The commentaries have both drawn out the implications of, and challenged, our definition and operationalization of innovation. In this response, we reply to these concerns, discuss the differences between our operationalization and the preexisting operationalization if innovation, and make suggestions for the advancement of the challenging and exciting field of animal innovation.

Ramsey et al. attempt to clarify methodological issues for identifying innovative behaviour. Their effort is seriously weakened by an underlying presumption that the behavior of primates is generally learned and that of non-primates is generally This presumption is based on a poor grasp of the non-primate literature and a flawed understanding of how learned behaviour is genetically assimilated.

Ecology is being introduced to Evolutionary Developmental Biology to enhance organism-, population-, species-, and higher-taxon-level studies. This exciting, bourgeoning troika will revolutionise how investigators consider relationships among environment, ontogeny, and phylogeny. Features are studied (and even defined) differently in ecology, development, and evolution. Form is central to development and evolution but peripheral to ecology. Congruence (i.e., homology) is applied at different hierarchical levels in the three disciplines. Function is central to ecology but peripheral to development. Herein, the supercategories form (‘isomorphic’ (...) or ‘allomorphic’), congruence (‘homologous’ or ‘homoplastic’), and function (‘adaptive’ or ‘nonadaptive’) are combined with two developmental mode (i.e., growth) categories (‘conformational’ or ‘nonconformational’) to provide a 16-class system for analysing features in studies in which ecology, development, and evolution are integrated. (shrink)