Summary 1.Millions of shorebirds migrate each year through a small number of highly productive staging areas where they often conflict with fisheries interests. Delaware Bay, USA, is a major shorebird stopover site where, in spring, many thousands of shorebirds undergo rapid mass gain by feeding on the eggs of commercially harvested horseshoe crabs Limulus polyphemus . 2. Environmental factors may cause deviations from the best migration schedule. We used within-year mass gain data from red knot Calidris canutus caught in Delaware Bay between 1998 and 2005 to determine the degree of flexibility individuals have to vary migration speed. 3. Mass gain by birds below 133 g was shown to comprise 15·3% fat (39 kJ g − 1 ), the remainder being lean mass (6 kJ g − 1 ). Above this critical level, fat comprised 83·9% of mass deposition. The rates of energy deposition (kJ d − 1 ) were therefore fundamentally different between the two states but were among the highest ever recorded among vertebrates (5-7 × basic metabolic rate). 4. A total of 36-62% of the variation in observed rates of energy deposition between 1998 and 2002 was explained by a year factor, date and mass at initial capture and interaction terms, such that light-weight birds at the end of May had rates of mass gain or energy deposition two to three times higher than birds of similar mass in mid-May, indicating that birds were attempting to achieve a certain mass by a certain date. In 2003 and 2005, this relationship broke down as a result of lower densities of eggs. 5. Synthesis and application. The maintenance of high densities of crab eggs required for high rates of mass gain in red knot requires severe cuts in, or the complete cessation of, the crab harvest, reduced human and raptor-related disturbance as well as management of beaches to provide sufficient crab-spawning habitat. These findings are widely applicable to other systems where harvesting activities come into conflict with migrating animals and show that certain sections of the population, in this case the long-distance migrants from South America, will be impacted more than short-distance migrants whose physiology may give them access to alternative food resources.

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As field determinations take much effort, it would be useful to be able to predict easily the coefficients describing the functional response of free-living predators, the function relating food intake rate to the abundance of food organisms in the environment. As a means easily to parameterise an individual-based model of shorebird Charadriiformes populations, we attempted this for shorebirds eating macro-invertebrates. Intake rate is measured as the ash-free dry mass (AFDM) per second of active foraging ; i.e. excluding time spent on digestive pauses and other activities, such as preening. The present and previous studies show that the general shape of the functional response in shorebirds eating approximately the same size of prey across the full range of prey density is a decelerating rise to a plateau, thus approximating the Holling type II (' disc equation ') formulation. But field studies confirmed that the asymptote was not set by handling time, as assumed by the disc equation, because only about half the foraging time was spent in successfully or unsuccessfully attacking and handling prey, the rest being devoted to searching.A review of 30 functional responses showed that intake rate in free-living shorebirds varied independently of prey density over a wide range, with the asymptote being reached at very low prey densities (<150/m x2 ). Accordingly, most of the many studies of shorebird intake rate have probably been conducted at or near the asymptote of the functional response, suggesting that equations that predict intake rate should also predict the asymptote.A multivariate analysis of 468 ' spot ' estimates of intake rates from 26 shorebirds identified ten variables, representing prey and shorebird characteristics, that accounted for 81% of the variance in logarithm-transformed intake rate. But four-variables accounted for almost as much (77.3%), these being bird size, prey size, whether the bird was an oystercatcher Haematopus ostralegus eating mussels Mytilus edulis, or breeding. The four variable equation under-predicted, on average, the observed 30 estimates of the asymptote by 11.6 %, but this discrepancy was reduced to 0.2% when two suspect estimates from one early study in the 1960s were removed. The equation therefore predicted the observed asymptote very successfully in 93% of cases.We conclude that the asymptote can be reliably predicted from just four easily measured variables. Indeed, if the birds are not breeding and are not oystercatchers eating mussels, reliable predictions can be obtained using just two variables, bird and prey sizes. A multivariate analysis of 23 estimates of the half-asymptote constant suggested they were smaller when prey were small but greater when the birds were large, especially in oystercatchers. The resulting equation could be used to predict the half-asymptote constant, but its predictive power has yet to be tested.As well as predicting the asymptote of the functional response, the equations will enable research workers engaged in many areas of shore...

As field determinations take much effort, it would be useful to be able to predict easily the coefficients describing the functional response of free-living predators, the function relating food intake rate to the abundance of food organisms in the environment. As a means easily to parameterise an individual-based model of shorebird Charadriiformes populations, we attempted this for shorebirds eating macro-invertebrates. Intake rate is measured as the ash-free dry mass (AFDM) per second of active foraging ; i.e. excluding time spent on digestive pauses and other activities, such as preening. The present and previous studies show that the general shape of the functional response in shorebirds eating approximately the same size of prey across the full range of prey density is a decelerating rise to a plateau, thus approximating the Holling type II (' disc equation ') formulation. But field studies confirmed that the asymptote was not set by handling time, as assumed by the disc equation, because only about half the foraging time was spent in successfully or unsuccessfully attacking and handling prey, the rest being devoted to searching.A review of 30 functional responses showed that intake rate in free-living shorebirds varied independently of prey density over a wide range, with the asymptote being reached at very low prey densities (<150/m x2 ). Accordingly, most of the many studies of shorebird intake rate have probably been conducted at or near the asymptote of the functional response, suggesting that equations that predict intake rate should also predict the asymptote.A multivariate analysis of 468 ' spot ' estimates of intake rates from 26 shorebirds identified ten variables, representing prey and shorebird characteristics, that accounted for 81% of the variance in logarithm-transformed intake rate. But four-variables accounted for almost as much (77.3%), these being bird size, prey size, whether the bird was an oystercatcher Haematopus ostralegus eating mussels Mytilus edulis, or breeding. The four variable equation under-predicted, on average, the observed 30 estimates of the asymptote by 11.6 %, but this discrepancy was reduced to 0.2% when two suspect estimates from one early study in the 1960s were removed. The equation therefore predicted the observed asymptote very successfully in 93% of cases.We conclude that the asymptote can be reliably predicted from just four easily measured variables. Indeed, if the birds are not breeding and are not oystercatchers eating mussels, reliable predictions can be obtained using just two variables, bird and prey sizes. A multivariate analysis of 23 estimates of the half-asymptote constant suggested they were smaller when prey were small but greater when the birds were large, especially in oystercatchers. The resulting equation could be used to predict the half-asymptote constant, but its predictive power has yet to be tested.As well as predicting the asymptote of the functional response, the equations will enable research workers engaged in many areas of shore...

By radio-tracking and recording the movements of flocks, the distribution of feeding Red Knots (Calidris canutus rufa) was studied day and night at a migration stopover site near San Antonio Oeste, Río Negro, Argentina in March and April 1998. By day, the birds fed in dense flocks of 500-4000 on an area of restinga or rock platform where there were beds of the small mussel Brachidontes rodriguezi. By night, this site was deserted, and the birds were found widely scattered over nearby sandflats. It was evident that the birds were feeding at night because variation in the signal strength of the radio-transmitters indicated that the birds were active. Also fresh knot droppings were found in an area which only became exposed by the tide after dark. The reason for the change in feeding distribution may be that the restinga is close to terrestrial habitats that harbor night-adapted predators. Therefore it is avoided at night. By day, it may be safer because better visibility means that predators can be identified more readily. Alternatively, it could be that feeding opportunities become available at night that are better than on the restinga, and this is why the birds feed elsewhere.