Despite considerable effort for surveillance of wild birds for avian influenza viruses (AIVs), empirical investigations of ecological drivers of AIV prevalence in wild birds are still scarce. Here we used a continental-scale dataset, collected in tropical wetlands of 15 African countries, to test the relative roles of a range of ecological factors on patterns of AIV prevalence in wildfowl. Seasonal and geographical variations in prevalence were positively related to the local density of the wildfowl community and to the wintering period of Eurasian migratory birds in Africa. The predominant influence of wildfowl density with no influence of climatic conditions suggests, in contrast to temperate regions, a predominant role for inter-individual transmission rather than transmission via long-lived virus persisting in the environment. Higher prevalences were found in Anas species than in non-Anas species even when we account for differences in their foraging behaviour (primarily dabbling or not) or their geographical origin (Eurasian or Afro-tropical), suggesting the existence of intrinsic differences between wildfowl taxonomic groups in receptivity to infection. Birds were found infected as often in oropharyngeal as in cloacal samples, but rarely for both types of sample concurrently, indicating that both respiratory and digestive tracts may be important for AIV replication.

Summary 1.Human interests often conflict with those of wildlife. In the coastal zone humans often exploit shellfish populations that would otherwise provide food for populations of shorebirds (Charadrii). There has been considerable debate on the consequences of shellfishing for the survival of shorebirds, and conversely the effects of shorebird predation on the shellfish stocks remaining for human exploitation. Until now, it has been difficult to determine the impact of current shellfishery practices on birds or to investigate how possible alternative policies would affect their survival and numbers. 2. One long-running contentious issue has been how to manage mussel Mytilus edulis and cockle Cerastoderma edule shellfisheries in a way that has least effect on a co-dependent shorebird, the oystercatcher Haematopus ostralegus , which also consumes these shellfish. This study used a behaviour-based model to explore the effects that the present-day management regimes of a mussel (Exe estuary, UK) and a cockle (Burry inlet, UK) fishery have on the survival and numbers of overwintering oystercatchers. It also explored how alternative regimes might affect the birds. 3. The model includes depletion and disturbance as two possibly detrimental effects of shellfishing and some of the longer-term effects on shellfish stocks. Importantly, model birds respond to shellfishing in the same ways as real birds. They increase the time spent feeding at low tide and feed in fields and upshore areas at other times. When shellfishing removes the larger prey, birds eat more smaller prey. 4. The results suggest that, currently, neither shellfishery causes oystercatcher mortality to be higher than it would otherwise be in the absence of shellfishing; at present intensities, shellfishing does not significantly affect the birds. However, they also show that changes in management practices, such as increasing fishing effort, reducing the minimum size of shellfish collected or increasing the daily quota, can greatly affect oystercatcher mortality and population size, and that the detrimental effect of shellfishing can be greatly increased by periods of cold weather or when prey are unusually scarce. By providing quantitative predictions of bird survival and numbers of a range of alternative shellfishery management regimes, the model can guide management policy in these and other estuaries.

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Interference is an important component of food competition but is often di.cult to detect and measure in natural animal populations[ Although interference has been shown to occur between oystercatchers Haematopus ostralegus L[ feeding on mussels Mytilus edulis L[\ four previous studies have not detected interference between oyster! catchers feeding on cockles Cerastoderma edule L[ In contrast\ this study detected interference between cockle!feeding oystercatchers in the Baie de Somme\ France[ Prey stealing "kleptoparasitism#\ one of the main causes of interference between mussel!feeders\ also occurred between oystercatchers in the Baie de Somme[ The kleptoparasitism rate was related to the natural variation in the food supply\ tending to be higher when cockles were rare[ Feeding rate was negatively related to competitor density\ so providing evidence for interference\ but\ as in mussel!feeders\ only above a threshold density of about 49Ð099 birds ha −0 [ The strength of interference at a _xed competitor density was related to the cockle food supply\ usually being greater when cockles were rare[ Previous studies probably failed to detect interference between cockle!feeders because competitor densities were too low\ or cockles were too abun! dant\ or because they were not conducted during late winter when interference is most intense[ The study shows that natural variation in the food supply can in~uence the strength of interference within an animal population and provides support for those behaviour!based interference models which predict that the strength of interference will be greatest when competitor densities are high and prey scarce[ anisms of interference have predicted that the strength of interference "de_ned as the proportional change in intake rate caused by a proportional change in den! sity# should vary with competitor density\ being insig! ni_cant at low densities but steadily increasing in intensity as density rises "Ruxton\ Gurney + de Roos 0881^Moody + Houston 0884^Stillman\ Goss!Cus! tard + Caldow 0886#[ Behaviour!based interference models also predict that interference will be more intense when prey are rare[ This occurs either because predators spend more time searching for prey when prey are scarce and while in this state may interfere with one another "Ruxton et al[ 0881^Moody + Hou! ston 0884^Moody + Ruxton 0885#\ and:or because prey stealing "and hence interference# is only pro_t! able when prey are scarce "Stillman et al[ 0886# [ Two 144 P[ Triplet\ R[A[ Stillman + J[D[ Goss!Custard

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...