Study area
The study was conducted in the Western Ghats biodiversity hotspot of southern India. The landscape (12.17°N, 75.80°E), located in Kodagu district of Karnataka state, is classified as mid-elevation evergreen forests (Pascal, 1986). Elevation spanned 650-1100 m ASL and mean annual precipitation ranged from ca. 2400-3800 mm across sites. The region has well-drained clayey and loamy type soils (Anonymous, 1998). The forests here were fragmented ca. 100 years ago with the expansion of coffee plantations, paddy fields and human settlements. For the last 40 years, both contiguous and fragmented forests were legally protected against land-use change, extraction and hunting. Local communities also protect fragments from conversion due to their cultural value as sacred groves (Osuri and Sankaran, 2016b).