Population trends from count data: Handling environmental bias, overdispersion and excess of zeroes

The assessment of population trends is a key point in wildlife conservation. Survey data collected over long period may not be comparable due to the presence of environmental biases (i.e. inadequate representation of the variability of environmental covariates in the study area). Moreover, count data may be affected by both overdispersion (i.e. the variance is larger than the mean) and excess of zero counts (potentially leading to zero inflation). The aim of this study was to define a modelling procedure to assess long-term population trends that addressed these three issues and to shed light on the effects of environmental bias, overdispersion, and zero inflation on trend estimates. To test our procedure, we used six bird species whose data were collected in northern Italy from 1992 to 2019. We designed a multi-step approach. First, using generalised additive models (GAMs), we implemented a full factorial design of models (eight models per species) taking or not into account the environmental bias (including or not including environmental covariates, respectively), overdispersion (using a negative binomial distribution or a Poisson distribution, respectively), and zero inflation (using or not using zero-inflated models, respectively). Models were ranked according to the Akaike Information Criterion. Second, annual population indices (median and 95% confidence interval of the number of breeding pairs per point count) were predicted through a parametric bootstrap procedure. Third, long-term population trends were assessed and tested for significance fitting weighted least square linear regression models to the predicted annual indices. To evaluate the effect of environmental bias, overdispersion, and zero inflation on trend estimates, an average discrepancy index was calculated for each model group. The results showed that environmental bias was the most important driver in determining different trend estimates, although overlooking overdispersion and zero inflation could lead to misleading results. For five species, zero-inflated GAMs resulted the best models to predict annual population indices. Our findings suggested a mutual interaction between zero inflation and overdispersion, with overdispersion arising in non-zero-inflated models. Moreover, for species having flocking foraging and/or colonial breeding behaviours, overdispersed and zero-inflated models may be more adequate. In conclusion, properly handling environmental bias, which may affect several data sets coming from long-term monitoring programs, is crucial to obtain reliable estimates of population trends. Furthermore, the extent to which overdispersion and zero inflation may affect trend estimates should be assessed by comparing different models, rather than presumed using statistical assumption.