"The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme" (1979), by Stephen J. Gould and Richard C. Lewontin

The Spandrels of San Marco and the Panglossian Paradigm:
A Critique of the Adaptationist Programme, hereafter called
The Spandrels, is an article written by Stephen J. Gould and
Richard C. Lewontin published in the Proceedings of the Royal
Society of London in 1979. The paper emphasizes issues with
what the two authors call adaptationism or the adaptationist
programme as a framework to explain how species and traits evolved. The paper
is one in a series of works in which Gould emphasized the
role of development in evolutionary theories. The article suggests
that constraints on how organisms can develop and constraints on how species can evolve from others play a
central role in explaining the how species and traits evolve. The
authors note that organisms from different species develop as
embryos through stages similar across species, genera, and higher
classes. Gould and Lewontin hypothesize that those stages
constrained the possible pathways of evolution and has therefore
guided the history of life. Throughout the paper, the authors rely on analogy of some parts of organisms to architectural structures called spandrels, marked in this image as 'a'."