Diversity,speciation, endemism and extinction in Devonian reef and level-bottom communities,eastern North America

Lower Devonian late Emsian (Bois Blanc and Clear Creek Limestones; Schoharie Formation) level-bottom communities in New York, Michigan and Illinois were moderately cosmopolitan and diverse and dominated by brachiopods and solitary rugose corals. Subsequently (Early Eifelian?), there was an important episode of cratonal patch reef building in New York (Edgecliff Member, Onondaga Limestone), southwestern Ontario (Formosa Reef Limestone, lower Detroit River Group), and the Hudson Bay Lowland (Kwataboahegan Formation) by highly diverse endemic communities. The Edgecliff reefs were built by corals whereas the Formosa and Kwataboahegan reefs were built primarily by stromatoporoids. The strong correlation between high diversity and high endemism during the reef-building episode suggests that these communities contained numberous, small species populations belonging to several major taxa — an example of rapid speciation by geographic isolation and genetic drift.     

Ordovician reefs of the Ural Mountains,Russia: A review

Summary The Upper Ordovician reefs of the Urals were formed at a subsiding shelf-margin during an early Late Ashgillian (Sur’ya time interval) regressive phase. Reefs of this age were studied in detail from the western slope of the Northern, the Subpolar and the Polar Urals with respect to lithofacies, biotic composition and paleogeographical patterns. The thickness of the reefs varies between 100 and 500 m. The backreef areas are characterized by lagoons with increased salinity and sabkha development. Microbial associations and a diverse algal flora (Cyanophyta, green and red algae and alga incertae sedis) are the main constituents of reefal boundstones. Tabulate and rugose corals, heliolitids, calcareous sponge-like fossils, bryozoans and problematic hydroids were also part of the reef communities. Each reef exhibits a characteristic framework-building association. Reef development was terminated by a rapid and abrupt sea-level rise at the end of the middle Upper Ashgillian connected with the global Late Ordovician glaciation.     

Post‐tectonic limitations on Early Devonian (Emsian) reef development in the Gobi‐Altai region,Mongolia

Pellegrini, A.F.A., Soja, C.M. & Minjin, C. 2011: Post‐tectonic limitations on Early Devonian (Emsian) reef development in the Gobi‐Altai region, Mongolia. Lethaia, Vol. 45, pp. 46–61. This study investigates a Lower Devonian (Emsian) carbonate sequence from the Chuluun Formation where it is exposed in the Gobi‐Altai region of southern Mongolia. Quantification of abundance patterns across guild, morphotype and general taxonomic levels was based on stratigraphical and thin‐section analyses. Comparison with other Emsian carbonate platforms allowed the factors that influenced community development in the wake of a tectonic perturbation to be determined. Our evidence reveals that potential reef‐building biotas preserved in the Chuluun Formation experienced rapid colonization of a newly submerged carbonate platform following an episode of tectonic uplift and the development of a coastal alluvial fan. Although critical reef‐building organisms were present, colonial corals and stromatoporoids exhibited limited vertical growth and showed no significant lateral expansion of individuals or biotic assemblages. Nor did those taxa experience significant increases in abundance, density, or size. We conclude that incomplete succession and the lack of reef development occurred most likely because of an unsuitable substrate, limited accommodation space and isolation that reduced colonization potential. □Community succession, Emsian, palaeoecology, reef suppression.     

Considérations paléobiogéographiques sur les Streptelasmatina (Coelenterata,Rugosa) solitaires du Dévonien des monts Cantabriques (NW de l'Espagne)

The study of numerous fossil solitary forms ofStreptelasmatina (Coelenterata, Rugosa) and some ones belonging to different suborders from the Devonian of the Cantabrian Mountains (NW Spain), shows the importance of such organisms concerning the relations among the major faunal provinces during the Devonian time.The analysis of the Devonian Rugose coral fauna(from Emsian to Frasnian) lets to establish the paleobiogeographical relations of the Cantabrian genera. Particularly, during the Lower Devonian (Emsian) the relationships between the Eastern of North America and Spain would be closer than it was supposed until now.The evidence on the world distribution of thestudied forms suggest possible migration roads.     

A new actinolepid arthrodire (Class Placodermi) from the Lower Devonian Sevy Dolomite, East-Central Nevada

A new genus and species of actinolepid arthrodire, Aleosteus eganensis , is described from the Lower Devonian (Late Emsian) Sevy Dolomite in the Egan Range of east-central Nevada. A cladistic analysis of the family Actinolepidae is presented for the first time and shows the Baltic Actinolepis species form the primitive sister-group of all other actinolepids. Rapid evolution of the Actinolepidae during the Lochkovian is reflected in the dispersal of the family around the Old Red Sandstone Continent at this time followed by the development of endemic faunas through the Lower Devonian and into the Middle Devonian.     

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

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

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.     

Oldest scleractinian coral reefs on the North American craton: Upper Triassic (Carnian), northeastern British Columbia,Canada

Bioclastic accumulations composed of crinoids, brachiopods, molluscs, spongiomorphs and scleractinian corals occur within Upper Triassic strata of the lower Baldonnel Formation at Pardonet Hill in northeastern British Columbia Canada. These small buildups (∼100 to 500 m³) have planar bases and broadly convex tops. These mounds are interpreted as small patch reefs composed of packstone, bioclastic floatstone/rudstone and carbonate breccia intercalated with mixed siliciclastic carbonate sediments deposited in a shallow subtidal setting (i.e. above fairweather wave base). Amalgamated hummocky cross-stratified to current ripple-laminated, quartz-dominated sandstone beds and numerous sharp-based, normally graded bioclastic (commonly encrinitic) packstone/grainstone — quartz–sandstone couplets characterize inter-reef lithologies.Conodont biostratigraphy indicates that the Pardonet Hill patch reefs occur within strata dated as earliest Upper Carnian (lower nodosus zone). The Pardonet Hill patch reefs originated and developed during an interval of regional sea level lowstand. Strata within which these patch reefs occur represent the westernmost migration of the Triassic shoreline in western Canada. Disappearance of coral reefs in the study area may have been affected by rapid marine transgression and failure of reef faunas to recolonize the new shore zone further to the east.The Pardonet Hill locality occurred on the western margin of the North American craton during the Triassic. Prior to their discovery reef-like structures dominated by corals in the western Panthalassa were limited to allochthonous terranes (now part of the Cordillera). The Pardonet Hill patch reefs occur at approximately 30° Triassic paleolatitude. In modern settings, this is at the extreme latitudinal margin of subtropical zooxanthellate reef development. The presence of benthic faunas characteristic of low-paleolatitude settings on the northwestern coast of Pangea has significant implications in paleotectonic and paleoenvironmental reconstructions.     

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.     

Devonian conodont biogeography: Quantitative analysis of provinciality

Cluster analysis of conodont faunas from each of 17 Lower‐lower Upper Devonian zones and subzones (data as reported by Klapper and Johnson, 1980) reveal changing patterns of provinciality.

Provinciality, expressed by a differentiation into western Laurussian and proto‐Tethyan biogeographic regions, is moderate in the lower Lochkovian but is low or absent in the upper Lochkovian‐lower Pragian. Provincialism returns in the Pragian and reaches its maximum development during the Emsian. Most Australian faunas are distinct from those of western Laurussia and the proto‐Tethys. Conodont faunas from suspect terranes of western North America display no unusual biogeographic affinities. Provincialism declines during the Eifelian and is only weakly developed in Givetian‐lower Frasnian faunas.

Changing global sea level during the Devonian may explain the development of Devonian conodont provinciality. As proposed by Klapper and Johnson (1980), low provinciality is associated with low stands of sea level. Endemic faunas develop in isolated epeiric seas during intermediate stages of sea level rise. High stands of sea level ultimately drown barriers to faunal exchange and prompt a return to low provinciality conditions.     

新疆乌恰县萨瓦亚尔顿金矿区早泥盆世牙形刺

在新疆乌恰县萨瓦亚尔顿金矿区首次发现了早泥盆世牙形刺,确认了Ｌｏｃｈｋｏｖｉａｎ阶（Ａｎ－ｃｙｒｏｄｅｌｌｏｉｄｅｓ ｏｍｕｓ）和Ｅｍｓｉａｎ阶（ｏｌｙｇａｎａｔｈｕｓ ｋｉｔａｂｉｃｕｓ,Ｐ．ｅｘｃａｖａｔｕｓ）的存在,并阐述了此次发现的意义和存在的问题。     

Subtle genetic connectivity between Mexican Caribbean and south-western Gulf of Mexico reefs: the case of the bicolor damselfish,Stegastes partitus

Efficient reef management strategies rely on detailed knowledge of biological exchange dynamics. At present, available connectivity information on Mexican Atlantic reefs is scarce, particularly concerning the Veracruz Reef System (VRS), which is located in the south-western Gulf of Mexico. This study used a hierarchically nested sampling design to evaluate the levels of genetic connectivity both within and between the Mexican Caribbean (MC) and VRS reef regions; all of the studied reefs are marine protected areas. Microsatellites were used as genetic markers, and bicolor damselfish (Stegastes partitus) recruits were used as a biological model. The paired genetic differentiation index between regions (Fst _(ENA) = 0.008) was lower than the global index (Fst _(ENA) = 0.027), suggesting that the stronger restrictions to gene flow may be located inside the regions rather than between them. The AMOVA results supported this explanation, as the differences were only non-significant between regions. In the VRS, Santiaguillo reef was associated with low genetic connectivity levels, whilst within the MC region the group formed by Chinchorro Bank and Cozumel exhibited a restriction to gene flow with Puerto Morelos, their northernmost reef. Despite their spatial separation, reefs from different regions (Puerto Morelos and Anegada de Adentro) showed the lowest, albeit significant, genetic difference, meaning that a subtle genetic connectivity exists at the regional scale. The detected composite flow pattern is likely related to self-recruitment and cohesive dispersal processes interacting with current patterns, which may favour genetic connections under specific conditions. The results presented here suggest that coral reef management in the Mexican Atlantic Ocean should consider large scale measures in addition to appropriate local actions to protect reef fish populations.     

Microbes and mass extinctions: paleoenvironmental distribution of microbialites during times of biotic crisis

Widespread development of microbialites characterizes the substrate and ecological response during the aftermath of two of the 'big five' mass extinctions of the Phanerozoic. This study reviews the microbial response recorded by macroscopic microbial structures to these events to examine how extinction mechanism may be linked to the style of microbialite development. Two main styles of response are recognized: (i) the expansion of microbialites into environments not previously occupied during the pre-extinction interval and (ii) increases in microbialite abundance and attainment of ecological dominance within environments occupied prior to the extinction. The Late Devonian biotic crisis contributed toward the decimation of platform margin reef taxa and was followed by increases in microbialite abundance in Famennian and earliest Carboniferous platform interior, margin, and slope settings. The end-Permian event records the suppression of infaunal activity and an elimination of metazoan-dominated reefs. The aftermath of this mass extinction is characterized by the expansion of microbialites into new environments including offshore and nearshore ramp, platform interior, and slope settings. The mass extinctions at the end of the Triassic and Cretaceous have not yet been associated with a macroscopic microbial response, although one has been suggested for the end-Ordovician event. The case for microbialites behaving as 'disaster forms' in the aftermath of mass extinctions accurately describes the response following the Late Devonian and end-Permian events, and this may be because each is marked by the reduction of reef communities in addition to a suppression of bioturbation related to the development of shallow-water anoxia.     

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.     

华南中泥盆世艾菲尔期浅海相四射珊瑚的组合特征和古生物地理区系

虽然在华南中泥盆世艾菲尔期(Eifelian)较深水或斜坡相地层中产有许多国际标准的牙形类带化石,但在广阔的浅海相地层中却很难寻觅到其踪影。华南浅海相沉积地区是否缺失艾菲尔阶?在野外众多剖面的实地考察中表明,从下泥盆统埃姆斯阶到中泥盆统吉维特阶都是连续沉积,中间并没有发现地层的缺失或间断现象。浅海相的艾菲尔阶与其上覆的吉维特阶和下伏的埃姆斯阶都是连续沉积,说明可能是由于海水太浅,不太适合那些国际标准分子的生存而已。艾菲尔期中期末发生一次生物灭绝事件(Mid-Eifelian event),favositids,heliolitids和许多古老类型的珊瑚在地球上灭绝。华南艾菲尔期四射珊瑚可以划分成下、上两个完全不同的组合:1)Utara-tuiasinen sis-Sociop hyllum minor组合(牙形类partitus带—costatus带);2)Columnaria spinosa-Dendrostella praerhenana组合(牙形类australis带—kocklianus带)。     

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.     

Late Triassic reefs from the Northwest and South Tethys: distribution, setting, and biotic composition

The paleolatitudinal distribution patterns during Ladinian and Carnian time are characterized by an increasing expansion of reefs from the northern to the southern hemisphere. The optimum of reef diversity and frequency in the Norian is associated with the development of extended attached or isolated carbonate platforms. Norian-Rhaetian sponge and coral reefs of the Northern Calcareous Alps developed (1) as reef belt composed of patch reefs in platform-edge positions facing the open-marine northwestern Tethys basins and (2) as patch reefs in intraplatform basins as well as in ramp positions.Carnian and Norian-Rhaetian sponge and coral reefs of the Arabian Peninsula are formed (1) as reef complexes at the margins of carbonate platforms on the tops of volcanic seamounts in the southern Tethyan ocean, as small biostromes on these isolated platforms, and (2) as transgressive reef complexes on the attached platform of the Gondwana margin. The Norian Gosaukamm reefal breccia of the NW Tethys is a counterpart of Jabal Wasa reefal limestone of the Gondwana margin with similarities in geological setting and biotic composition. Rhaetian coral biostromes of low diversity known from the Austrian Koessen basin resemble to the time equivalent Ala biostromes of the isolated Kawr platform in the southern Neo-Tethys by forming a discontinuous layer in shallow intraplatform basin setting.     

Jurassic reef events in the Moroccan Atlas: A review

In the Moroccan Atlas, sedimentary deposits provide important data on reef events that characterize the Jurassic period. Recent work allows us to enhance knowledge of the Jurassic reefs in the Atlas, in particular their age, character and palaeogeographic distribution. Numerous localities with sponge-microbial mud mounds, coral reefs, and lithiotid bioherms are recorded from the Middle and High Atlas regions. These different biogenic constructions occupy different palaeogeographic settings: on the top of tilted blocks within the basin center; in slightly deeper positions, at the basin platform junction; and on adjacent platforms in the middle of the coastal area. The main episodes of reef building span nearly 30 million years, as follows: (1) Sinemurian, (2) early Pliensbachian, (3) late Toarcian, (4) Aalenian–early Bajocian (pars), and (5) late Bajocian. These five distinct reef events can be linked with general fluctuations of sea level and tectonism, and have palaeoclimatic implications.     

Reassessing Devonotarbus, a phalangiotarbid arachnid from the Lower Devonian (Siegenian and Emsian) of the Rheinisches Schiefergebirge (SW Germany)

Seven further specimens of the hitherto only known Devonian phalangiotarbid Devonotarbus Poschmann, Anderson and Dunlop, 2005 are described. Thus, the genus is recorded from four different localities of Siegenian (Hombach and Burglahr) and Lower Emsian (Willwerath and Alken) age, respectively, but the specific identity of Siegenian and Emsian specimens remains unproven. The new material suggests that this Devonian phalangiotarbid possessed more than six eye lenses, tergites five and six fused into a diplotergite, and an almost terminally situated anal operculum. It thus notably deviates morphologically from its younger Permocarboniferous relatives, and a new family, Devonotarbidae nov. fam., is proposed to accommodate Devonotarbus.     

Bryozoans, corals, and microfacies of Lower Eifelian (Middle Devonian) limestones at Kierspe, Germany

The Lower Eifelian Meinerzhagener Korallenkalk (= upper Cultrijugatus Beds) at Kierspe, Sauerland, contains a rich reefal fauna. Eight bryozoan species are described, two of them are new: the cystoporate Fistuliporella kierspensis n. sp. and the trepostome Leptotrypella sophiae n. sp. The bryozoans from the Meinerzhagener Korallenkalk shows distinct similarities to the Lower–Middle Devonian of Spain (Santa Lucía Formation, Emsian–Eifelian), and to the Middle Devonian (Eifelian) of Transcaucasia. The coral fauna comprises five tabulate corals and one rugose coral that document a paleobiogeographic relationship between Central and Eastern Europe and Central Asia. The associated fauna is represented by brachiopods, ostracods, and echinoderms. The studied limestones also commonly contain calcimicrobes represented by three species. The faunal and microfacial characteristics indicate a shallow marine depositional environment just above the storm wave base, with a supposed depth of 20 m, within the photic zone. The nutrient regime was at least a mesotrophic. The upper boundary of the Cultrijugatus Beds coincides with the Chote?-Event that strongly affected brachiopods, whereas corals and bryozoans were insensitive to this event.