Bathymodiolus growth dynamics in relation to environmental fluctuations in vent habitats

The deep-sea mussel Bathymodiolus thermophilus is a dominant species in the East Pacific Rise (EPR) hydrothermal vent fields. On the EPR volcanically unstable area, this late colonizer reaches high biomass within 4-5 years on new habitats created by lava flows. The environmental conditions and growth rates characterizing the reestablishment of B. thermophilus populations are however largely unknown, leaving unconstrained the role of this foundation species in the ecosystem dynamics. A typical example from the vent field at 9°50'N that was affected by the last massive eruption was the Bio-9 hydrothermal vent site. Here, six years later, a large mussel population had reestablished. The von Bertalanffy growth model estimates the oldest B. thermophilus specimens to be 1.3 year-old in March 2012, consistent with the observation of scarce juveniles among tubeworms in 2010. Younger cohorts were also observed in 2012 but the low number of individuals, relatively to older cohorts, suggests limited survival or growth of new recruits at this site, that could reflect unsuitable habitat conditions. To further explore this asumption, we investigated the relationships between mussel growth dynamics and habitat properties. The approach combined sclerochronology analyses of daily shell growth with continuous habitat monitoring for two mussel assemblages; one from the Bio-9 new settlement and a second from the V-vent site unreached by the lava flow. At both vent sites, semi-diurnal fluctuations of abiotic conditions were recorded using sensors deployed in the mussel bed over 5 to 10 days. These data depict steep transitions from well oxygenated to oxygen-depleted conditions and from alkaline to acidic pH, combined with intermittent sulfide exposure. These semi-diurnal fluctuations exhibited marked changes in amplitude over time, exposing mussels to distinct regimes of abiotic constraints. The V-vent samples allowed growth patterns to be examined at the scale of individual life and compared to long-term records of habitat temperature and oceanographic mooring data in the years following the eruption. Both shell growth and habitat temperature at V-vent varied over the spring-neap tidal cycle and over longer periods of c.a. 60 days. The correlation of growth rate with temperature and, for some individuals, with current velocities supports the idea that tidal forcing impacts growth. Its influence on habitat conditions includes the spring-neap cycle, which is not reflected in current velocities but influences the venting rate. Additionally, it is expected that mesoscale eddies periodically passing across the ridge imprint shell growth through the influence of bottom current on the decimeter-thick mixing interface where mussels thrive. We conclude that diurnal-semidiurnal tidal fluctuations exert major abiotic constraints on B. thermophilus mussels and that low-frequency fluctuations act as significant determinants on growth. Finally, we postulate that the modulation of tidal fluctuations by large-scale hydrodynamic forcing ultimately constrains the capacity of this mussel species to form high biomass aggregations. This study indeed shows that the absence of these strong hydrodynamic drivers would limit the alternance of oxic and sulfidic conditions and significantly affect the growth rate of this species over time.