Early Ordovician reefs in South China (Chenjiahe section, Hubei Province): deciphering the early evolution of skeletal-dominated reefs

The Lower Ordovician (late Tremadocian–early Floian) Fenhsiang and the overlying Hunghuayuan Formations at the Chenjiahe section in the Three Gorges area of Hubei Province, South China, include four types of reef: microbe-dominated (lithistid sponge–stromatolite and lithistid sponge–calcimicrobial) reefs, and skeletal-dominated (lithistid sponge–bryozoan and bryozoan–pelmatozoan) reefs. The microbe-dominated reefs are characterized by the dominance of microbial sediments that encrusted and bound the surfaces of sponges to reinforce the reef frameworks. In contrast, the skeletal-dominated reefs are distinguished by bryozoans that encrusted frame-building sponges and pelmatozoans, and that grew downward to fill the open spaces available within the frameworks. A series of these reefs shows a temporal succession in reef type, with a decline in the lithistid sponge–stromatolite reefs and an increase in the lithistid sponges and receptaculitids within the lithistid sponge–calcimicrobial reefs in the Hunghuayuan Formation; the lithistid sponge–bryozoan reefs are common in both the Fenhsiang and Hunghuayuan Formations. These features of the Chenjiahe reefs are in marked contrast to other coeval reefs on the Yangtze Platform and elsewhere. Skeletal-dominated reefs first developed in the Three Gorges and adjacent areas, located on the central part of the platform. Likewise, lithistid sponges and receptaculitids first developed in the Three Gorges area and then expanded their range. In contrast, stromatolites declined over time, but remained abundant on a marginal part of the platform. The spatial–temporal distributions of these reefs on the Yangtze Platform reflect the initiation of the Great Ordovician Biodiversification Event and its consequences, although influenced by local environmental conditions. The Three Gorges area was a center for the development of skeletal-dominated reefs, which were established earlier here than elsewhere in the world. These reef types and their spatial–temporal successions provide invaluable clues to the earliest evolution of skeletal-dominated reefs and their ensuing development during the Middle–Late Ordovician.     

The oldest labechiid stromatoporoids from intraskeletal crypts in lithistid sponge–Calathium reefs

Early Ordovician (early Floian) reefs of South China include lithistid sponge–Calathium reefs with a three‐dimensional skeletal framework. These structures are among the first post‐Cambrian skeletal‐dominated reef structures and provides an opportunity to test how the novel metazoan builders changed the environments and increased topographic complexity within benthic communities. We document the oldest labechiid stromatoporoid (Cystostroma) in a lithistid sponge–Calathium reef of the Hunghuayuan Formation in southeastern Guizhou, South China. These earliest stromatoporoids may have originated in reefs, and we argue that the complex topography created by the hypercalcified sponge Calathium facilitated the emergence of stromatoporoids. Beyond Cystostroma, keratose sponges, Pulchrilamina (hypercalcified sponge) and bryozoans have also inhabited in the micro‐habitats (cavities and hard substrates) provided by Calathium. These findings suggest that ecosystem engineering by Calathium played an important role in the further diversification of reefs during the Ordovician.     

The oldest bryozoan reefs: a unique Early Ordovician skeletal framework construction

Adachi, N., Ezaki, Y. & Liu, J. 2011: The oldest bryozoan reefs: a unique Early Ordovician skeletal framework construction. Lethaia, Vol. 45, pp. 14–23. The oldest bryozoan reefs occur in the Lower Ordovician (late Tremadocian) Fenhsiang Formation of the Three Gorges area, South China. These reefs show a unique type of bryozoan (Nekhorosheviella) framework, and were constructed as follows: the first stage involved colonization by lithistid sponges, which acted as a baffler to trap sediments, providing bryozoans with a stable substrate for attachment. The bryozoans then grew as an encruser on the surfaces of sponges, showing a preferential downwards and lateral growth within the sponge scaffolding to avoid biological and physical disturbance. Finally, these biotic combinations among skeletal organisms formed a rigid, three‐dimensional skeletal framework. This mode of bryozoan growth in association with lithistid sponges is remarkable and unique in its growth direction, and the appearance of such reefs, just prior to the widespread development of skeletal‐dominated reefs as part of the Great Ordovician Biodiversification Event, provides an excellent example of the earliest attempts by skeletal organisms to form frameworks by themselves. This find significantly enhances our understanding of the initial stages of skeletal‐dominated reef evolution and the ensuing development of reefs during the Middle–Late Ordovician. □Bryozoa, Early Ordovician, lithistid sponge, Ordovician radiation, reef.     

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.     

Youngest early carboniferous (late Visean) shallow-water patch reefs in eastern Australia (Rockhampton Group, Queensland): Combining quantitative micro- and macro-scale data

Summary Although skeletal organisms have received most of the emphasis in studies on Phanerozoic roef history, the roles of non-skeletal (non-enzymatic) carbonates (e.g., synsedimentary cements, automicrite, microbialite, etc.) in reef framework construction are becoming increasingly better understood. One problem in understanding the role of non-enzymatic carbonates in reef construction has been the difficulty in recognizing them in reef facies. Whereas skeletal organisms commonly can be recognized and documented in the field, non-enzymatic carbonates may be recognizable only in thin section. This paper describes the application of a new sampling technique that allows the quantitative comparison of skeletal macrofauna and flora with associated non-enzymatic carbonates and other microfaunal/microfloral constituents. The technique involves the point counting of thin sections made from small diameter cores that are systematically recovered from grids and line transects that cover a reasonable area (m²) of reef facies. Small, shallow-water patch reefs are abundant in scattered oolitic intervals in the Lower Carboniferous strata of eastern Australia. The youngest known Carboniferous reefs in eastern Australia occur in uppermost Visean strata (limestone FC5) near the top of the Rockhampton Group, approximately 50 km west-northwest of Rockhampton, Queensland. The largest sampled reef was 15 m thick and 42 m in diameter, with synsedimentary relief above the sea floor of at least 2 m during the primary growth phase. The reef occurs within bioclasticoolitic grainstones representing a shallow shelf setting and consists of eight common framework microfacies: 1) coral boundstone; 2) bryozoan boundstone; 3) mixed crinoid-bryozoan boundstone; 4) tubular problematica boundstone; 5) sponge-automicrite boundstone; 6) encrusted thrombolite boundstone; 7) mixed automicrite boundstone; and 8) thrombolitic wackestone-packstone. Reef growth was initiated by automicrite-producing biofilms, sponges and a tubular problematic organism. Primary relief building was accomplished by automicrite-dominated frameworks and lithistid sponges, crinoids, and corals. Large cerioidAphrophyllum coral colonies had a heterogeneous distribution through the reef. The framework of the main relief-bearing portion of the reef consists on average of 44.4% automicrite and automicrite-bound detritus, excluding automicrite-bound sponge body fossils, and at most 19.6% skeletal organisms in growth position (minimum of 7.2%). If sponge body fossils are included as automicrite framework, because they are preserved only as a result of automicrite formation, the percentage of automicrite and bound sediment is 54.9%. A smaller sampled reef consisting of the same basic facies had 39.5% automicrite and automicrite-bound sediment in its fremework (50.2% including sponges) and, at most, 33.4% skeletal organisms in growth position (minimum of 22.7%). The greater volume of skeletal framework in the small reef reflects a greater proportion of large corals. Of framebuilding skeletal organisms, automicrite-preserved lithistid and other sponges and cerioid rugose corals provided the greatest volume. However, crinoid holdfasts were the most widespread skeletal framework components. The dominant framework facies are sponge-automicrite boundstone, encrusted thrombolite, boundstone, mixed automicrite boundstone, and coral boundstone. The reefs are similar in overall framework construction and ecological succession to slightly older Visean reefs in eastern Australia and to some of the late Visean reefs of northern England. Surprisingly, framework similarities also exist between the reefs and certain thrombolite-lithistid-coral reefs of the European Jurassic.     

Mud mounds: A polygenetic spectrum of fine-grained carbonate buildups

Summary This research report contains nine case studies (part II to X) dealing with Palaeozoic and Mesozoic mud mounds, microbial reefs, and modern zones of active micrite production, and two parts (I and XI) summarizing the major questions and results. The formation of different types ofin situ formed micrites (automicrites) in close association with siliceous sponges is documented in Devonian, Carboniferous, Triassic, Jurassic and Cretaceous mounds and suggests a common origin with a modern facies found within reef caves. Processes involved in the formation of autochthonous micrites comprise: (i) calcifying mucus enriched in Asp and Glu, this type presumably is linked to the formation of stromatolites, thrombolites and massive fabrics; (ii) protein-rich substances within confined spaces (e.g. microcavities) result in peloidal pockets, peloidal coatings and peloidal stromatolites, and (iii) decay of sponge soft tissues, presumably enriched with symbiotic bacteria, lead to the micropeloidal preservation of parts of former sponge bodies. As a consequence, there is strong evidence that the primary production of micrite in place represents the initial cause for buildup development. The mode of precipitation corresponds to biologically-induced, matrix-mediated mineralization which results in high-Mg-calcites, isotopically balanced with inorganic cements or equilibrium skeletal carbonates, respectively. If distinct automicritic fabrics are absent, the source or origin of micrite remains questionable. However, the co-occurring identifiable components are inadequate, by quantity and physiology, to explain the enhanced accumulation of fine-grained calcium carbonate. The stromatolite reefs from the Permian Zechstein Basin are regarded as reminiscent of ancestral (Precambrian) reef facies, considered the precursor of automicrite/sponge buildups. Automicrite/sponge buildups represent the basic Phanerozoic reef type. Analogous facies are still present within modern cryptic reef habitats, where the biocalcifying carbonate factory is restricted in space.     

The dorid nudibranchs Peltodoris lentiginosa and Archidoris odhneri as predators of glass sponges

The dorid nudibranchs Peltodoris lentiginosa and Archidoris odhneri were found on glass sponges (Porifera, Hexactinellida) during remotely operated vehicle surveys of three reefs in the Strait of Georgia, British Columbia, Canada. Eight nudibranchs were sampled from 2009 to 2011. Identification of sponge spicules found in their gut and fecal contents confirmed the nudibranchs to be predators of the reef‐forming hexactinellids Aphrocallistes vastus and Heterochone calyx, as well as of the demosponge Desmacella austini, which encrusts skeletons of the glass sponges. Four of five nudibranchs dissected for gut content analysis had stomachs containing sponge spicules. Counts from high‐definition video footage taken during systematic surveys done in 2009 showed that nudibranchs were found in only two of the three glass sponge reefs. These data provide the first quantitative evidence of a molluscan predator on glass sponges found outside of Antarctica, and establish the first trophic link between glass sponges and their associated community of animals in a sponge reef ecosystem on the western Canadian continental shelf.     

Cryptic growth strategies of the Cambrian coral Cambroctoconus: flexible modes of budding and growth in immediate response to available space

The Cambrian coral Cambroctoconus occurs selectively in the crypts of calcimicrobe Epiphyton reefs of the Zhangxia Formation (Miaolingian) in Shandong Province, North China. These cryptobionts preferentially grow laterally and/or downward, in some cases showing pendent growth from the ceilings of the reef framework. The upside-down growth is clearly shown by the downward-oriented aperture and the presence of fork-like holdfasts at its base. The coral aggregations reveal two cryptic growth strategies: one involves the prevalence of far smaller individuals in vertically narrow spaces; the other involves the predominance of modular individuals in large cavernous spaces. Offset individuals frequently appear projecting downward and are connected to each other in a chain-like fashion. Individuals often undergo rejuvenescence laterally and downward, when the growth directions may change. Therefore, the size and modularity of individuals and the direction of budding depend largely upon the cryptic microenvironments available at the time. The cryptobionts make the best use of vacant spaces by modifying the location of budding and altering their growth directions through rejuvenescence. The cryptic growth strategies found in Cambroctoconus indicate a certain degree of morphological variability in the modules. More importantly, they indicate that the individuals flexibly and exquisitely utilized the microbial crypts that were still predominant even during the middle Cambrian. These modes of cryptic growth demonstrate the age-specific exploitation of niches by sessile skeletal organisms that was facilitated by the development of a firm attachment structure.     

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     

Complex nitrogen cycling in the sponge Geodia barretti

Marine sponges constitute major parts of coral reefs and deep‐water communities. They often harbour high amounts of phylogenetically and physiologically diverse microbes, which are so far poorly characterized. Many of these sponges regulate their internal oxygen concentration by modulating their ventilation behaviour providing a suitable habitat for both aerobic and anaerobic microbes. In the present study, both aerobic (nitrification) and anaerobic (denitrification, anammox) microbial processes of the nitrogen cycle were quantified in the sponge Geodia barretti and possible involved microbes were identified by molecular techniques. Nitrification rates of 566 nmol N cm^?3 sponge day^?1 were obtained when monitoring the production of nitrite and nitrate. In support of this finding, ammonia‐oxidizing Archaea (crenarchaeotes) were found by amplification of the amoA gene, and nitrite‐oxidizing bacteria of the genus Nitrospira were detected based on rRNA gene analyses. Incubation experiments with stable isotopes (¹⁵NO₃^‐ and ¹⁵NH₄⁺) revealed denitrification and anaerobic ammonium oxidation (anammox) rates of 92 nmol N cm^?3 sponge day^?1 and 3 nmol N cm^?3 sponge day^?1 respectively. Accordingly, sequences closely related to ‘Candidatus Scalindua sorokinii’ and ‘Candidatus Scalindua brodae’ were detected in 16S rRNA gene libraries. The amplification of the nirS gene revealed the presence of denitrifiers, likely belonging to the Betaproteobacteria. This is the first proof of anammox and denitrification in the same animal host, and the first proof of anammox and denitrification in sponges. The close and complex interactions of aerobic, anaerobic, autotrophic and heterotrophic microbial processes are fuelled by metabolic waste products of the sponge host, and enable efficient utilization and recirculation of nutrients within the sponge–microbe system. Since denitrification and anammox remove inorganic nitrogen from the environment, sponges may function as so far unrecognized nitrogen sinks in the ocean. In certain marine environments with high sponge cover, sponge‐mediated nitrogen mineralization processes might even be more important than sediment processes.     

Clones or clans: the genetic structure of a deep‐sea sponge,Aphrocallistes vastus,in unique sponge reefs of British Columbia,Canada

Understanding patterns of reproduction, dispersal and recruitment in deep‐sea communities is increasingly important with the need to manage resource extraction and conserve species diversity. Glass sponges are usually found in deep water (>1000 m) worldwide but form kilometre‐long reefs on the continental shelf of British Columbia and Alaska that are under threat from trawling and resource exploration. Due to their deep‐water habitat, larvae have not yet been found and the level of genetic connectivity between reefs and nonreef communities is unknown. The genetic structure of Aphrocallistes vastus, the primary reef‐building species in the Strait of Georgia (SoG) British Columbia, was studied using single nucleotide polymorphisms (SNPs). Pairwise comparisons of multilocus genotypes were used to assess whether sexual reproduction is common. Structure was examined 1) between individuals in reefs, 2) between reefs and 3) between sites in and outside the SoG. Sixty‐seven SNPs were genotyped in 91 samples from areas in and around the SoG, including four sponge reefs and nearby nonreef sites. The results show that sponge reefs are formed through sexual reproduction. Within a reef and across the SoG basin, the genetic distance between individuals does not vary with geographic distance (r = ?0.005 to 0.014), but populations within the SoG basin are genetically distinct from populations in Barkley Sound, on the west coast of Vancouver Island. Population structure was seen across all sample sites (global F_ST = 0.248), especially between SoG and non‐SoG locations (average pairwise F_ST = 0.251). Our results suggest that genetic mixing occurs across sponge reefs via larvae that disperse widely.     

Middle Jurassic Porifera from Kachchh, western India

From the Late Bathonian sponge biofacies at Jumara Dome, Kachchh (western India) 11 species of ‘lithistid’, hexactinellinid and calcarean sponges are described. New taxa are the order Sigmatospirida, the genusJumarella, and the speciesJumarella astrorhiza, Mastosia rhytidodes, Radicispongia kraspedophora, andHexactinella prisca. The diverse sponge assemblage is associated with a rich fauna of epibenthic bivalves and brachiopods and formed meadows on fine-grained carbonate substrates. The sponge meadows grew on a carbonate ramp at the lower end of the photic zone, in quiet waters below storm wave base. The rate of sedimentation exceeded that of sponge production. This prevented the development of reef-like bodies. In contrast to Mesozoic sponge reefs, growth of the sponge meadows appears to have been confined to the regressive phases of small transgressive-regressive cycles.     

Retention efficiencies of the coral reef sponges Aplysina lacunosa, Callyspongia vaginalis and Niphates digitalis determined by Coulter counter and plate culture analysis

Sponges are the dominant organisms on many coral reefs and through feeding they may greatly reduce the concentration of suspended food particles. Retention efficiencies of the tubular sponges Aplysina lacunosa, Callyspongia vaginalis and Niphates digitalis were examined on a coral reef located in the Florida Keys. Replicate ambient and exhalant water samples were collected in situ from individuals of each species and analysed using two methods. Retention efficiencies of suspended particles (0.75-18 µm) examined using Coulter counter analysis were similar among the three sponge species, averaging 86%. For all sponges, particle retention decreased as particle size increased from 0.7 to 18 µm. Water samples plated on to Marine Agar produced 54 microbial types. Retention efficiencies of culturable microbes were similar among the three species, averaging 82%. This study suggests that the coral reef sponges Aplysina lacunosa, Callyspongia vaginalis and Niphates digitalis play an important role in the transfer of energy between the pelagic and benthic environments.     

Could some coral reefs become sponge reefs as our climate changes?

Coral reefs across the world have been seriously degraded and have a bleak future in response to predicted global warming and ocean acidification (OA). However, this is not the first time that biocalcifying organisms, including corals, have faced the threat of extinction. The end‐Triassic mass extinction (200 million years ago) was the most severe biotic crisis experienced by modern marine invertebrates, which selected against biocalcifiers; this was followed by the proliferation of another invertebrate group, sponges. The duration of this sponge‐dominated period far surpasses that of alternative stable‐ecosystem or phase‐shift states reported on modern day coral reefs and, as such, a shift to sponge‐dominated reefs warrants serious consideration as one future trajectory of coral reefs. We hypothesise that some coral reefs of today may become sponge reefs in the future, as sponges and corals respond differently to changing ocean chemistry and environmental conditions. To support this hypothesis, we discuss: (i) the presence of sponge reefs in the geological record; (ii) reported shifts from coral‐ to sponge‐dominated systems; and (iii) direct and indirect responses of the sponge holobiont and its constituent parts (host and symbionts) to changes in temperature and pH. Based on this evidence, we propose that sponges may be one group to benefit from projected climate change and ocean acidification scenarios, and that increased sponge abundance represents a possible future trajectory for some coral reefs, which would have important implications for overall reef functioning.     

An evolutionary fast‐track to biocalcification

The ability to construct mineralized shells, spicules, spines and skeletons is thought to be a key factor that fuelled the expansion of multicellular animal life during the early Cambrian. The genes and molecular mechanisms that control the process of biomineralization in disparate phyla are gradually being revealed, and it is broadly recognized that an insoluble matrix of proteins, carbohydrates and other organic molecules are required for the initiation, regulation and inhibition of crystal growth. Here, we show that Astrosclera willeyana, a living representative of the now largely extinct stromatoporid sponges (a polyphyletic grade of poriferan bauplan), has apparently bypassed the requirement to evolve many of these mineral‐regulating matrix proteins by using the degraded remains of bacteria to seed CaCO₃ crystal growth. Because stromatoporid sponges formed extensive reefs during the Paelozoic and Mesozoic eras (fulfilling the role that stony corals play in modern coral reefs), and fossil evidence suggests that the same process of bacterial skeleton formation occurred in these stromatoporid ancestors, we infer that some ancient reef ecosystems might have been founded on this microbial–metazoan relationship.     

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.