Study area
We conducted field observations around Ambodiala (commune Farahalana, Sambava District) and Antsikory (commune Ampanefena, Vohemar District), north-eastern Madagascar (Fig. 1). The climate in this part of Madagascar is tropical-humid with an average annual temperature of 25 °C and rainfall over 2133 mm data from ‘Chelsa’ climatologies (Karger et al. 2017). The landscape was formerly covered with humid evergreen forest (Du Puy and Moat 1996) but forests are nowadays fragmented (Vieilledent et al. 2018) and the landscape is dominated by smallholder agriculture.