Upper Permian (Murghabian) rugose corals from Oman (Ba’id area,Saih Hatat): Community structure and contributions to reefbuilding processes

Summary Coral-dominated communties are rare in Upper Permian reefs. The study of Murghabian rugose and tabulate corals from allochtonous carbonates (‘Oman exotics’) of the Hawasina Complex and autochthonous carbonates of the Saih Hatat area/Arabian Platform (Oman) provides evidence for a significant contribution of rugose corals to the formation of Late Permian reefs. The corals are described with respect to taxonomy, microfacies and community structure. 8 genera and 7 species were recognized.Monothecalis minor n.sp.,Praewentzelella regulare n.sp. andWentzelella katoi magna n.ssp. are new. The corals represent three communities: (1)Praewentzelella community (Hawasina Complex), (2) cerioid coral community (Hawasina Complex), and (3)Waagenophyllum community (Hawasina Complex and Saih hatat). The corals from the Hawasina Complex and the Saih Hatat flourished in significantly different environments: Rugosa from the Hawasina Complex are representatives of reefs, whereas their counterparts from the Saih Hatat lived in level-bottom communities. Coral-bearing reefal boundstones are characterized by a diverse assemblage of sphinctozoans, inozoans, chaetetids, bryozoans, crinoids,Tubiphytes, Archaeolithoporella and algae. These communities produced bafflestones or framestones and were part of a sponge reef complex. The level-bottom community of the Saih Hatat is low-diverse only comprising rugose and tabulate corals. These of isolated colonies locally acted as bafflers.     

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).     

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

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

Bioerosion in Late Permian Rugosa from reefal blocks (Hawasina Complex,Oman Mountains): implications for reef degradation

Summary Rugose corals are known from allochthonous Late Permian reefal blocks of the A1 Jil and Ba’id Formation (Hawasina Complex), Oman Mountains. In contrast to many Late Permian Rugosa found elsewhere in the Tethys, they occurred in sponge reefs and contributed to reef construction. The waagenophyllid warm water coral fauna is moderately diverse comprising cerioid, thamnasterioid, and fasciculate taxa. In contrast to sponges, chaetetids, and low-growing reefbuilders, the corals secreted diagenetically stable, most probably Mg-calcitic skeletons. Borings in coral skeletons are consequently well preserved providing important data for the interpretation of reef destructive processes. Thin-section analysis revealed three taxa of infaunal borers includingEntobia Bronn 1837, uncertain thallophyte borings, and borings of unknown bioeroders. Macroborers were more important than microborers, because of the dominance of clionid sponges. Good evidence exists also for the occurrence of two types of undetermined grazers which destroyed the coral surfaces. The amount and distribution of bioerosions is variable among different coral taxa. The fasciculate coralPraewentzelella regulare Flügel 1995 was the favorate substrate. Up to 33% of the calices were bored. Dendroid and compound corals were bored subordinately. Bioerosion of these colonies does not exceed 2%. There is good evidence for substrate preference amongst the borers. Major controlling factors affecting borer distribution are believed to be variations of skeletal density and gross morphology. The borer assemblage could not limit reef accretion significantly. Factors controlling boring activity might have been quality of substrate, sedimentation rate, rapid incrustation of substrates, and competition for food with reef constructors including sponges, chaetetids, and rugose corals.     

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’.     

Anthracoporella mounds in the Late Carboniferous Auernig Group,Carnic Alps (Austria)

Summary A heretofore undocumented example of skeletal mounds formed by the dasycladacean algaAnthracoporella spectabilis is described from mixed carbonate-clastic cycles (Auernig cyclothems) of the Late Carboniferous (Gzhelian) Auernig Group of the central Carnic Alps in southern Austria. The massive mound facies forms biostromal reef mounds that are up to several m thick and extend laterally over more than 100 m. The mound facies is developed in the middle of bedded limestones, which are up to 16 m thick. These limestones formed during relative sea-level highstands when clastic influx was near zero. The mound facies is characterized by well developed baffler and binder guilds and does not show any horizontal or vertical zonation. Within the massive mound faciesAnthracoporella is frequently found in growth position forming bafflestones and wackestones composed of abundantAnthracoporella skeletons which toppled in situ or drifted slightly.Anthracoporella grew in such profusion that it dominated the available sea bottom living space, forming ‘algal meadows’ which acted as efficient sediment producers and bafflers. BecauseAnthracoporella could not provide a substantial reef framework, and could not withstand high water turbulence, the biostromal skeletal mounds accumulated in shallow, quiet water below the active wave base in water depths less than 30 m. The massive mound facies is under- and overlain by, and laterally grades into bedded, fossiliferous limestones of the intermound facies, composed mainly of different types of wackestones and packstones. Individual beds containAnthracoporella andArchaeolithophyllum missouriense in growth position, forming “micromounds’. Two stages of mound formation are recognized: (1) the stabilization stage when bioclastic wackestones accumulated, and (2) the skeletal mound stage when the sea-bottom was colonized byAnthracoporella and other members of the baffler and binder guilds, formingAnthracoporella bafflestones and wackestones of the mound facies. A slight drop in sea-level led to the termination of the mound growth and accumulation of organic debris, particularly calcareous algae, fusulinids, crinoids and bryozoans, forming well bedded limestones, which overlie the mound facies     

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.     

Distribution and sediment production of large benthic foraminifers on reef flats of the Majuro Atoll,Marshall Islands

The distributions and population densities of large benthic foraminifers (LBFs) were investigated on reef flats of the Majuro Atoll, Marshall Islands. Annual sediment production by foraminifers was estimated based on population density data. Predominant LBFs were Calcarina and Amphistegina, and the population densities of these foraminifers varied with location and substratum type on reef flats. Both foraminifers primarily attached to macrophytes, particularly turf-forming algae, and were most abundant on an ocean reef flat (ORF) and in an inter-island channel near windward, sparsely populated islands. Calcarina density was higher on windward compared to leeward sides of ORFs, whereas Amphistegina density was similar on both sides of ORFs. These foraminifers were more common on the ocean side relative to the lagoon side of reef flats around a windward reef island, and both were rare or absent in nearshore zones around reef islands and on an ORF near windward, densely populated islands. Foraminiferal production rates varied with the degree to which habitats were subject to water motion and human influences. Highly productive sites (>10³ g CaCO₃ m⁻² year⁻¹) included an ORF and an inter-island channel near windward, sparsely populated islands, and a seaward area of a reef flat with no reef islands. Low-productivity sites (<10 g CaCO₃ m⁻² year⁻¹) included generally nearshore zones of lagoonal reef flats, leeward ORFs, and a windward ORF near densely populated islands. These results suggest that the distribution and production of LBFs were largely influenced by a combination of natural environmental factors, including water motion, water depth, elevation relative to the lowest tidal level at spring tide, and the distribution of suitable substratum. The presence of reef islands may limit the distribution and production of foraminifers by altering water circulation in nearshore environments. Furthermore, increased anthropogenic factors (population and activities) may adversely affect foraminiferal distribution and production.     

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.     

Microbial origin of travertine fabrics—two examples from Southern Germany (Pleistocene stuttgart travertines and miocene riedöschingen Travertine)

Summary In Southern Germany, two examples of travertines of different age and depositional morphology were examined in detail. Travertines are laminated carbonate rocks formed by precipitation from mineral and/or thermal waters. They include characteristic facies types, such as bushy layers (‘shrubs’) referred to calcification of branching microbes (‘Dichothrix’-morphotype), laminar microbial mats, peloidal layers, and gas bubble layers formed within the sediment. In travertines, microbial activity is the most important factor for carbonate precipitation. Tufas differ from travertines by their abundance of molds of higher plants (leaves, reed, moss, green algae). They may be associated with travertines, but do not exhibit strict travertine facies types. Tufas are common in normal fresh water environments. Contrary to travertines and tufas, calcareous sinters usually occur in restricted areas like spring fissures, caves, or in pores, where microbial activity is not totally absent, but not of paramount importance for precipitation. Pedogenetic processes, which can alter travertine deposits, are responsible for large-scale features such as tepee-structures, and some intraclastic layers, and microscopic structures like endolithic borings andMicrocodium. Travertines may also grade into lacustrine limestones with Characeae, ostracods, and aquatic gastropods.     

The western edge of the Mediterranean Pelagian Platform: A Messinian mixed siliciclastic–carbonate ramp in northern Tunisia

The marine Messinian deposits of Tunisia cover a narrow littoral strip some 300 km long between the northern Bizerte and Cap Bon areas and the central–eastern Sahel region. Litho- and biofacies analysis of six stratigraphic sections reveals the distinctive features of these deposits.The lower Messinian deposits are characterized by ubiquitous siliciclastics and abundant oolitic/bioclastic limestones organized in an eastward facing ramp. Westward (landward), this ramp changes into coastal lagoons, sometimes containing evaporites. Eastward, the ramp passes to the reefal Pelagian Platform extending as far as Lampedusa.Two main sedimentary cycles are distinguished: 1) an early Messinian siliciclastic retrogradational then oolitic/bioclastic progradational cycle (Beni Khiar Formation and lower Oued bel Khedim Formation); 2) a late Messinian brackish to continental cycle that probably accumulated in rapidly subsiding lagoons (Oued el Bir Formation and upper Oued bel Khedim Formation). The Tunisian early Messinian cycle is partly eustatically controlled, but the late Messinian cycle cannot be confidently correlated to other well-known Messinian series because of tectonic movements.The lower Messinian deposits of Tunisia are also characterized by abundant suspension-feeding organisms (molluscs and bryozoans) and rare corals, calcareous algae, echinoids, and larger benthic foraminifers. The proposed palaeoenvironmental model shows that the lower Messinian ramp of Tunisia was located on a current-protected margin and subjected to continent-derived sediment and nutrient supply. Eastward, nutrient influx diminished and a shallow-water isolated carbonate platform with coralgal facies developed between the western and the eastern Mediterranean basins. The main hydrological connection between these two basins occurred through a narrow seaway situated to the northeast of the Pelagian Platform, south of Sicily and Malta.     

Paleoecology of the cenozoic reefal foraminifers and algae — A brief review

The paleoecology of reefal foraminifers and algae assumes a considerable importance in determining and delineating sub-environments of ancient reefs, especially those of non-coral origin.A review of the ecologic distribution of the Cenozoic larger foraminifers in different biofacies of the reef-complex environment has revealed the following: (1) a prolific growth of “Alveolina” was possible in the back-reef region near the reef core; (2) Orbitolites and Marginopora preferred sheltered waters on the reef-flat and in the back-reef zones; (3) nummulitids and Discocyclina thrived in both fore- and back-reef shoal areas, but the species living in the former are much stouter than those living in the latter; (4) Heterostegina is and, in the geologic past, was a form, preferring quieter waters of the back-reef lagoons and reef-flat pools; (5) Pellatispira was a typical fore-reef form.Smaller foraminifers, as a whole, are dominant in back-reef lagoons. An abundance of miliolids indicates a sheltered environment prevailing in the reef-flat pools and back-reef zones, whereas reef flats, in general, are characterized by a paucity of smaller foraminifers. An increase in the number of nodosariids and globigerinids points to a fore-reef environment, the depth of which is indicated by the relative abundance of the latter group. Encrusting foraminifers are characteristic of the reef core and are important constituents of for-algal (foraminiferal + algal) reef complexes.Of the algae, the calcarous chlorophyte Halimeda is relatively more abundant in the sheltered parts of a reef-complex, especially the lagoons, where water is moderately agitated and clear; its sudden abundance in the geologic record indicates the advent of a reefal environment. An abundance of the calcareous chlorophyte Dasycladaceae indicates the shallow back-reef areas adjacent to the reef core. Articulated coralline algae are associated with reef-complexes but are varied in their adaptability and, hence, are widely distributed in different parts of the complex. Abundant crustose coralline algae almost certainly indicate a reef-core sub-environment; their skeletons are among the chief constructional units of the core. They increase in abundance towards the outer edge of the reef core and decrease away from it.     

The influence of prey communities on fish species assemblages on artificial reefs in Puget Sound,Washington

Synopsis Research on eleven artificial reefs in Puget Sound, Washington examined the relative importance of reef-produced prey items to recreationally important reef fish species assemblages. The colonization of potential prey items, and fish species assemblages to ten artificial reefs were examined for the reefs first two to five years, and observations were conducted on an eleventh reef during its forty-ninth productive year. Fish species became more abundant, or were seen more frequently on reef habitats whose substrates had successionally developed from barnalces to algal mats. Fish species most affected by this successional change foraged heavily on organisms which were associated with reef algae. Starfish and nudibranchs. who preyed on the barnacles, were identified as the ‘keystone’ predators of these subtidal reef habitats.     

Response of triassic reef coral communities to sea-level fluctuations, storms and sedimentation: Evidence from a spectacular outcrop (Adnet, Austria)

Summary The Upper Rhaetian coral limestone of Adnet, southeast of Salzburg Austria has been repeatedly referred to as one of the most spectacular examples of an ancient ‘autochthonous’ coral reef structure. The ‘Tropfbruch’ quarry is probably the best outcrop for interpreting the distributional patterns of biotic successions and communities of a late Triassic patch reef. Our study is based on the interpretation of a) outcrop photographs, b) reef maps resulting from quadrat transects, and c) the analysis of quantitative data describing the distribution and frequency of reef organisms and sediment. A new methodological approach (combination of reef mapping and photo-transects) is used to obtain quantitative field data which can be compared in greater detail with data from modern coral reefs investigated by corresponding quantitative surveys. Three unconformities and three well-defined ‘reef growth stages’ reflecting the vertical and lateral development of the reef structure were differrentiated using transects: Stage 1, representing the reef growth optimum, is characterized by laterally differentiated coral reef knobs with corals in growth position. Criteria supporting this interpretation are the extraordinary size of the corals, their preservation in situ and the great thickness of this interval. The massive coralPamiroseris grew under higher energy conditions at the rim of the reef knob, whereas branchingRetiophyllia colonies preferred less agitated water in the center. Vertical changes are reflected by an increase in frequency of the dasycladacean algaDiplopora adnetensis and by the decreasing size ofRetiophyllia. These sedimentological and biological criteria together with the unconformity above indicate a fall in the sea level as a major control mechanism. Stage 2, separated from stage 1 by an unconformity caused by partial subaerial exposure and karstification, is characterized by vertically stacked coral successions with diverse reef debris. Facies heterogeneity is reflected by differences in the diversity, taphonomy and packing density of reef-building organisms as well as by differences in sediment input from the platform. Water depths and accommodation space were lower, therefore minor sea level fluctuations had a stronger effect on the biotic composition. The high percentage of coral debris and corals reworked by storms and the increase in the input of platform sediment led to a reduction of reef growth. Stage 3, again separated at the base by an unconformity associated with karstification, is characterized by bioclastic sediments with isolated reefbuilders forming a level-bottom community. The distribution of different coral morphotypes suggests that sea level fluctuations were not the only controlling factor. Variations in the substrate were caused by differences in the input of platform sediment. The three-step development seen in Adnet documents the response of low-diverse coral associations to variations caused by small-scale sea level changes, storm activity and sedimentation. The vertical changes in reef community structures correspond to a sequence of ‘allogenic replacements’. The Adnet reef structure should not be regarded as a general model of Alpine Upper Rhaetian reefs, because of the particular setting of the patch reef. Only the ‘capping beds’ of the Upper Rhaetian Reef Limestone of the Steinplatte exhibit criteria similar to Adnet. Potential modern analogues of features seen in the coral communities of Adnet are the internal structure of theRetiophyllia thickets, the key role of branching corals within the communities, the scattered distribution and low and even diversity of corals subsequent to breaks in settlement, segration patterns of corals indicating ‘contact avoidance’, toppling of large coral colonies by intensive boring, and decreasing coral coverage from deeper and sheltered settings to more shallower water depths.     

The late jurassic ‘Massenkalk fazies’ of Southern Germany: calcareous sand piles rather than organic reefs

Summary Upper Jurassic (Malm δ to ζ1) massive limestones (‘algal-sponge-reefs, sponge-reefs, reef-complexes, reefs, algal-sponge-bioherms, biolithites, Massenkalk, bioherms, Stillwasser-Mudmounds’) were analyzed in the Southern Swabian Alb, the Southern Franconian Alb and in drilling wells in the Molasse basin (Southern Bavaria). This analysis was carried out within the frame of a multidisciplinary DFG-study with the objective of decifering the controls on the development of Upper Jurassic spongiolites, their three-dimensional distribution, their characteristic faunal composition, and the diagenetic trends of the different primary facies. The data base consists of detailed facies mapping in the areas of the Eybtal and the Blautal (1300 samples) as well as comparative studies in the Upper Donautal (Swabian Alb) and the Southern Franconian Alb (400 samples). All together about 500 thin sections were studied. The distribution of the most important components (ooids, intraclasts, peloids, corals, sponges, sponge spicules, cyanobacterial crusts, brachiopods, molluscs, echinoids, bryozoans, serpulids,Terebella, Tubiphytes), and diagenetic features (dolomite, dedolomite, silicification, stylolites, clay flasers, hematite patches) results in a spatial distribution pattern of facies types. The largest part (70 %) of the massive limestones consists of a peloid-lithoclast-ooid sand facies rich in completely or partly micritized ooids. These ooids, especially in beds of the Malm δ to ε, might be the clue to a reinterpretation of the water depth. True biogenic constructions occur (about 30 % of the volume; sponge-algalmudmounds, algal-sponge-boundstones, and brachiopod-algal-sponge-mounds) within and at the margins of this facies and are interpreted as platform sands. The spatial distribution of the buildups in relation to the sand facies was probably controlled by hydrodynamic conditions. In addition, zoned sponge-algal-mounds occur in intraplatform channels and nodular sponge-algal-mudmounds in the marly basin sediments between platform sand areas. Breccias and slumpings in beds older than the Malm ζ have to be reinterpreted. Most of the breccias found originated from the flanks of the sand platforms, reflecting the faunal composition of the algal-sponge-boundstones which stabilized the flanks. Breccias of this composition occur throughout the Malm δ-ζ1 and differ markedly in their composition from the sand facies. The boundary breccia (Malm ε/ζ1) is interpreted as marking a regressive maximum. The increasing growth of buildups, rich in brachiopods in the Malm ζ1, is ascribed to an increase of reef growth at the beginning of a transgression. Detailed facies analyses necessary for the reconstruction of the spatial distribution of different facies types are in progress. Most of the older data on faunal distributions cannot be used for detailed facies analysis because they differentiated only between massive facies and bedded facies. Therefore Upper Jurassic limestones of Southern Germany should be restudied in order to recognize the volumetric importance of sand facies and buildups within massive limestones.     

Changes in gastropod assemblages in freshwater habitats in the vicinity of Basel (Switzerland) over 87 years

Net pen fish farms generally enrich the surrounding waters and the underlying sediments with nutrients and organic matter, and these loadings can cause a variety of environmental problems, such as algal blooms and sediment anoxia. In this study we test the potential of biofiltration by artificial reefs for reducing the negative environmental impacts surrounding fish farms in the Gulf of Aqaba, Red Sea. Two triangular-shaped artificial reefs (reef volume 8.2 m³) constructed from porous durable polyethylene were deployed at 20 m; one below a commercial fish farm and the other 500 m west of this farm in order to monitor the colonization of these reefs by the local fauna and to determine whether the reef community can remove fish farm effluents from the water. Both reefs became rapidly colonized by a wide variety of organisms with potential for the removal of compounds released from the farms. Within the first year of this study fish abundances and the number of species reached 518–1185 individuals per reef and 25–42 species per reef. Moreover, numerous benthic algae; small sessile invertebrates (bryozoa, tunicates, bivalves, polychaetes, sponges, anemones) and large motile macrofauna (crustaceans, sea urchins, gastropods) settled on the reef surfaces. Depletion of chlorophyll a was measured in the water traversing the artificial reefs in order to assess the biofiltration capacity of the associated fauna. Chlorophyll a was significantly reduced to a level 15–35% lower than ambient concentrations. This reduction was greatest at intermediate current speeds (3–10 cm s^–1), but was not influenced by current direction. The reef structures served as a successful base for colonization by natural fauna and flora, thereby boosting the local benthic biodiversity, and also served as effective biofilters of phytoplankton.     

Vertical zonation of megabenthic taxa on a deep photosynthetic reef (50–140 m) in the Au’au Channel,Hawaii

This study surveyed several locations at depths between 50 and 140 m within the Au’au Channel, Hawaii to characterize the deep reef habitat and determine the depth distribution and relative abundance of the dominant, habitat forming megabenthic taxa. In the Au’au Channel, the depth distribution of megabenthic taxa exhibited a pattern of vertical zonation with relatively few taxa dominating each zone. Macroalgae particularly Halimeda spp. and to a lesser extent scleractinian corals Leptoseris spp. were dominant between 50 and 80 m; Leptoseris spp. were dominant between 80 and 90 m as macroalgae decreased in abundance; the invasive octocoral Carijoa riisei was dominant between 90 and 100 m primarily on rugose features; Antipathes spp. and Leptoseris spp. were dominant between 100 and 120 m on exposed fossil reef; and small wire corals were dominant between 120 and 140 m. In general, the percentage of live benthic cover decreased with depth, particularly below 90 m where a large majority of the area was uncolonized, soft substrata. The gradient of downwelling light intensity appears to play a major role in regulating the depth distribution of photosynthetic organisms, skilophilous organisms, and other benthic fauna, which compete for space with dominant photosynthetic species. The depth of the seasonal thermocline also appears to play an important role in limiting the distribution of tropical benthic species.     

Coral communities and reef growth in the southern Great Barrier Reef

 Fringing reef development is limited around 22° S along the inner Great Barrier Reef, although there is substantial development north and south of this latitude. This study examined the relationships among coral communities and the extent of reef development. Reefs were examined to determine coral composition, colony abundance, colony size and growth form between the latitudes 20°S and 23°S. Major reef framework builders (scler- actinian genus Acropora and families Faviidae and Poritidae) dominated reefs north and south of 22°S, but declined significantly at 22°S where foliose and encrusting corals (Turbinaria and Montipora spp.) were most common. Porites spp. were present at 22° S but had encrusting morphologies. Consistently high turbidity at this latitude, caused by a 10 m tidal range and strong tidal flows, resuspends silts from the shallow shelf, and appears to have precluded reef development throughout the Holocene, by limiting the abundance, stunting the growth, and shortening the life expectancies of reef framework corals. The distinctions between ‘natural’ and ‘human-induced’ degradation may be interpreted on the basis of the relationship between Holocene development and current benthic community longevity. A mismatch between substantial past reef building capacity (a broad and/or thick reef) and non-existent or limited present reef-building capacity could signify anything from a long-period, natural cycle to an unprecedented deterioration in ecosystem function caused by human influence. Accepted: 29 July 1996     

Autochthonous facies and allochthonous debris flows compared: Early oligocene carbonate facies patterns of the Lower Inn Valley (Tyrol, Austria)

Summary This study presents a microfacies analysis and palaco-environmental interpretations of Early Oligocene carbon ates from the Lower Inn Valley Tertiary (“Unterinntal-Terti?r”) of Austria. The well preserved biogenic components allow detailed investigations of component relationships and controlling ecological parameters. The carbonates are dominated by coralline algae, corals, small and large benthic foraminifers, bryozoans and lithoclasts. Bivalves, gastropods, echinoderms, brachiopods and serpulids are subordinate. The limestones are present as A) autochthonous carbonates transgressing directly above the Triassic basement and B) allochthonous debris flows within deeper-water marls. These carbonates are found within the Paisslberg Formation. The Werlberg Member within this formation, pertains to the autochthonous carbonates and larger debris flows. Five facies types are separated following fabric analysis and statistical treatment (correlation, cluster analysis, principal components analysis) of semi-quantitative data consisting of component frequencies of thin sections. Facies distribution patterns are principally controlled by variations in substrate characteristics, turbulence and light along a depth gradient. Reconstruction of facies pattern distribution reveal both lateral and proximal-distal facies trends: coral-coralline algal facies, coralline algal facies as well as foraminiferal facies were situated in shallower environments, laterally adjacent to each other. These grade distally into coralline algal-bryozoan facies, bryozoan facies and finally into mollusc rich marls. Debris flows consisting of reworked material from all of the known facies (bioclastic packstone facies) is restricted to the debris flow and possible represents transport induced differentiation of components and grain size within distal debris flows.     

Pleistocene coral reefs associated with claystones, Southwestern Taiwan

 Scleractinian coral reefs, when coexistent with siliciclastic sediments, usually occur in association with deltaic or coastal sands. Nevertheless, Pleistocene reef limestones in southwestern Taiwan are developed in association with thick claystones that were deposited in a deeper-water environment. These reef limestones are characterized by: (1) rapid transition from underlying claystones upward to reefal limestones, (2) lateral interfingering with open-shelf claystones, (3) being overlain by terrestrial deposits or exposed with no covering strata, and (4) being located in close association with anticlines. The authors propose that these reef limestones developed on anticlinal ridges raised above the adjacent sea floor by thrust-front migration in a foreland setting. Accepted: 21 April 1998