Ecology and Biogeography of Free-Living Nematodes Associated with Chemosynthetic Environments in the Deep Sea: A Review

Background

Here, insight is provided into the present knowledge on free-living nematodes associated with chemosynthetic environments in the deep sea. It was investigated if the same trends of high standing stock, low diversity, and the dominance of a specialized fauna, as observed for macro-invertebrates, are also present in the nematodes in both vents and seeps.

Methodology

This review is based on existing literature, in combination with integrated analysis of datasets, obtained through the Census of Marine Life program on Biogeography of Deep-Water Chemosynthetic Ecosystems (ChEss).

Findings

Nematodes are often thriving in the sulphidic sediments of deep cold seeps, with standing stock values ocassionaly exceeding largely the numbers at background sites. Vents seem not characterized by elevated densities. Both chemosynthetic driven ecosystems are showing low nematode diversity, and high dominance of single species. Genera richness seems inversely correlated to vent and seep fluid emissions, associated with distinct habitat types. Deep-sea cold seeps and hydrothermal vents are, however, highly dissimilar in terms of community composition and dominant taxa. There is no unique affinity of particular nematode taxa with seeps or vents.

Conclusions

It seems that shallow water relatives, rather than typical deep-sea taxa, have successfully colonized the reduced sediments of seeps at large water depth. For vents, the taxonomic similarity with adjacent regular sediments is much higher, supporting rather the importance of local adaptation, than that of long distance distribution. Likely the ephemeral nature of vents, its long distance offshore and the absence of pelagic transport mechanisms, have prevented so far the establishment of a successful and typical vent nematode fauna. Some future perspectives in meiofauna research are provided in order to get a more integrated picture of vent and seep biological processes, including all components of the marine ecosystem.     

Wood-fall associations from Late Cretaceous deep-water sediments of Hokkaido, Japan

Sunken wood (wood-fall) in the deep sea today is colonized and consumed by diverse invertebrate communities that show phylogenetic relationships to the chemotrophic fauna on whale carcasses, hydrothermal vents and hydrocarbon seeps. Here we document Late Cretaceous examples of wood-fall communities from deep-water sediments of the Yezo Group on Hokkaido, Japan, consisting of provannid, skeneiform, and patelliform gastropods, and thyasirid and nuculanid bivalves. These species are similar or identical to those found on plesiosaur bones and at hydrocarbon seeps in the same sediments, showing that many members of the modern chemotrophic deep-sea fauna colonized this range of habitats at least since Late Cretaceous time. We also document for the first time faecal chimneys in boreholes in these wood fragments, which were presumably built by xylophagain bivalves.     

Methane-flux-dependent lateral faunal changes in a Late Cretaceous chemosymbiotic assemblage from the Nakagawa area of Hokkaido, Japan

A Late Cretaceous carbonate body (2 m in maximum diameter) surrounded by clastic rocks, recently discovered in the Nakagawa area (Hokkaido, Japan), is interpreted as a methane‐seep deposit, on the basis of negative carbon isotopic composition (as low as ?43.5‰), variable sulphide sulphur isotopic composition, high carbonate content, and in situ fractures. It most likely formed owing to methane‐bearing pore‐water diffusion. We estimate that the concentration of methane decreased toward the margin of the carbonate body, and that only small carbonate concretions were precipitated at a certain distance from the methane‐seep centre. These spatial characteristics coincide well with the observed pattern of faunal distribution. The gastropod‐dominated association (indeterminate abyssochrysids and ataphrids and the acmaeid limpet Serradonta sp. are most common) co‐occurs with lucinid and thyasirid bivalves (Thyasira sp., Myrtea sp., and Miltha sp.), and was found within and just above the methane‐derived carbonate body. Acharax and Nucinella (solemyoid bivalves) are more typical of the peripheral part of the methane‐influenced sediments. We suggest that this pattern of faunal distribution reflects the decreasing concentration of methane and apparently also hydrogen sulphide when moving from the centre of discharge toward the periphery of the methane seep.     

Miocene ostracodes of cold seep settings from northern Apennines (Italy)

The Miocene Termina Formation of the northern Apennines (Italy) contains thin marly limestone or marly sandstone bodies rich in macrofauna mainly consisting of large lucinid clams. Modern representatives of this fauna occur in cold seep settings where they house chemosymbiotic bacteria. Moreover, these rocks record a δ¹³C-depletion confirming their origin influenced by a cold seep. The Miocene sections of Sasso delle Streghe and Sarsetta (near Modena) represent classic examples of cold seeps. These sections consist of marls and marly sandstones, respectively, with lucinids (Sasso delle Streghe) or bioclastic sands with lucinids (Sarsetta); both sections are capped by the Sandstones of Montebaranzone. They yield ostracodes that are mainly represented by deep-water filter-feeder species (Platycopa, i.e. Cytherella sp.) and numerous deposit-feeders (Podocopa: i.e. Neonesidea and Krithe), together with species frequently occurring in shallow water environments. These assemblages are able to colonize disaerobic environments such as cold seeps. Although some authors consider the filter-feeders as dominant taxa in disaerobic settings, our data do not provide evidence of significant differences in the ostracode assemblage composition between the deposits originated in seafloors with or without seepage influence. However, the number of specimens is greater in seafloors devoid of seepage influence.     

Cenozoic Methane-Seep Faunas of the Caribbean Region

We report new examples of Cenozoic cold-seep communities from Colombia, Cuba, the Dominican Republic, Trinidad, and Venezuela, and attempt to improve the stratigraphic dating of Cenozoic Caribbean seep communities using strontium isotope stratigraphy. Two seep faunas are distinguished in Barbados: the late Eocene mudstone-hosted ‘Joes River fauna’ consists mainly of large lucinid bivalves and tall abyssochrysoid gastropods, and the early Miocene carbonate-hosted ‘Bath Cliffs fauna’ containing the vesicomyid Pleurophopsis, the mytilid Bathymodiolus and small gastropods. Two new Oligocene seep communities from the Sinú River basin in Colombia consist of lucinid bivalves including Elongatolucina, thyasirid and solemyid bivalves, and Pleurophopsis. A new early Miocene seep community from Cuba includes Pleurophopsis and the large lucinid Meganodontia. Strontium isotope stratigraphy suggests an Eocene age for the Cuban Elmira asphalt mine seep community, making it the oldest in the Caribbean region. A new basal Pliocene seep fauna from the Dominican Republic is characterized by the large lucinid Anodontia (Pegophysema). In Trinidad we distinguish two types of seep faunas: the mudstone-hosted Godineau River fauna consisting mainly of lucinid bivalves, and the limestone-hosted Freeman’s Bay fauna consisting chiefly of Pleurophopsis, Bathymodiolus, and small gastropods; they are all dated as late Miocene. Four new seep communities of Oligocene to Miocene age are reported from Venezuela. They consist mainly of large globular lucinid bivalves including Meganodontia, and moderately sized vesicomyid bivalves. After the late Miocene many large and typical ‘Cenozoic’ lucinid genera disappeared from the Caribbean seeps and are today known only from the central Indo-Pacific Ocean. We speculate that the increasingly oligotrophic conditions in the Caribbean Sea after the closure of the Isthmus of Panama in the Pliocene may have been unfavorable for such large lucinids because they are only facultative chemosymbiotic and need to derive a significant proportion of their nutrition from suspended organic matter.     

Autochthonous death assemblages from chemoautotrophic communities at petroleum seeps: Palaeoproduction,energy flow,and implications for the fossil record

Death assemblages produced by chemoautotrophic communities at cold seeps represent a type of autochthonous accumulation that is difficult to differentiate from other heterotrophic autochthonous communities using taphonomic characteristics. We test the hypothesis that cold‐seep assemblages can be discriminated by unique biological or community attributes rather than taphonomic attributes. To test this hypothesis, we compared several cold seeps on the Louisiana upper continental slope to heterotrophic sites on the Louisiana slope and to a putative seep site in the middle‐late Campanian Pierre Shale near Pueblo, Colorado. Seep assemblages are characterized by a unique tier and guild structure, size‐frequency composition, and animal density that together identify the palaeoenergetics structure of these communities and distinguish them from the other assemblages of the shelf and slope. All seep assemblages were dominated by primary consumers, whereas the heterotrophic assemblage was dominated by carnivores. Carnivore dominance seems to be typical of shelf (or euhaline) death assemblages. Seep assemblages, in contrast, retain the theoretically‐expected rarity of predaceous forms in fossil assemblages. Epifauna and semi‐infauna dominate the tier structure of the heterotrophic assemblage as is typical for continental shelf and slope assemblages. The infaunal tier was unusually well represented in most petroleum seep assemblages. Local enrichment of food resources and the dominance of shelled primary consumers explain the guild and tier structure of seep assemblages. Hindcasting of energy demand (palaeoingestion) and an estimate of sedimentation rate confirms that energy demand by the community exceeds the supply from planktonic rain in seep communities. Thus, seep assemblages can be recognized using biological attributes where taphonomic analysis is ambiguous.     

A high‐stress shelly fauna associated with sponge mud‐mounds in the Coniacian Arnager Limestone of Bornholm,Denmark

The benthic macrofaunas of the Upper Cretaceous chalk of NW Europe show characteristically high species‐richnesses and commonly high densities. They are predominated by bivalves, brachiopods, polychaetes, echinoids, crinoids, asteroids, sponges and towards the end of the Cretaceous also by bryozoans. The mound‐bedded chalk of the Coniacian Arnager Limestone on the island of Bornholm in the Baltic Sea, Denmark, differs from this general picture. It was deposited on a small fault block adjacent to the main Bornholm block, which was emerged during much of the Mesozoic and thus occupied a much more proximal position than most other Upper Cretaceous chalks in NW Europe. The Arnager Limestone contains a unique, exceptionally rich and well‐preserved fauna of mainly hexactinellid, lyssacinosan sponges. The low mud‐mounds are interpreted as formed by baffling and trapping of fine sediment particles by the dense sponge thickets. In contrast, the associated shelly fauna is unusually sparse, of very low richness and extremely low density, except for inoceramid bivalves. It represents a strongly depauparated version of the shelly faunas of contemporaneous chalks in NW Europe. The rare specimens of non‐inoceramid shelly species are interpreted to represent occasional successful spatfalls of benthic species from the deeper‐water chalk farther offshore in the Baltic area. The sponge mud‐mounds of the Arnager Limestone show remarkable resemblances with modern sponge mounds recently discovered on the continental shelf of western Canada. They form an important link between the well‐known older Mesozoic sponge mud‐mounds or ‘reefs’ and the modern mounds and are among the youngest examples of Mesozoic sponge mounds.     

The contrasted evolutionary fates of deep‐sea chemosynthetic mussels (Bivalvia,Bathymodiolinae)

Bathymodiolinae are giant mussels that were discovered at hydrothermal vents and harboring chemosynthetic symbionts. Due to their close phylogenetic relationship with seep species and tiny mussels from organic substrates, it was hypothesized that they gradually evolved from shallow to deeper environments, and specialized in decaying organic remains, then in seeps, and finally colonized deep‐sea vents. Here, we present a multigene phylogeny that reveals that most of the genera are polyphyletic and/or paraphyletic. The robustness of the phylogeny allows us to revise the genus‐level classification. Organic remains are robustly supported as the ancestral habitat for Bathymodiolinae. However, rather than a single step toward colonization of vents and seeps, recurrent habitat shifts from organic substrates to vents and seeps occurred during evolution, and never the reverse. This new phylogenetic framework challenges the gradualist scenarios “from shallow to deep.” Mussels from organic remains tolerate a large range of ecological conditions and display a spectacular species diversity contrary to vent mussels, although such habitats are yet underexplored compared to vents and seeps. Overall, our data suggest that for deep‐sea mussels, the high specialization to vent habitats provides ecological success in this harsh habitat but also brings the lineage to a kind of evolutionary dead end.     

A GIANT LUCINID BIVALVE FROM THE EOCENE OF JAMAICA – SYSTEMATICS, LIFE HABITS AND CHEMOSYMBIOSIS (MOLLUSCA: BIVALVIA: LUCINIDAE)

Abstract:  The giant bivalve Lucina megameris Dall, 1901 , from the late Eocene White Limestone Group of Jamaica and by far the largest known species of the family Lucinidae, is placed in a new genus Superlucina . Apart from its large size, with a shell height exceeding 310 mm, it is distinguished from other genera, such as Pseudomiltha and Eomiltha by external shell characters and the extremely long and narrow, anterior adductor muscle scar. Features preserved on internal moulds suggest that, in common with living Lucinidae, S. megameris was chemosymbiotic with sulphide – oxidizing bacteria housed in the gills. Palaeoenvironmental evidence suggests a habitat in oligotrophic, shallow waters, probably in seagrass beds, with an associated molluscan fauna including large cardiids that may have been photosymbiotic. Superlucina is considerably larger than any living lucinid that range in size from 3 to 150 mm with most encompassed within 5–30 mm. From the Jurassic onwards, a few other large lucinids are known from cold seep sites, with several other records from possible shallow water seagrass beds.     

Cold Seep Epifaunal Communities on the Hikurangi Margin,New Zealand: Composition,Succession, and Vulnerability to Human Activities

Cold seep communities with distinctive chemoautotrophic fauna occur where hydrocarbon-rich fluids escape from the seabed. We describe community composition, population densities, spatial extent, and within-region variability of epifaunal communities at methane-rich cold seep sites on the Hikurangi Margin, New Zealand. Using data from towed camera transects, we match observations to information about the probable life-history characteristics of the principal fauna to develop a hypothetical succession sequence for the Hikurangi seep communities, from the onset of fluid flux to senescence. New Zealand seep communities exhibit taxa characteristic of seeps in other regions, including predominance of large siboglinid tubeworms, vesicomyid clams, and bathymodiolin mussels. Some aspects appear to be novel; however, particularly the association of dense populations of ampharetid polychaetes with high-sulphide, high-methane flux, soft-sediment microhabitats. The common occurrence of these ampharetids suggests they play a role in conditioning sulphide-rich sediments at the sediment-water interface, thus facilitating settlement of clam and tubeworm taxa which dominate space during later successional stages. The seep sites are subject to disturbance from bottom trawling at present and potentially from gas hydrate extraction in future. The likely life-history characteristics of the dominant megafauna suggest that while ampharetids, clams, and mussels exploit ephemeral resources through rapid growth and reproduction, lamellibrachid tubeworm populations may persist potentially for centuries. The potential consequences of gas hydrate extraction cannot be fully assessed until extraction methods and target localities are defined but any long-term modification of fluid flow to seep sites would have consequences for all chemoautotrophic fauna.     

松辽盆地晚白垩世介形类古生态学

通过化石组合和岩性指标分析松辽盆地上白垩统自下而上青山口组—姚家组—嫩江组一段介形类的古生态学。青山口组盐度具有偏咸性、微咸性到半咸性的变化,32个介形类种以栖居于温暖湿润气候带的浅湖微咸水类型占绝对优势,水体清澈安静,有机质丰富,底质为泥等细粒沉积物(个别的为粉砂沉积物),宜于介形类动物群生长且有利于壳体完好保存;其次为生活于半咸水的深湖—半深湖类型,底质为泥。姚家组—嫩江组一段见23个介形类种,栖居于温暖半湿润气候带,以半咸水的深湖、半深湖—浅湖区的类型为主,多泥底,偶见粉砂质底;其次为微咸水的浅湖—三角洲前缘型,泥或粉砂质底;仅3个种生活于温暖湿润的粉砂质底淡水浅湖区。温暖湿润的淡水三角洲分流平原带则难以产出介形类化石。     

A record of fossil shallow‐water whale falls from Italy

Twenty‐five Neogene–Quaternary whales hosted in Italian museum collections and their associated fauna were analysed for evidence of whale‐fall community development in shallow‐water settings. The degree of bone articulation, completeness of the skeleton and lithology of the embedding sediments were used to gather information on relative water depth, water energy, sedimentation rate and overall environmental predictability around the bones. Shark teeth and hard‐shelled invertebrates with a necrophagous diet in close association with the bones were used as evidence of scavenging. Fossil bone bioerosion, microbially mediated cementation and other mollusc shells in the proximity of the remains informed on past biological activity around the bones. The results are consistent with the hypothesis that shallow‐water whale falls differ from their deep‐water counterparts. Taphonomic pathways are more variable on the shelf and whale carcasses may not go through all steps of the ecological succession as recognised in the deep sea. Whilst the mobile scavenger and the enrichment opportunistic stages are well represented, chemosynthetic taxa typical of the sulphophilic stage were recovered only in one instance. The presence of a generalist fauna among the suspension feeding bivalves and carnivorous gastropods, and the extreme rarity of chemosynthetic taxa, suggest that predatory pressure rules out whale‐fall specialists from shallow shelf settings as in analogous cold seep and vent shallow‐water communities.     

Paläobiogeographie jurassischer Muschelfaunen: Beziehung zwischen Süd- und Nordrand der Tethys

Similarities of mid-Jurassic bivalve faunas between the European and the Ethiopian faunal province are very high at the genus-level. At the species-level, however, it is shown that during the Bathonian and Callovian 35% of the bivalves occurring in the Ethiopian faunal province are restricted to this province. In the region of Kachchh (W-India) in the same time-interval 25% of all bivalves are endemic. In the Ethiopian faunal province a clear tendency of increasing endemism from the Bathonian to the Tithonian/Lower Cretaceous at the genus-level and, even more obviously, at the species-level exists. Endemism and provincialism are most marked within the orders Arcoida, Trigonioida, and Nuculoida. The degree of endemism is lower within the Veneroida, but still very high. The orders Mytiloida, Pterioida, and Pholadomyoida hold the largest portion of cosmopolitan species. The rise of endemism and provincialism in Kachchh and m the Ethiopian faunal province from the Bathonian onwards can be explained only partly by the increasing broadening of the Tethys and its effect as an oceanic barrier. The steep increase of endemism in the Upper Jurassic of Kachchh is essentially caused by a radiation within the astartids and trigoniids, accompanied by a reduction of facies-types, due to a regional regression. The very southerly palaeogeographic position of India, the opening of the ‘South African Seaway’, and a change in the marine current system in the uppermost Jurassic led to an increasing differentiation of the Ethiopian faunal province in an ‘Ethiopian-Tethyan’ subprovince to the north and an ‘Ethiopian-Austral’ subprovince to the south. A migration of bivalves in mid-Jurassic times can be reconstructed along the southern margin of the Tethys mainly from east to west. On the other hand, an easternward migration of bivalves along the northern margin of the Tethys from Europe to China and Japan can be documented especially in the Upper Jurassic. This corroberates the existence of a clock-wise marine current system in the northern hemisphere in the Jurassic. The distribution patterns of bivalves in Kachchh and the Ethiopian faunal province are essentially characterized by ‘migration’ of bivalves. The opening of the ‘Hispanic Corridor’ in the Pliensbachian gave way to the immigration of East Pacific bivalves via the western Tethys as far as Kachchh and Madagascar. The dispersal ofPisotrigonia, Seebachia, Tendagurium, andMegacucullaea in the uppermost Jurassic/lowermost Cretaceous from Kachchh and East-Africa respectively to South-Africa and South-America documents the establishment of a ‘South-African Seaway’ and favours migration. However, ‘migration’ and ‘vicariance’ do not exclude each other. On the contrary, both are important mechanisms for creating distributional patterns of bivalves, although within different geological dimensions. Vicariance events produce faunal provinces which last for a long time and within this time-interval, migration seems to be the more important mechanism affecting palaeobiogeographic distribution of bivalves. There is no evidence that the distribution patterns of bivalves in Kachchh and in the Ethiopian faunal province are governed by eustatic sea-level changes. The dominating factors have been a change of the palaeogeographic constellation as a consequence of the break-up of Gondwana, and the local facies distribution. The number of bivalve species known from Europe is much larger than the number of species of the Ethiopian faunal province. A comparison of rarefaction curves of associations, however, shows that this is not a primary feature, but is a consequence of a greater number of different facies types and is due to a far more intense collecting activity in Europe. The bivalves of the Spiti Shales are unequivocal Ethiopian-Tethyan in character. The composition of the fauna indicates the deposition on the deeper shelf of the southern margin of the Tethys. All occurring ‘European’ faunal elements are not significant because of their more or less cosmopolitan distribution.     

Diversity in mussel beds at deep-sea hydrothermal vents and cold seeps

Remarkably little is known about fundamental distinctions (or similarities) between the faunas of deep‐sea hydrothermal vents and seeps. Low species richness at vents has been attributed to the transient nature of vent habitats and to toxic effects of hydrogen sulphide and heavy metals in vent effluents. Seeps are arguably more stable and more chemically benign than vents. They have also been regarded as more diverse, but until now there has not been a rigorous test of this hypothesis. We evaluated diversity indices for invertebrates associated with mussel beds at six vents and two seeps and found that invertebrate diversity was significantly higher at seeps than vents, although some vent mussel beds supported nearly the same diversity as seep mussel beds. Lower diversity at vents may be a consequence of a greater physiological barrier to invasion at vents than at seeps. Diversity was lowest where spacing between vents was greatest, suggesting that risks of extinction as a result of dispersal‐related processes may contribute to the pattern of diversity observed at vents.     

Phylogenetic placement of Cibicidoides wuellerstorfi (Schwager, 1866) from methane seeps and non‐seep habitats on the Pacific margin

Benthic foraminifera are among the most abundant groups found in deep‐sea habitats, including methane seep environments. Unlike many groups, no endemic foraminiferal species have been reported from methane seeps, and to our knowledge, genetic data are currently sparse for Pacific deep‐sea foraminifera. In an effort to understand the relationships between seep and non‐seep populations of the deep‐sea foraminifera Cibicidoides wuellerstorfi, a common paleo‐indicator species, specimens from methane seeps in the Pacific were analyzed and compared to one another for genetic similarities of small subunit rDNA (SSU rDNA) sequences. Pacific Ocean C. wuellerstorfi were also compared to those collected from other localities around the world (based on 18S gene available on Genbank, e.g., Schweizer et al., 2009). Results from this study revealed that C. wuellerstorfi living in seeps near Costa Rica and Hydrate Ridge are genetically similar to one another at the species level. Individuals collected from the same location that display opposite coiling directions (dextral and sinstral) had no species level genetic differences. Comparisons of specimens with genetic information available from Genbank (SSU rDNA) showed that Pacific individuals, collected for this study, are genetically similar to those previously analyzed from the North Atlantic and Antarctic. These observations provide strong evidence for the true cosmopolitan nature of C. wuellerstorfi and highlight the importance of understanding how these microscopic organisms are able to maintain sufficient genetic exchange to remain within the same species between seep and non‐seep habitats and over global distances.     

Chemosymbiotic bivalves from the mud volcanoes of the Gulf of Cadiz, NE Atlantic, with descriptions of new species of Solemyidae, Lucinidae and Vesicomyidae

The chemosymbiotic bivalves collected from the mud volcanoes of the Gulf of Cadiz are reviewed. Of the thirteen species closely associated with chemosynthetic settings two Solemyidae, Solemya (Petrasma) elarraichensissp. n. and Acharax gadiraesp. n., one Lucinidae, Lucinoma asapheussp. n., and one Vesicomyidae, Isorropodon megadesmussp. n. are described and compared to close relatives of their respective families. The biodiversity and distribution of the chemosymbiotic bivalves in the Gulf of Cadiz are discussed and compared to the available information from other cold seeps in the Eastern Atlantic and Mediterranean. Although there is considerable similarity at the genus level between seep/mud volcano fields in the Eastern Atlantic and Mediterranean, there is little overlap at the species level. This indicates a high degree of endemism within chemosymbiotic bivalve assemblages.     

Biogeography and Potential Exchanges Among the Atlantic Equatorial Belt Cold-Seep Faunas

Like hydrothermal vents along oceanic ridges, cold seeps are patchy and isolated ecosystems along continental margins, extending from bathyal to abyssal depths. The Atlantic Equatorial Belt (AEB), from the Gulf of Mexico to the Gulf of Guinea, was one focus of the Census of Marine Life ChEss (Chemosynthetic Ecosystems) program to study biogeography of seep and vent fauna. We present a review and analysis of collections from five seep regions along the AEB: the Gulf of Mexico where extensive faunal sampling has been conducted from 400 to 3300m, the Barbados accretionary prism, the Blake ridge diapir, and in the Eastern Atlantic from the Congo and Gabon margins and the recently explored Nigeria margin. Of the 72 taxa identified at the species level, a total of 9 species or species complexes are identified as amphi-Atlantic. Similarity analyses based on both Bray Curtis and Hellinger distances among 9 faunal collections, and principal component analysis based on presence/absence of megafauna species at these sites, suggest that within the AEB seep megafauna community structure is influenced primarily by depth rather than by geographic distance. Depth segregation is observed between 1000 and 2000m, with the middle slope sites either grouped with those deeper than 2000m or with the shallower sites. The highest level of community similarity was found between the seeps of the Florida escarpment and Congo margin. In the western Atlantic, the highest degree of similarity is observed between the shallowest sites of the Barbados prism and of the Louisiana slope. The high number of amphi-atlantic cold-seep species that do not cluster according to biogeographic regions, and the importance of depth in structuring AEB cold-seep communities are the major conclusions of this study. The hydrothermal vent sites along the Mid Atlantic Ridge (MAR) did not appear as “stepping stones” for dispersal of the AEB seep fauna, however, the south MAR and off axis regions should be further explored to more fully test this hypothesis.     

Kieselschwämme aus dem unterjurassischen Misonekalk der Trento-Plattform (Südalpen): Taxonomie und phylogenetische Relevanz

The Lower Jurassic Misone Limestone of the Trento Platform (Southern Alps, Italy) contains a siliceous sponge fauna which is here described. Besides the well-known Moroccan sponge carbonates, these Lower Jurassic spongioliths from the Trento Platform are presently the second mass occurrence of siliceous sponges, which is known from the southern margin of the Tethys. They differ from each other in regard of the composition of the sponge fauna and the absence of microbial crusts in the spongioliths of the Trento Platform. There, hexactinosans and lithistid demosponges occur in equal proportions. Sphinctozoans are another very characteristic element. Because of the richness in both sphinctozoans and siliceous sponges, the Trento occurrences may be considered as a transitional fauna between the late Palaeozoic-Triassic sponge fauna dominated by sphinctozoans and the post-Liassic sponge fauna dominated by more modern groups of siliceous sponges. Two new siliceous sponge genera with their species are established:Misonia baldensis n. gen. n. sp. (Hexactinosa) andBenacia princeps n. gen. n. sp. (lithistid Demospongiae). The rarity of siliceous sponge dominated spongioliths in the Early Jurassic is due to the restricted occurrence of low energy, deeper shelf areas.     

Palaeoecology and evolution of Mesozoic salinity-controlled benthic macroinvertebrate associations

Salinity-controlled benthic macroinvertebrate associations are typical of many Mesozoic marginally marine environments. They can be recognized by abiotic criteria (e.g., environmental setting, specific autigenic minerals), by biotic criteria (faunal composition, diversity, shell morphology, size-frequency histograms, taphonomic features, associated micro fauna and microflora), and by isotope geochemistry of shells. Although salinity-controlled associations must have been widespread in the European German Triassic, very little is known about their ecology. They appear to have been dominated by the bivalve Unionites and the brachiopod Lingula. In the Jurassic, brackish-water associations are characterized by bivalves, in particular neomiodontids, corbulids, mytilids, bakevelliids, isognomonids, and oysters. In the Cretaceous, in addition, corbiculid bivalves and gastropods become increasingly abundant. Salinity-controlled benthic macroinvertebrate associations can be used to reconstruct salinity regimes of ancient environments, but emphasis should be placed on an integrated sedimentological and ecological approach, as salinity is rarely the only parameter influencing faunal composition and diversity. Although the species composition of salinity-controlled benthic associations changes distinctly through time, the composition of morphotypes remains surprisingly constant throughout the Mesozoic and up to the Recent, evidence of a conservative evolution of benthic faunas within marginal marine high-stress environments. □ Salinity, benthic associations, palaeoecology, Mesozoic.     

Spitsbergen glacial bays macrobenthos – a comparative study

Macrobenthos was studied in seven glacial bays situated along the Spitsbergen coast between 77 and 79°N. The fauna was dominated by deposit-feeding or carnivorous polychaetes and bivalves. Only 4 of 118 species identified in the collected material occurred in all the west Spitsbergen localities examined (the polychaetes Chaetozone/Tharyx sp., Cossura longocirrata, Lumbrinereis fragilis s.l. (sensu lato), and the bivalve Thyasira flexuosa). Clustering of samples showed a difference between the faunas of east and west Spitsbergen; the latter formed two subgroups, localities open to Atlantic waters and those from inner fjord basins. The fauna in open basins was dominated by cosmopolitan species, whereas arctic elements shares were higher in inner basins and predominated in the fauna in Bettybukta (east Spitsbergen). This indicates arctic, relict character of the inner fjords sites. The biomass ranged from 6 to 310 g/m² and Shannon diversities from 0.49 to 2.54. Received: 16 July 1997 / Accepted: 9 February 1998