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Decoupled morphological and biomechanical evolution and diversification of the wing in bats

Bats use their forelimbs in different ways, flight being the most notable example of morphological adaptation. However, different behavioural specializations beyond flight have also been described in several bat lineages. Understanding the postcranial evolution during the locomotory and behavioural diversification of bats is fundamental to understanding bat evolution. We investigate whether different functional demands influenced the evolutionary trajectories of humeral cross-sectional shape and biomechanics. We found a strong ecological signal and no phylogenetic structuring in the morphological and biomechanical variation in humerus phenotypes. Decoupled modes of shape and biomechanical variation were consistently found, with size and diet explaining variation in shape and biomechanics respectively. We tested both hypothesis- and data-driven multivariate evolutionary models, revealing decoupled pathways of evolution across different sections of the humerus diaphysis. We found evidence for a complex evolutionary landscape where flight might have acted as an evolutionary constraint, while size- and diet-based ecological opportunities facilitated diversification. We also found shifts in adaptive regimes independent from the evolution of flight (i.e. terrestrial locomotion and upstand roosting). Our results suggest that complex and multiple evolutionary pathways interplay in the postcranium, leading to the independent evolution of different features and regions of skeletal elements optimised for different functional demands.