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.     

The role of microbes in reef-building communities of the Cannindah limestone (Mississippian), Monto region, Queensland, Australia

Reefs in the Cannindah Limestone at Old Cannindah Homestead, Monto region, Queensland, are exceptional in Eastern Australian Mississippian (Carboniferous) build-ups because of their largest dimension and differentiated microbial fabrics. Calcimicrobes and microbial carbonates, which represent a marine reefal environment occupied by both corals and sponges, are particularly abundant in the reef framework fabrics compared to other Mississippian build-ups in the world. They contributed significantly to the rigidity of the reefs on a crinoidal bank setting. Metazoans and calcimicrobes coexisted and played different roles in reef construction. Reef-building and cavity-dwelling microbes include Renalcis, Palaeomicrocodium, Girvanella, problematic Aphralysia, Ortonella, Shamovella-like, Rothpletzella-like, Wetheredella-like, and some problematic calcimicrobes, which occur in inter-corallite infillings of fasciculate rugose corals, in thrombolitic textures, in or within deposits between microdigitate stromatolite and laminated microbialites, and in reef cavities. Some reef intervals are entirely formed by Renalcis, Palaeomicrocodium, problematic calcimicrobes, and cement. Girvanella, as an encrusting calcimicrobe, generally bound bioclasts and micrite, or together with cement, formed boundstone. Microbial carbonates, including thrombolites, microencrusters, microdigitate stromatolite, laminated and tabular microbialite, irregular layers of self-encrusting vesicles, and microbial micrite, occur commonly in reef framestone and boundstone. The role of microbes and relevant microbial carbonates in the Cannindah reef limestone highlighted a significant account of microbial facies complexes associated with the Mississippian reefs.     

Patch reef development in the florigemma-Bank Member (Oxfordian) from the Deister Mts (NW Germany): a type example for Late Jurassic coral thrombolite thickets

Small reefal bioconstructions that developed in lagoonal settings are widespread in a few horizons of the Late Jurassic (Oxfordian) succession of the Korallenoolith Formation, exposed southwest of Hannover, Northwest Germany. Especially the florigemma-Bank Member, “sandwiched” between oolite shoal deposits, exposes a high variety of build-ups, ranging from coral thrombolite patch reefs, to biostromes and to coral meadows. The reefs show a distribution with gradual facies variations along an outcrop belt that extends about 30 km from the Wesergebirge in the NW to the Osterwald Mts in the SE.The patch reefs from the Deister Mts locality at the “Speckhals” are developed as coral-chaetetid-solenoporid-microbialite reefs and represent a reef type that was hitherto unknown so far north of its Tethyan counterparts. They are mainly built up by coral thickets that are preserved in situ up to 1.5 m in height and a few metres in diameter. They contain up to 20 coral species of different morphotypes but are chiefly composed of phaceloid Stylosmilia corallina and Goniocora socialis subordinately. The tightly branched Stylosmilia colonies are stabilized by their anastomosing growth. The coral branches are coated with microbial crusts and micro-encrusters reinforcing the coral framework. Encrusters and other biota within the thicket show a typical community replacement sequence: Lithocodium aggregatum, Koskinobullina socialis and Iberopora bodeuri are pioneer organisms, whereas the occurrence of non-rigid sponges represents the terminal growth stage. The latter are preserved in situ and seem to be characteristic so far poorly known constituents of the Late Jurassic cryptobiont reef dweller community. The distance and overall arrangement of branches seems to be the crucial factor for the manifestation of a (cryptic) habitat promoting such community replacement sequences. Widely spaced branches often lack any encrusting and/or other reef dwelling organisms, whereas tightly branched corals, as is St. corallina, stimulate such biota. Hence, such reefs are well suited for research on coelobites and community sequences of encrusting and cavity dwelling organisms.     

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.     

Composition and spatial distribution of microencrusters and microbial crusts in upper Jurassic–lowermost Cretaceous reef limestone (Torinosu Limestone,southwest Japan)

Tethyan microencrusters and microbial crusts, most of them previously unknown in Japanese Mesozoic biotas, are present in the uppermost Jurassic–lowermost Cretaceous Torinosu Limestone distributed in southwestern Japan. They construct reefal facies together with reef-forming metazoans. Bacinella irregularis and Lithocodium aggregatum are quantitatively most important, while subordinate constituents include Thaumatoporella parvovesiculifera, Koskinobullina socialis , Iberopora bodeuri , Girvanella sp. and Tubiphytes morronensis. They are especially common in the shallow-water reefal facies, but appear micritic in outcrops. Microencrusters and microbial crusts can only be recognized in thin sections, and they grow around the reef building metazoans and form bindstone. Each microencruster exhibits some specific spatial distribution associated with its paleoecology. Similarities with the taxonomic composition of the upper Jurassic Tethyan microencruster association imply that the community extended geographically at least to the Tethyan gateway where the Japanese Island Arc was located.     

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.     

Microbial impacts on the genesis of Lower Devonian reefal limestones, eastern Australia

Microbial contributions to reefal limestones are evident in eastern Australian Lower Devonian microbial frame/bindstones, red algal-microbial-stromatoporoid bindstones, and microbial-stromatoporoid bindstones. Varied microbialite textures, such as stromatolites, thrombolites, and leiolites, originated as accumulations and partial aggregations of calcimicrobes, peloids, and micrites, which also derived from microbial activities. In microbial frame/bindstones, calcimicrobes (e.g., Rothpletzella and Wetheredella) and dense micrite layers covered and bound underlying substrates. Stabilized substrates promoted the subsequent construction of layered, domal, and columnar frameworks, which were produced by combined accumulations and intermixed associations of calcimicrobes and micritic microbialites. Microbes flourished in the microbial-stromatoporoid bindstones and red algal-microbial-stromatoporoid bindstones during repeated growth interruptions of the framework-building skeletal organisms. Microbes bored into and eroded the skeletal frameworks to subsequently leave micritic envelopes, on which microbial and skeletal encrustations took place in turn. The importance of microbial colonization on the skeletal frameworks was first as subsidiary encrusters that helped to preserve them from erosion, and second as modifiers of the spaces suitable for succeeding encrusters. Partial aggregations of Renalcis filled in the interstices of the skeletal and microbial frameworks, thereby enhancing their rigidity.The microbial impacts on the genesis of reefal limestones are: (1) origination of components (calcimicrobes, peloids, and micrites); (2) formation of characteristic microbial textures; (3) main and subsidiary reef construction and encrustation; and (4) destruction of these components, textures, and structures, but also the protection of resultant constructions in turn. The Lower Devonian reefal limestones treated herein, surprisingly, preserve excellent records of a variety of microbial impacts. Similar effects may also have been common, although variable in preservation, in other ancient reefal deposits.     

A coral-microbialite patch reef from the late jurassic (florigemma-Bank, Oxfordian) of NW Germany (Süntel mountains)

Summary Anin situ Oxfordian patch reef from the Süntel hills (florigemma-Bank, Korallenoolith, NW-Germany) is described. It is composed of an autochthonous reef core overlain by a ‘parautochthonous’ biostrome. The exposed reefal area amounts to about 20 m in lateral and up to 4 m in vertical direction. Nearly all major marine reefal fossil associations from the Tethyal realm are present. In the reef core two facies can be distinguished: (1)Thamnasteria dendroidea thicket facies and (2) thrombolite facies. The first facies is composed of a thin branched autochthonous coral thicket mainly constructed ofTh. dendroidea colonies with only a minor portion ofStylosmilia. Frequently, theTh. dendroidea branches laterally coalesce bridge-like forming a delicate initial framework which was subsequently reinforced by thick microbial coatings, that make up approximately 80% of the rock volume. This facies is an excellent example for microbialite binding in reefal architecture. Additionally, several generations of micromorphic and partly cryptic encrusting organisms settled on theTh. dendroidea branches and microbialite crusts. They successively overgrow each other and fill the space between the coral branches in the thicket forming a characteristic community replacement sequence. Initial colonization of theThamnasteria dendroidea took place on an oncoidic/bioclastic hardground. During this early phase of reefal development, microbialites also played an important role in stabilizing and binding the reef body. The thrombolite facies (2) occupying nearly the same volume of the reef body as facies type (1) consists of a thrombolitic microbialitic limestone which fills the interstice between the coral colonies. It shows a considerably lower faunal diversity than theTh. dendroidea facies. Numerous cavities are interspersed in the thrombolithe and are almost completely filled with dolomitized allomicrite. In contrast, microbialite and allomicrite adjacent to the reef core rarely reveal any dolomitized areas. Above the reef core, mostly toppledSolenopora jurassica thalli occur together with a few massiveIsastrea colonies forming a parautochthnous biostrome. They are inhabited by a low diverse assemblage of encrusting organisms. Microbialites are only rarely present in this biostromal unit. The patch reef is developed within a lagoonal limemud facies both separated by a sharp interface. In contrast, continuous facies transition exists between theSolenopora biostrome and adjacent deposits which are characterized by micritic to pelmicritic limestone sometimes with lenses of oncoids. Debris derived from the patch reef is only sporadically intercalated in the reef surrounding lagoonal sediments. Gastropods, bivalves, and dasycladalean algae dominate the lagoonal biota. Up-section following theSolenopora biostrome nerinean gastropods become the most abundant species amounting to a ‘Nerinea-bed’. This horizon moderately elevates above the patch reef indicating, that is arose above the surrounding sea floor forming a relief. The patch reef established on a secondary hardground probably released by a minor transgression and a nondepositional regime. It grew up on a well-illuminated sea floor only a few meters below sea level. Only a low background sedimentation rate and modest water circulation are assumed during reefal growth. These features characterize an open marine lagoon. A subsequent shallowing upwards trend caused emergence of the early lithifiedflorigemma-Bank sediments. In the following erosional phase the reef core,Solenopora biostrome and ‘Nerinea-bed’ were sharply cut. Paleokarst phenomena (karst solution of the rocks, selective leaching of the aragonitic corals) truncate the surface of theflorigemma-Bank. Released by a transgressive sea level, the paleokarst surface is densely inhabited by marine boring and encrusting organisms (oysters, serpulids). Karst cavities are filled with an oncoid-bearing bioclastic limestone with a large portion of siliciclastics. Theflorigemma-Bank is overlain by the reddish bioclastic sandstone of the ‘Zwischenfl?zregion’.     

Cambrian Series 3 lithistid sponge–microbial reefs in Shandong Province,North China: reef development after the disappearance of archaeocyaths

The Cambrian Series 3 Zhangxia Formation in Shandong Province, North China, includes small‐scale lithistid sponge–microbial reefs. The lithistid sponges grew on oolitic and bioclastic sediments, which were stabilized by microbial activities. The relative abundances of microbial components (e.g. calcimicrobe Epiphyton and stromatolites) vary among the reefs. However, the microbial components commonly encrusted or bound the lithistid sponges, formed remarkable encrustations on the surfaces of the sponges. Epiphyton especially grew upward and downward. The lithistid sponges thus provided substrates for the attachment and development of microbes, and the microbes played essential roles as consolidators, by encrusting reef‐building sponges. Additionally, the lithistid sponges were prone to degradation via microbial activities and diagenetic processes, and were thus preserved as micritic bodies, showing faint spicular networks or abundant spicules. Such low preservation potential within the reef environment obscured the presence of the sponges and their widespread contribution as reef‐building organisms during the Cambrian. During the prolonged interval after the demise of archaeocyaths, purely microbial reefs, such as stromatolites and thrombolites have been considered to be the principal reef builders, in association with rare lithistid sponge–microbial associations. However, recent findings, including those from Shandong Province and Korea, suggest that the lithistid sponge‐bearing reefs were more extensive during the Epoch 3 to the Furongian than previously thought. These lithistid sponge–microbial reefs were precursors of the sponge–microbial reefs that dominated worldwide in the Early Ordovician.     

Millennial‐scale ocean acidification and late Quaternary decline of cryptic bacterial crusts in tropical reefs

Ocean acidification by atmospheric carbon dioxide has increased almost continuously since the last glacial maximum (LGM), 21 000 years ago. It is expected to impair tropical reef development, but effects on reefs at the present day and in the recent past have proved difficult to evaluate. We present evidence that acidification has already significantly reduced the formation of calcified bacterial crusts in tropical reefs. Unlike major reef builders such as coralline algae and corals that more closely control their calcification, bacterial calcification is very sensitive to ambient changes in carbonate chemistry. Bacterial crusts in reef cavities have declined in thickness over the past 14 000 years with largest reduction occurring 12 000–10 000 years ago. We interpret this as an early effect of deglacial ocean acidification on reef calcification and infer that similar crusts were likely to have been thicker when seawater carbonate saturation was increased during earlier glacial intervals, and thinner during interglacials. These changes in crust thickness could have substantially affected reef development over glacial cycles, as rigid crusts significantly strengthen framework and their reduction would have increased the susceptibility of reefs to biological and physical erosion. Bacterial crust decline reveals previously unrecognized millennial‐scale acidification effects on tropical reefs. This directs attention to the role of crusts in reef formation and the ability of bioinduced calcification to reflect changes in seawater chemistry. It also provides a long‐term context for assessing anticipated anthropogenic effects.     

A late-devonian (Famennian)Renalcis-Epiphyton reef at Zhaijiang, Guilin, South China

Summary Microbial reefs, together with stromatolitic mounds and ooid shoals, constitute massive limestones in Famennian platform marginal strata in Guilin, in sharp contrast to the well-known coral-stromatoporoid reefs in the Givetian and Frasnian. Microbes played a significant and important role as stabilizers in the Famennian carbonate deposits of Guilin. A reef at Zhaijiang was constructed byEpiphyton andRenalcis, and is representative of such carbonate buildups. The reef is situated 10 km west of Guilin and corresponds to a microbe-dominated platform margin carbonate complex. Organisms in the Zhaijiang microbial reef are low diversity and dominated by ostracods and two genera of microbes,Epiphyton andRenalcis. Other microbial genera such asSphaerocodium andWetheredella occur in most of reef facies in Guilin, but their role as reef builder is doubtful because they occur only in minor amounts. The same four genera occur in volumetrically significant amounts in the upper Devonian carbonate complexes of Alberta. Canada and Western Australia. However.Epiphyton is more abundant in the Guilin reefs. The Zhaijiang microbial reef developed above Famennian proximal slope faices, as suggested by reef architecture and paleogeographic setting. The facies sequence of the microbial reef can be divided into three parts. The lower part is composed of medium-bedded bioclastic grainstones with a few microbial framestone lithoclasts, representing a proximal slope facies. The middle part consists of thin-bedded mudstone and shale with limestone lenses that are thought to be low stand deposits. In some cross sections, mudstone and shale infilled tidal channels that developed in the bioclastic grainstones.Renalcis-Epiphyton framestone constitutes the upper part with massive stacking patterns. The reef is 35 m thick and over 50 m in width. Nine litho- and biofacies are recognized. Zhaijiang reef provides an example of a binder guild-dominated buildup in the almost vacant reef ecosystem of the Famennian and represents a characteristic kind of reef after the Frasnian/Famennian extinction.     

Extensive metazoan reefs from the Ediacaran Nama Group,Namibia: the rise of benthic suspension feeding

We describe new, ecologically complex reef types from the Ediacaran Nama Group, Namibia, dated at ~548 million years ago (Ma), where the earliest known skeletal metazoans, Cloudina riemkeae and Namacalathus, formed extensive reefs up to 20 m in height and width. C. riemkeae formed densely aggregating assemblages associated with microbialite and thrombolite, each from 30 to 100 mm high, which successively colonised former generations to create stacked laminar or columnar reef frameworks. C. riemkeae individuals show budding, multiple, radiating attachment sites and cementation between individuals. Isolated Namacalathus either intergrew with C. riemkeae or formed dense, monospecific aggregations succeeding C. riemkeae frameworks, providing a potential example of environmentally mediated ecological succession. Cloudina and Namacalathus also grow cryptically, either as pendent aggregations from laminar crypt ceilings in microbial framework reefs or as clusters associated with thrombolite attached to neptunian dyke walls. These reefs are notable for their size, exceeding that of the succeeding Lower Cambrian archaeocyath–microbial communities. The repeated colonisation shown by C. riemkeae of former assemblages implies philopatric larval aggregation to colonise limited favourable substrates. As such, not only were skeletal metazoans more important contributors to reef building in the Ediacaran, but there were also more variable reef types with more complex ecologies, than previously thought. Such an abundance of inferred suspension feeders with biomineralised skeletons indicates the efficient exploitation of new resources, more active carbon removal with a strengthened energy flow between planktic and benthic realms, and the rise of biological control over benthic carbonate production. These mark the prelude to the Cambrian Explosion and the modernisation of the global carbon cycle.     

Anisian (middle triassic) buildups of the Northern Dolomites (Italy): The recovery of reef communities after the permian/triassic crisis

Summary After the end-Permian crisis and a global ‘reef gap’ in the early Triassic, reefs appeared again during the early Middle Triassic. Records of Anisian reefs are rare in the Tethys as well as in non-Tethyan regions. Most Anisian reefs are known from the western part of the Tethys but there are only very few studies focused on biota, facies types and the paleogeographical situation of these reefs. From the eastern part of the Tethys, Anisian reefs, reefal buildups or potential reef-building organisms have been reported from different regions of southern China. Most of the Anisian reefs known from western and central Europe as well as from southern China seem to be of middle and late Pelsonian age. The study area is situated in the northern Dolomites (South Tyrol, Italy) southeast of Bruneck (Brunico). It comprises the area between Olang (Valdaora) and Prags (Braies). The study is based on detailed investigations of the regional geology, stratigraphy and lithofacies (R. Zühlke, T. Bechst?dt) as well as on a comprehensive inventory of Anisian reef organisms (B. Senowbari-Daryan, E. Flügel). These data are used in the discussion of the controls on the recovery of reefs during the early Middle Triassic. Most late Anisian reef carbonates studied are represented by allochthonous talus reef blocks of cubicmeter size. Small biostromal autochthonous mounds are extremely rare (Piz da Peres). The reef mounds as well as most of the reef blocks occur within the middle to late Pelsonian Recoaro Formation. They were formed on the middle reaches of carbonate ramps in subtidal depths, slightly above the storm wave base with only moderate water energy. Most lithotypes observed in the reef blocks correspond to sponge and/or algal bafflestones. Low-growing sessile organisms (Olangocoelia (sponge, alga?), sphinctozoan sponges, bryozoans, soleno-poracean algae, corals) and encrusting epibionts (sponges, porostromate algae, cyanophycean crusts, foraminifera, worms, microproblematica) created low cm-sized biogenic structures (bioconstructions) which baffled and bound sediment. Organic framework was only of minor importance; it is restricted to theOlangocoelia lithotype. Framework porosity was small in these reef mounds. Submarine carbonate cements, therefore, are only of minor importance s compared with Permian or Ladinian reefs. The relatively high number of lithotypes encountered in the reef blocks indicates a high biofacies diversity. Regarding the relative frequency, the diverse biota consist in descending order ofOlangocoelia, sponges (sphinctozoans, inozoans, siliceous sponges), bryozoans, porostromate algae and worm tubes. The sphinctozoans are characterized by small, mostly incrusting forms. The numerical diversity (species richness) is low compared with late Permian or Ladinian and late Triassic sphinctozoan faunas occurring within reefs. Following the sponges, monospecific bryozoans (Reptonoditrypa cautica Sch?fer & Fois) are the most common organisms in the reef limestones. Porostromate algae were restricted to areas within the bioconstructions not inhabited by sponges. The low-diverse corals had no importance in the construction of an organic framework. Surprisingly, microbial crusts are rare or even lacking in the investigated Anisian bioconstructions. This is in contrast to late Permian and Ladinian as well as Carnian reefs which are characterized by the abundance of specific organic crusts. The same comes true for‘Tubiphytes’ which is a common constituent in Permian, Ladinian and Carnian reef carbonates but is very rare in the Anisian of the Olang Dolomites. Instead of‘Tubiphytes’ different kinds of worm tubes (spirorbid tubes, Mg-calcitic tubes and agglutinated tubes) were of importance as epifaunal elements. Macrobial encrustations consisting of characteristic successions of sponges, bryozoans, algae, worm tubes and microproblematica seem to be of greater quantitative importance than in Ladinian reefs. Destruction of organic skeletons (predominantly of bryozoans) by macroborers (cirripedia?) is a common feature. The Anisian reef organisms are distinctly different from late Permian and from most Ladinian reef-builders. No Permian Lazarus taxa have been found. New taxa: Sphinctozoan sponges—Celyphia? minima n.sp.,Thaumastocoelia dolomitica n. sp.,Deningeria tenuireticulata n. sp.,Deningeria crassireticulata n. sp.,Anisothalamia minima n.g. n.sp., Inozoan sponges-Meandrostia triassica n.sp. Microproblematica-Anisocellula fecunda n.g. n.sp., Porostromate alga-Brandneria dolomitica n.g. n.sp. Most of our data are in agreement with the model described byFois & Gaetani (1984) for the recovery of reef-building communities during the Ansian but the biotic diversity seems to be considerably higher than previously assumed. Anisian deposition and the formation of the reef mounds within the Pelsonian Recoaro Formation of the Dolomites were controlled by the combined effects of synsedimentary tectonics and eustatic changes in sea-level. During several time intervals, especially the early Anisian (northern and western Dolomites: tectonic uplift), the early Pelsonian (eastern Dolomites: drowning) and the late Illyrian (wide parts of the Dolomites: uplift and drowning), the sedimentation was predominantly controlled by regionally different tectonic subsidence rates. The amount of terrigenous clastic input associated with synsedimentary tectonics (tectonic uplift of hinterlands) had a major influence on carbonate deposition and reef development. The re-appearance of reef environments in the Olang Dolomites was controlled by a combination of regional and global factors (paleogeographic situation: development of carbonate ramps; decreasing subsidence of horst blocks; reduced terrigenous input; moderate rise in sea-level).     

Encrusting micro-organisms and microbial structures in Upper Jurassic limestones from the Southern Carpathians (Romania)

Late Jurassic–Early Cretaceous ?tramberk-type reef limestones are known from some parts of the Southern Carpathians in Romania. The Upper Jurassic deposits mainly consist of massif reef limestones including a variety of microbialites associated with micro-encrusters. They played an important role in the formation and evolution of the reef frameworks and thus are of significant importance for deciphering the depositional environments. For our study, the most important encrusting organisms are Crescentiella morronensis, Koskinobullina socialis, Lithocodium aggregatum, Bacinella-type structures, Radiomura cautica, Perturbatacrusta leini, Coscinophragma sp., and crust-forming coralline sponges such as Calcistella. Based on microscopic observations, microbial contribution to reef construction is documented by the abundance of dense micrite, laminate structures, clotted, thrombolithic or peloidal microfabrics, constructive micritic cortices, biogenic encrustations and cement crusts, as well as by other types of microbial structures and crusts. Most of the investigated carbonate deposits can be classified as “coral-microbial-microencruster boundstones” which are characteristic for the Intra-Tethyan domain. Their paleogeographical significance is indicated by the presence of many features comparable with carbonate deposits of rimmed platform systems from the Northern Calcareous Alps or Central Apennines. Based on the distribution of the facies and facies associations within the carbonate sequences under study we can distinguish slope and external shelf margin environments. The microbial crusts, the encrusting micro-organisms, and in some cases the syndepositional cements have stabilized and bound the carbonates of the slope facies types. Subsequently, the stable substrate favored the installation of coral-microbial bioconstruction levels.     

Intraspecific variation in an Ediacaran skeletal metazoan: Namacalathus from the Nama Group,Namibia

Namacalathus hermanastes is one of the oldest known skeletal metazoans, found in carbonate settings of the terminal Ediacaran (~550–541 million years ago [Ma]). The palaeoecology of this widespread, goblet‐shaped, benthic organism is poorly constrained yet critical for understanding the dynamics of the earliest metazoan communities. Analysis of in situ assemblages from the Nama Group, Namibia (~548–541 Ma), shows that Namacalathus exhibited size variation in response to differing water depths, hydrodynamic conditions and substrate types. In low‐energy, inner ramp environments, Namacalathus attains the largest average sizes but grew in transient, loosely aggregating, monospecific aggregations attached to microbial mats. In high‐energy mid‐ramp reefs, Namacalathus spatially segregated into different palaeoecological habitats with distinct size distributions. In outer ramp environments, individuals were small and formed patchy, dense, monospecific aggregations attached to thin microbial mats. Asexual budding is common in all settings. We infer that variations in size distribution in Namacalathus reflect differences in habitat heterogeneity and stability, including the longevity of mechanically stable substrates and oxic conditions. In the Nama Group, long‐lived skeletal metazoan communities developed within topographically heterogeneous mid‐ramp reefs, which provided diverse mechanically stable microbial substrates in persistently oxic waters, while inner and outer ramp communities were often ephemeral, developing during fleeting episodes of either oxia and/or substrate stability. We conclude that Namacalathus, which forms a component of these communities in the Nama Group, was a generalist that adapted to various palaeoecological habitats within a heterogeneous ecosystem landscape where favourable conditions persisted, and was also able to opportunistically colonise transiently hospitable environments. These early skeletal metazoans colonised previously unoccupied substrates in thrombolitic reefs and other microbial carbonate settings, and while they experienced relatively low levels of interspecific competition, they were nonetheless adapted to the diverse environments and highly dynamic redox conditions present in the terminal Ediacaran.     

Facies and palaeoecology of Upper Jurassic (Middle Oxfordian) coral reefs in England

Summary This study documents the facies and fauna of Late Jurassic (Middle Oxfordian) coral reefs in England. Sedimentological and palaeoecological analysis of these reefs distinguishes three generic reef types: (1) small reef patches and thickets associated with siliciclastic deposits; (2) small reef patches and thickets associated with siliciclastic-free bioclastic grainstones and packstones; and (3) biostromal units associated with deep water facies. The depositional environments of these reef types are discussed. Two coral assemblages are identified: (1) the microsolenid assemblage; and (2) theThamnasteria, Isastraea, Fungiastraea andThecosmilia assemblage (Thamnasteria assemblage). TheThamnasteria assemblage developed in all shallow water environments in the study area, regardless of local environmental conditions. The fauna is very eurytopic,r-selected and can tolerate significant environmental fluctuations on short temporal scales (sub-seasonal). The main control on the development of the microsolenid assemblage was low light intensity, low background sedimentation rates and low hydrodynamic energy levels.     

Coral assemblages, biofacies, and ecological zones in the mid-Holocene reef deDosits of the Enriquillo Valley, Dominican Republic

Stemann, T. A. & Johnson, K. G. 1992 07 15: Coral assemblages, biofacies. and ecological zones in the mid-Holocene reef deposits of the Enriquillo Valley, Dominican Republic. A large, subaerially exposed mid-Holocene reef in the Enriquillo Valley (southwest Dominican Republic) provides an excellent opportunity to examine the relationship between reefal ecology and reefal deposits. Coral species richness and diversity in the Enriquillo reef are comparable to that found in the recent of the Caribbean, and ecological zonation comprised of a shallow-water branching coral zone and a deeper water mixed-coral zone is apparent. Similar zonation and diversity patterns have been recognized on living Caribbcan reefs with moderate wave exposure. Three statistically discrete biopdcies can be discriminated in the Enriquillo deposits using quadrat point-counting techniques commonly used to census modern reefs. They include a facies dominated by Acropora cervicornis, a low diversity assemblage with abundant, large colonies of Siderastrea siderea and Stephanocoenia intersepta, and a higher diversity assembbdge composed of various taxa including Montastraea spp., Colpophyllia spp., and Agaricia spp. Each facies can be recognized at scales of 1–3 m², though in some cases they extend for more than 20 m². In general, the A. cervicornis facies is spatially segregated from the other two biofacies. although neither the shallow nor the deep-water ecological zone is comprised of a single reef biofacies. Rather, the biofacies described here appear to represent distinct micro-environments resulting from ecological variation at a subzonal scale. Micro-environments of similar scale are most likely preserved in other reef deposits. Recognition of these subzonal biofacies may have important consequences for the stratigraphical and paleoccological interpretation of fossil reefs. Corals, biofacies, reef zonation, coral communities, fossil reefs.     

The Torres Reefs,North Queensland,Australia —strong tidal flows a modern control on their growth

High resolution seismic data from Torres Strait off northern Australia provide the first insights into the development of the striking platform reefs of the region. In the southern part of Torres Strait, where the sea floor is shallow and featureless, the Torres Reefs have developed as long, narrow platforms parallel to very strong east-west flowing tidal currents. The reefs have expanded since the end of the post-glacial marine transgression (approximately 6000 years ago) through preferential growth at each end of their platforms. In these localized areas sheltered from the strong tidal currents, patch reefs have developed. Sediments swept from the reef margin and the inter-reef channels are deposited in these current lee areas at the ends of the reefs. The combination of the patch reefs and reefal sediments provides the essentials for extension of the platform reefs. The seismic data confirm that modern reef expansion is occurring without there being a high substrate for coral colonization. The evolutionary sequence from an uncolonized seafloor to a mature platform reef envisages a synchronous process of patch reef development and sediment accumulation.     

Bioerosion in the Miocene Reefs of the northwest Red Sea,Egypt

Macroborings provide detailed information on the bioerosion, accretion and palaeoenvironment of both modern and fossil reefs. Dolomitized reefal carbonates in the Um Mahara Formation exhibit an outstanding example of spatially distributed, well‐preserved bioerosion structures in tropical to subtropical syn‐rift Miocene reefs. Ten ichnospecies belonging to five ichnogenera are identified; three belonging to the bivalve‐boring ichnogenus Gastrochaenolites, three attributed to the sponge‐boring ichnogenus Entobia, and four ichnospecies assigned to three worm‐boring ichnogenera Trypanites, Maeandropolydora and Caulostrepsis. The distribution of the reported borings is strongly linked to the palaeo‐reef zones. Two distinctive ichnological boring assemblages are recognized. The Gastrochaenolites‐dominated assemblage reflects shallower‐marine conditions, under water depths of a few metres, mostly in back‐reef to patch‐reef zones of a back‐reef lagoon. The Entobia‐dominated assemblage signifies relatively deeper marine conditions, mostly in reef core of the fringing Miocene reefs. These ichnological assemblages are attributed herein to the Entobia sub‐ichnofacies of the Trypanites ichnofacies. This ichnofacies indicates boring in hard carbonate substrates (such as corals, rhodoliths, carbonate cements and hardgrounds) during periods of non‐sedimentation or reduced sediment input.