Winfried Werner

Microbial crusts of the late jurassic: Composition, palaeoecological significance and importance in reef construction

SummaryUpper Jurassic reefs contain variable amounts of calcareous microbial crusts. In examples from Portugal, Spain and southern Germany they occur within coral biostromes and bioherms, mixed coral-siliceous sponge reefs, siliceous sponge meadows and mudmounds, and build up thrombolities with or without additional reef metazoans. The crusts are of paramount importance for the establishment and development of positive buildups. Commonly, reef growth starts with crusts which develop from a narrow base and rapidly expand laterally by rising above the sea floor. Reef associations with little or no microbial crust normally did not develop distinct relief.The basic microbial crust type is characterised by a dense to peloidal, mostly clotted, hence thrombolitic fabric which developed due to calcification triggered by microbes. Morphological evidence for this organic nature are positive relief, bridge-structures, and the shape and arrangement of peloids. The basic thrombolitic crust type is a eurytopic feature, equally occurring in settings of different bathymetry, waterenergy, salinity and oxygen/nutrient concentrations. However, the crusts also comprise additional micro-encrusters of variable abundance and diversity. The concurrent occurrence of these encrusters and diversity trends allows discrimination between crusts of different environments, particularly of different water depths. Microbial crusts from non-reefal marine oncoids show both similarities and differences with reefal crusts. For some of the mostly enigmatic micro-encrusters new clues to their nature could be detected. For instance, bubble-like structures, formerly interpreted as sporangia inLithocodium could be identified as the foraminiferBullopora aff.laevis, possibly living as a parasite or symbiont in theLithocodium algal tissue.Lithocodium andBacinella are regarded as different organisms.‘Tubiphytes’ morronensis clearly represents a symbiotic intergrowth between a nubeculinellid foraminifer and a microbe of unknown nature.The main prerequisite for the occurrence of microbial crusts is a cessation of background sedimentation which commonly can be tied to rises in sea level. This results in the development of crust-rich reefs. Fluctuations in oxygen and nutrient levels are indicated by dysaerobic bivalves and richness in authigenic glauconite, and led to the microbes outcompeting reefal metazoans, and to the development of thrombolites. Such thrombolites occur at very different depths which is interpreted to be related to a rise of dysaerobic waters due to climatic buffering and lowering of oceanic circulation during sea level rises.Microbial crusts in modern reefs are largely restricted to shaded, cryptic settings which contrasts with the wide distribution of crusts in Upper Jurassic reefs. Microbial crusts were increasingly replaced by coralline red algae since the Late Mesozoic, but despite their restricted modern habitat seem to still play an important, commonly overlooked role in the stabilisation of reef framework.

The origin of Jurassic reefs: Current research developments and results

SummaryIn 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.

Microbialite morphology, structure and growth: a model of the Upper Jurassic reefs of the Chay Peninsula (Western France)

Abstract During the Early Kimmeridgian, the northern margin of the Aquitaine Basin (Western France) is characterised by a significant development of coral reefs. The reef formation of the Chay Peninsula comprises two main reefal units, in which the microbial structures can contribute up to 70% of framework. The microbial crusts, which played an important role in the stabilisation and growth of the reef body, show the characteristic clotted aspect of thrombolitic microbialites. Corals are the main skeletal components of the build-ups. The bioconstructions of the Chay area are thus classified as coral-thrombolite reefs. Four main morpho-structural types of microbial crusts are distinguished: (1) pseudostalactitic microbialites on the roof of intra-reef palaeocaves; (2) mamillated microbialites, found either on the undersides or on the flanks of the bioherms; (3) reticular microbialites in marginal parts of the reefs and between adjacent bioconstructed units; and (4) interstitial microbialites in voids of bioclastic deposits. Thrombolitic crusts developed on various substrates such as corals, bivalves, or bioclasts. The thrombolites formed a dense, clotted and/or micropeloidal microbial framework, in which macro- and micro-encrusters also occur. Variations in accumulation rate strongly influenced the reef morphology, in particular its relief above the sediment surface. The coalescence of the coral-microbialite patches created numerous intra-reef cavities of metre-scale dimensions. The direction of microbial growth, which defined the macroscopic microbialite forms, strongly depended on the position within the reef framework but was also controlled by water energy, accumulation rate and light availability.

Palaeoecology of coralline sponge-coral meadows from the upper jurassic of Portugal

KurzfassungDieComophyllia polymorpha-Crispispongia cf.expansa Assoziation charakterisiert eine 5–10 m mächtige Lage bioklastreicher Mergel in der Alcobaça-Formation (Kimmendge) der Umgebung von Alcobaça (Estremadura, Portugal). Dominierende Elemente der Assoziation sind Korallen, Kalkschwämme (vorwiegend Vertreter der pharetroniden Calcarea) und Algen-/Cyanobakterienkrusten, gefolgt von Crinoiden, Muscheln und Brachiopoden. Sie lassen sich in Gerüstbildner, das Gerüst inkrustierende und bindende Organismen, Riffbewohner und Riffzerstörer unterteilen. Die Organismen bildeten Riffrasen im vollmarinen Flachwasser. Sedimentationsraten waren gering, das Energieniveau gering bis mäßig, abgesehen von seltenen Hochenergie-Ereignissen. Coralline Schwämme spielen, im Gegensatz zu dem hier diskutierten Vorkommen, in anderen jurassischen Riffen nur eine sehr untergeordnete Rolle; ihr gehäuftes Auftreten war offensichtlich an sehr flaches Wasser gebunden.AbstractTheComophyllia polymorpha-Crispispongia cf.expansa association of the Kimmeridgian Alcobaça Formation occurs in a 5–10 m thick unit that can be followed for at least 10 km in the vicinity of Alcobaça (Estremadura). Corals, coralline sponges (mainly Calcarea), cryptalgal crusts and, to a lesser extent, crinoids are the dominant constituents of the autochthonous community relic which can be grouped in framebuilders, framework encrusters, frame binders, reef-dwellers, and reef destroyers. These organisms formed low meadows in a shallow, fully marine environment subject to low rates of sedimentation and moderate to low energy levels punctuated by rare high energy events. The abundance of coralline sponges in reefs and reef-like communities is uncommon in the Jurassic and appears to be restricted to very shallow water environments.

Ecophenotypic Plasticity Versus Evolutionary Trends—Morphological Variability in Upper Jurassic Bivalve Shells from Portugal

Upper Jurassic marginal marine strata of the Lusitanian Basin (central Portugal) yield a rich benthic macro fauna from which three bivalve target taxa, i.e., Arcomytilus, Isognomon, and Eomiodon, were chosen for morphometric studies, because of their abundance both in space and time and their variability in shell shape. The shells have been analysed with regard to outline shape (Fourier shape analysis), dimensions, ornamentation (Arcumytilus) and ligament arrangement (Isognomon). Additionally, data on co-occurring fauna and palaeotemperatures calculated from &dgr;18O values have been recorded. The results of the morphometric analyses have been interpreted with regard to phytogeny and palaeoecology. In all target taxa, a distinct, rapid size increase at around the Early/Late Kimmeridgian boundary is evident. Potential causes for this process are discussed, and an increase in food availability is regarded the most likely scenario. In Isognomon rugosus, a distinct change in resilifer arrangement co-occurs with size increase, resulting in the evolution of an endemic species in the Lusitanian Basin, for which the name Isognomon lusitanicus is re-established. Like in several extant Mytilidae, morphological species characterisation in Arcomytilus turns out unsatisfactory, due to high intra-specific variability. However, Arcomytilus morrisii is still regarded as a valid species that evolved in the Lusitanian Basin. Despite high shape variability, Eomiodon securiformis is also considered to be a clearly distinguished species. For all target taxa morphologic variability is discussed with regard to environment, and variation between populations is delineated. The data suggest a weak correlation of facies and shell shape in Arcomytilus, while Isognomon lusitanicus seems to develop local varieties in different subbasins. Finally, the great morphologic plasticity of bivalves from rather distinct systematic entities is shown to result from different causes, thus demonstrating that careful studies of the involved species are a prerequisite to draw correct palaeoecological conclusions.

Reconstruction of palaeosalinity using carbon isotopes and benthic associations: a comparison

Carbon and oxygen isotopes were determined on 40 recrystallized shells of Late Jurassic bivalves from the Lusitanian Basin of Portugal. In contrast with the oxygen isotopes, which exhibited considerable diagenetic distortion, the carbon isotopes are thought to preserve a record of the salinity of the Jurassic marginal marine seas in which these bivalves lived. The reconstructed palaeosalinities range from 35%o (euhaline) to 5% (oligohaline). Comparing these values with the palaeosalinity reconstructed from a palaeoecological analysis of 17 stratigraphic levels within the basin, the independently derived values agree in most cases. Strongly differing values are explained as being due to biotic factors and to diagenetic distortion of the isotopic signal; they are less likely to be due to smallscale time-averaging or insufficient microstratigraphic sampling. On the whole, the carbon isotope analyses are thought to produce reasonable palaeosalinity values, although data from infaunal, originally aragonitic bivalves appear to be less reliable than those from epifaunal bivalves with a predominantly or exclusively calcitic shell. As diagenetic alteration of the carbon isotope signal is, however, unpredictable and biotic effects on the isotopic composition are insufficiently known, palaeosalinity reconstructions based on stable isotope data should be supported by palaeoecological data.

Salinity zonation of benthic associations in the Upper Jurassic of the Lusitanian Basin (Portugal)

Abstract Within the Upper Jurassic of the Lusitanian Basin, over 40 benthic associations and autochthonous assemblages can be recognized. Most of them inhabited marginal marine environments subject to a varying degree of environmental stress. The influence of environmental factors such as substrate and energy level on the distribution pattern is discussed. Most associations were subject to reductions in salinity which reflects the regressive nature of the sedimentary sequence. They can be arranged along salinity gradients ranging from fully marine to freshwater conditions.

A parautochthonous shallow marine fauna from the Late Burdigalian (early Ottnangian) of Gurlarn (Lower Bavaria, SE Germany): Macrofaunal inventory and paleoecology

This paper describes and illustrates a diverse parautochthonous macrofauna from a single Upper Burdigalian (lower Ottnangian) horizon at Gurlarn in Lower Bavaria (SE Germany). In total, 80 different taxa are recorded in specific or open nomenclature; some 50 % of these taxa are bryozoans, followed by bivalves (16 taxa), cirripedes (7 taxa), echinoderms, corals (5 taxa each), brachiopods, fish (4 taxa each), serpulids, and gastropods (3 taxa each). The presence of additional organisms was documented by actualistic comparison based on indirect evidence such as drillholes, bite marks, and specialized growth forms of bryozoan colonies. Analysis of autand synecological indicators suggests that the fauna thrived in a near-shore shallow marine setting at a water depth of 5-20 m. Based on particular faunal elements and overall faunal composition we hypothesize that the environment is characterized by three distinct but interfingering habitats, i.e. (1) rocky slopes and boulders, (2) seagrass meadows, and (3) bryozoan meadows. Because similar, albeit less wellpreserved, faunas occur at several localities along the northern coast of the early Ottnangian Molasse Sea, the parautochthonous assemblage from Gurlarn provides an excellent example for the structure of these typical biota.

Taxonomy and ecology of Juranomia calcibyssata gen. et sp. nov.-Awidespread anomiid bivalve from the Upper Jurassic of Portugal

Abstract Juranomia calcibyssata gen. et sp. nov. is a typical anomiidbivalve characterized by a byssal notch and crus supporting the ligament in the lower, right valve and by three interconnected muscle scars in the upper, left valve. The shells are fixed to the substrate by means of a calcified byssus. The species is abundant in marginal marine (brackish) environments of the Lusitanian Basin of Portugal during Kimmeridgian times.

A sequence analyzed from the basin to the platform : the Middle Oxfordian calcareous succession in southeastern France

Middle Oxfordian sedimentation is very homogeneous in southeastern France. It is characterized by a specific alternation of marls and fine-grained limestones (G. transversarium Zone). This work shows that sets of calcareous beds allow accurate stratigraphic correlations in various paleogeographic areas, from the shallow Jura platform to the deep Dauphinois basin. Following a rifting period, this sedimentation illustrates a doming period with decreasing subsidence and water depth. This corresponds to the establishment of a wide marine area more favourable to carbonate sedimentation. Considering sequence stratigraphy, some slight variations occur in the sediment record and lead to interesting comparisons between the platform and the basin. The geochemical evolution, mainly the manganese content, seems to be dependant of the activity of the Ligurian Tethys oceanic ridge. This fact shows that sedimentation in southeastern France could be significantly controlled by tectono-eustasy.