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.     

Local and regional genetic connectivity in a Caribbean coral reef fish

Coupled bio-physical models of larval dispersal predict that the Costa Rica–Panama (CR–PAN) reefs should constitute a demographically isolated region in the western Caribbean. We tested the hypothesis that CR–PAN coral reef fish populations would be isolated from Mesoamerican Barrier Reef System (MBRS) populations. To test that, we assessed population genetic structure in bicolor damselfish (Stegastes partitus) from both regions. Adult fish were genotyped from five reefs in CR–PAN and from four reefs along the MBRS at 12 microsatellite loci. Between-region F _ST (F _ST = 0.0030, P < 0.005) and exact test (x ² = 74.34, df = 18, P < 0.0001) results indicated that there is weak but significant genetic differentiation between regions, suggesting some restriction in connectivity along the Central American coastline, as predicted by bio-oceanographic models. Additionally, there is among-site genetic structure in the CR–PAN region, relative to the MBRS and between regions, suggesting higher self-recruitment within CR–PAN. This finding may be explained by differences in habitat characteristics.     

Influence of sediment nitrogen-availability on carbon and nitrogen dynamics in the seagrass Thalassia testudinum

Variations in tissue carbon (C), nitrogen (N) content, and non-structural carbohydrate (NSC) reserves in the turtle grass Thalassia testudinum Banks ex König were examined in relation to changes in sediment-N availability in Corpus Christi Bay (CCB) and lower Laguna Madre (LLM), Texas, USA, from May to October 1997. Under natural conditions, sediment pore-water NH⁺ ₄-concentrations were higher at CCB (100?μM) than at LLM (30?μM); this difference was reflected in a significantly higher leaf and rhizome N-content at CCB than at LLM. However, sediment NH⁺ ₄-enrichment using a commercial fertilizer resulted in significantly higher tissue N-content relative to controls at both sites. N enrichment also influenced plant carbon metabolism, as reflected by distinct increases in leaf C-content at both sites. Significant decreases in rhizome NSC-content was recorded during the first two months of the experiment, suggesting that C was reallocated from rhizomes to leaves to support stimulated leaf growth at both sites. At LLM, leaf growth-rates increased and leaf turnover-time decreased as a result of sediment NH⁺ ₄-enrichment. With respect to chlorophyll, concentrations did not change significantly at CCB, but increased steadily at LLM after the first month following fertilization. In general however, chlorophyll concentrations in control plots were significantly higher at CCB than that at LLM. These observations suggest that leaf function related to C-fixation is enhanced under higher sediment N conditions, as reflected in higher leaf growth-rates and increased blade chlorophyll-content. In contrast, under low-N conditions, below-ground tissue production is enhanced at the expense of the above-ground shoots and leaves, resulting in the high below: above-ground biomass ratios often observed in seagrass beds of oligotrophic environments.     

Resident mysids: community structure,abundance and small-scale distributions in a coral reef lagoon

Seasonal and diel variations in community structure and abundance of coral-reef lagoon mysids were examined at Davies Reef in the central region of the Great Barrier Reef (GBR) between June 1980 and May 1981. Twenty-five mysid species belonging to three subfamilies of the family Mysidae were captured during the study, with six new records for the GBR. The epibenthic mysid community differed from that in the overlying water, was faunistically uniform, but formed characteristic seasonal and diel groupings. The dominant epibenthic species were Erythrops sp., Anisomysis pelewensis, Doxomysis littoralis, A. laticauda, Prionomysis stenolepis, A. lamellicauda, and A. australis, five of which formed schools. Total mysid abundances ranged between 110 and 790 m^-3 with peak abundance in October. Schooling species occurred at local densities of up to 500 000 m^-3. Mysids were absent from shallow and midwater depths during the day, but were distributed throughout all depths at night with peak abundances in mid-water and deep layers. The dominant species in the water column at night were Pseudanchialina inermis, A. laticauda and Gastrosaccus indicus, in descending order of abundance. Lagoonal mysids contribute little to the food of sessile reef planktivores, as all but three species remain concentrated near or on the lagoon floor both day and night. The contribution of resident lagoon mysids to reef trophodynamics is probably through remineralization of lagoon detritus. Given the vast reef areas comprised of sandy lagoons, the large populations and relatively large size of lagoon mysids, this trophodynamic role may be of considerable importance.A.I.M.S. Contribution No. 477     

Thalassia testudinum productivity and grazing by green turtles in a highly disturbed seagrass bed

There has been an historical decline in the seagrass beds in Maho and Francis Bays, St. John, U.S. Virgin Islands: presently (1986) there are only five small seagrass beds in shallows water. These seagrass beds are highly disturbed by heavy boat usage and are intensively grazed by the green turtle Chelonia mydas L. Fifteen to 50 boats anchor each night in the bays: anchor scars cause a loss of up to 6.5 m² d^-1 or 1.8% yr^-1 of the seagrass beds. Seagrasses regrew into such scars only minimally within a period of 7 mo. The size of the green turtle population was estimated at 50 subadults and their feeding behavior was determined by direct observation and radiotelemetry. The behavior of the green turtles differed from other observations published on the species. Here, the turtles grazed all available Thalassia testudinum, their preferred seagrass food, rather than creating discrete grazing scars, and spent all their waking hours (9 h per day) feeding. Areal productivity of T. testudinum leaves (33 to 97 mg dry wt m^-2d^-1) in the bays was at least an order of magnitude lower than published values or than the productivity of another, lessdisturbed seagrass bed on St. John, despite having comparable leaf-shoot density. Leaf shoots were stunted, fragile, achlorotic, and had only two leaves as opposed to the five leaves per shoot more typically seen. The green turtle population was near the estimated carrying capacity of T. testudinum, based on the standing crop and productivity of T. testudinum and the grazing rate of the turtles. The effect of disturbance of T. testudinum from boats and turtles was assessed by excluding these with emergent fences. Within 3 mo of protection, the areal and shoot-specific productivity of T. testudinum leaves as well as leaf size increased significantly compared to unprotected areas. Conservation efforts are recommended in Maho Bays and Francis because seagrass productivity is low, disturbance rates are higher than recovery rates, the turtles cannot increase further their feeding rate in order to compensate for such factors, and there are few alternate sources of T. testudinum on the north shore of St. John.Contribution No. 175 from West Indies Laboratory, Teague Bay, Christiansted, St. Croix, U.S. Virgin Islands 00820, USA     

Nitrogen fixation (acetylene reduction) in the phyllosphere of Thalassia testudinum

N₂ fixation (C₂H₂ reduction) associated with the leaves of the sea grass Thalassia testudinum was investigated at 5 sites in South Florida (Biscayne Bay) and one site in the Bahamas (Bimini Harbor). Significant activities were correlated with the occurrence of a heterocystous blue-green alga (Calothrix sp.) on the leaves. C₂H₂ reduction was not stimulated by organic compounds, either aerobically or anaerobically in the light or dark. Therefore, other physiological types of microbes were not important in N₂ fixation. Diurnal and seasonal variations in N₂ fixation occurred, with maximal rates during the daytime and in the late spring and early summer. N₂ fixation was negligible at four stations in Biscayne Bay. At the fifth station, near Fowey Rock, about 5 kg N ha^-1 year^-1 was fixed. In the summer, the N₂ fixed per day (4–5 mg N m^-2) could provide 4 to 23% of the foliar productivity demands of T. testudinum at this site and the station in Bimini Harbor. N₂ fixation at the periphery of a sea-grass patch, near Fowey Rock, could provide 8 to 38% of the daily nitrogen requirement for leaf production, and thereby might compensate for a less effective trapping and recycling of nitrogen from dead leaves in such regions.     

Comparative species richness,composition, and abundance of invertebrates in Caribbean seagrass (Thalassia testudinum) meadows (Panamá)

The results of a year-long study in which epibenthic invertebrates were collected monthly from seagrass (Thalassia testudinum) meadows along the Caribbean coast of Panamá and the Panamá Canal Zone are described in this paper. Differences in species composition and abundance among sites were primarily due to the proximity of surrounding habitats, especially coral reefs, which contain a number of species that utilize the seagrass meadows. In contrast to many previous characterizations of tropical marine habitats, important seasonal fluctuations in both species number and abundance took place at each of the sites. Data on breeding activity among several species of decapod crustaceans indicate year-round reproduction, although considerable seasonal differences occur in the percentage of ovigerous females. These interspecific differences in observed reproductive output may be explained by differences in life-cycle length, a factor not often considered in discussions of seasonal breeding patterns in tropical marine invertebrates. Overall species composition was qualitatively similar to that reported in comparable studies of tropical and subtropical seagrass meadows elsewhere, although caridean shrimp and xanthid orab species were reduced in number and total abundance were much lower than in previous studies.     

Reef-basel herbivores and the distribution of two seagrasses (Syringodium filiforme and Thalassia testudinum) in the San Blas Islands (Western Caribbean)

Two seagrasses, manatee grass (Syringodium filiforme) and turtle grass (Thalassia testudinum), predominated in the areas bordering Ukkup Tupo, San Blas Islands, Western Caribbean. These seagrasses occupied the following three concentric zones extending outward from the reef: a near-reef turtle grass zone, an intermediate manatee grass zone and an off-reef turtle grass zone. Feeding experiments between January and March 1980 indicate that the absence of manatee grass close to the reef resulted from grazing by reef-associated herbivores, mainly day-active fishes and night-active sea urchins (Diadema antillarum). Grazing on manatee grass by fishes was approximately six times greater than grazing by sea urchins; thus, it appears that herbivorous fishes restrict the near-reef distribution of manatee grass at the study area. Where grazing was heaviest, the inner boundary of the manatee grass zone was farthest from the reef. The volume of manatee grass grazed during experiments was five times the volume of turtle grass consumed, strongly suggesting that the former species is a preferred food item. This is the first evidence for selective grazing on seagrasses.     

Modelling the dynamics of coral reef macroalgae using a Bayesian belief network approach

Macroalgae are a major benthic component of coral reefs and their dynamics influence the resilience of coral reefs to disturbance. However, the relative importance of physical and ecological processes in driving macroalgal dynamics is poorly understood. Here we develop a Bayesian belief network (BBN) model to integrate many of these processes and predict the growth of coral reef macroalgae. Bayesian belief networks use probabilistic relationships rather than deterministic rules to quantify the cause and effect assumptions. The model was developed using both new empirical data and quantified relationships elicited from previous studies. We demonstrate the efficacy of the BBN to predict the dynamics of a common Caribbean macroalgal genus Dictyota. Predictions of the model have an average accuracy of 55% (implying that 55% of the predicted categories of Dictyota cover were assigned to the correct class). Sensitivity analysis suggested that macroalgal dynamics were primarily driven by top–down processes of grazing rather than bottom–up nutrification. BBNs provide a useful framework for modelling complex systems, identifying gaps in our scientific understanding and communicating the complexities of the associated uncertainties in an explicit manner to stakeholders. We anticipate that accuracies will improve as new data are added to the model.     

Molecular genetic evidence for probable reticulate speciation in the coral genus Madracis from a Caribbean fringing reef slope

For many corals, the existence of morphologically distinct yet sympatric populations/species implies reproductive isolation. Conversely, the presence of many intermediate and overlapping morphologies combined with synchronous, mass spawning suggests incomplete reproductive isolation. In Madracis (Scleractinia: Astrocoeniina: Pocilloporidae), high levels of morphological plasticity among the five most commonly recognized species (M. mirabilis, M. senaria, M. decactis, M. formosa and M. pharensis) on Caribbean reefs led us to question species boundaries. Phylogenetic relationships were investigated at the intra-individual, inter-individual and inter-specific levels using the ITS1-5.8S-ITS2 region (ca. 613 bp) of the ribosomal DNA cistron. Inter-specific divergence was ca. 6%, while intra-individual and intra-specific divergences ranged from 0% to 4.9% and 3.3% to 3.5%, respectively. M. senaria and M. mirabilis formed monophyletic groups. M. formosa, M. decactis and M. pharensis formed a paraphyletic complex. High levels of intra-individual and intra-specific ITS polymorphism in the decactis-formosa-pharensis cluster may be the result of very recent speciation within the clade (i.e. maintenance of ancestral polymorphism and incomplete lineage sorting), or the result of repeated introgressive hybridization among the three taxa. Polymorphism parsimony of 89 sites, including nine that showed additivity, revealed a phylogenetic topology more consistent with inter-taxal hybridization. Results are discussed in terms of weak reproductive barriers, and phylogenetic fission and fusion under Veron's model of reticulate speciation in corals. Ecological studies involving Madracis should consider M. decactis, M. formosa and M. pharensis as a complex.     

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.     

Significance of pelagic bacteria as a trophic resource in a coral reef lagoon,One Tree Island,Great Barrier Reef

Bacterial numbers and frequency of dividing cells (FDC) were estimated in surface waters at seven stations along a transeet across One Tree lagoon and reefs at 2 h intervals over one tidal cycle in May 1984 and once a day on 8 d in December 1984/January 1985. Cell numbers ranged between 1.24 and 13.10x10⁵ ml^-1, with an overall mean value of 4.81x10⁵. There was a progressive depletion in standing stock from windward to leeward stations across the lagoon, also reflected in the FDC, which generally showed similar dynamic patterns to cell numbers throughout the study. Predator-free seawater (filtered through 2 m pore-size filters) and mucus-enriched incubations were also used to establish the relationship between the growth rate () and FDC for coral-reef populations. Average growth rates predicted from FDC values ranged from 0.062 to 0.174 h^-1, which is equivalent to doubling times of 5.74 to 16.00 h, with an overall mean value of 11.5 h. These fast doubling times suggest that bacteria respond rapidly to pulses of enrichment as they float over the lagoon and reefs, characteristically achieving about one doubling per tidal cycle. This probably ensures that the bulk of labile organic matter and dissolved nutrients is conserved within the immediate coral-reef environment. Estimates of bacterial production also suggest that they may contribute up to 40% of total picoplankton production and about 25% of total microplankton production. Pelagic bacteria are therefore potentially a major food resource for benthic filter-feeders, especially for those mainly dependent on the smaller (pico) components of the plankton.     

Evidence of limited gene flow in three species of coral reef fishes in the lagoon of New Caledonia

Allozyme variation was used to investigate the genetic structure of the coral reef fishes Stegastes nigricans, Epinephelus merra and Acanthurus triostegus around New Caledonia. Each species was sampled from each of three sites in the ≃1000 km circumference of the lagoon of New Caledonia. Allelic variation was recorded for each species at 14, 13 and 17 loci, respectively, and heterozygosity diversity (H s) was 0.082, 0.065 and 0.116, respectively. Analysis of genetic differentiation between sites produced inconsistent results between species, with spatial heterogeneity in two species (S. nigricans, F st  = 0.038; A. triostegus, F st = 0.049) and homogeneity in one species (E. merra, F st = 0.000). Hydrological and climatic data from the lagoon suggest that the eastern and western sides of the lagoon are isolated, since they lie in water masses of different origin. This may explain the genetic differentiation and restricted gene flow found at a local scale for S. nigricans and A. triostegus. Homogeneity in populations of E. merra is discussed in relation to its low genetic diversity and its reproductive behaviour. Received: 23 April 1997 / Accepted: 25 September 1997     

The costs and benefits of territory sharing for the Caribbean coral reef fish,Scarus iserti

Summary Along the Caribbean coast of Panama, groups of unrelated female striped parrotfish, Scarus iserti, co-defend a common feeding territory. Field manipulations of group size and composition were performed to examine the benefits and costs accrued by dominant fish within these shared territories. Dominant fish benefit from the presence of relatively large subordinates because they share in the defense of the territory. Removals of these fish caused increases in defense time and decreases in feeding time for dominant group members. Dominants benefit from the presence of small subordinates because they increase the foraging efficiency of dominants. Removals of smaller subordinates caused reductions in the feeding time of dominant fish, although no changes in defense time occurred. Concurrently, dominant fish reduce costs of resource depletion by displacing subordinate group members from good food patches. Dominance interactions within a group reduce the amount of time subordinates spend feeding (subordinate individuals fed at higher rates following the removal of a dominant) and limit a subordinate's access to high quality resources. This combination of benefits and reduced costs ultimately contributes to the economic defensibility of a striped parrotfish territory and has led to the evolution of group territorial behavior in the absence of kin selection and cooperative parental care.     

Life history characteristics of a coral reef sponge

Study of the life history characteristics of the common Red Sea sponge Mycale fistulifera (Poecilosclerida: Demospongiae) reveals several traits which may categorize it as an opportunistic species: (1) at least part of the population is reproductively active throughout the year, providing a continuous supply of larvae for settlement; (2) sponges may produce and release a large number (152±39 larvae d^-1) of brooded larvae; (3) released larvae are capable of fast settlement and metamorphosis (minutes to 30 hours post-release); (4) the turnover of the population is high and population size varies with time. From 48 initial sponges plus 94 recruits, only 25 remained alive after 14 months of observation; (5) most members of the population (>70%) have a small body size (<30 cm²); (6) sexual maturity may occur at an early age. These traits facilitate continuous establishment of M. fistulifera in new spaces on the reef; (7) M. fistulifera, preferred substrate is another opportunistic species, the coral Stylophora pistillata which it overgrows.     

Diurnal variation in chlorophyll fluorescence of Thalassia testudinum seedlings in response to controlled salinity and light conditions

Diurnal variability in chlorophyll fluorescence caused by dynamic irradiance conditions is an important issue when using pulse amplitude modulation fluorometry to measure physiological conditions of plants at the landscape scale. We examined the use of slopes and y-intercepts of diurnal effective photochemical efficiency of photosystem II (PSII) (ΔF/F _m′) versus photosynthetically active radiation (PAR) regressions in addition to direct measurements of maximum photochemical efficiencies of PSII (F _v/F _m) values to assess physiological status of Thalassia testudinum seedlings in a controlled mesocosm study. Seedlings were exposed to two light treatments (full sun and 50–70 % light reduction) and three salinity treatments (20, 35, and 50). Measurements were taken at 0600, 0900, 1200, 1500, 1800, and 2100 hours in order to assess the diurnal variation in photochemical efficiency of PSII and PAR, with measurements at 2100 providing F _v/F _m. Results indicated significant effects of light and salinity on regression y-intercepts and measured F _v/F _m values. Shaded seedlings had higher values for both parameters, suggesting low-light acclimation. The highest salinity treatment resulted in significant reductions for both parameters, suggesting stress. Stress was also indicated by significant reductions in both seedling leaf growth and mean differences between seedling leaves and media osmolalities in the hypersaline treatments (152.0 ± 26.4 vs. 630 ± 40.2 mmol kg^?1 for the control treatments). Slopes of ΔF/F _m′ versus PAR significantly differed with varying light treatments, with full sun seedlings exhibiting shallower slopes than shaded seedlings, indicating higher efficiency of dissipation of excess energy (photoprotection). These experimental results confirm field data suggesting that diurnal ΔF/F _m′ versus PAR regressions are responsive to changes in the physiological status of T. testudinum and that the y-intercepts of diurnal regressions may be used as a proxy for F _v/F _m.