Cross-shelf differences in the pattern and pace of bioerosion of experimental carbonate substrates exposed for 3 years on the northern Great Barrier Reef, Australia

Patterns of bioerosion of dead corals and rubbles on the northern Great Barrier Reef were studied by using blocks of the massive coral Porites experimentally exposed at six sites, located on an inshore–offshore profile, for 1 year and 3 years. Rates of microbioerosion by microborers, grazing by fish, and macrobioerosion by filter-feeding organisms were simultaneously evaluated using image analysis. Microbioerosion, grazing, and total bioerosion were lower at reefs near the Queensland coast than at the edge of the continental shelf (1.81 kg m⁻² and 6.07 kg m⁻² after 3 years of exposure respectively, for total bioerosion). The opposite pattern was observed for macrobioerosion. Bioaccretion was negligible. These patterns were evident after 1 year of exposure, and became enhanced after 3 years. Microborers were established and were the main agent of bioerosion after 1 year of exposure, and as the principal support for grazing, continued to be the main cause of carbonate loss after 3 years. Full grazing activity and establishment of a mature community of macroborers required more than 1 year of exposure. After 1 year, macroborers and grazers were the second most important agents of bioerosion on both inshore and offshore reefs. However, after 3 years, grazers became the main agents at all sites except at the inshore sites, where macroborers were the principal agents. Because the contribution of microborers, grazers, and macroborers to bioerosion varies in space and time, we suggest that the estimation of reef carbonate budgets need to take in account the activities of all bioerosion agents.     

Distribution patterns and bioerosion of the sea urchin Centrostephanus coronatus (Diadematoida: Diadematidae), at the reef of Playa Blanca, Colombian Pacific

Regular sea-urchins are one of the main bioeroding organisms affecting coral reefs around the world. The abundance, distribution and bioerosion rate of the sea-urchin Centrostephanus coronatus, were determined in different reef zones of Playa Blanca fringing reef (Gorgona Island, Colombian pacific coast) during 1997 and 1998. The erosion rates were determined calcinating the gut content of the sea-urchins to eliminate all organic components and preserve the inorganic portion of calcium carbonate. C. coronatus showed the highest densities towards the central zones of the reef (plain-crest and front) (12.4 ind/m2; range 0-48 ind/m2). The highest mean bioerosion rate was 0.103 kgCaCO3/m2/yr in the reef plain-crest (0-0.69 kgCaCO3/m2/yr). In the other zones, (back reef and reef front) the mean bioerosion rates were 0.071 (range 0-0.39) and 0.052 (range 0-0.31) kgCaCO3/m2/yr respectively. According to the present data, it can be seen that the destruction of coralline skeletons, produced in this reef by sea-urchins is rather low, compared with the abrasion caused by these organisms in other places of the world. However, the combined action of C. coronatus and other bioeroding organisms (borers and grazers). along with some adverse environmental factors to corals, can be causing a negative balance between normal processes of reef accretion-destruction in Gorgona Island reefs.     

Spatial and temporal patterns of non-colonial boring organisms (polychaetes,sipunculans and bivalve molluscs) in Porites at Lizard Island,Great Barrier Reef

A study of the spatial and temporal patterns of colonisation by non-colonial boring organisms to dead Porites coral substrate was conducted at Lizard Island, Great Barrier Reef over a 4 year period. These fluctuations were analysed for each group of borers, and most exhibited strong site preferences, with preferred sites being on the windward slope in 10 m and on the reef flat in 1 m. A lagoonal patch reef site exhibited consistently low colonisation. Most groups showed inter-year variations in colonisation with spring/early summer dominating. These variations are discussed in terms of what is known about their life histories. These results together with those of Kiene (in preparation) which document varying rates of bioerosion, at these sites over the same time period, demonstrate that variations in borer colonisation are responsible for the variations in rates of bioerosion calculated. Thus rates of bioerosion by borers will vary significantly between different reef environments.     

Comments on coral reef regeneration, bioerosion, biogeography, and chemical ecology: future directions

This paper represents a brief review of three processes operating on coral reefs and the results of studies of a fourth process, and how those results may be applied to the benefit of humankind. The areas are reef regeneration processes; bioerosion; dispersal, recruitment and biogeography of corals; and chemical ecology and natural products chemistry of reef organisms. Possible future directions for research will also be considered in each area. Regarding reef regeneration processes, coral reefs are degrading rapidly on a global scale due to over-fishing, fishing techniques causing habitat destruction, deforestation, mass mortalities of key reef species, nutrient enrichment and sedimentation. Seeding of reefs with the larvae of corals and other key reef organisms, such as echinoids, may help to promote and enhance reef regeneration in the future. Such techniques will be made possible by studies of the embryology, larval ecology, dispersal and recruitment processes, and related local physical oceanographic processes. Regarding bioerosion, both internal and external bioerosion are affected by grazers and predators. Bioerosion is also affected by nutrient enrichment, as shown through correlative studies (Great Barrier Reef) and studies of opportunity (Kaneohe Bay). Ongoing experiments such as ENCORE will help to answer questions about the role of dissolved nutrients in enhancing internal bioerosion. Questions still remain, however, regarding the role of particulates in promoting internal bioerosion and the resultant weakening of and negative growth in the reef framework. Regarding dispersal, recruitment and the biogeography of corals, it is now known that most species of coral reproduce via broadcasting, although there appear to be proportionally more brooders in the Caribbean than in the western Pacific. Differential extinctions in the western Pacific vs. the western Atlantic have contributed to the biogeographic distribution of corals we observe today and the concentric isoclines of species diversity in numerous reef organisms in the western Pacific. The role of reproductive mode in contributing to these patterns is, however, still not understood. Investigations into the roles of different larval longevities and reproductive modes may help us answer questions regarding their differential distribution and the potential effects of major perturbations such as global warming on future distributions. With respect to the chemical ecology of alcyonacean octocorals (soft corals), four functions have been determined thus far for secondary metabolites in this group, anti-predation, anti-competition (allelopathy), anti-fouling, and enhancement of reproductive success. Investigations of alcyonacean reproduction has revealed that it may be necessary for several secondary metabolites to be present simultaneously before a function may be realized or fully manifested. This raises questions regarding the manner in which novel compounds are tested by medical laboratories for bioactivity using a single compound. Simultaneously testing of multiple compounds derived from a single organism may be necessary in the future to reveal potential valuable synergistic bioactivity. Also, some novel secondary metabolites may have other valuable commercial applications, as is the case with the UV-absorbing compounds of corals and other reef organisms found on the Great Barrier Reef. In order to avoid overlooking medically or commercially valuable functions of these compounds, broader testing may be necessary.     

A Novel μCT Analysis Reveals Different Responses of Bioerosion and Secondary Accretion to Environmental Variability

Corals build reefs through accretion of calcium carbonate (CaCO₃) skeletons, but net reef growth also depends on bioerosion by grazers and borers and on secondary calcification by crustose coralline algae and other calcifying invertebrates. However, traditional field methods for quantifying secondary accretion and bioerosion confound both processes, do not measure them on the same time-scale, or are restricted to 2D methods. In a prior study, we compared multiple environmental drivers of net erosion using pre- and post-deployment micro-computed tomography scans (μCT; calculated as the % change in volume of experimental CaCO₃ blocks) and found a shift from net accretion to net erosion with increasing ocean acidity. Here, we present a novel μCT method and detail a procedure that aligns and digitally subtracts pre- and post-deployment μCT scans and measures the simultaneous response of secondary accretion and bioerosion on blocks exposed to the same environmental variation over the same time-scale. We tested our method on a dataset from a prior study and show that it can be used to uncover information previously unattainable using traditional methods. We demonstrated that secondary accretion and bioerosion are driven by different environmental parameters, bioerosion is more sensitive to ocean acidity than secondary accretion, and net erosion is driven more by changes in bioerosion than secondary accretion.     

Sponge borehole size as a relative measure of bioerosion and paleoproductivity

Bioerosion intensity has been proposed as a measure of paleoproductivity in fossil reefs, but it is difficult to measure directly because fossil corals are often incomplete and because it is difticult to infer the length of time a given coral was exposed to bioeroding organisms. Both nutrient availability and taphonomic factors can affect bioerosion intensity as measured in dead corals. Here, we examine these two effects separately using data from previous studies on bioerosion in modern and fossil corals. Size of individual sponge borings accurately reflects total bioerosion in modern massive and branching corals on the Great Barrier Reef. Total bioerosion in both massive and branching corals decreases outward across the continental shelf, paralleling trends in nutrient availability. Size of individual Cliothosa hancocki borings decreases across the shelf in branching Acropora but not in massive Porites. Fossil sponge borings Entobia convoluta and Uniglobites glomerata in massive corals from Oligocene and Miocene reefs in Puerto Rico are smallest in Oligocene shelf-edge reefs, intermediate in Oligocene patch reefs, and largest in Miocene patch reefs. Both facies-related influence, represented by Oligocene shelf-edge reefs vs. Oligocene patch reefs, and nutrient-related influence, represented by Oligocene vs. Miocene patch reefs, were reflected in the size of sponge boreholes. Size of sponge borings also varies among species of host corals, apparently in relation to skeletal architecture. Borehole size is inversely correlated with skeletal density as measured by the relative proportion of skeleton and pore space in transverse thin section. There is a weak positive correlation between borehole size and corallite diameter. These findings contradict reported positive correlations between total bioerosion and bulk density in modern corals. Borehole size appears accurately to reflect intensity of total internal bioerosion in fossil corals. Facies-controlled taphonomic overprints and influence of skeletal differences between coral species limit the use of sponge borehole size to a rough indicator of paleoproductivity in fossil coral reef environments.     

Drivers of region-wide declines in architectural complexity on Caribbean reefs

Severe declines in the cover of live hard coral on reefs have been reported worldwide, and in the Caribbean region, the architectural complexity of coral reefs has also declined markedly. While the drivers of coral cover loss are relatively well understood, little is known about the drivers of regional-scale declines in architectural complexity. We have used a dataset of 49 time series reporting reef architectural complexity to explore the effect of hurricanes, coral bleaching and fishing on Caribbean-wide annual rates of change in reef complexity. Hurricane impacts greatly influence reef complexity, with the most rapid rates of decline in complexity occurring at sites impacted during their survey period, and with lower rates of loss occurring at unimpacted sites. Reef architectural complexity did not change significantly following mass bleaching events (in a time frame of <5 years) or positive thermal anomalies. Although the rates of change in architectural complexity were similar in and out of marine protected areas (MPAs), significant declines in complexity were observed inside but not outside of MPAs, possibly because reductions in fishing can lead to increased bioerosion by herbivores within MPAs. Our findings suggest that major drivers of coral mortality, such as coral bleaching, do not influence reef architectural complexity in the short term (<5 years). Instead, direct physical impacts and reef bioerosion appear to be important drivers of the widespread loss of architecturally complex reefs in the Caribbean.     

Macroborers within coral framework at Discovery Bay, north Jamaica: species distribution and abundance, and effects on coral preservation

 Macroboring organisms are recognised as key agents of reef framework modification and destruction, and while recent studies in the Pacific have improved understanding of spatial variations in macroboring community structure, and rates of macroboring within individual reefs, comparable studies from the Caribbean are largely lacking. This study assesses the distribution of macroboring species and the degree of framework infestation across the reefs at Discovery Bay, north Jamaica. Although individual species of borers exhibit variable distributions across the reef, relative abundances of the main groups of macroborers (sponges, bivalves, worms) illustrate clear distributional trends. Sponges are dominant at fore-reef sites, while sipunculan and polychaete worms are only of importance at back-reef/lagoon and shallow fore-reef sites. Bivalves are locally important within back-reef and lagoon patch reef framework. Average percentages of internal bioerosion (macroboring) vary between sites, but are highest at back-reef and deep fore-reef sites. No systematic pattern of variation occurs within back-reef/lagoon samples, but a significant trend of increased macroboring is recognised with increased water depth on the fore-reef. In addition, significant differences in terms of the susceptibility of individual coral species are recognised. These factors are likely to result in biasing of the fossil record, with variable styles of preservation evident both between sites (i.e. with depth/environment) and within sites (i.e. between coral species). Accepted: 1 June 1998     

Effects of eutrophication on bioeroding sponge communities with the description of new West Indian sponges, Cliona spp. (Porifera: Hadromerida: Clionidae)

Abstract. Pieces of coral rubble ( Porites porites ), collected from across 3 fringing reefs that lie along a eutrophication gradient, were examined for the presence of clionid sponges. A similarity analysis of species composition showed that reef zone had less effect on clionid community composition than did other factors affecting the reefs as a whole. Except on the back reef, the Zones, distances, and depths within the reefs had no significant influence on the number of clionid invasions. Reef comparisons demonstrated that clionid abundance increased with increasing eutrophication. Clionids were found in 41% of the pieces collected from the most eutrophic site vs. 24% from the least eutrophic. Because clionids are the principal bio-eroders of coral reefs, any increase in their abundance will likely result in greater bioerosion rates. The mean abundance of Type 3 corals (in which fragmentation is the primary mode of propagation) is positively related to the frequency of boring sponge invasion. suggesting that increased bioerosion may be partly responsible for community shifts toward Type 3 corals in polluted waters. Cliona cf. vastifica , found for the first time in Barbados, flourishes on the most eutrophic reef and may become an important bioeroder under the highly eutrophic conditions that have begun to plague West Indian reefs. Two new species of Cliona (Porifera: Hadro-merida: Clionidae) are described.     

The role of modern and fossil cyanobacterial borings in bioerosion and bathymetry

Measurements on modern coral reefs at Lee Stocking Island (Bahamas) illustrate that boring cyanobacteria species make a major contribution to microboring bioerosion rates. Borings attributed to cyanobacteria also occur in fossil environments. Bioerosional studies on Permian and Triassic reefs show similar intensities to those observed on modern equivalents. The importance of borings assigned to cyanobacterial activity is even more apparent in paleobathymetry. Comparison of the bathymétrie ranges known from modern and fossil microborings demonstrates a preference of boring cyanobacteria for shallow marine environments. Furthermore, some traces are linked to distinct portions of the shallow euphotic zone. They significantly contribute to characterize typical microboring assemblages, which are used for paleodepth reconstructions. In contrast to these stenobathic species, one cyanobacterial species turned out to be eurybathic. It has been recorded as deep as the dysphotic zone but may even extend to the aphotic zone.     

Effects of grazing and damselfish territoriality on internal bioerosion of dead corals : indirect effects

An experiment was performed on Britomart Reef, Great Barrier Reef (central region), to determine the relationship between fish grazing, damselfish territoriality, and internal bioerosion of dead coral substratum. The damselfish Hemiglyphidodon plagiometopon Bleeker (Pomacentridae) was used for the study because it actively excludes herbivorous fish, particularly scarids and acanthurids, from its territories, creating undergrazed patches in the environment. Its territories simulated conditions of naturally reduced grazing. Freshly killed pieces of the plating coral Pachyseris speciosa Dana were placed under four experimental conditions: (1) within cages, excluding grazing fish; (2) within damselfish territories; (3) beneath shade tops to control for light; and (4) outside damselfish territories, fully exposed to grazers. Internal bioeroders were identified by pattern of substratum excavation and characterization of borings, and were quantified by digitizing x-ray radiographs of the substratum. Three major categories of borers were identified: Cliothosa hancocki Topsent, “other sponges” (of the Cliona viridis Schmidt species complex), and “worms” (including polychaetes and sipunculids). Variations in grazing pressure were found to significantly alter the taxonomic composition of the bioeroder community. Bioerosion by C. hancocki, a boring sponge with large exposed papillae, was found to increase significantly when grazing was reduced within damselfish territories. By contrast, other boring sponges of the C. viridis complex decreased in abundance; they were not affected by higher sedimentation in cages. The response of bioerosion by “worms” was less clear but increased slightly within damselfish territories. This was due primarily to a shift in taxonomic composition and dominance from polychaetes to sipunculids (particularly Cleosiphon aspergillum Quatrefages). The effects of grazing on the internal bioeroder community were often altered or obscured in the caged treatments; this was most likely due to caging artifacts such as increased sedimentation and decreased light. In general, bioerosion rates of the substratum P. speciosa were low in comparison to rates established or estimated for corals with less dense skeletons. Total internal bioerosion rates did not vary significantly with changes in grazing pressure. This study implies that, overall, reduced grazing pressure will lead to production of fine sediments derived from internal bioeroders. Under high grazing pressures, the addition of external bioerosion effected directly by grazers would also produce coarse sediment, resulting in an increase in total bioerosion rates (internal and external) and an increased contribution of both coarse and fine sediments to the reef environment.     

Large-scale spatial variability in bioerosion of experimental coral substrates on the Great Barrier Reef (Australia): importance of microborers

To study the effects of terrigenous inputs on rates of bioerosion by grazers and micro- and macroborers, two inshore sites, one mid-shelf site, two outer barrier sites, and one site located in the Coral Sea were selected along a cross-shelf transect (200 km from onshore to offshore) on the northern Great Barrier Reef. At each site, two grids were firmly attached to the substrate on which replicate experimental blocks, cut from live colonies of Porites sp., were laid in order to study the intra-site bioerosion variability as well as the variation between sites. After 1 year of exposure, rates of bioerosion were estimated using petrographic sections, scanning electron microscopy, and image analysis. Sites differed significantly. The lowest rates were found at the inshore sites (0.46ǂ.14 kg CaCO₃ m^-2 year^-1), while the highest rates were found at the outer barrier sites (3.6ǂ.52). The pattern of bioerosion exhibited at these six sites varied also, mainly due to different levels of bioerosion by grazers and microborers. Rates of macrobioerosion were low. We suggest that this was due to varying levels of terrigenous inputs, hydrographic patterns, and interactions between agents of bioerosion.     

Boring sponges,an increasing threat for coral reefs affected by bleaching events

Coral bleaching is a stress response of corals induced by a variety of factors, but these events have become more frequent and intense in response to recent climate‐change‐related temperature anomalies. We tested the hypothesis that coral reefs affected by bleaching events are currently heavily infested by boring sponges, which are playing a significant role in the destruction of their physical structure. Seventeen reefs that cover the entire distributional range of corals along the Mexican Pacific coast were studied between 2005/2006, and later between 2009/2010. Most of these coral reefs were previously impacted by bleaching events, which resulted in coral mortalities. Sponge abundance and species richness was used as an indicator of bioerosion, and coral cover was used to describe the present condition of coral reefs. Coral reefs are currently highly invaded (46% of the samples examined) by a very high diversity of boring sponges (20 species); being the coral reef framework the substrate most invaded (56%) followed by the rubbles (45%), and the living colonies (36%). The results also indicated that boring sponges are promoting the dislodgment of live colonies and large fragments from the framework. In summary, the eastern coral reefs affected by bleaching phenomena, mainly provoked by El Niño, present a high diversity and abundance of boring sponges, which are weakening the union of the colony with the reef framework and promoting their dislodgment. These phenomena will probably become even more intense and severe, as temperatures are projected to continue to rise under the scenarios for future climate change, which could place many eastern coral reefs beyond their survival threshold.     

Changing dynamics of Caribbean reef carbonate budgets: emergence of reef bioeroders as critical controls on present and future reef growth potential

Coral cover has declined rapidly on Caribbean reefs since the early 1980s, reducing carbonate production and reef growth. Using a cross-regional dataset, we show that widespread reductions in bioerosion rates—a key carbonate cycling process—have accompanied carbonate production declines. Bioerosion by parrotfish, urchins, endolithic sponges and microendoliths collectively averages 2 G (where G = kg CaCO₃ m⁻² yr⁻¹) (range 0.96–3.67 G). This rate is at least 75% lower than that reported from Caribbean reefs prior to their shift towards their present degraded state. Despite chronic overfishing, parrotfish are the dominant bioeroders, but erosion rates are reduced from averages of approximately 4 to 1.6 G. Urchin erosion rates have declined further and are functionally irrelevant to bioerosion on most reefs. These changes demonstrate a fundamental shift in Caribbean reef carbonate budget dynamics. To-date, reduced bioerosion rates have partially offset carbonate production declines, limiting the extent to which more widespread transitions to negative budget states have occurred. However, given the poor prognosis for coral recovery in the Caribbean and reported shifts to coral community states dominated by slower calcifying taxa, a continued transition from production to bioerosion-controlled budget states, which will increasingly threaten reef growth, is predicted.     

Sea urchin bioerosion on coral reefs: place in the carbonate budget and relevant variables

Two aspects of erosion by sea urchins (Echinoidea) in coral reef habitats are: the direct passage of reef framework material through the gut and the indirect effects through the weakening of the reef structure. Urchin bioerosion can equal or exceed reef carbonate production. The impact of urchins on reefs depends on three variables: species type, test size and population density. Large differences in bioerosion by urchins of the same test size occur between different species. Size differences between species in a sea urchin community, as well as size differences within a species along a reef, can be significant. Bioerosion per urchin increases enormously with size. Changes in population density, through time and space, result in significant changes in bioerosion. It is demonstrated how the interaction of these variables determines in-situ sea urchin bioerosion.     

Human activity selectively impacts the ecosystem roles of parrotfishes on coral reefs

Around the globe, coral reefs and other marine ecosystems are increasingly overfished. Conventionally, studies of fishing impacts have focused on the population size and dynamics of targeted stocks rather than the broader ecosystem-wide effects of harvesting. Using parrotfishes as an example, we show how coral reef fish populations respond to escalating fishing pressure across the Indian and Pacific Oceans. Based on these fish abundance data, we infer the potential impact on four key functional roles performed by parrotfishes. Rates of bioerosion and coral predation are highly sensitive to human activity, whereas grazing and sediment removal are resilient to fishing. Our results offer new insights into the vulnerability and resilience of coral reefs to the ever-growing human footprint. The depletion of fishes causes differential decline of key ecosystem functions, radically changing the dynamics of coral reefs and setting the stage for future ecological surprises.     

Bioerosion in Late Permian Rugosa from reefal blocks (Hawasina Complex,Oman Mountains): implications for reef degradation

Summary Rugose corals are known from allochthonous Late Permian reefal blocks of the A1 Jil and Ba’id Formation (Hawasina Complex), Oman Mountains. In contrast to many Late Permian Rugosa found elsewhere in the Tethys, they occurred in sponge reefs and contributed to reef construction. The waagenophyllid warm water coral fauna is moderately diverse comprising cerioid, thamnasterioid, and fasciculate taxa. In contrast to sponges, chaetetids, and low-growing reefbuilders, the corals secreted diagenetically stable, most probably Mg-calcitic skeletons. Borings in coral skeletons are consequently well preserved providing important data for the interpretation of reef destructive processes. Thin-section analysis revealed three taxa of infaunal borers includingEntobia Bronn 1837, uncertain thallophyte borings, and borings of unknown bioeroders. Macroborers were more important than microborers, because of the dominance of clionid sponges. Good evidence exists also for the occurrence of two types of undetermined grazers which destroyed the coral surfaces. The amount and distribution of bioerosions is variable among different coral taxa. The fasciculate coralPraewentzelella regulare Flügel 1995 was the favorate substrate. Up to 33% of the calices were bored. Dendroid and compound corals were bored subordinately. Bioerosion of these colonies does not exceed 2%. There is good evidence for substrate preference amongst the borers. Major controlling factors affecting borer distribution are believed to be variations of skeletal density and gross morphology. The borer assemblage could not limit reef accretion significantly. Factors controlling boring activity might have been quality of substrate, sedimentation rate, rapid incrustation of substrates, and competition for food with reef constructors including sponges, chaetetids, and rugose corals.     

Bioeroders in fossil reefs

Summary Boring algae, fungi and bacteria have been the most constant factor in bioerosion through earth history. Their record reaches back into the middle Precambrian. The only fossil reefs specifically researched for these microendoliths are of Triassic and Upper Jurassic age. Boring worms appear in reefs in the Lower Cambrian. Boring sponges and bivalves first appear also in the lower Paleozoic, but do not become abundant in reefs until the Triassic. Effective substrate excavating grazers are relatively young geologically: Patellids and substrate excavating Echinoids evolved in the Triassic but did not become important bioeroders until the Jurassic or Cretaceous. Scarid fishes are even younger, the oldest representatives having been found in the Miocene. Thus, it seems that the intensity of bioerosion changed significantly during earth history. This may have had consequences for diversity of reef organisms, quality and quantity of reef debris, for diagenesis and record of reef rock.     

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.     

Taphonomy of coral reefs: a review

This review summarises the major factors that affect the post-mortem history of skeletons in a coral reef environment. Skeletal material is traced from life, through death, breakdown, transport, burial and diagenesis to its final fossil form. The fact that most reef sediments are of skeletal composition poses problems of concentration or dilution of individual grain types in taphonomic analysis of reefs. Rates of supply of grains vary, not only with organism abundance and skeletal growth rates, but also with rates of physical and biological breakdown to transportable sediment. Physical and organic processes affect sedimentary structures and textures by mixing or segregating skeletal grains, though biogenic processes normally dominate in the protected setting of reef lagoons. The soft and hard substrates associated with reefs present different media for calcium carbonate accumulation and post-depositional disturbance, for example, loose sediments suffer bioturbation and rocks surfaces suffer bioerosion. The wide range of durability of skeletons and their susceptibility to diagenesis contribute further to the complexities of the preservation of coral reefs.