates was variable with the smallest transect length and width producing the least precise results. Higher density estimates were obtained for juvenile fish over a 1 m strip width than a 2 m width. A significant effect of disturbance on the densities of C. rainfordi and Chelmon rostratus was caused by the laying out of the transect tape. Consequently a technique of simultaneously censusing and delimiting transect boundaries was tested and found to give consistently higher density estimates. The potential effect of censusing at different times of the tide was examined and found to be insignificant for two species but quite complex for the other. Finally, a pilot sampling program was done at seven localities across One Tree Reef to determine the most efficient way of allocating sampling effort, for future census work.     

Latitudinal variation in coral communities in eastern Australia: a qualitative biophysical model of factors regulating coral reefs

The processes underlying the distributional limits of both corals and coral reefs can be elucidated by examining coral communities at high latitudes. Coral-dominated communities in eastern Australia cover a latitudinal range of >2,500 km, from the northern Great Barrier Reef (11°S) to South West Rocks (31.5°S). Patterns of coral species richness from 11 locations showed a clear separation between the Great Barrier Reef and subtropical sites, with a further abrupt change at around 31°S. Differences in community structure between the Great Barrier Reef and more southern sites were mainly attributable to higher cover of massive corals, branching Acropora, dead coral and coralline algae on the Great Barrier Reef, and higher cover of macroalgae and bare rock at more southern sites. The absence of some major reef-building taxa (i.e., staghorn Acropora and massive Porites) from most subtropical sites coincided with the loss of reef accretion capacity. Despite high cover of hard corals in communities at up to 31°S, only Lord Howe Island contained areas of reef accretion south of the Great Barrier Reef. Factors that have been hypothesized to account for latitudinal changes in coral community structure include water temperature, aragonite saturation, light availability, currents and larval dispersal, competition between corals and other biota including macroalgae, reduced coral growth rates, and failure of coral reproduction or recruitment. These factors do not operate independently of each other, and they interact in complex ways.     

Variation in the structure of epifaunal invertebrate assemblages among coral hosts

The high biodiversity of coral reefs is attributable to the many invertebrate groups which live in symbiotic relationships with other reef organisms, particularly those which associate with the living coral habitat. However, few studies have examined the diversity and community structure of coral-dwelling invertebrates and how they vary among coral species. This study quantified the species richness and composition of animals associated with four common species of branching corals (Acropora nasuta, A. millepora, Pocillopora damicornis, and Seriatopora hystrix) at Lizard Island in the northern Great Barrier Reef. One hundred and seventy-eight nominal species from 12 different phyla were extracted across 50 replicate colonies of each coral host. A single coral colony, approximately 20 cm in diameter, harbored as many as 73 individuals and 24 species. There were substantial differences in invertebrate species composition among coral hosts of different families as well as genera. Twenty-seven species (15% of all taxa collected) were found on only one of the four different coral species, which may potentially indicate some level of specialization among coral hosts. The distinct assemblages on different coral species, and the presence of potential specialists, suggests invertebrate communities will be sensitive to the differential loss of branching coral species resulting from coral reef degradation.     

Micro‐CT reconstruction reveals the colony pattern regulations of four dominant reef‐building corals

Colonies are the basic geometric building blocks of coral reefs. However, the forming regulations of both colonies and reefs are still not understood adequately. Therefore, in this study, we reconstructed 25 samples using high‐resolution micro‐computed tomography to investigate coral growth patterns and parameters. Our skeleton and canal reconstructions revealed the characteristics of different coral species, and we further visualized the growth axes and growth rings to understand the coral growth directions. We drew a skeleton grayscale map and calculated the coral skeleton void ratios to ascertain the skeletal diversity, devising a method to quantify coral growth. On the basis of the three‐dimensional (3D) reconstructions and growth parameters, we investigated the growth strategies of different coral species. This research increases the breadth of knowledge on how reef‐building corals grow their colonies, providing information on reef‐forming regulations. The data in this paper contain a large amount of coral growth information, which can be used in further research on reef‐forming patterns under different conditions. The method used in this study can also be applied to animals with porous skeletons.     

Spatial patterns in abundance of a damselfish reflect availability of suitable habitat

For species with metapopulation structures, variation in abundance among patches can arise from variation in the input rate of colonists. For reef fishes, variability in larval supply frequently is invoked as a major determinant of spatial patterns. We examined the extent to which spatial variation in the amount of suitable habitat predicted variation in the abundance of the damselfish Dascyllus aruanus, an abundant planktivore that occupies live, branched coral throughout the Indo-Pacific. Reef surveys established that size, branching structure and location (proximity to sand) of the coral colonies together determined the ”suitability” of microhabitats for different ontogenetic stages of D. aruanus. Once these criteria were known, patterns of habitat use were quantified within lagoons of five Pacific islands. Availability of suitable habitat generally was an excellent predictor of density, and patterns were qualitatively consistent at several spatial scales, including among different lagoons on the same island, among different islands and between the central (French Polynesia and Rarotonga) and western (Great Barrier Reef, Australia) South Pacific. A field experiment that varied the amount of suitable coral among local plots indicated that habitat for settlers accounted for almost all of the spatial variation in the number of D. aruanus that settled at that location, suggesting that spatial patterns of abundance can be established at settlement without spatial variation in larval supply. Surveys of four other species of reef-associated fish revealed that a substantial fraction of their spatial variation in density also was explained by availability of suitable reef habitat, suggesting that habitat may be a prevalent determinant of spatial patterns. The results underscore the critical need to identify accurately the resource requirements of different species and life stages when evaluating causes of spatial variation in abundance of reef fishes. Received: 18 May 1999 / Accepted: 9 January 1999     

Distribution and Abundance of Fauna on Living Tissues of Two Brazilian Hermatypic Corals (Mussismilia hispida (Verril 1902) and Siderastrea stellata Verril, 1868)

This study describes the distribution and abundance patterns of the associate fauna on the living surface of the corals Siderastrea stellata Verril, 1868 and Mussismilia hispida (Verril 1902) using a non-destructive method, on the northern coast of Rio de Janeiro State. For each coral species, infestation density and proportions of infested colonies, colonies attached and unattached to the substrate were estimated. A total of 474 colonies of S. stellata and 452 colonies of M. hispida were examined. The barnacle Ceratoconcha floridana (Pilsbry, 1931) was the dominant coral associate found, followed by gall-crabs of the family Cryptochiridae Paulson, 1875 and the bivalve Lithophaga bisulcata (d’Orbigny, 1842). Both coral species presented similar patterns of infestation dominance. S. stellata colonies were more commonly infested and showed a greater mean infestation density of 0.62 ind/cm² at Armação dos Búzios, whereas M. hispida colonies had infestation densities of only 0.20 ind/cm². Infestation density does not appear to impact negatively on corals of Armação dos Búzios. A clear negative relationship between the number of associates in the coral colony and coral size was found. Evidently abundance and frequency of occurrence of associated fauna is highly related to coral community structure and composition and the results highlight the importance of local scale studies.     

Geographic and habitat partitioning of genetically distinct zooxanthellae (Symbiodinium) in Acropora corals on the Great Barrier Reef

Intra- and intercolony diversity and distribution of zooxanthellae in acroporid corals is largely uncharted. In this study, two molecular methods were applied to determine the distribution of zooxanthellae in the branching corals Acropora tenuis and A. valida at several reef locations in the central section of the Great Barrier Reef. Sun-exposed and shaded parts of all colonies were examined. Single-stranded conformational polymorphism analysis showed that individual colonies of A. tenuis at two locations harbour two strains of Symbiodinium belonging to clade C (C1 and C2), whereas conspecific colonies at two other reefs harboured a single zooxanthella strain. A. valida was found to simultaneously harbour strains belonging to two distinct phylogenetic clades (C and D) at all locations sampled. A novel method with improved sensitivity (quantitative polymerase chain reaction using Taqman fluorogenic probes) was used to map the relative abundance distribution of the two zooxanthella clades. At two of the five sampling locations both coral species were collected. At these two locations, composition of the zooxanthella communities showed the same pattern in both coral species, i.e. correlation with ambient light in Pioneer Bay and an absence thereof in Nelly Bay. The results show that the distribution of genetically distinct zooxanthellae is correlated with light regime and possibly temperature in some (but not all) colonies of A. tenuis and A. valida and at some reef locations, which we interpret as acclimation to local environmental conditions.