Benthic jellyfish dominate water mixing in mangrove ecosystems

Water mixing is a critical mechanism in marine habitats that governs many important processes, including nutrient transport. Physical mechanisms, such as winds or tides, are primarily responsible for mixing effects in shallow coastal systems, but the sheltered habitats adjacent to mangroves experience very low turbulence and vertical mixing. The significance of biogenic mixing in pelagic habitats has been investigated but remains unclear. In this study we show that the upside-down jellyfish Cassiopea sp. plays a significant role with respect to biogenic contributions to water column mixing within its shallow natural habitat (< 2 m deep). The mixing contribution was determined by means of high-resolution flow velocimetry methods in both the laboratory and in the natural environment. We demonstrate that Cassiopea sp. continuously pumps water from the benthos upward in a vertical jet with flow velocities on the scale of centimeters per second. The volumetric flow rate was calculated to be 212 l h-1 for average sized animals (8.6 cm bell diameter), which translates to turnover of the entire water column every 15 minutes for a median population density (29 animals m-2). In addition, we found Cassiopea sp. are capable of releasing porewater into the water column at an average rate of 2.64 ml h-1 per individual. The release of nutrient-rich benthic porewater combined with strong contributions to water column mixing, suggest a role for Cassiopea sp. as an ecosystem engineer in mangrove habitats. Significance StatementWater mixing is a critical process for aquatic life. Coastal mangrove habitats are vital nurseries for commercially and ecologically important species, but these sheltered habitats experience little water mixing. The upside-down jellyfish, Cassiopea sp., occurs in circumtropical mangrove habitats at high densities. They are epibenthic and pulse nearly continuously, producing a vertical current that transports hundreds of liters of seawater per hour. This results in turnover of the entire water column every 15 minutes for an average population. Additionally, Cassiopea sp. can greatly expedite the transport of nutrient-rich water from sediments into the water column. Thus, Cassiopea sp. represents a previously unrecognized ecosystem engineer that can affect primary productivity, nutrient distribution, and alter new habitats as their range is expanding.