Origines and evolutions of several hydrothermal vent lineages

De par leur originalité, certains organismes de la faune hydrothermale ont été classés dans des nouveaux groupes taxonomiques de haut rang. De précédentes études moléculaires ont permis d’en réassigner certains dans des lignées connues, occasionnant parfois d’importantes réductions de rangs taxonomiques. Par ailleurs, en phylogénie moléculaire, il est difficile d’optimiser à la fois l’échantillonnage taxonomique et le nombre de marqueurs. Cette thèse illustre cette limitation, mais fournit des avancées sur la compréhension de l’origine et l’évolution de trois groupes hydrothermaux. Chez les vers Polynoidae, l’approche multigène appliquée sur un grand nombre d’espèces suggère au moins deux événements de colonisation de ce milieu. Toutefois, le manque de résolution des marqueurs pour les nœuds profonds entrave la compréhension de l’histoire de ces colonisations. Une limitation similaire avait empêché de replacer les familles de crevettes Alvinocarididae et de crabes Bythograeidae dans leur infra-ordre respectifs (Caridea et Brachyura). Deux approches de recherche et d’identification de marqueurs sont donc testées pour ces groupes. La première, basée sur le séquençage de génome mitochondrial (facilement généralisable), résout les relations profondes des Brachyura et place les espèces de Bythograeidae disponibles proches des Xanthidae. La seconde, basée sur le séquençage de transcriptomes permet d’identifier des marqueurs suffisamment conservés pour résoudre les relations inter-familles chez les Caridea. Cette approche est moins généralisable, mais les marqueurs identifiés pourront a posteriori être recherchés dans un échantillonnage large par capture à l’aide de sonde.The originality of the hydrothermal vents fauna led to the classification of some organisms under new high taxonomic ranks. However, previous molecular studies reassigned them to known lineages, leading to major reductions in such ranking. Classically in phylogenetic studies, optimizing both taxonomic sampling and molecular markers is challenging. This Ph.D project illustrates this limitation, but still provides breakthroughs in the understanding of the origin and evolution of three hydrothermal taxa. In Polynoidae worms, the multigenic approach, led on a large taxonomic and ecological sampling, indicates at least two colonization events of hydrothermal vents. However, the limited resolution of these markers for deep nodes prevented the clear understanding of such events. A similar limitation was previously encountered for Alvinocarididae shrimp and Bythograeidae crabs families in their respective infra-orders (Caridea and Brachyura). Here, two approaches aiming to search and identify markers were tested on these groups. The first one, based on the sequencing of the mitochondrial genome (easily generalizable), resolves deep nodes in Brachyura, and places the available Bythograeidae species near the Xanthidae. The second, based on transcriptome sequencing, allows the identification of molecular markers conserved enough to resolve inter-familial relationships in Caridea. Although this approach is less generalizable, the identified markers could be targeted a posteriori on a wide taxonomic scale using marker-specific probes

Origines et évolution de lignées hydrothermales

The originality of the hydrothermal vents fauna led to the classification of some organisms under new high taxonomic ranks. However, previous molecular studies reassigned them to known lineages, leading to major reductions in such ranking. Classically in phylogenetic studies, optimizing both taxonomic sampling and molecular markers is challenging. This Ph.D project illustrates this limitation, but still provides breakthroughs in the understanding of the origin and evolution of three hydrothermal taxa. In Polynoidae worms, the multigenic approach, led on a large taxonomic and ecological sampling, indicates at least two colonization events of hydrothermal vents. However, the limited resolution of these markers for deep nodes prevented the clear understanding of such events. A similar limitation was previously encountered for Alvinocarididae shrimp and Bythograeidae crabs families in their respective infra-orders (Caridea and Brachyura). Here, two approaches aiming to search and identify markers were tested on these groups. The first one, based on the sequencing of the mitochondrial genome (easily generalizable), resolves deep nodes in Brachyura, and places the available Bythograeidae species near the Xanthidae. The second, based on transcriptome sequencing, allows the identification of molecular markers conserved enough to resolve inter-familial relationships in Caridea. Although this approach is less generalizable, the identified markers could be targeted a posteriori on a wide taxonomic scale using marker-specific probes.De par leur originalité, certains organismes de la faune hydrothermale ont été classés dans des nouveaux groupes taxonomiques de haut rang. De précédentes études moléculaires ont permis d’en réassigner certains dans des lignées connues, occasionnant parfois d’importantes réductions de rangs taxonomiques. Par ailleurs, en phylogénie moléculaire, il est difficile d’optimiser à la fois l’échantillonnage taxonomique et le nombre de marqueurs. Cette thèse illustre cette limitation, mais fournit des avancées sur la compréhension de l’origine et l’évolution de trois groupes hydrothermaux. Chez les vers Polynoidae, l’approche multigène appliquée sur un grand nombre d’espèces suggère au moins deux événements de colonisation de ce milieu. Toutefois, le manque de résolution des marqueurs pour les nœuds profonds entrave la compréhension de l’histoire de ces colonisations. Une limitation similaire avait empêché de replacer les familles de crevettes Alvinocarididae et de crabes Bythograeidae dans leur infra-ordre respectifs (Caridea et Brachyura). Deux approches de recherche et d’identification de marqueurs sont donc testées pour ces groupes. La première, basée sur le séquençage de génome mitochondrial (facilement généralisable), résout les relations profondes des Brachyura et place les espèces de Bythograeidae disponibles proches des Xanthidae. La seconde, basée sur le séquençage de transcriptomes permet d’identifier des marqueurs suffisamment conservés pour résoudre les relations inter-familles chez les Caridea. Cette approche est moins généralisable, mais les marqueurs identifiés pourront a posteriori être recherchés dans un échantillonnage large par capture à l’aide de sonde

Assembly of the mitochondrial genome of the hydrothermal vent crab Segonzacia mesatlantica and detection of potential nuclear pseudogenes

International audienceWe assembled the mitogenome of the Bythograeid crab Segonzacia mesatlantica, using long-range amplification of the mitochondrial genome. The mitogenome is 15,521 base pair long (33.8% A, 21.7% C, 10.5% G, 34%T) with 13 protein-coding genes, 2 ribosomal RNA genes, 22 transfer RNAs and a 624 bp AT-rich region. The gene arrangement is similar to other Brachyuran species. A whole genome shotgun sequencing approach revealed the presence of mitochondrial pseudogenes in the nuclear genome. This fifth mitogenome for a species of Bythograeidae should help resolve the puzzling question of the evolutionary origin of a family limited to deep-sea hydrothermal vents

Rapid colonisation by nematodes on organic and inorganic substrata deployed at the deep-sea Lucky Strike hydrothermal vent field (Mid-Atlantic Ridge)

Despite the fragmented nature of hydrothermal vent (HV) fields, nascent vent sites are rapidly colonized by a pool of regional species. While succession of large animals at vents is relatively well established, we lack information on the associated meiofauna, in particular, on nematodes. The aim of the present study is to investigate the process of colonisation after 9 months of organic (wood and bone) and inorganic (slate) substrata by nematode assemblages deployed at the Eiffel Tower hydrothermal edifice on the Lucky Strike vent field on the Mid-Atlantic Ridge (MAR), at varying distances from visible hydrothermal activity. Abundance, biomass and diversity of colonising nematodes were compared with the results from an earlier similar two-year experiment. Near the vents, nematodes preferred inorganic substrata while in areas not influenced by vent activity, organic substrata were preferred. Nematode females dominated at almost all sites while numerous females at the ovigerous stage and juveniles were reported near the vent emissions, suggesting that nematode populations reproduce well after just 9 months. Our nine-month experiment on the MAR showed that the type of substratum influenced significantly the composition of colonising nematodes, while after two years, the community structure was instead influenced by hydrothermal activity

Ecology and trophic role of Oncholaimus dyvae sp. nov. (Nematoda: Oncholaimidae) from the lucky strike hydrothermal vent field (Mid-Atlantic Ridge)

International audienceBackground: Nematodes are an important component of deep-sea hydrothermal vent communities, but only few nematode species are able to cope to the harsh conditions of the most active vent sites. The genus Oncholaimus is known to tolerate extreme geothermal conditions and high sulphide concentrations in shallow water hydrothermal vents, but it was only occasionally reported in deep-sea vents. In this study, we performed morphological, genetic and ecological investigations (including feeding strategies) on an abundant species of Oncholaimus recently discovered at Lucky strike vent field on the Mid-Atlantic Ridge at 1700 m water depth. Results: We described this species as Oncholaimus dyvae sp. nov.. This new species differs from all other members of the genus by the combination of the following characters: body length (up to 9 mm), the presence of a long spicule (79 mu m) with a distally pointed end, a complex pericloacal setal ornamentation with one precloacal papilla surrounded by short spines, and a body cuticule with very fine striation shortly posterior to the amphid opening. Overall, O. dyvae sp. nov. abundance increased with increasing temperature and vent emissions. Carbon isotopic ratios suggest that this species could consume both thiotroph and methanotrophic producers. Furthermore sulfur-oxidizing bacteria related to Epsilonproteobacteria and Gammaproteobacteria were detected in the cuticle, in the digestive cavity and in the intestine of O. dyvae sp. nov. suggesting a potential symbiotic association. Conclusions: This study improves our understanding of vent biology and ecology, revealing a new nematode species able to adapt and be very abundant in active vent areas due to their association with chemosynthetic micro-organisms. Faced by the rapid increase of anthropogenic pressure to access mineral resources in the deep sea, hydrothermal vents are particularly susceptible to be impacted by exploitation of seafloor massive sulfide deposits. It is necessary to document and understand vent species able to flourish in these peculiar ecosystems