The distribution of molluscan assemblages and their postmortem fate on coral reefs in the Gulf of Aqaba (northern Red Sea)

Molluscan assemblages were studied on fringing reefs (reef flats, Millepora-fringing reefs, fringing reefs with massive corals) and fore-reef hard substrata (coral patches, coral carpets and small patch reefs) in the Gulf of Aqaba at water depths ranging from the intertidal to 26 m. A total of 1,665 molluscan individuals from 51 taxa was counted on 44 transects, which covered 220 m2 at eight diving sites. The most important molluscs in the assemblage were the parasitic gastropod Coralliophila neritoidea, the encrusting gastropod Dendropoma maxima and the coral-associated bivalve Pedum spondyloideum. The dead assemblage, in contrast, was dominated by encrusting bivalves (Ostreoidea, Chamoidea, Spondylidae) and the coral-predating gastropod Drupella cornus. Distinct molluscan assemblages inhabit each of the three fringing reef-habitats and most of the important depth-related community changes occurred within the uppermost 5 m. In contrast, the three deeper fore-reef habitats are characterized by a more uniform molluscan composition. Molluscan assemblages were more dependent on substrata and their coral associations than on water depth. Comparisons with other published studies indicate that reefoidal hard substrata in the northern Red Sea are largely characterized by similar species-abundance patterns. The minor differences to other Red Sea studies probably reflect the northern, isolated position of the Gulf of Aqaba, the lack of certain molluscan habitats, and the differential impact of anthropogenic influences. Strong differences between living and dead assemblages in Aqaba are similar to those observed in other regions and are due to distinct biases in the dead assemblage. Molluscs closely associated with living corals (mostly bivalves and Dendropoma) can easily be overgrown after death and are thus undetectable in visual censuses. Some gastropod taxa are preferentially transported into surrounding soft-substrata postmortem or redistributed by hermit crabs. Such complex relationships between ecology and taphonomy are crucial in evaluating the quality of the molluscan fossil record in coral reef environments. The comparison of our results with literature data documents an increase in coral predators during the last two decades in the northern Red Sea. Due to the greater mollusc biodiversity in the shallower Aqaba reef habitats, damage to this coral reef zone would have the greatest impact on the overall mollusc community.     

A note on the breeding season of three ascidians on coral reefs at Galeta in the Caribbean Sea

Three Caribbean ascidians were sampled from April, 1972 until June, 1973 and the breeding season determined by the presence of larvae in the colonies. In Eudistoma sp. A, larvae occurred from early June until mid-November, in Trididemnum solidum from mid-June until the end of December, and in Didemnum sp. C, throughout the year with occasional interruptions.     

The trophic status of herbivorous fishes on coral reefs

Estimates of feeding rates, alimentary tract structure and temporal patterns of food processing obtained from twelve species of nominally herbivorous fishes on the northern Great Barrier Reef were compared. These included members of the families Acanthuridae, Scaridae and Kyphosidae. Based on an analysis of diet and short-chain fatty acid (SCFA) profiles from a previous study we initially partitioned the twelve species into four dietary categories, as follows: (a) Category 1: herbivores with a diet of macroscopic brown algae and high SCFA profiles in the hindgut region (Naso unicornis, Kyphosus vaigiensis); (b) Category 2: herbivores feeding on turfing and filamentous red and green algae with moderate SCFA profiles in the hindgut region (N. tonganus, K. cinerascens, Zebrasoma scopas, Acanthurus lineatus); (c) Category 3: zooplankton feeders with moderate SCFA profiles (N. vlamingii, N. brevirostris); (d) Category 4: species feeding on detrital and sedimentary materials with low levels of SCFA (Chlorurus microrhinos, Scarus schlegeli, Ctenochaetus striatus, A. olivaceus). The purpose of this comparison was to determine whether measures of feeding activity, alimentary tract structure, and food processing were concordant with diet. A dichotomy in feeding rates was observed. Species with a diet of algae and zooplankton (categories 1–3) had slower feeding rates than those feeding on detrital aggregates and sediment (category 4). The pattern of food processing also followed the same dichotomy with species of categories 1–3 retaining food in the alimentary tract overnight and commencing the feeding day with substantial amounts of food in the intestine and hindgut. Category-4 species commenced the feeding day with empty alimentary tracts suggesting a rapid turnover of gut contents. Within the herbivorous and zooplankton-feeding species neither alimentary tract structure nor food processing mode were predicted by diet or SCFA profiles. A hindgut fermentation chamber was present in K. vaigiensis but not in N. unicornis, a species with high levels of SCFA in the hindgut region and a diet of brown macroscopic algae. In contrast N. vlamingii, with a diet dominated by animal matter, retained large amounts of food material in a hindgut chamber over the entire feeding cycle. In tropical perciform fishes, herbivory and fermentation are not associated with the alimentary tract structures that characterise herbivorous terrestrial vertebrates. Estimates of the abundance of the different groupings of nominally herbivorous fishes indicated that the dominant elements in the reef grazing and browsing fauna were consumers of detrital and sedimentary materials. These could not be classified as herbivores. Members of this group were dominant in all habitats investigated. Explicitly herbivorous taxa were a minority component in all habitats investigated.Communicated by G.F. Humphrey, Sydney     

The trophic status of herbivorous fishes on coral reefs

Dietary analyses of 17 species of nominally herbivorous fishes on the northern Great Barrier Reef were carried out to resolve the questions to what extent diet in nominally herbivorous reef fishes is dominated by living plant material and whether short-chain fatty acid profiles in the alimentary tract predict diet. The fishes included members of the families Acanthuridae, Scaridae and Kyphosidae. The analyses revealed consumption of a wide range of food resources including macroscopic algae, turfing algae, pelagic and planktonic animal matter, organic detritus and sediments. In only 7 of the 17 species was living algae the dominant dietary item. Gastrointestinal short-chain fatty acids (SCFA) profiles reflecting the fermentative activities of gut symbionts were used as a framework for the initial analysis of dietary patterns. The feeding patterns implied by the SCFA profiles were validated by direct observation of food material through gut content analysis. We identified four main trophic groupings differentiated by fermentation profiles and diet. These were: (1) species with low levels of SCFA in gut samples (mean mM 9.2ǂ.8) but a high proportion of isovalerate (mean 15.4%dž.7): Acanthurus nigricauda, A. olivaceus, Ctenochaetus striatus, Chlorurus microrhinos, C.sordidus, Scarus schlegeli; (2) species that fed on turfing, filamentous and small thallate algae and displayed moderate levels of SCFA (mean mM 21.9DŽ.9): Acanthurus nigricans, A.lineatus, Naso tuberosus; (3) species that fed on large thallate (macroscopic) algae with high levels of SCFA (mean mM 41.2Dž.5) with very low proportions of isovalerate (mean 0.5%ǂ.1): Kyphosus cinerascens, K.vaigiensis, Naso unicornis, Zebrasoma scopas; and (4) species with diets dominated by planktonic animal material with moderate levels of SCFA (mean mM 21.3ǃ.5): Naso annulatus, N. brevirostris, N. hexacanthus, N. vlamingii. We conclude that a wide range of dietary items are taken by the 17 nominal herbivores, and that dietary groupings do not reflect taxonomic relationships. There was evidence of convergence in feeding modes and diet between phylogenetically distinct taxa and divergence within particular lineages.     

Parrotfish predation on cryptic sponges of Caribbean coral reefs

Some sponge species that live in crevices in the reef frame appear to be restricted to their cryptic habitat by predation. When cryptic sponges were excavated, on Guigalotupo reef, San Blas, Panama, exposing them to potential predators, they were eaten by fishes that are generally considered to be herbivores, primarily parrotfishes of the genus Sparisoma: S.aurofrenatum (Cuvier & Valenciennes), S.viride (Bonnaterre), and S.chrysopterum (Bloch & Schneider). Of the 9150 bites observed to be taken by these species during paired (i.e., with sponges versus without sponges) trials conducted in defined feeding areas during 1986, 1987, and 1988, 72% (i.e. 6581 bites) were on cryptic sponges, even though these were only offered during half of the total observation time and never constituted more than 7% of the cover of the feeding observation areas. Individual parrotfish returned over and over to take bites of the exposed cryptic sponges until they were entirely consumed. They vigorously chased each other away from the sponges, but exhibited no such defense of their usual algal foods. A total of 18 sponge species were tested. Of the cryptic and semi-cryptic sponge species tested, only one of six was rejected by the parrotfish. Two of these six sponge species were consistently consumed entirely, and two were consumed entirely whenever their surfaces were sliced off with a razor blade, demonstrating that these sponges concentrate defenses against predators in their surfaces. One semi-cryptic species and one semi-exposed species were fed upon, but not entirely consumed. By contrast, 11 of 12 of the exposed and semi-exposed species were rejected. Cryptic sponges grew out of their cavities in the reef only when protected by seasonally thick mats of macroalgae or by cages that excluded fish. Received: 10 January 1997 / Accepted: 4 February 1997     

Diversity and stability of herbivorous fishes on coral reefs

Biodiversity may provide insurance against ecosystem collapse by stabilizing assemblages that perform particular ecological functions (the "portfolio effect"). However, the extent to which this occurs in nature and the importance of different mechanisms that generate portfolio effects remain controversial. On coral reefs, herbivory helps maintain coral dominated states, so volatility in levels of herbivory has important implications for reef ecosystems. Here, we used an extensive time series of abundances on 35 reefs of the Great Barrier Reef of Australia to quantify the strength of the portfolio effect for herbivorous fishes. Then, we disentangled the contributions of two mechanisms that underlie it (compensatory interactions and differential responses to environmental fluctuations ["response diversity"]) by fitting a community-dynamic model that explicitly includes terms for both mechanisms. We found that portfolio effects operate strongly in herbivorous fishes, as shown by nearly independent fluctuations in abundances over time. Moreover, we found strong evidence for high response diversity, with nearly independent responses to environmental fluctuations. In contrast, we found little evidence that the portfolio effect in this system was enhanced by compensatory ecological interactions. Our results show that portfolio effects are driven principally by response diversity for herbivorous fishes on coral reefs. We conclude that portfolio effects can be very strong in nature and that, for coral reefs in particular, response diversity may help maintain herbivory above the threshold levels that trigger regime shifts.     

Modeling the beta diversity of coral reefs

Quantifying the beta diversity (species replacement along spatiotemporal gradients) of ecosystems is important for understanding and conserving patterns of biodiversity. However, virtually all studies of beta diversity focus on one-dimensional transects orientated along a specific environmental gradient that is defined a priori. By ignoring a second spatial dimension and the associated changes in species composition and environmental gradients, this approach may provide limited insight into the full pattern of beta diversity. Here, we use remotely sensed imagery to quantify beta diversity continuously, in two dimensions, and at multiple scales across an entire tropical marine seascape. We then show that beta diversity can be modeled (0.852 > or = r2 > or = 0.590) at spatial scales between 0.5 and 5.0 km2, using the environmental variables of mean and variance of depth and wave exposure. Beta diversity, quantified within a "window" of a given size, is positively correlated to the range of environmental conditions within that window. For example, beta diversity increases with increasing variance of depth. By analyzing such relationships across seascapes, this study provides a framework for a range of disparate coral reef literature including studies of zonation, diversity, and disturbance. Using supporting evidence from soft-bottom communities, we hypothesize that depth will be an important variable for modeling beta diversity in a range of marine systems. We discuss the implications of our results for the design of marine reserves.     

Biomass of suspended bacteria over coral reefs

The biomass of bacteria suspended in water flowing over coral reefs at Lizard Island and Yonge Reef (Northern Great Barrier Reef) was estimated by measurement of muramic acid. Values ranged from 20 mg C m^-3 in the open water up to about 60 mg C m^-3 over the reef flat. Direct counts of total numbers of free bacteria were made for comparison. Values of around 2.0x10⁹ cells g^-1 muramic acid showed that there was a good agreement between direct counts and muramic acid content of free bacteria in the open water. In samples containing suspended particulate matter, ratios of direct counts to muramic acid concentration were lower, because bacteria on particles could not be counted. Thus, these ratios were used to indicate the proportions of bacteria attached to particles. Changes in the biomass and numbers of bacteria were determined in water masses identified either by a drogue or fluorescein, as they moved across the reefs. In the zone on the outside of the reef, the number of free bacteria decreased compared to open sea water, but total biomass increased, showing that particulate matter containing bacteria was thrown up into suspension. About 50% of bacteria were attached to particles. Water flowing over the reef flats contained much particulate material with bacteria attached. Bacteria constituted between about 5 and 20% of particulate organic carbon.     

Development of contemporary Eastern Pacific coral reefs

An overview of oceanographic conditions prevailing in the tropical Eastern Pacific Ocean suggests that the entire region is environmentally marginal for coral-reef development. The principal features of this environment are a strong, permanent, shallow thermocline and an annual north-south migration of the Intertropical Convergence resulting in wet and dry seasons. Along tropical Eastern Pacific continental margins structural coral reefs are best developed in the Gulf of Chiriquí off western Panamá. These reefs are relatively small, with reef formation taking place at a maximum depth of roughly 10 m. All of the reefs are judged to have formed since sea level approached its present height some 5,000 years ago. A study of the physical environment in the Gulf of Chiriquí revealed the following. Seasonal differences in surface temperatures were small but significant (P0.01), with the dry-season median (Md) of 28.9°C higher than the rainy season one of 28.0°C. At all times, surface-water temperatures were within the range considered optimal for coral growth. There were also significant (P<0.05) seasonal differences in the depths of the 25°, 20°, and 18°C isotherms. The first was shallowest (Md=18.5 m) during the rainy season due to vertical mixing, while the latter two were shallowest (Mds=31.5 and 33.0 m, respectively) during the dry season due to a generalized shoaling of the thermocline. All three isotherms are closely associated with the thermocline and showed remarkable variability in depth, most likely connected with internal waves. Salinities were reduced down to depths greater than 20 m and for distances of more than 50 km from the coast. Seasonal differences were slight (0.7% S) but statistically significant (P<0.01). Turbidity during the rainy season reduces the amount of light reaching the bottom at 10 m depth roughly by a factor of three compared to the dry season. Even the dry season amount is only about one half as much as would be expected to reach the same depth on the seaward reef of a Westen Pacific atoll. These conditions of cool water a short distance below the surface, reduced salinities, and high seasonal turbidity combine to make the region a poor one for coral-reef formation. The history of the Eastern Pacific coral fauna is traced from the Cretaceous to the Holocene. The present fauna is of Indo-Western Pacific origin, having become established following (1) the final closure of the connections between the Caribbean and Eastern Pacific (Pliocene), (2) movement of the northern Line Islands by sea-floor spreading into the path of the North Pacific Equatorial Countercurrent (Pliocene), and (3) the loss of all Eastern Pacific hermatypes during the Pleistocene.     

Herbivory by the Caribbean king crab on coral patch reefs

Caribbean coral reefs are increasingly dominated by macroalgae instead of corals due to several factors, including the decline of herbivores. Yet, virtually unknown is the role of crustacean macrograzers on coral reef macroalgae. We examined the effect of grazing by the Caribbean king crab (Mithrax spinosissimus) on coral patch reef algal communities in the Florida Keys, Florida (USA), by: (1) measuring crab selectivity and consumption of macroalgae, (2) estimating crab density, and (3) comparing the effect of crab herbivory to that of fishes. Mithrax prefers fleshy macroalgae, but it also consumes relatively unpalatable calcareous algae. Per capita grazing rates by Mithrax exceed those of most herbivorous fish, but Mithrax often occurs at low densities on reefs and its foraging activities are reduced in predator-rich environments. Therefore, the effects of grazing by Mithrax tend to be localized and when at low density contribute primarily to spatial heterogeneity in coral reef macroalgal communities.     

Catastrophe and the life span of coral reefs

A strong earthquake in the western Caribbean in 2009 had a catastrophic impact on uncemented, unconsolidated coral reefs in the central sector of the shelf lagoon of the Belizean barrier reef. In a set of 21 reef sites that had been observed prior to the earthquake, the benthic assemblages of 10 were eradicated, and one was partially damaged, by avalanching of their slopes. Ecological dynamics that had played out over the previous 23 years, including the mass mortalities of two sequentially dominant coral species and a large increase in the cover of an encrusting sponge, were instantaneously rendered moot in the areas of catastrophic reef-slope failure. Because these prior dynamics also determined the benthic composition and resilience of adjacent sections of reef that remained intact, the history of disturbance prior to the earthquake will strongly influence decadal-scale recovery in the failed areas. Geological analysis of the reef framework yielded a minimum return time of 2000-4000 years for this type of high-amplitude event. Anthropogenic degradation of ecosystems must be viewed against the backdrop of long-period, natural catastrophes, such as the impact of strong earthquakes on uncemented, lagoonal reefs.     

Malthusian overfishing and efforts to overcome it on Kenyan coral reefs

This study examined trends along a gradient of fishing intensity in an artisanal coral reef fishery over a 10-year period along 75 km of Kenya's most populated coastline. As predicted by Malthusian scenarios, catch per unit effort (CPUE), mean trophic level, the functional diversity of fished taxa, and the diversity of gear declined, while total annual catch and catch variability increased along the fishing pressure gradient. The fishery was able to sustain high (approximately 16 Mg x km(-2) x yr(-1)) but variable yields at high fishing pressure due to the dominance of a few productive herbivorous fish species in the catch. The effect of two separate management strategies to overcome this Malthusian pattern was investigated: fisheries area closure and elimination of the dominant and most "competitive" gear. We found that sites within 5 km of the enforced closure showed significantly lower total catch and CPUE, but increased yield stability and trophic level of catch than predicted by regression models normalized for fishing effort. Sites that had excluded illegal beach seine use through active gear management exhibited increased total catch and CPUE. There was a strong interaction between closure and gear management, which indicates that, for closures to be effective at increasing catch, there must be simultaneous efforts at gear management around the periphery of the closures. We propose that Malthusian effects are responsible for the variation in gear and catch and that active management through reduced effort and reductions in the most competitive gear have the greatest potential to increase the functional and trophic diversity and per-person productivity.     

Stable carbon isotope ratios in Red Sea barnacles (Cirripedia) as an indicator of their food source

The stable isotope ratio ¹²C/¹³C was used to investigate the source of carbon in free-living barnacles and in coral-inhabiting barnacles from the Red Sea. The δ¹³C of most of the barnacles collected on the open shore ranges between −17.5 and −19.7‰, indicating relative enrichment of light carbon originating from the open-sea phytoplankton. Those collected in closed habitats showed heavier isotopic composition. The δ¹³C of the coral-inhabiting barnacles ranges from −14.1 to −16.7‰, suggesting that the carbon contribution of open-sea plankton to these barnacles is less important than it is to free-living barnacles. We hypothesize that coral organic matter and zooxanthellae expelled by the host coral contribute carbon to the barnacle, and that a mixture of this relatively heavy carbon with carbon from other sources is responsible for the high values of δ¹³C in coral barnacles. Received: 28 February 1997 / Accepted: 16 September 1997     

Habitat biodiversity as a determinant of fish community structure on coral reefs

Increased habitat diversity is often predicted to promote the diversity of animal communities because a greater variety of habitats increases the opportunities for species to specialize on different resources and coexist. Although positive correlations between the diversities of habitat and associated animals are often observed, the underlying mechanisms are only now starting to emerge, and none have been tested specifically in the marine environment. Scleractinian corals constitute the primary habitat-forming organisms on coral reefs and, as such, play an important role in structuring associated reef fish communities. Using the same field experimental design in two geographic localities differing in regional fish species composition, we tested the effects of coral species richness and composition on the diversity, abundance, and structure of the local fish community. Richness of coral species overall had a positive effect on fish species richness but had no effect on total fish abundance or evenness. At both localities, certain individual coral species supported similar levels of fish diversity and abundance as the high coral richness treatments, suggesting that particular coral species are disproportionately important in promoting high local fish diversity. Furthermore, in both localities, different microhabitats (coral species) supported very different fish communities, indicating that most reef fish species distinguish habitat at the level of coral species. Fish communities colonizing treatments of higher coral species richness represented a combination of those inhabiting the constituent coral species. These findings suggest that mechanisms underlying habitat-animal interaction in the terrestrial environment also apply to marine systems and highlight the importance of coral diversity to local fish diversity. The loss of particular key coral species is likely to have a disproportionate impact on the biodiversity of associated fish communities.     

Light harvesting by wavelength transformation in a symbiotic coral of the Red Sea twilight zone

We report an extraordinary depth range for Leptoseris fragilis (Milne Edwards and Haime), a reef building coral of the Red Sea living in cytosymbiosis with zooxanthellae. The coral harbours an as yet unknown pigment system. We suggest that the heterotrophic host — the coral — provides its photoautotrophic symbionts with additional light. The supplementary light is provided by host pigments which transform light of short wavelengths into suitable wavelengths for photosynthesis, thus amplifying and increasing the transfer of photoassimilates from the zooxanthellae to the host.     

Sexual reproduction and settlement of the coral reef spongeChalinula sp. from the Red Sea

Characteristics of the sexual reproduction and larval settlement of the haplosclerid spongeChalinula sp., which inhabits the shallow waters (1 to 6 m) of Eilat, Red Sea, were investigated from September 1985 through to November 1987. This species was found to be a simultaneous hermaphroditic brooder, hence gonochorism is not the rule in the order Haplosclerida. Brooding always takes place in special brooding chambers. While the oocytes in the brooding chambers are among the largest known in sponges (355±37µm), the spermatic cysts distributed in the choanosome are among the smallest known for this phylum (average 26±7µm).Chalinula sp. breeds throughout the year and in experiments most larvae (74%) settled within 1 to 8 h post-release, generally within 4.5 h. Metamorphosis from larval shape to a sessile sponge lasts 1 to 6 h. Thus, larvae had a short swimming period, settled fast, and metamorphosed rapidly (within 1 to 6 h). The large size of the larvae may contribute to their ability to rapidly reorganize their body shape into that of a sessile sponge. In addition, the existence of already differentiated choanocyte chambers in the larvae, facilitates fast construction of the water filtration system in the newly settled sponges. The reproductive and larval characteristics ofChalinula sp. enable the larvae to settle on any vacant space in the reef, which may explain its abundance in the Red Sea.     

Estimating the footprint of pollution on coral reefs with models of species turnover

Ecological communities typically change along gradients of human impact, although it is difficult to estimate the footprint of impacts for diffuse threats such as pollution. We developed a joint model (i.e., one that includes multiple species and their interactions with each other and environmental covariates) of benthic habitats on lagoonal coral reefs and used it to infer change in benthic composition along a gradient of distance from logging operations. The model estimated both changes in abundances of benthic groups and their compositional turnover, a type of beta diversity. We used the model to predict the footprint of turbidity impacts from past and recent logging. Benthic communities far from logging were dominated by branching corals, whereas communities close to logging had higher cover of dead coral, massive corals, and soft sediment. Recent impacts were predicted to be small relative to the extensive impacts of past logging because recent logging has occurred far from lagoonal reefs. Our model can be used more generally to estimate the footprint of human impacts on ecosystems and evaluate the benefits of conservation actions for ecosystems.     

Appraisal of visual assessments of habitat complexity and benthic composition on coral reefs

Visual assessments of topographic habitat structure and benthos on coral reefs were appraised using quantitative data collected from 16 replicate surveys within each of 21 sites on Seychelles reefs. Results from visual assessments of reef benthos were similar to those obtained using techniques frequently used to assess benthic complexity and composition. Visual estimates of habitat topography were correlated with rugosity, reef height and holes of 10–70 cm diameter, whilst visual estimates of benthic composition were very similar to those obtained from line intercept transects. Visual estimates of topography correlated strongly with species richness of fish communities and explained 42% of the variation in these data. The relationship between visual estimates of topography and species richness is strongest with fish 10–30 cm total length (TL), abundance of fish within this size category also correlating positively with topographic visual assessments. Visual techniques are prone to observer bias, however with regular training they can be used to quickly provide a reliable and effective means of assessing habitat complexity and benthos on coral reefs.     

The basics of acidification: baseline variability of pH on Australian coral reefs

Ocean acidification is one of the key threats facing coral reef ecosystems, but there are few estimates of spatial and temporal variability in pH among reef habitats. The present study documents levels of spatial variability in pH among coral reef habitats (9 to 10), among locations separated by 100’s km of latitude and between east (Great Barrier Reef, GBR) and west (Ningaloo Reef) coasts of Australia. Differences were found in pH between inshore and offshore waters along Ningaloo Reef (means 8.45, 8.53, respectively). Replicate assessments here ranged from 8.22 to 8.64. On the GBR, the range of values over all habitats and replicates was 0.39 pH units (7.98 to 8.37). There were minor but significant differences of 0.05 pH units between 5 consecutive days for habitats on average. Highest pH was recorded in filamentous algal beds maintained by the damselfish Dischistodus perspicillatus. Lowest pH was found in water extracted from sand-dwelling goby holes. While there were marked changes in pH over a 48-h sampling period among 4 habitats at Lizard Island (GBR), there was little evidence of a diel trend. Understanding how pH varies at scales that are relevant to organisms that live on shallow coral reefs is crucial for the design and interpretation of experiments that test the effects on organisms of the changes in water chemistry predicted to affect oceans in the future.