Tubiphytes Maslov, 1956 and description of similar organisms from Triassic reefs of the Tethys

Tubiphytes Maslov (Nigriporella Rigby, Shamovella Rauser-Chernoussova) and similar organisms belong to one of the most abundant, enigmatic fossils reported from Carboniferous to Cretaceous rocks of numerous localities worldwide. Tubiphytes has been referred to many distinctly different organisms of the Tethys occurring in shallow water, particularly reef biotopes. Paleozoic Tubiphytes were revised by Senowbari-Daryan and Flügel, and Jurassic-Cretaceous species by Senowbari-Daryan et al. In this paper, the Triassic representatives of Tubiphytes and Tubiphytes-like organisms are described. The morphological and structural differences between the Paleozoic and Triassic Tubiphytes are shown and their systematic position is discussed. The following new species are described: Tubiphytes carnicus, T. ramosus, T. alcicornis, and Plexoramea cylindrica. The new genus Carniphytes is proposed for Tubiphytes multisiphonatus Schäfer and Senowbari-Daryan. Tubiphytes gracilis Schäfer and Senowbari-Daryan is transferred to the genus Plexoramea Mello. All described taxa are relatively abundant in Ladinian-Carnian reefs (e.g., “Wetterstein limestone” of the Alps) and in shallow-water carbonates of the northwestern Tethys. Some of them occur rarely in Anisian or in Norian-Rhaetian reefs.     

Upper Triassic reefs of the Oman Mountains: Data from the South Tethyan margin

Summary The Upper Triassic reefal limestones of the Oman Mountains were investigated with respect to their microfacies, palaeontology and community structure. The reef fauna described and figured for the first time occurs in parautochthonous slope deposits of the Arabian platform (Sumeini Group) and in allochthonous reefal blocks (‘Oman Exotics’, Hawasina Complex). The ‘Oman Exotics’ are tectonically dislocated blocks, derived from isolated carbonate platforms on seamounts in the Hawasina basin or in the South Tethys Sea. The lithofacies and fauna of these blocks comprise a cyclic platform facies with megalodonts, reef and reef debris facies. The reefal limestones are dated as Norian/Rhaetian by benthic foraminiferal associations (Costifera, Siculocosta, Galeanella) and typical encrusting organisms (Alpinophragmium, Microtubus). Some small ‘Oman Exotics’ are of Carnian age. The shallow-marine organisms include scleractinian corals of different growth forms, ‘sphinctozoans’, ‘inozoans’ chaetetids, spongiomorphids, disjectoporids and solenoporacean algae as the main reef builders, various encrusters like microbes, foraminifers, sponges and many different problematical organisms for the stabilisation of the reef framework and a group of dwellers including benthic foraminifers, gastropods, bivalves and a few dasycladacean algae. The reef communities are characterized by the coverage of organisms and distributional pattern. Analogies with the coeval reef deposits from the European part of the Tethys have been recognized. Some species, now collected in Oman, were also reported from American and Asian localities.     

Norian-Rhaetian reefs in Argolis Peninsula,Greece

Summary Upper Triassic to Lower Jurassic shallow-water carbonate sequences of the ‘Pantokrator limestones’ are widely distributed in the Argolis Peninsula, southern Greece. Within this sequence are some reef or reefal structures. In the Mavrovouni Mountains, near Sarmeika, 6 km SE of the ancient theatre of Epidavros (Argolis Peninsula), a Norian-Rhaetian reef complex has been identified. This is the first well-documented Norian-Rhaetian reef in Greece. The main reef builders are coralline sponges (‘sphinctozoans,’ ‘inozoans’, and sclerosponges), followed by dendroid, cerioid, and solitary corals, and algae. The reef type corresponds to a ‘sponge-coral reef’.     

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).     

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.     

Foraminifera and their associations of a possibly Rhaetian section of the Nayband Formation in central Iran,northeast of Esfahan

The following paper describes the foraminiferal fauna and associated faunal assemblages of the bedded and reef carbonates of the Upper Triassic (most probably Rhaetian) Nayband Formation, which are exposed in a section south of the small town of Bagher-Abad, northeast of Esfahan. Foraminifers are extremely rare in sponge- or coral-dominated bioconstructions and in the bedded carbonates of the Nayband Formation in central Iran. Some carbonate beds are composed of bioclastic wackstone/packstone. These are exposed in the solenoporacean horizon at the uppermost part of the section. Here, the aulotortid- and trocholinid-type foraminifers are relatively abundant. The following foraminiferal taxa with different abundances were found within the carbonates of the investigated section: Trocholina umbo Frentzen, T. turris Frentzen, T. gracilis Blau, Aulotortus tumidus (Kristan-Tollmann), Aulotortus tenuis (Kristan), Aulotortus friedli (Kristan-Tollmann), Coronipora etrusca (Pirini), Semiinvoluta clari Kristan, Turrispirillina? licia variabilis Blau, Galeanella? laticarinata Al-Shaibani, Carter and Zaninetti, Ophthalmidium sp., Agathammina sp., “Sigmoilina” schaeferae Zaninetti, Planiinvoluta carinata Leischner, Planiinvoluta sp., Nubecularia sp., Endothyra sp., Paeolituonella sp. and some sessile agglutinated and nodosariid types. All mentioned taxa are very rare, except the involutinid and trocholinid types. The following species are described as new: Trocholina blaui nov. sp., Spirilina? iranica nov. sp., and Coronipora serraforma nov. sp. Trocholina blaui is usually attached to solenoporacean thalli. Four foraminiferal associations, which are named after the occurrence of the abundant species, were distinguished as Aulotortus tumidus association, Aulotortus friedli association, Trocholina umbo association, and Trocholina blaui association. Aulotortid types and Trocholina umbo were found within the bioclastic wackstone/packstone carbonates. Trocholina blaui is abundant in solenoporacean framestones. The foraminiferal association of investigated carbonates contains a mixed fauna, known from Upper Triassic–Liassic in the Tethyan realm. Carbonates of the whole investigated section are dated—due to occurrence of the genus Aulotortus, with species A. tumidus (Kristan-Tollmann), A. tenuis (Kristan), and A. friedli (Kristan-Tollmann)—as Upper Triassic (most probably Rhaetian). The “typical” foraminifers occurring in the reef biotopes in the northwestern Tethys are either missing or extremely rare in the Iranian bioconstructions.     

Calcareous algae from the Triassic (Anisian reef boulders and Norian reef limestones) of Karaburun, western Turkey

Invertebrate Anisian and Norian reef boulders were found in the Gerence and Güvercinlik Formations, respectively, exposed in northeastern part of the Karaburun Peninsula, western Turkey. Halimedacean green algae and solenoporacean red algae, usually associated with other reef building organisms are described in this paper. The following taxa were determinated: halimedaceans: Egericodium hungaricum Flügel, Velledits, Senowbari-Daryan and Riedel, Aternasus irregularis n. gen., n. sp.; solenoporaceans: Solenopora cf. alcicornis Ott, Solenopora triasina Vinassa de Regny, Solenopora vachardi n. sp., Solenopora concentrica n. sp., Solenopora paraconcentrica n. sp., Tauristorea parallela Senowbari-Daryan and Link, Tauristorea discursa n. sp., and Parachaetetes cassianus (Flügel). The monospecific genus Aternasus n. gen. is the most abundant alga within the Anisian reef boulders. The solenoporaceans are represented by several taxa but the individual species are less abundant. E. hungaricum, T. parallela Senowbari-Daryan and Link, and S. triasina Vinassa de Regny are described from the Norian reef limestones embedded in Güvercinlik Formation, all other taxa from the Anisian reef boulders embedded in the Gerence Formation.     

Bioeroders in fossil reefs

Summary Boring algae, fungi and bacteria have been the most constant factor in bioerosion through earth history. Their record reaches back into the middle Precambrian. The only fossil reefs specifically researched for these microendoliths are of Triassic and Upper Jurassic age. Boring worms appear in reefs in the Lower Cambrian. Boring sponges and bivalves first appear also in the lower Paleozoic, but do not become abundant in reefs until the Triassic. Effective substrate excavating grazers are relatively young geologically: Patellids and substrate excavating Echinoids evolved in the Triassic but did not become important bioeroders until the Jurassic or Cretaceous. Scarid fishes are even younger, the oldest representatives having been found in the Miocene. Thus, it seems that the intensity of bioerosion changed significantly during earth history. This may have had consequences for diversity of reef organisms, quality and quantity of reef debris, for diagenesis and record of reef rock.     

<Emphasis Type="Italic">Crescentiella</Emphasis>-microbial-cement microframeworks in the Upper Jurassic reefs of the Crimean Peninsula

In the Upper Jurassic reef successions of the Crimean Peninsula (Sudak and Jalta areas), the microencruster Crescentiella morronensis (Crescenti), microbialites, and multiple generations of cements, form microframeworks. They were observed in two stages of the carbonate platform evolution, in the Middle–Upper Oxfordian, and in the Upper Kimmeridgian–Tithonian. Generally, in both stages, the features of the microframeworks are similar and consist of densely packed Crescentiella associated with microbialites and branched colonies of the sclerosponge Neuropora lusitanica Termier. The difference between the occurrences of the two stages is the variable amount of nubecularid foraminifera and enigmatic tube-shaped structures forming the central cavities of Crescentiella. The Crescentiella-microbial-cement microframeworks formed under phreatic conditions in the upper slope and seaward marginal depositional settings where intensive synsedimentary cementation took place. They formed in the initial stages of long cycles of restoration and blooming of the reefs. The late Jurassic examples resemble the Permian algae-microbial-cement reefs as well as the Triassic Tubiphytes and cement crust-dominated reefs. Concurrently, all these examples formed a transitional facies zone between typical slope facies to shallow subtidal platform margin facies characterized by high taxonomic diversity of calcified sponges, corals, and microencrusters forming the principal part of the reefs.     

Paleoecology of Middle Ordovician stromatoporoid mounds in Vermont

Fossiliferous mounds of carbonate mud are a distinctive facies in the middle Chazy Group (Crown Point Formation) at Isle La Motte, Lake Champlain. The mounds are surrounded by bedded calcarenite of spar-cemented pelmatozoan debris. Channels which cut into the mounds during mound growth are filled with the same calcarenite. The mud-free intermound rocks and the mound biota suggest agitated, normal marine shallow-water environments. The principal lime-secreting organisms within the mounds are stromatoporoids, calcareous algae, tabulate corals, sponges, and bryozoans. Each mound is dominated in terms of biomass by one of three groups: stromatoporoids, calcareous algae, and bryozoans. Most of the mound biota first appear at the base of the Crown Point Formation. In the lower Crown Point Formation the organisms increase in number and species. Both changes in the biota are related to periods of shallowing of the Chazy sea which are also reflected in the character of the carbonate sands.     

<Emphasis Type="Italic">Bevocastria magna</Emphasis> n. sp., eine neue Cyanobakterie aus den obertriassischen Riffkalken des Taurus-Gebirges (Türkei)

Bevocastria magna n. sp., an extremely large nodular species of the genusBevocastria, is described from Upper Triassic (Norian) reef limestones of a locality south of the town of Dereköy (SW Antalya, Turkey). The nodule ofB. magna looks like skeletons of chaetetid sponges which occur commonly at the same locality.B. magna is the first report of the genus from the Triassic deposits of Turkey.     

四川华蓥山地区晚二叠世生物礁中“板状水螅”化石的初步研究

所讨论的“板状水螅”是一类分类位置尚有争议的化石，为华蓥山地区上二叠统生物礁的主要造礁生物之一。这里描述了３属３种．其中包括２新属和３新种．它们是Ｐｓｅｕｄｏｐａｌａｅｏａｐｌｙｓｉｎａｈｕａｙｉｎｇｅｎｓｉｓ，Ｐｈｒａｇｍｏｒｐｈａａｓｉａｔｉｃａ和Ｃｎｉｄｏｐｏｒａｔｕｂｅｒｃｕｌｏｓａ。     

Solenoporaceen aus den obertriassischen (Nor) Riffkalken des Taurusgebirges (Antalya-Gebiet, Südtürkei)

From the Upper Triassic (Norian) reef limestones north and west of Antalya (Taurus Mts., Turkey) the red algae (Solenoporaceans)Solenopora undata n. sp.,Parachaetetes clatratus n. sp.,Parachaetetes riedeli n. sp.,Parachaetetes tauricus n. sp., andTauristorea parallela n. gen., n. sp. are described.Tauristorea is characterized by cells orientated parallel to the Substrate (vertically to the growth direction of the alga). InSolenopora undata andParachaetetes clatratus some cavity structures occur, whose function as reproductive organs are discussed.     

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.     

Microproblematica,calcareous algae,and microbialites at the Frasnian-Famennian boundary interval in the Šumbera section (Moravian Karst,Czech Republic) and their significance in the context of the Kellwasser Crisis

The Kellwasser Crisis represents one of the most severe extinction events in the Phanerozoic. The ?umbera section (Moravo-Silesian Basin, Moravian Karst) spans the Upper rhenana to Palmatolepis minuta minuta (or younger) conodont zones and corresponds to the upper part of a carbonate ramp with material derived from shallower areas. A rich association of calcareous algae, cyanobacteria, and microproblematica, comprising renalcids (Izhella), Girvanella, Rectangulina cf. tortuosa, Rothpletzella, “solenoporaceans”, “Keega”-like microfossils, udoteacean alga Paralitanaia, volvocean and radiospherid calcispheres, palaeoberesellids, Umbellina bella, and Wetheredella, is similar in diversity to associations known from China, Australia, Canada, and Belgium. The ?umbera section represents a unique locality where changes in shallow-water biota at the Frasnian-Famennian boundary are well calibrated by detailed conodont biostratigraphy. We track in detail the evolution of shallow-water biota and the features that are often discussed in the context of “anachronistic” facies, such as flourishing microbial structures, enhanced early marine cementation, and flat-pebble conglomerates. Some features which could be indicative of environmental stress such as flat-pebble conglomerates developed during the Frasnian part of the Kellwasser Crisis (Upper rhenana to linguiformis Zones), whereas a renalcid boom started at the Frasnian-Famennian boundary.     

A late-surviving Triassic protomonaxonid sponge from the Paris Biota (Bear Lake County,Idaho, USA)

Protomonaxonid sponges are a major group of Cambrian and Ordovician fossils in exceptionally preserved (especially Burgess Shale-type) faunas, but are rare thereafter. Rare examples of apparent surviving lineages are known from the late Palaeozoic and Mesozoic, but by this time more derived groups of sponges have generally displaced them in at least shallow-water (shelf depth) ecosystems. The early Spathian (Early Triassic) Paris Biota includes abundant material of a new leptomitid protomonaxonid, Pseudoleptomitus advenus Botting nov. gen., nov. sp., distinguished by having an unbundled longitudinal skeleton and very weak transverse component. This is the first post-Ordovician leptomitid known, and indicates long-term survival of the group in unknown environments. Its occurrence near storm wave base is similar to the preferred environment of earlier examples of the family, suggesting either ecological rarity or taphonomic reasons for their ∼200-million-year absence from later Palaeozoic rocks.     

Upper Triassic reef facies in the Asher-Atlit-1 borehole, Northern Israel: Microfacies, cement stratigraphy and paleogeographic implications

Summary  The late Triassic succession of the Asher-Atlit 1 borehole is over 1000 m thick, and is composed of reefal and associated facies interbedded with volcanics of Norian age. Only borehole cuttings are available. Microfacies classification and cement stratigraphy determined by optical and CL microscopy, allowed discrimination of six episodes of reef establishment, progradation, shallowing, and termination. Organic buildups are constructed of reef-building biota (sponges, possible corals, encrusting organisms) typical for the late Triassic of the Tethys. Reef-associated facies include fore-slope, central reef, ooid shoal, lagoonal, and supratidal environments. Cement zoning patterns trace diagenetic signatures which range from early neomorphic skeletal replacements and original marine cements, via characteristic burial sequences; depositional and diagenetic sequences are terminated by marginal marine intra- or supratidal conditions, and subaerial exposure with pedogenic overprints. Volcanic episodes tend to be associated with termination of carbonate sedimentation episodes, while volcanic quiescence and subsidence permit vertical progradation of reefal and associated facies. The depositional and progradational environment, rapid rate of sedimentation, periodicity, association with volcanics, and regional considerations, suggest a depositional setting on the rifted shelf-margin of the nascent Neo-Tethys, with a possible eustatic overprint.     

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

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

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.