Macroevolutionary integration underlies limb modularity in the origin of avian flight

The origin of flight in avian dinosaurs has been historically an ideal framework for proposing the evolutionary relationship between form and function in limb proportions under the hypothesis of specialized locomotor modules. However, other hypotheses suggest that the evolution of the forelimb and the hindlimb is strongly influenced by shared developmental constraints, entailing that limbs evolve in an integrated manner and keeping the scientific debate open. To assess this, we used an alternative morphometric approach to compare and statistically analyse limb morphological covariation in a phylogenetically broad context across non-avian maniraptoran theropods and modern birds. Our results show that the maniraptoran lineage shares a strong covariation between limb proportions, a pattern indicating that consistent morphological integration has constrained the forelimb and hindlimb evolutionary transformation. Different evolutionary grades within Maniraptora, both volant and non-volant lineages, display distinct and weaker covariation patterns, suggesting the emergence of independent evolutionary trends within such underlying patterns of integration. These findings are consistent with a developmental hypothesis in which the evolutionary transformation of limbs in maniraptoran dinosaurs was influenced by its serial homology, underscored by shared developmental programmes. Thus, limb evolution was not solely driven by modular (functional) specialization for flight.