Beyond Spandrels: Stephen J. Gould, EvoDevo, and the Extended Synthesis

  • Gerd B. Müller
Conference paper


In evolutionary biology, the term “spandrel” infallibly elicits the memory of Steve Gould. It has become a standard in referring to constructional byproducts and developmental constraints. More often than not, these were regarded as lesser facets of evolutionary change, with priority given to population dynamics and the workings of natural selection. But the fundamental criticism, in the spandrels paper and other works of Gould, of the absence of organism level factors in the standard Modern Synthesis account, also helped trigger the EvoDevo revolution and important reconceptualizations of evolutionary theory. Recent versions of theory expansion include many of Gould’s propositions but also theoretical changes emerging from other fields, such as genomics, non-genetic inheritance, niche construction, and others. These amount not merely to a numerical addition of factors to be taken into account, but also initiate major shifts in theory structure. As a consequence, today’s extended frameworks of evolutionary theory entail a significant increase in explanatory capacity and predictive power.


Natural Selection Evolutionary Theory Niche Construction Modern Synthesis Developmental Constraint 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



I thank the Istituto Veneto di Science, Lettere ed Arti for the thoughtfulness to organize a meeting commemorating Stephen J. Gould. I am particularly grateful to the president of the Istituto, Gian Antonio Danieli, for the invitation to participate, as well as to Elena Gagliasso, Alessandro Minelli, Telmo Pievani, and Maria Turchetto for their enthusiasm regarding evolutionary theory.


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© Springer-Verlag Italia 2013

Authors and Affiliations

  1. 1.Department of Theoretical BiologyUniversity of Vienna and Konrad Lorenz Institute for Evolution and Cognition ResearchVienna and AltenbergAustria

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