Biology & Philosophy

, 24:215 | Cite as

San Marco and evolutionary biology



Gould and Lewontin use San Marco, Venice, to criticise the adaptationist program in biology. Following their lead, the architectural term “spandrel” is now widely used in biology to denote a feature that is a necessary byproduct of other aspects of the organism. I review the debate over San Marco and argue that the spandrels are not necessary in the sense originally used by Gould and Lewontin. I conclude that almost all the claims that Gould makes about San Marco are wrong and that it is reasonable to view the architectural spandrel as an adaptation. The spandrels example has not provided a good illustration of why adaptive explanations should be avoided. In fact, it can be used as an example of how adaptive explanations can be dismissed even when there is evidence in their favour. I also discuss the use of the concept of a spandrel in biology.


Adaptation Constraint Mosaic Necessity San Marco Spandrel Symmetry 



I thank Lutz Fromhage, Robert Mark, Andrew Phillips and Sean Rands for helpful comments on previous versions of the ms and Tim Colborn for producing the figures. This work was supported by a Leverhulme Research Fellowship.


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Copyright information

© Springer Science+Business Media B.V. 2008

Authors and Affiliations

  1. 1.School of Biological SciencesUniversity of BristolBristolUK

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