Temporal and spatial variation in bacterial production in the water column over a coral reef

Production and doubling times of the bacterial populations in the water around and over the reefs at Lizard Island, Great Barrier Reef were measured during summer and winter, 1982 and 1983. Bacterial productivity, determined from the rate of tritiated thymidine incorporation into DNA, was high over the reef flats and a Thalassia hemprichii sand flat (28 to 58 g Cl^-1 d^-1). Bacterial growth rates increased during the day and fell at night over the reef flats and seagrass bed. Growth rates were slower over the reef front and in open water. Doubling times ranged from about 2 d in the open water to about 3 h over the reef flat in summer. As numbers did not increase, grazing was probably intense on the reef flats. Growth rates were much slower in winter. The main source of organic nutrient used by the bacteria was probably mucus released following photosynthesis in the corals. The cyanobacterium Synechococcus sp. was sometimes very numerous, especially in summer when 2×10⁸ cells l^-1 were recorded in one water mass. The number of bacteria was also very high in summer, with values ranging from 1×10⁹ to 2.5×10⁹l^-1.     

Zooplankton abundance and grazing at Davies Reef,Great Barrier Reef,Australia

Zooplankton abundance and grazing on autotrophic and heterotrophic particulate matter were measured along a transect across Davis Reef (18°5S; 147°39E) and in the back-reef lagoon over tidal and diel cycles during austral winter (August 1984). Zooplankton entering the reef from the surrounding shelf waters decreased in abundance over the reef flat, presumably because of predation. Within the reef lagoon, maximum daytime densities of pelagic copepods occurred during high water, suggesting an external input. At night, water-column zooplankton biomass increased by a factor of 2 to 3 due to the emergence of demersal reef zooplankton. Zooplankton grazing rates on heterotrophic particulate matter (bacteria + detritus and Protozoa) compared to phytoplankton were higher on the reef flat than on the fore-reef or lagoon. Within the lagoon, zooplankton grazing rates on heterotrophic material were maximum during high water, coincident with maximum tidal concentrations of particulate organic carbon. The combined demersal and pelagic zooplankton community were often able to crop 30% of the daily primary production by >2µm phytoplankton. However, >50% of phytoplankton biomass was in cells <2µm, presumably unavailable to these zooplankton. Our particulate production and ingestion measurements, together with zooplankton carbon demand extrapolated from respiration estimates, suggest that the zooplankton community of Davies Reef derives much of its nutrition from detritus.Joint contribution from the University of Maryland, Center for Environmental and Estuarine Studies (No. 2015), and the Microbial Ecology on a Coral Reef Workshop (MECOR No. 19)     

Rates of nitrogen fixation on coral reefs across the continental shelf of the central Great Barrier Reef

This study was undertaken in 1981 to determine whether there were major variations in potential rates of nitrogen fixation on apparently bare coralline substrate from reefs across the continental shelf of the central Great Barrier Reef. Nitrogen fixation, measured as rates of ethylene production (nmol cm^-2h^-1), was significantly lower on substrata from two inner-shelf reefs, (0.46 and 1.07) than on two middle-shelf reefs (2.10 and 2.97) and on two outer-shelf reefs (3.20 and 3.81). By contrast, algal biomass (mg cm^-2) on experimental substrate was significantly higher on inner-shelf reefs (80.8 and 59.4) than on middleshelf (27.1 and 23.8) and outer-shelf reefs (26.4 and 22.4). The rate of nitrogen fixation was positively correlated with the proportion of bare substratum and significantly higher concentrations of dissolved inorganic nitrogen were found in waters over the reefs than in water flowing onto those reefs. The abundance of algal-grazing fishes was reported previously to be significantly lower on inner-shelf reefs. It is suggested that this cross-shelf variation in the activity of algal-grazing fishes may be a determinant of the observed cross-shelf variations in potential nitrogen fixation.Contribution No. 233 from the Australian Institute of Marine Science     

Nitrogen excretion by some demersal macrozooplankton in Heron and One Tree Reefs,Great Barrier Reef,Australia

Nitrogen excretion rates of demersal macrozooplankton were measured together with nitrogen concentrations in the water column and sediments in lagoons of Heron Reef and One Tree Reef, Great Barrier Reef, Australia, during August and November 1991. Excretion rates increased with body weight, and weight-specific excretion rates of the demersal macrozooplankton were comparable to those of pelagic zooplankton and meiofauna in the Great Barrier Reef. Values of demersal macrozooplankton abundance from previous studies and excretion rates from this study were combined to estimate fluxes of ammonium from demersal macrozooplankton in coral reef lagoons. The estimated fluxes in the water column and sediments were 12 M NH₄ m^-2 d^-1 and 34 M NH₄ m^-2d^-1, respectively. These fluxes were compared with reported fluxes of ammonium in coral reef lagoons in the Great Barrier Reef, Australia. The estimated flux from the demersal macrozooplankton in the water column was 29 and 9% of those reported for microheterotroph regeneration and phytoplankton utilization, respectively. It was 10% of the reported advective flux during periods of low advection and 13% of the maximum efflux from sediments computed from diffusion models. The estimated flux from the demersal macrozooplankton in the sediments exceeded those reported for meiofauna, and was 5 to 32% and 2 to 13% of those reported for ammonification and utilization in sediments, respectively. The potential importance of demersal macrozooplankton in mediating sediment-water column exchanges in the absence of diffusive effluxes and when they swarm is discussed.     

Detrital pathways in a coral reef lagoon

Coral reef lagoons are generally regarded as zones of net heterotrophy reliant on organic detritus generated in more productive parts of the reef system, such as the seaward reef flat. The abundance and biomass of sediment infauna were measured seasonally for one year (1986) within the lagoon of Davies Reef, central Great Barrier Reef, to test the hypothesis that macrofaunal biomass and production of coral reef lagoons would decrease with distance from the reef flat and would change seasonally. In general, there were no simple relationships between infaunal standing stock or production and distance from the reef flat or season. Bioturbation by callianassid shrimps negatively affected the abundance of smaller infauna, suggesting a community limited by biogenic disturbance rather than by supply of organic material. Polychaetes and crustaceans were dominant amongst the smaller infauna (0.5 to 2mm) while larger animals (> 2 mm) were mostly polychaetes and molluscs. Mean biomass of infauna at both sites and all seasons was 3 181 mg C m^?2. The smaller animals (0.5 to 2 mm) contributed about 40% of total macrofaunal respiration and production although they represented only 15% of the total macrofaunal biomass. The biomass of macrofauna was about equal to that of the bacteria and meiofauna, while respiration represented 10 to 20% of total community respiration. Consumption by macrofauna accounts for only 3 to 11% of total organic inputs to sediment, with a further 14 to 17% being lost by macrofaunal respiration.     

Ecological studies of the coral predator Acanthaster planci in the south pacific

Ecological surveys involving over 500 man-days between 1966 and 1969 indicate that the coral-eating sea star, Acanthaster planci, is a normal component of the coral reef community throughout the tropical Pacific, and that its abundance in the past has probably been underestimated. The sea star is not uncommon in certain environments, particularly back-reef and lagoon slopes. Sheltered, inner reefs are generally preferred over less protected reefs. Recently reported population explosions of A. planci at Guam and on the Great Barrier Reef of Australia appear to be isolated, widely-separated, local infestations of unknown cause. The infestation on the Great Barrier Reef has not spread beyond the area off Cairns and Innisfail. Approximately 40 of the more than 1000 reefs comprising the Great Barrier Reef complex have been infested heavily.     

Planktonic bacteria in the Humber Estuary; Seasonal variation in population density and heterotrophic activity

Seasonal variation in population density and heterotrophic activity (as maximum potential rate, V _max, for glucose mineralization) of bacteria attached to suspended solids, of free bacteria and of total bacteria was examined in the water of the Humber Estuary, north-east England. At the principal site, which was studied over 2 yr, values of density and activity of attached and total bacteria showed marked seasonal periodicity, being low in summer and high from autumn to spring. This was a consequence of density of attached bacteria being dependent on the concentration of suspended solids, which was high in winter and low during summer. Values of density and activity of free bacteria, which were less than those of attached bacteria, showed no seasonal pattern but fluctuated irregularly and were independent of concentration of suspended solids. At two further sites, which were studied over 1 yr only, the density of bacteria varied seasonally in a similar manner, but the activity of attached and total bacteria showed no distinct seasonal pattern — this may, however, be due to limitations of the data rather than to fundamental differences between the sites studied.     

Lipid content of some Caribbean corals in relation to depth and light

Three procedures for the extraction of coral lipids were compared and a rapid and effective method for future use is suggested. This method was used to measure the lipid content of the branching coral Porites porites (Pallas) and the massive corals Montastrea annularis (Ellis and Solander) and Siderastrea siderea (Ellis and Solander) during July and August 1991. P. porites and M. annularis were sampled on two fringing reefs, each characterised by different water quality affecting light transmission, and at depths of down to 30 m on a barrier reef on the west coast of the island. m. annularis contained, on average, 29% of dry weight as lipid, and there were no significant differences in lipid levels between corals sampled on either fringing reef at 3 and 6 m, or between fringing reefs and the barrier reef at 13, 20 or 30 m depth. Five samples were also taken from a single massive colony of S. siderea at 3 m on a fringing reef and at 13, 20 and 30 m depth on the barrier reef. Values for lipid ranged from 26 to 35% of dry tissue weight. P. porites from 3 and 6 m depth on both fringing reefs contained the same amount of lipid (11% of dry tissue weight). However, at 13 m depth on the barrier reef this coral contained significantly less lipid (8.5% of dry tissue weight). This difference may be attributable to a higher nutritional intake by P. porites on the fringing reefs.     

In situ swimming and settlement behaviour of larvae of an Indo-Pacific coral-reef fish, the coral trout Plectropomus leopardus (Pisces: Serranidae)

Late larvae of the serranid coral trout Plectropomus leopardus (Lacepède), captured in light traps, were released during the day both in open water and adjacent to two reefs, and their behaviour was observed by divers at Lizard Island, northern Great Barrier Reef. Coral trout larvae (n = 110) were present in light-trap catches from 18 November to 3 December 1997, including new moon (30 November). The swimming speed of larvae in open water or when swimming away from reefs was significantly greater (mean 17.9 cm s⁻¹) than the speed of larvae swimming towards or over reefs (mean 7.2 cm s⁻¹). Near reefs, larvae swam at average depths of 2.7 to 4.2 m, avoiding 0 to 2 m. In open water, swimming depth varied with location: larvae >1 km east of Lizard Island swam steeply downward to >20 m in 2 to 4 min; larvae >1 km west oscillated between 2.6 and 13 m; larvae 100 to 200 m east of Lizard Island oscillated between 0.8 and 15 m. Nearly all larvae swam directionally in open water and near reefs. In open water, the average swimming direction of all larvae was towards the island, and 80% (4 of 5) swam directionally (p < 0.05, Rayleigh's test). Larvae swam directionally over the reef while looking for settlement sites. The frequency of behaviours by larvae differed between two reefs of different exposure and morphology. Depending on site, 26 to 32% of larvae released adjacent to reefs swam to open water: of these, some initially swam towards or over the reef before swimming offshore. In some cases, offshore-swimming seemed to be due to the presence of predators, but usually no obvious cause was observed. Depending on the reef, 49 to 64% of the larvae settled. Non-predatory reef residents aggressively approached 19% of settlers. Between 5 and 17% of the larvae were eaten while approaching the reef or attempting to settle, primarily by lizardfishes but also by wrasses, groupers and snappers. A higher percentage of larvae settled in the second week of our study than in the first. Average time to settlement was short (138 s ± 33 SE), but some larvae took up to 15 min to settle. Average settlement depth was 7.5 to 9.9 m, and differed between locations. No settlement took place on reef flats or at depths <4.2 m. Larvae did not appear to be selective about settlement substrate, but settled most frequently on live and dead hard coral. Late-stage larvae of coral trout are capable swimmers with considerable control over speed, depth and direction. Habitat selection, avoidance of predators and settlement seem to rely on vision. Received: 7 July 1998 / Accepted: 26 January 1999     

Coral recruitment: a spatio-temporal analysis along the coastline of Eilat,northern Red Sea

Recruitment rates of stony corals to artificial substrates were monitored for 2 years at 20 sites along the coast of Eilat, northern Red Sea, to compare with those recorded at other coral reef locations and to assess variation in recruitment at several spatial scales. Coral recruitment was low compared to that observed on the Great Barrier Reef in Australia, but was similar to levels reported from other high-latitude reef locations. Pocilloporids were the most abundant coral recruits in all seasons. Recruitment was twofold higher during the first year than during the second year of study. There was considerable spatial variability, with the largest proportion of variance, apart from the error term, attributable to differences between sites, at a scale of 10² m. Spearmans ranked correlation showed consistency in spatial patterns of recruitment of pocilloporid corals between years, but not of acroporid corals. During spring, when only the brooding pocilloporid coral Stylophora pistillata reproduces at this locality, most coral recruitment occurred at central and southern sites adjacent to well-developed coral reefs. During summer, recruitment patterns varied significantly between years, with wide variation in the recruitment of broadcasting acroporid corals at northern sites located distant from coral reefs. Settlement was low at all sites during autumn and winter. This work is the first detailed analysis of coral recruitment patterns in the Red Sea, and contributes to the understanding of the spatial and temporal scales of variation in this important reef process.Communicated by O. Kinne, Oldendorf/Luhe     

Seston composition of the bottom waters of Great Lameshur Bay,St. John,US Virgin Islands

Seston composition [particulate organic carbon (POC), particulate nitrogen (PN), phyto- and microzooplankton numbers and biomass] was investigated in the bottom waters of Great Lameshur Bay, St. John, Virgin Islands (USA), in October, 1970 during Tektite II Mission 17–50. Mean values of 67.4 g POC · I^-1 and 7.2 g PN · I^-1 were determined. A mean phytoplankton carbon content of 42.2 g · I^-1 and zooplankton carbon content of 5.5 g · I^-1 were calculated from counts. The phytoplankton consisted mainly of dinoflagellates 71.2% phytoplankton carbon. Copepods were the dominant zooplankters (61.8% zooplankton carbon), followed by larvaceans (30.9% zooplankton carbon). Organic carbon content of counted zooplankton faecal pellets ranged between 0.4 and 1.6 g · I^-1, and amounted probably to about 15% of the total zooplankton carbon value. Plankton and detritus components as possible food for coral-reef animals are discussed. The ratio carbon: nitrogen of suspended particles is compared to that of sedimented matter.     

Indications from photosynthetic components that iron is a limiting nutrient in primary producers on coral reefs

Because iron is not available generally in oxygenated sea water, it may be a limiting factor in marine primary production. This hypothesis was tested in the context of Davies Reef, Latitude 18°50′S (one of the coral reefs in the central region of the Great Barrier Reef system). Samples were collected for study in the period August, 1980 to March, 1981. Sea water around the reef contained ≦2x10^-6 M Fe, surface sediments from the reef contained 66±26 (1 SD) ppm total Fe, and interstitial water near the surface contained ≧5x10^-7 M Fe. Thus, Fe constituted a trace component of the reef environment, but limited Fe should be available to algae associated with the sediments. Specific biochemical analyses to test the Fe status of benthic photosynthetic organisms were carried out with a common blue-green alga, Phormidium sp., and a ubiquitous symbiotic dinoflagellate, Gymnodinium microadriaticum (zooxanthellae). The blue-green alga contained the electron transport protein, flavodoxin, which is found only in Fe-deficient organisms. Supporting evidence for Fe stress in this organism included chlorosis in the presence of plentiful biliprotein, and very low extractable photosynthetic cytochrome, c-553. The latter observations were shown to be the result of Fe deficiency in laboratory cultures of a blue-green alga, Synechococcus sp. These cultures showed that production of flavodoxin is not a universal response of algae to Fe stress, but that lowered cellular concentrations of Fe-containing proteins involved in photosynthesis probably is universal. The zooxanthellae from a soft coral, Sinularia sp., had three-fold lower total Fe and ferredoxin (an electron transport protein), than the same alga from a clam, Tridacna maxima. Thus, some algae in symbiotic associations may also suffer Fe-deficiency. It was concluded that the degree and extent of Fe-stress in primary producers on a coral reef may influence growth rates, biomass, and distribution of species.     

Environmental controls of patch-reef growth and development

The effects of predominant currents on the morphology of coral reefs has been studied in the leeward patch reefs of Anegada, British Virgin Islands. Measurements of current velocity and direction and sediment characteristics together with mapping of approximately 31 patch reefs within a 2 km² area suggest that even under relatively low but constant current velocities (surface current 20 cm sec^-1) the major reef-building corals show definite distribution patterns on the reef. The currents also appear to be responsible, in part, for marked orientation and elongation of patch reefs in a north-west/ south-east direction and for a gradation of patch-reef type across the study area.     

Spatial and temporal variation in coral recruitment on the Great Barrier Reef: Implications for dispersal hypotheses

Over 15 000 coral recruits were counted on settlement plates from three mid-shelf reefs and six fringing reefs in the northern section of the Great Barrier Reef during two summers (1986 and 1987) and one winter (1987). The density of coral recruits on some settlement plates from a fringing reef was up to 4.88 cm^–2, the highest value ever reported. Mean density of recruits was greater on fringing reefs (81.1 recruits/settlement plate) than on mid-shelf reefs (15.6 recruits/settlement plate), but there was greater spatial variation in abundance of recruits between the fringing reef sites. Other differences between the mid-shelf reefs and the fringing reefs were that different taxa were dominant, and that settlement orientation differed, with mid-shelf recruits settling preferentially on horizontally oriented surfaces and fringingreef recruits preferring vertical surfaces. Of the three midshelf reefs, Green Island reef recorded the highest recruitment rate for each of the two summers, despite having a depauperate adult coral population following predation by the asteroidAcanthaster planci. This suggests that coral larvae frequently travel between reefs. In contrast with an earlier study, there was no consistent difference in abundance of recruits between forereef and backreef locations. Overall, the results indicated great spatial variation in the availability of coral larvae, both on the scale of whole reefs and within-reef habitats.     

Among-habitat variation in herbivory on Sargassum spp. on a mid-shelf reef in the northern Great Barrier Reef

Herbivory is widely acknowledged as a key process determining the benthic community structure and resilience of coral reefs. Despite numerous studies that have examined herbivory across reef gradients in the Caribbean, few studies have directly quantified this process on Pacific reefs. Bioassays of two species of erect macroalgae (Sargassum swartzii and S. cristaefolium) were used to quantify variation in grazing intensity across seven habitats of varying depth and wave exposure on a mid-shelf reef in the northern Great Barrier Reef. Removal rates of Sargassum varied significantly among habitats, with both species displaying broadly similar patterns. The shallow habitats on the exposed aspect of the reef (i.e. reef crest, flat and back reef) experienced the highest reductions in mass (81.4–91.6% day⁻¹) for both S. swartzii and S. cristaefolium, while the deeper exposed habitats (reef slope and base) displayed the lowest reductions (3.8–13.4% day⁻¹) over a 24 h period. In contrast, the grazing intensity varied between the two species in the three habitats on the leeward aspect of the reef. Reductions in mass remained relatively high for S. swartzii on the patch reef and sheltered reef base and flat (62.7–76.5% day⁻¹) but were considerably lower for S. cristaefolium (37.9–63.5% day⁻¹) across the same habitats. Surprisingly, the rates of removal of Sargassum displayed no relationship with the density or biomass of roving herbivorous fishes or those species known to consume erect macroalgae, either collectively or independently. These results suggest that the relationship between browsing rates and herbivorous fish biomass is complex and may be driven by species that are underestimated in visual surveys. Direct quantification of browsing intensity using assays revealed a different pattern to inferences based on herbivore densities and highlights the potential difficulties of evaluating ecosystem processes based on visual census data alone.     

The kinetics and possible significance of nitrate uptake by several algal-invertebrate symbioses

Nitrification in the coral reef community at Lizard Island, Great Barrier Reef, Australia, elevated the nitrate concentration to above that of the nearby open ocean water. Reef corals and a zooxanthellae-bearing foraminiferan were shown to take up nitrate from nitrate-enriched seawater; a lag period was absent, indicating that the responsible enzymes did not require induction. The relationship of nitrate uptake to seawater nitrate concentration could be described by a hyperbola with a non-zero intercept on the abscissa. Corals are opportunistic in acquiring nitrogen; in addition to gaining particulate nitrogen from ingested food, they acquire dissolved nitrogen in the form of nitrate, ammonia and urea.     

Detrital pathways in a coral reef lagoon

Coral reef lagoons have generally been regarded as sinks for organic matter exported from more productive reef front and reef flat zones. The object of this study was to examine the importance of detritus as a carbon source for benthic communities in the lagoon at Davies Reef, central Great Barrier Reef. We report the results of seasonal measurements, taken in 1986, of bacterial numbers and production, protozoan numbers, community primary production and respiration in the sediments of Davies Reef lagoon. Deposition rates of organic matter in the lagoon were also measured. Deposition rates (±1 SE) of carbon ranged from 9.2 (±1.5) to 140.7 (±10.3) mg Cm^-2d^-1. Deposition rates were highest in winter and spring, lowest in summer. Rates of bacterial production ranged from 4.7 (±0.2) pmol thymidine incorporated g^-1 dry wt (DW) h^-1 in winter to 23.5 (±1.0) pmol thymidine incorporated g^-1 DW h^-1 in spring. The number of ciliates ranged from 65 (±10) to 356 (±50) cm^-3 through the year and the number of large (20 m) flagellates from 38 (±7) to 108 (±16) cm^-3. There were no clear relationships between the sediment organic content, detrital input or temperature and the rates of bacterial processes, community metabolism or the standing stocks of microbes in the lagoon. The relative significance of detritus and in situ primary production as sources of carbon in the lagoon varied with season. In summer and autumn, detritus was less important than primary production as a source of carbon (4 to 27% of total carbon input). In winter and spring, detritus input became more significant in supply of carbon to the sediments (32 to 67% of the total carbon input). The lagoon does not simply act as a sink for carbon exported from the reef flat. We calculate that only 5% of the net reef flat primary production reached lagoon sediments in summer, but nearly 40% in winter.     

Genetic differentiation of bicolor damselfish (Eupomacentrus partitus) populations in the Florida Keys

Electrophoretic variation in proteins encoded by 23 loci revealed substantial genetic differentiation among populations of bicolor damselfish (Eupomacentrus partitus) collected from four coral reefs in the Florida Keys, USA, during 1986–1988. Genetic differentiation was concentrated between a sample collected from Little Grecian Rocks Reef (LGR) and the remaining samples, including fish from a reef only 600 m distant (Grecian Rocks Reef). Genetic distinction of the LGR sample derived from significantly heterogenous allelic frequencies at six of eight polymorphic loci. Aco-1 (aconitase); Ada (adenosine deaminase); Gpi-2 (glucosephosphate isomerase); Ldh-2 and Ldh-3 (lactate dehydrogenase); and Me-1 (malic enzyme); nevertheless, differentiation at cytosolic aconitase (Aco-1) far exceeded that observed for other loci (fixation index, F _ST=0.482), and differences in Aco-1 allele frequencies were largely responsible for large genetic distances (0.20) between LGR and the other reefs. Paradoxically, estimates of numbers of migrants exchanged between reefs per generation (mN _e=17.47) indicated the potential for extensive gene flow. The extent of genetic differentiation among these populations is evaluated relative to models of population genetic structure based on equilibrium between gene flow and natural selection or genetic drift.     

A link between biologically imported particulate organic nutrients and the detritus food web in reef communities

Previous work with planktivorous fishes has shown that they import particulate organic and inorganic material to reefs in the form of fecal pellets, which, in part, are deposited in crevices on the reef where these fishes shelter during their inactive period. Since these feces do not accumulate in fish shelters, we predicted that some of the feces could be rapidly consumed by reef detritivores. We examined the attractiveness of fish feces to potential reef detritivores by placing traps baited with planktivorous fish feces, along with unbaited control traps, in crevices on rocky reefs at Santa Catalina Island, California, USA, between June 1982 and November 1983, and on coral reefs at St. Croix, U.S. Virgin Islands, during June 1983. Significantly more animals (the majority being crustaceans) were trapped in the baited traps compared to the unbaited controls on both reefs. There was also a significant association between the presence of trapped animals and fish feces at Santa Catalina Island (p = 0.009); this association was not quite as strong in St. Croix (p = 0.069). The consumption by shrimp of feces marked with carmine particles and, in turn, the predation on these shrimp by a reef fish demonstrates the links between this type of imported particulate organic and inorganic material and the food web of the reef community.Please address all correspondence and requests for reprints to Dr. Miller at California State University, Long Beach