Major environmental drivers determining life and death of cold-water corals through time

Cold-water corals (CWCs) are the engineers of complex ecosystems forming unique biodiversity hotspots in the deep sea. They are expected to suffer dramatically from future environmental changes in the oceans such as ocean warming, food depletion, deoxygenation, and acidification. However, over the last decades of intense deep-sea research, no extinction event of a CWC ecosystem is documented, leaving quite some uncertainty on their sensitivity to these environmental parameters. Paleoceanographic reconstructions offer the opportunity to align the on- and offsets of CWC proliferation to environmental parameters. Here, we present the synthesis of 6 case studies from the North Atlantic Ocean and the Mediterranean Sea, revealing that food supply controlled by export production and turbulent hydrodynamics at the seabed exerted the strongest impact on coral vitality during the past 20,000 years, whereas locally low oxygen concentrations in the bottom water can act as an additional relevant stressor. The fate of CWCs in a changing ocean will largely depend on how these oceanographic processes will be modulated. Future ocean deoxygenation may be compensated regionally where the food delivery and food quality are optimal.

Cold-water corals are the engineers of complex ecosystems forming unique biodiversity hot spots in the deep sea, but there is substantial uncertainty regarding their sensitivity to environmental parameters. A synthesis of six paleoceanographic reconstructions reveals that food supply exerted the strongest impact on coral vitality during the past 20,000 years, whereas locally low oxygen concentrations can act as a stressor.     

Seasonal controls on the diet,metabolic activity,tissue reserves and growth of the cold-water coral Lophelia pertusa

Coral Reefs - Vast cold-water coral (CWC) reefs occur in temperate regions, where strong seasonality in temperature and light leads to a short but highly productive spring period. How CWCs respond...     

Tidal downwelling and implications for the carbon biogeochemistry of cold‐water corals in relation to future ocean acidification and warming

Cold‐water coral (CWC) reefs are recognized as ecologically and biologically significant areas that generate habitats and diversity. The interaction between hydrodynamics and CWCs has been well studied at the Mingulay Reef Complex, a relatively shallow area of reefs found on the continental shelf off Scotland, UK. Within ‘Mingulay Area 01’ a rapid tidal downwelling of surface waters, brought about as an internal wave, is known to supply warmer, phytoplankton‐rich waters to corals growing on the northern flank of an east‐west trending seabed ridge. This study shows that this tidal downwelling also causes short‐term perturbations in the inorganic carbon (C_T) and nutrient dynamics through the water column and immediately above the reef. Over a 14 h period, corresponding to one semi‐diurnal tidal cycle, seawater pH overlying the reef varied by ca. 0.1 pH unit, while pCO₂ shifted by >60 μatm, a shift equivalent to a ca. 25 year jump into the future, with respect to atmospheric pCO₂. During the summer stratified period, these downwelling events result in the reef being washed over with surface water that has higher pH, is warmer, nutrient depleted, but rich in phytoplankton‐derived particles compared to the deeper waters in which the corals sit. Empirical observations, together with outputs from the European Regional Shelf Sea Ecosystem Model, demonstrate that the variability that the CWC reefs experience changes through the seasons and into the future. Hence, as ocean acidification and warming increase into the future, the downwelling event specific to this site could provide short‐term amelioration of corrosive conditions at certain times of the year; however, it could additionally result in enhanced detrimental impacts of warming on CWCs. Natural variability in the C_T and nutrient conditions, as well as local hydrodynamic regimes, must be accounted for in any future predictions concerning the responses of marine ecosystems to climate change.     

Bottlenecks to coral recovery in the Seychelles

Processes that affect recovery of coral assemblages require investigation because coral reefs are experiencing a diverse array of more frequent disturbances. Potential bottlenecks to coral recovery include limited larval supply, low rates of settlement, and high mortality of new recruits or juvenile corals. We investigated spatial variation in local abundance of scleractinian corals in the Seychelles at three distinct life history stages (recruits, juveniles, and adults) on reefs with differing benthic conditions. Following widespread coral loss due to the 1998 bleaching event, some reefs are recovering (i.e., relatively high scleractinian coral cover: ‘coral-dominated’), some reefs have low cover of living macrobenthos and unconsolidated rubble substrates (‘rubble-dominated’), and some reefs have high cover of macroalgae (‘macroalgal-dominated’). Rates of coral recruitment to artificial settlement tiles were similar across all reef conditions, suggesting that larval supply does not explain differential coral recovery across the three reef types. However, acroporid recruits were absent on macroalgal-dominated reefs (0.0 ± 0.0 recruits tile^?1) in comparison to coral-dominated reefs (5.2 ± 1.6 recruits tile^?1). Juvenile coral colony density was significantly lower on macroalgal-dominated reefs (2.4 ± 1.1 colonies m^?2), compared to coral-dominated reefs (16.8 ± 2.4 m^?2) and rubble-dominated reefs (33.1 ± 7.3 m^?2), suggesting that macroalgal-dominated reefs have either a bottleneck to successful settlement on the natural substrates or a high post-settlement mortality bottleneck. Rubble-dominated reefs had very low cover of adult corals (10.0 ± 1.7 %) compared to coral-dominated reefs (33.4 ± 3.6 %) despite no statistical difference in their juvenile coral densities. A bottleneck caused by low juvenile colony survivorship on unconsolidated rubble-dominated reefs is possible, or alternatively, recruitment to rubble-dominated reefs has only recently begun. This study identified bottlenecks to recovery of coral assemblages that varied depending on post-disturbance habitat condition.     

Into the depth of population genetics: pattern of structuring in mesophotic red coral populations

Deep-sea reef-building corals are among the most conspicuous invertebrates inhabiting the hard-bottom habitats worldwide and are particularly susceptible to human threats. The precious red coral (Corallium rubrum, L. 1758) has a wide bathymetric distribution, from shallow up to 800 m depth, and represents a key species in the Mediterranean mesophotic reefs. Several studies have investigated genetic variability in shallow-water red coral populations, while geographic patterns in mesophotic habitats are largely unknown. This study investigated genetic variability of C. rubrum populations dwelling between 55 and 120 m depth, from the Ligurian to the Ionian Sea along about 1500 km of coastline. A total of 18 deep rocky banks were sampled. Colonies were analyzed by means of a set of microsatellite loci and the putative control region of the mitochondrial DNA. Collected data were compared with previous studies. Both types of molecular markers showed high genetic similarity between populations within the northern (Ligurian Sea and Tuscan Archipelago) and the southern (Tyrrhenian and Ionian seas) study areas. Variability in habitat features between the sampling sites did not affect the genetic variability of the populations. Conversely, the patchy distribution of suitable habitats affected populations’ connectivity within and among deep coral banks. Based on these results and due to the emphasis on red coral protection in the Mediterranean Sea by international institutions, red coral could be promoted as a ‘focal species’ to develop management plans for the conservation of deep coralligenous reefs, a reservoir of marine biodiversity.     

Trophic structure of cold-water coral communities revealed from the analysis of tissue isotopes and fatty acid composition

The trophic structure of cold-water coral reef communities at two contrasting locations, the 800-m deep Belgica Mounds (Irish margin) and the 300-m deep Træna reefs (Norwegian Shelf), was investigated using stable isotope (δ¹³C and δ¹⁵N) and fatty-acid composition analysis. A broad range of specimens, with emphasis on (commercial) fish specie's, and organic matter sources were sampled using a variety of tools. Irrespective of the environmental and geographical setting, the δ¹⁵N values indicated that the food web encompasses roughly 1.5 to 3 trophic levels. Mobile echinoderms, i.e. sea urchins and sea stars, had highest δ¹⁵N values, indicative of a high trophic position in the food web. The fraction of bacterial fatty acids in reef fauna was generally low (<5%), indicating that enhanced bacterial production in the water column through seafloor seepage of nutrients (‘hydraulic theory’) does not form a significant energy pathway into the food web. The high fraction of algal and essential fatty acids in reef fauna and fish at both locations indicates a close coupling with surface productivity, but the transport mechanism depends on the hydrographic setting. At Træna, Calanus copepods and euphausiids form an additional link between primary production and fish, which is largely absent at Belgica Mounds. At Belgica Mounds, the reef community is primarily supported by phytodetritus, as evidenced by the high contribution of algal fatty acids in faunal tissue and seasonal chlorophyll a deposition and marine snow at the reef. The environmental setting of cold-water coral reefs influences the structure of the associated food web.     

Coral—the world's most diverse symbiotic ecosystem

Zooxanthellate corals (i.e. those harbouring Symbiodinium) are the main builders of the world's shallow‐water marine coral reefs. They represent intimate diverse symbioses between coral animals, single‐celled photosynthetic dinoflagellates (Symbiodinium spp.), other microscopic eukaryotes, prokaryotes and viruses. Crabs and other crustaceans, worms, sponges, bivalves and hydrozoans, fishes, sea urchins, octopuses and sea stars are itinerant members of these ‘rainforests of the sea’. This review focuses on the biodiversity of scleractinian coral animals and their best studied microscopic epi‐ and endosymbionts. In relation to coral‐associated species diversity, Symbiodinium internal transcribed spacer region sequence types tally 10²–10³ or up to ~15 different operational taxonomic units (OTUs, or putative species at the 97% sequence identity level; this cut‐off was chosen based on intragenomic sequence diversity observed in monoclonal cultures) and prokaryotes (mostly bacterial) total 10²–10⁴ OTUs. We analysed all publically accessible 16S rRNA gene sequence data and found Gammaproteobacteria were extremely abundant, followed by Alphaproteobacteria. Notably, Archaea were poorly represented and ‘unassigned OTUs’ were abundant in data generated by high‐throughput DNA sequencing studies of corals. We outline and compare model systems that could be used in future studies of the coral holobiont. In our future directions, we recommend a global coral sampling effort including substantial attention being paid to method of coral tissue acquisition, which compartments (mucus, tissue, skeleton) to explore, broadening the holobiont members considered and linking biodiversity with functional investigations.     

Behavioural and developmental responses of predatory coral reef fish to variation in the abundance of prey

Ecological theory suggests that the behaviour, growth and abundance of predators will be strongly influenced by the abundance of prey. Predators may in turn play an important role in structuring prey populations and communities. Responses of predators to variation in prey abundance have most commonly been demonstrated in low-diversity communities where food webs are relatively simple. How predators respond in highly diverse assemblages such as in coral reef habitats is largely unknown. This study describes an experiment that examined how the movement, diet and growth of the coral reef piscivore, Cephalopholis boenak (Serranidae) responded to variation in the abundance of its prey. Predator densities were standardised on small patch reefs made from the lagoonal reef-building coral, Porites cylindrica. These patch reefs exhibited natural variation in the abundance and community structure of multiple species of prey. However, our experiment generated a relatively simple predator–prey relationship, with C. boenak primarily responding to the most abundant species of prey. Three responses of predators were observed: aggregative, functional and developmental. Thirty-one per cent of individuals moved between patch reefs during the experiment, all from areas of relatively low to high prey density. Feeding rates were higher on patch reefs of high prey density, while growth rates of fish that remained on low prey density reefs throughout the experiment were lower. Growth rates of C. boenak on the experimental reefs were also much higher than for those living on natural patch reefs over the same time period, corresponding with overall differences in prey abundance. These results suggest that local abundance, feeding rate and growth of C. boenak were closely linked to the abundance of their main prey. This combination of predatory responses is a potential mechanism behind recent observations of density-dependent mortality and population regulation of prey in coral reef fish communities.     

Coral and sea urchin assemblage structure and interrelationships in Kenyan reef lagoons

Patterns of hard coral and sea urchin assemblage structure (species richness, diversity, and abundance) were studied in Kenyan coral reef lagoons which experienced different types of human resource use. Two protected reefs (Malindi and Watamu Marine National Parks) were protected from fishing and coral collection, but exposed to heavy tourist use. One reef (Mombasa MNP) received protection from fishermen for one year and was exploited for fish and corals prior to protection and was defined as a transitional reef. Three reefs (Vipingo, Kanamai, and Diani) were unprotected and experienced heavy fishing and some coral collection. Protected and unprotected reefs were distinct in terms of their assemblage structure with the transitional reef grouping with unprotected reefs based on relative and absolute abundance of coral genera. Protected reefs had slightly higher (p<0.01) coral cover (23.6 ± 8.3 % ± S.D.) than unprotected reefs (16.7 ± 8.5), but the transitional reef had the highest coral cover (30.8 ± 6.4) which increased by 250% since measured in 1987: largely attributable to a large increase inPorites nigrescens cover. Protected reefs had higher coral species richness and diversity and a greater relative abundance ofAcropora, Montipora andGalaxea than unprotected reefs. The transitional reef had high species richness, but lower diversity due to the high dominance ofPorites. Sea urchins showed the opposite pattern with highest diversity in most unprotected reefs. Coral cover, species richness, and diversity were negatively associated with sea urchin abundance, but the relative abundance ofPorites increased with sea urchin abundance to the point wherePorites composed >90% of the coral cover at sites with the highest sea urchin abundance. Effects of coral overcollection was only likely for the genusAcropora (staghorn corals). A combination of direct and indirect effects of human resource use may reduce diversity, species richness, and abundance of corals while increasing the absolute abundance of sea urchins and the relative cover ofPorites.     

Incorporating fine-scale seascape composition in an assessment of habitat quality for the giant sea anemone Stichodactyla gigantea in a coral reef shore zone

Habitat loss due to land reclamation often occurs in sandy coral reef shore zones. The giant sea anemone Stichodactyla gigantea, which harbors the false clown anemonefish Amphiprion ocellaris, both of which are potentially flagship species, inhabit these places. To assess habitat quality for S. gigantea, we examined correlative associations between the number and the body size of S. gigantea and the amount of habitat types in fine-scale seascape composition quantified from an enlarged section of a high-resolution (1/2,500) color aerial photograph of the shallow shore zone of Shiraho Reef, Ishigaki Island, Japan. This study confirmed that anemones were most abundant at the edges of dense seagrass beds characterized by shallow sandy bottoms, rock beds, and sparse seagrass beds, while they were less abundant in coral patch reefs. However, anemones inhabiting coral patch reefs were significantly larger and their rate of disappearance over 3 years was lower than those inhabiting other habitats. This suggests that coral patch reefs may be more suitable habitats supporting larger animals and greater persistence of S. gigantea. The visual census techniques applied here, combined with aerial photography and image-analysis software, may be useful as a simple analytical tool for local assessment of suitable habitats for relatively small-bodied marine fauna in shallow-water seascapes.     

Foraging plasticity in obligate corallivorous Melon butterflyfish across three recently bleached reefs

Because obligate corallivorous butterflyfish feed exclusively on coral polyps, they are particularly sensitive to changes in coral cover and its spatial distribution. To understand how such differences in coral cover influence obligate corallivores, we examined the densities and foraging behavior of Melon butterflyfish Chaetodon trifasciatus across three reefs in the Lakshadweep archipelago. These reefs suffered differential bleaching mortality after the 2010 El Niño Southern Oscillation, resulting in wide variation in coral cover and community composition. Despite these differences, C. trifasciatus were able to persist at similar densities across reefs. However, our analysis of high‐resolution video recordings of multiple focal fish revealed that time budgets, bite rates, and diet selectivity differed significantly. Fish in resource‐poor reefs spent more time moving between coral patches and less time foraging than ones in relatively resource‐rich reefs. We also found that fish in resource‐poor reefs had higher bite rates and were less selective in their foraging. Our results provide novel insights into how obligate corallivores cope with even large differences in resource availability. At a time when we are rapidly losing corals to repeated climate‐induced bleaching events, this flexibility may represent a critical mechanism that enables persistence of obligate corallivores in resource‐poor reefs, even if it does not guarantee longer‐term survival.     

Water column contributions to coral reef productivity: overcoming challenges of context dependence

Coral reefs are declining at an unprecedented rate. Effective management and conservation initiatives necessitate improved understanding of the drivers of production because the high rates found in these ecosystems are the foundation of the many services they provide. The water column is the nexus of coral reef ecosystem dynamics, and functions as the interface through which essentially all energy and nutrients are transferred to fuel both new and recycled production. Substantial research has described many aspects of water column dynamics, often focusing on specific components because water column dynamics are highly spatially and temporally context dependent. Although necessary, a cost of this approach is that these dynamics are often not well linked to the broader ecosystem or across systems. To help overcome the challenge of context dependence, we provide a comprehensive review of this literature, and synthesise it through the perspective of ecosystem ecology. Specifically, we provide a framework to organise the drivers of temporal and spatial variation in production dynamics, structured around five primary state factors. These state factors are used to deconstruct the environmental contexts in which three water column sub-food webs mediate ‘new’ and ‘recycled’ production. We then highlight critical pathways by which global change drivers are altering coral reefs via the water column. We end by discussing four key knowledge gaps hindering understanding of the role of the water column for mediating coral reef production, and how overcoming these could improve conservation and management strategies. Throughout, we identify areas of extensive research and those where studies remain lacking and provide a database of 84 published studies. Improved integration of water column dynamics into models of coral reef ecosystem function is imperative to achieve the understanding of ecosystem production necessary to develop effective conservation and management strategies needed to stem global coral loss.     

Fitness consequences of habitat variability,trophic position,and energy allocation across the depth distribution of a coral-reef fish

Environmental clines such as latitude and depth that limit species’ distributions may be associated with gradients in habitat suitability that can affect the fitness of an organism. With the global loss of shallow-water photosynthetic coral reefs, mesophotic coral ecosystems (~30–150 m) may be buffered from some environmental stressors, thereby serving as refuges for a range of organisms including mobile obligate reef dwellers. Yet habitat suitability may be diminished at the depth boundary of photosynthetic coral reefs. We assessed the suitability of coral-reef habitats across the majority of the depth distribution of a common demersal reef fish (Stegastes partitus) ranging from shallow shelf (SS, <10 m) and deep shelf (DS, 20–30 m) habitats in the Florida Keys to mesophotic depths (MP, 60–70 m) at Pulley Ridge on the west Florida Shelf. Diet, behavior, and potential energetic trade-offs differed across study sites, but did not always have a monotonic relationship with depth, suggesting that some drivers of habitat suitability are decoupled from depth and may be linked with geographic location or the local environment. Feeding and diet composition differed among depths with the highest consumption of annelids, lowest ingestion of appendicularians, and the lowest gut fullness in DS habitats where predator densities were highest and fish exhibited risk-averse behavior that may restrict foraging. Fish in MP environments had a broader diet niche, higher trophic position, and higher muscle C:N ratios compared to shallower environments. High C:N ratios suggest increased tissue lipid content in fish in MP habitats that coincided with higher investment in reproduction based on gonado-somatic index. These results suggest that peripheral MP reefs are suitable habitats for demersal reef fish and may be important refuges for organisms common on declining shallow coral reefs.

     

Coral reef conservation in Bali in light of international best practice,a literature review

Bali, Indonesia sits within the coral triangle and is internationally recognised for its high coral reef diversity. The health of Bali’s marine ecosystems has declined in recent decades, and this is thought to be due to threats from climate change, destructive fishing practices, pollution, outbreaks coral eating invertebrates, coral disease and unsustainable tourism. As a response, multiple conservation strategies have been introduced by the island’s communities, non-government organisations and governments, with the aim of preventing further decline, as well as restoring already degraded coral reefs. This literature review provides an in-depth analysis of the tools used to conserve Bali’s coral reefs, and compares them to those used in other countries. In light of international ‘best practice’ in coral reef conservation, this review makes suggestions on how Bali could better conserve its coral reef ecosystems. These include (1) increasing its designation of official Marine Protected Areas (MPAS) and strengthening management of existing ones, (2) creating an MPA network, (3) substantially reducing marine plastic pollution, (4) continuing artificial reef construction in degraded habitats, (5) continuing to develop Bali as an ecotourism destination, (6) increasing engagement in global science to inform marine conservation decision-making, and (7) developing more marine monitoring programmes.     

Distribution and abundance of fish in deep-sea coral habitats

Experimental fishing with long-lines and gillnets was conducted on the continental shelf off southwestern Norway between 150 and 350 m depth. Abundance and distribution of redfish (Sebastes marinus L., 1758), ling (Molva molva L., 1758), and tusk (Brosme brosme Ascanius, 1772) were quantified in Lophelia pertusa (L., 1758) coral reefs and in non-coral habitats. The largest catches of redfish were made with long-line fleets set in coral reef habitats. Ling and tusk were also most numerous in coral habitats, although not statistically significant. Fish caught in coral habitats tended to be larger in size than in non-coral habitats. The diet of redfish, tusk and ling included the same prey groups in all habitats, but they differed at the species level. Lophelia-reefs may provide a profitable feeding place for tusk. For the planktivorous Sebastes, on the other hand, their affinity to the reefs seems primarily to be related to the physical structure offered by the reefs.     

Physiological performance of the cold-water coral Dendrophyllia cornigera reveals its preference for temperate environments

Cold-water corals (CWCs) are key ecosystem engineers in deep-sea benthic communities around the world. Their distribution patterns are related to several abiotic and biotic factors, of which seawater temperature is arguably one of the most important due to its role in coral physiological processes. The CWC Dendrophyllia cornigera has the particular ability to thrive in several locations in which temperatures range from 11 to 17 °C, but to be apparently absent from most CWC reefs at temperatures constantly below 11 °C. This study thus aimed to assess the thermal tolerance of this CWC species, collected in the Mediterranean Sea at 12 °C, and grown at the three relevant temperatures of 8, 12, and 16 °C. This species displayed thermal tolerance to the large range of seawater temperatures investigated, but growth, calcification, respiration, and total organic carbon (TOC) fluxes severely decreased at 8 °C compared to the in situ temperature of 12 °C. Conversely, no significant differences in calcification, respiration, and TOC fluxes were observed between corals maintained at 12 and 16 °C, suggesting that the fitness of this CWC is higher in temperate rather than cold environments. The capacity to maintain physiological functions between 12 and 16 °C allows D. cornigera to be the most abundant CWC species in deep-sea ecosystems where temperatures are too warm for other CWC species (e.g., Canary Islands). This study also shows that not all CWC species occurring in the Mediterranean Sea (at deep-water temperatures of 12–14 °C) are currently living at their upper thermal tolerance limit.     

ECOLOGY AND MORPHOLOGY OF RECENT CORAL REEFS

1. The classical ‘coral reef problem’ concerned the geological relationships of reefs as major topographical features; modern coral studies consider reefs both as complex biological systems of high productivity and as geological structures forming a framework for and being modified by coral growth. 2. Deep borings in reefs have conclusively confirmed the general arguments of Darwin, that oceanic reefs developed by progressive subsidence of their foundations. Darwin failed to take account of Pleistocene changes in sea level and their effect on the present surface features of reefs. Daly's alternative ‘glacial control theory’ was based on false assumptions concerning marine erosion rates during glacial periods, but if sea level during the Holocene was higher than at present, as Daly also supposed, the effects on reef features would be profound. 3. Reefs are complex biological systems in tropical seas, dominated by scleractinian corals. Coral faunas are larger and more diverse in the Indo-Pacific than in the Atlantic. Hermatypic corals are restricted to shallow water by the light requirements of their symbiotic algae, but temperature is a major control of worldwide distributions. Temperature, salinity and sediment tolerances of corals are wider than formerly supposed, and corals can survive brief emersion except when it coincides with heavy rainfall. Water turbulence is an important ecological control, but difficult to measure. 4. The trophic status of corals is still unclear, but in spite of their anatomical and physiological specialization as carnivores it is likely that they derive some nutrient substances from zooxanthellae. Suggestions that filamentous algae in coral heads play a major part in the economy of the corals have not been supported by later work, but biomass pyramids constructed on the basis by Odum and Odum remain the only ones available. Most reefs are apparently autotrophic, with 1500–3500 g. Carbon being fixed per m.² per year. 5. Few animals eat corals, which may account for their success. Important predators are fish and the echinoderm Acanthaster. Quantitative estimates of biogenic erosion of organic skeletons on reefs are high. Fish affect not only corals but other invertebrates, algae and marine phanerogams. 6. Corals may be killed by ‘dark water’, intense rain or river floodwaters, earth movements, human interference and especially hurricanes. Reef recovery after hurricanes may take 10–20 years. 7. In addition to fringing, barrier and atoll reefs, intermediate types are recognised. The main types may consist of linear reefs or faros. Smaller lagoon reefs include pinnacles, patches and platforms, and submerged knolls. Complex cellular or mesh reef patterns are also found. 8. Reefs are conspicuously zoned, both laterally in response to changing exposure to waves to form windward and leeward reefs, and transversely, as a result of steep environmental gradients across reef flats from sea to lagoon. Topographic and ecological zones may be characterized by particular coral species, but these vary widely from reef to reef. A major distinction can be made between reefs with and without algal ridges, which are common on open-ocean trade-wind reefs, in the Indo-Pacific, but are absent on Caribbean reefs and on Indo-Pacific reefs in more sheltered waters. gorgonians are common on Caribbean reefs, alcyonaceans in the Indo-Pacific. 9. Much of the difficulty in comparing reefs stems from the lack of uniformity in surveying methods. Problems of describing the complex three-dimensional patterns of organisms on reefs have yet to be solved, and hence little progress has been made in explanation of these patterns. Explanation in terms of simple environmental controls is inadequate. 10. Understanding the distribution of corals is made more difficult both by taxo-nomic problems and by the plasticity of growth form in different situations. 11. Growth of corals and reefs may be estimated by measuring the growth of individual colonies, measuring rates of calcium carbonate deposition in the skeleton, measuring topographic change on the reef and deducing net rates of reef growth from geological evidence. Massive corals may increase in diameter by 1 cm./year, branches of branching corals may increase in length by 10 cm./year. Study of deposition rates shows variation within colonies, between species, in light and dark, and seasonally. Rates of reef growth extrapolated from colony measurements reach 2–5 cm./year, and contrast with figures as low as 0–02 cm/year averaged over 70 million years from borehole data. Both colony growth rates and geological data suggest worldwide variations in rates of reef growth. 12. In spite of clear evidence of long-continued subsidence, present surface features of reefs, often only thinly veneered by modern corals, have been much affected by recent sea level fluctuations. Many slightly raised reefs at 2–10 m. above sea level date at 90–160 thousand years B.P.; there is evidence for a sea level at about the present level at 30–35 thousand years B.P.; and controversy continues over whether sea level has stood higher than the present at any time since the last sea level rise began about 20,000 years ago. Evidence from many reefs suggests a slightly higher sea level in the last 4000 years, but on other reefs such evidence is lacking. 13. Several reef features (submerged terraces, groove-spur systems, algal ridge, reef flat, reef blocks and reef islands) have been interpreted either as relict features dating from a higher sea level in the last 5000 years, or contemporary features developed in response to present processes. In some cases the evidence is equivocal; in others it is clear that diverse features are being grouped together under the same name. If such features are referable to a higher sea level, this may have been of last Interglacial or even Interstadial age rather than Holocene. 14. A reef consists of a rigid framework defining several major depositional environments within and around it. Sediments are of biological, mainly skeletal origin, except in unusual environments such as the Bahama Banks. The characteristics of sediments derived from organisms depend partly on the breakdown patterns of particular skeletons, partly on transportation and sorting processes. Fine sediments may be either detrital, or physicochemical precipitates. 15. Organisms affect sediments after deposition, by disturbance, transportation and probably comminution. Fish and holothurians have been studied in detail. 16. While new theories of coral reefs are proposed from time to time, the need is less for new theories than for standardised procedures to ensure comparability of reef studies and the identification of variations in reefs both on local and regional scales. While reefs as biological systems adjust relatively rapidly to changes, reefs as geological systems adjust much more slowly. Because of the magnitude and recency of Pleistocene fluctuations in sea level, many biological features of reefs are out of phase with inherited geological features, and this had led to much controversy.     

Are artificial reefs surrogates of natural habitats for corals and fish in Dubai,United Arab Emirates?

Artificial reefs are often promoted as mitigating human impacts in coastal ecosystems and enhancing fisheries; however, evidence supporting their benefits is equivocal. Such structures must be compared with natural reefs in order to assess their performance, but past comparisons typically examined artificial structures that were too small, or were immature, relative to the natural reefs. We compared coral and fish communities on two large (>400,000 m³) and mature (>25 year) artificial reefs with six natural coral patches. Coral cover was higher on artificial reefs (50%) than in natural habitats (31%), but natural coral patches contained higher species richness (29 vs. 20) and coral diversity (H′ = 2.3 vs. 1.8). Multivariate analyses indicated strong differences between coral communities in natural and artificial habitats. Fish communities were sampled seasonally for 1 year. Multivariate fish communities differed significantly among habitat types in the summer and fall, but converged in the winter and spring. Univariate analysis indicated that species richness and abundance were stable throughout the year on natural coral patches but increased significantly in the summer on artificial reefs compared with the winter and spring, explaining the multivariate changes in community structure. The increased summer abundance on artificial reefs was mainly due to adult immigration. Piscivores were much more abundant in the fall than in the winter or spring on artificial reefs, but had low and stable abundance throughout the year in natural habitats. It is likely that the decreased winter and spring abundance of fish on the artificial reefs resulted from both predation and emigration. These results indicate that large artificial reefs can support diverse and abundant coral and fish communities. However, these communities differ structurally and functionally from those in natural habitats, and they should not be considered as replacements for natural coral and fish communities.     

Caribbean mesophotic coral ecosystems are unlikely climate change refugia

Deeper coral reefs experience reduced temperatures and light and are often shielded from localized anthropogenic stressors such as pollution and fishing. The deep reef refugia hypothesis posits that light‐dependent stony coral species at deeper depths are buffered from thermal stress and will avoid bleaching‐related mass mortalities caused by increasing sea surface temperatures under climate change. This hypothesis has not been tested because data collection on deeper coral reefs is difficult. Here we show that deeper (mesophotic) reefs, 30–75 m depth, in the Caribbean are not refugia because they have lower bleaching threshold temperatures than shallow reefs. Over two thermal stress events, mesophotic reef bleaching was driven by a bleaching threshold that declines 0.26 °C every +10 m depth. Thus, the main premise of the deep reef refugia hypothesis that cooler environments are protective is incorrect; any increase in temperatures above the local mean warmest conditions can lead to thermal stress and bleaching. Thus, relatively cooler temperatures can no longer be considered a de facto refugium for corals and it is likely that many deeper coral reefs are as vulnerable to climate change as shallow water reefs.     

Review of coral reef ecosystem remote sensing

Coral reef communities face unprecedented pressures at local, regional and global scales as a consequence of climate change and anthropogenic disturbance. Remote sensing, from satellites or aircraft, is possibly the only means to measure the effects of such stresses at appropriately large spatial scales. In the past 30 years, remote sensing of coral reefs has made rapid progress. However, the current technology is still not mature enough to monitor complicated coral reef ecosystems. Compared with foreign research in this field, our work lags far behind. There are still deficiencies in many aspects, such as basic data collection, theoretical research and platform construction. In our nation, it is even unclear how coral reefs disperse and where they may be unhealthy. In this paper, general characteristics of coral reef ecosystems and spectral features of different reef benthos have been summarized, based initially on a review of relevant literature in recent years. Based on the spectral separability of different reef types or benthos, remote sensing can be used to monitor two aspects of coral reefs: (1) Measurement of the ecological properties of reefs. (2) Health assessment of the coral reef ecosystem. In the first part, optical remote sensing methods are widely used to map reef geomorphology and habitats or biotopes. The investigation of geomorphologic zonation has proven to be one of the most successful applications, as different geomorphologic zones are associated with characteristic benthic community structures and occur at spatial scales of tens to hundreds of meters, they are amenable to remote detection by moderate to high resolution sensors. With more and more attention on the ecological problems of coral reefs, a number of studies have used high resolution sensors to map reef communities. The number of classes distinguishable depends on many factors, including the platforms, resolution (spectral, spatial and temporal resolution) and environmental conditions (water depth, water clarity, surface roughness, etc.). Compared with deep water color remote sensing, or terrestrial remote sensing, three techniques for the measurement of reef ecological properties are examined in this paper: (1) Coral reef classification system using remote sensing. (2) Techniques of sea surface correction and water column correction. (3) Techniques of coral reef information extraction from images. In terms of the complexity of coral reef ecosystems, the current techniques still need further improvement or optimization. In the health assessment of coral reef ecosystems, there are two ways to carry out the monitoring using remote sensing: (1) Monitoring the pigment or symbiotic zooxanthellae contents in corals. (2) Measuring the environmental properties of reefs. The first way is theoretically feasible, but difficult to achieve in practice. Currently, most reef health assessments are carried out by measuring environmental parameters, including sea surface temperature, solar radiation, ultraviolet radiation, water color, wind speed and direction, rainfall, ocean acidification, sea level, etc., of which sea surface temperature has been routinely measured by NOAA to monitor coral bleaching. In addition to the contents above, this article puts forward five main prospects for development in the future: (1) Establishment of a coral reef classification system using remote sensing. (2) Satellite launch for monitoring coral reefs. (3) Theoretical and methodological development. (4) Establishment of a spectral database for different reef benthos. (5) Integrated application of multi-source remote sensing data. It is hoped that the information provided here will be a reference for subsequent similar studies.