How dung beetles respond to a human-modified variegated landscape in Mexican cloud forest: A study of biodiversity integrating ecological and biogeographical perspectives

Aim: To analyse how the dung beetles (Scarabaeinae) respond to a modified, variegated landscape, taking into account the biogeographical peculiarities of the Mexican Transition Zone. Location: This study covers cloud forest (CF) of the Sierra Norte de Puebla mountain range and part of the Sierra Madre Oriental mountain range (Mexico). Methods: We applied proportional sampling based on the landscape variegation model with Scarabaeinae as the indicator group, and using two approaches: structural units (vegetation type) and spatial units (windows). We used two measures richness and Shannon diversity and applied multiplicative diversity partitioning to obtain independent alpha and beta diversities for the landscape, windows and vegetation types. We grouped species by biogeographical distribution pattern for the biogeographical analysis and by whether they were originally from CF. Results: The transformation of CF into secondary forest, pastures and other types of vegetation increases the Scarabaeinae diversity of the landscape, in vegetation types and windows. This increase is the result of species arriving from the tropical lowlands. However, the original dung beetle community of the CF dominates at different scales in the number of species, abundance and biomass. With increasing habitat modification, beta diversity increases in the windows, and species with the Tropical Palaeoamerican distribution pattern increase in abundance in vegetation types and windows. Main conclusions: The variegated character of the landscape explains well the distribution and diversity of this dung beetle community. The peculiar characteristics of the Mexican Transition Zone have an effect owing to the overlap of fauna with different biogeographical origins. The conversion of fragmented landscapes to variegated landscapes could be a conservation goal in human-modified mountain landscapes. Sampling proportional to the area of different types of vegetation and the use of windows offer an alternative experimental design in variegated landscapes.