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.     

A new apparatus for force measurement in marine bioadhesion

In an experimental study on the effect of parrotfish (probably Scarus taeniurus) grazing on the structure of benthic reef communities, fishes in densities of 0.6 to 1.5 parrotfish per m² or 9 to 17 g wet weight of fish per m² of feeding surface were found to have an optimum effect, resulting in the greatest benthic species richness and biomass on 2-dimensional surfaces. The presence of refuges (3-dimensional habitats), however, has a greater impact on bemthic community structure (number of species and biomass) than does just the density of parrotfish in such an experimental system. Coral recruitment is enhanced by the presence of refuges and, like coralline algae, is more successful under increased grazing pressure. These optimum densities of parrotfishes relate well to observed field densities where, in a collection from a Hawaiian patch reef, there were 1.1 fish or 10.8 g wet weight of parrotfish per square meter of collection area. The success of coralline algae and corals under high grazing pressure may have important consequences for the stability and structure of modern coral reefs.     

Population structure and feeding deterrence in three shallow-water antarctic soft corals

Alcyonium paessleri and Clavularia frankliniana are numerically abundant soft corals in the nearshore (12 to 33 m depth) benthic communities of eastern McMurdo Sound. They are much less abundant in western McMurdo Sound where a third species, Gersemia antarctica, co-occurs in low numbers. The body tissues of these three species are comprised mainly of organic material (53 to 70% dry wt), which is primarily dervied from NaOH-soluble protein and refractory material. The energetic contents of the whole-body tissues of A. paessleri, C. frankliniana and G. antarctica are 15.9, 17.3, and 14.5 kJ g^-1 dry wt, respectively. The mean biomass per individual is 1.81, 0.008, and 45 g dry wt for each respective species. Based on population densities of 7.3, 1337.3, and 0.04 soft corals m^-2 for A. paessleri, C. frankliniana and G. antarctica, respectively, the population energetic densities are estimated to be 210.1, 185.1, and 26.1 kJ m^-2. Despite the relatively rich energetic content of the tissue and apparent vulnerability to predators, very little predation occurs on these soft corals. Two potential predators, the antarctic sea stars Perknaster fuscus and Odontaster validus, exhibited significant chemotactic defensive tube-foot retractions to hexane, chloroform, methanol, and aqueous methanol extracts of each soft coral. In addition, wholebody tissue of each soft coral was rejected by the demersal fish Pseudotrematomus bernacchii and the cryopelagic fish Pagothenia borchgrevinki. In contrast, whole soft-coral tissues sequentially extracted in four increasingly polar solvents were readily ingested by these antarctic fishes, indicating that sclerites do not play a significant role in deterring predators. Our results indicate that these antarctic soft corals contain bioactive compounds which deter common predatory seastars and fishes.     

Larvae of a colonial ascidian use a non-contact mode of substratum selection on a coral reef

The rate at which larvae successfully recruit into communities of marine benthic invertebrates is partially dependent upon how well larvae avoid benthic predators and settle on appropriate substrata. Therefore, to be able to predict recruitment success, information is needed on how larvae search for settlement sites, whether larvae preferentially settle on certain substrata, and the extent to which there are adequate cues for larvae to find these substrata. This article describes how larvae of the colonial ascidian Diplosoma similis find settlement sites on a coral reef. Direct field observations of larval settlement were made on a fringing reef in Kaneohe Bay, Oahu, Hawaii, between September 1985 and April 1986. A comparison of the substrata that larvae contacted prior to settlement relative to the percentage cover of these substrata on the study reef suggests that larvae are using a non-contact mode of substratum identification to locate suitable settlement sites. This mode of substratum identification allowed 74% of larvae to evade predation by benthic organisms who would otherwise have eaten larvae if they had been contacted. Of those larvae that evaded predation, 88% subsequently settled on the same two substrata upon which most adults are found (dead coral or the green alga Dictyosphaeria cavernosa). This pattern of settlement was probably a result of active selection, since the two substrata cover only 14.4% of the reef's surface and currents had little effect on the direction in which larvae swam. An important contributing factor to the high success rate of larval settlement on suitable substrata was the lack of any temporal decay in substratum preference. It is concluded that for Diplosoma similis larval supply is a sufficient predictor of larval settlement rate. However, for marine invertebrates whose larvae are passively dispersed and exhibit a greater temporal decay in substratum preference, larval settlement should generally have a greater dependency on spatial variation in the abundance of benthic predators and suitable substrata.     

Reproductive cycles of tropical sea cucumbers (Echinodermata: Holothuroidea) in southern Taiwan

Reproductive cycles of Afrocucumis africana (Dendrochirotida), Actinopyga echinites, Holothuria leucospilota, H. cinerascens, H. difficilis (Aspidochirotida), Synaptamaculata, Opheodesoma grisea, Patinapta taiwaniensis and Polycheira rufescens (Apodida), representing three orders of intertidal holothurians in southern Taiwan, were determined by gonad index and histological examination from March 1990 through September 1991. All nine species of holothurians have annual spawning periods lasting 2 to 4 mo in spring or summer. The early spawning of H. cinerascens and P. rufescens (suspension feeders) suggests that their feeding mode is related to food resources and may influence the reproductive period. In the direct-developing dendrochirotid (suspension feeder), gonad development occurs in cool months, and gametes are released at the beginning of warm months. In indirect-developing aspidochirotid and apodid (deposit feeder), gamete release occurs in late spring or summer and appears to be correlated with summer phytoplankton growth. The holothurians in the present study display spawning periodicity and duration similar to those of temperate species. Four species of holothurians lengthened spawning season with decreased latitude.     

Variation in benthic communities of eastern Caribbean coral reefs in relation to surface sediment composition

The effects of sedimentation on coral reefs are commonly studied at local scales, but larger-scale patterns have been elusive, making it difficult to determine the role of sedimentation in region-wide changes in these ecosystems. We examined the relationships between characteristics of reef-associated surface sediment and benthic composition of 22 reefs around 11 islands of the eastern Caribbean. The terrigenous fraction in surface sediment increased with proximity to a clear source of sediment input. The percent cover of live coral, macroalgae, and turf algae decreased with higher terrigenous sediment fraction, while sponge cover increased. Sites with sediment containing high and low terrigenous fraction differed in coral species assemblages. In particular, the cover of Montastraea annularis complex decreased with increasing terrigenous sediment fraction. The proportion of fine-grained sediment had no effect on benthic composition. These results suggest that sedimentation may play a role in shaping coral reef communities at a regional scale.     

Ultraviolet photosensitivity and feeding in larval and juvenile coral reef fishes

The ability of young coral reef fishes to feed using solely ultraviolet-A (UV-A) radiation during ontogeny was examined using natural prey in experimental tanks. Larvae and juveniles of three coral reef fish species (Pomacentrus amboinensis, Premnas biaculeatus and Apogon compressus) are able to feed successfully using UV-A radiation alone during the later half of the pelagic larval phase. The minimum UV radiation intensities required for larval feeding occur in the field down to depths of 90–130 m in oceanic waters and 15–20 m in turbid inshore waters. There was no abrupt change in UV sensitivity after settlement, indicating that UV photosensitivity may continue to play a significant role in benthic juveniles on coral reefs. Tests of UV sensitivity in the field using light traps indicate that larval and juvenile stages of 16 coral reef fish families are able to detect and respond photopositively to UV wavelengths. These include representatives from families that are unlikely to possess UV sensitivity as adults due to the UV transmission characteristics of the ocular media. Functional UV sensitivity may be more widespread in young coral reef fishes than in the adults, and may play a significant role in detecting zooplanktonic prey.     

Biology of small juveniles of the tropical holothurian Actinopyga echinites: growth,mortality, and habitat preferences

Several morphological, physiological and ecological experiments on the general biology of small juveniles (drained body weight=0.09 to 17.34 g) of Actinopyga echinites (Jäger 1833) (Echinodermata, Holothuroidea) were conducted between August 1991 and July 1992 at Bise reef flat, Okinawa, southern Japan. Supplementary experiments were carried out at the laboratory. The experiments were designed with a view to potential stock enhancement projects of A. echinites and other commercially exploited tropical sea cucumbers of the coral reef zone. During the enclosure experiments, the average percentage of drained body weight to fresh body weight of the juveniles was 48.3%. The specimens displayed a growth rate of 1500% during the 11-mo period. Their drained body weight increased from 0.87 to 12.82 g. The juveniles' internal and skeletal morphology differed considerably from the morphology of adult A. echinites. Two new types of skeletal spicules were discovered. Individual growth of all spicule types monitored differed notably, and shrinkage was observed for the two newly discovered types. Relative frequency of the spicule types within the skeletons changed with increasing body weights of the individuals. The juveniles of A. echinites displayed a strong habitat preference for plate-like substrate types such as eroded limestone or dead coral plates. Skeletons of Acropora spp. were accepted at a much lower rate. Weight-frequency distributions of the specimens on varying substrate types and between various times of the day showed significant differences. Cryptic behaviour of the juveniles was observed at all times of the day. Natural mortality of the holothurians, excluding predation and minor dislodgement effects, was low at 0.6% mo^–1. When predation effects were introduced to the experiments, mortality reached a rate of 0.3% mo^–1. The average percentage of predation effects within the natural mortality total was 76.8%. Juvenile A. echinites exhibited a maximum short-distance migration speed of 9 cm h^–1. This was two orders of magnitude slower than the speed observed in adults (900 cm h^–1) when differences in total body length were taken into consideration. The holothurians had two activity peaks during the 4-h periods monitored, around sunrise and sunset, respectively.     

Temporal genetic variation in subpopulations of bicolor damselfish (Stegastes partitus) inhabiting coral reefs in the Florida Keys

In 1986 we observed significant genetic heterogeneity among samples of bicolor damselfish (Stegastes partitus) from geographically proximate coral reefs in the Florida Keys. This result was in contrast to results of virtually all previous studies of coral reef fishes. To assess temporal stability of localized genetic differentiation, we resampled reefs in 1989. Within approximately two generations, allele frequencies at the most differentiated locus,ACO-1 ^*, had changed by as much as 0.65, resulting in almost complete homogeneity between previously differentiated subpopulations. Estimates of 10.78 migrants per generation suggested that high gene flow is the most likely factor responsible for the significant change in allele frequencies. We attribute the original genetic differentiation to a population bottleneck, possibly caused by environmental perturbations such as hurricanes and consequent genetic drift. For reef fishes with pelagic egg and/or larval stages, a growing body of data suggests that: (1) populations are geographically extensive gene pools, and (2) rates of dispersal appear to be high enough to allow for continual repopulation of isolated and/or perturbed coral reef communities.     

Dynamics of parrotfish grazing scars

Parrotfishes exhibit a range of feeding modes. These species vary in both feeding morphology and behaviour, but the vast majority of species leave distinctive scars on the substratum when feeding. Although the role of parrotfishes in reef resilience is well documented, the basis of this role and the effect of their grazing scars on the benthic community structure remain unclear. This study evaluated the dynamics of grazing scars of large adult Scarus rivulatus and Chlorurus microrhinos on an inshore reef in the Great Barrier Reef (GBR). These species represent the most abundant scraping and excavating parrotfish species on inshore reefs. Grazing scars of each species were marked, measured and observed for seven consecutive days. S. rivulatus grazing scars were smaller in area and volume and more rapidly reoccupied by algae than those of C. microrhinos. However, because of the higher abundance and feeding frequency of S. rivulatus at the study site, this species had higher algal removal rates than C. microrhinos. These species appear to play distinctly different functional roles in shaping the benthic community of inshore GBRs. S. rivulatus is primarily responsible for algal dynamics dominated by vegetative regrowth. In contrast, C. microrhinos opens relatively large areas which remain clear for several days. These scars may represent settlement sites which are relatively free from algae and sediment. This study provides new information on the differences between scraping and excavating parrotfishes and, in a system with just one abundant large excavating species, emphasizes the potential for low functional redundancy in high diversity coral reef systems.     

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.     

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.     

Video-monitored predation by Caribbean reef fishes on an array of mangrove and reef sponges

Although predation by fishes is thought to structure benthic invertebrate communities on coral reefs, evidence to support this claim has been difficult to obtain. We deployed an array of eight sponge species on Conch Reef (16 m depth) off Key Largo, Florida, USA, and used a remote video-camera to record fish activity near the array continuously during five daylight periods (6 h for 1 d, at least 11.5 h for 4 d) and one night period (11 h). Of the eight sponge species, four were from adjacent reefs (Agelas wiedenmayeri, Geodia neptuni, Aplysina fistularis, and Pseudaxinella lunaecharta), and four were from a nearby mangrove habitat (Chondrosia collectrix, Geodia gibberosa, Halichondria sp., andTedania ignis). Each species of reef sponge was chosen to match the corresponding mangrove species in form and color (black, brown, yellow, and red, respectively). Predation events only occurred during daylight hours. Tallies of the number of times fishes bit sponges revealed intense feeding by the expected species of sponge-eating fishes, such as the angelfishHolacanthus bermudensis, H. tricolor, andPomacanthus arcuatus, the cowfishLactophrys quadricornis, and the filefishCantherhines pullus, but surprisingly also by the parrotfishSparisoma aurofrenatum andS. chrysopterum. Of 35 301 bites recorded, 50.8% were taken by angelfish, 34.8% by parrotfish, and 13.7% by trunkfish and filefish. Mangrove sponges were preferred by all reef fishes; 96% of bites were taken from mangrove species, with angelfish preferringChondrosia collectrix and parrotfish preferringGeodia gibberosa. Fishes often bit the same sponge repetitively, and frequently consumed entire samples within 30 min of their deployment. Sponge color did not influence fish feeding. Two of the four mangrove sponge-species deployed on the array were also found living in cryptic habitats on adjacent reefs and were rapidly consumed by fishes when exposed. Our results demonstrate the importance of fish predation in controlling the distribution of sponges on Caribbean reefs.     

Extreme genetic diversity and temporal rather than spatial partitioning in a widely distributed coral reef fish

Mitochondrial control region (HVR-1) sequences were used to identify patterns of genetic structure and diversity in Naso vlamingii, a widespread coral reef fish with a long evolutionary history. We examined 113 individuals from eight locations across the Indo-Pacific Ocean. Our aims were to determine the spatial scale at which population partitioning occurred and then to evaluate the extent to which either vicariance and/or dispersal events have shaped the population structure of N. vlamingii. The analysis produced several unexpected findings. Firstly, the genetic structure of this species was temporal rather than spatial. Secondly, there was no evidence of a barrier to dispersal throughout the vast distribution range. Apparently larvae of this species traverse vicariance barriers that inhibit inter-oceanic migration of other widespread reef fish taxa. Thirdly, an unusual life history and long evolutionary history was associated with a population structure that was unique amongst coral reef fishes in terms of the magnitude and pattern of genetic diversity (haplotype diversity, h = 1.0 and nucleotide diversity π = 13.6%). In addition to these unique characteristics, there was no evidence of isolation by distance (r = 0.458, R ² = 0.210, P = 0.078) as has also been shown for some other widespread reef species. However, some reductions in gene flow were observed among and within Ocean basins [Indian–Pacific analysis of molecular variance (AMOVA), Φ _st = 0.0766, P < 0.05; West Indian–East Indian–Pacific AMOVA Φ _st = 0.079, P < 0.05]. These findings are contrasted with recent studies of coral reef fishes that imply a greater degree of spatial structuring in coral reef fish populations than would be expected from the dispersive nature of their life cycles. We conclude that increased taxon sampling of coral reef fishes for phylogeographic analysis will provide an extended view of the ecological and evolutionary processes shaping coral reef fish diversity at both ends of the life history spectrum.     

Coral Reef Habitats as Surrogates of Species, Ecological Functions, and Ecosystem Services

Abstract: Habitat maps are often the core spatially consistent data set on which marine reserve networks are designed, but their efficacy as surrogates for species richness and applicability to other conservation measures is poorly understood. Combining an analysis of field survey data, literature review, and expert assessment by a multidisciplinary working group, we examined the degree to which Caribbean coastal habitats provide useful planning information on 4 conservation measures: species richness, the ecological functions of fish species, ecosystem processes, and ecosystem services. Approximately one‐quarter to one‐third of benthic invertebrate species and fish species (disaggregated by life phase; hereafter fish species) occurred in a single habitat, and Montastraea‐dominated forereefs consistently had the highest richness of all species, processes, and services. All 11 habitats were needed to represent all 277 fish species in the seascape, although reducing the conservation target to 95% of species approximately halved the number of habitats required to ensure representation. Species accumulation indices (SAIs) were used to compare the efficacy of surrogates and revealed that fish species were a more appropriate surrogate of benthic species (SAI = 71%) than benthic species were for fishes (SAI = 42%). Species of reef fishes were also distributed more widely across the seascape than invertebrates and therefore their use as a surrogate simultaneously included mangroves, sea grass, and coral reef habitats. Functional classes of fishes served as effective surrogates of fish and benthic species which, given their ease to survey, makes them a particularly useful measure for conservation planning. Ecosystem processes and services exhibited great redundancy among habitats and were ineffective as surrogates of species. Therefore, processes and services in this case were generally unsuitable for a complementarity‐based approach to reserve design. In contrast, the representation of species or functional classes ensured inclusion of all processes and services in the reserve network.     

Microbial RNA and DNA synthesis in marine sediments

A technique for measuring rates of RNA and DNA synthesis in sedimentary microbial communities has been adapted from methods developed for marine and freshwater microplankton research. The procedure measures the uptake, incorporation and turnover of exogenous [2, ³H]-adenine by benthic microbial populations. With minor modification, it is applicable to a wide range of sediment types. Measurement of nucleic acid synthesis rates are reported from selected benthic marine environments, including coral reef sediments (Kaneohe Bay, Oahu, Hawaii), intertidal beach sands (Oahu and southern California) and California borderland basin sediment (San Pedro Basin), and comparisons are made to selected water-column microbial communities. Biomass-specific rates of nucleic acid synthesis in sediment microbial communities were comparable to those observed in water-column assemblages (i.e., 0.02 to 2.0 pmol deoxyadenine incorporated into DNA [ng ATP]^-1 h^-1 and 0.2 to 8.9 pmol adenine incorporated into RNA [ng ATP]^-1 h^-1). DNA synthesis rates were used to calculate carbon production estimates ranging from 2 g C cm^-3 h^-1 in San Pedro Basin sediment (880 m water depth) to 807 g C cm^-3 h^-1 in coral reef sediment from the Kaneohe Bay. Microbial community specific growth rate, (d^-1), estimated from DNA synthesis rates in surface sediments ranged from 0.1 in San Pedro Basin to 4.2 in Scripps Beach (La Jolla, California) intertidal sand.     

Exploited species impacts on trophic linkages along reef-seagrass interfaces in the Florida Keys

The removal of fish biomass by extensive commercial and recreational fishing has been hypothesized to drastically alter the strength of trophic linkages among adjacent habitats. We evaluated the effects of removing predatory fishes on trophic transfers between coral reefs and adjacent seagrass meadows by comparing fish community structure, grazing intensity, and invertebrate predation potential in predator-rich no-take sites and nearby predator-poor fished sites in the Florida Keys (USA). Exploited fishes were more abundant at the no-take sites than at the fished sites. Most of the exploited fishes were either omnivores or invertivores. More piscivores were recorded at no-take sites, but most (approximately 95%) were moderately fished and unexploited species (barracuda and bar jacks, respectively). Impacts of these consumers on lower trophic levels were modest. Herbivorous and smaller prey fish (< 10 cm total length) densities and seagrass grazing diminished with distance from reefs and were not negatively impacted by the elevated densities of exploited fishes at no-take sites. Predation by reef fishes on most tethered invertebrates was high, but exploited species impacts varied with prey type. The results of the study show that, even though abundances of reef-associated fishes have been reduced at fished sites, there is little evidence that this has produced cascading trophic effects or interrupted cross-habitat energy exchanges between coral reefs and seagrasses.     

Quantitative distribution of herbivorous reef fishes in the Gulf of Aqaba (Red Sea)

The distribution of the main herbivorous fishes (Acanthuridae, Scaridae, Siganidae) was studied across a coral reef of the Jordanian coast in the Gulf of Aqaba (Red Sea). Visual counts were realized by diving along transects (200 m long and 5 m wide), parallel to the shore, at 10 stations located from the lagoon to 40 m deep on the outer reef slope. Herbivorous reef fishes reach their highest abundance on the reef front, where 234 fishes were counted per 1,000 m². Their density decreases on the reef flat, with an average of 150 fish 1,000 m^-2, and is lowest on the outer reef slope (69 fish 1,000 m^-2). Surgeonfishes form 63% of the herbivorous ichthyofauna, parrotfishes 35%, and rabbitfishes 2%. Families and species display different distributions according to biota. The Acanthuridae dominate on the reef flat, whereas the Scaridae are more numerous on the outer reef slope. The evolution of the social structure of the main species was observed: the adults generally school in the lagoon and on the reef flat, but are mainly solitary on the reef slope. The distribution of juvenile individuals is more restricted: they are concentrated on the reef front and on the upper part of the reef slope.This study is part of a cooperation programme between the University of Nice (France) and the University of Jordan, to study the ecology of the coral reefs and the surrounding waters of the Jordanian coast (Gulf of Aqaba, Red Sea)     

Differences in the structures of fish assemblages inThalassia testudinum beds in Guadeloupe,French West Indies,and their ecological significance

The structures of fish assemblages in twoThalassia testudinum beds in Guadeloupe, French West Indies, one adjacent to mangroves and the other adjacent to coral reefs, were compared between January 1983 and May 1984. The aim of the study was to compare the influences of mangroves and coral reefs on the utilization of seagrass beds by fishes through examination of species composition, catch rate, size of fishes and temporal changes. The two fish assemblages were similar in terms of the number of species they had in common (nearly 44% of the total number of species collected) and the great abundance of juveniles. They both comprised species that usually inhabit other habitats, i.e., estuaries, open waters or coral reefs. Estuary-associated species (e.g. Gerreidae) were the most abundant species in the seagrass bed near the mangroves, while small pelagic species (e.g. Clupeidae) were the most abundant species in the seagrass bed near the coral reefs. The seagrass bed near the mangroves was preferentially utilized as a nursery area by small juveniles of various species (e.g. Clupeidae, Sparidae, Gerreidae, and at least one coral reef species,Ocyurus chrysurus). The abundance of these species varied frequently, suggesting successive arrivals and departures of juveniles over time. The seagrass bed near the coral reefs was characteristically utilized by fishes that are more able to avoid predation, i.e., fishes that forage over seagrass beds at night and shelter in or near the coral reefs during the day (large juveniles of coral reef species and adults of schooling pelagic species, respectively). The constant migrations of these fishes between the coral reefs and seagrass beds explained the relative stability of the structure of the fish assemblage in the seagrass bed over time. Thus, the two seagrass beds were not equivalent habitats for fishes. The distinct ecological influences of the mangroves (as a nursery for small juveniles) and coral reefs (as a shelter for larger fishes) on the nearby seagrass beds was clearly reflected by the distinct utilizations of these seagrass beds by fishes.     

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.