Scale and intensity of intertidal habitat use by knots Calidris canutus in the Western Wadden Sea in relation to food, friends and foes Piersma, Theunis; Hoekstra, Rinke; Dekinga, Anne; Koolhaas, Anita; Wolf, Pim; Battley, Phil; Wiersma, Popko Take-down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim.Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons the number of authors shown on this cover page is limited to 10 maximum. ABSTRACTIn August-October 1988-1992 we studied the distribution and abundance of knots Calidris canutus around Griend in the western Wadden Sea, and the extent to which these can be explained by benthic prey availability and presence of avian predators. Numbers in the nonbreeding season showed monthly averages of 10000 to 25000 birds. Over 100000 knots were recorded on three occasions. Knots feed in large flocks, individual birds usually experiencing 4000 to 15000 flock-mates. The Siberian-breeding/west-African wintering canutus subspecies passed through in late July and early August. Otherwise the Greenlandic/Canadian breeding islandica subspecies was present. Over the period 1964-1992 there were no clear trends in the number of knots, but canutus-knots were particularly abundant in July-August 1991, whereas in 1992 both subspecies were absent. Macoma balthica was the preferred prey of both subspecies. Hydrobia ulvae, Mytilus edulis and Cerastoderma edule were eaten when Macoma was absent close to the surface of the sediment. As Macoma buried deeper from July onwards, canutus faced better average feeding conditions than islandica later in the year. The spatial distribution of knots feeding on the intertidal flats around Griend was best explained by the harvestable biomass of the prevalent prey species in a particular year and season, Le. Macoma (main prey when their harvestable biomass densities were greater than ca 0.8 g AFDM per m 2) and Cerastoderma, and by the avoidance of situations where they run the risk of attack by bird-eating birds. Flocks of knots covered most of the intertidal flats in the western Dutch Wadden Sea in a couple of tidal cycles. This is about 800 km 2, much larger than the equivalent area used by knots on their wintering grounds in Mauritania (10-15 km2), a difference that is correlated with prey spectrum, prey availability and predictability.

Summary1. There is world-wide concern about the effects of bottom-dredging on benthic communities in soft sediments. In autumn 1988, almost a third of the 50-km 2 intertidal system around the island of Griend in the western Dutch Wadden Sea was suction-dredged for edible cockles Cerastoderma edule and this study assessed subsequent effects. An adjacent area not directly touched by this fishery and an area from which the mussel Mytilus edulis beds were removed, served as reference areas. 2. Sediment characteristics, together with the total stock size and settlement densities of Cerastoderma , Baltic tellin Macoma balthica and soft-shelled clam Mya arenaria , were documented during 11 successive autumns before (August-September 1988) and after (August-September 1989-98) the suction-dredging event in fished and unfished areas. Four other areas in the Dutch Wadden Sea, where changes in densities of juvenile bivalves from 1992 to 1998 were measured, served as additional reference locations. 3. Between 1988 and 1994, median sediment grain size increased while silt was lost from sediments near Griend that were dredged for cockles. The initial sediment characteristics were re-attained by 1996. 4. After the removal of all Mytilus and most Cerastoderma , the abundance of Macoma declined for 8 years. From 1989 to 1998, stocks of Cerastoderma , Macoma and Mytilus did not recover to the 1988 levels, with the loss of Cerastoderma and Macoma being most pronounced in the area dredged for cockles. Declines of bivalve stocks were caused by particularly low rates of settlement in fished areas until 1996, i.e. 8 years after the dredging. 5. A comparison of settlement in the short (1992-94) and medium term (1996-98) after cockle-dredging in several fished and unfished areas spread over the entire Dutch Wadden Sea, showed a significant negative effect of dredging on subsequent settlement of Cerastoderma . Macoma also declined, but not significantly. 6. We conclude that suction-dredging of Cerastoderma had long-lasting negative effects on recruitment of bivalves, particularly the target species, in sandy parts of the Wadden Sea basin. Initially, sediment reworking by suction-dredging (especially during autumn storms) probably caused losses of fine silts. Negative feedback processes appeared to follow that prevented the accumulation of fine-grained sediments conducive to bivalve settlement.

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. University of California Press and American Ornithologists' Union are collaborating with JSTOR to digitize, preserve and extend access to The Auk.ABSTRACT.-Captive Red Knots (Calidris canutus) fed soft food pellets developed atrophied stomachs, and were reluctant to eat their usual hard-shelled mollusc prey. An interspecific comparison among shorebirds showed that wild Red Knots and other intact-mollusc-eating species have gizzards with relatively great mass but very small proventriculi. Within six different shorebird species, the heavier individuals usually had the heavier stomachs as well, but in Red Knots and Bar-tailed Godwits (Limosa lapponica) we identified heavy premigrant individuals with reduced stomach masses, suggesting a reallocation of protein reserves before long-distance flights. In both species reduced stomach mass was associated with a relatively soft diet. We were unable to show that during adjustment of stomachs to hard-shelled prey, such prey are broken down to smaller fragments. We attribute this to the counteractive influence of the pylorus during adjustment. We summarize the suggested stomach/diet interactions as a network of causal relationships and feedback loops involving the type of diet and gizzard mass. We identify two basic modifiers of gizzard mass: one working via endurance training and disuse atrophy; and another involving endocrine and/or neural mechanisms. It is likely that, in the course of their annual cycle, shorebirds are prevented from achieving maximal digestive performance owing to seasonal changes in feeding habitats and diet enforced by their long-distance migrations.

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