Seasonality of bioirrigation by the maldanid polychaete Clymenella torquata and related oxygen dynamics in permeable sediments

Benthic organisms in coastal sediments affect elemental cycling and control benthic-pelagic coupling through particle reworking and ventilation of their burrows. Bioirrigation and associated porewater advection create intermittently oxic regions within sediments. The spatio-temporal patterns of such biogenic redox oscillations likely respond to seasonal factors, but quantitative information on the seasonality of bioirrigator behaviors and associated redox dynamics is scarce. We examined bioirrigation by the maldanid polychaete Clymenella torquata and its impacts on sediment oxygenation patterns in permeable sediments using high-resolution planar optode oxygen imaging. In sediment mesocosms with reconstructed summer-collected sediment, the durations of pumping and resting varied inversely with temperature. The average durations of pumping and resting increased from 4 min/4 min at 21 degrees C, to 6 min/6 min at 12 degrees C, to 15 min/14 min at 5 degrees C. In intact cores collected in summer, irrigation patterns (3.5 min/3.5 min) were similar to those observed at 21 degrees C during the temperature ramp. Pumping and resting durations in intact cores collected in winter at 6 degrees C averaged 9 min/26 min, significantly different from patterns at comparable temperatures in the temperature ramp. Pumping patterns strongly affected the temporal patterns of redox dynamics in surrounding sediments. In addition, temperature strongly affected burrow irrigation depth (exclusively within the top -10 cm at 21 degrees C, and down to -20 cm at 5-6 degrees C with an apparent transition at -15 degrees C), indicating that the zone with dynamic redox conditions migrates vertically on a seasonal basis. The differences in pumping patterns between in- and out-of-season experiments and the effect of temperature on irrigation depth underscore the importance of conducting experiments with bioturbators in-season and at field temperatures. The observed seasonal differences in bioirrigation patterns and associated spatio-temporal redox dynamics suggest that rates and pathways of redox-sensitive diagenetic processes and benthic chemical fluxes in permeable sediments likely show considerable seasonal variation.