A tribute to Hubert Saint Girons: niche separation between Vipera aspis and V. berus on the basis of distribution models

Interspecific competition in contact areas is a major topic in ecological studies. A number of studies were carried out on European vipers, focusing on contact areas between two or three species characterized as appropriate by ecological needs more or less similar. The aim of this study is to extend the analysis of this topic to the case of Vipera aspis and V. berus in an alpine area of northern Italy, by comparing suitability models to verify which ecological factors affect their occurrence and to assess a possible niche separation. Potential distribution was modelled using the maximum entropy method, using six non-correlated ecogeographical variables as predictors. The models fitted well for both species (mean AUC = 0.926; 87.4% of testing data correctly classified). The most informative variables were: habitat, altitude and solar radiation for the asp viper; altitude and habitat for the adder. Deciduous woods, meadows and urban areas had a positive effect on V. aspis distribution as wetlands, meadows and rocks vegetation did on V. berus. However, the variable best separating the species was the elevation, the adder occurring more frequently at higher altitude than the asp viper. Our data showed that the two vipers were mutually exclusive, as already observed by Saint Girons in 1975. Vipera aspis is more thermophilic and lives at low altitude, while Vipera berus lives under cool and humid areas typical of alpine pastures. A similar pattern were found in the contact areas between European vipers belonging to the V. aspis and Pelias group respectively.