Region‐wide temporal and spatial variation in Caribbean reef architecture: is coral cover the whole story?

The architectural complexity of coral reefs is largely generated by reef‐building corals, yet the effects of current regional‐scale declines in coral cover on reef complexity are poorly understood. In particular, both the extent to which declines in coral cover lead to declines in complexity and the length of time it takes for reefs to collapse following coral mortality are unknown. Here we assess the extent of temporal and spatial covariation between coral cover and reef architectural complexity using a Caribbean‐wide dataset of temporally replicated estimates spanning four decades. Both coral cover and architectural complexity have declined rapidly over time, with little evidence of a time‐lag. However, annual rates of change in coral cover and complexity do not covary, and levels of complexity vary greatly among reefs with similar coral cover. These findings suggest that the stressors influencing Caribbean reefs are sufficiently severe and widespread to produce similar regional‐scale declines in coral cover and reef complexity, even though reef architectural complexity is not a direct function of coral cover at local scales. Given that architectural complexity is not a simple function of coral cover, it is important that conservation monitoring and restoration give due consideration to both architecture and coral cover. This will help ensure that the ecosystem services supported by architectural complexity, such as nutrient recycling, dissipation of wave energy, fish production and diversity, are maintained and enhanced.     

Benthic N2 fixation in coral reefs and the potential effects of human‐induced environmental change

Tropical coral reefs are among the most productive and diverse ecosystems, despite being surrounded by ocean waters where nutrients are in short supply. Benthic dinitrogen (N₂) fixation is a significant internal source of “new” nitrogen (N) in reef ecosystems, but related information appears to be sparse. Here, we review the current state (and gaps) of knowledge on N₂ fixation associated with coral reef organisms and their ecosystems. By summarizing the existing literature, we show that benthic N₂ fixation is an omnipresent process in tropical reef environments. Highest N₂ fixation rates are detected in reef‐associated cyanobacterial mats and sea grass meadows, clearly showing the significance of these functional groups, if present, to the input of new N in reef ecosystems. Nonetheless, key benthic organisms such as hard corals also importantly contribute to benthic N₂ fixation in the reef. Given the usually high coral coverage of healthy reef systems, these results indicate that benthic symbiotic associations may be more important than previously thought. In fact, mutualisms between carbon (C) and N₂ fixers have likely evolved that may enable reef communities to mitigate N limitation. We then explore the potential effects of the increasing human interferences on the process of benthic reef N₂ fixation via changes in diazotrophic populations, enzymatic activities, or availability of benthic substrates favorable to these microorganisms. Current knowledge indicates positive effects of ocean acidification, warming, and deoxygenation and negative effects of increased ultraviolet radiation on the amount of N fixed in coral reefs. Eutrophication may either boost or suppress N₂ fixation, depending on the nutrient becoming limiting. As N₂ fixation appears to play a fundamental role in nutrient‐limited reef ecosystems, these assumptions need to be expanded and confirmed by future research efforts addressing the knowledge gaps identified in this review.     

Coral reef primary productivity. A hierarchy of pattern and process

The abundance of interdisciplinary studies of coral reef metabolism means that coral reefs are amongst the best understood of marine benthic communities in terms of the determinants of primary production and nutrient fluxes, and their variation, at different levels of integration. Recent work has extended our understanding of the variation in coral reef productivity at different spatial and temporal scales. Nutrient limitation of production at scales ranging from the organism to the ecosystem is one of the most important topics of research at present, the relevance of which extends well beyond the tropics.     

High macroalgal cover and low coral recruitment undermines the potential resilience of the world's southernmost coral reef assemblages

Coral reefs are under increasing pressure from anthropogenic and climate-induced stressors. The ability of reefs to reassemble and regenerate after disturbances (i.e., resilience) is largely dependent on the capacity of herbivores to prevent macroalgal expansion, and the replenishment of coral populations through larval recruitment. Currently there is a paucity of this information for higher latitude, subtropical reefs. To assess the potential resilience of the benthic reef assemblages of Lord Howe Island (31°32'S, 159°04'E), the worlds' southernmost coral reef, we quantified the benthic composition, densities of juvenile corals (as a proxy for coral recruitment), and herbivorous fish communities. Despite some variation among habitats and sites, benthic communities were dominated by live scleractinian corals (mean cover 37.4%) and fleshy macroalgae (20.9%). Live coral cover was higher than in most other subtropical reefs and directly comparable to lower latitude tropical reefs. Juvenile coral densities (0.8 ind.m(-2)), however, were 5-200 times lower than those reported for tropical reefs. Overall, macroalgal cover was negatively related to the cover of live coral and the density of juvenile corals, but displayed no relationship with herbivorous fish biomass. The biomass of herbivorous fishes was relatively low (204 kg.ha(-1)), and in marked contrast to tropical reefs was dominated by macroalgal browsing species (84.1%) with relatively few grazing species. Despite their extremely low biomass, grazing fishes were positively related to both the density of juvenile corals and the cover of bare substrata, suggesting that they may enhance the recruitment of corals through the provision of suitable settlement sites. Although Lord Howe Islands' reefs are currently coral-dominated, the high macroalgal cover, coupled with limited coral recruitment and low coral growth rates suggest these reefs may be extremely susceptible to future disturbances.     

Modelling three-dimensional flow over spur-and-groove morphology

Spur-and-groove (SAG) morphology characterizes the fore reef of many coral reefs worldwide. Although the existence and geometrical properties of SAG have been well documented, an understanding of the hydrodynamics over them is limited. Here, the three-dimensional flow patterns over SAG formations, and a sensitivity of those patterns to waves, currents, and SAG geometry were characterized using the physics-based Delft3D-FLOW and SWAN models. Shore-normal shoaling waves over SAG formations were shown to drive two circulation cells: a cell on the lower fore reef with offshore flow over the spurs and onshore flow over the grooves, except near the seabed where velocities were always onshore, and a cell on the upper fore reef with offshore surface velocities and onshore bottom currents, which result in depth-averaged onshore and offshore flow over the spurs and grooves, respectively. The mechanism driving this flow results from the net of the radiation stress gradients and pressure gradient, which is balanced by the Reynolds stress gradients and bottom friction that differ over the spur and over the groove. Waves were the primary driver of variations in modelled flow over SAG, with the flow strength increasing for increasing wave heights and periods. Spur height, SAG wavelength, and the water depth at peak spur height were the dominant influences on the hydrodynamics, with spur heights directly proportional to the strength of SAG circulation cells. SAG formations with shorter SAG wavelengths only presented one circulation cell on the shallower portion of the reef, as opposed to the two circulation cells for longer SAG wavelengths. SAG formations with peak spur heights occurring in shallower water had stronger circulation than those with peak spur heights occurring in deeper water. These hydrodynamic patterns also likely affect coral and reef development through sediment and nutrient fluxes.

     

PROCESSES OF ORGANIC PRODUCTION ON CORAL REEFS

1. The first quantitative studies of production on coral reefs were those of Sargent & Austin who showed that productivity on reefs was considerably higher than in surrounding waters. This high production occurred in spite of nutrient limitation and low productivity of offshore waters. Their conclusions have since been confirmed by numerous other workers in both the Atlantic and the Pacific. 2. Primary production on reefs has been studied by flow respirometry, measuring changes in oxygen or carbon dioxide concentrations in water flowing over reefs. Production of benthic organisms has also been measured in situ by light and dark bottle methods and by radioactive tracer techniques. Production values obtained by the various methods are not identical but their use in combination is to be recommended. 3. Rates of gross primary production on reefs vary between 300–5000 gC/m²/yr. These rates are higher than general oceanic values and as high as those of the most productive marine communities. 4. Sources of primary production include fleshy macrophytes, calcareous algae, filamentous algae on the coral skeletons or calcareous rock, marine grasses and the zooxanthellae within coral tissue. Production values from the various sources fall within the range of production of reefs as a whole. 5. Concentrations of nitrogen and phosphorus in waters flowing over reefs are consistently low. There is evidence to suggest that both these nutrients are recycled rapidly on the reef and that nitrogen is fixed by bacteria and primary producers. 6. In many instances the mass of detritus over coral reefs exceeds the biomass of zooplankton. While the quantitative significance of detritus as food for corals and other benthic organisms has not been evaluated, there is a growing body of evidence to show that this may be the key to understanding secondary production. 7. Opinions differ on the adequacy of zooplankton in satisfying the food requirements of corals and other benthic invertebrates on reefs. The weight of evidence suggests that while there is a removal of zooplankton by benthic organisms, the total biomass carried over the reef is too small to support the energy needs of secondary production. 8. Bacteria are a potential source of energy for secondary production on reefs and are implicated in nitrogen fixation, decomposition and biogeochemical cycling. 9. There is an abundance of sessile invertebrates other than corals on reefs but there are few quantitative data on their importance in secondary production. 10. The biomass of fish on reefs may be very high but the quantitative significance of grazing and predation is not fully established. 11. Studies on the growth of corals themselves have been based on measurements of skeletal accretion. These methods do not lead directly to estimates of reef organic production. Growth rates of corals vary considerably between and within species. 12. Estimates of reef growth have been made from measurements of coral growth and from the flux of calcium carbonate. There is less quantitative information on erosion caused by mechanical damage, by boring organisms and by human pollution. 13. Hydrographic factors influence growth and form of reefs and there is some evidence to show that production is enhanced by conservation of water in lagoonal areas.     

Coral Reefs at the Northernmost Tip of Borneo: An Assessment of Scleractinian Species Richness Patterns and Benthic Reef Assemblages

The coral reefs at the northernmost tip of Sabah, Borneo will be established under a marine protected area: the Tun Mustapha Park (TMP) by the end of 2015. This area is a passage where the Sulu Sea meets the South China Sea and it is situated at the border of the area of maximum marine biodiversity, the Coral Triangle. The TMP includes fringing and patch reefs established on a relatively shallow sea floor. Surveys were carried out to examine features of the coral reefs in terms of scleractinian species richness, and benthic reef assemblages following the Reef Check substrate categories, with emphasis on hard coral cover. Variation in scleractinian diversity was based on the species composition of coral families Fungiidae (n = 39), Agariciidae (n = 30) and Euphylliidae (n = 15). The number of coral species was highest at reefs with a larger depth gradient i.e. at the periphery of the study area and in the deep South Banggi Channel. Average live hard coral cover across the sites was 49%. Only 7% of the examined reefs had > 75% hard coral cover, while the majority of the reef sites were rated fair (51%) and good (38%). Sites with low coral cover and high rubble fragments are evidence of blast fishing, although the observed damage appeared old. Depth was a dominant factor in influencing the coral species composition and benthic reef communities in the TMP. Besides filling in the information gaps regarding species richness and benthic cover for reef areas that were previously without any data, the results of this study together with information that is already available on the coral reefs of TMP will be used to make informed decisions on zoning plans for conservation priorities in the proposed park.     

The influence of coral reef benthic condition on associated fish assemblages

Accumulative disturbances can erode a coral reef's resilience, often leading to replacement of scleractinian corals by macroalgae or other non-coral organisms. These degraded reef systems have been mostly described based on changes in the composition of the reef benthos, and there is little understanding of how such changes are influenced by, and in turn influence, other components of the reef ecosystem. This study investigated the spatial variation in benthic communities on fringing reefs around the inner Seychelles islands. Specifically, relationships between benthic composition and the underlying substrata, as well as the associated fish assemblages were assessed. High variability in benthic composition was found among reefs, with a gradient from high coral cover (up to 58%) and high structural complexity to high macroalgae cover (up to 95%) and low structural complexity at the extremes. This gradient was associated with declining species richness of fishes, reduced diversity of fish functional groups, and lower abundance of corallivorous fishes. There were no reciprocal increases in herbivorous fish abundances, and relationships with other fish functional groups and total fish abundance were weak. Reefs grouping at the extremes of complex coral habitats or low-complexity macroalgal habitats displayed markedly different fish communities, with only two species of benthic invertebrate feeding fishes in greater abundance in the macroalgal habitat. These results have negative implications for the continuation of many coral reef ecosystem processes and services if more reefs shift to extreme degraded conditions dominated by macroalgae.     

Great Barrier Reef butterflyfish community structure: the role of shelf position and benthic community type

The extent to which fish communities are structured by spatial variability in coral reef habitats versus stochastic processes (such as larval supply) is very important in predicting responses to sustained and ongoing habitat degradation. In this study, butterflyfish and benthic communities were surveyed annually over 15 years on 47 reefs (spanning 12° of latitude) of the Great Barrier Reef (GBR). Spatial autocorrelation in the structure of butterflyfish communities versus key differences in reef habitats was investigated to assess the extent to which the structure of these fish communities is influenced by habitat conditions. Benthic communities on each of the 47 reefs were broadly categorised as either: 1. Poritidae/Alcyoniidae, 2. mixed taxa, 3. soft coral or 4. Acropora-dominated habitats. These habitat types most reflected increases in water clarity and wave exposure, moving across the GBR shelf from coastal to outer-shelf environments. In turn, each habitat type also supported very distinct butterflyfish communities. Hard coral feeders were always the dominant butterflyfish species in each community type. However, the numerically dominant species changed according to habitat type, representing spatial replacement of species across the shelf. This study reveals clear and consistent differences in the structure of fish communities among reefs associated with marked differences in habitat structure.     

Holocene coral reef accretion in Hawaii: a function of wave exposure and sea level history

 In the high Hawaiian Islands, significant accretion due to coral reef growth is limited by wave exposure and sea level. Holocene coral growth and reef accretion was measured at four stations off Oahu, Hawaii, chosen along a gradient in wave energy from minimum to maximum exposures. The results show that coral growth of living colonies (linear extension) at optimal depths is comparable at all stations (7.7–10.1 mm/y), but significant reef accretion occurs only at wave sheltered stations. At wave sheltered stations in Hanauma Bay and Kaneohe Bay, rates of long term reef accretion are about 2.0 mm/y. At wave exposed stations, off Mamala Bay and Sunset Beach, reef accretion rates are virtually zero in both shallow (1 m) and deeper (optimal) depths (12 m). At wave sheltered stations, such as Kaneohe Bay and Hanauma Bay, Holocene reef accretion is on the order of 10–15 m thick. At wave exposed stations, Holocene accretion is represented by only a thin veneer of living corals resting on antecedent Pleistocene limestone foundations. Modern coral communities in wave exposed environments undergo constant turnover associated with mortality and recruitment or re-growth of fragmented colonies and are rarely thicker than a single living colony. Breakage, scour, and abrasion of living corals during high wave events appears to be the major source of mortality and ultimately limits accretion to wave sheltered environments. Depth is particularly important as a modulator of wave energy. The lack of coral reef accretion along shallow open ocean coastlines may explain the absence of mature barrier reefs in the high Hawaiian Islands. Accepted: 14 May 1998     

Towards an understanding of resilience in isolated coral reefs

In 1998, seawater temperature anomalies led to unprecedented levels of coral bleaching on reefs worldwide. We studied the direct effects of this thermal event on benthic communities and its indirect effects on their associated coral reef fish communities at a group of remote reefs off NW Australia. Long‐term monitoring of benthic and fish assemblages on these reefs allowed us to compare the responses of these communities to coral bleaching using a data series that included 4 years before, and 6 years following, this bleaching event. While bleaching mortality was evident to >30 m depth, it was patchy among the shallower survey sites with decreases in live coral cover ranging from 30% to 90% across seven surveyed locations Within 2 years of the bleaching, hard coral recovery had begun at all sites and by 2003 reef‐wide coral cover had increased to ～39% of its preimpact levels. We exploited this pattern of differential survival of corals among sites, the associated changes in these benthic communities, and their patterns of recovery, to better understand links between benthic community dynamics and their associated fish communities. Temporal changes in the resident fish communities strongly reflected the differential shifts in the benthic communities, but were lagged by 12–18 months. Five years after the bleaching event, the fish communities on five of the seven surveyed locations showed evidence of recovery, however, none had regained their preimpact structures. Analyses of these communities by taxonomic family revealed a range of responses to the disturbance reflective of their life‐histories and trophic and habitat affiliations. The slow but recognizable recovery of this isolated reef system has parallels with other relatively isolated systems that displayed resilience to the 1998 bleaching event, e.g. the Chagos archipelago, but it also contrasts sharply with low levels of resilience documented in other isolated reef systems subject to the same disturbance, e.g. the Seychelles. In this context, our results highlight the significant knowledge gaps remaining in understanding the resilience of these ecosystems to disturbance.     

Benthic Reef Primary Production in Response to Large Amplitude Internal Waves at the Similan Islands (Andaman Sea,Thailand)

Coral reefs are facing rapidly changing environments, but implications for reef ecosystem functioning and important services, such as productivity, are difficult to predict. Comparative investigations on coral reefs that are naturally exposed to differing environmental settings can provide essential information in this context. One prevalent phenomenon regularly introducing alterations in water chemistry into coral reefs are internal waves. This study therefore investigates the effect of large amplitude internal waves (LAIW) on primary productivity in coral reefs at the Similan Islands (Andaman Sea, Thailand). The LAIW-exposed west sides of the islands are subjected to sudden drops in water temperature accompanied by enhanced inorganic nutrient concentrations compared to the sheltered east. At the central island, Ko Miang, east and west reefs are only few hundred meters apart, but feature pronounced differences. On the west lower live coral cover (-38 %) coincides with higher turf algae cover (+64 %) and growth (+54 %) compared to the east side. Turf algae and the reef sand-associated microphytobenthos displayed similar chlorophyll a contents on both island sides, but under LAIW exposure, turf algae exhibited higher net photosynthesis (+23 %), whereas the microphytobenthos displayed reduced net and gross photosynthesis (-19 % and -26 %, respectively) accompanied by lower respiration (-42 %). In contrast, the predominant coral Porites lutea showed higher chlorophyll a tissues contents (+42 %) on the LAIW-exposed west in response to lower light availability and higher inorganic nutrient concentrations, but net photosynthesis was comparable for both sides. Turf algae were the major primary producers on the west side, whereas microphytobenthos dominated on the east. The overall primary production rate (comprising all main benthic primary producers) was similar on both island sides, which indicates high primary production variability under different environmental conditions.     

Differential modification of seawater carbonate chemistry by major coral reef benthic communities

Ocean acidification (OA) resulting from uptake of anthropogenic CO₂ may negatively affect coral reefs by causing decreased rates of biogenic calcification and increased rates of CaCO₃ dissolution and bioerosion. However, in addition to the gradual decrease in seawater pH and Ω _a resulting from anthropogenic activities, seawater carbonate chemistry in these coastal ecosystems is also strongly influenced by the benthic metabolism which can either exacerbate or alleviate OA through net community calcification (NCC = calcification – CaCO₃ dissolution) and net community organic carbon production (NCP = primary production ? respiration). Therefore, to project OA on coral reefs, it is necessary to understand how different benthic communities modify the reef seawater carbonate chemistry. In this study, we used flow-through mesocosms to investigate the modification of seawater carbonate chemistry by benthic metabolism of five distinct reef communities [carbonate sand, crustose coralline algae (CCA), corals, fleshy algae, and a mixed community] under ambient and acidified conditions during summer and winter. The results showed that different communities had distinct influences on carbonate chemistry related to the relative importance of NCC and NCP. Sand, CCA, and corals exerted relatively small influences on seawater pH and Ω _a over diel cycles due to closely balanced NCC and NCP rates, whereas fleshy algae and mixed communities strongly elevated daytime pH and Ω _a due to high NCP rates. Interestingly, the influence on seawater pH at night was relatively small and quite similar across communities. NCC and NCP rates were not significantly affected by short-term acidification, but larger diel variability in pH was observed due to decreased seawater buffering capacity. Except for corals, increased net dissolution was observed at night for all communities under OA, partially buffering against nighttime acidification. Thus, algal-dominated areas of coral reefs and increased net CaCO₃ dissolution may partially counteract reductions in seawater pH associated with anthropogenic OA at the local scale.     

Assessing the accuracy of acoustic seabed classification for mapping coral reef environments in South Florida (Broward County, USA)

The Atlantic coast of Broward County, Florida (USA) is paralleled by a series of progressively deeper, shore-parallel coral reef communities. Two of these reef systems are drowned early Holocene coral reefs of 5 ky and 7 ky uncorrected radiocarbon age. Despite the case of access to these reefs, and their major contribution to the local economy, accurate benthic habitat maps of the area are not available. Ecological studies have shown that different benthic communities (i.e. communities composed of different biological taxa) exist along several spatial gradients on all reefs. Since these studies are limited by time and spatial extent, acoustic surveys with the QTCView V bottom classification system based on a 50 kHz transducer were used as an alternative method of producing habitat maps. From the acoustic data of a 3.1 km(2) survey area, spatial prediction maps were created for the area. These were compared with habitat maps interpreted from in situ data and Laser Airborne Depth Sounder (LADS) bathymetry, in order to ground-truth the remotely sensed data. An error matrix was used to quantitatively determine the accuracy of the acoustically derived spatial prediction model against the maps derived from the in situ and LADS data sets. Confusion analysis of 100 random points showed that the system was able to distinguish areas of reef from areas of rubble and sand with an overall accuracy of 61%. When asked to detect more subtle spatial differences, for example, those between distinct reef communities, the classification was only about 40% accurate. We discuss to what degree a synthesis of acoustic and in situ techniques can provide accurate habitat maps in coral reef environments, and conclude that acoustic methods were able to reflect the spatial extent and composition of at least three different biological communities.     

A geospatial assessment of the relationship between reef flat community calcium carbonate production and wave energy

The ability of benthic communities inhabiting coral reefs to produce calcium carbonate underpins the development of reef platforms and associated sedimentary landforms, as well as the fixation of inorganic carbon and buffering of diurnal pH fluctuations in ocean surface waters. Quantification of the relationship between reef flat community calcium carbonate production and wave energy provides an empirical basis for understanding and managing this functionally important process. This study employs geospatial techniques across the reef platform at Lizard Island, Great Barrier Reef, to (1) map the distribution and estimate the total magnitude of reef community carbonate production and (2) empirically ascertain the influence of wave energy on community carbonate production. A World-View-2 satellite image and a field data set of 364 ground referencing points are employed, along with data on physical reef characteristics (e.g. bathymetry, rugosity) to map and validate the spatial distribution of the four major community carbonate producers (live coral, carbonate sand, green calcareous macroalgae and encrusting calcified algae) across the reef platform. Carbonate production is estimated for the complete reef platform from the composition of these community components. A synoptic model of wave energy is developed using the Simulating WAves Nearshore (SWAN) two-dimensional model for the entire reef platform. The relationship between locally derived measures of carbonate production and wave energy is evaluated at both the global scale and local scale along spatial gradients of wave energy traversing the reef platform. A wave energy threshold is identified, below which carbonate production levels appear to increase with wave energy and above which mechanical forcing reduces community production. This implies an optimal set of hydrodynamic conditions characterized by wave energy levels of approximately 300 J m^?2, providing an empirical basis for management of potential changes in community carbonate production associated with climate change-driven increases in wave energy.     

The effects of top–down versus bottom–up control on benthic coral reef community structure

While climate change and associated increases in sea surface temperature and ocean acidification, are among the most important global stressors to coral reefs, overfishing and nutrient pollution are among the most significant local threats. Here we examined the independent and interactive effects of reduced grazing pressure and nutrient enrichment using settlement tiles on a coral-dominated reef via long-term manipulative experimentation. We found that unique assemblages developed in each treatment combination confirming that both nutrients and herbivores are important drivers of reef community structure. When herbivores were removed, fleshy algae dominated, while crustose coralline algae (CCA) and coral were more abundant when herbivores were present. The effects of fertilization varied depending on herbivore treatment; without herbivores fleshy algae increased in abundance and with herbivores, CCA increased. Coral recruits only persisted in treatments exposed to grazers. Herbivore removal resulted in rapid changes in community structure while there was a lag in response to fertilization. Lastly, re-exposure of communities to natural herbivore populations caused reversals in benthic community trajectories but the effects of fertilization remained for at least 2 months. These results suggest that increasing herbivore populations on degraded reefs may be an effective strategy for restoring ecosystem structure and function and in reversing coral–algal phase-shifts but that this strategy may be most effective in the absence of other confounding disturbances such as nutrient pollution.     

Coral recruitment and juvenile mortality as structuring factors for reef benthic communities in Biscayne National Park, USA

Coral communities of Biscayne National Park (BNP) on offshore linear bank-barrier reefs are depauperate of reef corals and have little topographic relief, while those on lagoonal patch reefs have greater coral cover and species richness despite presumably more stressful environmental regimes closer to shore. We hypothesized that differences in rates of coral recruitment and/or of coral survivorship were responsible for these differences in community structure. These processes were investigated by measuring: (1) juvenile and adult coral densities, and (2) size-frequency distributions of smaller coral size classes, at three pairs of bank- and patch-reefs distributed along the north-south range of coral reefs within the Park. In addition, small quadrats (0.25 m²) were censused for colonies <2 cm in size on three reefs (one offshore and one patch reef in the central park, and one intermediate reef at the southern end), and re-surveyed after 1 year. Density and size frequency data confirmed that large coral colonies were virtually absent from the offshore reefs, but showed that juvenile corals were common and had similar densities to those of adjacent bank and patch reefs. Large coral colonies were more common on inshore patch reefs, suggesting lower survivorship (higher mortality) of small and intermediate sized colonies on the offshore reefs. The more limited small-quadrat data showed similar survivorship rates and initial and final juvenile densities at all three sites, but a higher influx of new recruits to the patch reef site during the single annual study period. We consider the size-frequency data to be a better indicator of juvenile coral dynamics, since it is a more time-integrated measurement and was replicated at more sites. We conclude that lack of recruitment does not appear to explain the impoverished coral communities on offshore bank reefs in BNP. Instead, higher juvenile coral mortality appears to be a dominant factor structuring these communities. Accepted: 9 September 1999     

The dynamics of architectural complexity on coral reefs under climate change

One striking feature of coral reef ecosystems is the complex benthic architecture which supports diverse and abundant fauna, particularly of reef fish. Reef‐building corals are in decline worldwide, with a corresponding loss of live coral cover resulting in a loss of architectural complexity. Understanding the dynamics of the reef architecture is therefore important to envision the ability of corals to maintain functional habitats in an era of climate change. Here, we develop a mechanistic model of reef topographical complexity for contemporary Caribbean reefs. The model describes the dynamics of corals and other benthic taxa under climate‐driven disturbances (hurricanes and coral bleaching). Corals have a simplified shape with explicit diameter and height, allowing species‐specific calculation of their colony surface and volume. Growth and the mechanical (hurricanes) and biological erosion (parrotfish) of carbonate skeletons are important in driving the pace of extension/reduction in the upper reef surface, the net outcome being quantified by a simple surface roughness index (reef rugosity). The model accurately simulated the decadal changes of coral cover observed in Cozumel (Mexico) between 1984 and 2008, and provided a realistic hindcast of coral colony‐scale (1–10 m) changing rugosity over the same period. We then projected future changes of Caribbean reef rugosity in response to global warming. Under severe and frequent thermal stress, the model predicted a dramatic loss of rugosity over the next two or three decades. Critically, reefs with managed parrotfish populations were able to delay the general loss of architectural complexity, as the benefits of grazing in maintaining living coral outweighed the bioerosion of dead coral skeletons. Overall, this model provides the first explicit projections of reef rugosity in a warming climate, and highlights the need of combining local (protecting and restoring high grazing) to global (mitigation of greenhouse gas emissions) interventions for the persistence of functional reef habitats.     

In-Situ Effects of Simulated Overfishing and Eutrophication on Benthic Coral Reef Algae Growth,Succession, and Composition in the Central Red Sea

Overfishing and land-derived eutrophication are major local threats to coral reefs and may affect benthic communities, moving them from coral dominated reefs to algal dominated ones. The Central Red Sea is a highly under-investigated area, where healthy coral reefs are contending against intense coastal development. This in-situ study investigated both the independent and combined effects of manipulated inorganic nutrient enrichment (simulation of eutrophication) and herbivore exclosure (simulation of overfishing) on benthic algae development. Light-exposed and shaded terracotta tiles were positioned at an offshore patch reef close to Thuwal, Saudi Arabia and sampled over a period of 4 months. Findings revealed that nutrient enrichment alone affected neither algal dry mass nor algae-derived C or N production. In contrast, herbivore exclusion significantly increased algal dry mass up to 300-fold, and in conjunction with nutrient enrichment, this total increased to 500-fold. Though the increase in dry mass led to a 7 and 8-fold increase in organic C and N content, respectively, the algal C/N ratio (18±1) was significantly lowered in the combined treatment relative to controls (26±2). Furthermore, exclusion of herbivores significantly increased the relative abundance of filamentous algae on the light-exposed tiles and reduced crustose coralline algae and non-coralline red crusts on the shaded tiles. The combination of the herbivore exclusion and nutrient enrichment treatments pronounced these effects. The results of our study suggest that herbivore reduction, particularly when coupled with nutrient enrichment, favors non-calcifying, filamentous algae growth with high biomass production, which thoroughly outcompetes the encrusting (calcifying) algae that dominates in undisturbed conditions. These results suggest that the healthy reefs of the Central Red Sea may experience rapid shifts in benthic community composition with ensuing effects for biogeochemical cycles if anthropogenic impacts, particularly overfishing, are not controlled.     

Variable Responses of Benthic Communities to Anomalously Warm Sea Temperatures on a High-Latitude Coral Reef

High-latitude reefs support unique ecological communities occurring at the biogeographic boundaries between tropical and temperate marine ecosystems. Due to their lower ambient temperatures, they are regarded as potential refugia for tropical species shifting poleward due to rising sea temperatures. However, acute warming events can cause rapid shifts in the composition of high-latitude reef communities, including range contractions of temperate macroalgae and bleaching-induced mortality in corals. While bleaching has been reported on numerous high-latitude reefs, post-bleaching trajectories of benthic communities are poorly described. Consequently, the longer-term effects of thermal anomalies on high-latitude reefs are difficult to predict. Here, we use an autonomous underwater vehicle to conduct repeated surveys of three 625 m² plots on a coral-dominated high-latitude reef in the Houtman Abrolhos Islands, Western Australia, over a four-year period spanning a large-magnitude thermal anomaly. Quantification of benthic communities revealed high coral cover (>70%, comprising three main morphospecies) prior to the bleaching event. Plating Montipora was most susceptible to bleaching, but in the plot where it was most abundant, coral cover did not change significantly because of post-bleaching increases in branching Acropora. In the other two plots, coral cover decreased while macroalgal cover increased markedly. Overall, coral cover declined from 73% to 59% over the course of the study, while macroalgal cover increased from 11% to 24%. The significant differences in impacts and post-bleaching trajectories among plots underline the importance of understanding the underlying causes of such variation to improve predictions of how climate change will affect reefs, especially at high-latitudes.