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.     

Upper Permian coral reef and colonial rugose corals in northwest Hunan, South China

Summary The roles of Permian colonial corals in forming organic reefs have not been adequately assessed, although they are common fossils in the Permian strata. It is now known that colonial corals were important contributors to reef framework during the middle and late Permian such as those in South China, northeast Japan, Oman and Thailand. A coral reef occurs in Kanjia-ping, Cili County, Hunan, South China. It is formed by erect and unscathed colonies ofWaagenophyllum growing on top of one anotherin situ to form a baffle and framework. Paleontological data of the Cili coral reef indicates a middle to late Changhsing age (Late Permian), corresponding to thePalaeofusulina zone. The coral reef exposure extends along the inner platform margin striking in E-S direction for nearly 4 km laterally and generally 35 to 57 m thick. The Cili coral reef exhibits a lateral differentiation into three main reef facies; reef core facies, fore-reef facies, and marginal slope facies. The major reef-core facies is well exposed in Shenxian-wan and Guanyin-an sections where it rests on the marginal slope facies. Colonial corals are dispersed and preserved in non-living position easward. Sponges become major stabilizing organisms in the eastern part of Changhsing limestone outcrop in Kanjia-ping, but no read sponge reefs were formed. Coral reefs at Cili County in Human are different distinctly from calcisponge reefs in South China in their palaeogeography, lithofacies development, organic constitutuents, palaeoecology and diagenesis. The Cili coral reef also shows differences in age, depositional facies association, reef organisms and diagenesis from coral reefs in South Kitakami of Japan, Khorat Plateau of Thailand, and Saih Hatat of Oman. Although some sponge reefs and mounds can reach up to the unconformable Permian/Triassic boundary, coral reef at Kanjia-ping, Cili County, is the latest Permian reef known. This reef appears to had been formed in a palaeoenvironment that is different from that of the sponge reefs and provides an example of new and unique Permian reef type in South China, and could help us to: 1) understand the significance of colonial corals in Permian carbonate buildups; 2) evaluate the importance of coral community evolution prior to the collapse of reef ecosystems at the Permian/Triassic boundary; 3) better understand the effects of the biotic extinction events in Palaeotethys realm; 4) look for environmental factors that may have controlled reefs through time and space, and 5) provide valuable data for the study of Permian palaeoclimate and global evolutionary changes of Permian reefs and reef community.     

珊瑚礁区碳循环研究进展

珊瑚礁是海洋中生产力水平最高的生态系统之一,其碳循环受到有机碳代谢(光合作用/呼吸作用)和无机碳代谢(钙化/溶解)两大代谢过程的共同作用,过程十分复杂.珊瑚礁植物的光合作用保证了有机碳的有效补充,动物摄食及微生物降解等生物过程驱动了珊瑚礁区有机碳高效循环,只有不超过7％的有机碳进入沉积物,而向大洋区水平输出的有机碳通量变化幅度较大,主要受到水动力条件的影响.珊瑚礁区碳酸盐沉积(无机碳代谢)是全球碳酸盐库的重要组成部分,年累积量达到全球CaCO3年累积量的23%～26%,是影响大气CO2浓度的重要组成;珊瑚礁是大气CO2源或汇则取决于净有机生产力与净无机生产力的比值(ROI),当ROI <0.6时,珊瑚礁区是大气CO2的源,反之,则是大气CO2的汇.     

Microbial carbonates and corals on the marginal French Jura platform (Late Oxfordian,Molinges section)

Molinges was located on an Upper Jurassic ramp system of low-energy regime that developed at the southern margin of the French Jura platform. The sedimentary succession is characterized by the transition from a mixed siliciclastic-carbonate to a carbonate depositional setting that occurred during a long-term shallowing-upward trend. The disappearance of siliciclastics is explained by a climatic change, from humid and cold to drier and warmer conditions, previously identified in Late Oxfordian adjacent basins. The base of the section shows marl-limestone alternations of outer ramp. In its middle part, the section displays oncolitic marls, coral-microbialite beds and oncolitic limestones that deposited in a mid ramp position. Finally, the upper section part is made of oolitic limestones of inner ramp. In outer- to mid-ramp settings submitted to terrigenous inputs, the stacking pattern of deposits and facies evolution allow the identification of elementary, small-, medium-, and large-scale sequences. Small amplitudes of sea-level variations probably controlled rapid shifts of facies belts and reef window occurrences. In small-scale sequences, the coral beds developed during periods of sea-level rise. The decreasing rate of sea-level rise is marked by the downramp shift of the oncolitic limestone belt that led to the demise of coral-microbialite beds. These bioconstructions are mainly represented by thin biostromes in which corals never reach great sizes. The coral assemblages mainly include the genera Enallhelia, Dimorpharaea, Thamnasteria, and some solitary forms (Montlivaltia and Epistreptophyllum). They suggest relatively low-mesotrophic conditions in marine waters during the edification of the primary framework. Relatively cold water temperatures and periods of more elevated nutrient contents are probably responsible of the reduced coral development and the formation of a large amount of microbialites.     

Growth models of Bajocian coral-microbialite reefs of Chargey-lès-Port (eastern France): palaeoenvironmental interpretations

Very large amount of microbialites, up to 70% of the reef volume takes part in the edification of Lower Bajocian coral reefs in the Chargey-lès-Port quarry (Haute-Saône, France). Such high amounts of microbialites were unknown within bioconstructions of Middle Jurassic age. Along the 16 m-thick section, seven successive biohermal or biostromal units developed on a shallow platform. Bioconstructions display a first coral growth phase with either constratal or superstratal growth fabrics. Coral fauna is relatively poorly diversified and is dominated by massive forms (Isastrea, Thamnasteria, and Periseris) or branched phaceloid (Cladophyllia) and ramose (Dendraraea) colonies. Corals can be heavily encrusted by microbialites of diverse forms and fabrics (leiolitic, thrombolitic, and stromatolitic). According to the coral growth fabrics, microbialite crusts developed on top of or at the underside of coral colonies, forming a coral-microbialite elementary unit. Microbialites show a multiphase development: (i) directly at the coral surface, a first and mm-scale microbialite layer locally developed; (ii) a second, cm-scale microbialite layer (up to 8 cm thick) covered the entire coral reef framework and assumed the main building role; and (iii) a third, mm- to cm-scale, laminated microbialite layer may also be observed onlapping previous reef structures, before having been progressively buried under sediments. Contemporaneously to the coral growth phase, the first microbialite layer developed on dead portions of coral colonies. The transition between coral growth and microbialite development (i.e., second layer of microbialites) is interpreted as a result of a coral reef crisis, probably reflecting more nutrient-rich conditions. The passage to a stromatolitic (third) layer suggests a control of the accumulation rate. Composition and architecture of coral-microbialite reef units of Chargey-lès-Port highlight the relations between high-frequency fluctuating environmental factors (mainly accumulation rate and trophic conditions) and reef development.     

Two decades of carbonate budget change on shifted coral reef assemblages: are these reefs being locked into low net budget states?

The ecology of coral reefs is rapidly shifting from historical baselines. One key-question is whether under these new, less favourable ecological conditions, coral reefs will be able to sustain key geo-ecological processes such as the capacity to accumulate carbonate structure. Here, we use data from 34 Caribbean reef sites to examine how the carbonate production, net erosion and net carbonate budgets, as well as the organisms underlying these processes, have changed over the past 15 years in the absence of further severe acute disturbances. We find that despite fundamental benthic ecological changes, these ecologically shifted coral assemblages have exhibited a modest but significant increase in their net carbonate budgets over the past 15 years. However, contrary to expectations this trend was driven by a decrease in erosion pressure, largely resulting from changes in the abundance and size-frequency distribution of parrotfishes, and not by an increase in rates of coral carbonate production. Although in the short term, the carbonate budgets seem to have benefitted marginally from reduced parrotfish erosion, the absence of these key substrate grazers, particularly of larger individuals, is unlikely to be conducive to reef recovery and will thus probably lock these reefs into low budget states.     

Caribbean reef coral diversity during the early to middle Miocene: an example from the Anguilla Formation

Reefal units in the early to middle Miocene of Anguilla consist of small, irregular lenses of variable coral composition which developed on a shallow, isolated offshore carbonate platform. They are composed of three distinct coral biofacies (branched, mound-shaped, and platy), which are haphazardly distributed in association with inter-reef sands. These units most probably formed as patch reefs across a broad, shallow area that was exposed to moderate energy conditions and periodically affected by storms. No evidence supports the existence of a more extensive barrier reef system. Comparisons with Oligocene and Mio-Pliocene reefs suggest that during the early to middle Miocene, Caribbean reefs were generally smaller in size (<100 m³) and lower in diversity (21 species in Anguilla, 42 species in total across the Caribbean) than Caribbean reefs during the late Oligocene or during the ate Miocene to early Pliocene (71 species in the Dominican Republic, 80 species total across the Caribbean). The early to middle Miocene Caribbean reef coral fauna was dominated by nine widespread species that occur in deposits of similar age in both Anguilla and Panama. More than half of the fauna consisted of Oligocene relicts. Of the 21 genera that first appeared in the Caribbean during Miocene time, 14 had first occurrences after the middle Miocene, as barrier reef systems became more prevalent across the central Caribbean.     

Microbialites and micro-encrusters in shallow coral bioherms (Middle to Late Oxfordian, Swiss Jura mountains)

Summary Benthic microbial crusts (microbialites or microbolites) are an important component of Middle to Upper Oxfordian shallow-water coral bioherms in the Swiss Jura. They display stromatolitic (laminated), thrombolitic (clotted), and leiolitic (structureless) fabrics, which are distributed heterogeneously throughout the studied sections. The bioherms can be subdivided into coral-microbialite facies, microbialite-dominated facies, and sediment matrix. Macroscopic and microscopic study reveals that microbialitic encrustations commonly occur in two layers. The first one is directly in contact with the substrate and composed of leiolite (locally stromatolite) and a well-diversified micro-encruster fauna; the second one fills the remaining porosity partly or completely with thrombolite and low-diversity micro-encrusters. The growth of the first layer accompanies the growth of the coral reef and thus formed under the same environmental conditions. The second layer is the result of a moving encrustation front filling the remaining porosity (micro- and macrocavities) inside the reef, below the living surface. Both layers play an important role in early cementation. Phototrophic cyanobacteria probably intervene in the formation of the first encrustation zone, whereas heterotrophic bacteria associated to acidic, Ca²⁺-binding macromolecules in biofilms are thought to contribute to the thrombolite inside the reef body. When coral growth cannot take pace with microbialite development, the thrombolite from reaches the surface of the construction and finally covers the reef. The result is a thick interval of thrombolite, which can be interpreted as being related to an ecological crisis in coral-reef evolution. A semi-quantitative analysis of the relative abundance of microbialite types and associated micro-encrusters permits to better constrain the processes leading to a reef crisis. Four micro-encruster associations can be distinguished, and each follows an evolutionary trend in the studied section:Terebella-Tubiphytes dominated,Serpula-Berenicea dominated,Litho-codium dominated, andBacinella dominated. These trends are interpreted to reflect changes in environmental conditions. Bioerosion generally is at its maximum before and after abundant growth of microbialite. According to microbialite-bioerosion relationships and shifts in micro-encruster associations, we propose that the evolution towards a coral-reef crisis involves four main phases: (1) An oligotrophic to low mesotrophic phase when low water turbidity and good oxygenation allow phototrophic metabolisms. This leads to a maximum of coral diversity and development of light-dependent micro-encrusters. (2) A low-mesotrophic phase when increased turbidity and slack water circulation reduce the photic zone and favor heterotrophic micro- and macrofauna. Bioerosion through bivalves increases. (3) A high-mesotrophic phase when environmental conditions are so bad that only microbiatite can be produced. (4) A eutrophic phase when carbonate production is inhibited by high nutrient input and clay flocculation as a result of increased terrestrial run-off. The observed evolutionary trends are not directly linked to changes in bathymetry, but sea-level fluctuations played an important role in opening and closing the depositional environments on the shallow platform. Climatic changes contributed in modulating the influx of siliciclastics and nutrients, and the alkalinity of the water. Demise of coral reefs generally coincides with low sea level and humid climate. Sea-level and climatic fluctuations and, consequently, the crises in reef growth are linked to orbital cycles in the Milandkovitch frequency band.     

Late Triassic reefs from the Northwest and South Tethys: distribution, setting, and biotic composition

The paleolatitudinal distribution patterns during Ladinian and Carnian time are characterized by an increasing expansion of reefs from the northern to the southern hemisphere. The optimum of reef diversity and frequency in the Norian is associated with the development of extended attached or isolated carbonate platforms. Norian-Rhaetian sponge and coral reefs of the Northern Calcareous Alps developed (1) as reef belt composed of patch reefs in platform-edge positions facing the open-marine northwestern Tethys basins and (2) as patch reefs in intraplatform basins as well as in ramp positions.Carnian and Norian-Rhaetian sponge and coral reefs of the Arabian Peninsula are formed (1) as reef complexes at the margins of carbonate platforms on the tops of volcanic seamounts in the southern Tethyan ocean, as small biostromes on these isolated platforms, and (2) as transgressive reef complexes on the attached platform of the Gondwana margin. The Norian Gosaukamm reefal breccia of the NW Tethys is a counterpart of Jabal Wasa reefal limestone of the Gondwana margin with similarities in geological setting and biotic composition. Rhaetian coral biostromes of low diversity known from the Austrian Koessen basin resemble to the time equivalent Ala biostromes of the isolated Kawr platform in the southern Neo-Tethys by forming a discontinuous layer in shallow intraplatform basin setting.     

塔里木板块塔中区上奥陶统良里塔格组的生物礁类型

塔里木板块塔中Ⅰ号坡折带附近上奥陶统良里塔格组取芯井段中可识别多种生物礁灰岩类型,包括珊瑚骨架/障积岩、海绵骨架/绑结岩、苔藓虫绑结岩、钙藻障积岩、钙质菌藻障积/绑结岩等礁灰岩类,藉此可归纳出珊瑚礁、珊瑚-钙藻礁、层孔虫礁、层孔虫-钙藻礁、珊瑚-层孔虫-钙藻礁、苔藓虫礁丘、钙藻礁丘、灰泥丘和微生物礁等生物建造单元。这些礁体的时空分布模式与古环境分异相关联,纵向上具有灰泥丘向珊瑚-层孔虫-钙藻礁至苔藓虫礁丘和钙藻礁的群落结构更替趋势;空间分布则向台地北缘,即I号坡折带延伸显示由低能带灰泥丘向高能带珊瑚-层孔虫-钙藻礁的相变,而且高能带珊瑚-层孔虫-钙藻主体礁和环其周缘相对低能带的钙藻礁丘、灰泥丘等在一定范围内构成造礁群落结构分异。     

Plaeocene reef sediments from the maiella carbonate platform,Italy

Summary Upper Cretaceous and Paleocene reef limestones from the Maiella carbonate platform show how reefs evolved during a time of faunal turn-over. Biostratigraphy and facies analysis of the reef limestones reveal the details of reef growth, composition, and age. Rudists disappeared as reef builders from the Maiella platform shortly before the Cretaceous/Tertiary boundary. Small coral-algal reefs became established in the Danian to Late Thanetian. These scleractinian-red algal dominated boundstones and framestones represent two periods of reef sedimentation and the subsequent interruption of reef growth by emersion and erosion, controlled primarily by fluctuations of relative sea-level. The coral-algal reefs evolved as the taxonomic composition of reef organisms changed. The Paleocene reef sediments are preserved as large slide blocks and as boulders redeposited from the shallow-water platform onto the slope during the course of the Paleocene.     

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.     

Carboniferous reef succession of the Panthalassan open-ocean setting: example from Omi Limestone, central Japan

Summary The Carboniferous-Permian (Visean-Midian) Omi Limestone in the Akiyoshi Terrane, central Japan is a large carbonate unit developed on a seamount in the Panthalassa Ocean. As the seamount subsided during Carboniferous and Permian time, the carbonate deposition at the top of a seamount was almost continous. Terrigenous siliciclastic sediments are absent, because the seamount was situated in an open-ocean setting. The lower part of this seamount-type limestone records a nearly continuous Carboniferous reef succession. Sedimentary facies in the Carboniferous part of the Omi Limestone are generally highly diverse, but their diversity varies in each age. The Upper Carboniferous part consists of highly diversified facies including fore reef, reef front, reef crest, sand shoal, and lagoon facies, while a simple facies assemblage, composed only of fore reef, reef front, and sand shoal facies, occurs in the Lower Carboniferous. The Carboniferous reef succession consists of four phases characterized, in ascending order, by the coralbryozoan-crinoid community, problematic skeletal organism-microencruster community, chaetetid-microencruster community, and calcareous algal community. The first phase, comprising the coral-bryozoan-crinoid community, occurs in theEndothyra spp. Zone to theEostaffella kanmerai Zone (Visean to Serpukhovian). This community acted only as sediment-bafflers and/or contributors. The second phase, represented by the problematic skeletal organism-microencruster community, is developed in theMillerella sp. Zone to theAkiyoshiella ozawai Zone (Bashkirian to lowermost Moscovian), and the third phase, comprising the chaetetid-microencruster community, occurs in the overlyingFusulinella biconica Zone (Lower Moscovian). These two communities are characterized by highly diversified reef-building organisms that had the ability to build rigid frameworks. Calcareous algae and incertae sedis such asHikorocodium, solenoporaceans and phylloid algae characterize the fourth phase, which occurs in theBeedeina sp. Zone (Upper Moscovian). The changes of the reef communities were sucessive for a long period of more than 40 m.y., and each community was distributed in various environments. In addition, the continuous subsidence of the isolated seamount resulted in environmental stability. These properties indicate that this succession represents the biotic evolution of reef-building organisms. The problematic skeletal organism-microencruster community and chaetetid-microencruster community of the Late Carboniferous formed wave-resistant and rigid frameworks along with abundant submarine cements. The growth of these reef frameworks resulted in the formation of highly diversified sedimentary facies comparable to those of a modern reef complex. Such reefs are also recognized in the seamount-type Akiyoshi Limestone, but rare on Carboniferous Pangean shelves. Therefore, the formation of these types of reefs appear to be characteristic of open-ocean seamount settings, which differed from epicontinental shelf settings in having no siliciclastic input, being exposed to relatively strong openocean waves and swells, and probably more environmental stability resulting from the relatively continuous subsidence of the seamount.     

Changes of palaeozonation patterns within Miocene coral reefs, Gebel Abu Shaar, Gulf of Suez, Egypt

The Gebel Abu Shaar represents the southern end of a tilted fault-block consisting of Precambrian basement upon which a mixed siliciclastic-carbonate platform developed during mid-Miocene time. The Miocene sequence contains coral reefs and reef facies. Palaeozonation patterns of the Miocene coral reefs were assessed on the well-preserved outcrops of the Gebel Abu Shaar by recording both qualitative and quantitative data of reef-building assemblages. The various patterns of palaeozonation were analysed and compared at different spatial and temporal scales, including both within-reef and between-reef variations. The palaeozonal changes recorded within these fossil reefs are similar to those described from present-day living coral reefs and involve contraction or extension, fusion or splitting, replacement and omission of reef-builder zones. Depending on the scale concerned, the nature and importance of changes affecting the palaeozonation pattern vary. Spatial variations of the palaeozonal pattern appear mainly controlled by local changes of ecological conditions while temporal variations are related to regional and global environmental changes.     

A modern-type Kimmeridgian reef (Ota Limestone,Portugal): Implications for Jurassic reef models

Upper Jurassic reefs rich in microbial crusts generally appear in deeper (sponge—‘algal’ crust reefs) or in very shallow but protected settings (coral or coral-coralline sponge meadows with ‘algal’ crusts). Upper Jurassic high-energy reefs (coral reefs and coral-stromatoporoid reefs) normally lack major participation of microbial crusts but rather represent huge bioclastic piles with only minor framestone patches preserved. An exception to this rule is represented by the high-energy, coral-‘algal’ Ota Reef from the Kimmeridgian of the Lusitanian Basin (Portugal). The narrow Ota Reef tract rims a small intra-basinal carbonate platform exhibiting perfect facies zonation (from W to E: Reef tract, back reef sands, peritidal belt, low-energy shallow lagoon). The reef is dominated by massive corals (Thamnasteria, Microsolena, Stylina). Complete preservation of coral framework is rare: like other Upper Jurassic high-energy reefs, the Ota Reef is very rich in debris; however, this debris is largely stabilized by algal and microbial crusts, what contrasts the other examples and gives the Ota Reef the appearance of a typical modern high-energy coral-melobesioid algal reef. Further similarities to modern reefs are the likely existence of a spur-and-groove system, the perfect sheltering of inner platform areas and the occurrence of small islands, as indicated by local blackenings and early vadose and karstic features.     

A late Devonian (Frasnian) coral-bafflestone reef at Houshan in Guilin, South China

Summary Givetian to early Carboniferous sediments of South China are characterized by carbonates. Middle and Late Devonian strata are best developed in the Guilin area. Reefs and organic shoals are recorded by various lithofacies types indicating the existence of an extended carbonate platform and a change of the composition of reef communities in time. Starting in the late Devonian, stromatoporoids and corals were replaced by algae that subsequently played an important role together with stromatoporoids, receptaculitids and fasciculate rugose corals in reef communities. In Houshan, 5 km west of Guilin, a coral-bafflestone reef occurs in the Frasnian strata, situated near an offshore algal-stromatoporoid reef. The coral reef was formed in a back-reef area adjacent to the inner platform margin. The coral-bafflestone reef is unique among the late Devonian reefs of South China with regard to the biotic composition. The reef is composed of fasciculate colonies ofSmithiphyllum guilinense n. sp. embedded within in packstones and wackestones. The height of colonies reaches 1 m. The community is low-diverse. The species ofSmithiphyllum occurring in the Frasnian reef complexes of Guilin exhibit a distinct facies control:Smithiphyllum guilinense occurs in or near to margin facies and formed bafflestone, constituting a coral reef whereasSmithiphyllum occidentale Sorauf, 1972 andSmithiphyllum sp.—characterized by small colonies with thin corallites—are restricted to the back-reef and marginal slope facies. The bush-like coral colonies baffled sediments. Algae and stromatoporoids (mainlyStachyodes) are other reef biota. Reef-dwelling organisms are dominated by brachiopods. The reefs are composed from base to top of five lithofacies types: 1) cryptalgal micrite, 2) peloidal packstone, 3) stromatactis limestone, 4) coral-bafflestone, and 5) pseudopeloidal packstone. The reef complex can be subdivided into back-reef subfacies, reef flat and marginal subfacies, and marginal fore-slope subfacies. The Houshan coral-bafflestone reef is not a barrier reef but a coral patch reef located near the inner margin of a carbonate platform.     

Early (Series 2) Cambrian archaeocyathan reefs of southern Labrador as a locus for skeletal carbonate production

Pruss, S.B., Clemente, H. & Laflamme, M. 2012: Early (Series 2) Cambrian archaeocyathan reefs of southern Labrador as a locus for skeletal carbonate production. Lethaia, Vol. 45, pp. 401–410. Archaeocyathan reefs, the first reefs produced by animals, are prominent, global features of early Cambrian successions. However, microbialites – the dominant reef components of the Proterozoic – were still abundant in most archaeocyathan reefs. Although such reefs were a locus for carbonate production, it is unclear how much carbonate was produced skeletally. This analysis of well‐known early Cambrian archaeocyathan patch reefs of the Forteau Formation, southern Labrador, demonstrates that skeletal carbonate was abundantly produced in these archaeocyathan reefs, although only about half was produced by archaeocyathans. Trilobites, echinoderms and brachiopods contributed substantially to the total carbonate budget, particularly in grainstone facies flanking the reefs. Through point count analysis of samples collected from the reef core and flanking grainstones, it can be demonstrated that skeletal material was most abundant in grainstone facies, where animals such as trilobites and echinoderms contributed significantly to carbonate production. In contrast, microbial fabrics were more abundant than skeletal fabrics in the reef core, although archaeocyathan material was more abundant than other skeletal debris. Similar to modern reefs, these reefs created a variety of habitats that allowed for the proliferation of skeletal organisms living on and around the reef, thereby promoting skeletal carbonate production through ecosystem engineering. □Archaeocyatha, bioherms, carbonates, calcification, point count analysis     

Distribution and evolution of Carboniferous reefs in South China

Reefs are sensitive proxies for palaeontological, palaeoenvironmental, and palaeogeographical changes during geological history. In South China, after the collapse of the reef ecosystem during the Frasnian-Famennian and Hangenberg mass extinction events, Carboniferous reefs underwent evolutionary episodes of recovery, decline, and turnover, which were controlled by changes of reef-builders abundance, sedimentary facies, relative sea level, and even global climate. In Tournaisian times, only a few Waulsortian-like banks have been found in Liuzhou, Guangxi without metazoan reefs, which were caused by the lack of reef-builders, such as colonial rugose corals and bryozoans, and the dominant non-carbonate facies (shale, mud stone and sandstone) driven by low relative sea level. The absence of mud mounds in the early Viséan was attributed to the regression event during the Tournaisian-Viséan boundary. During Viséan times, bryozoan-coral reefs in Huishui, Guizhou and Tianlin, Guangxi occurred during a time of increasing biodiversity and carbonate facies resulting from relative sea-level rise. The number of potential reef-builders as colonial rugose coral and bryozoan genera significantly increased in Viséan times in South China. The reef abundance declined during Serpukhovian times in South China and the controlling factors were decreasing abundance of potential reef-builders and developing non-carbonate facies due to a relative sea-level fall. The sedimentary facies were characterized by shale, mud stone, sandstone, and dolostone during this time. A distinct change in reef types occurred after the Mississippian-Pennsylvanian boundary, when phylloid algae and red algae reefs (distributed in Ziyun, Guizhou and Beibuwan, Guangxi) replaced metazoan reefs and became the dominant role in reef ecosystem. This reef turnover event may be triggered by the dramatic relative sea-level fall during the mid-Carboniferous, and continued low relative sea level in South China and global flourish of phylloid and red algae during Pennsylvanian times. Grainstone and dolomitic limestone were the main composition of the platform sedimentary facies in South China during Pennsylvanian times. In addition, global climate cooling and warming, resulted from the waxing and waning of Gondwana glaciation, may also influence the reef evolution in South China, as evidenced from the consistent transgression and regression events and reef evolutionary pattern between South China and globe during the Carboniferous.     

Guadalupian algae-sponge reefs in siliciclastic environments—the reefs at Lengwu (South China) compared with the reef at Iwaizaki (Japan)

Summary Guadalupian reefs occur locally in Guangxi, Guizhou, Yunnan and Western Zhejiang, South China. Two types of Guadalupian reefs can be recognized, one is developed in carbonate platforms, e.g. those in the juncture areas of Guangxi, Yunnan and Guizhou; the other occurs in a littoral clastic shelf. The Lengwu reef in Western Zhejiang is a representative of the latter type, which is a major topic of this paper. Lengwu algae-sponge reef, more than one hundred meters in thickness, are composed mainly of sponges, hydrozoans, algae, bryozoans, microbes and lime mud. Reef limestones sit on the mudstone interbedded with fine sandstone of the proximal prodelta facies and are overlain by coarse clasts of the delta front sediments. Lengwu reef displays a lens-shaped relief, dipping and thinning from the reef core, which is remarkably different from the surrounding sediments, showing a protruding relief. Sponges and microbe/algae form bafflestone, bindstone and framestone of the reef core facies. Fore-reef facies is characterized by lithoclastic rudstone and bioclastic packstone. Reef limestone sequence is composed of three cycles and controlled by sea level changes and sediment influx. Such reef is unique among the Guadalupian reefs in South China, but seems similar in some aspects to Iwaizaki reef limestones of south Kitakami in Japan. Algae and microbes growing around sponges to form rigid structure in Lengwu reef are a typical feature, which is distinctly different to Guadalupian reefs in a stable platform facies of Guizhou, Yunnan and Guangxi, South China.     

Coral adaptive capacity insufficient to halt global transition of coral reefs into net erosion under climate change

Projecting the effects of climate change on net reef calcium carbonate production is critical to understanding the future impacts on ecosystem function, but prior estimates have not included corals' natural adaptive capacity to such change. Here we estimate how the ability of symbionts to evolve tolerance to heat stress, or for coral hosts to shuffle to favourable symbionts, and their combination, may influence responses to the combined impacts of ocean warming and acidification under three representative concentration pathway (RCP) emissions scenarios (RCP2.6, RCP4.5 and RCP8.5). We show that symbiont evolution and shuffling, both individually and when combined, favours persistent positive net reef calcium carbonate production. However, our projections of future net calcium carbonate production (NCCP) under climate change vary both spatially and by RCP. For example, 19%–35% of modelled coral reefs are still projected to have net positive NCCP by 2050 if symbionts can evolve increased thermal tolerance, depending on the RCP. Without symbiont adaptive capacity, the number of coral reefs with positive NCCP drops to 9%–13% by 2050. Accounting for both symbiont evolution and shuffling, we project median positive NCPP of coral reefs will still occur under low greenhouse emissions (RCP2.6) in the Indian Ocean, and even under moderate emissions (RCP4.5) in the Pacific Ocean. However, adaptive capacity will be insufficient to halt the transition of coral reefs globally into erosion by 2050 under severe emissions scenarios (RCP8.5).