Combining functional traits and phylogeny to disentangling Amazonian butterfly assemblages on anthropogenic gradients

Environmental gradients consist of sequential changes in the physical and structural characteristics of a region. These allow us to follow species responses and tolerances under different habitat conditions. Among them, forest fragmentation and succession comprise the most common examples of forest gradients, where organismal responses require distinct morphological, physiological, and behavioral adaptations. However, environmental changes can impose ecological and evolutionary constraints that act on species traits, as well as on local species assemblies through their phylogenetic history. In this study, we evaluated the differences in species distribution and composition on fruit-feeding butterfly assemblages along forest fragmentation and succession gradients. We combine functional and phylogenetic methods for determining butterfly assemblages, and inferred species resistance and resilience according to habitat changes in tropical forests. We used a database of 471 fruit-feeding butterflies of 60 species sampled from different environments in the central Amazon rainforest. A total of 13 functional traits were measured, and a phylogenetic tree was obtained for the sampled species. The trait–environment relationship was analyzed along both forest fragmentation and succession gradients, controlling for phylogenetic signal on species distribution and functional composition when necessary. Several traits presented phylogenetic signal, and phylogeny was also driving butterfly species distribution along the successional gradient. After controlling for phylogeny, individual characteristics related to flight speed (thoracic weight) and antipredatory strategies (camouflage) increased in early-successional forests, with large butterflies (body length) prevailing in primary forests. No clear functional and phylogenetic pattern was identified for the fragmentation gradient. Our results are consistent with the idea that butterflies may be employing distinct functional strategies to attenuate habitat change effects. Larger butterflies, with lower dispersal ability, are preferentially susceptible to local extinctions in the early-successional environments, mainly when forested habitat and its resources become spatially restricted. In addition, several anti-predatory strategies related to conspicuous colors may be losing their functionality in open areas, where not being distinctive against the background becomes the primary defense against predation