Habitat complexity and consumer-mediated positive feedbacks on a Caribbean coral reef

Theoretical and empirical evidence suggest that positive feedbacks can increase resilience in ecological communities. On Caribbean coral reefs, there have been striking shifts from physically complex communities with high coral cover to relatively homogenous communities dominated by macroalgae, which have persisted for decades. However, little is known about positive feedbacks that may maintain coral reef community states. Here, I explore a potential consumer-mediated feedback on a Jamaican reef by examining how grazing by a keystone herbivore ( Diadema antillarum ) is enhanced by physical structure, which offer refugia from predation. Surveys revealed that habitat complexity and Diadema density were positively related. Increasing habitat complexity by adding physical structure significantly decreased macroalgal cover and increased the proportion of urchins in algal habitats in field manipulations. Experimental increases in urchin density also decreased macroalgal cover, but did not affect the proportion of urchins in algal habitats. These results suggest that the low habitat complexity of macroalgal-dominated reefs may inhibit an urchin-mediated shift to coral dominance and that positive feedbacks must be considered in reef restoration efforts.     

Microbial to reef scale interactions between the reef-building coral Montastraea annularis and benthic algae

Competition between reef-building corals and benthic algae is of key importance for reef dynamics. These interactions occur on many spatial scales, ranging from chemical to regional. Using microprobes, 16S rDNA pyrosequencing and underwater surveys, we examined the interactions between the reef-building coral Montastraea annularis and four types of benthic algae. The macroalgae Dictyota bartayresiana and Halimeda opuntia, as well as a mixed consortium of turf algae, caused hypoxia on the adjacent coral tissue. Turf algae were also associated with major shifts in the bacterial communities at the interaction zones, including more pathogens and virulence genes. In contrast to turf algae, interactions with crustose coralline algae (CCA) and M. annularis did not appear to be antagonistic at any scale. These zones were not hypoxic, the microbes were not pathogen-like and the abundance of coral-CCA interactions was positively correlated with per cent coral cover. We propose a model in which fleshy algae (i.e. some species of turf and fleshy macroalgae) alter benthic competition dynamics by stimulating bacterial respiration and promoting invasion of virulent bacteria on corals. This gives fleshy algae a competitive advantage over corals when human activities, such as overfishing and eutrophication, remove controls on algal abundance. Together, these results demonstrate the intricate connections and mechanisms that structure coral reefs.     

40 Years of benthic community change on the Caribbean reefs of Curaçao and Bonaire: the rise of slimy cyanobacterial mats

Over the past decades numerous studies have reported declines in stony corals and, in many cases, phase shifts to fleshy macroalgae. However, long-term studies documenting changes in other benthic reef organisms are scarce. Here, we studied changes in cover of corals, algal turfs, benthic cyanobacterial mats, macroalgae, sponges and crustose coralline algae at four reef sites of the Caribbean islands of Curaçao and Bonaire over a time span of 40 yr. Permanent 9 m² quadrats at 10, 20, 30 and 40 m depth were photographed at 3- to 6-yr intervals from 1973 to 2013. The temporal and spatial dynamics in the six dominant benthic groups were assessed based on image point-analysis. Our results show consistent patterns of benthic community change with a decrease in the cover of calcifying organisms across all sites and depths from 32.6 (1973) to 9.2% (2013) for corals and from 6.4 to 1% for crustose coralline algae. Initially, coral cover was replaced by algal turfs increasing from 24.5 (1973) to 38% around the early 1990s. Fleshy macroalgae, still absent in 1973, also proliferated covering 12% of the substratum approximately 20 yr later. However, these new dominants largely declined in abundance from 2002 to 2013 (11 and 2%, respectively), marking the rise of benthic cyanobacterial mats. Cyanobacterial mats became the most dominant benthic component increasing from a mere 7.1 (2002) to 22.2% (2013). The observed increase was paralleled by a small but significant increase in sponge cover (0.5 to 2.3%). Strikingly, this pattern of degradation and phase change occurred over the reef slope down to mesophotic depths of 40 m. These findings suggest that reefs dominated by algae may be less stable than previously thought and that the next phase may be the dominance of slimy cyanobacterial mats with some sponges.     

Multiple anthropogenic stressors exert complex,interactive effects on a coral reef community

Multiple natural and anthropogenic stressors impact coral reefs across the globe leading to declines of coral populations, but the relative importance of different stressors and the ways they interact remain poorly understood. Because coral reefs exist in environments commonly impacted by multiple stressors simultaneously, understanding their interactions is of particular importance. To evaluate the role of multiple stressors we experimentally manipulated three stressors (herbivore abundance, nutrient supply, and sediment loading) in plots on a natural reef in the Gulf of Panamá in the Eastern Tropical Pacific. Monitoring of the benthic community (coral, macroalgae, algal turf, and crustose coralline algae) showed complex responses with all three stressors impacting the community, but at different times, in different combinations, and with varying effects on different community members. Reduction of top–down control in combination with sediment addition had the strongest effect on the community, and led to approximately three times greater algal biomass. Coral cover was reduced in all experimental units with a negative effect of nutrients over time and a synergistic interaction between herbivore exclosures and sediment addition. In contrast, nutrient and sediment additions interacted antagonistically in their impacts on crustose coralline algae and turf algae so that in combination the treatments limited each other’s effects. Interactions between stressors and temporal variability indicated that, while each stressor had the potential to impact community structure, their combinations and the broader environmental conditions under which they acted strongly influenced their specific effects. Thus, it is critical to evaluate the effects of stressors on community dynamics not only independently but also under different combinations or environmental conditions to understand how those effects will be played out in more realistic scenarios.     

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.     

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.     

Monitoring of the Meager Shoal coral reef, Cahuita National Park, Costa Rica (CARICOMP site)

The coral reefs at Cahuita National Park, Caribbean coast of Costa Rica, specifically at the CARICOMP site Meager Shoal, have been monitored since 1999. Complete data sets from 2000 and 2004 have shown that live coral cover has increased less than 3 % (from 15 to 17 %), but non-coralline algae cover has increased much (63 to 74 %) and coralline algae cover has decreased (17 to 5 %) significantly. The proportion of affected colonies by diseases, injuries and bleaching decreased from 24 % in 2000 to 10 % in 2004, but the difference was not statistically significant. Densities of the urchin Diadema antillarum increased, and are probably help to maintain the macroalgae biomass low, while those of Echinometra viridis decreased significantly. The coral reef at Cahuita National Park continues to be impacted by chronic terrigenous sediments and does not show a significant recovery since the late 1970's.     

The relationship between habitat structure and fish faunas on New Zealand reefs

Reef fish abundances were sampled at 11 shallow reef localities extending over 1000 km of coastline in northern New Zealand. Sampling was restricted to the 4–10-m depth stratum and included six coastal and five island localities. These were either coralline reef flats dominated by echinoids, or algal reefs with high densities of laminarian and fucoid algae. Reefs dominated by macroscopic algae supported large numbers of small fishes, mainly labrids, and few large benthic-feeding fishes. Echinoid-dominated reefs supported a different fish fauna with more large benthic-feeding species. Additional sampling of echinoiddominated reefs and algal stands in deeper water provided confirmation of these findings. A second sampling programme was carried out at a series of eight sites within a single locality covering 5 km of coastline. These spanned a moderate exposure gradient and ranged from algal dominated reefs to typical coralline reef flats with high densities of grazing invertebrates. The relationship between habitat structure and reef fish species composition and size frequency was similar to that of the large-scale sampling programme. Thirdly, observations on reef fish foraging and feeding patterns within a single reef site suggested that larger benthic-feeding reef fishes were less likely to feed within macroscopic algal stands. Experimental reductions of grazing invertebrates designed to produce brown algal stands on echinoid-dominated reef flats supported these observations. Larger individuals capable of removing echinoids and grazing gastropods did not frequent or feed in laminarian and fucoid algal stands. This pattern is discernible at several spatial scales. Our conclusion is that the type of shallow reef habitat, echinoid- as opposed to algal-dominated, will have an important rôle in determining the associated reef fish fauna.     

Responses of algal communities to gradients in herbivore biomass and water quality in Marovo Lagoon, Solomon Islands

Settlement tiles were used to characterise and quantify coral reef associated algal communities along water quality and herbivory gradients from terrestrial influenced near shore sites to oceanic passage sites in Marovo Lagoon, the Solomon Islands. After 6 months, settlement tile communities from inshore reefs were dominated by high biomass algal turfs (filamentous algae and cyanobacteria) whereas tiles located on offshore reefs were characterised by a mixed low biomass community of calcareous crustose algae, fleshy crustose algae and bare tile. The exclusion of macrograzers, via caging of tiles, on the outer reef sites resulted in the development of an algal turf community similar to that observed on inshore reefs. Caging on the inshore reef tiles had a limited impact on community composition or biomass. Water quality and herbivorous fish biomass were quantified at each site to elucidate factors that might influence algal community structure across the lagoon. Herbivore biomass was the dominant driver of algal community structure. Algal biomass on the other hand was controlled by both herbivory and water quality (particularly dissolved nutrients). This study demonstrates that algal communities on settlement tiles are an indicator capable of integrating the impacts of water quality and herbivory over a small spatial scale (kilometres) and short temporal scale (months), where other environmental drivers (current, light, regional variability) are constant.     

168 Cell Wall Proteomics of the Green Alga Haematococcus Pluvialis (Chlorophyceae)

Coral reefs of US-held islands in the central Pacific Ocean are among the most pristine in the world and represent over 93% of the reef systems under United States jurisdiction. The remote location of many islands has limited past algal research, resulting in incomplete understanding of species diversity, quantity, and ecology. Starting in 2000, the Coral Reef Ecosystem Investigation (CREI) began rapid ecological assessments on many Pacific island reefs to monitor ecological changes in reef biota over time. During the past year, algal efforts have concentrated on the French Frigate Shoals (Northwestern Hawaiian Islands) where we have increased the number of algal species reported by 1000%. Additionally, species new to science, including Acrosymphyton brainardii and Scinaia huismanii, have been described. Quantitative field sampling using a photoquadrat method is revealing species of the green algae Halimeda and Microdicyton to be ecological dominants in many areas during late summer/early autumn. Preliminary analyses with Primer software show species composition and abundance of all benthic organisms to differ significantly between most field sites sampled. Additional benthic habitat mapping of Pacific island reefs by CREI researchers is breaking the long-held paradigm that macroalgal cover is minimal in healthy tropical reef systems. Videotape analyses of benthic communities often find over 50% algal cover from 1 to 20 meter depths in many locations. Common ratios of macroalgae, turf algae, and crustose coralline algae to corals, other benthic organisms and substrate types on US Pacific reefs are being calculated for the first time.     

Large-scale associations between macroalgal cover and grazer biomass on mid-depth reefs in the Caribbean

Since the 1970s, macroalgae have become considerably more abundant on many Caribbean reefs and overfishing of grazing fishes has been implicated as a contributory factor. We explored relationships between algal cover and grazers (biomass of herbivorous fishes and abundance of the sea-urchin Diadema antillarum) on mid-depth reefs (12-15 m) in 19 areas at seven locations in Jamaica, Barbados, Belize, Grand Cayman and Cuba, between April 1997 and April 1998. Diadema antillarum density was never >0.01 m^-2, while herbivorous fish biomass (acanthurids and scarids ⁸ cm total length) varied from 2-5 g m^-2 in Jamaica to 17.1 g m^-2 in Barbados, and was strongly correlated, negatively with macroalgal cover and positively with 'cropped' substratum (sum of 'bare', turf and crustose-coralline substrata) cover. However, overfishing of herbivorous fishes alone cannot explain the widespread abundance of macroalgae, as even on lightly fished reefs, macroalgal cover was mostly >20%. Herbivorous fish populations on those reefs were apparently only able to maintain approximately 40-60% of reef substratum in cropped states, but due to low space-occupation by coral and other invertebrates, 70-90% of substratum was available to algae. The abundance of macroalgae on lightly fished reefs may therefore be a symptom of low coral cover in combination with the continuing absence of Diadema antillarum.     

167 Algal Monitoring Studies on Remote Tropical Pacific Reefs

Coral reefs of US‐held islands in the central Pacific Ocean are among the most pristine in the world and represent over 93% of the reef systems under United States jurisdiction. The remote location of many islands has limited past algal research, resulting in incomplete understanding of species diversity, quantity, and ecology. Starting in 2000, the Coral Reef Ecosystem Investigation (CREI) began rapid ecological assessments on many Pacific island reefs to monitor ecological changes in reef biota over time. During the past year, algal efforts have concentrated on the French Frigate Shoals (Northwestern Hawaiian Islands) where we have increased the number of algal species reported by 1000%. Additionally, species new to science, including Acrosymphyton brainardii and Scinaia huismanii, have been described. Quantitative field sampling using a photoquadrat method is revealing species of the green algae Halimeda and Microdicyton to be ecological dominants in many areas during late summer/early autumn. Preliminary analyses with Primer software show species composition and abundance of all benthic organisms to differ significantly between most field sites sampled. Additional benthic habitat mapping of Pacific island reefs by CREI researchers is breaking the long‐held paradigm that macroalgal cover is minimal in healthy tropical reef systems. Videotape analyses of benthic communities often find over 50% algal cover from 1 to 20 meter depths in many locations. Common ratios of macroalgae, turf algae, and crustose coralline algae to corals, other benthic organisms and substrate types on US Pacific reefs are being calculated for the first time.     

Re-evaluating the health of coral reef communities: baselines and evidence for human impacts across the central Pacific

Numerous studies have documented declines in the abundance of reef-building corals over the last several decades and in some but not all cases, phase shifts to dominance by macroalgae have occurred. These assessments, however, often ignore the remainder of the benthos and thus provide limited information on the present-day structure and function of coral reef communities. Here, using an unprecedentedly large dataset collected within the last 10 years across 56 islands spanning five archipelagos in the central Pacific, we examine how benthic reef communities differ in the presence and absence of human populations. Using islands as replicates, we examine whether benthic community structure is associated with human habitation within and among archipelagos and across latitude. While there was no evidence for coral to macroalgal phase shifts across our dataset we did find that the majority of reefs on inhabited islands were dominated by fleshy non-reef-building organisms (turf algae, fleshy macroalgae and non-calcifying invertebrates). By contrast, benthic communities from uninhabited islands were more variable but in general supported more calcifiers and active reef builders (stony corals and crustose coralline algae). Our results suggest that cumulative human impacts across the central Pacific may be causing a reduction in the abundance of reef builders resulting in island scale phase shifts to dominance by fleshy organisms.     

Local and regional scale recovery of Diadema promotes recruitment of scleractinian corals

The phase change from coral to macroalgal dominance on many Caribbean reefs was exacerbated by the mortality of the echinoid Diadema antillarum in 1983–1984, and until recently, this sea urchin has remained rare on reefs throughout the western Atlantic. By the late 1990s, Diadema started to reappear in large numbers on some Jamaican reefs, and by 2000, the high densities were correlated with significantly greater abundances of juvenile corals. Here, we show that dense populations of Diadema now occur over a multi-kilometre-wide scale at six locations scattered along a 4100 km arc across the entire Caribbean. In all cases, these dense populations are found in shallow water (< 6 m depth) on outer reef communities and are associated with reduced macroalgal cover and enhanced coral recruitment. We conclude that population recovery of Diadema is occurring at both local and regional scales, and that grazing by this echinoid is creating conditions favouring the recruitment of corals.     

Natural inducers for coral larval metamorphosis

 Coral gametes from Acropora millepora (Ehrenberg, 1834) and from multi-species spawning slicks provided larvae for use in metamorphosis assays with a selection of naturally occurring inducer chemicals. Four species of crustose coralline algae, one non-coralline crustose alga and two branching coralline algae induced larval metamorphosis. However, one additional species of branching coralline algae did not produce a larval response. Metamorphosis was also observed when larvae were exposed to skeleton from the massive coral Goniastrea retiformis (Lamarck, 1816) and to calcified reef rubble, demonstrating metamorphosis is possible in the absence of encrusting algae. Chemical extracts from these algae and the coral skeleton, obtained using either decalcification or simple methanol extraction procedures, also contained active inducers. These results extend the number of crustose algal species known to induce coral metamorphosis, suggest that some inducers may not necessarily be strongly associated with the calcified algal cell walls, and indicate that inducer sources in reef habitats may be more diverse than previously reported. Accepted: 21 May 1999     

Interactions between herbivorous fish guilds and their influence on algal succession on a coastal coral reef

Herbivory is an important mechanism affecting algal succession, particularly on coral reefs where the relationship between algae and corals is largely controlled by herbivores. However, different functional groups of herbivores may have contrasting effects on succession, which may explain different trajectories of coral reef recovery after disturbance. Here, the effects of different herbivore groups (roving herbivores = foragers and territorial damselfish = farmers) were isolated by a multi-factorial experiment carried out on a coastal coral reef with high macroalgal cover, high farmer densities and relatively low forager abundance. The effects of foragers and farmers were distinguished by monitoring algal succession on settlement tiles placed inside and outside exclusion cages, with orthogonal treatments established inside and outside damselfish territories (with appropriate cage controls). Within 12 months, algal assemblages on ungrazed tiles inside exclusion cages proceeded rapidly from fine filamentous turfs, to corticated algae, to tough erect (e.g. Amphiroa spp.) and foliose (e.g. Peyssonnellidae) calcified algae. Farmers had a dramatic impact on succession, essentially arresting the development of the algal community at a point where it was dominated by palatable filamentous algae of the genus Polysiphonia. Fleshy macroalgae such as Sargassum spp. were excluded from farmer territories. In contrast, foragers did not suppress fleshy macroalgae, but rather, appeared to decelerate succession and promote a relatively diverse assemblage. In contrast to forager-dominated reefs, farmer territories did not appear to function solely as forager exclusion areas or promote algal diversity as a result of intermediate grazing pressure. The relatively strong effects of farmers observed here may represent a future scenario for coral reefs that are increasingly subject to overfishing of large grazing fishes.     

Black reefs: iron-induced phase shifts on coral reefs

The Line Islands are calcium carbonate coral reef platforms located in iron-poor regions of the central Pacific. Natural terrestrial run-off of iron is non-existent and aerial deposition is extremely low. However, a number of ship groundings have occurred on these atolls. The reefs surrounding the shipwreck debris are characterized by high benthic cover of turf algae, macroalgae, cyanobacterial mats and corallimorphs, as well as particulate-laden, cloudy water. These sites also have very low coral and crustose coralline algal cover and are call black reefs because of the dark-colored benthic community and reduced clarity of the overlying water column. Here we use a combination of benthic surveys, chemistry, metagenomics and microcosms to investigate if and how shipwrecks initiate and maintain black reefs. Comparative surveys show that the live coral cover was reduced from 40 to 60% to <10% on black reefs on Millennium, Tabuaeran and Kingman. These three sites are relatively large (>0.75 km²). The phase shift occurs rapidly; the Kingman black reef formed within 3 years of the ship grounding. Iron concentrations in algae tissue from the Millennium black reef site were six times higher than in algae collected from reference sites. Metagenomic sequencing of the Millennium Atoll black reef-associated microbial community was enriched in iron-associated virulence genes and known pathogens. Microcosm experiments showed that corals were killed by black reef rubble through microbial activity. Together these results demonstrate that shipwrecks and their associated iron pose significant threats to coral reefs in iron-limited regions.     

Effects of coral reef benthic primary producers on dissolved organic carbon and microbial activity

Benthic primary producers in marine ecosystems may significantly alter biogeochemical cycling and microbial processes in their surrounding environment. To examine these interactions, we studied dissolved organic matter release by dominant benthic taxa and subsequent microbial remineralization in the lagoonal reefs of Moorea, French Polynesia. Rates of photosynthesis, respiration, and dissolved organic carbon (DOC) release were assessed for several common benthic reef organisms from the backreef habitat. We assessed microbial community response to dissolved exudates of each benthic producer by measuring bacterioplankton growth, respiration, and DOC drawdown in two-day dark dilution culture incubations. Experiments were conducted for six benthic producers: three species of macroalgae (each representing a different algal phylum: Turbinaria ornata – Ochrophyta; Amansia rhodantha – Rhodophyta; Halimeda opuntia – Chlorophyta), a mixed assemblage of turf algae, a species of crustose coralline algae (Hydrolithon reinboldii) and a dominant hermatypic coral (Porites lobata). Our results show that all five types of algae, but not the coral, exuded significant amounts of labile DOC into their surrounding environment. In general, primary producers with the highest rates of photosynthesis released the most DOC and yielded the greatest bacterioplankton growth; turf algae produced nearly twice as much DOC per unit surface area than the other benthic producers (14.0±2.8 µmol h⁻¹ dm⁻²), stimulating rapid bacterioplankton growth (0.044±0.002 log10 cells h⁻¹) and concomitant oxygen drawdown (0.16±0.05 µmol L⁻¹ h⁻¹ dm⁻²). Our results demonstrate that benthic reef algae can release a significant fraction of their photosynthetically-fixed carbon as DOC, these release rates vary by species, and this DOC is available to and consumed by reef associated microbes. These data provide compelling evidence that benthic primary producers differentially influence reef microbial dynamics and biogeochemical parameters (i.e., DOC and oxygen availability, bacterial abundance and metabolism) in coral reef communities.     

Hyperspectral and Physiological Analyses of Coral-Algal Interactions

Space limitation leads to competition between benthic, sessile organisms on coral reefs. As a primary example, reef-building corals are in direct contact with each other and many different species and functional groups of algae. Here we characterize interactions between three coral genera and three algal functional groups using a combination of hyperspectral imaging and oxygen microprofiling. We also performed in situ interaction transects to quantify the relative occurrence of these interaction on coral reefs. These studies were conducted in the Southern Line Islands, home to some of the most remote and near-pristine reefs in the world. Our goal was to determine if different types of coral-coral and coral-algal interactions were characterized by unique fine-scale physiological signatures. This is the first report using hyperspectral imaging for characterization of marine benthic organisms at the micron scale and proved to be a valuable tool for discriminating among different photosynthetic organisms. Consistent patterns emerged in physiology across different types of competitive interactions. In cases where corals were in direct contact with turf or macroalgae, there was a zone of hypoxia and altered pigmentation on the coral. In contrast, interaction zones between corals and crustose coralline algae (CCA) were not hypoxic and the coral tissue was consistent across the colony. Our results suggest that at least two main characteristic coral interaction phenotypes exist: 1) hypoxia and coral tissue disruption, seen with interactions between corals and fleshy turf and/or some species of macroalgae, and 2) no hypoxia or tissue disruption, seen with interactions between corals and some species of CCA. Hyperspectral imaging in combination with oxygen profiling provided useful information on competitive interactions between benthic reef organisms, and demonstrated that some turf and fleshy macroalgae can be a constant source of stress for corals, while CCA are not.     

A unique algal ridge system in the Exuma Cays, Bahamas

 An algal ridge system discovered along the Exuma Cays, Bahamas constructs bioherms to a thickness of at least 1.5 m and is associated with modern intertidal stromatolites. These algal ridges are unique because they grow in atypical environments characterized by relatively low wave energy, high rates of sedimentation and low rates of herbivory. They also are composed primarily of the branching crustose coralline alga, Neogoniolithon strictum, which heretofore was not known to form algal ridges. Lateral growth rates of crusts, vertical growth rates of branches and survivorship of transplanted N. strictum were greatest in the shallow fore reef zone of the algal ridge. The alga is also capable of surviving and growing when covered with sediment for at least 100 days. Under such conditions it transforms from a branched to an unbranched morphology. Parrotfish grazing, which is said to limit the abundance of branched corallines and algal ridges, was two orders of magnitude lower than in published accounts from other reef systems of the Caribbean and one order of magnitude less than that found on nearby coral reefs of the west Exuma Sound. Neogoniolithon strictum, a delicate and open-branched coralline, persisted for over a year without grazing damage when transplanted to a depth of 2.3 m. This algal ridge-building coralline becomes a well-indurated limestone following submarine lithification of sediment that infills the open branch framework. As a result, N. strictum ridges are comparable to the dense frameworks associated with most algal ridges. Observations of N. strictum -associated bioherms along Central America suggest this ridge system may exist elsewhere under conditions similar to those described for the Bahamas. Accepted: 23 March 1996