Interpreting the History - Specialized Research AI at MIT

Interpreting the History
of Evolutionary Biology
through a Kuhnian Prism:
Sense or Nonsense?

Koen B. Tanghe
Department of Philosophy and Moral
Sciences, Universiteit Gent, Belgium

Lieven Pauwels
Department of Criminology,
Criminal Law and Social Law,
Universiteit Gent, Belgium

Alexis De Tiège
Department of Philosophy and Moral
Sciences, Universiteit Gent, Belgium

Johan Braeckman
Department of Philosophy and Moral
Sciences, Universiteit Gent, Belgium

Traditionally, Thomas S. Kuhn’s The Structure of Scientiﬁc Revolutions
(1962) is largely identiﬁed with his analysis of the structure of scientiﬁc revo-
lutions. Here, we contribute to a minority tradition in the Kuhn literature by
interpreting the history of evolutionary biology through the prism of the entire
historical developmental model of sciences that he elaborates in The Structure.
This research not only reveals a certain match between this model and the history
of evolutionary biology but, more importantly, also sheds new light on several
episodes in that history, and particularly on the publication of Charles Darwin’s
On the Origin of Species (1859), the construction of the modern evolutionary
synthesis, the chronic discontent with it, and the latest expression of that discon-
tent, called the extended evolutionary synthesis. Lastly, we also explain why this
kind of analysis hasn’t been done before.

We would like to thank two anonymous reviewers for their constructive review, as well as the
editor Alex Levine.

Perspectives on Science 2021, vol. 29, no. 1
© 2021 by The Massachusetts Institute of Technology

https://doi.org/10.1162/posc_a_00359

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Evolutionary Biology through a Kuhnian Prism

Introduction

1.
About a century ago, the American anthropologist Franz Boas (1911) ex-
plained that our experiences of the world are inﬁnitely diverse and that, con-
sequently, reductive linguistic categorization is necessary in order to be able to
think and talk about it. Evidently, our categorization of the world doesn’t
end there but is supplemented with second-order, cognitive abstractions and
categorizations. Examples are typologies of earthquakes or human diseases
and developmental schemes of civilizations or sciences. An example of a
developmental scheme of sciences is that of the Canadian sociologist and
historian of science Yves Gingras (1991). He distinguishes three steps in the
sociological development of scientiﬁc disciplines: the emergence of a new prac-
tice, its institutionalization, and the shaping of a social identity. Méthot (2016)
uses this simple tripartite framework in his analysis of the history and histori-
ography of virology.

Here, we want to apply the developmental model of sciences that Thomas S.
Kuhn elaborated in his The Structure of Scientiﬁc Revolutions (1962) to the history
of evolutionary biology. Of course, as its title indicates, scientiﬁc revolutions
form the main part of Kuhn’s book. Or, as he put it himself (Kuhn [1959]
1977, p. 226): the emphasis of Structure is on “the importance to scientiﬁc
development of ‘revolutions’.” It cannot be denied, though, that it also offers
a “schematic description of scientiﬁc development” (Kuhn 1962, p. 160) or
of “the essential structure of a science’s continuing evolution” (Kuhn 1962,
p. 160) and that his analysis of the structure of scientiﬁc revolutions merely
forms a part of that broader developmental scheme.1

In a ﬁrst section (section 2), we argue that this scheme can be discerned in
the history of evolutionary biology, albeit only partly: on the one hand, it
encompasses a phase that cannot, or not yet, be discerned in the history of
evolutionary biology (a Kuhnian revolution), on the other hand, that history
started (in earnest) with a very important achievement (the publication of
Charles Darwin’s On the Origin of Species in 1859) that, strangely, doesn’t really
ﬁt in Kuhn’s original scheme. Our revelation of this imperfect match between
Kuhn’s developmental model and the standard historiography of evolutionary
biology is, in itself, a not very relevant or maybe even trivial exercise in the
philosophy of the history of science.2 Our analysis doesn’t end there, though: as

1. Greene points out that Structure was the ﬁrst attempt to “construct a generalized pic-
ture of the process by which a science is born and undergoes change and development
(…)”([1971] 1981, p. 30).

2. The standard historiography of evolutionary biology is also known as ‘Synthesis His-
toriography’ since it tells the story of this history in ways that “turn out right for the Modern
Synthesis” (Stoltzfus 2017, p. 4). Stoltzfus and Cable (2014) develop an alternative historiog-
raphy. They claim that a Mendelian-Mutationist synthesis emerged in the early twentieth cen-
tury and that contemporary evolutionary thinking seems closer to this synthesis than to the

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Perspectives on Science

we will show in section 3, a Kuhnian x-ray scan of the history of evolutionary
biology yields several new insights in that history. Lastly, section 4 is dedicated
to explaining why the history of evolutionary biology hasn’t been interpreted in
terms of Kuhn’s developmental scheme before.

2. An Imperfect Match
The broad outline of Kuhn’s scheme is well known: a pre-paradigmatic phase
in the historical development of a science is followed by the emergence of a
paradigm, a normal science phase and a phase of extraordinary science and rev-
olution. The main distinction in The Structure is that between this revolutionary
phase and normal science. This is the ‘essential tension’ in scientiﬁc research
(see, e.g., Marcum 2015, pp. 46–7 or Bird 2000, ch. 2). The pre-paradigmatic
phase, by contrast, is only brieﬂy discussed in chapter two, “The Route to
Normal Science.”3 It is characterized by the coexistence of a number of com-
peting schools which each interpret a natural phenomenon in a different way.
The bickering of adherents of these different pre-paradigmatic schools is not
entirely fruitless, though. They are genuine scientists, but much of their efforts
are devoted to combatting and trying to convince opponents, rather than to
cumulative research.

A normal science phase starts with the emergence of a paradigm: a new
theoretical framework that inspires the work of a majority of the practitioners
of a speciﬁc discipline because it solves important problems and offers research
opportunities and particularly because it provides them with standard exam-
ples of good scientiﬁc practice. Or, as Kuhn put it himself:

[These works were] sufﬁciently unprecedented to attract an enduring
group of adherents away from competing modes of scientiﬁc activity.
Simultaneously, it was sufﬁciently open-ended to leave all sorts
of problems for the redeﬁned group of practitioners to resolve.
Achievements that share these two characteristics I shall henceforth

modern evolutionary synthesis. Consequently, it is the Mendelian-Mutationist synthesis that
should be considered as the foundation of modern evolutionary biology. For another example of
a criticism of the Synthesis-dominated historiography of evolutionary biology, see Amundson
(2005).
3.

In this chapter, Kuhn points out that the study of light and of electricity was charac-
terized by fundamental disagreements before the publication of the work of, respectively, Isaac
Newton and Benjamin Franklin (and his immediate successors). These disagreements also
characterize (1) “the study of motion before Aristotle and of statics before Archimedes, the
study of heat before Black, of chemistry before Boyle and Boerhaave, and of historical geology
before Hutton” (Kuhn 1962, p. 15) and (2) the study of life phenomena before the appearance
of unspeciﬁed biologists. He is less certain about the social sciences: “it remains an open ques-
tion what parts of social science have yet acquired such paradigms at all. History suggests that
the road to a ﬁrm research consensus is extraordinarily arduous” (Kuhn 1962, p. 15).

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Evolutionary Biology through a Kuhnian Prism

refer to as ‘paradigms,’ a term that relates closely to ‘normal science’.
By choosing it, I mean to suggest that some accepted examples of
actual scientiﬁc practice (…) provide models from which spring
particular coherent traditions of scientiﬁc research (1962, p. 10).4

This paradigm concept was inspired by the fundamental difference between
internally divided social scientists and more united practitioners of the
natural sciences that Kuhn noticed during a stay in 1958–1959 at the Center
of Advanced Studies in the Behavioral Sciences (see Kuhn 1962, p. x). How-
ever, by 1974, he had changed this origin story of his central term. It was now
not coined to account for the difference between the social sciences and the
natural sciences but, more in general, “to account for the consensus necessary
among scientists in a specialty in order for them to pursue their research goals
effectively” (Wray 2011, p. 49). Also, Kuhn now claimed that it was “the
sense of ‘paradigm’ as standard example that led originally to [his] choice
of that term” (Kuhn 1977, p. xviii): a group’s unproblematic and common
or paradigmatic conduct of research was facilitated by those shared examples
of successful practice (i.e., paradigms in the speciﬁc meaning of exemplars).
They could provide what a paradigmatic group “lacked in rules” (Kuhn
[1974] 1977, p. 318).5

In the postscript to the second edition of Structure, he had already claimed
that the term paradigm had been inspired by the exemplar component “of a
group’s shared commitments” (Kuhn [1962] 1970, p. 187). He introduced
this distinction between exemplars and a group’s “shared commitments” (i.e.,
“the entire constellation of beliefs, values, techniques and so on shared by

4. This is also how he deﬁnes paradigms in the preface of Structure: paradigms are “univer-
sally recognized scientiﬁc achievements that for a time provide model problems and solutions to
a community of practitioners” (p. viii). The term ‘achievement’ or ‘achievements’ appears 28
times in the ﬁrst edition and 3 times in the postscript to the second edition. However, the idea
that such achievements or paradigms (sensu lato) deﬁne a scientiﬁc tradition by providing exem-
plars to a community of scientists seems to be more applicable to the practice of the physical
sciences than to that of the life sciences (see, in this respect, Hacking 2012, p. xxiii; see also note
9; for examples of exemplar-based accounts of episodes in the history of biology, see Creager
2016 and Shan 2020).

In the same essay, he claimed that the distinction between what he had called a pre- and
a post-paradigmatic phase in the development of a science, was “both typical and important”
(Kuhn [1974] 1977, p. 295), but that it could “be discussed without reference to the ﬁrst
achievement of a paradigm” (Kuhn [1974] 1977, p. 295n4). Because, “[w]hatever paradigms
may be, they are possessed by any scientiﬁc community, including the schools of the so-called
pre-paradigm period” (Kuhn [1974] 1977, p. 295n4). However, he also pointed out that the
transition to the post-paradigmatic phase usually happens “in the aftermath of some notable
scientiﬁc achievement” (Kuhn [1974] 1977, p. 295n4) (i.e., a paradigm in the broadest
meaning of the term). It probably shouldn’t surprise us that the term paradigm “practically
vanished from Kuhn’s writing beginning around 1980” (Marcum 2015, p. 177).

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Perspectives on Science

members of a given community” [p. 175]) or disciplinary matrix in response
to criticisms of his multifarious semantic use of the term paradigm in the ﬁrst
edition (Shapere 1964; Masterman 1970; see also Hoyningen-Huene [1989]
1993, pp. 131–32). Exemplars were the main components of such a matrix
and “the most novel and least understood aspect” (Kuhn [1962] 1970, p. 187)
of Structure.

Whatever the true origin of the term, according to the ﬁrst edition of Struc-
ture, paradigms are important theoretical achievements (paradigmatic theo-
ries) that develop out of one of the pre-paradigmatic theories, or originate as a
synthesis of some of these theories. They subsequently inspire and guide the
research of a previously divided community of scientists: “Except with the
advantage of hindsight, it is hard to ﬁnd another criterion that so clearly pro-
claims a ﬁeld a science, than the creation of a paradigm that proved able to
guide the whole group’s research” (Kuhn 1962, p. 22). It is only then that
scientists engage in the kind of esoteric and technical research that scientists
typically do. Symptomatic for this professionalization is that scientists in-
creasingly communicate this research to their peers in specialized journals,
rather than in books, directed at a general public. Normal science can also
encompass reformulations of paradigms. These are sometimes required to
solve theoretical problems. Kuhn refers more particularly to eighteenth-
and nineteenth-century reformulations of Newton’s mechanical theory “in
an equivalent but logically and aesthetically more satisfying form” (1962, p.
33). Lastly, when a paradigm is confronted with serious or signiﬁcant anoma-
lies, a crisis ensues which, in some cases, is followed by its replacement by a new
paradigm. Ruse distinguishes four dimensions in this revolutionary process:
the new paradigm unites a new group of scientists (the sociological dimen-
sion), provides them with a new Weltanschauung (the psychological dimension)
and ditto methodology and data (the epistemological dimension) and even
changes the world (the ontological dimension): “it is not simply a question
of the world seeming to change for us, but rather the world really does change”
(1978, p. 246).

In what follows, we will show that at least three of Kuhn’s four phases
can be clearly distinguished in the history of evolutionary biology: the pre-
paradigmatic phase (section 2.1), the emergence of a paradigm (in the gen-
eral meaning of ‘paradigmatic theory’) (section 2.2) and a normal science
phase (section 2.3).6 Lastly, we will brieﬂy discuss the question whether we
are currently witnessing a Kuhnian crisis and the start of a Kuhnian rev-
olution (section 2.4).

6. Our previous analysis of these phases (Tanghe et al. 2018) is here slightly modiﬁed

and also in various ways extended.

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Evolutionary Biology through a Kuhnian Prism

The Pre-Paradigmatic Phase

2.1.
Charles Darwin (1859) managed to put the notion of evolution (as it later came
to be known) on the late-nineteenth century scientiﬁc agenda, but he did not
succeed at all in creating a single, united community of evolutionary biologists.
Put differently, Darwin’s original theory of evolution never became a paradigm.
Instead, there emerged a large number of not always sharply delineated pre-
paradigmatic schools of evolutionists (Bowler 1983) which each interpreted
the phenomenon of evolution in a different way and focused on speciﬁc biolog-
ical phenomena that suited their theory best.7 For example, neo-Darwinists
were preoccupied with adaptations, mutationists with discontinuous variations
and orthogenesists or adherents of the idea of straight-line evolution with (pre-
sumed) trends in the fossil record. All of these theories “competed for status”
(Largent 2009, p. 3). The standard term for this pre-paradigmatic phase in the
history of evolutionary biology is “the eclipse of Darwinism.”

The Paradigm

2.2.
The pre-paradigmatic chaos in evolutionary biology came to an end after the
Second World War, when the long genesis of the modern evolutionary syn-
thesis (henceforth MS) culminated in its coronation, at the 1947 Princeton
Conference on Genetics, Palaeontology and Evolution, as the ﬁrst paradigm
of evolutionary biology (Smocovitis 1996). Provine speaks in respect with this
genesis of an “evolutionary constriction” (1989, p. 61): one pre-paradigmatic
approach of evolution, that of population genetics, came out victorious and all
other alternatives lost all credibility among a majority of biologists. To be
precise, population genetics became the “formalized core of the MS theory”
(Müller 2017, p. 2).

The main reason why population genetics was victorious is that it offered
a convincing, mathematical solution for a major pre-paradigmatic problem:
the question whether the natural selection of continuous variations can cause
major adaptive transmutations of populations. The Cambridge mathemati-
cian H. T. J. Norton was the ﬁrst to make use of the Hardy-Weinberg equi-
librium principle (i.e., the relative frequency of alleles remains the same in a
population, unless this equilibrium is disturbed by forces like selection) and
to show, in an appendix to R. C. Punnett’s Mimicry in Butterﬂies (1915), the
efﬁcacy of natural selection. The ﬁnding that even a slight advantage would
lead to a substantial increase in the frequency of a favorable allele over a few
generations “came as a surprise to most people, and it stimulated a ﬂow of

7. Bowler (1983) distinguishes three schools: Lamarckism or neo-Lamarckism, ortho-
genesis and the mutation theory. Elsewhere (Bowler [1989] 2009), he speaks of four schools:
Lamarckism or neo-Lamarckism, orthogenesis, the mutation theory and Weismann’s neo-
Darwinism.

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Perspectives on Science

new research” (Young 1993, p. 190). Population genetics thus also intro-
duced “a new and more rigid deﬁnition of the ﬁeld” (Kuhn 1962, p. 19):
“a change in the genetic composition of populations” (Dobzhansky 1937,
p. 11). This became the primary or default deﬁnition of evolution for many
evolutionary biologists and an important inspiration for evolutionary research
(Michod 1981, Charlesworth and Charlesworth 2017), although it was
“never totally accepted” (de Jong and Scharloo 1988, p. 1). The MS was,
nevertheless, welcomed as a breath of fresh air since “it was soon appreciated
that a basis had now been laid on which future evolutionary studies could be
safely built” (Young 1993, p. 218).

The Post-Paradigmatic Phase: Normal Science

2.3.
It is undeniable that the study of evolution became more professional after the
establishment of the MS as a paradigm. The fuzzy, philosophical concept of
“progress” was expelled from evolutionary theory (Ruse 2009a). The phrase
“evolutionary biology,” ﬁrst coined in 1881 in Grant Allen’s Vignettes From
Nature (he spoke of an evolutionary biologist), “increasingly became an ac-
cepted disciplinary appellation” (Smocovitis 1996, p. 163). In 1945, the So-
ciety for the Study of Evolution (SSE), replaced the Society for the Study of
Speciation. Specialized journals, dedicated to evolution, were founded in sev-
eral countries. In 1946, the SSE, on the occasion of its First Annual Meeting,
established a research journal in the ﬁeld of evolution: Evolution: An Interna-
tional Journal of Organic Evolution.8 The journal American Naturalist turned from
experimental biology to evolutionary biology. Courses in evolutionary biology
blossomed and important evolutionists moved back from museums into uni-
versity departments.

Most importantly, the MS marked the start of a period of normal science.
Dickins and Rahman (2012, p. 2914), for example, speak in respect with the
period after the emergence of the MS of “many years of normal scientiﬁc activity
exploring the hypothesis-space that it created.” Normal science consists, in
general terms, of the realisation of the promise of success in the investigation
of nature that largely explains the attraction of a paradigm. It encompasses an
empirical and a theoretical dimension and is achieved by extending the knowl-
edge of those facts that the paradigm displays as particularly revealing or
relevant (“factual determinations”), by matching of facts with theory, and
through a further articulation of the paradigm itself. It is through normal

8. The original subtitle was: “An International Journal of Evolutionary Biology.” The SSE
changed it in 1947. Smocovitis (1996, p. 163n199) couldn’t ﬁnd a reason, in the archives, for
this change. She believes that palaeontologists, who occupied positions in geology depart-
ments, found it difﬁcult to adopt the term “evolutionary biology.” Mayr’s remark that some
older members of the SSE wanted the word “organic” included in the journal’s title offers an
alternative or additional explanation. See: http://www.evolutionsociety.org/history.html.

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Evolutionary Biology through a Kuhnian Prism

science that a paradigm acquires the precision that is necessary to reveal the
anomalies that can lead to a crisis and, eventually, a revolution. Empirical
normal science can be thought of as forcing “nature into the preformed and
relatively inﬂexible box that the paradigm supplies” (Kuhn 1962, p. 24),
whereas theoretical normal science (i.e., the modiﬁcation, reformulation and
further articulation of the paradigm) can be thought of as extending and also
modifying that “relatively inﬂexible” box.

The MS clearly inspired these two general kinds of Kuhnian research.9
Examples of empirical research are the application of the models of mathe-
matical population genetics in the ﬁeld and in the laboratory (e.g., Lewontin
et al. 1981), the study of swift changes, through natural selection, of body
and beak size in Galápagos ﬁnches in response to changes in their food supply
(Grant and Grant 2003, 2008), and the empirical study of cladogenesis
through geographical isolation (allopatry; Mayr 1942). The MS has, secondly,
also been greatly extended and substantially modiﬁed (i.e., theoretical normal
science), especially in the last few decades.

Some examples of important modiﬁcations of the MS are Kimura’s neutral-
ism (1968, 1991), the exploration of alternative speciation models (i.e., next to
allopatry) (Mallet 2001), and the acknowledgment that macroevolution is not
always merely an extrapolation of microevolutionary processes—one of the
tenets of the original MS (Erwin 2000). Lastly, Lu and Bourrat (2017) distin-
guish three kinds of extensions of the MS: classical population genetics has
been generalized to quantitative genetics for continuous traits and is now also
better suited to account for the evolution of microorganisms and plants. A
third category of extensions consists of progress in various biological subﬁelds.
Two examples are the theory of major evolutionary transitions (Maynard Smith
and Szathmáry 1999) and Bill Hamilton’s (1964) neo-Darwinian explanation
(i.e., kin selection) of altruism.

2.4. A Kuhnian Crisis?
Some scholars suggest that evolutionary biology is in the thralls of a Kuhnian
crisis and that the so-called extended evolutionary synthesis (henceforth EES)
(e.g., Pigliucci and Müller 2010) is about to replace the MS. For example, Dick-
ins and Rahman write:

The MS is now a commonly agreed framework adhered to by many
biologists. But one might expect a gradual accumulation of glitches, of

9. Kuhn’s detailed account of ‘the nature of normal science’ (ch. 3) is structured around
these two kinds of research. They are each subdivided in three subcategories. However, these
subcategories are clearly very much inspired by the practice of the physical sciences and, there-
fore, not always applicable to the practice of evolutionary biology (for example experimental
efforts to articulate a paradigm by determining universal constants).

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ﬁndings that do not quite ﬁt with the common framework, and as this
accumulation increases more suitable theories will begin to be sought,
theories that might encompass the glitches as well as reorganize our
previously accepted facts. We would anticipate a paradigm shift. It is
this form of analysis that informs those seeking an extended synthesis.
(2012, p. 2914)

That remains to be seen though. As the eminent Kuhn scholar Hoyningen-
Huene writes in his Kuhnian analysis of the chemical revolution: “Normal
science ceases and a phase of extraordinary science begins when significant
anomalies indicate that there are very serious problems concerning the
currently accepted theoretical and/or experimental framework” (2008, p. 105),
and “[t]he term ‘signiﬁcant’ just used to qualify anomalies is of extreme impor-
tance,” since most anomalies are “an omnipresent concomitant of normal sci-
ence” (2008, p. 105). Are evolutionary biologists currently really faced with
signiﬁcant anomalies that indicate that there are “very serious problems” with
the MS? We doubt it. Telling, in this respect, is that most adherents of the EES
themselves do not believe that this model is an emerging, new paradigm.
Rather, they tend to interpret it as an alternative, Lakatosian research pro-
gram (see, e.g., Pigliucci and Finkelman 2014; Laland et al. 2015; see also
Pievani 2012).10 In the following section, we will propose an alternative and
new Kuhnian interpretation of the EES.

3. New Insights and Contributions
As pointed out above, the identiﬁcation of a certain correspondence between
phases in Kuhn’s developmental scheme and stages in the history of evolution-
ary biology is, in itself, quite trivial. The more pertinent question is whether a
Kuhnian analysis of that history is heuristically rewarding. We believe that this
is indeed the case: such an analysis particularly provides us with a better insight
in the so-called eclipse of Darwinism (3.1), in the still poorly understood gen-
esis of the MS (3.2), and in the relatively turbulent normal science period in the
history of evolutionary biology (3.3), including the precise nature of the EES.
As we shall see, the extreme multidisciplinarity of the science of evolutionary
biology (Welch 2017) left a strong mark on each one of these three Kuhnian
phases in its history.

The Misnomer, Called the Eclipse of Darwinism

3.1.
We will return to the place (or lack thereof ) of Darwin’s On the Origin in
Kuhn’s (original) scheme and the implications of our Kuhnian analysis for

10. Müller (2017) is an example of a more ambitious proponent of an EES. He claims
that the term (or acronym) ‘EES’ is not meant as a simple extension of the MS, “as sometimes
wrongly implied, but to indicate a comprehensive new synthesis (…)” (p. 9).

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the historical status of his theory of evolution in section 4.1. (see particularly
note 17). Here, we want to focus on what followed the publication of this
seminal work. Largent portrays the phrase ‘eclipse of Darwinism’ as a deter-
ministic and Whiggish metaphor that has served the purposes of certain bi-
ologists and historians, at the “expense of our understanding of the research,
conclusions, and worldviews of early twentieth-century American evolutionists”
(2009, p. 7). It constructed the MS as an inevitable, discontinuous and “predict-
able solution to the problem of darkness” (2009, p. 4) that supposedly had pla-
gued evolutionary biology after the publication of Origin (1859). He refers more
particularly to Vernon Lyman Kellogg’s book Darwinism To-Day (1907) which
has erroneously been interpreted from the perspective of this metaphor but
which, in reality, “heralded the primacy of Darwinian natural selection (…)”
(2009, p. 15). The eclipse metaphor is a lingering relic that is harmful to our
ability accurately to depict twentieth-century evolutionary biology and should
be replaced with “a new term and a new conception of the work done in evo-
lutionary biology between 1880 and 1940, one that analyzes early twentieth-
century evolutionary biology on its own terms, not merely in the context of what
followed” (2009, p. 18).

As we explained previously (Tanghe et al. 2018), the phrase ‘eclipse of
Darwinism’ is indeed somewhat Whiggish or anachronistic as we can only
describe this episode as an eclipse with the beneﬁt of hindsight. It is in several
ways very misleading. Firstly, as pointed out above, Darwinism was never the
dominant interpretation of evolution (i.e., a paradigm); secondly, it did never
completely ‘disappear’ (there were always Darwinists or ultra-Darwinists);
thirdly, the eclipse metaphor ignores the fact that, during the so-called eclipse,
Darwinism was one of several competing pre-paradigmatic theories (Junker
2008, p. 496); and, fourthly, Darwinism also didn’t reappear unchanged in
the twentieth century, like the sun after a literal eclipse. We shouldn’t overes-
timate the negative impact this metaphor has had on modern historians, though.
In 1983, Bowler (1983, p. ix) already claimed that his thesis that the eclipse rep-
resented a crucial phase in the development of modern evolutionism “has become
widely accepted.” Still, “pre-paradigmatic phase” seems to us to be a better phrase
since Kuhn’s characterization of this period is a more accurate description of what
evolutionists did between 1860 and 1947 than what is suggested by the phrase
‘eclipse of Darwinism’ (or by Largent’s alternative term “interphase”): they did
sometimes important research but within the context of different and quarrelling,
pre-paradigmatic schools of thought.

However, in one important way, the pre-paradigmatic phase in the history
of evolutionary biology was unlike pre-paradigmatic phases of other sciences:
the multidisciplinarity of this science led to a tentative association between
pre-paradigmatic approaches of evolution and speciﬁc biological disciplines.
They each tended to approach evolution from the speciﬁc perspective of their

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own disciplinary specialty, which, as we shall see, highly complicated the
construction of the MS. Some Mendelists, for example, were drawn to de
Vries’ mutationism because they studied the transmission of discrete and
clearly deﬁned characters, ﬁeld naturalists were attracted to Lamarckism be-
cause it seemed a good explanation for the adaptive patterns they observed
in the ﬁeld, whereas many paleontologists were adherents of orthogenesis
because of the linear patterns they discerned in the fossil record. Put dif-
ferently: the speciﬁc interpretation of evolution of each one of these disci-
plines was distorted or lopsided by their speciﬁc, disciplinary perspective.
The so-called “population genetics approach to evolution” (Michod 1981,
p. 3) which interprets evolution in terms of changes in the relative frequency
of alleles, can, as already indicated, be considered one of these pre-paradigmatic
theories or approaches, even though we normally don’t interpret it as such from
our modern perspective, since it was this approach of evolution that eventually
won the pre-paradigmatic competition.11

The Complex Construction of the MS

3.2.
The question of how the MS was constructed, remains “one of the most vexing
problems in the history of biology” (Smocovitis 1996, p. xii; Amundson 2005,
section 8.3), despite the existence of a veritable ‘Synthesis Industry’ (e.g.,
Mayr and Provine 1980; Mayr 1982; Smocovitis 1996; Bowler [1989]
2009; Gould 2002). From a Kuhnian perspective, its construction is, like
the pre-paradigmatic phase, classic and atypical at the same time.

We already know in what way the genesis of the MS can be called a Kuhnian
process: it is based on one of the pre-paradigmatic approaches of evolution
(population genetics). However, due to the highly multidisciplinary nature
of evolutionary biology, its construction was at the same time also atypical.
It is not difﬁcult to understand why: since the victorious evolutionary approach
was that of one particular biological discipline, the construction of the MS also
had to encompass a synthesis of information from other biological disciplines
(hence its name). Theodosius Dobzhansky’s Genetics and the Origin of Species
(1937) started this synthesis process and inspired a small number of other
biologists from various disciplines (Huxley 1942; Mayr 1942; Simpson
1944; Rensch 1947; Stebbins 1950) to elaborate a more or less coherent inter-
pretation of evolution (i.e., the MS). This heterogeneous set of books in its turn
inspired many other biologists and started a neo-Darwinian research tradition.
They clearly constitute an example of a Kuhnian “universally recognized sci-
entiﬁc achievement” and at the same time also form yet another reﬂection of the

11. Another pre-paradigmatic theory that is not always recognized as such is August

Weismann’s neo-Darwinism.

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Evolutionary Biology through a Kuhnian Prism

extreme disciplinary heterogeneity of evolutionary biology: Kuhn’s paradigms
are, typically, elaborated in single books.12

The Turbulent Normal Science Period

3.3.
This complex, binary construction of the MS (constriction and synthesis) helps
to explain why it has turned out to be somewhat schizophrenic in its perfor-
mance as a paradigm: functional and dysfunctional at the same time.

3.3.1. A Run-of-the-Mill Paradigm. Above, we saw how the MS facili-
tated the start of a still ongoing phase of normal science. It is in that Kuhnian
light that we can understand a mystery, pondered by John Maynard Smith.
He wondered why a major characteristic of evolutionary biology since 1960
has been the attempt to develop Darwinian explanations of seemingly anom-
alous biological phenomena such as sex, ageing, sexual ornamentation, and,
“most important of all, cooperation” (Maynard Smith 1994, p. x; see also
Mayr 2004, p. 139). From a Kuhnian perspective, there is no mystery what-
soever: as long as scientists still discuss the fundamental nature of the natural
phenomenon that they study (electricity, light, evolution, …), there does not
exist much interest in that kind of esoteric puzzles. It is only once a paradigm
emerges that they attract widespread attention or that they become the ‘main
business’ of scientists and thus can inspire a crisis and a scientiﬁc revolution.
One of the main Kuhnian puzzles in evolutionary biology, next to coop-
eration, is the existence of sex. In the preface to his Sex and Evolution (1975),
George Williams even claimed that the prevalence of sexual reproduction in
higher plants and animals is “inconsistent with current evolutionary theory”
and that, consequently, there was “a kind of crisis at hand in evolutionary
biology” (p. v). With his book, he wanted to propose “minimal modiﬁcations
of the theory in order to account for the persistence of so seemingly maladap-
tive a character” (p. v). Sex may have been (and still be) the “queen of prob-
lems in evolutionary biology” (Bell 1982, p. 19), but it is (contra Williams)
not serious enough to have inspired (or inspire) a true, Kuhnian crisis. Rather,
it is an example of the “omnipresent concomitant of normal science” that, as
we saw, most anomalies are. The problem is not that it cannot be explained
but rather that we are still not certain which evolutionary explanation(s) is (or
are) correct (see, e.g., Barton and Charlesworth 1998).

As stated above, a paradigm implies “a new and more rigid deﬁnition of the
ﬁeld. Those unwilling or unable to accommodate their work to it must proceed
in isolation (…)” (Kuhn 1962, p. 19). Entire schools can thus endure in

12. Kuhn refers to “Aristotle’s Physica, Ptolemy’s Almagest, Newton’s Principia and
Opticks, Franklin’s Electricity, Lavoisier’s Chemistry, and Lyell’s Geology—these and many other
works served for a time implicitly to deﬁne the legitimate problems and methods of a re-
search ﬁeld for succeeding generations of practitioners” (1962, p. 10).

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“increasing isolation from professional schools” (Kuhn 1962, p. 19). Examples
of such groups are or were adherents of astrology (once an integral part of
astronomy) and of “romantic” chemistry (i.e., alchemy). Something similar
happened to traditional or “naïve” group selectionists (i.e., adherents of the idea
that prosocial behaviors will automatically spread in populations because they
beneﬁt groups), once the ﬁrst generation of biologists, educated in the new par-
adigm, started their career. The fathers of population genetics (Ronald Fisher,
John B. S. Haldane, and Sewall Wright) had each considered the question of
group selection or multilevel selection, but only brieﬂy. Consequently, group
selection did not really form an integral part of the original MS. In the wake of
its emergence, individual selection even became the norm. As Borrello puts it:

Essentially, the neo-Darwinian paradigm held that all evolution is due to
the accumulation of small genetic changes, guided by natural selection.
The theory of group selection clearly did not ﬁt within this deﬁnition,
and the era of peaceful coexistence came to a decisive end. (2005, p. 45)

Ironically, V. C. Wynne-Edwards (1962), who is generally seen as one of the
main modern proponents of the idea of naïve group selection, was also inspired
by the MS (Borrello 2003): he saw in its focus on populations a way of thinking
about evolution that was in line with his own research interests (in populations
and population regulation). It is particularly through reactions to his work that
individual-level adaptation or selection was turned into “a core concept of the
modern synthesis” (Borrello 2003, p. 531) and group selection into something
of a (non-paradigmatic) heresy (to many evolutionists, it still is suspect or of
little importance, in spite of its resurrection, in recent decades, see, e.g.,
Borrello 2005). Within-group (individual) selection was assumed to be almost
always stronger than between-group selection. For example, in a letter in
Nature, entitled “Group Selection and Kin Selection” (1964), Maynard
Smith directly challenged group selection. “I will contrast group selection,”
he wrote, “with something I will call kin selection. Kin selection has been
treated by Haldane and Hamilton” (quoted in Segerstraele 2001, p. 63).

Likewise, in the preface to the 1996 edition of Adaptation and Natural
Selection: A Critique of Some Current Evolutionary Thought ([1966] 1996),
Williams refers to “what seemed to be a pervasive inconsistency in the use
of the theory of natural selection” (p. ix). Adaptive changes are, in theory, only
caused by the selection of individuals. However, in practice, many biologists
continuously spoke of group adaptations, which evolved through group
selection and required individual organisms to forgo their own interests for
the greater good of the group or the species. A lecture by the renowned
ecologist and termite specialist A. E. Emerson about beneﬁcial death, the idea
that senescence evolved for the beneﬁt of the species, which Williams
attended while on a teaching fellowship at the University of Chicago, was

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Evolutionary Biology through a Kuhnian Prism

the “triggering event” (p. ix). If that was acceptable biology, he rather wanted
to become an insurance salesman. In the introduction, he wrote that, with
some minor qualiﬁcations, “it can be said that there is no escape from the
conclusion that natural selection, as portrayed in elementary texts and in most
of the technical contributions of population geneticists, can only produce
adaptations for the genetic survival of individuals” (pp. 7–8).

Dawkins, who was “greatly inﬂuenced” by “Williams’s great book”
(Dawkins 1976, p. 11) wanted with his own more popular book The Selﬁsh Gene
(1976) to “examine the biology of selﬁshness and altruism” (p. 1)—i.e., pro-
pound recently published ideas about the evolution of social and particularly
altruistic behaviors—, and to falsify the “erroneous assumption” (p. 2) behind
books like Konrad Lorenz’ On Agression (1966) and Robert Ardrey’s The Social
Contract (1970) “that the important thing in evolution is the good of the species
(or the group) rather than the good of the individual (or the gene).” The reason
why he sounded so “evangelist” in The Selﬁsh Gene, he later explained (Dawkins
1979), is that he wanted to rebut the still popular group selection mechanism
by bringing in gene selectionism as an alternative.

3.3.2. Not-So-Normal Normal Science. In some ways, though, evolution-
ary biology was and/or is an atypical paradigmatic science. For example, its
professionalization proceeded surprisingly slowly (Antonovics 1987).13 Also,
the MS was soon challenged by an ever-increasing plethora of alternative evo-
lutionary models (Huneman and Walsh 2017). Or, as Bowler puts it: the foun-
ders of the MS, who had hoped that their paradigm would inspire a long
period of what Thomas Kuhn called normal science, “were to be disappointed.
The diversity of new theories has never matched that of the ‘eclipse of Darwin-
ism’, but it has been possible to articulate divergent perspectives within the
narrower frame of reference established by the synthesis” ([1989] 2009, p.
347). Evidently, not all proponents of those ‘divergent perspectives’ would
agree with that last claim. Some scholars even believe that the original MS
has ceased to exist and that “there is no longer a single, unifying ‘Darwinian
evolutionary theory’” (Kutchera 2013, p. 544).

In any case, as we explained elsewhere in more detail (Tanghe et al. 2018),
this unusual degree of disagreement among modern evolutionists can, once
again, only be explained by the extreme multidisciplinarity of evolutionary

13. Antonovics (1987) even believes that the MS “had little direct effect on the progress
of evolutionary biology as a discipline (…)” (p. 321). After the foundation of the journal
Evolution: An International Journal of Organic Evolution (1946), no other evolutionary journals
appeared until the 1970s; for a long time, textbooks were absent and evolutionary biology
was, as late as 1987, rarely thought of as a discipline in its own right (i.e., there was a scarcity
of international congresses, departments or university programs and institutional organiza-
tions, exclusively dedicated to evolution, as well as a painful lack of funding). See, in this
respect, Ruse (2009a).

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biology. It not only, and evidently, sharply increases the chance of disagree-
ments (among practitioners of various evolutionary disciplines) but was,
through the complex construction of the MS, also antecedent to four prob-
lematic and contentious features of this paradigm. First, since the approach of
evolution that became the basis of the MS was that of one evolutionary dis-
cipline (population genetics), the MS was inevitably somewhat lopsided (i.e.,
quite gene-centric). As aforementioned, it even identiﬁed evolution with its
genetic dimension and, more particularly, with changes in allele frequencies.
Dobzhansky (1937, p. 11) put it as follows: “Since evolution is a change in the
genetic composition of populations, the mechanisms of evolution constitute
problems of population genetics.”

Furthermore, the additional synthesis of data and concepts from a variety
of other biological disciplines around this genecentric approach of evolution
led to two other problems. That synthesis was not only incomplete—or, as
Eldredge (1985) put it, unﬁnished—but also imperfect. It was incomplete
because not all life sciences (fully) participated in the synthesis—some of
course only came into being after the establishment of the MS—and it was
imperfect because of two related reasons: practitioners of several biological
disciplines believed that their ﬁeld should have made a more substantial con-
tribution to the MS and the MS did not resolve the old, pre-paradigmatic
conﬂict between an organism-focused and a gene-focused approach to the
study of (evolving) life (Mayr 1982, pp. 540–50), as soon became painfully
clear in the comments of some of its main architects (see, e.g., Mayr 1959,
p. 13). Lastly, due to its convoluted construction (a lopsided constriction,
followed by a problematic synthesis), the MS is also surprisingly fuzzy, as
has often been pointed out: it is a ‘moving target’ and meant and means dif-
ferent things to different people and, particularly, to practitioners of different
biological disciplines (see, e.g., Craig 2015, p. 255).

This brings us back to the EES. As we pointed out above, only a minority
of its proponents sees it as a potential new paradigm. It can be conceived as an
alternative Lakatosian research program but it can also, and maybe more
appropriately, be interpreted as a classic, Kuhnian reformulation of the
modern, extended MS by more organism-focused biologists. This interpreta-
tion is corroborated by Müller’s (2017) remark that the new way of thinking
about evolution of the EES is historically rooted in the not-so-new “organicist
tradition” (p. 8). The EES is, in any case, at least partly inspired by a strong
aversion, among organism-focused biologists, for the baked-in, greedy gene-
centrism of the MS. This is the theoretical problem that they want to solve
through their reformulation.

Laland et al., for example, argue that “important drivers of evolution, ones
that cannot be reduced to genes, must be woven into the very fabric of evolu-
tionary theory” (2014, p. 161). Genes are not causally privileged as programs or

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blueprints, but are rather “parts of the systemic dynamics of interactions that
mobilize self-organizing processes in the evolution of development and entire
life cycles” (Müller 2017, p. 7). Also, organism-focused biological disciplines
like Laland’s ethology or Müller’s developmental biology and the somatic,
ecological and epigenetic phenomena that they study, occupy a much more
important place in the EES than they do in the MS. The current friction
between the MS- and the EES-community can, in this sense, be seen as a con-
tinuation or resurgence of the old conﬂict between an organism-focused and a
gene-focused approach to the study of (evolving) life.

4. Why Was This Kuhnian Analysis Not Done Before?
If, as we have argued, Kuhn’s developmental model of sciences is partly dis-
cernible in the history of evolutionary biology and if a Kuhnian analysis of
that history is heuristically interesting or rewarding, why then has this kind
of analysis not been done before? We already know one possible reason: there
doesn’t seem to be place in Kuhn’s scheme for the publication of On the Origin
of Species (1859) (4.1). Furthermore, a Kuhnian analysis of the history of evo-
lutionary biology also constitutes an unusual application of The Structure (4.2)
and an odd, philosophical kind of historiography to boot (4.3). This multi-
faceted explanation involves a deep dive in the Kuhn literature but much of
that more esoteric information will be discussed in extended footnotes and
can be ignored by readers who are only interested in the gist of this section.

4.1. On the Origin of Species (1859): A Kuhnian Anomaly
Several scholars have investigated whether On the Origin of Species (1859) can be
considered an example of a Kuhnian revolution.14 The standard result of these
studies is almost invariably negative: Charles Darwin’s theory of evolution may
have been hugely successful and inﬂuential but most scholars do not believe
that it caused a Kuhnian revolution.15 This is corroborated by the present anal-

14. Kuhn seems to have believed so. In his preface, he points out that “Far more historical
evidence is available than I have had space to exploit below. Furthermore, that evidence comes
from the history of biological as well as of physical science. My decision to deal here exclusively
with the latter was made partly to increase this essay’s coherence and partly on grounds of
present competence” (Kuhn 1962, p. xi). However, in later chapters, he does refer to evidence
from the history of biology. For example, he refers to parts of biology “—the study of heredity,
for example—” where “the ﬁrst universally received paradigms are still more recent (…)”
(p. 15). In various places, he also refers to Darwin and/or On the Origin of Species (1859) (Kuhn
[1962] 1970, pp. 20, 151, 171–2, 180). It seems clear that he considered the emergence of the
idea of evolution through natural selection to be an example of a scientiﬁc revolution (see also
table 1 in Wray 2011, pp. 18–19).

15. Greene ([1971] 1981, p. 31), for example, examines “the developments leading up
to the Darwinian revolution in natural history to see to what extent they ﬁt the pattern of
historical development described in Kuhn’s book.” His conclusion is that the Darwinian

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ysis as it implies that On the Origin merely initiated the pre-paradigmatic phase
in the history of evolutionary biology and simply does not ﬁt within Kuhn’s
original scheme (since that scheme doesn’t mention the various ways in which
pre-paradigmatic phases in the history of scientiﬁc disciplines start).

This implication of our analysis, in turn, is perfectly in line with yet another
strand in the Kuhn literature, namely the criticism that his developmental
scheme does not “capture some crucial episodes in the history of science
(…)” (Politi 2018, p. 2284).16 Bird refers in this respect to the discovery of
the structure of DNA. It was “clearly revolutionary in [its] consequence for

revolution “overthrew the static view of nature and natural history but failed to establish a
clear-cut paradigm in its place” (p. 54), and that “nothing approaching a ‘Darwinian’ para-
digm became established until the 1930s (…)” (p. 53). He set the pattern for subsequent
studies. For example, Mayr (1972, p. 988) claims that “the Darwinian revolution does not
conform to the simple model of the scientiﬁc revolution, as described, for instance, by T. S.
Kuhn” (see also Mayr 1994, 2004, p. 165; Wilkins 1996, p. 696; Hodge 2005; Smocovitis
2012). Ruse has developed a somewhat more nuanced view. Kuhn’s analysis taken head on
“fails on the Darwinian example” (Ruse 2005, p. 13) but in some ways, it is legitimate to call
the publication and success of On the Origin (1859) a (Kuhnian) revolution. Darwin’s notion of
evolution through natural selection was paradigmatic and constituted a different world view
(Ruse 1989). There were “different paradigms if you will” (Ruse 2009b, p. 10045), “but note
that it is not just a question of evolution or not evolution, and certainly not of selection or not
selection.” There is also a kind of incommensurability between functionalism (which stresses
the function of organic structures) and formalism (which emphasizes the form of organic struc-
tures). Darwin was a functionalist but Thomas H. Huxley, for example, was a hardline
formalist: “I would argue that in a real sense we have Kuhnian paradigm differences operating
here. Different visions, unable to bridge the gap” (Ruse 2009b, p. 10045), even though this
sense of paradigm “does not ﬁt exactly with the senses of paradigm found in the Structure of
Scientiﬁc Revolutions” (Ruse 2009b, p. 10045). Lastly, the philosopher Massimo Pigliucci
(2012) believes that the only time in the history of biology when a Kuhnian transition occurred
was “when Darwin’s original theory replaced the dominant ‘paradigm’ of the day, Paley-style
natural theology” (p. 46) although this “paradigm shift did not really happen within the con-
ﬁnes of an established science” (p. 56) (see also Pigliucci 2009). This is indeed correct and also
highly problematic as genuine paradigm shifts per deﬁnition happen within the conﬁnes of an
established or mature science (since the creation of a paradigm is the hallmark of a mature
science).

16. One of three traditional criticisms of Kuhn’s central distinction between normal
science and revolutionary science is that these two categories “fail to provide us with the con-
ceptual resources necessary to understand the variety of changes that occur in science” ( Wray
2011, p. 24). The other two criticisms are that the various changes that Kuhn regards as
revolutionary (1) are a mixed lot and do not belong in one and the same category and (2)
are not different in kind from changes that occur during the normal science phase (see Wray
2011, pp. 21–4; see also Casadevall and Fang 2016). Wray (2011, pp. 29–33) defends Kuhn’s
later, revised account of scientiﬁc revolutions against these criticisms. In this analysis, Kuhn
doesn’t refer to paradigms anymore but to conceptual taxonomies or lexicons. Thus, he came
full circle as he ﬁrst had assigned the functions of a paradigm to “language and pre-
dispositions” (Reisch 2016, p. 23). For a discussion of the applicability (or non-applicability)
of that revised account to the history of biology, see Keller (2012).

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biochemistry and molecular genetics” but “simply does not ﬁt Kuhn’s descrip-
tion of scientiﬁc development—it originated in no crisis and required little or
no revision of existing paradigms even though it brought into existence major
new ﬁelds of research” (Bird 2000, p. 60).

We agree with Politi that the discovery of the structure of DNA was, con-
tra Bird, not revolutionary in its consequence for molecular genetics “simply
because, before such a discovery, there was not such a thing as ‘molecular ge-
netics’” (Politi 2018, p. 2284). Rather, Watson’s and Crick’s discovery was
revolutionary “precisely in virtue of its role in the creation and establishment
of molecular biology (…)” (Politi 2018, p. 2284). Likewise, the discovery of
the deep history of Earth (Rudwick 2005), subatomic particles, the true
nature of viruses (Méthot 2016) and evolution was revolutionary in virtue of
the role it played in the establishment of geology, quantum mechanics, virology
and evolutionary biology, respectively.17 Consequently, scientiﬁc discoveries can
be revolutionary because of the impact they have within an established science
or because of the seminal role they play in the emergence of a new science.

Undoubtedly, the aforementioned fact that Kuhn treats the period in the
history of a science that precedes the emergence of a paradigm and the normal
science phase (annex scientiﬁc revolutions) in a somewhat stepmotherly
fashion, is an important reason why he did not foresee a separate phase or
speciﬁc place in his developmental scheme for these important, primordial
occurrences in the history of some sciences. When, in his later work and in
the context of his evolutionary philosophy of science (Marcum 2015, ch. 6),

17. Evidently, Earth was already empirically studied before the discovery of its deep his-
tory but that was not geology in the modern meaning of the word. It is only when it became
clear that Earth has a deep history that the scientiﬁc study of that long and convoluted history
(i.e., geology) could start in earnest (Rudwick 2005). As we saw in note 3, Kuhn (1962, p. 15)
refers in this respect to the work of the Scottish geologist James Hutton and portrays him as the
father of historical geology. See, however, Rudwick (2005, p. 172): “A sense of the history of the
earth, whatever its source may have been, certainly did not come from Hutton.” Likewise,
evolutionary biology could only emerge as a new scientiﬁc ﬁeld once Darwin put the notion
of evolution on the late-nineteenth century scientiﬁc agenda. Of course, the idea that popu-
lations can undergo certain transmutations was centuries old. By contrast, the idea of evolution
(i.e., life as we know it has developed from a very simple beginning) only dates from the 18th
century. By scientiﬁcally consolidating that hitherto vague, unorthodox and philosophical
notion, On the Origin played, without a shred of a doubt, a crucial role in the emergence of
the science of evolutionary biology. It is not only a milestone in the history of (evolutionary)
biology but maybe even a unique achievement in the history of science as it, at the same time,
also identiﬁed the mechanism behind adaptive evolution (i.e., natural selection). Also, it was
‘consilient’—i.e., it connected and explained data from a wide array of areas (Ruse 1975)—,
reshaped the way we see the world (Langdon 2016) and inspired the concrete research of some
biologists or naturalists (Bowler 1996). That, however, is not enough to assign it the status of
(new) paradigm. It merely marked the true start of the history of evolutionary biology and
subsequently became an important source of inspiration for the MS.

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he ﬁnally did devote special attention to the birth of new sciences, he focused
on specialization or specialty formation (the equivalent of biological specia-
tion, see Wray 2011, ch. 7; Nickles 2017, sections 4 and 6.2; Politi 2018).18
In his Rothschild lecture, he even seemed to replace the notion of scientiﬁc
revolutions with the notion of ‘scientiﬁc speciation’. The episodes that “he once
described as scientiﬁc revolutions,” he declared, “are intimately associated with
(…) speciation” (Kuhn, cited in Nye 2012, p. 561). Also, as we shall see in the
next subsection, his developmental scheme wasn’t even meant to be complete
or perfect.

4.2. An Unorthodox Use of Structure
There are at least three reasons why an interpretation of the history of evolu-
tionary biology through the prism of Kuhn’s historical developmental model
constitutes an unusual way of applying The Structure to the study of science. To
begin with, that model was a “highly schematic sketch” (Kuhn, cited in Si-
gurdsson 2016, p. 27) that wasn’t meant to be applied this way:19

I’ve always said, assimilate this point of view and this way of doing it,
and then see what it does for you when you try to write a history, but

18.

In Structure (1962, p. 169), Kuhn had already suggested that increasing specializa-
tion plays an important role in the historical development of sciences but he saw it as a con-
sequence of scientiﬁc progress. Later, he regarded specialization as the cause of scientiﬁc
progress and emphasized its importance as an alternative for scientiﬁc revolutions within ex-
isting sciences, although he never made this idea the focus of a speciﬁc paper or book ( Wray
2015, pp. 177–80). Nickles (2017) points out that he still gave little attention to the reverse
process of hybridizations between various ﬁelds. Also, some sciences originated partly or
completely in pre-scientiﬁc practices (particularly natural history and natural philosophy).

19. As Hacking (2012, p. xi) puts it: Kuhn “thought of Structure as more of a book
outline than a book.” Structure originated as a contribution to the International Encyclopedia of
Uniﬁed Science. Therefore, Kuhn presented his views “in an extremely condensed and schematic
form” (Kuhn 1962, pp. xlii–xliii). In what seems to have been the ﬁrst version of the ﬁrst
chapter of Structure, “Kuhn reckoned the monograph would have only ﬁve chapters (and about
eighty pages, according to the editors’ suggestion)” (Pinto de Oliveira 2017, p. 748). Unfor-
tunately, he never wrote a longer version of his theory. Instead, in his later work, he focused on a
few components of his 1962 sketch, mainly the notion of paradigms, the notion of incommen-
surability, and the passage about evolutionary epistemology (Renzi 2009): Structure truly “con-
tinued to dominate Kuhn’s thinking until the end of his life (…)” (Marcum 2015, p. 53). As
Hoyningen-Huene (2015, p. 185) puts it: “A really comprehensive account of these develop-
ment (sic) remains to be written (…).” Kuhn’s own account, The Plurality of Worlds: An Evo-
lutionary Theory of Scientiﬁc Development, remained unwritten (see Hoyningen-Huene 2015).
Holton (2016, p. 35) believes that “he was not able to publish the work. And that, in my view,
was a chief source of Tom’s internal state of dismay, especially in his last decade, as he was
trying to reach the new, high professional identity level he had set for himself.” This maybe
also explains why the one surviving possible editor of the manuscript, designated by Kuhn,
James Conant, still announces the work as “soon to be completed,” a quarter of a century after
Kuhn passed away. See: https://philosophy.uchicago.edu/faculty/conant.

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don’t go out looking at history to see whether this is true or false, to test
the ideas. The only test of the ideas, at least at this level of development,
is going to be whether having assimilated those ideas, you see the
material usefully different. But it’s not going to be “Can you always
locate the paradigm, can you always tell the difference between a
revolution and a normal development?” It’s not meant to be applied
that way. (Kuhn, cited in Sigurdsson 2016, p. 27)

What did Kuhn mean by “this point of view and this way of doing” history of
science? With Structure, he wanted to contribute to a “historiographic revolu-
tion in the study of science, though one that is still in its early stages” (Kuhn
1962, p. 3; see also Pinto de Oliveira 2020). In the introduction, he pointed
out that the point of departure of the traditional, Whiggish history of science
was the constellation of facts and theories in modern science textbooks. The
history of science then constituted of the piecemeal and gradual addition of
items to this modern constellation and the main task of historians was to
determine when each item had been added and by whom. Science was com-
pared to “an ever-growing ediﬁce to which each scientist strives to add a few
stones or a bit of mortar” (Kuhn, cited in Pinto de Oliveira 2017, p. 748). By
contrast, according to the revolutionary new image of the history of science
that he wanted to help elaborate, science did “not progress by adding stones
to an initially incomplete structure, but by tearing down one habitable struc-
ture and rebuilding to a new plan with the old materials and, perhaps, new
ones besides” (Kuhn, cited in Hufbauer 2012, p. 459).20

He wanted to contribute to this new image in two different ways. The ﬁrst
way was by elaborating a non-Whiggish developmental model of sciences and
thus promoting an non-Whiggish way of doing history of science. The reason
why scientiﬁc revolutions occupied a central place in that model is of course
that they were at the heart of the new, non-gradual image of science (although
it should immediately be added that Kuhn believed that most science is of
the ‘normal’, non-revolutionary kind).21 In what seems to have been the ﬁrst

20. He had ﬁrst come into contact with that “new image” in 1947, as a graduate student
in physics. That was the year when, at Bernard Cohen’s recommendation, he read Alexandre
Koyré’s Études Galiléennes (1939). Koyré was the scholar who Kuhn deemed, more than any
other, “responsible for (…) the historiographical revolution” (Kuhn 1962, p. 67). For Koyré’s
signiﬁcant inﬂuence on Kuhn, see, e.g., Omodeo (2016). 1947 was also the year that James B.
Conant, president of Harvard University, invited Kuhn to become an assistant for his course on
the history of science for upper-level undergraduates. For the special relationship between
Kuhn and Conant (and the Cold War political/ideological background of Structure), see,
e.g., Fuller (2000), Reisch (2016) and Omodeo (2016).

21. As Bird (2000, p. 33) puts it: “It is appropriate then that despite the title, The
Structure of Scientiﬁc Revolutions, half of that book does not concern revolutionary science at
all.” One can indeed state that Kuhn’s title is misleading. Scientiﬁc revolutions may be the

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draft of the ﬁrst chapter of Structure, he even introduced his main argument by
making a comparison with revolutions in the history of art.22 This is a second
reason why our application of Structure is unusual: most applications have
focused on the question whether Kuhn’s all-important description of the
structure of scientiﬁc revolutions ﬁts a particular revolution (e.g., the Darwinian
revolution, the chemical revolution or the plate tectonics revolution), rather
than on the question whether a particular science follows his developmental
scheme in its historical development (see Marcum 2015, ch. 7).

A second goal of Kuhn was to delineate the philosophical consequences of
the new image of science for he believed that the old, positivist philosophy
that had been associated with the Whiggish, development-by-accumulation
historiography had to be replaced with a new philosophy of science.23 He
even thought of The Structure of Scientiﬁc Revolutions (1962) “as addressed pri-
marily to philosophers” (Kuhn 1993, p. xii).24 It is also generally deemed to

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22.

quintessential part of Structure and many of its chapters (5 out of 13) are indeed devoted to their
analysis, but his title does, nevertheless, not reﬂect very well the true purpose and content of his
book. Consequently, we don’t entirely agree with Hacking’s claim that “Structure and revo-
lution are rightly put up front in the book’s title” (2012, p. x). Nevertheless, it may have
contributed to its success. See, in this respect, Gavroglu (2016). He believes that Kuhn’s title,
and more particularly “what ‘happened’ to” the words ‘science’ and ‘revolution’ was “absolutely
decisive for the book’s success” (p. 51) outside academia. See also Nye (2016).

See Pinto de Oliveira (2017). It differed signiﬁcantly from the ﬁrst chapter of Struc-
ture, though. The title was: ‘What are Scientiﬁc Revolutions?’. He eventually abandoned the
art/science analogy (as a central element of his argument) because he was afraid he knew too
little of art as activity. That is a shame because as Kuhn ([1969] 1977, p. 340) put it himself:
Structure must be seen as a “belated product” of his “discovery of the close and persistent par-
allels” between science and art. Pinto de Oliveira (2017, pp. 758–60) discusses two passages in
Structure that highlight this analogy. Kuhn also referred to it in the postscript to the second
edition of Structure ([1962] 1970, p. 208) (see also Kuhn [1969] 1977). It helps to explain why
his book was so inﬂuential outside the domain of the history and philosophy of science. As
Pinto de Oliveira puts it: “Kuhn gives back to the other disciplines, with ‘added value’, the
idea of development with breaks” (2017, p. 761). The analogy also seems to have inﬂuenced
Feyerabend, “the other main name of the contemporary historically oriented philosophy of
science (…)” (Pinto de Oliveira 2017, p. 762). It is probably no coincidence that it was also
the two fathers of this analogy, Kuhn and Feyerabend, “who put the word incommensurable on
the table” (Hacking 2012, p. xxx).

23. As late as 1991, Kuhn pointed out, in his Robert and Maurine Rothschild Distin-
guished Lecture (“The Trouble with the Historical Philosophy of Science”) that “the problem
with the historiographic revolution was that it was unable to provide a philosophy of science to
replace the one it demolished and to account for the growth of scientiﬁc knowledge” (Kuhn,
cited in Marcum 2015, p. 25).

24. He admitted, though, that “limitations of space have drastically affected my treatment
of the philosophical implications of this essay’s historically oriented view of science” (Kuhn
1962, p. xliv). This also explains why his later attempts to develop it further were directed
“exclusively to the book’s philosophical aspects” and particularly to the “underpinnings of in-
commensurability (…)” (Kuhn 1993, p. xii). For Kuhn’s philosophical goals, see Bird (2012a).

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Evolutionary Biology through a Kuhnian Prism

have been more inﬂuential among philosophers than among historians
(Reingold 1980; Cohen 1985, p. 27; Wray 2012; Bird 2012a; Marcum
2015). This is a third and last reason why a Kuhnian interpretation of the
history of evolutionary biology can be said to be unusual or unorthodox.25

4.3. An Odd Kind of Historiography
Of course, that does not mean that our historiographical use or application of
Kuhn’s historical developmental model is completely new. It has, for exam-
ple, already been pointed out that it “ﬁts the history of the development of
geology quite well” (O’Hara 2018, p. 258). Frankel (1978), for example, in-
terprets the history of geology through the prism of Kuhn’s model. In a sim-
ilar vein, Melchert observes (or hopes) in a paper, entitled “Leaving behind
Our Preparadigmatic Past: Professional Psychology as a Uniﬁed Clinical Sci-
ence” that “[t]he regular introduction of new theoretical orientations was a
main feature of the pre-paradigmatic era in psychology, but that era has come
to an end” (2016, p. 21).

Unfortunately, though, scholars who investigate whether Kuhn’s historical
developmental model can be discerned in the history of a speciﬁc science, often
misunderstand it. Méthot, for example, points out that the science of virology
“developed its own paradigm in the mid-twentieth century by breaking with
bacteriology, along the lines van Helvoort suggested” (2016, p. 7). This was
preceded by “deep controversies in virus research in the ﬁrst half of the
twentieth century” (2016, p. 7) and followed by a period of normal science.
However, he does not seem to understand that this implies that the science of
virology broadly followed Kuhn’s developmental model (pre-paradigmatic
phase, paradigm, normal science) since he next wonders whether this process

25.

It should be added, though, that Structure was highly instrumental in the historical
turn in philosophy of science. As Michael Friedman (1993, p. 37) put it: since the publication
of Structure, “careful and sensitive attention to the history of science must remain absolutely
central in any serious philosophical consideration of science” (see also Bird 2000, p. viii). As a
matter of fact, Kuhn was professor of history of science at Berkeley when he published Structure,
even though he taught in both the philosophy and the history department. Most of Kuhn’s
own publications before 1962 were in the history of science. In 1968, six years after the pub-
lication of Structure, he still called himself “a practicing historian of science (…) I am a member
of the American Historical, not the American Philosophical, Association” (Kuhn, cited in
Hacking 2012, p. x). This explains why, of the 150 footnotes in the ﬁrst edition of Structure,
only 13 include references to philosophers, the vast bulk of the other references are to historians
(Bird 2000, p. x). To be precise, sixty percent of the sources (76/127), cited in Structure, and
ninety percent of the most cited sources, are works in the history of science ( Wray 2015). There
are only 13 citations of philosophical sources. Structure also became “the most widely read book
on the history of science” (Blum et al. 2016, p. 1) and thus “the second major milestone, after
the ﬁrst publication of the journal Isis in 1912, to mark the rise of the history of science to a
ﬁeld enjoying broad recognition beyond the narrow community of its practitioners” (Blum
et al. 2016, p. 1).

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(i.e., the establishment of virology as a mature science) can “really be called a
‘revolution’ (…) in the Kuhnian sense?”26

A second, and more fundamental problem, associated with using Kuhn’s
historical developmental model in the study of the history of speciﬁc sciences,
is that this is very much at odds with the modern historiography of science.
As Arabatzis (2016, p. 196) puts it, “Kuhn’s grand narrative of scientiﬁc de-
velopment was not well received by historians of science (…)” (see, e.g., also
Kitcher 2012, pp. 532–3). Daston explains why: the very idea of “looking for
overarching regularities in the history of science seems bizarre (…)”(2012,
p. 498). And: “Since roughly the 1990s, the focus (…) has shifted from
the streamlined to the dense and detailed; the professed aim has been to
‘complexify’ rather than simplify and to reveal variability rather than unifor-
mity” (2012, p. 498). That is why “Most historians of science no longer
believe that any kind of structure could possibly do justice to their subject
matter” (2016, p. 117).

Ironically, this is partly due to the inﬂuence of Structure.27 Its publication
was “a signal event leading to the widespread rejection of presentism in the
history of science” (Kremer 2013, p. 301) and “the starting point for a long
conversation that contributed to the view held by many historians of science
today, that science is a historically rooted and culturally embedded set of
practices” (Smocovitis 2012, p. 569). Nye (2012, p. 560) speaks of a shift,
particularly in the decades after the publication of The Structure, to the study
of the history of “the ﬁne structure of scientiﬁc practice and scientiﬁc ideas
(…).” Kuhn had always been “unwavering in his conviction that the future of
the history of science lay in history departments. (…) It would become
historicist, in the sense of situating science in its speciﬁc context, just as
the history of art (…) embedded styles and genres in particular periods
and places” (Daston 2016, pp. 117–118).28 It is in this sense that he left a

26.

See, for example, also Weimer and Palermo (1973) or Bowler (1983, p. 11). Bowler
calls Structure the best-known attempt to create a “general scheme describing the emergence of
a new theory,” sketches the emergence of a paradigm and its replacement with a new theory
and subsequently points out that there has already been some debate wether the Darwinian
revolution ﬁts this pattern.

27. As Findlen puts it, “there is an ironic coda to the inspiration that Kuhn offered to
pioneers of microhistory by writing a grand narrative, in light of the function of anomalies in
destabilizing the explanatory power of prevalent scientiﬁc theories. He inadvertently played a
role in inspiring microanalysis as a means of rethinking commonly received historical narra-
tives. By the 1990s the petite histoire had replaced the longue durée” (2005, p. 233). For an in-
vestigation of the possibility of reconstructing new ‘big pictures’ from the mass of modern,
microhistoriographical detail, see Secord (1993).

28. However, as Daston (2016) also points out: this conviction stood in sharp contrast
with his “search for structures” (p. 124). The historicisation of the history of science also

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Evolutionary Biology through a Kuhnian Prism

“remarkably substantial” (Blum et al. 2016, p. 2) legacy in the history of
science or a “stronger” legacy than sometimes now claimed (Nye 2016,
p. 292).29

By thus helping to push the history of science in an externalist, historicist
and petite histoire direction, Structure and its developmental model became, to
modern historians, a relic of a foregone era. Arabatzis (2016) categorizes it as a
contribution to “the philosophy of history of science” (p. 192), an endeavor
which “has been relatively neglected” (p. 196) by modern historians (see, e.g.,
also Bird 2000, pp. viii, 29, Bird 2012b; Shapin 2015, p. 11). Bird calls it a
work in “theoretical history” (2000) or in the “philosophy of the history of
science” (2012b). Consequently, it is in two different ways philosophical (but
also historical): it belongs to the archaic ﬁeld of the philosophy of history of
science but also aimed at developing a new philosophy of science (a ﬁeld that
it historicized). It is no wonder, then, that, ever since 1962, scholars have
struggled to categorize Structure and that even Kuhn (1993, p. xii) himself
was “often at a loss for response,” when asked what ﬁeld his book dealt with.

Conclusion

5.
There are important reasons why the here presented Kuhnian analysis of the
history of evolutionary biology hasn’t been done before. Firstly, the match
between Kuhn’s model and that history is imperfect; secondly, it is, for
several reasons, an unusual way of applying Kuhn’s theory to the study of
(a) science; and lastly and maybe most importantly, modern, complexifying,
and petite histoire (as opposed to ‘big history’) historians of science are generally
not interested anymore in the search for overarching regularities in the history
of science (to the extent that they are still interested in exploring its internal
history at all).

It cannot be denied, though, that Kuhn’s developmental scheme is to a cer-
tain extent discernible in the history of evolutionary biology and that a Kuhnian
interpretation of that history yields very intriguing insights. It sheds somewhat
new light on On the Origin of Species (1859), gives us a better and less misleading
term for the so-called ‘eclipse of Darwinism’, and particularly provides us with a

resulted in the denial, by historians, of the splendid isolation or autonomy which Kuhn saw as
a deﬁning characteristic of science. On Kuhn’s internalism, see Bird (2012b).

29. For concrete examples of the sometimes-subtle ways in which Kuhn inspired histo-
rians, see Buchwald (2016) and Rudwick (2016). Structure also transformed a growing concern
for the Scientiﬁc Revolution of the seventeenth century into “a research program directed
toward individual small-scale revolutions in the sciences” (Cohen 1985, pp. 403, 23; see also
Daston 2012, p. 497). Thanks to Kuhn, historians also learned to appreciate the ‘losers’ of
scientiﬁc revolutions (they often resisted new paradigms for good reasons) and the tradition,
tacit knowledge and rigid forms of training that are antecedent to the practice of science
(Arabatzis 2016, p. 196).

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Perspectives on Science

new way of thinking about the genesis and nature of the MS, about the reasons
for the chronic discontent with this paradigm and about the epistemic status of
the EES. Our Kuhnian, comparative analysis has also revealed that the extreme
multidisciplinarity and heterogeneity of the science of evolutionary biology is an
underestimated ﬁl rouge and complicating factor in its rich and complex history.
In conclusion and in answer to the question that heads this paper: interpreting
that history through a Kuhnian prism may be somewhat unorthodox but it
most deﬁnitely makes sense.

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