Estimating species richness:: calibrating a large avian monitoring programme

1. Species richness is the most widely used measure for the diversity of a biological community. Unfortunately, the number of species counted is usually a biased measure, as not all species present may be detected. Use of species counts as a proxy for true species richness requires the assumption of constant (over space and time) species detectability. This index assumption is hardly ever tested and, if violated, comparisons over time, space or other dimensions, for example different habitats, will be distorted. In monitoring programmes one therefore needs to know the proportion of species present that are detected and how this proportion is affected by external factors. 2. We used capture-recapture techniques to calibrate the Swiss breeding bird survey, where species richness is recorded annually in c. 270 1-km(2) quadrats during two to three visits and interest is focused on annual trends and regional comparisons. Hitherto, analysis has been restricted to species counts, while species detectability and its determinants are not known. We used the interpolated jackknife estimator to compute mean species detectability for 268 quadrats in 2001-03 and tested determinants of detectability related to space, time, observer, survey effort and biology. 3. Mean species detectability averaged 0.89 (SD 0.06, range 0.72-1.00), with no significant difference among years and significant, but small, regional differences. Observers differed, but surprisingly not in relation to their experience in a quadrat. Detectability was positively related to mean visit duration. Larger communities had a lower mean species detectability. A slight violation of population closure because of staggered arrival of migrants did not introduce any measurable bias into our results. 4. Synthesis and applications. Species detectability in the Swiss programme was high and varied little in relation to recognized sources of heterogeneity. Nevertheless, increased standardization should be considered for mean visit duration. While these results are pleasing for the Swiss programme and show that using counts as indices of species diversity need not always induce serious bias, conditions in other programmes, and in the future in the Swiss programme, may be quite different. Both in monitoring programmes and in ecological studies, as a way of risk minimization, species richness ought to be rigorously estimated whenever possible to avoid detection of spurious effects because of changes in species detectability.