Patch reef development in the florigemma-Bank Member (Oxfordian) from the Deister Mts (NW Germany): a type example for Late Jurassic coral thrombolite thickets

Small reefal bioconstructions that developed in lagoonal settings are widespread in a few horizons of the Late Jurassic (Oxfordian) succession of the Korallenoolith Formation, exposed southwest of Hannover, Northwest Germany. Especially the florigemma-Bank Member, “sandwiched” between oolite shoal deposits, exposes a high variety of build-ups, ranging from coral thrombolite patch reefs, to biostromes and to coral meadows. The reefs show a distribution with gradual facies variations along an outcrop belt that extends about 30 km from the Wesergebirge in the NW to the Osterwald Mts in the SE.The patch reefs from the Deister Mts locality at the “Speckhals” are developed as coral-chaetetid-solenoporid-microbialite reefs and represent a reef type that was hitherto unknown so far north of its Tethyan counterparts. They are mainly built up by coral thickets that are preserved in situ up to 1.5 m in height and a few metres in diameter. They contain up to 20 coral species of different morphotypes but are chiefly composed of phaceloid Stylosmilia corallina and Goniocora socialis subordinately. The tightly branched Stylosmilia colonies are stabilized by their anastomosing growth. The coral branches are coated with microbial crusts and micro-encrusters reinforcing the coral framework. Encrusters and other biota within the thicket show a typical community replacement sequence: Lithocodium aggregatum, Koskinobullina socialis and Iberopora bodeuri are pioneer organisms, whereas the occurrence of non-rigid sponges represents the terminal growth stage. The latter are preserved in situ and seem to be characteristic so far poorly known constituents of the Late Jurassic cryptobiont reef dweller community. The distance and overall arrangement of branches seems to be the crucial factor for the manifestation of a (cryptic) habitat promoting such community replacement sequences. Widely spaced branches often lack any encrusting and/or other reef dwelling organisms, whereas tightly branched corals, as is St. corallina, stimulate such biota. Hence, such reefs are well suited for research on coelobites and community sequences of encrusting and cavity dwelling organisms.     

Late Moscovian (mid-Pennsylvanian) rugose corals from Wadi Araba (Egypt,Eastern Desert): Taxonomy,palaeoecology and palaeobiogeography

An important late Moscovian rugose coral association is described from the Rod El Hamal Formation of the Wadi Araba area (northern Eastern Desert, Egypt). The upper part of the formation yielded a moderately abundant but poorly diversified coral fauna composed of large dissepimented solitary rugose corals. In total, six species belong to the Bothrophyllidae and the Geyerophyllidae, including three new species: Bothrophyllum suezensis, Bothrophyllum cylindricum, and Amygdalophylloides omarai. Three undeterminable Bothrophyllidae taxa are left in open nomenclature. The corals were attached to the soft substrate by talons and rootlets, either rooted and growing upward as mud-stickers or as secondary recliners. They show frequent rejuvenations and constrictions. Encrustation and/or bioerosion are scarce. Carbonate microfacies indicate a habitat in littoral and inner neritic zones adjacent to a low terrigenous hinterland. Colonial taxa are missing and tabulate corals are rare. The corals from the Rod El Hamal Formation are the only example of a late Moscovian coral fauna on the southern margin of the Palaeotethys, that is to say along the northern margin of Gondwana. The association shows similarities with northwestern Spain (Cantabrian Mountains), the Donets and Moscow basins, and thus a general attribution to the Palaeotethys realm.     

Calcareous algae (“Porostromata”, rhodophyta dasycladales) and microproblem-atica with algal affinity from the NW German Korallenoolith Formation (Oxfordian,Süntel Mountains)

Summary In the florigemma-Bank Member of the Late Jurassic (Oxfordian) NW German Korallenoolith Formation moderately diverse assemblages of calcareous algae are present. The following taxa have been identified: “Porostromata”, Solenopora jurassica Brown, Solenopora helvetica Peterhans, Pycnoporidium lobatum Yabe and Toyama, Otternstella cf. lemmensis (Bernier), Salpingoporella? suentelensis n. sp., Salpingoporella annulata Carozzi, Rajkaella cf. iailaensis (Maslov), Anisoporella? cretacea (Dragastan), Terquemella div. spec., Iberopora bodeuri Granier and Berthou, Bacinella irregularis Radoicic and Koskinobullina socialis Cherchi and Schroeder. Except for the well known but rarely illustrated Goniolina geometrica (Roemer), that is ubiquitous in the Korallenoolith succession, all specimens were recovered during the last years in the course of the investigations concerning the reefal bioconstructions of the Korallenoolith Formation. Most of the taxa described here are reported for the first time from the Lower Saxony Basin. The sedimentary succession of the florigemma-Bank Member exposes up to eight different lithofacies types that are mainly characterized by different algal associations. The lithofacies types are described and discussed concerning their depositional environments and their floral content. The occurrence of floral elements, that are generally restricted to tropical/subtropical environments by temperature and solar radiation in palaeolatitudes as far north as the Lower Saxony Basin, has not yet been observed and adds further detail to a presumed shift towards subtropical climates in the Middle Oxfordian of this area. Similar algal associations at comparable palaeolatitudes have not been reported from the European Oxfordian (e.g. “Corallian”, southern England) yet.     

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.     

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

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

Danian cold‐water corals from the Baunekule facies,Faxe Formation,Denmark: a rare taphonomic window of a coral mound flank habitat

Well‐preserved cold‐water corals are comparatively rare in the fossil record. This is partly due to the very low fossilization potential of the predominantly aragonitic corals but also due to the fact that coral ecosystems of deep water are a geologically young development. A Middle Danian cold‐water coral mound complex is well exposed in Faxe Quarry, Denmark. The coral mounds are intercalated with bryozoan mounds of various sizes and form the Faxe Formation. The coral limestone displays large variations in diagenesis, and this complicates the palaeoecological reconstructions. However, the Baunekule facies from the Faxe Formation contain a well‐preserved originally aragonitic and calcitic fauna. The aragonitic skeletons have been recrystallized to calcite during early diagenesis and the excellent preservation makes taxonomic identifications straightforward. A diverse fauna of ten scleractinian coral species, nine stylasterine coral species and seven octocoral species has been described from the Baunekule facies. The fossil fauna represents an ecological niche between the dead coral framework and coral rubble on a flank of a growing Dendrophyllia coral mound with multiple colonization events. The diversity and relative abundance of the fossil scleractinian corals are comparable to the modern settings in the NE Atlantic and Mediterranean. The distribution and diversity of the octocorals and the stylasterine corals are suggested to represent coral gardens as described from modern setting in the NE Pacific. The presence of a diverse and abundant stylasterine fauna suggests a stable palaeoenvironment, probably in a bathymetric depth range of 200–400 metre.     

Distribution and Abundance of Fauna on Living Tissues of Two Brazilian Hermatypic Corals (Mussismilia hispida (Verril 1902) and Siderastrea stellata Verril, 1868)

This study describes the distribution and abundance patterns of the associate fauna on the living surface of the corals Siderastrea stellata Verril, 1868 and Mussismilia hispida (Verril 1902) using a non-destructive method, on the northern coast of Rio de Janeiro State. For each coral species, infestation density and proportions of infested colonies, colonies attached and unattached to the substrate were estimated. A total of 474 colonies of S. stellata and 452 colonies of M. hispida were examined. The barnacle Ceratoconcha floridana (Pilsbry, 1931) was the dominant coral associate found, followed by gall-crabs of the family Cryptochiridae Paulson, 1875 and the bivalve Lithophaga bisulcata (d’Orbigny, 1842). Both coral species presented similar patterns of infestation dominance. S. stellata colonies were more commonly infested and showed a greater mean infestation density of 0.62 ind/cm² at Armação dos Búzios, whereas M. hispida colonies had infestation densities of only 0.20 ind/cm². Infestation density does not appear to impact negatively on corals of Armação dos Búzios. A clear negative relationship between the number of associates in the coral colony and coral size was found. Evidently abundance and frequency of occurrence of associated fauna is highly related to coral community structure and composition and the results highlight the importance of local scale studies.     

Lower Jurassic corals from the Pamir Mountains,Central Asia

The paper describes new Lower Jurassic corals from the South-Eastern Pamir Mountains (Tajikistan) and interprets their relationships with contemporaneous West Tethyan corals. Taxonomic similarities with Pliensbachian European and North African faunas indicate a Pliensbachian age for this fauna, which was previously considered to be of Hettangian/Sinemurian age. Together with the taxa earlier described from the Pamirs, this fauna consists of 30 species of 25 genera, including Triassic holdover genera such as Stylophyllopsis, Phacelostylophyllum and Eocomoseris. The bulk of the fauna represents new Jurassic genera: Alichurastrea, Eomicrophyllia, Guembelastreomorpha, Gurumdynia, Pinacomorpha, Protostephanastrea, Psenophyllia, Sedekastrea and Stylimorpha. Earlier coral studies of the region concerned the genera: Archaeosmilia Melnikova, 1975, Archaeosmiliopsis Melnikova, 1975, Cylismilia Roniewicz, 1988, Pachysmilia Melnikova, 1989, and Prodonacosmilia Melnikova in Melnikova and Roniewicz, 1976. Two species that were considered to belong to the genus Cylismilia, are redescribed and reclassified in the genera Psenophyllia, and Archaeosmilia Melnikova, respectively.     

新疆准噶尔盆地周边晚石炭世的珊瑚*

新疆北部准噶尔盆地周边石炭系广泛出露, 其中上石炭统发育海相火山—沉积序列, 包括西准噶尔地区的哈拉阿拉特组、阿腊德依克赛组, 东准噶尔地区的石钱滩组以及盆地南部的祁家沟组、奥尔吐组等。本文研究在上述各地层的灰岩层或夹层、碎屑岩中获得的四射珊瑚9属17种和床板珊瑚2属3种, 对石钱滩组中的3个新种进行了详细的系统古生物学描述。上述珊瑚化石的地质时代主要为巴什基尔期至莫斯科期。在珊瑚动物群的组成上, 哈拉阿拉特组中新发现Caninophyllum属分子, 结合盆地东、西部共同产出的床板珊瑚Cystodendropora 属分子, 说明整个准噶尔盆地周边的动物群面貌大致类似, 而西准噶尔地区的化石丰度、分异度显著低于盆地东部和东南部。综合前人资料, 目前在准噶尔盆地周边发现的珊瑚化石有约一半为地方性分子。本文认为研究区珊瑚化石的分布和组成特征与晚石炭世准噶尔洋走向封闭或半封闭的演化趋势密切相关。     

河北唐山西域山晚寒武世长山组的树形笔石

本文对东秦岭南部湖北郧西地区早石炭世四射珊瑚进行了研究,系统描述了11科中的24属和39种,其中包含2新属和21新种。本区早石炭世四射珊瑚拟划分为4个组合带:范家坪组上部Palaeosmilia-Lithostrotion组合带,下部Yuanophyllum组合带。袁家沟组上部Shennongia majus组合带,下部Zaphrentoides组合带。范家坪组是新建的组名。文中也讨论了Cystophrentis的分布时限,及其与Shennongia的亲缘关系。此外,Neoclisiophyllum在本区与岩关期的Shennongia共生,出现的时间早于华南各地。     

Rugose corals from the upper Viséan (Carboniferous) of the Jerada Massif (NE Morocco): taxonomy,biostratigraphy, facies and palaeobiogeography

The Viséan succession of the Jerada Massif contains a relatively diverse rugose coral fauna, which comprises 12 genera. Only two of these (Siphonodendron and Lithostrotion) are reported to include more than one species. Most taxa show distinctive facies dependencies. Small solitary corals are found in shaly environments of the Oued Es-Sassi Formation. On the northern flank a coral patch reef formed on an oolitic shoal in the Koudiat Es-Senn Formation. Its core mainly consists of Lithostrotion vorticale colonies, whereas in its surroundings literally hundreds of specimens of Siphonophyllia samsonensis occur. Associated with these dominant taxa occur colonial and further large solitary taxa (Palaeosmilia, Dibunophyllum). Tizraia and Pareynia are restricted to microbial-dominated buildup facies. Their presence might be strongly controlled by the development of this buildup type, because further occurrences in Algeria, Morocco, and Belgium are all in the same facies. The coral fauna at Jerada is a typical Late Viséan assemblage for the Western European Province. The Eastern Moroccan Meseta may be an important pathway within the province for the connection between the Central Saharan basins and NW Europe. The biostratigraphic ages of the coral fauna partly contradict ages based on carbonate microfossils; the coral ages are slightly older and typical Brigantian coral taxa are absent.     

Importance of live coral habitat for reef fishes

Live corals are the key habitat forming organisms on coral reefs, contributing to both biological and physical structure. Understanding the importance of corals for reef fishes is, however, restricted to a few key families of fishes, whereas it is likely that a vast number of fish species will be adversely affected by the loss of live corals. This study used data from published literature together with independent field based surveys to quantify the range of reef fish species that use live coral habitats. A total of 320 species from 39 families use live coral habitats, accounting for approximately 8 % of all reef fishes. Many of the fishes reported to use live corals are from the families Pomacentridae (68 spp.) and Gobiidae (44 spp.) and most (66 %) are either planktivores or omnivores. 126 species of fish associate with corals as juveniles, although many of these fishes have no apparent affiliation with coral as adults, suggesting an ontogenetic shift in coral reliance. Collectively, reef fishes have been reported to use at least 93 species of coral, mainly from the genus Acropora and Porities and associate predominantly with branching growth forms. Some fish associate with a single coral species, whilst others can be found on more than 20 different species of coral indicating there is considerable variation in habitat specialisation among coral associated fish species. The large number of fishes that rely on coral highlights that habitat degradation and coral loss will have significant consequences for biodiversity and productivity of reef fish assemblages.     

Effects of the fish anesthetic, clove oil (eugenol), on coral health and growth

Ecological research within coral reefs often requires the use of anesthetics to immobilize organisms. It is therefore important to consider the effect of these chemicals on the surrounding flora and fauna, particularly to the corals themselves. We quantified the effects of clove oil, a commonly used fish anesthetic, on the growth and occurrence of bleaching in three species of corals: Acropora striata, Pocillopora verrucosa, and Porites australiensis. We compared coral responses to five treatments: a gradient of four clove oil concentrations (0-28%) in seawater, and one concentration of clove oil (14%) in ethanol. Each week, we assessed the presence of bleaching, and then applied the treatment. We measured growth over the duration of the 6-week experiment using the buoyant weight technique. Growth and bleaching showed a dose response to clove oil exposure, and the use of ethanol as a solvent had an additional deleterious effect, as also suggested by observed changes in concentrations of eugenol following field application. Overall, growth was reduced by 37.6% at the highest concentration (28% clove oil in seawater) relative to the control (0% clove oil). The reduction in growth was nearly as great (35.3% of the control) at half the concentration of clove oil (14%) when dissolved in ethanol. These results suggest the repeated use of clove oil (even without a solvent) can deleteriously affect corals.     

Dimethylsulfoniopropionate,superoxide dismutase and glutathione as stress response indicators in three corals under short-term hyposalinity stress

Corals are among the most active producers of dimethylsulfoniopropionate (DMSP), a key molecule in marine sulfur cycling, yet the specific physiological role of DMSP in corals remains elusive. Here, we examine the oxidative stress response of three coral species (Acropora millepora, Stylophora pistillata and Pocillopora damicornis) and explore the antioxidant role of DMSP and its breakdown products under short-term hyposalinity stress. Symbiont photosynthetic activity declined with hyposalinity exposure in all three reef-building corals. This corresponded with the upregulation of superoxide dismutase and glutathione in the animal host of all three species. For the symbiont component, there were differences in antioxidant regulation, demonstrating differential responses to oxidative stress between the Symbiodinium subclades. Of the three coral species investigated, only A. millepora provided any evidence of the role of DMSP in the oxidative stress response. Our study reveals variability in antioxidant regulation in corals and highlights the influence life-history traits, and the subcladal differences can have on coral physiology. Our data expand on the emerging understanding of the role of DMSP in coral stress regulation and emphasizes the importance of exploring both the host and symbiont responses for defining the threshold of the coral holobiont to hyposalinity stress.     

New data about deep-water solitary ahermatypic scleractinians of the shelf of Vietnam

This paper reports data on deep-water solitary ahermatypic corals that were collected on the shelf of Vietnam. Among these, 28 species belonging to 16 genera and eight families were identified; 9 species are first records for the South China Sea, and 2 species (Deltocyathus radiatus sp. n. and Flabellum septodentatus sp. n.) are described as new to science. Dendrophyllia sp. (33 indiv. in sample) and Truncatoflabellum crassum (28 indiv. in sample) formed massive aggregations. The taxonomically richest genera were Flabellum, Truncatoflabellum, and Caryophyllia. Coral density and species diversity were highest at depths of 90–230 and 285–390 m, respectively. Deep-water corals of Vietnam are integral with the coral fauna of the Indo-Polynesian Province. The data we obtained contribute to the knowledge of the composition and chorology of the worldwide fauna of solitary corals.     

新疆准噶尔盆地周边晚石炭世的珊瑚*

新疆北部准噶尔盆地周边石炭系广泛出露, 其中上石炭统发育海相火山—沉积序列, 包括西准噶尔地区的哈拉阿拉特组、阿腊德依克赛组, 东准噶尔地区的石钱滩组以及盆地南部的祁家沟组、奥尔吐组等。本文研究在上述各地层的灰岩层或夹层、碎屑岩中获得的四射珊瑚9属17种和床板珊瑚2属3种, 对石钱滩组中的3个新种进行了详细的系统古生物学描述。上述珊瑚化石的地质时代主要为巴什基尔期至莫斯科期。在珊瑚动物群的组成上, 哈拉阿拉特组中新发现Caninophyllum属分子, 结合盆地东、西部共同产出的床板珊瑚Cystodendropora 属分子, 说明整个准噶尔盆地周边的动物群面貌大致类似, 而西准噶尔地区的化石丰度、分异度显著低于盆地东部和东南部。综合前人资料, 目前在准噶尔盆地周边发现的珊瑚化石有约一半为地方性分子。本文认为研究区珊瑚化石的分布和组成特征与晚石炭世准噶尔洋走向封闭或半封闭的演化趋势密切相关。     

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.     

Dietary shift in corallivorous Drupella snails following a major bleaching event at Koh Tao, Gulf of Thailand

The island Koh Tao in the western Gulf of Thailand suffered severe coral bleaching in 2010. Its mushroom coral fauna of 20 species was surveyed during the bleaching in 2010 and after the bleaching in 2011. Multi-species assemblages of free-living mushroom corals occurred around the island, two of which were invaded by corallivorous Drupella snails after the bleaching. Previously these gastropods were known to mainly consume branching corals and hardly any mushroom corals. The snails were found preying on four fungiid species, three of which were susceptible to bleaching. The dietary shift became apparent after populations of preferred prey species (Acroporidae and Pocilloporidae) had died during the bleaching event. It seems that bleaching mortality reduced the availability of preferred prey, causing the corallivores to switch to less preferred species that occur in dense aggregations.     

Growth Dynamics of the Threatened Caribbean Staghorn Coral Acropora cervicornis: Influence of Host Genotype,Symbiont Identity,Colony Size,and Environmental Setting

Background

The drastic decline in the abundance of Caribbean acroporid corals (Acropora cervicornis, A. palmata) has prompted the listing of this genus as threatened as well as the development of a regional propagation and restoration program. Using in situ underwater nurseries, we documented the influence of coral genotype and symbiont identity, colony size, and propagation method on the growth and branching patterns of staghorn corals in Florida and the Dominican Republic.

Methodology/Principal Findings

Individual tracking of> 1700 nursery-grown staghorn fragments and colonies from 37 distinct genotypes (identified using microsatellites) in Florida and the Dominican Republic revealed a significant positive relationship between size and growth, but a decreasing rate of productivity with increasing size. Pruning vigor (enhanced growth after fragmentation) was documented even in colonies that lost 95% of their coral tissue/skeleton, indicating that high productivity can be maintained within nurseries by sequentially fragmenting corals. A significant effect of coral genotype was documented for corals grown in a common-garden setting, with fast-growing genotypes growing up to an order of magnitude faster than slow-growing genotypes. Algal-symbiont identity established using qPCR techniques showed that clade A (likely Symbiodinium A3) was the dominant symbiont type for all coral genotypes, except for one coral genotype in the DR and two in Florida that were dominated by clade C, with A- and C-dominated genotypes having similar growth rates.

Conclusion/Significance

The threatened Caribbean staghorn coral is capable of extremely fast growth, with annual productivity rates exceeding 5 cm of new coral produced for every cm of existing coral. This species benefits from high fragment survivorship coupled by the pruning vigor experienced by the parent colonies after fragmentation. These life-history characteristics make A. cervicornis a successful candidate nursery species and provide optimism for the potential role that active propagation can play in the recovery of this keystone species.     

ökologie und Taxonomie koralleninkrustierender Bryozoen des norddeutschen Malm

In the Late Oxfordian Korallenoolith of northern Germany, eight bryozoan taxa encrusting corals have been found, six of which had not been previously recorded in the area. In most cases, they can only be described using open nomenclature since the gonozooids are not known and the type material of existing species is too poorly preserved. “Stationary” bereniciform bryozoans were relatively good competitors adapted to a predictable environment. For this reason they thrived in the calm-water facies. However, the environment was disturbed frequently as proved by the joint occurrence with Stomatoporidae. These “mobile” taxa could grow fast on strongly three-dimensional surfaces, thus being able to survive even in highly agitated facies by escaping into cryptic habitats. The distribution of individual taxa is thus governed by the type and frequency of perturbations of their habitats. Bryozoans had a binding function in the fauna related to corals. Because of their low number and poor coverage, they mostly had little importance, however. This is due to the high water energy or high rate of sedimentation, which characterized the environments of the sampled localities. Thus, ideal habitats were not available. Nevertheless, the characteristic bryozoan associations of the European Jurassic became established together with typical associated faunal elements.