Holocene reef building on eastern St. Croix, US Virgin Islands: Lang Bank revisited

New core and seismic data suggest that widespread reef building started on Lang Bank by 8,900 CalBP and was dominated by Acropora palmata for the next three millennia. Accretion rates averaged 5.81 m ky^?1, a rate that was sufficient for reefs to keep pace with rising sea level on the bank throughout their history. Seismic data show a deep platform interior that was flooded well in advance of reef building along the elevated rim. As a result, those reefs were buffered from sediment stress by their higher positions and active water flow to the west. A. palmata disappeared from the shallow margin by 6,350 yr ago, and reef building on Lang Bank largely ceased by 5,035 CalBP. The reasons for these dramatic events are unclear. Water depth over the reefs was generally shallower than when they started to build, and sea level was slowing dramatically. The new data described here show that reefs flourished on Lang Bank throughout the hiatus suggested by earlier studies (10–7 kyrs BP), and the ultimate demise of shelf-edge reefs is clearly not associated with either poor water quality or sudden sea-level rise. In addition, accretion rates from eastern St. Croix and throughout the Caribbean were well below the high values (≥10 m ky^?1) that have been widely assumed. These data collectively argue against models that require extreme environmental or oceanographic phenomena to drown reefs on Lang Bank where reef building was too fast to be outpaced by Holocene sea-level rise. This also bears on more generalized Caribbean models that depend on the presumed reef history on eastern St. Croix.     

Numerical modeling of the impact of sea-level rise on fringing coral reef hydrodynamics and sediment transport

Most climate projections suggest that sea level may rise on the order of 0.5–1.0 m by 2100; it is not clear, however, how fluid flow and sediment dynamics on exposed fringing reefs might change in response to this rapid sea-level rise. Coupled hydrodynamic and sediment-transport numerical modeling is consistent with recent published results that suggest that an increase in water depth on the order of 0.5–1.0 m on a 1–2 m deep exposed fringing reef flat would result in larger significant wave heights and setup, further elevating water depths on the reef flat. Larger waves would generate higher near-bed shear stresses, which, in turn, would result in an increase in both the size and the quantity of sediment that can be resuspended from the seabed or eroded from adjacent coastal plain deposits. Greater wave- and wind-driven currents would develop with increasing water depth, increasing the alongshore and offshore flux of water and sediment from the inner reef flat to the outer reef flat and fore reef where coral growth is typically greatest. Sediment residence time on the fringing reef flat was modeled to decrease exponentially with increasing sea-level rise as the magnitude of sea-level rise approached the mean water depth over the reef flat. The model results presented here suggest that a 0.5–1.0 m rise in sea level will likely increase coastal erosion, mixing and circulation, the amount of sediment resuspended, and the duration of high turbidity on exposed reef flats, resulting in decreased light availability for photosynthesis, increased sediment-induced stress on the reef ecosystem, and potentially affecting a number of other ecological processes.     

Acropora palmata reef framework: A reliable indicator of sea level in the western atlantic for the past 10,000 years

Summary A minimum sea-level curve for the past 10,000 years has been constructed on the basis of radiocarbon dates of Acropora palmata (Lamarck) samples from the shallow-water framework of both relict and modern reefs of the tropical western Atlantic. A. palmata framework is a reliable reference for reconstructing the history of late Quaternary sea levels owing to its restricted depth range (<1 to 5 m), the lack of postdepositional transport of A. palmata framework, the ease of obtaining uncontaminated samples, and the minimal compaction of A. palmata reef facies. The minimum sea-level curve constructed in this study is useful not only in evaluating the reliability of present and future Holocene sea-level curves for the western Atlantic, but also in estimating paleo-water depths in the study of Holocene reef history of this area.     

Kenyan coral reef lagoon fish: effects of fishing,substrate complexity,and sea urchins

Population density, number of species, diversity, and species-area relationships of fish species in eight common coral reef-associated families were studied in three marine parks receiving total protection from fishing, four sites with unregulated fishing, and one reef which recently received protection from fishing (referred to as a transition reef). Data on coral cover, reef topographic complexity, and sea urchin abundance were collected and correlated with fish abundance and species richness. The most striking result of this survey is a consistent and large reduction in the population density and species richness of 5 families (surgeonfish, triggerfish, butterflyfish, angelfish, and parrotfish). Poor recovery of parrotfish in the transition reef, relative to other fish families, is interpreted as evidence for competitive exclusion of parrotfish by sea urchins. Reef substrate complexity is significantly associated with fish abundance and diversity, but data suggest different responses for protected versus fished reefs, protected reefs having higher species richness and numbers of individuals than unprotected reefs for the same reef complexity. Sea urchin abundance is negatively associated with numbers of fish and fish species but the interrelationship between sea urchins, substrate complexity, coral cover, and management make it difficult to attribute a set percent of variance to each factor-although fishing versus no fishing appears to be the strongest variable in predicting numbers of individuals and species of fish, and their community similarity. Localized species extirpation is evident for many species on fished reefs (for the sampled area of 1.0 ha). Fifty-two of 110 species found on protected reefs were not found on unprotected reefs.     

Ramp morphology controlling the facies differentiation of a Late Ordovician reef complex at Bachu,Tarim Block,NW China

A carbonate ramp in the shallow‐marine northwestern part of the Central Tarim Uplift, Bachu, NW China, exhibits an extraordinary Late Ordovician reef complex along the Lianglitag Mountains, exposed for a distance of about 25 km. Seven localities within the ‘Middle Red Limestone’ of the Upper Member of the Lianglitag Formation (Katian, Late Ordovician) illustrated the changes in biofacies and lithofacies: northern, seaward‐directed patch reefs are replaced towards the south by coeval grain banks. The patch reef units are dominated by microbial and calcareous algal components. The reefs at the northernmost locality are knoll‐shaped, kalyptra‐shaped or irregularly shaped with sizes of individual reefs increasing from about 2 m in height and diameter. Stratigraphically upward, reefs notably expand to larger structures by several mounds coalescing; they are generally about 10 m thick and tens of metres in lateral extent. The maximum thickness of the main patch reef is more than 30 m, and its diameter is around 100 m. The reefal units turn into biostromes with gentler relief southward and still further south grade into banks composed of peloids and coated grains. The southernmost locality is still a shallow‐water bank, and the coastline is not documented in the study area. The present evidence indicates that the Late Ordovician palaeo‐oceanography provided a number of environments for the optimal growth of carbonate build‐ups; microbial‐calcareous algal communities could thrive in areas where the innovative metazoan reef frameworks consisting of corals and stromatoporoids did not play a significant role. The ramp morphology, especially changes in water depth, controlled the configuration of the reef complex.     

Increased sea level promotes coral cover on shallow reef flats in the Andaman Sea, eastern Indian Ocean

Sea level in the Indian Ocean is subject to considerable temporal and spatial variabilities. During the period 1960–2009 at Phuket, Thailand, in the NE Indian Ocean, mean sea level increased by 2.7 mm y⁻¹. Regular monitoring of coral cover on fringing reef flats at Phuket since 1979 revealed a sensitive response of this habitat to both transient sea-level depressions and sea-level elevation. Since 1987 when more frequent sampling began, coral cover was positively correlated with the mean sea level experienced over the preceding months. Changing mean sea level explained a high proportion of the observed variation in cover, with overall increasing sea levels and a lack of negative sea-level anomalies promoting cover on the outer reef flats. Concomitantly, there have been no changes in reef community structure or any apparent shifts in zonation patterns across the reef. While future benefits of continued increases in mean sea level on reef flats in the region will be constrained by the frequency and intensity of sea-level depressions associated with the Indian Ocean Dipole, and bleaching events, the overall picture for these shallow reefs is a positive one as they respond to increasing sea level and show rapid recovery from environmental disturbances.     

LIDAR optical rugosity of coral reefs in Biscayne National Park,Florida

The NASA Experimental Advanced Airborne Research Lidar (EAARL), a temporal waveform-resolving, airborne, green wavelength LIDAR (light detection and ranging), is designed to measure the submeter-scale topography of shallow reef substrates. Topographic variability is a prime component of habitat complexity, an ecological factor that both expresses and controls the abundance and distribution of many reef organisms. Following the acquisition of EAARL coverage over both mid-platform patch reefs and shelf-margin bank reefs within Biscayne National Park in August 2002, EAARL-based optical indices of topographic variability were evaluated at 15 patch reef and bank reef sites. Several sites were selected to match reefs previously evaluated in situ along underwater video and belt transects. The analysis used large populations of submarine topographic transects derived from the examination of closely spaced laser spot reflections along LIDAR raster scans. At all 15 sites, each LIDAR transect was evaluated separately to determine optical rugosity (Ro_tran), and the average elevation difference between adjacent points (Av(E_ap)). Further, the whole-site mean and maximum values of Ro_tran and Av(E_ap) for the entire population of transects at each analysis site, along with their standard deviations, were calculated. This study revealed that the greater habitat complexity of inshore patch reefs versus outer bank reefs results in relative differences in topographic complexity that can be discerned in the laser returns. Accordingly, LIDAR sensing of optical rugosity is proposed as a complementary new technique for the rapid assessment of shallow coral reefs.     

Deep reefs are not refugium for shallow‐water fish communities in the southwestern Atlantic

1. The deep reef refugia hypothesis (DRRH) predicts that deep reef ecosystems may act as refugium for the biota of disturbed shallow waters. Because deep reefs are among the most understudied habitats on Earth, formal tests of the DRRH remain scarce. If the DRRH is valid at the community level, the diversity of species, functions, and lineages of fish communities of shallow reefs should be encapsulated in deep reefs.
2. We tested the DRRH by assessing the taxonomic, functional, and phylogenetic diversity of 22 Brazilian fish communities between 2 and 62 m depth. We partitioned the gamma diversity of shallow (<30 m) and deep reefs (>30 m) into independent alpha and beta components, accounted for species’ abundance, and assessed whether beta patterns were mostly driven by spatial turnover or nestedness.
3. We recorded 3,821 fishes belonging to 85 species and 36 families. Contrary to DRRH expectations, only 48% of the species occurred in both shallow and deep reefs. Alpha diversity of rare species was higher in deep reefs as expected, but alpha diversity of typical and dominant species did not vary with depth. Alpha functional diversity was higher in deep reefs only for rare and typical species, but not for dominant species. Alpha phylogenetic diversity was consistently higher in deep reefs, supporting DRRH expectations.
4. Profiles of taxonomic, functional, and phylogenetic beta diversity indicated that deep reefs were not more heterogeneous than shallow reefs, contradicting expectations of biotic homogenization near sea surface. Furthermore, pairwise beta‐diversity analyses revealed that the patterns were mostly driven by spatial turnover rather than nestedness at any depth.
5. Conclusions. Although some results support the DRRH, most indicate that the shallow‐water reef fish diversity is not fully encapsulated in deep reefs. Every reef contributes significantly to the regional diversity and must be managed and protected accordingly.

     

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.     

Holocene reef evolution in a macrotidal setting: Buccaneer Archipelago,Kimberley Bioregion,Northwest Australia

This study uses information derived from cores to describe the Holocene accretion history of coral reefs in the macrotidal (up to 11 m tidal range) Buccaneer Archipelago of the southern Kimberley coast, Western Australia. The internal architecture of all cored reefs is broadly similar, constituting well-preserved detrital coral fragments, predominantly branching Acropora, in a poorly sorted sandy mud matrix. However, once the reefs reach sea level, they diverge into two types: low intertidal reefs that maintain their detrital character and develop relatively narrow, horizontal or gently sloping reef flats at approximately mean low water spring, and high intertidal reefs that develop broad coralline algal-dominated reef flats at elevations between mean low water neap and mean high water neap. The high intertidal reefs develop where strong, ebb-dominated, tidal asymmetry retains seawater over the low tide and allows continued accretion. Both reef types are ultimately constrained by sea level but differ in elevation by 3–4 m.     

The impact of the Mid-Pleistocene Transition on the composition of submerged reefs of the Maui Nui Complex,Hawaii

The submarine reef terraces (L1–L12) of the Maui Nui Complex (MNC—the islands of Lanai, Molokai, Maui and Kahoolawe) in Hawaii provide a unique opportunity to investigate the impact of climate and sea level change on coral reef growth by examining changes in reef development through the Mid-Pleistocene Transition (900–800 ka). We present an analysis of the biological and sedimentary composition of the reefs that builds directly on recently published chronological and morphological data. We define nine distinct limestone facies and place them in a spatial and stratigraphic context within 12 reef terraces using ROV and submersible observations. These include oolitic, two coral reef, two coralline algal nodule, algal crust, hemi-pelagic mud, bioclastic and peloidal mud facies. These facies characterise environments from high energy shallow water coral reef crests to low energy non-reefal deep-water settings. Combining the bottom observations and sedimentary facies data, we report a shift in the observed sedimentary facies across the submerged reefs of the MNC from dominant shallow coral reef facies on the deep reefs to coralline algae dominated exposed outcrop morphology on the shallower reefs. We argue that this shift is a reflection of the change in period and amplitude of glacioeustatic sea level cycles (41 kyr and 60–70 m to 100 kyr and 120 m) during the Mid-Pleistocene Transition (MPT, ~ 800 ka), coupled with a slowing in the subsidence rate of the complex. The growth of stratigraphically thick coral reef units on the deep Pre-MPT reefs was due to the rapid subsidence of the substrate and the shorter, smaller amplitude sea level cycles allowing re-occupation and coral growth on successive cycle low-stands. Longer, larger amplitude sea level cycles after the MPT combined with greater vertical stability at this time produced conditions conducive to deep-water coralline algae growth which veneered the shallower terraces. Additionally, we compare reef development both within the MNC, and between the MNC and Hawaii. Finally we suggest that climatic forcings such as sea-surface temperature and oceanographic currents may also have influenced the distribution of coral species within the sample suite, e.g., the disappearance of the Acropora genus from the Maui Nui Complex in the Middle Pleistocene.     

Shifting ecological baselines and the demise of Acropora cervicornis in the western North Atlantic and Caribbean Province: a Pleistocene perspective

 In recent years, marine scientists have become increasingly alarmed over the decline of live coral cover throughout the Caribbean and tropical western Atlantic region. The Holocene and Pleistocene fossil record of coral reefs from this region potentially provides a wealth of long-term ecologic information with which to assess the historical record of changes in shallow water coral reef communities. Before fossil data can be applied to the modern reef system, critical problems involving fossil preservation must be addressed. Moreover, it must be demonstrated that the classic reef coral zonation patterns described in the early days of coral reef ecology, and upon which “healthy” versus “unhealthy” reefs are determined, are themselves representative of reefs that existed prior to any human influence. To address these issues, we have conducted systematic censuses of life and death assemblages on modern “healthy” patch reefs in the Florida reef tract that conform to the classic Caribbean model of reef coral zonation, and a patch reef in the Bahamas that is currently undergoing a transition in coral dominance that is part of a greater Caribbean-wide phenomenon. Results were compared to censuses of ancient reef assemblages preserved in Pleistocene limestones in close proximity to each modern reef. We have determined that the Pleistocene fossil record of coral reefs may be used to calibrate an ecological baseline with which to compare modern reef assemblages, and suggest that the current and rapid decline of Acropora cervicornis observed on a Bahamian patch reef may be a unique event that contrasts with the long-term persistence of this taxon during Pleistocene and Holocene time. Accepted: 19 May 1998     

Oldest scleractinian coral reefs on the North American craton: Upper Triassic (Carnian), northeastern British Columbia,Canada

Bioclastic accumulations composed of crinoids, brachiopods, molluscs, spongiomorphs and scleractinian corals occur within Upper Triassic strata of the lower Baldonnel Formation at Pardonet Hill in northeastern British Columbia Canada. These small buildups (∼100 to 500 m³) have planar bases and broadly convex tops. These mounds are interpreted as small patch reefs composed of packstone, bioclastic floatstone/rudstone and carbonate breccia intercalated with mixed siliciclastic carbonate sediments deposited in a shallow subtidal setting (i.e. above fairweather wave base). Amalgamated hummocky cross-stratified to current ripple-laminated, quartz-dominated sandstone beds and numerous sharp-based, normally graded bioclastic (commonly encrinitic) packstone/grainstone — quartz–sandstone couplets characterize inter-reef lithologies.Conodont biostratigraphy indicates that the Pardonet Hill patch reefs occur within strata dated as earliest Upper Carnian (lower nodosus zone). The Pardonet Hill patch reefs originated and developed during an interval of regional sea level lowstand. Strata within which these patch reefs occur represent the westernmost migration of the Triassic shoreline in western Canada. Disappearance of coral reefs in the study area may have been affected by rapid marine transgression and failure of reef faunas to recolonize the new shore zone further to the east.The Pardonet Hill locality occurred on the western margin of the North American craton during the Triassic. Prior to their discovery reef-like structures dominated by corals in the western Panthalassa were limited to allochthonous terranes (now part of the Cordillera). The Pardonet Hill patch reefs occur at approximately 30° Triassic paleolatitude. In modern settings, this is at the extreme latitudinal margin of subtropical zooxanthellate reef development. The presence of benthic faunas characteristic of low-paleolatitude settings on the northwestern coast of Pangea has significant implications in paleotectonic and paleoenvironmental reconstructions.     

Reductions in the mitochondrial DNA diversity of coral reef fish provide evidence of population bottlenecks resulting from Holocene sea-level change

This study investigated the influence of reproductive strategy (benthic or pelagic eggs) and habitat preferences (lagoon or outer slope) on both diversity and genetic differentiation using a set of populations of seven coral reef fish species over different geographic scales within French Polynesia. We hypothesized that a Holocene sea-level decrease contributed to severe reduction of population size for species inhabiting lagoons and a subsequent decrease of genetic diversity. Conversely, we proposed that species inhabiting stable environments, such as the outer slope, should demonstrate higher genetic diversity but also more structured populations because they have potentially reached a migration-genetic drift equilibrium. Sequences of the 5' end of the mitochondrial DNA (mtDNA) control region were compared among populations sampled in five isolated islands within two archipelagos of French Polynesia. For all the species, no significant divergences among populations were found. Significant differences in mtDNA diversity between lagoonal and outer-slope species were demonstrated both for haplotype diversity and sequence divergence but none were found between species with different egg types. Pairwise mismatch distributions suggested rapid population growth for all the seven species involved in this study, but they revealed different distributions, depending on the habitat preference of the species. Although several scenarios can explain the observed patterns, the hypothesis of population size reduction events relative to Holocene sea-level regression and its consequence on French Polynesia coral reefs is the most parsimonious. Outer-slope species have undergone a probable weak and/or old bottleneck (outer reefs persisted during low sea level, leading to reef area reductions), whereas lagoonal species suffered a strong and/or recent bottleneck since Holocene sea-level regression resulted in the drying out of all the atolls that are maximum 70 meters deep. Since present sea level was reached between 5000 and 6000 years ago, different demographic events (bottlenecks or founder events) have lead to the actual populations of lagoons in French Polynesia.     

Indo-Pacific and Atlantic spurs and grooves revisited: the possible effects of different Holocene sea-level history,exposure, and reef accretion rate in the shallow fore reef

Shallow fore-reef areas worldwide are usually characterized by spurs and grooves. A comparison of examples from the three world oceans suggests that Indo-Pacific spurs and grooves are shaped predominantly by erosion, whereas western Atlantic spur and groove systems are largely a product of constructive processes. I propose that this difference is caused by regional differences in Holocene sea-level change, which controlled exposure to waves and currents, and reef-accretion rates. The transgressive–regressive sea-level curve in the Indo-Pacific realm, i.e., the Mid-to-Late Holocene sea-level fall in these areas has maintained high-energy conditions in the shallow fore reef. Higher exposure to waves and currents favors erosion and leads to a dominance of crustose coralline algae that have relatively slow growth rates. In the western Atlantic, the transgressive Holocene sea level has caused Mid-to-Late Holocene deepening and has maintained accommodation space for reef accretion. Fast-growing acroporid corals thrive under lower exposure and are more common than coralline algae. The fossil record of the spur and groove system is rather poor, which is probably a consequence of the need of excellent, three-dimensional outcrops for identification.     

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

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

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.     

The origin of Jurassic reefs: Current research developments and results

Summary In order to elucidate the control of local, regional and global factors on occurrence, distribution and character of Jurassic reefs, reefal settings of Mid and Late Jurassic age from southwestern Germany, Iberia and Romania were compared in terms of their sedimentological (including diagenetic), palaeoecological, architectural, stratigraphic and sequential aspects. Upper Jurassic reefs of southern Germany are dominated by siliceous sponge—microbial crust automicritic to allomicritic mounds. During the Oxfordian these form small to large buildups, whereas during the Kimmeridgian they more frequently are but marginal parts of large grain-dominated massive buildups. Diagenesis of sponge facies is largely governed by the original composition and fabric of sediments. The latest Kimmeridgian and Tithonian spongiolite development is locally accompanied by coral facies, forming large reefs on spongiolitic topographic elevations or, more frequently, small meadows and patch reefs within bioclastic to oolitic shoal and apron sediments. New biostratigraphic results indicate a narrower time gap between Swabian and Franconian coral development than previously thought. Palynostratigraphy and mineralostratigraphy partly allow good stratigraphic resolution also in spongiolitic buildups, and even in dolomitised massive limestones. Spongiolite development of the Bajocian and Oxfordian of eastern Spain shares many similarities. They are both dominated by extensive biostromal development which is related to hardground formation during flooding events. The Upper Jurassic siliceous sponge facies from Portugal is more localised, though more differentiated, comprising biostromal, mudmound and sponge-thrombolite as well as frequent mixed coral-sponge facies. The Iberian Upper Jurassic coral facies includes a great variety of coral reef and platform types, a pattern which together with the analysis of coral associations reflects the great variability of reefal environments. Microbial reefs ranging from coralrich to siliceous sponge-bearing to pure thrombolites frequently developed at different water depths. Reef corals even thrived within terrigeneous settings. In eastern Romania, small coral reefs of various types as well as larger siliceous sponge-microbial crust mounds grew contemporaneously during the Oxfordian, occupying different bathymetric positions on a homoclinal ramp. Application of sequence stratigraphic concepts demonstrates that onset or, in other cases, maximum development of reef growth is related to sea level rise (transgressions and early highstand) which caused a reduction in allochthonous sedimentation. The connection of reef development with low background sedimentation is corroborated by the richness of reefs in encrusting organisms, borers and microbial crusts. Microbial crusts and other automicrites can largely contribute to the formation of reef rock during allosedimentary hiatuses. However, many reefs could cope with variable, though reduced, rates of background sedimentation. This is reflected by differences in faunal diversities and the partial dominance of morphologically adapted forms. Besides corals, some sponges and associated brachiopods show distinct morphologies reflecting sedimentation rate and substrate consistency. Bathymetry is another important factor in the determination of reefal composition. Not only a generally deeper position of siliceous sponge facies relative to coral facies, but also further bathymetric differentiation within both facies groups is reflected by changes in the composition, diversity and, partly, morphology of sponges, corals, cementing bivalves and microencrusters. Criteria such as authigenic glauconite, dysaerobic epibentic bivalves,Chondrites burrows or framboidal pyrite in the surrounding sediments of many Upper Jurassic thrombolitic buildups suggest that oxygen depletion excluded higher reefal metazoans in many of these reefs. Their position within shallowing-upwards successions and associated fauna from aerated settings show that thrombolitic reefs occurred over a broad bathymetric area, from moderately shallow to deep water. Increases in the alkalinity of sea water possibly enhanced calcification. Reefs were much more common during the Late Jurassic than during the older parts of this period. Particularly the differences between the Mid and Late Jurassic frequencies of reefs can be largely explained by a wider availability of suitable reef habitats provided by the general sea level rise, rather than by an evolutionary radiation of reef biota. The scarcity of siliceous sponge reefs on the tectonically more active southern Tethyan margin as well as in the Lusitanian Basin of west-central Portugal reflects the scarcity of suitable mid to outer ramp niches. Coral reefs occurred in a larger variety of structural settings. Upper Jurassic coral reefs partly grew in high latitudinal areas suggesting an equilibrated climate. This appears to be an effect of the buffering capacity of high sea level. These feedback effects of high sea level also may have reduced oceanic circulation particularly during flooding events of third and higher order, which gave rise to the development of black shales and dysaerobic thrombolite reefs. Hence, the interplay of local, regional and global factors caused Jurassic reefs to be more differentiated than modern ones, including near-actualistic coral reefs as well as non-actualistic sponge and microbial reefs.