Multi-day water residence time as a mechanism for physical and biological gradients across intertidal flats

Abstract

Tidal flats with shallow-sloping bathymetry under meso-to macrotidal conditions allow organisms to occupy similar tidal elevations at different distances from subtidal channels. As water floods or ebbs across such tidal flats during a single tidal cycle, upstream organisms may modify water properties such as chlorophyll concentration, while physiochemical properties may change due to close association with sediments. Here we report evidence for an additional mechanism establishing cross-shore gradients: multi-day water residence times, in the sense that even if water completely drains into subtidal channels at low tide, a large fraction returns to the flats on the next high tide. We applied circulation modeling and empirical measurements of water properties and benthic secondary production to a 1-km-wide tidal flat in Willapa Bay, Washington, USA. From the circulation model, water parcels on this intertidal flat have residence times up to 2 d, that is, water found on the flat at one high tide returns to the intertidal zone for a median of 4 successive semidiurnal high tides. Modeled residence times generally increased towards shore. Four empirical datasets showed cross-shore gradients consistent with modeled residence times: Salinity time series lagged towards shore; water column chlorophyll declined towards shore at fixed stations (near-bottom) and in surface transects more than could be explained by benthic suspension-feeding during a single transit of water; and oyster (Magallana = Crassostrea gigas) condition declined 25% over 0.5 km from channel to shore, independent of tidal elevation. One environmental measurement was more consistent with within-tide change, as water temperatures warmed towards shore on afternoon flood tides but showed no tidal-cycle lags. Taken together, these patterns suggest that multi-day water residence times can contribute to environmental heterogeneity from channel to shore on tidal flats, acting orthogonally to well-recognized estuarine gradients in residence time from ocean to river.