'Dinosaur-bird' macroevolution, locomotor modules and the origins of flight

The dinosaurian origin of birds is one of the best documented events that palaeontology has contributed to the understanding of deep time evolution. This transition has been studied on multiple fossils using numerous multidisciplinary resources, including systematics, taxonomic, anatomical, morphological, biomechanical and molecular approaches. However, whereas deep time origins and phylogenetic relationships are robust, important nuances of this transition's dynamics remain controversial. In particular, the fossil record of several maniraptoran groups clearly shows that aerial locomotion was developed before an 'avialization' (i.e., before the first divergence towards avialans), thus earlier than presumed. Although aspects as important as miniaturization and the acquisition of several anatomical and morphological modifications are key factors determining such evolutionary transition, understanding this macroevolutionary trend also involves to seize the evolution of developmental systems, which requires assessing the morphological expression of integration and modularity of the locomotor apparatus throughout time. This is so because, as it happened in other flying vertebrate taxa such as pterosaurs and bats, the transformation of the maniraptoran forelimbs into flying locomotor modules must not only have involved a gradual anatomical transformation, but also a complete developmental re-patterning of the integration scheme between them and the hindlimbs. Here, we review the most relevant aspects of limb morphological transformation during the so-called 'dinosaur-bird' transition to stress the importance of assessing the role of modularity and morphological integration in such macroevolutionary transition, which ultimately involves the origins of flight in dinosaurs.