Systematics and evolution of coastal and deepwater Hydrozoa from the NE Atlantic

The study of the Portuguese Hydrozoa fauna has been abandoned for more than half a century, except for the Azores archipelago. One of the main aims of this Ph.D. project was to contribute new hydrozoan records leading to a more accurate perception of the actual hydrozoan diversity found in Portuguese waters, including the archipelagos of Azores and Madeira, and neighbouring geographical areas, for habitats ranging from the deep sea to the intertidal. Shallow water hydroids from several Portuguese marine regions (including the Gorringe Bank) were sampled by scuba-diving. Deep-water hydroids, from the Azores, Madeira, Gulf of Cadiz and Alboran Sea, were collected by researchers of different institutions during several oceanographic campaigns. Occasional hydroid sampling by scuba-diving was performed in the UK, Malta and Spain. Over 300 hydroid species were identified and about 600 sequences of the hydrozoan ‘DNA barcode’ 16S mRNA were generated. The families Sertulariidae, Plumulariidae, Lafoeidae, Hebellidae, Aglaopheniidae, Campanulinidae, Halopterididae, Kirchenpaueriidae, Haleciidae and Eudendriidae, were studied in greater detail. About 350 16S sequences were generated for these taxa, allowing phylogenetic, phylogeographic and evolutionary inferences, and also more accurate taxonomic identifications. Phylogenetic analyses integrated molecular and morphological characters. Subsequent results revealed: particularly high levels of cryptic biodiversity, polyphyly in many taxonomic groups, pairs of species that were synonymous, the identity of several varieties as valid species, and highlighted phylogeographic associations of hydroids in deep and shallow-water areas of the NE Atlantic and W Mediterranean. It was proved that many (but not all) marine hydroid species with supposedly widespread vertical and/or horizontal geographical distributions, correspond in fact to complexes of cryptic taxa. This study further revealed that, in the NE Atlantic, shallow environments sustain higher hydrozoan diversity and abundance, but the importance of bathyal habitats as a source of phylogenetic diversity was also revealed. The Azorean seamounts were shown to be particularly important in the segregation of populations of hydroids with reduced dispersive potential. The bathyal habitats of the Gulf of Cadiz proved to harbour a considerably high number of cryptic species, which may mainly be a consequence of habitat heterogeneity and convergence of various water masses in the Gulf. The main causes proposed for speciation and population divergence of hydroids were: species population size, dispersal mechanisms and plasticity to inhabit different environmental conditions, but also the influence of oceanic currents (and its properties), habitat heterogeneity, climate change and continental drift. Higher phylogenetic resolution obtained for the family Plumulariidae revealed particularly that glacial cycles likely facilitated population divergence, ultimately speciation, and also faunal evolutionary transitions from deep to shallow waters.

An overview of chemosynthetic symbioses in bivalves from the North Atlantic and Mediterranean Sea

Deep-sea bivalves found at hydrothermal vents, cold seeps and organic falls are sustained by chemosynthetic bacteria that ensure part or all of their carbon nutrition. These symbioses are of prime importance for the functioning of the ecosystems. Similar symbioses occur in other bivalve species living in shallow and coastal reduced habitats worldwide. In recent years, several deep-sea species have been investigated from continental margins around Europe, West Africa, eastern Americas, the Gulf of Mexico, and from hydrothermal vents on the Mid-Atlantic Ridge. In parallel, numerous, more easily accessible shallow marine species have been studied. Herein we provide a summary of the current knowledge available on chemosymbiotic bivalves in the area ranging west-to-east from the Gulf of Mexico to the Sea of Marmara, and north-to-south from the Arctic to the Gulf of Guinea. Characteristics of symbioses in 53 species from the area are summarized for each of the five bivalve families documented to harbor chemosynthetic symbionts (Mytilidae, Vesicomyidae, Solemyidae, Thyasiridae and Lucinidae). Comparisons are made between the families, with special emphasis on ecology, life cycle, and connectivity. Chemosynthetic symbioses are a major adaptation to ecosystems and habitats exposed to reducing conditions. However, relatively little is known regarding their diversity and functioning, apart from a few “model species” on which effort has focused over the last 30 yr. In the context of increasing concern about biodiversity and ecosystems, and increasing anthropogenic pressure on oceans, we advocate a better assessment of the diversity of bivalve symbioses in order to evaluate the capacities of these remarkable ecological and evolutionary units to withstand environmental change.