Evidence for density-dependent survival in adult cormorants from a combined analysis of recoveries and resightings.

1. The increasing population of cormorants (Phalacrocorax carbo sinensis) in Europe since 1970 has led to conflicts with fishery interests. Control of cormorant populations is a management issue in many countries and a predictive population model is needed. However, reliable estimates of survival are lacking as input for such a model 2. Capture-recapture estimates of survival of dispersive species like cormorants suffer from an unknown bias due to permanent emigration from the study area. However, a combined analysis of resightings and recovery of dead birds allows unbiased estimates of survival and emigration. 3. We use data on 11 000 cormorants colour-ringed as chicks in the Danish colony Vorsø 1977-97 to estimate adult survival and colony fidelity. Recent statistical models allowing simultaneous use of recovery and resighting data are employed. We compensate for variation in colour-ring quality, and study the effect of population size and winter severity on survival, as well as of breeding success on fidelity by including these factors as covariates in statistical models. 4. Annual adult survival fluctuated from year to year (0·74-0·95), with a mean of 0·88. A combination of population size in Europe and winter temperatures explained 52-64% of the year-to-year variation in survival. Differences in survival between sexes was less than 1%. Cormorants older than ≈ 12 years experienced lower survival, whereas second-year birds had survival similar to adults. Colony fidelity declined after 1990 from nearly 1 to ≈ 0·90, implying 10% permanent emigration per year. This change coincided with a decline in food availability. 5. Apparently, survival was more severely affected by winter severity when population size was high. This could be caused by saturation of high-quality wintering habitat, forcing some birds to winter in less good habitat where they would be more vulnerable to cold winters. There was thus evidence for density dependence in adult survival, at least in cold winters. 6. The high population growth rate sustained by European Ph. c. sinensis in the 1970s and 1980s can partly be accounted for by unusually high survival of immature and adult birds, probably caused by absence of hunting, low population density and high food availability.