Variation in numbers and behaviour of waders during the tidal cycle: implications for the use of estuarine sediment flats

Estuarine sediment flats are essential feeding areas for waders, but their exploitation is constrained by the movements of tides. In this cyclic environment the exposure period of sediment flats decreases several fold from upper to lower flats, and the moving tidal waterline briefly creates particular conditions for waders and their prey. This study attempts to determine how the exposure period and the movement of the tide line influence the use of space and food resources by waders across the sediment flats. Wader counts and observations of feeding behaviour were carried out in all phases of the tidal cycle, in plots forming a transect from upper to lower flats, thus representing a gradient of exposure periods. Pecking, prey intake, and success rates varied little along the gradient. Some species actively followed the tide line while foraging, whereas others are evenly spread over the exposed flats. Black-tailed Godwit, Dunlin and Avocet were 'tide followers', whereas Grey Plover, Redshank and Bar-tailed Godwit were 'non-followers'. Densities of 'followers' near the tide line were up to five times higher than elsewhere. Species differed markedly in the way they used space on the flats, but in general the rate of biomass acquisition (in grams of ash-free dry weight per time exposed) was much higher in lower flats. However, this preference was insufficient to counter the much longer exposure of the upper flats, so the total amount of biomass consumed on the latter was greater. Therefore, it was in these upper flats that waders fulfilled most of their energetic needs. Consequently, upper flats are of particular importance for the conservation of wader assemblages, but because they are usually closer to shore they tend to suffer the highest pressure from disturbance and land reclamation.