Accounting for Multiple Foundation Species in Oyster Reef Restoration Benefits

Many coastal habitat restoration projects are focused on restoring the population of a single foundation species to recover an entire ecological community. Estimates of the ecosystem services provided by the restoration project are used to justify, prioritize, and evaluate such projects. However, estimates of ecosystem services provided by a single species may vastly under‐represent true provisioning, as we demonstrate here with an example of oyster reefs, often restored to improve estuarine water quality. In the brackish Chesapeake Bay, the hooked mussel Ischadium recurvum can have greater abundance and biomass than the focal restoration species, the eastern oyster Crassostrea virginica. We measured the temperature‐dependent phytoplankton clearance rates of both bivalves and their filtration efficiency on three size classes of phytoplankton to parameterize an annual model of oyster reef filtration, with and without hooked mussels, for monitored oyster reefs and restoration scenarios in the eastern Chesapeake Bay. The inclusion of filtration by hooked mussels increased the filtration capacity of the habitat greater than 2‐fold. Hooked mussels were also twice as effective as oysters at filtering picoplankton (1.5–3 µm), indicating that they fill a distinct ecological niche by controlling phytoplankton in this size class, which makes up a significant proportion of the phytoplankton load in summer. When mussel and oyster filtration are accounted for in this, albeit simplistic, model, restoration of oyster reefs in a tributary scale restoration is predicted to control 100% of phytoplankton during the summer months.     

Fish attraction to artificial reefs not always harmful: a simulation study

The debate on whether artificial reefs produce new fish or simply attract existing fish biomass continues due to the difficulty in distinguishing these processes, and there remains considerable doubt as to whether artificial reefs are a harmful form of habitat modification. The harm typically associated with attraction is that fish will be easier to harvest due to the existing biomass aggregating at a newly deployed reef. This outcome of fish attraction has not progressed past an anecdotal form, however, and is always perceived as a harmful process. We present a numerical model that simulates the effect that a redistributed fish biomass, due to an artificial reef, has on fishing catch per unit effort (CPUE). This model can be used to identify the scenarios (in terms of reef, fish, and harvest characteristics) that pose the most risk of exploitation due to fish attraction. The properties of this model were compared to the long‐standing predictions by Bohnsack (1989) on the factors that increase the risk or the harm of attraction. Simulations revealed that attraction is not always harmful because it does not always increase maximum fish density. Rather, attraction sometimes disperses existing fish biomass making them harder to catch. Some attraction can be ideal, with CPUE lowest when attraction leads to an equal distribution of biomass between natural and artificial reefs. Simulations also showed that the outcomes from attraction depend on the characteristics of the target fish species, such that transient or pelagic species are often at more risk of harmful attraction than resident species. Our findings generally agree with Bohnsack's predictions, although we recommend distinguishing “mobility” and “fidelity” when identifying species most at risk from attraction, as these traits had great influence on patterns of harvest of attracted fish biomass.     

Guidelines for evaluating performance of oyster habitat restoration

Restoration of degraded ecosystems is an important societal goal, yet inadequate monitoring and the absence of clear performance metrics are common criticisms of many habitat restoration projects. Funding limitations can prevent adequate monitoring, but we suggest that the lack of accepted metrics to address the diversity of restoration objectives also presents a serious challenge to the monitoring of restoration projects. A working group with experience in designing and monitoring oyster reef projects was used to develop standardized monitoring metrics, units, and performance criteria that would allow for comparison among restoration sites and projects of various construction types. A set of four universal metrics (reef areal dimensions, reef height, oyster density, and oyster size–frequency distribution) and a set of three universal environmental variables (water temperature, salinity, and dissolved oxygen) are recommended to be monitored for all oyster habitat restoration projects regardless of their goal(s). In addition, restoration goal‐based metrics specific to four commonly cited ecosystem service‐based restoration goals are recommended, along with an optional set of seven supplemental ancillary metrics that could provide information useful to the interpretation of prerestoration and postrestoration monitoring data. Widespread adoption of a common set of metrics with standardized techniques and units to assess well‐defined goals not only allows practitioners to gauge the performance of their own projects but also allows for comparison among projects, which is both essential to the advancement of the field of oyster restoration and can provide new knowledge about the structure and ecological function of oyster reef ecosystems.     

Interactions between coral restoration and fish assemblages: implications for reef management

Corals create complex reef structures that provide both habitat and food for many fish species. Because of numerous natural and anthropogenic threats, many coral reefs are currently being degraded, endangering the fish assemblages they support. Coral reef restoration, an active ecological management tool, may help reverse some of the current trends in reef degradation through the transplantation of stony corals. Although restoration techniques have been extensively reviewed in relation to coral survival, our understanding of the effects of adding live coral cover and complexity on fishes is in its infancy with a lack of scientifically validated research. This study reviews the limited data on reef restoration and fish assemblages, and complements this with the more extensive understanding of complex interactions between natural reefs and fishes and how this might inform restoration efforts. It also discusses which key fish species or functional groups may promote, facilitate or inhibit restoration efforts and, in turn, how restoration efforts can be optimised to enhance coral fish assemblages. By highlighting critical knowledge gaps in relation to fishes and restoration interactions, the study aims to stimulate research into the role of reef fishes in restoration projects. A greater understanding of the functional roles of reef fishes would also help inform whether restoration projects can return fish assemblages to their natural compositions or whether alternative species compositions develop, and over what timeframe. Although alleviation of local and global reef stressors remains a priority, reef restoration is an important tool; an increased understanding of the interactions between replanted corals and the fishes they support is critical for ensuring its success for people and nature.     

Comparisons of macrofaunal assemblages on restored and non-restored oyster reefs in mesohaline regions of Chesapeake Bay in Maryland

Maryland's recently created oyster restored reefs provide us with a unique opportunity to observe the abundance and species composition of macrofauna assemblages on unexploited reefs with high concentrations of mature oysters and undisturbed reef architecture. They might thus be used to better understand the magnitude of losses to reef dwelling macrofauna communities, and the associated loss of ecological functions resulting from reef destruction. We sampled reef macrofaunal assemblages on restored plots at four restored oyster reefs and adjacent non-restored plots located outside restored boundaries. We then compared the effects of study site location, and habitat quality (restored versus non-restored) on macrofaunal density using thirteen response variables. Density of macrofauna was an order of magnitude higher on restored reefs, epifaunal density was more than twice as high on restored reefs and sessile macrofaunal density was two orders of magnitude higher on restored reefs. Three out of the five dominant taxonomic groups were much more abundant on restored plots. Mean amphipod density was 20 times higher on restored plots and densities of xanthid crabs and demersal fish were both four times greater on restored plots. Two out of four functional feeding groups: suspension feeders and carnivore/omnivores, were more abundant on restored plots. Since reef macrofauna include many important fish prey species, oyster reef restoration may have the potential to augment fish production by increasing fish prey densities and fish foraging efficiency.     

Effects of time and harvest on genetic diversity of natural and restored oyster reefs

Oysters serve as keystone species and ecosystem engineers in estuaries due to their fundamental role of providing services to the surrounding environment and to humans. Globally, however, oysters have precipitously declined in numbers over the last century. To remedy this drastic decrease, many coastal areas have initiated oyster restoration projects. In the Indian River Lagoon (IRL) of Florida, where oyster loss was primarily the result of recreational boat wake dislodgment, researchers have put in place a unique method to supplement natural recruitment of oysters. This method consists of deploying stabilized shell attached to mesh mats. Larval oysters thus have substrate on which to settle and three‐dimensional reef habitats have been reestablished in historical footprints. This restoration project has proven to be successful, shown by 9 years of data collection on growth, recruitment, and survivability. In this study, we sought to determine the length of time required for newly restored oyster reefs to reach equivalent levels of genetic diversity as undisturbed, natural (reference) oyster reefs. Additionally, we determined if recreational harvesting impacted the genetic diversity of these reference reefs. Using nine microsatellite loci, we found that restored oyster reefs accumulated as much genetic diversity as natural reefs as quickly as 1 month after stabilized shells were deployed. We likewise found that harvesting did not impact genetic diversity in oyster reefs in the IRL. These results are encouraging, and are a valuable addition to understanding the importance of oyster reef restoration on the ecosystem.     

Diet and Habitat use by Bluefish, Pomatomus Saltatrix, in a Chesapeake Bay Estuary

Bluefish, Pomatomus saltatrix, are recreationally valuable finfish along the Atlantic seaboard and in the Chesapeake Bay. Diet and habitat use patterns for bluefish life history intervals in Chesapeake Bay estuaries are poorly described although it is widely acknowledged that this apex piscivorous species relies on estuarine habitat for feeding and nursery grounds after oceanic spawning and inshore migration of larvae. Bluefish diet, distribution, and abundance patterns were examined in relation to oyster reef, oyster bar, and sand bottom habitat in the Piankatank River, Virginia. Bluefish from all sites were predominantly piscivorous and were more abundant at reef sites than non-reef sites. Bluefish caught in association with the oyster reef consumed a wider diversity of prey items than fish from other sites; diet diversity may contribute to bluefish success during periods when small pelagic food fish abundance is reduced. Bluefish estuarine habitat use is positively correlated with the presence of oyster shell habitat and the complex trophic communities centering on oyster reefs.     

Opening of an MPA to fishing: Natural variations in the structure of a coral reef fish assemblage obscure changes due to fishing

Marine Protected Areas (MPAs) are known to enhance diversity, density and biomass of coral reef fishes and to modify the size and trophic structures of these fish assemblages. Opening to fishing has the opposite effects, but on a much shorter time scale. The present study compares the evolution of the fish assemblages of two adjacent reef zones, both initially MPAs, one of them being afterwards opened to fishing. The study was conducted on Aboré Reef, a New Caledonian barrier reef (SW Pacific) which constituted a 148 km² marine protected area, of which 69 km² are within the lagoon. Two surveys of a coral reef fish assemblage, using underwater visual censuses, were performed, the first one was conducted in July 1993 following 5 years of protection from fishing, the second one was conducted in July 1995; part of the reef having been opened to fishing activity in September 1993. This study examined the effects of two factors on these fish communities: time (1993 vs. 1995) and zone (reefs protected from fishing vs. unprotected reefs); the interaction of these two factors indicating an effect of either protection or opening to fishing. Diversity (species/transect), density and biomass were tested for all species together (377 species), then according to diet, size and commercial use. There was a significant decrease over time of most values in both fished and unfished areas, the decrease being greater in the zone opened to fishing. The magnitude of decrease over time was within the range of known time variations from other studies in New Caledonia and other Pacific locations. However, this decrease was so large that it prevented the detection of effects linked to protection. Only some minor effects could be detected for 16 species with no specific pattern according to diet, size or use. The density and biomass of species of low commercial value were also affected by opening to fishing. Relative changes in diversity could be better detected than relative changes in density or biomass. This study demonstrates that on a short-term basis (2 years), natural variations can be of larger magnitude than changes that may be induced by management options, especially when fishing pressure is not very high.     

A restoration suitability index model for the eastern oyster (Crassostrea virginica) in the Mission-Aransas Estuary, TX, USA

Oyster reefs are one of the most threatened marine habitats on earth, with habitat loss resulting from water quality degradation, coastal development, destructive fishing practices, overfishing, and storm impacts. For successful and sustainable oyster reef restoration efforts, it is necessary to choose sites that support long-term growth and survival of oysters. Selection of suitable sites is critically important as it can greatly influence mortality factors and may largely determine the ultimate success of the restoration project. The application of Geographic Information Systems (GIS) provides an effective methodology for identifying suitable sites for oyster reef restoration and removes much of the uncertainty involved in the sometimes trial and error selection process. This approach also provides an objective and quantitative tool for planning future oyster reef restoration efforts. The aim of this study was to develop a restoration suitability index model and reef quality index model to characterize locations based on their potential for successful reef restoration within the Mission-Aransas Estuary, Texas, USA. The restoration suitability index model focuses on salinity, temperature, turbidity, dissolved oxygen, and depth, while the reef quality index model focuses on abundance of live oysters, dead shell, and spat. Size-specific Perkinsus marinus infection levels were mapped to illustrate general disease trends. This application was effective in identifying suitable sites for oyster reef restoration, is flexible in its use, and provides a mechanism for considering alternative approaches. The end product is a practical decision-support tool that can be used by coastal resource managers to improve oyster restoration efforts. As oyster reef restoration activities continue at small and large-scales, site selection criteria are critical for assisting stakeholders and managers and for maximizing long-term sustainability of oyster resources.     

Rapid development of a restored oyster reef facilitates habitat provision for estuarine fauna

Oyster reef restoration has become a principal strategy for ameliorating the loss of natural Crassostrea virginica populations and increasing habitat provision. In 2014, a large‐scale, high‐relief, 23‐ha subtidal C. virginica reef was restored at the historically productive Half Moon Reef in Matagorda Bay, TX, using concrete and limestone substrates. Encrusting and motile fauna were sampled seasonally until 17 months postrestoration at the restored reef and at adjacent unrestored sites. Restored oysters developed rapidly and were most abundant 3 months postrestoration, with subsequent declines possibly due to interacting effects of larval settlement success on new substrate versus post‐settlement mortality due to competitors and predators. Oyster densities were 2× higher than in a restored oyster population in Chesapeake Bay that was reported to be the largest reestablished metapopulation of native oysters in the world. Resident fauna on the restored reef were 62% more diverse, had 433% greater biomass, and comprised a distinct faunal community compared to unrestored sites. The presence of three‐dimensional habitat was the most important factor determining resident faunal community composition, indicating that substrate limitation is a major hindrance for oyster reef community success in Texas and other parts of the Gulf of Mexico. There were only minor differences in density, biomass, and diversity of associated fauna located adjacent (13 m) versus distant (150 m) to the restored reef. The two substrate types compared had little influence on oyster recruitment or faunal habitat provision. Results support the use of reef restoration as a productive means to rebuild habitat and facilitate faunal enhancement.     

Effects of marine reserves versus nursery habitat availability on structure of reef fish communities

No-take marine fishery reserves sustain commercial stocks by acting as buffers against overexploitation and enhancing fishery catches in adjacent areas through spillover. Likewise, nursery habitats such as mangroves enhance populations of some species in adjacent habitats. However, there is lack of understanding of the magnitude of stock enhancement and the effects on community structure when both protection from fishing and access to nurseries concurrently act as drivers of fish population dynamics. In this study we test the separate as well as interactive effects of marine reserves and nursery habitat proximity on structure and abundance of coral reef fish communities. Reserves had no effect on fish community composition, while proximity to nursery habitat only had a significant effect on community structure of species that use mangroves or seagrass beds as nurseries. In terms of reef fish biomass, proximity to nursery habitat by far outweighed (biomass 249% higher than that in areas with no nursery access) the effects of protection from fishing in reserves (biomass 21% lower than non-reserve areas) for small nursery fish (≤ 25 cm total length). For large-bodied individuals of nursery species (>25 cm total length), an additive effect was present for these two factors, although fish benefited more from fishing protection (203% higher biomass) than from proximity to nurseries (139% higher). The magnitude of elevated biomass for small fish on coral reefs due to proximity to nurseries was such that nursery habitats seem able to overrule the usually positive effects on fish biomass by reef reserves. As a result, conservation of nursery habitats gains importance and more consideration should be given to the ecological processes that occur along nursery-reef boundaries that connect neighboring ecosystems.     

Marine reserves and reproductive biomass: a case study of a heavily targeted reef fish

Recruitment overfishing (the reduction of a spawning stock past a point at which the stock can no longer replenish itself) is a common problem which can lead to a rapid and irreversible fishery collapse. Averting this disaster requires maintaining a sufficient spawning population to buffer stochastic fluctuations in recruitment of heavily harvested stocks. Optimal strategies for managing spawner biomass are well developed for temperate systems, yet remain uncertain for tropical fisheries, where the danger of collapse from recruitment overfishing looms largest. In this study, we explored empirically and through modeling, the role of marine reserves in maximizing spawner biomass of a heavily exploited reef fish, Lethrinus harak around Guam, Micronesia. On average, spawner biomass was 16 times higher inside the reserves compared with adjacent fished sites. Adult density and habitat-specific mean fish size were also significantly greater. We used these data in an age-structured population model to explore the effect of several management scenarios on L. harak demography. Under minimum-size limits, unlimited extraction and all rotational-closure scenarios, the model predicts that preferential mortality of larger and older fish prompt dramatic declines in spawner biomass and the proportion of male fish, as well as considerable declines in total abundance. For rotational closures this occurred because of the mismatch between the scales of recovery and extraction. Our results highlight how alternative management scenarios fall short in comparison to marine reserves in preserving reproductively viable fish populations on coral reefs.     

Human,Oceanographic and Habitat Drivers of Central and Western Pacific Coral Reef Fish Assemblages

Coral reefs around US- and US-affiliated Pacific islands and atolls span wide oceanographic gradients and levels of human impact. Here we examine the relative influence of these factors on coral reef fish biomass, using data from a consistent large-scale ecosystem monitoring program conducted by scientific divers over the course of >2,000 hours of underwater observation at 1,934 sites, across ~40 islands and atolls. Consistent with previous smaller-scale studies, our results show sharp declines in reef fish biomass at relatively low human population density, followed by more gradual declines as human population density increased further. Adjusting for other factors, the highest levels of oceanic productivity among our study locations were associated with more than double the biomass of reef fishes (including ~4 times the biomass of planktivores and piscivores) compared to islands with lowest oceanic productivity. Our results emphasize that coral reef areas do not all have equal ability to sustain large reef fish stocks, and that what is natural varies significantly amongst locations. Comparisons of biomass estimates derived from visual surveys with predicted biomass in the absence of humans indicated that total reef fish biomass was depleted by 61% to 69% at populated islands in the Mariana Archipelago; by 20% to 78% in the Main Hawaiian islands; and by 21% to 56% in American Samoa.     

Faunal utilization of constructed intertidal oyster (Crassostrea rivularis) reef in the Yangtze River estuary,China

An intertidal oyster reef (～260 ha) was created by planting hatchery-reared seed oysters (Crassostrea rivularis) on an artificial concrete modular reef in the Deepwater Navigation Channel Regulation Project of the Yangtze River estuary. We examined the development of reef communities (oyster, barnacle and motile epibenthic macrofauna), characterized nekton use and assessed the habitat value of the constructed reef. The C. rivularis oyster population showed a rapid exponential increase with time and reached maximum density (3410 ± 241 ind./m²) and biomass (3175 ± 532 g/m²) after one year of restoration. The barnacle Balanus albicostatus was the most abundant sessile macrofauna and had a significantly greater density in the high intertidal zone than in the low intertidal zone (P < 0.05). The reef also supported diverse motile epibenthic macrofauna (11 mollusks, 11 crustaceans, 4 annelids and 2 fishes), and the reef-associated communities were numerically dominated by Neanthes japonica, Perinereis aibuhitensis, Nerita yoldi and Littorinopsis intermedia. A total of 50 nekton species (31 fishes, 9 shrimps and 10 crabs) utilized the constructed intertidal oyster reef, and grass shrimp Palaemon spp. dominated the nekton communities in term of abundance. Since the constructed intertidal oyster reef supports a variety of reef communities and abundant nektons, it should be recognized as an important and protective fish habitat in the Yangtze River estuary.     

江苏海门蛎岈山牡蛎礁生态现状评价

基于2013—2014年间的生态调查结果,评价了江苏海门蛎岈山牡蛎礁的生态现状。无人机航拍结果显示,江苏海门蛎岈山分布有750个潮间带区牡蛎礁斑块,总面积约为201519.37 m2;与2003年相比,海门蛎岈山牡蛎礁面积约下降了38.8%。活体牡蛎的平均盖度约为66%,2013年5和9月熊本牡蛎Crassostrea sikamea的平均密度分别为(2199±363)个/m2和(2894±330)个/m2。2013年5月海门蛎岈山熊本牡蛎种群的平均肥满度(CI)和性腺指数(GI)分别为(9.76±0.95)%和(1013±82)mg/g,均显著低于浙江象山港养殖的熊本牡蛎种群(P0.05)。海门蛎岈山熊本牡蛎的单倍体多样性和核苷酸多样性指数分别为0.119和0.00028,均高于长江口野生种群和浙江象山港养殖种群。海门蛎岈山熊本牡蛎种群受到尼氏单孢子虫(Haplosporidium nelson)的轻度浸染,其感染率(17.2%)低于浙江象山港养殖群体(47.3%)。泥沙沉积和人类捕捞是江苏海门蛎岈山牡蛎礁面临的主要胁迫因子,今后牡蛎礁恢复的重点是增加附着底物的数量。     

Edge effects influence the composition and density of reef residents on subtidal restored oyster reefs

Within estuarine and coastal ecosystems globally, extensive habitat degradation and loss threaten critical ecosystem functions and necessitate widescale restoration efforts. There is abundant evidence that ecological processes and species interactions can vary with habitat characteristics, which has important implications for the design and implementation of restoration efforts aimed at enhancing specific ecosystem functions and services. We conducted an experiment examining how habitat characteristics (presence; edge vs. interior) influence the communities of resident fish and mobile invertebrates on restored oyster (Crassostrea virginica) reefs. Similar to previous studies, we found that restored reefs altered community composition and augmented total abundance and biomass relative to unstructured sand habitat. Community composition and biomass also differed between the edge and interior of individual reefs as a result of species-specific patterns over small spatial scales. These patterns were only weakly linked to oyster density, suggesting that other factors that vary between edge and interior (e.g. predator access or species interactions) are likely more important for community structure on oyster reefs. Fine-scale information on resident species' use of oyster reefs will help facilitate restoration by allowing decision makers to optimize the amount of edge versus interior habitat. To improve the prediction of faunal use and benefits from habitat restoration, we recommend investigations into the mechanisms shaping edge and interior preferences on oyster reefs.     

Evaluating Intertidal Oyster Reef Development in South Carolina Using Associated Faunal Indicators

Eastern oyster (Crassostrea virginica) habitat is increasingly being restored for the ecosystem services it provides rather than solely as a fishery resource. Community‐based projects with the goal of ecological restoration have successfully constructed oyster reefs; however, the habitat benefits of these restoration efforts are usually not assessed or reported. In this study, we examined oyster habitat development at five community‐based oyster restoration sites in South Carolina using oyster population parameters, resident fauna densities, and sedimentation (percent sediment coverage) as assessment metrics. All sites included multiple‐aged reefs (1–3 years old) at the time of the fall 2004 sampling. Resident crabs and mussels were abundant at all five sites and crab assemblages were related to the size structure of the oyster microhabitat. Scorched mussel (Brachidontes exustus) abundances were most frequently correlated with oyster and other resident species abundances. Associations among oysters and resident crabs and mussels were not evident when analyses were conducted with higher level taxonomic groupings (e.g., total number of crabs, mussels, or oysters), indicating that species‐level identifications improve our understanding of interactions among reef inhabitants and oyster populations. Community‐based restoration sites in South Carolina provide habitat for mussels and resident crabs, in some cases in the absence of dense populations of relatively large oysters. Monitoring programs that neglect species‐level identifications and counts of mussels and crabs may underestimate the successful habitat provision that can arise independent of large, dense oyster assemblages.     

Determinants of reef fish assemblages in tropical Oceanic islands

Diversity patterns are determined by biogeographic, energetic, and anthropogenic factors, yet few studies have combined them into a large‐scale framework in order to decouple and compare their relative effects on fish faunas. Using an empirical dataset derived from 1527 underwater visual censuses (UVC) at 18 oceanic islands (five different marine provinces), we determined the relative influence of such factors on reef fish species richness, functional dispersion, density and biomass estimated from each UVC unit. Species richness presented low variation but was high at large island sites. High functional dispersion, density, and biomass were found at islands with large local species pool and distance from nearest reef. Primary productivity positively affected fish richness, density and biomass confirming that more productive areas support larger populations, and higher biomass and richness on oceanic islands. Islands densely populated by humans had lower fish species richness and biomass reflecting anthropogenic effects. Species richness, functional dispersion, and biomass were positively related to distance from the mainland. Overall, species richness and fish density were mainly influenced by biogeographical and energetic factors, whereas functional dispersion and biomass were strongly influenced by anthropogenic factors. Our results extend previous hypotheses for different assemblage metrics estimated from empirical data and confirm the negative impact of humans on fish assemblages, highlighting the need for conservation of oceanic islands.     

Guidelines for evaluating performance of oyster habitat restoration should include tidal emersion: reply to Baggett et al.

Baggett et al. (2015) identified a set of three universal environmental variables to be monitored for evaluating all oyster habitat restoration projects: salinity, temperature, and dissolved oxygen. Perhaps evidencing a bias toward subtidal reefs, this set of parameters omits another first‐order environmental factor, tidal emersion. Intertidal oyster reefs can be the dominant reef habitat in estuaries, with clear zonation in oyster performance across the intertidal exposure gradient. Therefore, we propose to include tidal emersion as a fourth universal environmental parameter when designing and evaluating oyster restoration projects to better encompass the whole environmental spectrum along which reefs occur.