演化生物学
Gould responded that critics ignore that "later" selective value is a separate issue from "origination" as necessary consequences of structure; he summarised his use of the term 'spandrel' in 1997: "Evolutionary biology needs such an explicit term for features arising as byproducts, rather than adaptations, whatever their subsequent exaptive utility ... Causes of historical origin must always be separated from current utilities; their conflation has seriously hampered the evolutionary analysis of form in the history of life." Gould cites the masculinized genitalia of female hyenas and the brooding chamber of some snails as examples of evolutionary spandrels.