Mineralization of particulate organic matter derived from coral-reef organisms in reef sediments of the Gulf of Aqaba

In situ and laboratory incubation experiments in a fringing reef in the Gulf of Aqaba were performed to study degradation rates of particulate organic matter in reef sediments. Coral mucus, clam eggs, and zooxanthellae were used as model particulate organic compounds for these experiments. Aerobic and anaerobic mineralization rates were calculated by dissolved inorganic carbon (DIC) and O₂ fluxes from the sediments under different particulate organic matter additions. Fast enhancement (approximately twofold) of O₂ and DIC fluxes were found with the addition of coral mucus and clam eggs compared with control incubations without addition. Most of the degradation is believed to have occurred anaerobically rather than aerobically (DIC:O₂ ratios were 4.3-28.1). Higher degradation rates of coral mucus and clam eggs were estimated in carbonate sediment than in silicate sediment (1.2-1.6-fold), which was attributed to the different physical and chemical properties of both sediments. Our study shows the significance of the reef sediment as a suitable site for microbial degradation of particulate organic material excreted from different reef community organisms. This may increase the regeneration of nutrients in the reef environment necessary to sustain high biological productivity.     

It's a dirty job but someone has to do it: The role of marine benthic macrofauna in organic matter turnover and nutrient recycling to the water column

Benthic macrofaunal populations through their feeding, bioturbation, burrow construction and sediment irrigation activities have profound influences on organic matter inputs to marine sediments (biodeposition) and on the vertical distribution of deposited organic matter within the sediment. These effects in turn influence the rates and pathways of organic matter mineralisation, and element cycles. Similarly, bioturbation, burrow construction and burrow irrigation are major determinants of sediment-water column fluxes of oxygen and nutrients.

In this review, I discuss the influences of the different benthic macrofaunal feeding (functional) groups on mineralisation processes and sediment-water column fluxes of particulate and dissolved nutrients. How these effects influence diagenic processes, the balance between aerobic and anaerobic processes, and the redox status of the surficial sediments. Finally, I discuss some of the limitations of the predominantly laboratory techniques which have been used to study “macrofaunal effects” and how this hinders the inclusion of the effects in quantitative sediment biogeochemical models.     

Coral mucus stable isotope composition and labeling: experimental evidence for mucus uptake by epizoic acoelomorph worms

Mucus released by scleractinian corals can act as an important energy and nutrient carrier in coral reef ecosystems, and a distinct isotopic signature would allow following the fate of this material. This study investigates the natural C and N stable isotopic signatures of mucus released by four scleractinian coral genera (Acropora, Fungia, Pocillopora and Stylophora) in comparison with those of suspended particulate organic matter (POM) in seawater of a Northern Red Sea fringing coral reef near Aqaba, Jordan. The natural δ¹³C and δ¹⁵N signatures of coral mucus differed significantly from seawater POM for the majority of seasonal comparisons, but were inappropriate for explicit tracing of mucus in the coral reef food web. Thus, a labeling technique using stable isotope tracers (¹³C and ¹⁵N) was developed that produced δ¹³C values of up to 122 ± 5‰ (mean ± SE) and δ¹⁵N of up to 2,100 ± 151‰ in mucus exuded by Fungia corals. ¹³C and ¹⁵N-enriched compounds were rapidly (within 3 h) and light-dependently transferred from the endosymbiotic zooxanthellae to the mucus-producing coral host. The traceability of ¹⁵N-labeled mucus was examined by evaluating its uptake and potential utilization by epizoic acoelomorph Waminoa worms naturally occurring on a range of scleractinian coral taxa. This tracer experiment resulted in uptake of coral mucus by the coral-associated acoelomorphs and further demonstrated the possibility to trace stable isotope-labeled coral mucus by revealing a new trophic pathway in coral reef ecosystems.     

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.     

Effect of tidal currents,seston, and bottom sediments on growth of Mercenaria mercenaria: results of a field experiment

Tidal currents, seston, and sediments separately influence growth of the hard clam, Mercenaria mercenaria, but it is uncertain how these factors may interact. A 3×3 factorial field experiment, carried out in Great Sound, a coastal lagoon in Southern New Jersey, USA, between May and September 1986, determined the relative effects of three sediment types and three site-specific seston/tidal current regimes on the individual growth of M. mercenaria. Analysis of variance of the change in shell length after 15 wk (differences in initial and final lengths) demonstrated a significant difference (P=0.0064) in growth among sites, but no significant differences (P=0.1331) for growth in different sediments, although trends were evident. Effects of sites were independent of sediment type (P=0.2621). Shell growth rates differed by 10.7% between the slowest and fastest sites, but only differed by 5.7% between sediment types, with fastest growth in sand and slowest in mud. Tidal current speeds and four measures of seston (chlorophyll a, particulate inorganic and organic matter, PIM and POM, and energy content) were measured >20 times in near-bottom waters at each site. Horizontal fluxes of POM exhibited higher correlation coefficients with growth rates, than did seston concentrations or current speeds alone. We attribute significant site differences to differences in horizontal seston fluxes fluxes among sites. We suggest that horizontal seston fluxes may be a major factor affecting individual growth of suspension-feeding bivalves.No. 216 of the Jackson Estuarine Laboratory Contribution Series     

It's a dirty job but someone has to do it: The role of marine benthic macrofauna in organic matter turnover and nutrient recycling to the water column

Benthic macrofaunal populations through their feeding, bioturbation, burrow construction and sediment irrigation activities have profound influences on organic matter inputs to marine sediments (biodeposition) and on the vertical distribution of deposited organic matter within the sediment. These effects in turn influence the rates and pathways of organic matter mineralisation, and element cycles. Similarly, bioturbation, burrow construction and burrow irrigation are major determinants of sediment-water column fluxes of oxygen and nutrients.

In this review, I discuss the influences of the different benthic macrofaunal feeding (functional) groups on mineralisation processes and sediment-water column fluxes of particulate and dissolved nutrients. How these effects influence diagenic processes, the balance between aerobic and anaerobic processes, and the redox status of the surficial sediments. Finally, I discuss some of the limitations of the predominantly laboratory techniques which have been used to study “macrofaunal effects” and how this hinders the inclusion of the effects in quantitative sediment biogeochemical models.     

Diffusive boundary layers and photosynthesis of the epilithic algal community of coral reefs

The effects of mass transfer resistance due to the presence of a diffusive boundary layer on the photosynthesis of the epilithic algal community (EAC) of a coral reef were studied. Photosynthesis and respiration of the EAC of dead coral surfaces were investigated for samples from two locations: the Gulf of Aqaba, Eilat (Israel), and One Tree Reef on the Great Barrier Reef (Australia). Microsensors were used to measure O₂ and pH at the EAC surface and above. Oxygen profiles in the light and dark indicated a diffusive boundary layer (DBL) thickness of 180–590 μm under moderate flow (~0.08 m s^?1) and >2,000 μm under quasi-stagnant conditions. Under light saturation the oxygen concentration at the EAC surface rose within a few minutes to 200–550% air saturation levels under moderate flow and to 600–700% under quasi-stagnant conditions. High maximal rates of net photosynthesis of 8–25 mmol O₂ m^?2 h^?1 were calculated from measured O₂ concentration gradients, and dark respiration was 1.3–3.3 mmol O₂ m^?2 h^?1. From light–dark shifts, the maximal rates of gross photosynthesis at the EAC surface were calculated to be 16.5 nmol O₂ cm^?3 s^?1. Irradiance at the onset of saturation of photosynthesis, E _k, was <100 µmol photons m^?2 s^?1, indicating that the EAC is a shade-adapted community. The pH increased from 8.2 in the bulk seawater to 8.9 at the EAC surface, suggesting that very little carbon in the form of CO₂ occurs at the EAC surface. Thus the major source of dissolved inorganic carbon (DIC) must be in the form of HCO₃ ^?. Estimates of DIC fluxes across the DBL indicate that, throughout most of the daytime under in situ conditions, DIC is likely to be a major limiting factor for photosynthesis and therefore also for primary production and growth of the EAC.     

Seafloor ecosystem functioning: the importance of organic matter priming

Organic matter (OM) remineralization may be considered a key function of the benthic compartment of marine ecosystems and in this study we investigated if the input of labile organic carbon alters mineralization of indigenous sediment OM (OM priming). Using ¹³C-enriched diatoms as labile tracer carbon, we examined shallow-water sediments (surface and subsurface layers) containing organic carbon of different reactivity under oxic versus anoxic conditions. The background OM decomposition rates of the sediment used ranged from 0.08 to 0.44 μmol C ml_ws⁻¹ day⁻¹. Algal OM additions induced enhanced levels of background remineralization (priming) up to 31% and these measured excess fluxes were similar to mineralization of the added highly degradable tracer algal carbon. This suggests that OM priming may be important in marine sediments.     

Coral cavity sponges depend on reef-derived food resources: stable isotope and fatty acid constraints

The diet of cavity sponges on the narrow fringing reefs of Curaçao, Caribbean was studied. The origin and resources of the bulk food of these sponges, i.e., dissolved organic matter (DOM), were identified using stable carbon and nitrogen isotopes and fatty acid biomarkers. We found that phytoplankton and its derived DOM from the adjacent open sea and from reef overlying water is not the main source of food for most of the sponges examined nor is bacterioplankton. Interestingly, dual stable isotope signatures (δ¹³C_org, δ¹⁵N_org) and fatty acid biomarkers appoint coral mucus and organic matter derived from crustose coralline algae (CCA) as probable food sources for encrusting sponges. Mucus-derived DOM may contribute up to 66% to the diet of examined sponges based on results of dual isotope mixing model analysis. The contribution of CCA (as purported representative for benthic algae) was smaller with values up to 31%. Together, mucus- and CCA-derived substrates contributed for 48–73% to the diet of sponges. The presence of the exogenous fatty acid 20:4ω6 in sponges, which is abundant in coral mucus of Madracis mirabilis and in CCA, highlights these reef-derived resources as sources of nutrition for DOM feeding cavity sponges. The relatively high concentrations of exogenous 20:4ω6 in all sponges examined supports our hypothesis that the bulk of the food of the cavity sponge community is reef-derived. Our results imply that cavity sponges play an important role in conserving food and energy produced within the reef.     

Exponential Collapse of Benthic Depurative Capacity in A Eutrophic Tropical Marine Ecosystem: A Threshold Response to Organic Nitrogen Loading

Microbial decomposition of particulate organic matter in sediments can substantially modulate eutrophication of nearshore marine environments. Flux budgets for nitrogen compounds and quantification of rates of microbial transformation can provide important information on the process of eutrophication. This survey documents sediment nitrogen budgets for a eutrophic nearshore marine environment at La Parguera on southwest coast of Puerto Rico, including consideration of the organic fraction in addition to microbial transformations.

Sediments of the inshore channel at La Parguera denote the input of heavy organic loads with low redox potentials and high ammonium contents. Benthic fluxes of dissolved organic nitrogen are below those recorded for ammonium. These observations suggest that nearshore tropical sediments subject to heavy nitrogen loads act as a sink for organic nitrogen and that ammonification is a key process in the release of nitrogen from sediments to the water column. Nitrification and denitrification rates are low overall and inversely related to the redox potential. Depuration of excess nitrogen through denitrification is ineffective in these sediments. in contrast to more robust temperate environments, our work demonstrates that tropical marine systems are particularly susceptible to eutrophication given their limited capacity for depuration of excess nitrogen.     

Effects of mussel (Perna canaliculus) biodeposit decomposition on benthic respiration and nutrient fluxes

Suspension-feeding bivalves increase the quantity and quality of sedimenting organic matter through the production of faeces and pseudofaeces that are remineralised in coastal sediments and thus increase sediment oxygen demand and nutrient regeneration. Bivalves are intensively cultivated worldwide; however, no bivalve biodeposit decay rates are available to parameterise models describing the environmental effects of bivalve culture. We examined sediment biogeochemical changes as bivalve biodeposits age by incubating coastal sediments to which we added fresh mussel (Perna canaliculus) biodeposits and measured O₂ and nutrient fluxes as well as sediment characteristics over an 11-day period. Biodeposits elevated organic matter, chlorophyll a, phaeophytin a, organic carbon and nitrogen concentrations in the surface sediments. Sediment oxygen consumption (SOC) increased significantly (P=0.016) by ∼1.5 times to 1,010 μmol m⁻² h⁻¹ immediately after biodeposit addition and remained elevated compared to control cores without additions for the incubation period. This increase is in the range of observed in situ oxygen demand enhancements under mussel farms. To calculate a decay rate for biodeposits in sediments we fitted a first-order G model to the observed increase in SOC. The significant model fit (P=0.001, r ²=0.72) generated a decay rate of 0.16 day⁻¹ (P=0.033, SE=0.05) that corresponds to a half-life time of 4.3 day. This decay rate is 1–2 orders of magnitude higher than published decay rates of coastal sediments without organic enrichment but similar to rates of decaying zooplankton faecal pellets. NH₄⁺ release increased rapidly on the day of biodeposit addition (P=0.013) and reached a maximum of 144 μmol m⁻² h⁻¹ after 5 days which was 3.6 times higher compared to control cores. During this period NH₄⁺ release was significantly (P<0.001 to P=0.043) higher in the cores with biodeposit additions than in control cores.     

Productivity of bacteria and microalgae and the effect of grazing by holothurians in sediments on a coral reef flat

Bacterial productivity in sandy sediments on reef flats at Lizard Island, Great Barrier Reef was determined from the rate of incorporation of tritiated thymidine into DNA. The study was conducted during January 1982 and July 1983. A small diurnal increase occurred in sediments having a dense population of microalgae. Bacterial production was 120 to 370 mg C m^-2 d^-1 in summer on reef flats, which was equivalent to 30–40% of primary production by benthic microalgae. In winter, rates of primary production by benthic microalgae and secondary production by bacteria were about one-half to one-fifth of those in summer. There was much variation in production, due to patchiness in the distribution of benthic microbes, especially microalgae. Doubling times for the bacteria in surface sediment were 1 to 2 d in summer and 4 to 16 d in winter on the reef flats. These high productivity values for bacteria indicated that a net input of organic matter to the sediment was needed to support the growth of bacteria. Sediment bacteria thus have a very important role in transforming organic matter on the reef flats. Grazing by Holothuria atra depressed both primary production and bacterial production. It was estimated that these holothurians ate about 10 to 40% of bacterial carbon produced each day in summer, and thus have an important role in the carbon cycle. Harpacticoid copepods were numerically important components of the benthic meiofaunal community and probably had a significant impact on bacterial density as grazers.     

Ingestion and incorporation of coral-mucus detritus by reef zooplankton

The copepod Acartia tonsa and the reef mysid Mysidium integrum ingest stained coral mucus. Ingestion rates determined with radioisotope-labeled mucus ranged from 4 to 81% body carbon · 24 h^-1 for the copepods and I to 70% body carbon · 24 h^-1 for the mysids. Incorporation was measured by comparing the organic composition of fecal material and by the incorporation of isotope-labeled mucus. A. tonsa incorporated 47% of ingested ash-free material, 68% of carbon and 36% of nitrogen. M. integrum incorporated 44% of ingested ash-free matter, 57% of carbon and 55% of nitrogen. Incorporation estimates using ¹⁴C-labeled mucus were 65% and 39% for incorporation by A. tonsa and M. integrum respectively. A. tonsa and M. integrum incorporated both the mucus substrate and the epiphytic bacteria of the mucus-detritus.     

Microbial reaction rates and bacterial communities in sediment surrounding burrows of two nereidid polychaetes (Nereis diversicolor and N. virens)

The effects of infaunal mode of life on sediment properties, microbial reaction rates, as well as abundance and composition of bacterial communities were studied in sediment surrounding burrows (mucus lining, oxidised wall, ambient anoxic and surface sediment) of two closely related, but behaviourally different, nereidid polychaete worms: the facultative suspension-feeder Nereis (Hediste) diversicolor and the obligate deposit-feeder Nereis (Neanthes) virens. Burrow sediment of the two species was collected from two adjacent (50 m distance) shallow sandy locations (Kertinge Nor, Denmark). The burrow lining and wall of both polychaete species were enriched in organic matter originating from mucous secretions by the inhabitants and phytoplankton trapped through irrigation. This was more evident for N. diversicolor that shows a significantly higher irrigation rate than N. virens. Both the organic matter mineralisation rates (based on anaerobic incubations) and bacterial abundance were higher along the burrow linings and walls. However, accumulation of porewater TCO₂ and dissolved organic carbon in sediments adjacent to burrows increased most rapidly in the presence of N. diversicolor, suggesting higher heterotrophic activity associated with this species. Surprisingly, bacterial abundance was lower around burrows of N. diversicolor than those from N. virens indicating that burrow environments from the first species harbour a more active bacterial community. Molecular fingerprints of the 16S rRNA gene from bacterial communities showed that the composition of the burrow linings and walls resembled the ambient anoxic sediment rather than the oxic sediment surface. On the other hand, the bacterial fingerprints of the sediment surrounding the burrows of the two polychaete species were markedly different suggesting either a site-specific difference in sediment parameters or a significant species-specific impact of the burrow inhabitants.     

Benthic oxygen uptake and carbon cycling under aphotic and resource-limiting conditions in a submarine cave

To establish relationships between organic input to the benthos and decreases in benthic population biomass and density, benthic oxygen uptake was measured in an oligotrophic submarine cave in the northwestern Mediterranean Sea (Marseille, France), on seven separate occasions in 1987, using an in situ bell-jar respirometer. Oxygen uptake was measured in both the outer twilight section and the dark inner section of the cave during an annual survey (seven recording periods from February 1987 to November 1988). The mean annual benthic oxygen uptake was 80.9 litres O₂ m^–2 yr^–1 for the twilight outer section and 15.5 litres O₂ m^–2 yr^–1 for the dark inner section. The results are discussed and the biogeochemical budget for particulate organic carbon at the sediment-water interface calculated. Respiration rates (expressed as carbon equivalents), together with previously published data on vertical fluxes and burial of organic carbon, revealed that anaerobic pathways accounted for 14% and aerobic pathways for 86% of the total benthic metabolism in the outer part of the cave. In the inner section of the cave, degradation of organic carbon occurred only through aerobic degradation, indicating a strongly carbon-limited ecosystem. The low respiration rates recorded in the dark section were similar to values recorded for some oligotrophic deep-sea environments (1 000 to 2 000 m). Such budgets are essential preliminary steps in order to accurately model benthic metabolic pathways. The determination of annual fluxes linked to the acquisition of long-term data will yield better knowledge of the recycling processes at the water-sediment interface.     

Experimental field study of the effects of crude oil,drill cuttings and natural biodeposits on microphyto- and macrozoobenthic communities in a Mediterranean area

Effects of an experimental pollution by neogenous and fossil organic matter on microphyto- and macrozoobenthic communities were studied in situ over a 1 yr period (July 1989 to May 1990) in a shallow microtidal bay (Gulf of Fos, south coast of France). Three experimental enclosures of 1 m² non-defaunated sediments were covered with 1 cm of polluted defaunated sediments. The sediment in one enclosure contained natural biodeposits with a high organic matter content (BD), that in a second enclosure contained Arabian light crude oil (BAL), and the sediment in the third enclosure contained diesel oil-based cuttings (CUT). Pollution by contaminants did not prevent microphytobenthos from colouizing sediments. Population changes over time were quite similar in all enclosures, except in CUT, where a four times higher chlorophyll a content appeared to be related to a decreased number of grazers and consequently lower grazing rates of animals. Toxicity to the fauna was immediate in the case of BAL and occurred within 3 mo at CUT. Opportunistic species settled in all contaminated sediments; this occurred quite rapidly in BD and BAL which recovered within 3 mo to levels comparable with control sediments. In CUT, natural populations had not recovered after 1 yr, whereas a quasi-monospecific population of Capitella capitata was still present. During the first 3 mo, the oxygen demand of the sediment was higher in oil-contaminated sediments than in controls. On the whole, the changes in fluxes and organism assemblages in our weakly tidal area appear to be consistent with other findings in macrotidal seas.     

Effects of diet,ultraviolet exposure,and gender on the ultraviolet absorbance of fish mucus and ocular structures

Ultraviolet (UV) radiation can be damaging to fish skin and ocular components. Coral reef fishes are regularly exposed to potentially harmful radiation. It was recently discovered that tropical marine fishes possess UV-absorbing compounds in their mucus. This experiment demonstrates significant effects of both diet and ultraviolet exposure on the UV-absorbing compounds in the mucus of a tropical wrasse, Thalassoma duperrey. Fish that are exposed to UV radiation increase the UV absorbance of their mucus only if UV-absorbing compounds are provided in their diet. Fish that are protected from UV radiation decrease the UV absorbance of their mucus regardless of diet. Mucus from female T. duperrey absorbed less UV and females had higher rates of skin damage than males. Females sequester UV-absorbing compounds in their pelagic eggs as well as their epithelial mucus, whereas males do not sequester these compounds in the testes. Spectral transmission through the whole eye was not affected by diet or UV manipulations, but corneal tissue transmission decreased significantly in the UV-exposed individuals. These results demonstrate that coral reef fish can adapt to UV exposure, so long as UV-absorbing compounds are available in the diet.Communicated by P.W. Sammarco, Chauvin     

Exponential Collapse of Benthic Depurative Capacity in A Eutrophic Tropical Marine Ecosystem: A Threshold Response to Organic Nitrogen Loading

Microbial decomposition of particulate organic matter in sediments can substantially modulate eutrophication of nearshore marine environments. Flux budgets for nitrogen compounds and quantification of rates of microbial transformation can provide important information on the process of eutrophication. This survey documents sediment nitrogen budgets for a eutrophic nearshore marine environment at La Parguera on southwest coast of Puerto Rico, including consideration of the organic fraction in addition to microbial transformations.

Sediments of the inshore channel at La Parguera denote the input of heavy organic loads with low redox potentials and high ammonium contents. Benthic fluxes of dissolved organic nitrogen are below those recorded for ammonium. These observations suggest that nearshore tropical sediments subject to heavy nitrogen loads act as a sink for organic nitrogen and that ammonification is a key process in the release of nitrogen from sediments to the water column. Nitrification and denitrification rates are low overall and inversely related to the redox potential. Depuration of excess nitrogen through denitrification is ineffective in these sediments. in contrast to more robust temperate environments, our work demonstrates that tropical marine systems are particularly susceptible to eutrophication given their limited capacity for depuration of excess nitrogen.     

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.     

Mucous sheet formation on poritid corals: An evaluation of coral mucus as a nutrient source on reefs

The production, release, and subsequent consumption of coral mucus on reefs has been portrayed as a potential pathway for the transfer of coral and zooxanthellae production to other reef organisms. However, reported mucus production rates and analyses of nutritional value vary widely. Poritid corals provide a test system to measure mucus production because they produce mucous sheets which can be collected quantitatively. Fluid mucus and mucous sheets were collected fromPorites astreoides, P. furcata, P. divaricata during 1986 and 1987 on reefs in the San Blas Islands, Panama, La Parguera, Puerto Rico and the Florida Keys, USA. Mucus samples were collected from Indo-pacific poritids (P. australiensis, P. lutea, P. lobata, andP. murrayensis) on the Great Barrier Reef during 1985. Biochemical analyses of the fluid mucous secretions, and the derivative mucous sheet, indicate that the mucus is primarily a carbohydrateprotein complex.Porites fluid mucus had a mean caloric content of 4.7 cal mg^–1 ash-free dry weight (AFDW), while mucous sheets contained 3.5 cal mg^–1 AFDW. Sixty-eight percent of the mucous sheet was ash, while fluid mucus was 22% ash. The high ash and low organic contents suggest that mucous sheets have a low nutritional value. C:N ratios varied (range 6.9 to 13.7 for fluid mucus, and 4.8 to 5.9 for mucous sheets), but were similar to typical C:N ratios for marine organisms. Bacterial numbers and chlorophyll a concentrations were higher on mucous sheets than in the surrounding water. Although bacteria aggregate on mucous sheets, bacteria accounted for less than 0.1% of the carbon and nitrogen content of the mucous sheet. Lower C:N ratios in aged mucus, i.e. mucous sheets versus fluid mucus, were attributed to a loss of carbon rather than an increase in nitrogen. Mucous sheet production accounts for a small proportion (< 2% gross photosynthesis) of published values for coral production. In the San Blas Islands, Panama,P. astreoides produced mucous sheets at a rate of 1.5 g C m^–2 y^–1 and 0.3 g N m^–2 y^–1.P. astreoides andP. furcata produced mucous sheets with a lunar periodicity and may provide approximately monthly pulses of carbon and nitrogen to the reef food-web. However, the low annual production rates suggest that mucous sheets make a small contribution to overall energy flow on coral reefs.