“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” is an article authored by Stephen Jay Gould and Richard Charles
Lewontin published in the Proceedings of the Royal Society of London in 1979. The article
suggests that developmental constraints and phyletic constraints play a central role in explaining
the history of evolution. What they mean by this is, is that the possible paths that evolution could
have followed have been constrained by an organism’s or a phyla’s conserved developmental
stages. Many times, the early developmental stages of organisms are highly conserved because
small changes in early development tend to result in large detrimental defects in later
development. Gould and Lewontin hypothesize that this has constrained the possible pathways of
evolution and has therefore guided the history of life. Historians, biologists, and philosophers
have pointed to this work as important in instigating the philosophical argument about adaptation
and its role in the course of evolution. The authors use the intricate designs inside the spaces in
between arches inside of the rectangles of architectural structures, which are called spandrels, as
an analogy to adaptationist interpretations (see picture below for an example of a spandrel). The
paper emphasizes shortcomings of what the two authors call adaptationism or the adaptationist
programme and it is one in a series of works in that Gould emphasized the study of development
as important for establishing complete evolutionary theories of animal form and function.

Figure 1: Illustration of a Spandrel from Chambers's twentieth century dictionary of the English language, 1903.

1
 Biologists Gould and Lewontin both worked at Harvard University in Cambridge,
Massachusetts, from 1967 until 2002 when Gould died of lung cancer. When the Spandrels paper
was published in 1979 Gould had already published his influential work Ontogeny and
Phylogeny, which explores the relationship between development and ancestry. During this
same period, Lewontin was working on the importance of developmental constraints on the
diversification of life. Although they held different views on many matters, the two shared an
interest in discrediting biological theories they felt would have detrimental consequences to
society.
As leading philosopher of biology Elliot Sober explains, Gould and Lewontin wrote
“The Spandrels” paper for two main reasons: to critique a methodology and to make a claim
about nature. They wanted to challenge the methodology of using adaptation stories to explain
traits but they also wanted to warn against a trend towards determinism in biology. Proponents of
biological determinism, which is the belief that many human traits are determined primarily by
their genetic makeup, claimed that evolution could be used to explain human social behaviors
such as altruism, aggression, and gender behavior. This worried Gould and Lewontin because in
the past, individuals with similar viewpoints in biology used biological determinism to justify
civil rights violations based on innate criminality or poor intellect. A good example was the
eugenics movement at the beginning of the twentieth century when a large number of people
were sterilized against their will in order to prevent those who were seen as weak or feeble
minded from reproducing. But on top of their concerns about biological determinism, the authors
also observed that biologists continually disregarded non-selective forces like phyletic and
developmental constraints when describing the history of life on earth. They thought these forces
have been important in shaping the diversity of life.
The adaptationist program as described by Gould and Lewontin, is a program used to
describe the causes of form and function in animals. The tendency of this program is to view
organisms as a collection of separate parts, also called atomizing the organism, and to assume
each part has been optimized by natural selection. Adaptationist program followers then generate
a unique adaptation story for each part to explain its origins, which Gould and Lewontin saw as
making up just-so stories. The authors argue that this explanatory method is often in error
because it ignores constraints, which limit the number of possible forms organisms can take on.
Furthermore, the authors claim that constraint theories of this kind often provide better

2
explanation of structure and function than adaptationist theories do. According to them,
developmental constraints in particular hold the most power over which pathways evolution has
been able to follow.
The Spandrels paper has six sections. The first three sections describe Gould and
Lewontin’s interpretation of the adaptationist program and how those who support that program
view the development of the diversity of life. The last three offer an alternative approach to
understanding how life has developed by using phyletic and developmental constraints as the
center of analysis rather than adaptation.
In the introduction, Gould and Lewontin point out the shortcomings of the adaptationist
program using an analogy. The authors point to the spandrels in the church of St. Mark’s
Cathedral in Venice, Italy, that are beautifully decorated as an example of how something that
looks so intricately designed may lead us to believe that architects purposefully created the
spandrels for the decorations themselves. However, this is an inverted explanation of the proper
sequence of causes according to Gould and Lewontin. The reality, they note, is that the spandrels
are a product of placing a domed ceiling upon a square room with arched doorways. The authors
say that the spandrels are only afterthoughts; the architectural arches that the building needs to
stand in the first place created the spaces that the architects then used for design. Anytime there
is a domed ceiling on a square room with arched doorways one will get spandrels, and it is only
after the architecture has determined the spandrels that the artist can decorate them to look so
intricate and beautiful. This example is analogous to the situation in evolutionary biology when
biologists try to explain features of organisms in terms of adaptation. Maybe, Gould and
Lewontin hypothesize, developmental constraints are the cause of certain features and can
provide more insight into the origin of traits than adaptation can; just as the arches and
architecture provide more insight into the origin of spandrels than the intricate artwork inside
them can.
In sections two and three, Gould and Lewontin describe how adaptationists see natural
selection as an all-powerful force that can overcome any constraint in order to produce
adaptation. They think adaptationists see natural selection as the only important causal factor of
organic form and function and scientists who use this tactic may claim to recognize constraints
but in reality they rarely see them as important. The authors compare the adaptationists view of
organisms to that of the character of Dr. Pangloss, which is meant to parody the philosopher

3
Gottfried Wilhelm von Leibniz in a story by Voltaire. In the story, Dr. Pangloss claims that our
world is the best of all possible worlds and is portrayed as naive. Like Dr. Pangloss,
adaptationists are also being naive. Gould and Lewontin say that instead of seeing the reality of
the integrated whole organism, adaptationists naively atomize organisms into their traits and then
assign optimality stories. If an optimality story is falsified, the adaptationists will merely come
up with another optimality story and fail to explore different possibilities that do not involve
adaptation.
In the fourth section, Gould and Lewontin describe how Charles Darwin’s view is similar
to their view and not that of the adaptationists. They point out how Darwin consistently showed a
pluralistic approach to the mechanisms of evolution, which took into account several alternatives
to natural selection. Darwin did not claim that natural selection was the only important factor
affecting the evolution of organisms, Gould and Lewontin argue. The authors highlight Darwin’s
anger at his critics who claimed he thought there was only one mechanism of evolution.
In the fifth section Gould and Lewontin highlight several instances where adaptation is
not the primary cause of form and function. According to the authors selection can occur
without adaptation and adaptation without selection. They point to genetic drift, which is a
mechanism of evolution that happens due to random sampling, as a force that requires neither
selection nor adaptation. Furthermore, selection without adaptation can occur in the hypothetical
scenario where a mutation doubles the amount of offspring an organism has in a birthing event.
The mutation will spread through the population quickly (a selection) and yet if resources remain
the same these individuals will leave no more reproducing offspring then before, because the
organisms will just experience a much higher rate in the mortality of their young due to lack of
resources resulting in no adaptation. Also, adaptation without selection occurs in situations
where the environment itself physically causes adaptive traits. For example, the currents in the
ocean physically change sponges and corals. These changes, although not selected for, are
actually beneficial to the survival of the organisms. Gould and Lewontin highlight the difficulties
in distinguishing the origin of a part based on adaptation because one part could have been
originally adapted for a different function than it serves today and could have undergone multiple
adaptations for different functions throughout time.
The final section describes a view of how species change with developmental constraints
as the center of analysis. It illustrates the importance of integrating developmental constraints

4
with adaptive explanations into evolutionary biology. Gould and Lewontin discuss the concept of
the Bauplan (body plan), which is analogous to a blue print that an organism must stick to
because its ancestors evolved on a certain developmental pathway that has become necessary for
survival. For instance humans are not well adapted to being bipedal, which means walking on
two legs, because the majority of the human Bauplan has evolved for quadrupedal locomotion.
According to proponents of the Bauplan, organisms must be looked at as an integrative whole,
not as a collection of parts. Adaptationist explanations only describe superficial changes to the
Bauplan. What is more interesting than small adaptive changes, according to Gould and
Lewontin, is identifying the developmental pathways that have created different Bauplans in the
first place.
Gould and Lewontin believed that developmental constraints might be the best
explanation for the types of evolutionary pathways life has followed. They point to Karl Ernst
von Baer’s laws of embryology as important to the argument. Von Baer’s laws state that early
developmental stages are highly conserved; meaning that there are not many changes made early
on in the development of an organism. This is because the differentiation of organ systems is an
extremely delicate process. Changes made early in development will accumulate and affect all
systems in later stages of development so that small changes early on cause detrimental defects
later. The authors say this leads to developmental constraints; there are a limited number of
developmental pathways an organism can follow without resulting in death. Therefore organisms
have become constrained to a particular evolutionary pathway.
Gould and Lewontin conclude by emphasizing that the adaptationist’s view is not
compatible with the idea of an integrative organism. At beginning developmental stages, an
organism cannot be pulled apart piece by piece into different traits that can then each be
described for a specific purpose. This is because all systems are dependent upon one another.
Small changes to the Bauplan can be explained by adaptation but adaptation cannot explain the
different developmental pathways that evolution has followed. Understanding why these
pathways were followed will help us to understand why organisms look the way they do in the
present.
“The Spandrels of San Marco and the Panglossian Paradigm” is an important paper in the
history of developmental biology. It prompted Stephen Jay Gould and Elisabeth Vrba to publish a
paper, which suggested a new vocabulary in biology in order to distinguish between different

5
kinds of adaptation. They coined the term exaptation, which is an adaptation that has been
remodeled by natural selection for a new use. Moreover, it was one in a series of works by
Stephen Jay Gould that has been pointed out as important to the integration of evolution and
development in biology.
M. Elizabeth Barnes

Sources:
Amundson, Ron. “Two Concepts of Constraint: Adaptationism and the Challenge from
Developmental Biology.” Philosophy of Science 61 (1994): 556−578
Davidson, Thomas. Chambers's twentieth century dictionary of the English language. London:
W. & R. Chambers, 1903.
Dawkins, Richard “Universal Darwinism.” In: Evolution from molecules to man, eds. D. S.
Bendall, 403–425. Cambridge: Cambridge University Press, 1983.
Bendall. Cambridge University Press.Forber, P. “Spandrels and a pervasive problem of
evidence,” Biology and Philosophy, 24 (2009): 247–266.
Gould, Stephen Jay and Lewontin, Richard Charles. “The spandrels of San Marco and the
Panglossian paradigm: a critique of the adaptationist programme.” Proceedings of the
Royal Society of London 205 (1979): 581–598
Gould, Stephen Jay. “The evolutionary biology of constraint.” Daedalus 109 (1980): 39−52.
Maynard Smith, John et. al. “ Developmental constrains and evolution.” Quarterly Review of
Biology 60 (1985): 265−287
Mayr, Ernst. “How to Carry Out the Adaptationist Program?” The American Naturalist 121
(1983): 324−334
Mckitrick, Mary. “Phylogentetic Constraint in Evolutionary Theory: Has It Any Explanatory
Power?” Annual Review of Ecological Systems 24 (1993): 307−330.
Pigliucci, Massimo and Kaplan, Johnathan. “The fall and rise of Dr. Pangloss: adaptationism and
the Spandrels paper 20 years later.” Trends In Ecology and Evolution 15 (2000): 66–70.
Sober, Elliot “Evolution and optimality: feathers, bowling balls, and the thesis of adaptationism,”
Philosophic Exchange 26 (1996): 41–55.
Sober, Elliot. Philosophy of Biology, 2nd ed. Boulder: Westview, 2000. 121−147.

6
Orzack, Steven Hecht and Forber, Patrick, "Adaptationism." The Stanford Encyclopedia of
Philosophy. http://plato.stanford.edu/archives/win2012/entries/adaptationism/ (accessed
January 15, 2013).