Coral Community Adaptability to Environmental Change at the Scales of Regions, Reefs and Reef Zones

SYNOPSIS. Projected global increases in temperature, sea level,storminess and atmospheric carbon dioxide (CO₂) are likely tocause changes in reef coral communities which the present humangeneration will view as deleterious. It is likely coral communitytrajectories will be influenced as much by the reduction inintervals between extreme events as the projected increasesin means of environmental parameters such as temperature, atmosphericCO₂ and sea-level. Depressed calcification rates in corals causedby reduced aragonite saturation state of water may increasevulnerability of corals to storms. Moreover, reduction in intervalsbetween storms and other extreme events causing mass mortalityin corals (coral predators, diseases, bleaching) are likelyto more frequently "set back" reef coral communities to earlysuccessional stages or alternate states characterized by non-calcifyingbenthos (plants, soft corals, sponges). The greater the areaand the longer the duration of dominance of putative "coral/corallinealgae" zones of coral reefs by non-calcifying stages, the lesswill be the reefs capacity to accrete limestone bulk lockedup in the big skeletal units of late successional stages (i.e.,very large old corals). Averaged over decades to centuries,the effects of such changes on the coral community's carryingcapacity for other biota such as fish are unpredictable. A "shiftingsteady-state mosaic" null model may provide a useful conceptualtool for defining a baseline and tracking changes from thatbaseline through time.     

Ecological persistence interrupted in Caribbean coral reefs

The recent mass mortality of Caribbean reef corals dramatically altered reef community structure and begs the question of the past stability and persistence of coral assemblages before human disturbance began. We report within habitat stability in coral community composition in the Pleistocene fossil record of Barbados for at least 95 000 years despite marked variability in global sea level and climate. Results were consistent for surveys of both common and rare taxa. Comparison of Pleistocene and modern community structure shows that Recent human impacts have changed coral community structure in ways not observed in the preceding 220 000 years.     

Coral associations of the Pleistocene Ironshore Formation,Grand Cayman

The 125-ka sea level, which was approximately 6 m above present-day sea level, led to the partial flooding of many Caribbean islands. On Grand. Cayman, this event led to the formation of the large Ironshore Lagoon that covered most of the western half of the island and numerous, small embayments along the south, east, and north coasts. At that time, at least 33 coral species grew in waters around Grand Cayman. This fauna, like the modern coral fauna of Grand Cayman, was dominated byMontastrea annularis, Porites porites, Acropora polmata, andA. cervicornis. Scolymia cubensis andMycetophyllia ferox, not previously identified from the Late Pleistocene, are found in the Pleistocene patch reefs.Madracis mirabilis, Colpophyllia breviserialis, Agaricia tenuifolia, A. lamarcki, A. undata, Millepora spp., Mycetophyllia reesi, M. aliciae, andM. danaana, found on modern reefs, have not been identified from the Late Pleistocene reefs. Conversely,Pocillopora sp. cf.P. palmata, which is found in Late Pleistocene reefs, is absent on the modern reefs around Grand Cayman. The corals in the Ironshore Formation of Grand Cayman have been divided into 10 associations according to their dominant species, overall composition, and faunal diversity. Many of these associations are similar to the modern associations around Grand Cayman. Each of the Pleistocene coral associations, which can be accurately located on the known Late Pleistocene paleogeography of Grand Cayman, developed in distinct environmental settings. Overall trends identified in the modern settings are also apparent in the Late Pleistocene faunas. Thus, the diversity of the coral faunas increased from the interior of the Ironshore Lagoon to the reef crest. Similarly, the coral diversity in the Pleistocene patch reefs was related to the size of the reefs and their position relative to breaks in the barrier reef. The barrier reef included corals that are incapable of sediment rejection; whereas the patch reefs lacked such corals.     

Reef coral diversity and global change

Regional anthropogenic processes such as pollution, dredging, and overfishing on coral reefs currently threaten the biodiversity of stony corals and other reef-associated organisms. Global climate change may interact with anthropogenic processes to create additional impacts on coral diversity in the near future. In order to predict these changes, it is necessary to understand the magnitude and causes of variation in scleractinian coral diversity throughout their 240 million year history. The fossil record documents long periods of speciation in corals, interrupted repeatedly by events of mass extinction. Some of these events relate clearly to changes in global climate. Diversity in reef corals has increased since their last period of extinction at the end of the Cretaceous (65 My bp ), and is still rising. During the last 8 million years, the fragmentation of the once pantropical Tethys Sea separated corals into two major biogeographical provinces. Periods of glaciation also have caused major changes in sea level and temperature. Accumulated evidence supports the theory that relative habitat area and changing patterns of oceanic circulation are mainly responsible for the two observed centres of recent coral diversity at the western tropical margins of the Atlantic and Pacific oceans. At predicted rates of climate change in the near future, coral reefs are likely to survive as an ecosystem. Increases in sea level may actually benefit corals and lead to regional increases in diversity if new habitat area on back reefs is opened to increased water circulation and thus coral dispersal. Rising temperature may cause higher rates of coral mortality and even local extinction in isolated, small populations such as those on oceanic islands. The effects of increases in ultraviolet radiation (UV) are largely unknown, but likely to be negative. UV may damage planktonic coral propagules in oceanic surface waters and thus decrease rates of gene flow between coral populations. This may result in increased local extinctions, again with the strongest impact on widely separated reefs with small coral populations. The largest threats to coral diversity are regional anthropogenic impacts, which may interact with global climate change to exacerbate rates of local species extinctions. Centres of high reef coral diversity coincide with human population centres in south-east Asia and the Caribbean, and thus the greatest potential for species loss lies in these geographical areas.     

Character release following extinction in a Caribbean reef coral species complex

Abstract The Pleistocene extinction of the widespread organ‐pipe Montastraea coral had measurable morphological and ecological effects on surviving lineages of the Montastraea“annularis” species complex. Extinction of the organ‐pipe Montastraea occurred after more than 500,000 years of dominance in the shallow‐water reef habitat of Barbados. Extinction resulted in a morphological shift of the columnar Montastraea lineage from thick to thin columns in modern reef environments. Pleistocene colonies of the columnar morphotype sympatric with organ‐pipe Montastraea showed greater column widths than those in allopatry. We subjected our data to a number of criteria for interpreting the morphological shift as character release following lifting of competitive pressure after extinction. The morphological differences do not appear to be due either to chance or to physical properties of the marine environment. Differential local extinction and recolonization of four members of the species complex did not occur on Barbados, so that the species coexisted and appear to have coevolved between more than 600,000 and 82,000 years ago. The morphological shift is related to coral growth form and growth rate, and thus reflects the acquisition of a primary resource in corals‐light. Character release occurred at the same oceanic Caribbean island (Barbados) where environments have fluctuated with similar variance throughout the period of coexistence. Not only has competition among living members of the Montastraea“annularis” species complex been convincingly demonstrated, but trends in relative abundance among fossil members of the species complex strongly suggest that a competitive hierarchy was operating during their Pleistocene coexistence on Barbados. We also observed an ecological analogue to character release on another Caribbean island, Curaçao. The distribution and abundance of living columnarM. annularis s.s. and massive M. faveolata from the leeward reef crest in Curaçao is greater now than in the Pleistocene, when organ‐pipe Montastraea dominated this shallow‐water reef habitat. Extinction of the faster growing, shallow‐water organ‐pipe Montastraea resulted in higher abundance of the columnar Montastraea lineage in shallow‐water habitats, where it shifted its morphology to one adapted to high light levels. The species extinction released surviving lineages from a competitive network that had resulted in lower rank abundance in the Pleistocene community and enhanced abundance of both columnar M. annularis s.s. and M. faveolata in modern communities. Full validation of our interpretation of character release must await experiments that demonstrate whether phenotypic differences between populations have a genetic basis. However, we believe the results of this study point to the important, yet heretofore neglected, role that biological interactions have played in the evolution of closely related reef coral species.     

Photosynthesis and Calcification at Cellular, Organismal and Community Levels in Coral Reefs: A Review on Interactions and Control by Carbonate Chemistry

SYNOPSIS. Photosynthesis and calcification in zooxanthellatescleractinian corals and coral reefs are reviewed at severalscales: cellular (pathways and transport mechanisms of inorganiccarbon and calcium), organismal (interaction between photosynthesisand calcification, effect of light) and ecosystemic (communityprimary production and calcification, and air-sea CO₂ exchanges). The coral host plays a major role in supplying carbon for thephotosynthesis by the algal symbionts through a system similarto the carbon-concentrating mechanism described in free livingalgal cells. The details of carbon supply to the calcificationprocess are almost unknown, but metabolic CO₂ seems to be asignificant source. Calcium supply for calcification is diffusionalthrough oral layers, and active membrane transport only occursbetween the calicoblastic cells and the site of calcification.Photosynthesis and calcification are tightly coupled in zooxanthellatescleractinian corals and coral reef communities. Calcificationis, on average, three times higher in light than in darkness.The recent suggestion that calcification is dark-repressed ratherthan light-enhanced is not supported by the literature. Thereis a very strong correlation between photosynthesis and calcificationat both the organism and community levels, but the ratios ofcalcification to gross photosynthesis (0.6 in corals and 0.2in reef communities) differ from unity, and from each otheras a function of level. The potential effect of global climatic changes (pCO₂ and temperature)on the rate of calcification is also reviewed. In various calcifyingphotosynthetic organisms and communities, the rate of calcificationdecreases as a function of increasing pCO₂ and decreasing calciumcarbonate saturation state. The calculated decrease in CaCO₃,production, estimated using the scenarios considered by theInternational Panel on Climate Change (IPCC), is 10% between1880 and 1990, and 9–30% (mid estimate: 22%) from 1990to 2100. Inadequate understanding of the mechanism of calcificationand its interaction with photosynthesis severely limits theability to provide an accurate prediction of future changesin the rate of calcification.     

Vulnerability of global coral reef habitat suitability to ocean warming,acidification and eutrophication

Coral reefs are threatened by global and local stressors. Yet, reefs appear to respond differently to different environmental stressors. Using a global dataset of coral reef occurrence as a proxy for the long‐term adaptation of corals to environmental conditions in combination with global environmental data, we show here how global (warming: sea surface temperature; acidification: aragonite saturation state, Ω_arag) and local (eutrophication: nitrate concentration, and phosphate concentration) stressors influence coral reef habitat suitability. We analyse the relative distance of coral communities to their regional environmental optima. In addition, we calculate the expected change of coral reef habitat suitability across the tropics in relation to an increase of 0.1°C in temperature, an increase of 0.02 μmol/L in nitrate, an increase of 0.01 μmol/L in phosphate and a decrease of 0.04 in Ω_arag. Our findings reveal that only 6% of the reefs worldwide will be unaffected by local and global stressors and can thus act as temporary refugia. Local stressors, driven by nutrient increase, will affect 22% of the reefs worldwide, whereas global stressors will affect 11% of these reefs. The remaining 61% of the reefs will be simultaneously affected by local and global stressors. Appropriate wastewater treatments can mitigate local eutrophication and could increase areas of temporary refugia to 28%, allowing us to ‘buy time’, while international agreements are found to abate global stressors.     

热带珊瑚礁区海参的生境选择与生态作用

珊瑚礁生态系统是一个高生产力、高生物多样性的特殊海洋生态系统,具有为生物提供栖息地、参与生物地球化学循环、防浪护岸、指示水体污染程度等生态功能。珊瑚礁生态系统的突出特点是其生境异质性很高,各种各样的生境斑块为种类繁多、习性各异的游泳和底栖生物提供栖息场所,这些礁栖生物通过参与各项生态过程而形成各种特定的功能群,共同完成重要的生态功能。在热带珊瑚礁生态系统中,海参是大型底栖动物区系的重要一员。种类繁多的海参具有各自不同的生境选择特征,通过摄食、运动等行为活动发挥着改良底质、促进有机物矿化和营养盐再生等生态作用。近几年来,全球热带海参受人类过度捕捞和珊瑚礁退化的影响而面临资源衰退、物种多样性丧失等问题,深入认识其生态学功能、加强热带海参资源保护迫在眉睫。综述了国内外热带珊瑚礁海参的基础生态学研究进展：海参对珊瑚礁生境斑块呈现显著的偏好选择特征以及种间差异和季节变动,不同生境斑块的食物质量、底质类型和水动力条件是影响海参生境偏好的重要因素;海参通过生物扰动可以改变珊瑚礁生境沉积物的含水量、渗透性、颗粒组成、再矿化率、无机营养物质释放速率以及孔隙水的化学梯度,并增加沉积物中的溶氧浓度、促进溶解...     

The Physiological Mechanisms of Acclimatization in Tropical Reef Corals

SYNOPSIS. The ability of scleractinian corals to survive changesthat are predicted in the global environment over the next centurywill lie in their physiological mechanisms of acclimatization.Corals display rapid modifications in behavior, morphology andphysiology enabling them to photoacclimate to changing lightconditions, a scenario that demonstrates considerable biologicalflexibility. Here we argue that the acclimatization mechanismsin corals are fundamentally similar to those exhibited by otherinvertebrate taxa. We discuss protein metabolism as a mechanismunderlying acclimatization responses in reef corals, and explorethe relationship between protein turnover, metabolic rate, growthrate, and acclimatization capacity. Our preliminary analysessuggest that corals with low growth rates (µCa/mgN/h)and high metabolic rates (µO₂/cm²/hr), such as the massivespecies, acclimatize more effectively than those with high growthrates and low metabolic rates, a feature that is characteristicof branching species. We conclude that studies of protein turnover,combined with temporally relevant investigations into the dynamicaspects of coral dinoflagellate symbioses will provide considerableinsight into why corals exhibit such a high level of variationin response to the same environmental challenge. Furthermore,a more detailed understanding of acclimatization mechanismsis essential if we are to predict how a coral assemblage willrespond to present and future environmental challenges.     

Importance of live coral habitat for reef fishes

Live corals are the key habitat forming organisms on coral reefs, contributing to both biological and physical structure. Understanding the importance of corals for reef fishes is, however, restricted to a few key families of fishes, whereas it is likely that a vast number of fish species will be adversely affected by the loss of live corals. This study used data from published literature together with independent field based surveys to quantify the range of reef fish species that use live coral habitats. A total of 320 species from 39 families use live coral habitats, accounting for approximately 8 % of all reef fishes. Many of the fishes reported to use live corals are from the families Pomacentridae (68 spp.) and Gobiidae (44 spp.) and most (66 %) are either planktivores or omnivores. 126 species of fish associate with corals as juveniles, although many of these fishes have no apparent affiliation with coral as adults, suggesting an ontogenetic shift in coral reliance. Collectively, reef fishes have been reported to use at least 93 species of coral, mainly from the genus Acropora and Porities and associate predominantly with branching growth forms. Some fish associate with a single coral species, whilst others can be found on more than 20 different species of coral indicating there is considerable variation in habitat specialisation among coral associated fish species. The large number of fishes that rely on coral highlights that habitat degradation and coral loss will have significant consequences for biodiversity and productivity of reef fish assemblages.     

Regional and global climate risks for reef corals: Incorporating species-specific vulnerability and exposure to climate hazards

Climate change is driving rapid and widespread erosion of the environmental conditions that formerly supported species persistence. Existing projections of climate change typically focus on forecasts of acute environmental anomalies and global extinction risks. The current projections also frequently consider all species within a broad taxonomic group together without differentiating species-specific patterns. Consequently, we still know little about the explicit dimensions of climate risk (i.e., species-specific vulnerability, exposure and hazard) that are vital for predicting future biodiversity responses (e.g., adaptation, migration) and developing management and conservation strategies. Here, we use reef corals as model organisms (n = 741 species) to project the extent of regional and global climate risks of marine organisms into the future. We characterise species-specific vulnerability based on the global geographic range and historical environmental conditions (1900–1994) of each coral species within their ranges, and quantify the projected exposure to climate hazard beyond the historical conditions as climate risk. We show that many coral species will experience a complete loss of pre-modern climate analogs at the regional scale and across their entire distributional ranges, and such exposure to hazardous conditions are predicted to pose substantial regional and global climate risks to reef corals. Although high-latitude regions may provide climate refugia for some tropical corals until the mid-21st century, they will not become a universal haven for all corals. Notably, high-latitude specialists and species with small geographic ranges remain particularly vulnerable as they tend to possess limited capacities to avoid climate risks (e.g., via adaptive and migratory responses). Predicted climate risks are amplified substantially under the SSP5-8.5 compared with the SSP1-2.6 scenario, highlighting the need for stringent emission controls. Our projections of both regional and global climate risks offer unique opportunities to facilitate climate action at spatial scales relevant to conservation and management.     

Spatial dynamics of benthic competition on coral reefs

The community structure of sedentary organisms is largely controlled by the outcome of direct competition for space. Understanding factors defining competitive outcomes among neighbors is thus critical for predicting large-scale changes, such as transitions to alternate states within coral reefs. Using a spatially explicit model, we explored the importance of variation in two spatial properties in benthic dynamics on coral reefs: (1) patterns of herbivory are spatially distinct between fishes and sea urchins and (2) there is wide variation in the areal extent into which different coral species can expand. We reveal that the size-specific, competitive asymmetry of corals versus fleshy algae highlights the significance of spatial patterning of herbivory and of coral growth. Spatial dynamics that alter the demographic importance of coral recruitment and maturation have profound effects on the emergent structure of the reef benthic community. Spatially constrained herbivory (as by sea urchins) is more effective than spatially unconstrained herbivory (as by many fish) at opening space for the time needed for corals to settle and to recruit to the adult population. Further, spatially unconstrained coral growth (as by many branching coral species) reduces the number of recruitment events needed to fill a habitat with coral relative to more spatially constrained growth (as by many massive species). Our model predicts that widespread mortality of branching corals (e.g., Acropora spp) and herbivorous sea urchins (particularly Diadema antillarum) in the Caribbean has greatly reduced the potential for restoration across the region.     

Climate‐change refugia: shading reef corals by turbidity

Coral reefs have recently experienced an unprecedented decline as the world's oceans continue to warm. Yet global climate models reveal a heterogeneously warming ocean, which has initiated a search for refuges, where corals may survive in the near future. We hypothesized that some turbid nearshore environments may act as climate‐change refuges, shading corals from the harmful interaction between high sea‐surface temperatures and high irradiance. We took a hierarchical Bayesian approach to determine the expected distribution of 12 coral species in the Indian and Pacific Oceans, between the latitudes 37°N and 37°S, under representative concentration pathway 8.5 (W m^?2) by 2100. The turbid nearshore refuges identified in this study were located between latitudes 20–30°N and 15–25°S, where there was a strong coupling between turbidity and tidal fluctuations. Our model predicts that turbidity will mitigate high temperature bleaching for 9% of shallow reef habitat (to 30 m depth) – habitat that was previously considered inhospitable under ocean warming. Our model also predicted that turbidity will protect some coral species more than others from climate‐change‐associated thermal stress. We also identified locations where consistently high turbidity will likely reduce irradiance to <250 μmol m^?2 s^?1, and predict that 16% of reef‐coral habitat ≤30 m will preclude coral growth and reef development. Thus, protecting the turbid nearshore refuges identified in this study, particularly in the northwestern Hawaiian Islands, the northern Philippines, the Ryukyu Islands (Japan), eastern Vietnam, western and eastern Australia, New Caledonia, the northern Red Sea, and the Arabian Gulf, should become part of a judicious global strategy for reef‐coral persistence under climate change.     

Sporadic Disturbances in Fluctuating Coral Reef Environments: El Nino and Coral Reef Development in the Eastern Pacific

The disastrous effects of the intense 1982–83 El Niño-SouthernOscillation (ENSO) bring new insight into the long-term developmentof eastern Pacific coral reefs. The 1988–83 ENSO sea surfacewarming event caused extensive reef coral bleaching (loss ofsymbiotic zooxanthellae), resulting in up to 70–95% coralmortality on reefs in Costa Rica, Panama, Colombia and Ecuador.In the Galapagos Islands (Ecuador), most coral reefs experienced>95% coral mortality. Also, several coral species experiencedextreme reductions in population size, and local and regionalextinctions. The El Niño event spawned secondary disturbances,such as increased predation and bioerosion, that continue toimpact reef-building corals. The death of Pocillopora colonieswith their crustacean guards eliminated coral barriers now allowingthe corallivore Acanthaster planci access to formerly protectedcoral prey. Sea urchins and other organisms eroded disturbedcorals at rates that exceed carbonate production, potentiallyresulting in the elimination of existing reef buildups. In otherreefbuilding regions following extensive, catastrophic coralmortality, rapid recovery often occurs through the growth ofsurviving corals, recruitment of new corals from nearby sourcepopulations, and survival of consolidated reef surfaces. Inthe eastern Pacific, however, the return of upwelling conditionsand the survival of coral predators and bioeroders hamper coralreef recovery by reducing recruitment success and eroding coralreef substrates. Thus, coral reef growth that occurs betweendisturbance events is not conserved. Repeated El Niñodisturbances, which have occurred throughout the recent geologichistory of the eastern Pacific, prevent coral communities fromincreasing in diversity and limit the development and persistenceof significant reef features. The poor development of easternPacific coral reefs throughout Holocene and perhaps much ofPleistocene time may result from recurrent thermal disturbancesof the intensity of the 1982–83 El Niño event.     

Disappearance of <Emphasis Type="Italic">Acropora</Emphasis> from the Marquesas (French Polynesia) during the last deglacial period

The major reef-building coral genus Acropora has never been recorded, living or fossil, from the Marquesas Islands in the central Pacific Ocean, which are characterized by limited modern reef formations. During the “Musorstom 9” cruise in 1997, investigations of marine platforms representing drowned reef systems revealed for the first time the presence of two Acropora species as fossils at seven Marquesas islands. The predominant species was Acropora valida, which was widespread in the archipelago and dated between 7.4 and 48.6 ka, providing evidence of an earlier Pacific distribution pattern broader than previously observed. It is proposed that disappearance of Acropora after 7.4 ka was linked to climatic events probably ENSO events controlling the distribution of corals and coral reefs in the eastern Pacific without excluding alternatively the effects of an increase in sea-level rise.     

Regional patterns of evolutionary turnover in Neogene coral reefs from the central Indo-West Pacific Ocean

The Indo-Pacific is an area of intense ecological interest, not least because of the region’s rich biodiversity. Important insights into the origins, evolutionary history, and maintenance of Indo-Pacific reef faunas depend upon the analysis of faunal occurrences derived from detailed stratigraphic sections. We investigated Neogene origination and extinction patterns derived from a combination of new coral occurrences and previously published records from the central Indo-West Pacific Ocean (cIWP, Indonesia, Papua New Guinea and Fiji). Two faunal turnover events were observed. In the first, an increase in generic richness of Scleractinia from the cIWP during the middle Miocene (17–14 Ma) coincided with both large-scale sea level fluctuations and the great Mid-Miocene collision event. We raise the hypothesis that Mid-Miocene origination was facilitated by habitat and population fragmentation associated with tectonism and sea level fall. The second, subsequent, turnover event was characterized by an overall lowering of generic diversity throughout the late Miocene and Pliocene (7–3 Ma), and was followed by a pronounced pulse of extinction at the Pliocene–Pleistocene boundary (~2.6 Ma). With the exception of the onset of Pleistocene sea-level cycles and the onset of northern hemisphere glaciation around 2.5 Ma, which might explain increased extinction during this time interval, there are no tectonic, eustatic, climatic or oceanographic events that neatly coincide with this second episode of Neogene coral taxonomic turnover. Our results reveal a total of 62 genera, including synonyms, from the Miocene to the Pleistocene. Neither episode of turnover among coral genera is exactly coincident with turnover in the Atlantic thus regional environmental change is found to drive Neogene reef dynamics.     

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.     

Coral Reef Habitat Response to Climate Change Scenarios

Coral reef ecosystems are threatened by both climate change and direct anthropogenic stress. Climate change will alter the physico-chemical environment that reefs currently occupy, leaving only limited regions that are conducive to reef habitation. Identifying these regions early may aid conservation efforts and inform decisions to transplant particular coral species or groups. Here a species distribution model (Maxent) is used to describe habitat suitable for coral reef growth. Two climate change scenarios (RCP4.5, RCP8.5) from the National Center for Atmospheric Research’s Community Earth System Model were used with Maxent to determine environmental suitability for corals (order Scleractinia). Environmental input variables best at representing the limits of suitable reef growth regions were isolated using a principal component analysis. Climate-driven changes in suitable habitat depend strongly on the unique region of reefs used to train Maxent. Increased global habitat loss was predicted in both climate projections through the 21^st century. A maximum habitat loss of 43% by 2100 was predicted in RCP4.5 and 82% in RCP8.5. When the model is trained solely with environmental data from the Caribbean/Atlantic, 83% of global habitat was lost by 2100 for RCP4.5 and 88% was lost for RCP8.5. Similarly, global runs trained only with Pacific Ocean reefs estimated that 60% of suitable habitat would be lost by 2100 in RCP4.5 and 90% in RCP8.5. When Maxent was trained solely with Indian Ocean reefs, suitable habitat worldwide increased by 38% in RCP4.5 by 2100 and 28% in RCP8.5 by 2050. Global habitat loss by 2100 was just 10% for RCP8.5. This projection suggests that shallow tropical sites in the Indian Ocean basin experience conditions today that are most similar to future projections of worldwide conditions. Indian Ocean reefs may thus be ideal candidate regions from which to select the best strands of coral for potential re-seeding efforts.     

Habitat-specific environmental conditions primarily control the microbiomes of the coral Seriatopora hystrix

Reef-building corals form complex relationships with a range of microorganisms including bacteria, archaea, fungi and the unicellular microalgae of the genus Symbiodinium, which together form the coral holobiont. These symbionts are known to have both beneficial and deleterious effects on their coral host, but little is known about what the governing factors of these relationships are, or the interactions that exist between the different members of the holobiont and their environment. Here we used 16S ribosomal RNA gene amplicon sequencing to investigate how archaeal and bacterial communities associated with the widespread scleractinian coral Seriatopora hystrix are influenced by extrinsic (reef habitat and geographic location) and intrinsic (host genotype and Symbiodinium subclade) factors. Bacteria dominate the microbiome of S. hystrix, with members of the Alphaproteobacteria, Gammaproteobacteria and Bacteriodetes being the most predominant in all samples. The richness and evenness of these communities varied between reef habitats, but there was no significant difference between distinct coral host lineages or corals hosting distinct Symbiodinium subclades. The coral microbiomes correlated to reef habitat (depth) and geographic location, with a negative correlation between Alpha- and Gammaproteobacteria, driven by the key members of both groups (Rhodobacteraceae and Hahellaceae, respectively), which showed significant differences between location and depth. This study suggests that the control of microbial communities associated with the scleractinian coral S. hystrix is driven primarily by external environmental conditions rather than by those directly associated with the coral holobiont.