The history of Devonian-Carboniferous reef communities: Extinctions,effects, recovery

Summary Analysis of the taxonomic composition, diversity and guild structure of five “typical” reef and mud mound communities ranging in age from Late Devonian-Early Carboniferous indicates that each of these aspects of community organization changed dramatically in relation to three extinction events. These events include a major or mass extinction at the end of the Frasnian; reef communities were also effected by less drastic end-Givetian and mid-late Famennian extinctions of reef-building higher taxa. Peak Paleozoic generic diversities for reef-building stromatoporoids and rugose corals occurred in the Eifelian-Givetian; reef-building calcareous algal taxa were longranging with peak diversity in the Devonian. These three higher taxa dominated all reef-building guilds (Constructor, Binder, Baffler) in the Frasnian and formed fossil reef communities with balanced guild structures. The extinction of nearly all reef-building stromatoporoids and rugose corals at the end of the Frasnian and the survival of nearly all calcareous algac produced mid-late Famennian reef communities dominated by the Binder Guild. Despite the survival of most calcareous algae and tabulate corals, the mid-late Famennian extinction of all remaining Paleozoic stromatoporoids and nearly all shelf-dwelling Rugosa brought the already diminished Devonian reef-building to a halt. These Devonian extinctions differ from mass extinctions by the absence of a statistically significant drop in taxonomic diversity and by their successional and cumulative effects on reef communities. Tournaisian mud mounds contain communities markedly different from the frame-building communities in Late Devonian and Visean reefs. Mound-building biotas consist of an unusual association dominated by erect, weakly skeletonized members of the Baffler Guild (chiefly fenestrate Bryozoa; Pelmatozoa) and laterally expanded, mud-binding algae/stromatolites and reptant Bryozoa. The initial recovery to reefs with skeletal frameworks in the Visean was largely due to the re-appearance of new species of abundant colonial rugose corals (Constructor Guild) and fenestrate Bryozoa. This Frasnian-Visean evolution in the taxonomic composition and structure of the reef-building guilds is also expressed by abrupt changes in biofacies and petrology of the reef limestones they produced. Thus, “typical” Frasnian reef limestones with balanced guild structures are framestones-boundstones-bafflestones, Famennian reefs are predominantly boundstones, Tournaisian mud mounds are bafflestones and Visean reefs are bafflestones-framestones.     

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.     

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).     

First record of coralline demosponges in the Pleistocene: implications for reef ecology

We report the first discovery of coralline sponges from Pleistocene reef limestones of Vanuatu. Sponges of the genus Acanthochaetetes were identified from two reef terraces of Middle and Late Pleistocene age. As these sponges document cryptic habitats in modern coral reefs, they may be index fossils of cryptic habitats in the Pleistocene as well, thereby providing clues on growth conditions in fossil reefs. The small size of the discovered specimens may be attributed to the transient nature of their cryptic habitats, either due to reef growth or the occurrence of an unusual event.     

Coral reef encruster communities and carbonate production in cryptic and exposed coral reef habitats along a gradient of terrestrial disturbance

Encrusting calcareous organisms such as bryozoans, crustose coralline algae (CCA), foraminiferans, and serpulid worms are integral components of tropical framework-building reefs. They can contribute calcium carbonate to the reef framework, stabilise the substrate, and promote larval recruitment of other framework-building species (e.g. coral recruits). The percentage cover of encrusting organisms and their rates of carbonate production (g m⁻² year⁻¹) were assessed at four sites within a coastal embayment, along a gradient of riverine influence (high-low). As the orientation and type of substrate is thought to influence recruitment of encrusting organisms, organisms recruiting to both natural (the underside of platy corals) and experimental substrates were assessed. The effect of substrate exposure under different levels of riverine influence was assessed by orientating experimental substrates to mimic cryptic and exposed reef habitats (downwards-facing vs upwards-facing tiles) at each site. Cryptic experimental tiles supported similar encruster assemblages to those recruiting to the underneath (cryptic side) of platy corals, suggesting that tiles can be used as an experimental substrate to assess encruster recruitment in reef systems. Encruster cover, in particular CCA, and carbonate production was significantly higher at low-impact (clear water), high wave energy sites when compared to highly riverine impacted (turbid water), low wave energy sites. Cryptically orientated substrates supported a greater diversity of encrusting organisms, in particular serpulid worms and bryozoans. The inverse relationships observed between riverine inputs and encrusters (total encruster cover and carbonate production) have implications for both the current and future rates and styles of reefal framework production.     

Cryptic serpulid-microbialite bioconstructions in the Kakoskali submarine cave (Cyprus,Eastern Mediterranean)

The biostalactites from the Kakoskali cave in Cyprus represent a new example of the complex biotic relationships between skeletal organisms and microbial communities in building bioconstructions of cryptic marine environments. Biostalactites are mainly constituted of polychaetes of the family Serpulidae and, to a lesser degree, foraminifers and bryozoans. Within the skeletal framework of these organisms, two types of microcrystalline calcite (micrite) have been recognized: autochthonous and detrital micrite. The autochthonous fraction is syndepositionally lithified and occurs as clotted peloidal and, subordinately, aphanitic (structureless) textures, suggesting the presence of heterotrophic microbial activities thriving on decaying metazoan organic matter. This fraction is limited to the protected portions of the bioconstructions, especially in the inner and lower parts. The presence of iron and manganesiferous oxidizing bacteria is suggested by the deposition of ferromanganesiferous crusts and Frutexites-like structures. These microbial-induced biomineralizations are the main evidence of carbonatogenetic and Fe–Mn, autotrophic and chemoheterotrophic, bacterial activities. The Kakoskali cave is frequently visited by divers who, during their immersions, resuspend the fine bottom sediment, which later covers the surface of the bioconstructions, disturbing the delicate equilibrium of the biotic association. This perturbation, which is also caused by strong waves and currents, during winter months, reflects on the bioconstruction morphologies, community composition, and colonization pattern. Bioconstructions exhibit an upper smooth surface, produced by few taxa (e.g., polychaetes, foraminifers), hosting a low number of living individuals, and a lower comparably rough surface, colonized by a more abundant community showing a higher species richness. The ratio surface roughness/smoothness is related to micrite sediment type: the upper part is mainly characterized by loose detrital micrite while the internal and lower parts by syndepositional cemented autochthonous micrite.     

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.     

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.     

Calcification by Reef-Building Sclerobionts

It is widely accepted that deteriorating water quality associated with increased sediment stress has reduced calcification rates on coral reefs. However, there is limited information regarding the growth and development of reef building organisms, aside from the corals themselves. This study investigated encruster calcification on five fore-reefs in Tobago subjected to a range of sedimentation rates (1.2 to 15.9 mg cm⁻² d⁻¹). Experimental substrates were used to assess rates of calcification in sclerobionts (e.g. crustose coralline algae, bryozoans and barnacles) across key reef microhabitats: cryptic (low-light), exposed (open-horizontal) and vertical topographic settings. Sedimentation negatively impacted calcification by photosynthesising crustose coralline algae in exposed microhabitats and encrusting foram cover (%) in exposed and cryptic substrates. Heterotrophs were not affected by sedimentation. Fore-reef, turbid water encruster assemblages calcified at a mean rate of 757 (SD ±317) g m⁻² y⁻¹. Different microhabitats were characterised by distinct calcareous encruster assemblages with different rates of calcification. Taxa with rapid lateral growth dominated areal cover but were not responsible for the majority of CaCO₃ production. Cryptobiont assemblages were composed of a suite of calcifying taxa which included sciaphilic cheilostome bryozoans and suspension feeding barnacles. These calcified at mean rates of 20.1 (SD ±27) and 4.0 (SD ±3.6) g m⁻² y⁻¹ respectively. Encruster cover (%) on exposed and vertical substrates was dominated by crustose coralline algae which calcified at rates of 105.3 (SD ±67.7) g m⁻² y⁻¹ and 56.3 (SD ±8.3) g m⁻² y⁻¹ respectively. Globally, encrusting organisms contribute significant amounts of carbonate to the reef framework. These results provide experimental evidence that calcification rates, and the importance of different encrusting organisms, vary significantly according to topography and sediment impacts. These findings also highlight the need for caution when modelling reef framework accretion and interpreting results which extrapolate information from limited data.     

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.     

Microbial Biomineralization in Biotic Crusts from a Pleistocene Marine Cave (NW Sicily,Italy)

Biotic crusts occurring in the Early Pleistocene Rumena Cave, in NW Sicily, have been analyzed from a geomicrobiological point of view. The crusts consist largely of scleractinians and of subordinate bryozoans and serpuloideans, all typical of submarine cave biota. Encrustations document a blind cave in a shadowed setting, or possibly below the fair weather swell zone. Autochthonous and, subordinately, detrital fractions were observed within the skeletal framework of biotic crusts. The syndepositional lithified fraction occurs mainly as very fine-grained laminations. Clotted peloidal and aphanitic (structureless) textures occur in the micrites as well. Autochthonous micrite is always associated with a significant amount of organic matter remains. In caves from the Plemmirio area in SE Sicily, the autochthonous microbial micrite, occurring in the bioconstructions, contains bacterial lipid biomarkers, including abundant compounds derived from sulfate-reducing bacteria. It is likely that a similar microbial mediation was involved in the formation of the autochthonous micrite present in the biotic crusts of the Rumena Cave.     

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 role of bryozoans in fossil reefs—an example from the Middle Devonian of the Western Sahara

Stenolaemate bryozoans with their stable calcitic skeletons play a significant role in reef building. In the Middle Devonian Sabkhat Lafayrina reef complex (Western Sahara), bryozoans are abundant and diverse. Although they do not form part of the principal framework of reefs, bryozoans are involved significantly in reef growth, especially in the initial stage. In this way, bryozoans are important with respect to initiating reef growth. They contribute greatly to sediment stabilization, making it possible for principal reef builders to grow on hardened and stabilized substrates, and also play sediment-baffling and sediment-filling roles. The aim of this study is to document the diversity of bryozoans in a Middle Devonian reef complex and to estimate their potential for initiation and contribution to reef structures.     

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.     

Palaeoecology Of A Late Devonian Back Reef: Canning Basin, Western Australia

Back‐reef ecologies within the celebrated mixed carbonate‐siliciclastic Late Devonian (late Frasnian) Pillara Limestone of Windjana Gorge, in the Canning Basin, Western Australia, are re‐interpreted as being dominated by microbial communities. Proposed microbialites are expressed as weakly‐laminated, fenestral micrite, that show unsupported primary voids, peloidal textures, disseminated bioclastic debris, and traces of microfilaments. These grew as either extensive free‐standing mounds or columns, often intergrown with encrusting metazoans, or thick post‐mortem encrustations upon skeletal benthos. In some cases, microbial encrustations are inferred to have developed in protected cavities formed by progressive burial of the reef. The calcimicrobe Shuguria also shows a preferentially cryptic habit, encrusting either primary cavities formed by skeletal benthos, microbialite, or the ceilings of mm‐sized fenestrae within microbialite. A further calcimicrobe, Rothpletzella, formed columns up to 0.3 m high in areas enriched by very coarse siliciclastic sediment. Stromatoporoid sponges with a diverse range of morphologies also formed in situ growth fabrics. Monospecific thickets of closely‐aggregating dendroid stromatoporoid sponges (Stachyodes costulata), and platy‐laminar forms (?Hermatostroma spp.) were common, as were remarkably large stromatoporoids (Actinostroma spp.) that grew as isolated individuals up to 5 m in diameter. Such sponges showed impressive powers of regeneration from partial mortality, and individual clones may have been capable of substantial longevities of up to 500 years. Actinostroma spp. showed highly complex growth forms including platy‐multicolumnar (A. windjanicum), and a hitherto undescribed inferred whorl‐forming foliaceous morphology (Actinostroma sp.) reminiscent of the modern photosymbiotic coral Acropora palmata. These complex morphologies formed abundant primary cavities, previously thought to be only rarely developed in association with stromatoporoids.key words : Late Devonian, Canning Basin, reefs, palaeoecology, microbialite.     

Facies and biota of Anisian to Carnian carbonate platforms in the Northern Calcareous Alps (Tyrol and Bavaria)

Summary During the Middle and early Late Triassic carbonate ramps and rimmed platforms developed at the northwestern margin of the Tethys ocean. In the Northern Calcareous Alps, Anisian stacked homoclinal ramps evolved through a transitional stage with distally steepened ramps to huge rimmed platforms of Late Ladinian to Early Carnian age. Middle Triassic to early Late Triassic facies and biota of basin, slope and platform depositional systems are described. Special emphasis is given to foraminifers, sponges, microproblematic organisms and algae. The Ladinian to early Carnian reef associations are characterized by the abundance of segmented sponges, microproblematica, biogenic crusts and synsedimentary cements. Among the foraminifers, recifal forms likeHydrania dulloi andCucurbita infundibuliformis (Carnian in age) are reported from the Northern Calcareous Alps for the first time. Some sphinctozoid sponges likeParavesicocaulis concentricus were known until now only from the Hungarian and Russian Triassic.     

Will increased storm disturbance affect the biodiversity of intertidal,nonscleractinian sessile fauna on coral reefs?

Relatively little is known about how the future effects of climatic change, including increases in sea level, temperature and storm severity and frequency, will impact on patterns of biodiversity on coral reefs, with the notable exception of recent work on corals and fish in tropical reef ecosystems. Sessile invertebrates such as ascidians, sponges and bryozoans occupying intertidal rubble habitats on coral reefs contribute significantly to the overall biodiversity and ecosystem function, but there is little or no information available on the likely impacts on these species from climate change. The existing strong physical gradients in these intertidal habitats will be exacerbated under predicted climatic change. By examining the distribution and abundance of nonscleractinian, sessile invertebrate assemblages exposed to different levels of wave action and at different heights on the shore around a coral reef, we show that coral reef intertidal biodiversity is particularly sensitive to physical disturbance. As physical disturbance regimes increase due to more intense storms and wave action associated with global warming, we can expect to see a corresponding decrease in the diversity of these cryptic sessile assemblages. This could impact negatively on the future health and productivity of coral reef ecosystems, given the ecosystem services these organisms provide.     

Taphonomy of modern deep,cold‐temperate water coral reefs

Deep‐water corals are widely distributed along the cold‐temperate northeastern Atlantic continental margin. Despite the widespread occurrence of these aphotic coral constructions in deep shelf settings, the processes of framework formation and postmortem alterations which result in different preservational styles are still poorly known. Detailed mapping surveys on probably one of the largest Lophelia reef structures were carried out on the Sula Ridge, Mid‐Norwegian Shelf in 270 to 300 m depth. Side scan sonar records and camera surveys yield information at various scales of resolution on the reef complex which is more than 9 km long and up to 45 m high. Living Lophelia colonies effectively prevent colonization by other organisms and are successful in the rejection of passing detrital material from the soft tissue. In a healthy condition the coral is able to encrust repetitively attached organisms by selectively secreted sclerenchyme layers, thus, this defensive reaction results in the thickening of the skeleton. Early postmortem alteration in Lophelia colonies is introduced by the formation of a biofilm and Dodgella (fungi) infestation. The biofilm is associated with selective Fe‐Mn precipitation on the coral skeleton. This is the zone of intense attachment of sessile invertebrates such as serpulids, brachiopods, foraminifers and encrusting bryozoans. More advanced taphonomic stages show an increasing dominance in sponges which reduce the interskeletal framework porosity significantly. In addition, boring sponges excavate the thickly calcified Lophelia skeletons, thus leading to in situ collapsing structures on the sea floor. It is the intensity of sediment trapping biofilms and sponge colonization and the amount of imported detrital particles predominantly from the pelagial zone that control the generation of a pure coral rubble facies or the preservation of collapsed but mud‐rich detrital mounds.