Mass transfer limitation of photosynthesis of coral reef algal turfs

Algal turfs are the major primary producing component on many coral reefs and this production supports higher levels in the complex reef trophic web. Rates of metabolism of algal turfs are related positively to water motion, consistent with limitation by the diffusion of a substance through a boundary layer. Based on engineering mass transfer theory, we hypothesized that photosynthesis of algal turfs is controlled by rates of mass transfer and responses of photosynthesis to increasing flow speed should be predicted by engineering correlations. This hypothesis was tested in ten experiments where photosynthesis was estimated in a flume/respirometer from changes in dissolved oxygen at eight flow speeds between 0.08 and 0.52 m/s. Flow in the flume and over the reef at Kaneohe Bay, Oahu, Hawaii was estimated using hot-film thermistor and electromagnetic current meters. Rates of photosynthesis were related positively to flow in all experiments and plots of the log of the average Sherwood number (Sh _meas) versus log Reynolds number (Re _D) for each experiment are lower than predicted for mass transfer through a turbulent boundary layer. Algal turf-covered plates are characterized as hydrodynamically transitional to fully rough surfaces and the lower than predicted slopes suggest that roughness reduces rates of mass transfer. A negative correlation between algal turf biomass and slopes of the log Sh _meas−log Re _D plots suggests that mass transfer to algal turfs is affected significantly by the physical structure of the algal community. Patterns of photosynthesis based on changes in dissolved oxygen and dissolved inorganic carbon concentrations (DIC) indicate that the flow speed effect is not the result of increased flux of oxygen from the algal turfs, and combined with the short response time to flow speed, suggest that DIC may limit rates of photosynthesis. Although there are differences between flow in the flume and flow over algal turfs on the reef, these results suggest that photosynthesis is controlled, at least in part, by mass transfer. The chemical engineering approach provides a framework to pose further testable hypotheses about how algal canopy height, flow oscillation, turbulence, and substratum roughness may modulate rates of metabolism of coral reef algal turfs.     

Nitrogen fixation on a coral reef

Acetylene reduction was used to assess nitrogen fixation on all major substrates at all major areas over a period of 1 to 6 yr (1980–1986) at One Tree Reef (southern Great Barrier Reef). Experiments using ¹⁵N₂ gave a ratio of 3.45:1.0 for C₂H₂ reduced:N₂ fixed. Acetylene reduction was largely light-dependent, saturated at 0.15 ml C₂H₂ per ml seawater, and linear over 6 h. High fixation was associated with two emergent cyanophyte associations, Calothrix crustacea and Scytonema hofmannii, of limited distribution. Subtidally, the major contribution to nitrogen fixation came from well-grazed limestone substrates with an epilithic algal community in the reef flat and patch reefs (3 to 15 nmol C₂H₄ cm^-2 h^-1). Similar substrates from the outer reef slope showed lower rates. Nitrogen fixation on beach rock, intertidal coral rubble, reef crest and lagoon sand was relatively small (0.3 to 1.0 nmol C₂H₄ cm^-2 h^-1). Seasonal changes in light-saturated rates were small, with slight reduction only in winter. Rates are also reported for experimental coral blocks (13 to 39 nmol cm^-2 h^-1) and for branching coral inside and outside territories of gardening damselfish (3 to 28 nmol cm^-2 h^-1). This work supports the hypothesis that the high nitrogen fixation on the reef flat and patch reefs of the lagoon (34 to 68 kg N ha^-1 yr^-1) is because these subtidal areas support highly disturbed communities with the greatest abundance of nitrogen-fixing cyanophyte algae. It is calculated from a budget of all areas that One Tree Reef has an annual nitrogen fixation rate of 8 to 16 kg N ha^-1 yr^-1.     

Photorespiration and productivity rates of a coral reef algal turf assemblage

In studies conducted from 1982 to 1983, productivity (¹⁴C uptake) of a coral reef algal-turf assemblage was unaffected by oxygen concentration but decreased when pH rose, probably in response to declining CO₂ and HCO ₃ ^- supplies. Release of prefixed ¹⁴C was substantially lower in the light than in the dark and was unaffected by oxygen concentration. Release of organic prefixed ¹⁴C was greater in the light than in darkness. Total CO₂ compensation-points were low, showing no consistent response to oxygen or the photorespiratory inhibitor alpha-hydroxy-2-pyridine-methanesulfonic acid (HPMS). Oxygen has little if any influence upon turf productivity, which was high in comparison to other benthic algae. Decreases in net carbon-fixation rates of this turf more likely result from decreased photosynthesis than increased photorespiration, which is either not significant in turf metabolism under natural conditions or is compensated by efficient refixation of respired carbon.This paper describes research by J. M. Hackney in partial fulfillment of the requirements for the Ph.D. degree at Georgetown University     

Two contrasting effects of predation on species richness in coral reef habitats

The species richness of sessile organisms on settlement panels on a coral reef was measured by the slope of a regression of log_e number of species against log_e area of sample. At a well illuminated site where panels were colonised by algae, the species richness of algae was 19% smaller on surfaces grazed by fishes than on protected surfaces. At a second site in a cave, the species richness of animals on grazed surfaces was 20% greater than on protected surfaces. These results are discussed in the light of differences between the sites. The contrasting effects of predation at the two sites are probably the result of more selective predation at the cave site than at the other site.     

The effects of mating system on male mate choice in a coral reef fish

Summary We show how mate limitation appears to be critical in determining whether or not males exercise mate choice among available females. Thalassoma bifasciatum is a Caribbean reef fish with two distinct mating patterns: group-spawning and pair-spawning. In both mating systems, female fecundity is variable and size dependent, and female availability is high. However, sperm competition among group-spawning males apparently limits the number of effective matings in which a male may engage, whereas territorial pair-spawning males have little or no such limitation. Group-spawning males should be discriminating in their choice of mates and our data confirm this: there is strong evidence for assortative mating in group-spawns, with more large males joining in mating groups around large females. In contrast, pair-spawning males show no indication of mate preferences, and spawn with all females who arrive at their territories.     

Acute effects of removing large fish from a near-pristine coral reef

Large animals are severely depleted in many ecosystems, yet we are only beginning to understand the ecological implications of their loss. To empirically measure the short-term effects of removing large animals from an ocean ecosystem, we used exclosures to remove large fish from a near-pristine coral reef at Palmyra Atoll, Central Pacific Ocean. We identified a range of effects that followed from the removal of these large fish. These effects were revealed within weeks of their removal. Removing large fish (1) altered the behavior of prey fish; (2) reduced rates of herbivory on certain species of reef algae; (3) had both direct positive (reduced mortality of coral recruits) and indirect negative (through reduced grazing pressure on competitive algae) impacts on recruiting corals; and (4) tended to decrease abundances of small mobile benthic invertebrates. Results of this kind help advance our understanding of the ecological importance of large animals in ecosystems.     

Speciation of nitrogen in the coral reef sedimentary environment of Lakshadweep archipelago,Indian ocean

This paper focuses on the spatial distributional profiles of different species of the important micronutrient element, nitrogen (nitrite, nitrate and total nitrogen) in various coral-reef sedimentary environment of Lakshadweep Archipelago. The relative abundance of the three forms of nitrogen was in the order, total-N???nitrate-N???nitrite-N. Relatively very low levels of nitrite in the different microenvironments of the islands are an indication of a higher rate of nitrification, so as to produce thermodynamically most stable form of nitrogen, namely nitrate under the condition of well-oxygenated shallow coastal/lagoon waters. A lagoon-ward enrichment pattern of total nitrogen in the lagoon transects of Agathy, Minicoy, Kadamath and Kiltan Islands also reflected the fact that the rate and space available for nitrogen fixation in shallow zones of the lagoon are high. Further, the nitrogenous waste materials produced from the reefs and surrounding environments have limited exchange with the sea, and all these factors, together with the organic nitrogen retention capacity of sediment types, contribute to total nitrogen.     

Siliceous sponge spicules in coral reef sediments

Experimental etching with hydrofluoric acid indicated that silica deposition occurs in a recognizable pattern in common sponge microscleres. The postdepositional alteration of these spicules has previously been generally unrecognized or misinterpreted in the literature. Early stages of postdepositional etching of sponge spicules were observed in the acid insoluble fraction of sediments from the West Atlantic barrier reef near Carrie Bow Cay, Belize. Preliminary data on silica distribution in the Belize barrier reef show that concentrations in fine sediment (<0.25 mm) increase landward of the main reef tract. Sponge spicules are the main component of particulate silica in sediments of the reef and fore-reef where sponge populations abound, whereas grains prevail in the back-reef lagoon deposits. Recycling of locally dissolved silica appears to be important for the growth of many off-shore reef sponges.     

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.     

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.     

Increased behavioural lateralization in parasitized coral reef fish

Preferential use of one side of the body for cognitive or behavioural tasks (lateralization) is common in many animals, including humans. However, few studies have demonstrated whether lateralization is phenotypically plastic, and varies depending on the ecological context. We studied lateralization (measured as a turning preference) in the bridled monocle bream (Scolopsis bilineatus). This coral reef fish is commonly infected by a large, ectoparasitic isopod (Anilocra nemipteri) that attaches to the left or right side of its host’s head. Fish that were parasitized showed no turning bias with respect to the side on which the parasite had attached. On average, however, parasitised fish were significantly more lateralized (i.e. had a strong side bias) than unparasitized fish. The extent of lateralization declined significantly when we experimentally removed the parasite. Our results indicate that lateralization can vary with the ecological context. One possible explanation is that lateralization shortens the response time until fish flee after encountering a predator. A stronger side bias might be advantageous for parasitized individuals to overcome their recently documented lower maximum swimming speed.     

Diurnal space utilization in coral reef fish communities

The diurnal use of space by 25 resident species of coral reef fishes was investigated along three depth (10 to 40 m) transects over an 18 month period. Emphasis was placed on the small, cryptic members of the communities. Three aspects of the use of space — temporal utilization, occupancy and time span — are defined and quantified for each of the species. In addition, an estimate of the diurnal home range volume of each species is provided. The territorial pomacentrids showed the highest level of temporal utilization, holding the same territories throughout the study. The consistent presence of the holocentrid species within the reef infrastructure had the potential to influence the space available to other species. Some species residing in tube-worm holes and sponges also exhibited constant use of space.     

Diel patterns of abundance of presettlement reef fishes and pelagic larvae on a coral reef

Most presettlement reef fish settled at night at One Tree Island, Great Barrier Reef. Fish were sampled day and night using channel nets located on the reef crest, and a plankton-mesh purse-seine net in the lagoon (1992–1994). Catches of fish at night were generally tens to hundreds of times greater than those taken during the day. Preflexion fish, as well as postflexion and pelagic juveniles, were taken in greater numbers at night. Preflexion forms were a combination of those that had hatched from demersal eggs and later stages that had been transported over the reef crest. Highest numbers of postflexion and pelagic juvenile forms of Apogonidae, Blenniidae, Gobiesocidae, Gobiidae, Labridae, Lutjanidae, Mugiloididae, Mullidae, Pomacentridae, Scaridae, Serranidae and Tripterygiidae were found at night. Observations, while SCUBA diving, and purse-seine samples in the lagoon indicated that the only resident larvae were of the genera Spratelloides and Hypoatherina; most of the fishes caught in nets, therefore, were immigrants. Patch reefs, sampled for new settlers early in the morning and late in the day, indicated that the majority of apogonids (Apogon doederleini, >95%) settled at night. Although greater numbers of pomacentrids were found in morning counts (e.g. Pomacentrus wardi), if data were converted to an hourly rate, many pomacentrids showed a similar hourly rate of settlement day and night. Depth-stratified sampling in waters near One Tree Island (to 20 m) indicated that some taxa rise to the surface at night. This behaviour, perhaps combined with avoidance of diurnal predators may explain on-reef movement of potential settlers soon after dark. Studies on settlement cues, therefore, need to focus on night-related phenomena. Received: 3 March 2000 / Accepted: 20 June 2000     

Microhabitat utilisation patterns in cryptobenthic coral reef fish communities

The cryptobenthic reef fish communities from four microhabitats at Orpheus Island, central Great Barrier Reef are described. Eighty-four 0.4m² samples yielded a total of 368 individuals from 42 species in eight families, with a mean density of 11 individuals m^–2 (±1.7SE) and 2.9 species 0.4 m^–2 (±0.2SE). Caves contained the highest number of both individuals (120) and species (26), followed by sand/rubble, soft coral, and open reefs. Microhabitat associations included cave and soft coral specialists. Site fidelity in 71 tagged individuals of 4 species was high, with a mean recapture rate of 53% (±8.4SE) remaining within the ~0.4 m² sampling area after a 48-h period. Behavioural observations also reflect this limited movement, with the dominant mode of behaviour in 7 species being a motionless state (67.5% ±11.6SE), followed by feeding (21.8% ±8.7SE), hiding (6.3% ±1.6SE), and swimming (4.4% ±1.5SE). Two distinct behavioural groups are identified: (1) sedentary forms, characterised by long periods of immobility (5 species); and (2) winnowers, characterised by long feeding bouts (2 species). The fine-scale partitioning of microhabitats, restricted home ranges, and sedentary behaviour of many cryptobenthic reef fish species suggest that this reef fish community exhibits similar patterns of habitat utilisation to their larger reef-fish counterparts, but at a much finer scale.     

Predator biomass,prey density,and species composition effects on group size in recruit coral reef fishes

Group incidence and size are described for recruit parrotfishes, wrasses, and damselfishes on Hawaiian reefs over 3 years (2006–2008) at sites spanning the archipelago (20–28°N, 155–177°W). Coral-poor and coral-rich areas were surveyed at sites with both low (Hawaii Island) and high (Midway Atoll) predator densities, facilitating examination of relations among predator and recruit densities, habitat, and group metrics. Predator and recruit densities varied spatially and temporally, with a sixfold range in total recruit densities among years. Group (≥2 recruits) metrics varied with time and tracked predator and recruit densities and the proportion of schooling species. Groups often included heterospecifics whose proportion increased with group size. A non-saturating relationship between group size and recruit density suggests that the anti-predator benefits of aggregation exceeded competitive costs. Grouping behavior may have overarching importance for recruit survival–even at high recruit densities–and merits further study on Hawaiian reefs and elsewhere.     

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     

Impact of a kaolin clay spill on a coral reef in Hawaii

On April 27, 1980, the Greek freighter Anangel Liberty went aground on the reef at French Frigate Shoals, a National Wildlife Refuge in the Hawaiian Islands. The vessel was refloated with no major damage or fuel spillage after 2 200 tons (2 200 000 kg) of koalin cargo had been jettisoned on the reef. Huge plumes of suspended clay raised major concern over the possibility of widespread ecological damage. However, field investigations conducted 14 d after the kaolin was dumped revealed that environmental impact waw very minor and highly localized; it was evident that most of the kaolin had been suspended and removed from the area. The only significant damage was a 2 to 3 m deep channel plowed through the reef by the freighter. Within 50 m of both sides of the channel, some coral was smothered and colonies of Pocillopora spp. were alive but slightly bleached. Beyond 50 m there were no apparent impact, nor did any clay settle on the bottom. This incident illustrates that some events which initially appear to have potential pollutant impact do not produce significant and irreversible environmental changes and emphasizes the need to analyze such events on a case-by-case basis.Hawaii Institute of Marine Biology Contribution No. 610     

Toward pristine biomass: reef fish recovery in coral reef marine protected areas in Kenya.

Identifying the rates of recovery of fish in no-take areas is fundamental to designing protected area networks, managing fisheries, estimating yields, identifying ecological interactions, and informing stakeholders about the outcomes of this management. Here we study the recovery of coral reef fishes through 37 years of protection using a space-for-time chronosequence of four marine national parks in Kenya. Using AIC model selection techniques, we assessed recovery trends using five ecologically meaningful production models: asymptotic, Ricker, logistic, linear, and exponential. There were clear recovery trends with time for species richness, total and size class density, and wet masses at the level of the taxonomic family. Species richness recovered rapidly to an asymptote at 10 years. The two main herbivorous families displayed differing responses to protection, scarids recovering rapidly, but then exhibiting some decline while acanthurids recovered more slowly and steadily throughout the study. Recovery of the two invertebrate-eating groups suggested competitive interactions over resources, with the labrids recovering more rapidly before a decline and the balistids demonstrating a slower logistic recovery. Remaining families displayed differing trends with time, with a general pattern of decline in smaller size classes or small-bodied species after an initial recovery, which suggests that some species- and size-related competitive and predatory control occurs in older closures. There appears to be an ecological succession of dominance with an initial rapid rise in labrids and scarids, followed by a slower rise in balistids and acanthurids, an associated decline in sea urchins, and an ultimate dominance in calcifying algae. Our results indicate that the unfished "equilibrium" biomass of the fish assemblage > 10 cm is 1100-1200 kg/ha, but these small parks (< 10 km2) are likely to underestimate pre-human influence values due to edge effects and the rarity of taxa with large area requirement, such as apex predators, including sharks.     

Interactions during feeding among certain coral reef fishes in Elat

From July 1983 to March 1984 coral reef fishes in Elat (Red Sea) were fed in novel feeding situations. Thalassoma rüppelli had the shortest latency to first feed, Chaetodon paucifasciatus had the longest latency, while Sufflamen albicaudatus, Coris aygula and Lethrinus sp. had intermediate latencies. The mean number of T. rüppelli feeding at the beginning of experiments was higher than that of the other species. The difference decreased rapidly and disappeared within 90 s of the beginning. Latencies to first feed, and the number of fish feeding, were not correlated with the number of fish of each species in the study area. Aggression was directed predominantly by S. albicaudatus and by C. aygula towards Lethrinus sp. and T. rüppelli. It is suggested that these species which suffer more disturbances during their feeding and which receive more aggression are more likely to approach and feed more quickly in novel feeding situations.