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.     

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 micrite-dominated norian carbonate platform from Northern Calabria (Southern Italy)

Summary  This paper is focused on the facies associations of an Early-Middle Norian stratigraphic succession cropping outin the Northern Calabria (Buonvicino, Cosenza). These carbonate deposits, pertaining to the Verbicaro Unit, represent a dolomitized platform characterized by a clear dominance of automicrites over skeletal metazoans. On the basis of the chronostatigraphic data and sedimentary evolution, two stratigraphic units have been distinguished: the Lower Unit (object of the present paper) and the Upper Unit. The Lower Unit consists of an Early-Middle Norian high-relief prograding carbonate platform. The Upper Unit is represented by Middle p.p.-Upper Norian basinal deposits. Three main facies associations, indicative of different depositional settings, have been identified. The inner platform facies association is characterized by automicrite (cauliflower-columnar subordinate planar stromatolites) associated with detrital carbonates (intraclastic breccia, bioclastic grainstone and packstone). Apeculiar facies is represented by megalodontid-bearing beds. Mud-cracked horizons and low-developed teepees occur quite frequently. The margin facies association is dominated by automicrite (planar to low relief stromatolites associated with thrombolitic fenestral boundstone) and detrital carbonates with subaerial exposure features. The slope facies association includes detrital carbonates (breccia/megabreccia) associated with serpulid/sphinctozoan bioconstructions and automicrite (planar stromatolites/thrombolitic boundstone). The whole carbonate body is completely dolomitized; nevertheless the morphology and microarchitecture of carbonate components (cements, grains and automicrite) is still clearly recognizable. The dolomite Mg content ranges from 40 to 48 mole%, sometimes reaching the stoichiometric value. Cements, primary and late, represent a minor component of the rock volume; they occur more frequently on the margin and upper slope setting. Quantitative tacies analyses lead to the following conclusions:

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.     

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.     

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.     

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).     

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     

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.     

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

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

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.     

Coral Community Adaptability to Environmental Change at the Scales of Regions, Reefs and Reef Zones

SYNOPSIS. Projected global increases in temperature, sea level,storminess and atmospheric carbon dioxide (CO₂) are likely tocause changes in reef coral communities which the present humangeneration will view as deleterious. It is likely coral communitytrajectories will be influenced as much by the reduction inintervals between extreme events as the projected increasesin means of environmental parameters such as temperature, atmosphericCO₂ and sea-level. Depressed calcification rates in corals causedby reduced aragonite saturation state of water may increasevulnerability of corals to storms. Moreover, reduction in intervalsbetween storms and other extreme events causing mass mortalityin corals (coral predators, diseases, bleaching) are likelyto more frequently "set back" reef coral communities to earlysuccessional stages or alternate states characterized by non-calcifyingbenthos (plants, soft corals, sponges). The greater the areaand the longer the duration of dominance of putative "coral/corallinealgae" zones of coral reefs by non-calcifying stages, the lesswill be the reefs capacity to accrete limestone bulk lockedup in the big skeletal units of late successional stages (i.e.,very large old corals). Averaged over decades to centuries,the effects of such changes on the coral community's carryingcapacity for other biota such as fish are unpredictable. A "shiftingsteady-state mosaic" null model may provide a useful conceptualtool for defining a baseline and tracking changes from thatbaseline through time.     

Cyclone effects on coral reef habitats in New Caledonia (South Pacific)

The impacts of the unusually strong Cyclone Erica (March 2003) on coral reef habitats at a site located on the northwest coast of New Caledonia (South Pacific) were assessed using a 6-year data set (2002–2007). We examined the interannual variations of key variables describing reef habitats (live hard and soft corals, dead corals in place, coral debris, algae and relative proportion of mechanically vulnerable and resistant live hard corals). The cyclone-induced disturbances of habitats differed according to three reef types: patch reefs, barrier reefs far from passes (more than 3 km from the nearest pass) and barrier reefs near passes (less than 3 km from the nearest pass). Short-term mechanical damage was detected on the three-dimensional structure of reef habitats with a notable shift from a community dominated by mechanically vulnerable corals to one dominated by resistant corals on barrier reefs far from passes. The history of habitats and their pre-disturbance characteristics, in link with local hydrodynamics, was found to influence their short-term susceptibility to extreme events such as cyclones. However, the most significant effects appeared in the midterm (within 2 years after the cyclone) as the cover of live hard corals significantly decreased by approximately 45% between 2002 and 2004 on all reef types. The short- and midterm disturbances of coral reef habitats are discussed with regard to published temporal variations in reef fish assemblages, underlining the delayed effects of this cyclonic event on fish as well as benthic habitats. Coral reef habitats and live corals had shown significant patterns of recovery 4 years after the cyclone, followed by similar recovery in fish community, suggesting good resilience in a face of this major natural disturbance in an area under moderate anthropogenic pressure.     

Tolerance of Sponge Assemblages to Temperature Anomalies: Resilience and Proliferation of Sponges following the 1997–8 El-Ni?o Southern Oscillation

Coral reefs across the world are under threat from a range of stressors, and while there has been considerable focus on the impacts of these stressors on corals, far less is known about their effect on other reef organisms. The 1997–8 El-Niño Southern Oscillation (ENSO) had notable and severe impacts on coral reefs worldwide, but not all reef organisms were negatively impacted by this large-scale event. Here we describe how the sponge fauna at Bahia, Brazil was influenced by the 1997–8 ENSO event. Sponge assemblages from three contrasting reef habitats (reef tops, walls and shallow banks) at four sites were assessed annually from 1995 to 2011. The within-habitat sponge diversity did not vary significantly across the study period; however, there was a significant increase in density in all habitats. Multivariate analyses revealed no significant difference in sponge assemblage composition (ANOSIM) between pre- and post-ENSO years for any of the habitats, suggesting that neither the 1997–8 nor any subsequent smaller ENSO events have had any measurable impact on the reef sponge assemblage. Importantly, this is in marked contrast to the results previously reported for a suite of other taxa (including corals, echinoderms, bryozoans, and ascidians), which all suffered mass mortalities as a result of the ENSO event. Our results suggest that of all reef taxa, sponges have the potential to be resilient to large-scale thermal stress events and we hypothesize that sponges might be less affected by projected increases in sea surface temperature compared to other major groups of reef organisms.     

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

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

Rugose corals and associated carbonate microfossils from the Brigantian (Mississippian) of Castelsec (Montagne Noire, southern France)

The disused quarry east of Castelsec offers a view of shallow-marine carbonates of the poorly known Uppermost Mississippian of the Montagne Noire. At Castelsec, sections are studied in two characteristic facies types (bioclastic wackestone and microbial dominated boundstone) of the Upper Mississippian. The succession is rich in rugose corals and carbonate microfossils. Six genera with seven species belonging to a rugose coral fauna consisting of at least eight genera with several species are described herein; Dibunophyllum castelsecensis sp. nov. is described as new. Twenty-seven carbonate microfossils of different groups have been identified. The Castelsec succession is Brigantian in age, based on the stratigraphic occurrence of rugose corals, foraminifers, and calcareous algae observed in both sections. The rugose coral fauna shows relationships with the well-known fauna of northwestern Europe and the Ouralian-Asian Province. Typical elements of northwestern Europe are missing at Castelsec and vice versa. This differentiation between north and south is interpreted as responses to different palaeolatitudes and tectonic settings.     

Sponge-mediated coral reef growth and rejuvenation

Sponges mediate consolidation of Porites furcata rubble on shallow Caribbean reefs by quickly adhering to rubble and stabilizing it until carbonate secreting organisms can grow and consolidate it to the reef. Experimental investigations demonstrate that the entire cycle from (1) temporary binding of rubble by sponges, through (2) rubble consolidation by encrusting coralline algae, to (3) colonization of consolidated rubble by corals, can be completed within 10 months. Bound rubble both adds to vertical reef growth and also provides stable substrata for colonization by corals. Corals that colonize stabilized rubble are damaged less and survive better than on unstable rubble. Rubble that is not temporarily stabilized by sponges does not become bound to the reef, because continuous movement disturbs the consolidation process, and does not provide suitable substrata for settlement and growth of corals. Sponge-mediated consolidation of rubble may increase rates of reef growth and enhance reef recovery after damage. This new role for sponges in reef growth is not obvious from examination of the internal fabric of a reef frame. Spongemediated consolidation may help to explain geographic and temporal differences in growth and morphology among shallow reefs of ramose corals.     

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.     

Biotic structure and morphology of patch reefs from South China (Ningqiang Formation, Telychian, Llandovery, Silurian)

Summary Ningqiang Formation (late Telychian, Llandovery, Silurian), characterized by nearly 3000 m of shales in tercalated with carbonates, is situated between Ningqiang (S. Shaanxi Province) to Guangyuan (N. Sichuan Province) adjacent to the northwest margin of the Yangtze Platform. The high diversity “Xiushan Fauna”, and abundant reef development, illustrate a relatively warm and persistent shallo marine environment in these early Silurian sediments. The sequence shows reef radiation after recovery from the end Ordovician mass extinction envents. Multiple horizons of reef-building occurred within a relatively short geological interval and resulted in more than 30patch reefs up to 200 m in diameter and 1–50 m vertically, composed of abundant fossils. Reef biota include frame-building corals, stromatoporoids, bryozoans, and microbialites, and reef-associated oranisms such as crinoids, brachiopods, trilobites, gastropods, nautiloids and ostracods. Three reefrelated biotic associations are recognised: a) reefs dominated by framework with crinoids and microbia; b) reefs dominated by only crinoids and microbia; and c) crinoiddomainated facies. Seven representative reef examples illustrate different morphologies and growth styles. A high terrigenous debris input and shallow epicontinental ramp, which lacked obvious topographic variation, were major controls which resulted in rather simple reefs; sedimentation was apparently the main constraint on lateral and vertical extension of reefs, and prevented large-scale reef complexes developing.