RT Journal Article T1 Macroevolutionary integration underlies limb modularity in the origin of avian flight A1 Nebreda, Sergio M. A1 Hernández Fernández, Manuel A1 Marugán Lobón, Jesús AB 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. PB The Royal Society Publishing YR 2025 FD 2025-05-07 LK https://hdl.handle.net/20.500.14352/120047 UL https://hdl.handle.net/20.500.14352/120047 LA eng NO Nebreda, S.M., Hernández Fernández, M. & Marugán-Lobón, J. (2025) Macroevolutionary integration underlies limb modularity in the origin of avian flight. Biol. Lett. 21: 20240685 http://doi.org/10.1098/rsbl.2024.0685 NO Ministerio de Ciencia, Iinnovación y Universidades NO Natural History Museum of Los Angeles County NO Ministerio de Economía y Competitividad DS Docta Complutense RD 19 dic 2025