Vertical and horizontal distributions of coral‐reef fish larvae in open water immediately prior to reef colonization

To explore the vertical and horizontal distributions of fish larvae near the end of their pelagic period, six light traps were set up over four lunar months at different depths (sub‐surface, midwater and bottom) and different habitat types (reef slope: 50 m horizontal distance from the reef crest; frontier zone: 110 m horizontal distance; sandy zone: 200 m horizontal distance) on the outer reef slope of Moorea Island, French Polynesia. The highest captures were in sub‐surface traps on the reef slope and the frontier zone, and in bottom traps on the sandy zone and the frontier zone. It is hypothesized that fish larvae move towards the surface near the reef slope to avoid reef‐based planktivores and to get into a favourable position for surfing over the reef crest.     

Near- bottom depletion of zooplankton over coral reefs: I: diurnal dynamics and size distribution

Changes in the near-bottom abundance of zooplankton on scales of centimeters to meters and hours to seasons are of great importance to corals and other benthic zooplanktivores. Our objective was to characterize such spatio-temporal changes over several coral reefs in the Gulf of Aqaba, Red Sea. Using arrays of underwater pumps, we found a substantial depletion of zooplankton near the bottom. Vertical gradients in zooplankton abundance were steeper during the night than day, mostly due to a greater nocturnal increase in zooplankton biomass higher in the water column. On average, the layer <1 m above bottom (mab) was depleted by 2.6±2.2 mg m^–3 (46±35%) and 1.4±1.4 mg m^–3 (37±43%) during night and day, respectively. A long time series of bi-weekly samples at 0.5 mab, lasting 1.5 years, indicated a doubling of the biomass during the night with no apparent seasonality. The diel change was due to an increase in the abundance of only large (>200 m) zooplankters around dusk and their disappearance in the morning. Diurnal predation by zooplanktivorous fish, sediment resuspension by benthivorous fish and zooplankton behavior appear to control the dynamics of suspended particles over the reef, creating sharp vertical gradients and a remarkable diel cycle in the ratio between nutritious plankton and inorganic particles.     

In situ observation of settlement behaviour in larvae of coral reef fishes at night

The swimming behaviour of 534 coral reef fish larvae from 27 species was explored at Moorea Island (French Polynesia) while they searched for a suitable settlement habitat, on the first night of their lagoon life. Most larvae swam actively (74%) and avoided the bottom (77%). A significant relationship was highlighted between the vertical position of larvae in the water column and the distance they travelled from lagoon entrance to settlement habitat: larvae swimming close to the surface settled farther away on the reef than bottom-dwelling larvae.     

Spatiotemporal distribution of nocturnal coral reef fish juveniles in Moorea Island,French Polynesia

This study aimed to investigate the spatial structure of nocturnal fish communities at settlement on coral reefs in Moorea Island lagoon, French Polynesia; and the temporal consistency of habitat selection between winter (April to June 2001) and summer (November 2001). The Moorea lagoon was divided into 12 habitat zones (i.e., coral reef zones), which were distinct in terms of depth, wave exposure, and substratum composition. Nocturnal visual censuses among the 12 habitats found that the recently settled juveniles of 25 species recorded were dispatched among three communities spatially distributed according to the distance from the reef crest (reef crest, barrier reef, and fringing reef communities). This spatial communities structure of nocturnal juveniles was consistent in both winter and summer and would be explained primarily by a current gradient in Moorea lagoon (current speed was high near the reef crest and decreased towards the beach) and by the topographic characteristics of reef zones. Among the 25 species, 13 were recorded in both winter and summer. A comparison of the spatial distribution between summer and winter for 13 species that occurred during both seasons found that only 4 differed between the two seasons. For these species, habitat selection would be organized primarily by some stochastic processes such as inter- and intraspecific competition, predation, and food availability. Overall, the present study allowed us to highlight that most nocturnal coral reef fish juveniles at Moorea Island exhibited striking patterns in their distribution and current and topographic characteristics of reef zones might exert considerable influence on the distribution of fishes.     

Distributions of coral reef macroalgae in a back reef habitat in Moorea,French Polynesia

On tropical reefs where macroalgae are subjected to continuous herbivore pressure, spatial refuges typically are identified as large-scale, landscape interfaces that limit foraging behavior. However, algal distributions and community assemblages may also rely on the availability of smaller scale spatial refuges within the reef. The results of this study demonstrate that the patterns of macroalgal distribution across the back reef of Moorea, French Polynesia, are maintained by herbivores interacting with the small-scale structural complexities of the coral reef landscape. Although the majority of space available for colonization is composed of exposed surfaces, macroalgae rarely are found in the open. Instead, macroalgal occurrence is highest in the protected narrow crevices and hole microhabitats provided by massive Porites spp. coral heads. These distributions are determined initially by post-settlement mortality of young algal recruits in exposed habitats. Rates of consumption for two of the most common macroalgal species found in refuges across the back reef, Halimeda minima and Amansia rhodantha, indicate that algal recruits in exposed habitats are limited by herbivory. While algal abundance and community structure are highly dependent upon herbivore grazing, the availability of small-scale spatial refuges ultimately shapes the distinct community patterns and distributional boundaries of coral reef macroalgae in the back reefs of Moorea.     

Spatial variation in mechanical properties of coral reef substrate and implications for coral colony integrity

The physical structure of coral reefs plays a critical role as a barrier to storm waves and tsunamis and as a habitat for living reef-building and reef-associated organisms. However, the mechanical properties of reef substrate (i.e. the non-living benthos) are largely unknown, despite the fact that substrate properties may ultimately determine where organisms can persist. We used a geo-mechanical technique to measure substrate material density and strength over a reef hydrodynamic gradient. Contrary to expectation, we found a weak relationship between substrate strength and wave-induced water flow: flow rates decline sharply at the reef crest, whereas substrate properties are relatively constant over much of the reef before declining by almost an order of magnitude at the reef back. These gradients generate a novel hump-shaped pattern in resistance to mechanical disturbances for live corals, where colonies closer to the back reef are prone to dislodgement because of poorly cemented substrate. Our results help explain an intermediate zone of higher taxonomic and morphological diversity bounded by lower diversity exposed reef crest and unstable reef back zones.     

A geochemical model for coral reef formation

The conspicuous growth of a reef crest and the resulting differentiation of reef topography into a moat (shallow lagoon), crest and slope have long attracted the interest of scientists studying coral reefs. A geochemical model is here proposed for reef formation, taking into account diffusion-limited and light-enhanced calcification. First, to obtain data on net photosynthesis and calcification rates in the field, a typical coral community was cultured in situ on a reef flat. Using these data, equations including parameters for calcification were then developed and applied in computer simulations to model the development over time of reef profiles and the diffusion of carbon species. The reef topography simulated by the model was in general agreement with reef topography observed in nature. The process of reef growth as shown by the modeling was as follows. Increases in the shore-to-offshore gradients of the concentrations of carbonate species result from calcification by reef biota, giving a lower rate of growth on near-shore parts of the reef than on those further offshore. As a result, original topography is diversified into moat and reef crest for the first time. Reef growth on the reef crest is more rapid than in the inshore moat area, because more light is available at the crest. Reef growth on the near-shore side of the reef is further inhibited by damming of carbon-rich seawater on the seaward side of the reef by the reef crest. Over time, the topographic expression of the reef crest and moat becomes progressively more clearly defined by these geochemical processes.     

Diel migration patterns of demersal reef zooplankton

Demersal zooplankton reside in or near the reef substrata and usually migrate into the water column at night. There is no single pulse of migratory activity. The zooplankton rise at variable rates throughout the night, with a peak activity usually during the second hour after sunset. This temporal pattern is a reflection of the behavior of the dominant (80–90% of night samples) cyclopoid, Oithona colcarva Bowman.Not all zooplankton taxa exhibit the same diel migratory patterns. Harpacticoids, another Oithona sp., copepod nauplii, barnacle nauplii, and appendicularians are most abundant during the day. Isopods show a peak of activity also during the second hour after sunset while polychaetes are most abundant during the first hour. The behavior of the other groups studied (the cyclopoid Corycaeus sp., other cyclopoids, ostracods, amphipods, tanaids, decapods, mysids, and chaetognaths) was less easily defined.The migration of many species in a pulse during the period of least planktivore activity and migration during the day of small species and juvenile members of larger species suggests that visual predators have an important influence on the migratory behavior of reef zooplankton.     

Diel variation of zooplankton in the tropical coral-reef water of Tioman Island,Malaysia

Zooplankton was sampled at 3-h intervals for a 48-h period from a coral reef of Tioman Island, Malaysia. It was size-fractionated into three size classes: 100–200, 200–335, and >335 μm using different sieves with different mesh sizes. Total zooplankton (>100 μm) abundance and biomass in the water column were high later at night (0300 h), not just after sunset as previously described in other studies. Only the largest size-fraction (>335 μm) of zooplankton significantly differed in biomass and abundance between day and night. The increase in the large zooplankton later in the night is suggested to be caused by the advection of pelagic species into the reef. This work has provided a measurement of the variation of zooplankton community over coral reef that can exist on a scale of hours.     

Sedimentation on three caribbean atolls: Glovers reef,lighthouse reef and turneffe Islands,belize

Summary The chief mode of carbonate sedimentation on the Belizean atolls Glovers Reef, Lighthouse Reef and Turneffe Islands is the accumulation of organically-derived particles. Variations in the distribution of the composition and grain-sizes of surface sediments, collected along transects across the atolls, are environmentally controlled. Two major sediment types may be distinguished. (1) Reef and fore reef sediments are dominated by fragments of coral, coralline algae andHalimeda. Mean grain-sizes range from 1–2 mm. (2) Back reef sediments contain more mollusk fragments, more fine-grained sediment (<125 μm) and appear to have fewerHalimeda fragments. In addition, sediments from inner platforms and shallow lagoonal parts of Glovers and Lighthouse Reefs comprise non-skeletal grains, namely fecal pellets. Sediments from lagoonal patch reefs may contain up to 20% coral fragments. Mean grain-sizes range from 0.1–1 mm and are finest on the inner platform and lagoon floor of the back reef environment. Within the reef and fore reef environments, it is not possible to distinguish sub-environments on the basis of textural and compositional differences of the sediments. Sediments from patch reefs contrast with those from back reef lagoons and inner platforms and are similar in terms of grain-sizes and compositions to reef and fore reef surface sediments. Non-skeletal grains forming in shallow parts of the back reef in Glovers and Lighthouse Reefs are interpreted to be indurated by interstitial precipitation of calcium carbonate from warm, supersaturated water flushing the sediment. The lack of hardened non-skeletal particles in the back reef sediments of Turneffe Islands is most probably due to the abundance of muddy, organic-rich sediment in the well-protected lagoon. Fine sediment is less permeable and organic films prevent cement overgrowth on particles.     

Sediment suppresses herbivory across a coral reef depth gradient

Sediments are a ubiquitous feature of all coral reefs, yet our understanding of how they affect complex ecological processes on coral reefs is limited. Sediment in algal turfs has been shown to suppress herbivory by coral reef fishes on high-sediment, low-herbivory reef flats. Here, we investigate the role of sediment in suppressing herbivory across a depth gradient (reef base, crest and flat) by observing fish feeding following benthic sediment reductions. We found that sediment suppresses herbivory across all reef zones. Even slight reductions on the reef crest, which has 35 times less sediment than the reef flat, resulted in over 1800 more herbivore bites (h⁻¹ m⁻²). The Acanthuridae (surgeonfishes) were responsible for over 80 per cent of all bites observed, and on the reef crest and flat took over 1500 more bites (h⁻¹ m⁻²) when sediment load was reduced. These findings highlight the role of natural sediment loads in shaping coral reef herbivory and suggest that changes in benthic sediment loads could directly impair reef resilience.     

Flow through a rough,shallow reef

This paper presents a simple model of wave-driven flow through a coral reef that is characterized by a shallow, wide reef crest and a deeper, but frictional lagoon. Assuming that the dominant momentum balances are between quadratic drag and barotropic pressure gradients, along-lagoon and cross-reef flows are coupled through continuity and through a setup of the water surface in the lagoon that varies in the along-reef direction. Scaling of the governing equations shows that this flow is governed by a single parameter P that expresses the competing effects of cross-reef and along-lagoon drag. When P < 1, the cross-reef flow is nearly constant, whereas when P > 1, only that portion of the reef closest to the pass through the reef crest through which fluid exists the lagoon supports cross-reef flows.     

Near-bottom depletion of zooplankton over a coral reef II: relationships with zooplankton swimming ability

Intense predation by corals and associated fauna can generate vertical gradients of their zooplankton prey. The goal of this study was to characterize the small-scale distribution of zooplankton above the coral reef of Eilat, Israel. Four vertical arrays, each consisting five underwater pumps attached 0.5–3 m apart on a taut mooring, were deployed for 10 days at two sites over the reef slope. A distinct layer of depleted zooplankton was repeatedly found in the 1.5 m high benthic boundary layer. The gradient was sharpest for strong swimmers (copepods and polychaeta), intermediate for weak swimmers (nauplii, mollusks and appendicularia), and lacking for passive taxa (eggs and foraminifera). Spatio-temporal changes in the abundance of copepods and polychaetes were highly correlated, more so in the water aloft (Pearson r>0.9) than near the bottom (r>0.8). The spatio-temporal correlations between weak swimmers and between passive taxa were much weaker (r<0.8 and r<0.4, respectively). Flow and shear stress did not affect the distribution patterns. The correspondence between the zooplankton distribution and their swimming ability indicates that bottom avoidance contribute to the formation of depleted layer over the reef.     

Comparative predation rates on larval snappers (Lutjanidae) in oceanic, reef, and nearshore waters

The life cycle of most reef fishes involves pelagic larvae entering the nearshore environment to settle to benthic substrates. Settlement is considered to be highly risky as larvae encounter high rates of predation mortality associated with shallow nearshore habitats. This potential bottleneck may be particularly significant for many tropical snapper (Lutjanidae) species which bypass the reef to settle to very nearshore seagrass areas. To test the hypothesis that predation-related mortality increases as naïve late-stage fish larvae leave the offshore oceanic environment and enter the nearshore to settle, relative nocturnal predation rates on tethered late-stage snapper larvae were measured in oceanic, coral reef, and nearshore surface waters of the lower Florida Keys, USA. Both relative predation rate and probability of predation in oceanic areas seaward of the reef was significantly greater than over reef or nearshore seagrass/hardbottom habitats. This surprising result may be due to differences in the density or spatial distribution of potential predators between deep offshore (near flotsam at the surface) and shallow nearshore environments (demersal). These findings suggest that successful late-stage snapper larvae should avoid surface waters in deep oceanic areas and move upward in the water column as they pass over the reef and other shallow nearshore environments prior to settlement.     

Pelagic to demersal transition in a coral‐reef fish,the orbicular batfish Platax orbicularis

Behavioural and ecological observations were made on young, reared Platax orbicularis in Opunohu Bay, Moorea, French Polynesia, during their transition from the pelagic, dispersive stage to the reef‐orientated demersal stage. Seventy‐two young P. orbicularis (17–75 mm standard length, L_S) were released in the pelagic zone and 20 (40–70 mm L_S) adjacent to the reefs. Swimming speed was slow (mean 5·2 cm s^?1) and independent of size. An ontogenetic descent was observed: the smallest P. orbicularis swam at the surface, medium‐sized P. orbicularis swam in midwater (mean 5–13 m) and the largest P. orbicularis swam to the bottom, where many lay on their sides. Platax orbicularis swam southerly on average, away from the ocean and into the bay. Smaller P. orbicularis were more likely to swim directionally than larger individuals. Young P. orbicularis released near reef edges swam at similar, but more variable speeds (mean 6·6 cm s^?1). About half of those released near reefs swam away, but fewer swam away from an inshore fringing reef than from a patch reef near the bay mouth. Many P. orbicularis swam up the slope onto the reef top, but the little settlement observed was near the reef base. Average, near‐reef swimming direction was also southerly. Some reef residents, in particular the triggerfish Balistapus undulatus, harassed young P. orbicularis.     

A modern-type Kimmeridgian reef (Ota Limestone,Portugal): Implications for Jurassic reef models

Upper Jurassic reefs rich in microbial crusts generally appear in deeper (sponge—‘algal’ crust reefs) or in very shallow but protected settings (coral or coral-coralline sponge meadows with ‘algal’ crusts). Upper Jurassic high-energy reefs (coral reefs and coral-stromatoporoid reefs) normally lack major participation of microbial crusts but rather represent huge bioclastic piles with only minor framestone patches preserved. An exception to this rule is represented by the high-energy, coral-‘algal’ Ota Reef from the Kimmeridgian of the Lusitanian Basin (Portugal). The narrow Ota Reef tract rims a small intra-basinal carbonate platform exhibiting perfect facies zonation (from W to E: Reef tract, back reef sands, peritidal belt, low-energy shallow lagoon). The reef is dominated by massive corals (Thamnasteria, Microsolena, Stylina). Complete preservation of coral framework is rare: like other Upper Jurassic high-energy reefs, the Ota Reef is very rich in debris; however, this debris is largely stabilized by algal and microbial crusts, what contrasts the other examples and gives the Ota Reef the appearance of a typical modern high-energy coral-melobesioid algal reef. Further similarities to modern reefs are the likely existence of a spur-and-groove system, the perfect sheltering of inner platform areas and the occurrence of small islands, as indicated by local blackenings and early vadose and karstic features.     

Framework of barrier reefs threatened by ocean acidification

To date, studies of ocean acidification (OA) on coral reefs have focused on organisms rather than communities, and the few community effects that have been addressed have focused on shallow back reef habitats. The effects of OA on outer barrier reefs, which are the most striking of coral reef habitats and are functionally and physically different from back reefs, are unknown. Using 5‐m long outdoor flumes to create treatment conditions, we constructed coral reef communities comprised of calcified algae, corals, and reef pavement that were assembled to match the community structure at 17 m depth on the outer barrier reef of Moorea, French Polynesia. Communities were maintained under ambient and 1200 μatm pCO₂ for 7 weeks, and net calcification rates were measured at different flow speeds. Community net calcification was significantly affected by OA, especially at night when net calcification was depressed ~78% compared to ambient pCO₂. Flow speed (2–14 cm s^?1) enhanced net calcification only at night under elevated pCO₂. Reef pavement also was affected by OA, with dissolution ~86% higher under elevated pCO₂ compared to ambient pCO₂. These results suggest that net accretion of outer barrier reef communities will decline under OA conditions predicted within the next 100 years, largely because of increased dissolution of reef pavement. Such extensive dissolution poses a threat to the carbonate foundation of barrier reef communities.     

Near-bottom depletion of zooplankton over coral reefs: III: vertical gradient of predation pressure

Many diurnal planktivorous fish in coral reefs efficiently consume zooplankton drifting in the overlying water column. Our survey, carried out at two coral reefs in the Red Sea, showed that most of the diurnal planktivorous fish foraged near the bottom, close to the shelters from piscivores. The planktivorous fish were order of magnitude more abundant near (<1.5 m) the bottom than higher in the water column. The predation pressure exerted by these fish was assessed by measuring the consumption of brine shrimps tethered at different heights above the bottom on a vertical line which was pulled over the reef. Below 1.5 m above bottom, the shrimps survival probability sharply decreased toward the bottom. Higher in the water column, survivorship was nearly 100% with little vertical variation. Our results indicate that near-bottom depletion of zooplankton in coral reefs is likely due to intense predation at that boundary layer. Risk of predation by piscivorous fish apparently restricts planktivorous fish to forage near the bottom, with a distribution pattern greatly deviating from ideal-free distribution.     

Influences of wind-wave exposure on the distribution and density of recruit reef fishes at Kure and Pearl and Hermes Atolls,Northwestern Hawaiian Islands

This paper describes the results of a field survey designed to test the prediction that the density of benthic juveniles of shallow-reef fishes is greater on wind-wave “exposed” sectors of a pair of isolated oceanic atolls (Kure, Pearl and Hermes) at the far northwestern end of the Hawaiian Islands, an archipelago in which east-northeasterly trade winds dominate onshore water flow and transport by surface currents. The densities of recruits (juveniles ≤5 cm total length) were higher overall on windward versus leeward sectors of carbonate rock-rubble back reefs at both atolls, and the pattern was stronger for smaller (likely younger, more recently settled) recruits of four of the five most abundant species and the remainder pooled as an “Other” taxon. The windward-leeward disparity was four-fold greater at Pearl Hermes (the atoll with a three-fold longer perimeter) than at Kure. Resident predator biomass also was correlated with recruit densities, but habitat (benthic substratum) effects were generally weak. The distribution and abundance of recruits and juveniles of the primarily endemic reef fishes on shallow back reefs at these atolls appear partly influenced by relative rates of water flow over windward vs. leeward sectors of barrier reef and by the size, shape, and orientation of habitat parcels that filter out postlarval fishes with relatively weak swimming capabilities like labroids. Whole-reef geomorphology as well as fine-scale habitat heterogeneity and rugosity should be considered among the suite of many factors used to interpret observed spatial patterns of post-settlement juvenile fish distribution at atolls and perhaps some other tropical reefs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Performance of remote acoustic receivers within a coral reef habitat: implications for array design

Remote monitoring technologies are increasingly being implemented in the marine environment to better understand the movement patterns of taxa. Coral reefs are no exception. However, there is a paucity of information relating to the performance of acoustic receivers on coral reefs. Our results suggest that the detection performance of acoustic receivers may be significantly impacted by the unique nature of the reef environment. This study assessed the performance of passive acoustic receivers on a typical inner-shelf fringing reef, Orpheus Island, on the Great Barrier Reef, Australia. The detection range and diel performance variability of acoustic receivers was assessed using two parallel lines of 5 VR2W receivers spanning 125?m, deployed on the reef base and reef crest. Two 9-mm acoustic transmitters were moored at opposite ends of each receiver line. The working detection range for receivers was found to be approximately 90?m for the transmitter moored on the reef base and just 60?m for the transmitter moored on the reef crest. However, the detection range on the reef crest increased to 90?m when just the reef crest receivers were considered, highlighting importance of optimal receiver deployment. No diel patterns in receiver performance or detection capacities were detected, suggesting that no corrections are required when interpreting nocturnal versus diurnal activity patterns. We suggest that studies aiming for complete coverage of a site within a reef environment will require receivers in close (<100?m) proximity, and that the placement depth of receivers must be a major consideration, with shallow receivers exhibiting a greater detection range than those on the reef slope. Our results highlight the challenges imposed by coral reefs for acoustic telemetry and the importance of receiver placement for studies conducted within these habitats.