Philosophy of Biology

Philosophy of biology is the branch of philosophy of science that deals with

biological knowledge. It can be practiced not only by philosophers, but also by scientists
who reflect on their own work. The distinctive mark of philosophy of biology is the effort
to achieve generalizations about biology, up to various degrees. For instance, philosophy
of biology makes biology relevant to classic issues in philosophy of science such
as causation and explanation, chance, progress, history, and reductionism. It also works to
characterize how knowledge is acquired and modified in different areas of biology, and
sometimes to clarify the criteria that demarcate science from non-science.
Philosophy also performs constructive criticism of biology. For example, it has an
important role in analyzing cases of “naturalization”—when science becomes able to
study issues that traditionally were the exclusive domain of philosophy. The life sciences
and their objects are changing and growing exponentially. A challenge for philosophy of
biology is thus to keep the pace, not only with new knowledge modifying long-standing
ideas (for example, the “Tree of Life”), but also with new scientific practices and
unprecedented kinds of data. Accordingly, philosophy of biology is constantly provoked
in shifting its own methods and attention. In some cases, philosophy of biology can aid
the life sciences to reach their goals, by means of conceptual analysis, linguistic analysis,
and epistemological analysis.
Hybridizations and intersections between scientific fields are particularly conducive to
philosophical considerations. Contemporary examples are ‘EvoDevo’ (the recent
integration between development and evolution) and ‘cultural evolution’ (an approach
to cultural change inspired by evolutionary biology). Theses and analyses of philosophy
of biology are often entwined with history of biology and with the history of evolution.
Finally, philosophy of biology can elaborate messages and general views out of biology,
and has a crucial role in caring for how science is publicly interrogated and
communicated.

Table of Contents
1. Introduction
2. General Issues in Philosophy of Biology
1. General Problems in Philosophy of Science, as Seen in Biology
2. A General Picture of Biology
3. A General Picture of Science
4. Generalization as a Possible Distinctive Feature of Philosophy with
Respect to Biology
b. Philosophy Flanking Biology
1. Clarifying Taxonomy, Classification, Systematics, Phylogeny,
Homology
2. Formulating Natural Selection
b. Who Can Do Philosophy of Biology?
1. Philosophical Biologists
1. Mayr and Population Thinking
 2. Gould and Adaptationism
ii. Philosophical Issues Naturalized
1. An Example: The Biology of Morality
2. Philosophy Versus Naturalization?
b. Philosophy Bringing the Life Sciences out of Their Research Context
i. Philosophy of Biology at Intersections
ii. Biology’s Critical Friend
iii. Developing Messages from Biology
b. Scientifically Up-to-Date Philosophy
i. Questioning Influential Ideas
ii. Understanding New Scientific Practices
iii. Rethinking the Philosophical Approach from New Ways of Doing
Science
b. History and Philosophy of Biology
c. Conclusion
d. References and Further Reading
i. Cited Examples
ii. Classics
1. First Generation
2. Second Generation
ii. Contemporary
1. Reviews
2. Some Monographs
ii. Anthologies and Textbooks
iii. Journals
1. Dedicated
2. Generalist
ii. Organizations
iii. Online Resources
1. Introduction
According to several reconstructions of the history of philosophy of biology, the field
emerged gradually in the 1960s with a first generation of self-identified philosophers of
biology, especially Morton Beckner, David Lee Hull, Marjorie Grene, Kenneth Schaffner,
Michael Ruse, and William C. Wimsatt. As an explanation for such branching of
philosophy of science, some philosophers put forth the decline of logical positivism in the
1960s and 1970s. For others, logical positivism did not actually decline, and anyway it
had never suppressed philosophy of biology (Callebaut 1993). At times, the ‘official’
chronology gets questioned. For Byron (2007), proper philosophy of biology was already
there in early philosophy of science, since the 1930s, as shown by a bibliometrical
analysis. The most quoted philosopher in this article is David Lee Hull (1935-2010). He
is a noncontroversially important figure in the founding generation of philosophers of
biology. His meta-reflective papers “What philosophy of biology is not” (1969) and
“Recent philosophy of biology” (2002) are particularly useful.
Philosophy of biology turned into a professional subdiscipline since the mid-1970s, with
a ‘second generation’ of philosophers, the most cited being Ronald Amundson, John
Beatty, Robert N. Brandon, Richard Burian, Lindley Darden, David J. Depew, John
Dupré, James R. Griesemer, Philip Kitcher, Elisabeth A. Lloyd, Alexander Rosenberg,
Elliott Sober, and Bruce H. Weber. Some of them were experienced philosophers who
progressively shifted to biological issues. The first journal partially devoted to
philosophy of biology – History and Philosophy of the Life Sciences – began to be published
in 1979, and in the mid-1980s the discipline was fully established. Specialized journals
flourished. In the early 2000s, a growing number of scholars, institutions, and journals
specialized in philosophy of biology, and the discipline gained more and more room in
scientific books, journals, and conferences (see the resources at the end of the article).
As we shall see, philosophy of biology provides accounts of biological knowledge, asking:
how are explanation, causation, evidence and other epistemological primitives
elaborated in the explanations that are typical of biology, such as natural selection,
genetic drift, and homology? Does biology differ from other sciences? How? And how do
we understand the epistemological diversity across different branches of the biological
sciences? Philosophy of biology also considers whether biology may contribute to
redefine classical demarcations   of science from other forms of knowledge and human
creation.

Philosophy of biology can be seen as a possible aid for scientific advancement in the life
sciences. Contributions of philosophers were widely appreciated by scientists, for
example, in the areas of classification, taxonomy, and related activities, and in the
abstract formulation of natural selection in the development of biology after Darwin.
Scientists themselves may reflect philosophically on their own field of research,
justifying and correcting their practices, or denouncing biases and transformations in
their own community. Concepts, such as ‘adaptation’ or ‘species,’ are underlain by
complex, inferential structures that can be revealed and sometimes criticized by
philosophical analysis. Multiple and conflicting meanings may be uncovered and
systematized to help the progress of science and to develop more general messages.

Phenomena studied by biology make this science particularly sensible and interesting
for philosophy. Humans are organisms, and quite a few fields of biology have potential
or direct implications for our self-understanding. Interesting philosophical debates have
stemmed, for example, since the 1970s from the provocative proposal of a
‘sociobiological synthesis’; such synthesis claims to provide evolutionary explanations
for human prosocial (and anti-social) behaviors that were traditionally covered by
ethics. Philosophy overcame mere self-defensive attitudes, and its important role lied in
epistemological analysis and in deep reflections on the limits and conditions of
naturalization, which may be understood as the transition of a problem into the domain
of empirical science. Neurobiology offers a particularly fertile ground for reflections
about how human phenomena can be related to, or even explained by, biology. And how
should a philosophical field like moral philosophy take biology into account? (For more
on the topic of the naturalization of morality, for example, see ethics.)
Philosophy of biology may study and support the interaction among different life
sciences, as in the case of evolutionary developmental biology, where workers claim to
be reuniting genetics and evolution with embryology, recomposing a historical divide in
biology. How do different research traditions integrate or replace each other? This
question illuminates classic issues such as progress and scientific change with new light.
Philosophy of biology also monitors the natural hybridization of biology with extra-
biological fields, such as cultural transmission, and enriches the debate among scientists
where extreme positions often pop out: does biology offer more rigorous methods to
replace the failing methods of the social sciences? Are we facing, instead, a case of
mutual inspiration? Or methodological integration? Which reciprocal prejudices are
well-grounded? And how can they be overcome for fruitful scientific collaborations?

Philosophy of biology also has a mandatory critical role towards biology. For example, it
can unveil the progressionist, anthropomorphic, and anthropocentric biases that affect
scientists as human beings who live immersed in a society and in a cultural
environment. Critical attention must be particularly high when scientific classifications
of humans (for example, through measures such as IQ or ethnicity) may lead to justify
and increase social discrimination, segregation or oppression.

Philosophy of biology may also develop ways of thinking up from biological research,
providing an inspiring and readable encompassing view of the living world that will
hardly be found in any standard, scientific publication. Furthermore, philosophy of
biology is called upon to work on the interface between science and society, contributing
to both the common misunderstandings and the best strategies for citizens to become
conscious and informed, as they are called to decide what kind of research and
intervention will be allowed or actively pursued by society.

It is hard for philosophy of biology to keep pace with the fast development of biological
knowledge. But the effort of following the moving frontier of knowledge allows
philosophy of biology to study the fall of influential ideas, such as the universal Tree of
Life, and the rise of new scientific practices, such as intensive computer modeling.
Philosophy also has the unsettling opportunity to constantly rethink its own approach,
avoiding drifting too far away from scientific practice so as to become detached. In this
dynamic, philosophy of biology is also well integrated with history of science, so that it is
often hard to distinguish between the two. An analysis of the relationship between
molecular biology and Mendelian genetics, for example, is intertwined with the
historical account of the birth and early development of molecular biology in the 1980s.
In turn, the philosophical framing of genetics and developmental biology as either
ontology-based disciplines or research styles transforms radically the way in which the
history of the two fields is told.

Philosophy of biology belongs to philosophy, therefore, no fixed procedure or protocol

constrains its research (what is philosophy?). Philosophy of biology consists in free and
critical — although rigorous and informed — thought on biological knowledge as the
latter develops through time. However, as a mature and recognized field with its own
interconnected practicing community, philosophy of biology seems to feature some
methodological principles:
 Philosophy of biology is supposed to be scientifically informed and up-to-date,
capturing how recent research modifies established knowledge and creates new
scientific practices. In turn, these novelties transform philosophy’s problems and
approaches, especially in the current explosive growth of biology.
  Philosophy and biology are not always clearly distinct. Scientific work can
routinely require, for example, conceptual or epistemological On the other hand,
philosophy can turn out to be effective in setting up scientific research projects.
However, philosophy can be characterized by its leaning towards
generalities about biology, namely general philosophical problems, general
characterizations of fields and approaches within biology, or conceptualizations
of biology as a whole or even of science as a whole.
 Philosophy of biology should try to be understandable and possibly useful to
biologists. Its tools — conceptual analysis, epistemology, traditions of thinking
and debates — should be put to use for improving scientific research.
 Biologists can do philosophy of biology. This happens, for example, when they
become interested in general features of biology and try to contribute with
principles derived from their work or when they think about the inferential
patterns employed by themselves and their colleagues. Also, scientists can do
philosophy and speak to philosophy when particular objects of philosophical
study, such as human morality, get naturalized (see below).
 Philosophy of biology cares for working across disciplinary contexts. For
example, it studies novel contacts between previously separated fields, develops
general views of the living world from some aspect of the life sciences, or reveals
complex connections between science and the socio-cultural context in which it is
carried out. It also takes advantage of its knowledge for monitoring and assisting
how science is publicly communicated and interrogated.
 Philosophy of biology is increasingly seen as one piece with history of biology,
since philosophical and historical theses are mutually necessary, and their results
reverberate reciprocally.
These six methodological principles are usually tacit, but sometimes they are made
explicit by philosophers of biology, who may also disagree on some of them. The
principles will be presented here by means of exemplar studies. Any set of examples is
anyhow partial and biased, since philosophy of biology is a huge field full of fascinating
topics, growing exponentially along with biology. For a more complete picture, the
interested reader will have to navigate the resources listed at the end of the article, such
as philosophy of biology journals or programs of conferences such as the biennial
meetings of the International Society for the History, Philosophy, and Social Studies of
Biology, the main reference society of the field. A number of textbooks in philosophy of
biology are available, often in the form of anthologies. A list of all these resources is
provided at the end for further reading.
Given the vastness of the philosophy of biology literature, this article can only indicate
some of the main topics and the richness of discussions. The examples in this article are
mainly focused on evolutionary biology. Evolutionists such as Ernst Mayr (1904-2005)
and Stephen Jay Gould (1942-2002), two of the most influential authors, are extensively
treated in this article though they are not universally representative. The predominance
of evolution can be justified not only by the author’s specialization, but also by the fact
that—as many philosophers of biology have critically stressed in recent times—
evolutionary theory has long been the main target of philosophy of biology. Only in the
last few decades has this situation changed radically (Müller-Wille 2007, Pradeu 2009).
Philosophy of biology is already tackling an enormous range of topics in the most
disparate fields, from biomedicine to community ecology, from neurobiology to
microbiology and microbial ecology, and from chemistry and biochemistry to
exobiology. To look for some specific area, the interested reader is, once again,
encouraged to venture into the journals and online resources.

2. General Issues in Philosophy of Biology

Philosophers of science (though not always under this fairly recent name) have reflected
for centuries on explanation, causation, correlation, chance, and many other general topics
concerning science or knowledge in general. Important philosophers contributed to
concepts like reduction vs. multiple realizability, and provided theories of explanation that
describe what a scientific explanation is. During the second half of the 20th century,
philosophy of science adopted a pluralistic strategy, considering the diversity of
scientific disciplines and methods and striving to understand their differences along
with their common aspects. With this pluralization, the complex task remained of
finding a satisfying description of science as an endeavor which—unitary or not—is
distinct from other forms of knowledge.
The study of living beings offers a universe of occasions for philosophy of science to
advance the reflection. For instance, explanations by natural selection and by drift (see
below, 2.a) can be seen as instances of causal explanations that nonetheless bring about
new reflections and conceptual puzzles on the classic issues of causation, randomness,
and ontology of processes. Homology explanations, another typical feature of biology,
explain the properties or the variability of a biological character by citing the ancestral
character or organ, and the causal factors that historically modify the descendants of
that ancestral organ. In trying to account for biological sciences, philosophy of biology
may take concepts from philosophy of science, such as causal explanation or reduction,
and find new putative cases of them in the life sciences or locate failures of reduction in
biology. Other times, philosophy of biology may need to tailor new concepts to
accommodate biology. In fact, some kinds of explanation seem peculiar to biology or to
historical sciences. While chasing the peculiarities of biology, philosophy of biology also
has some general research goals among its aims:
1. Often, philosophy of biology scours biology in search of new insights on general
philosophical problems about science, such as the “problem of induction,” or the
realism vs. instrumentalism dichotomy. Additionally, new general problems arise
from the particular forms that explanation, causation, or reduction take in
biology.
2. Sometimes philosophy of biology seeks general characterizations of particular
fields, practices, or ways of thinking within the life sciences Other times, the goal
seems to be a general picture of biology, especially by contrast to other sciences,
such as classical mechanics.
Sometimes philosophy of biology suggests general views of science, descriptions and
characterizations of science with all the complexity, differentiation, and plurality that it
exhibits in the contemporary world. In fact, the classical task of a demarcation of true
science from other forms of knowledge has lost importance under the effect of philosophy
of ‘special’ sciences like biology (Fodor 1974).
a. General Problems in Philosophy of Science, as Seen
in Biology
Natural selection is a major biological explanation for the features of organisms.
Inherited traits, originated by cumulative retention of random variation, are there
because of the positive contribution they have brought to their bearers in past
generations, in terms of survival and reproduction. Yet, the explanatory structure of
natural selection is very complex, and implies a reflection on concepts like causality and
randomness. In a classic book, Sober (1984) pointed out that only a few of the traits that
get selected are in fact explanatory, specifically those traits that are selected for. Other
traits are free riders that are somehow correlated with traits that are selected for and
thus are preserved in the population without actively contributing to fitness. Thus, there
is selection of such free rider traits that are not causally relevant to survival and
reproduction. Hearts are positively selected; heartthrob is also selected, but not selected
for; efficiency in pumping blood is selected for; the existence of heartthrob is thus
explained by natural selection, but heartthrob is not explanatory per se; it undergoes
selection of, not selection for. The idea of free riders on selection was already considered
by Darwin, but philosophers of biology spelled out its consequences for the explanatory
structure of natural selection. Causal relationships are the core of some theories of
explanation. Sober proposed rethinking the idea of causality in light of evolutionary
biology, and this is an example of how classic philosophical categories can be modified
in their application to biology: “We must show that by considering evolutionary theory,
old problems can be transformed and new problems brought into being. It remains to be
seen, I think, how radically the philosophy of science will be reinterpreted” (Sober 1984:
7; see Matthen and Ariew 2009, Ramsey 2013).
Randomness and chance are very important in biology. Natural selection is not random:
“it requires randomness as its ‘input’…but the ‘output’ of natural selection is decidedly
nonrandom, the differential survival and reproduction of the variants that are better
adapted” (Rosenberg and McShea 2008: 21). Philosophers worked, for example, on the
meaning of “random mutation,” a concept considered essential to Darwinism as opposed
to Lamarckism (Merlin 2010). Random does not necessarily mean lacking deterministic
causes. Rather, random mutation points to the fact that the usefulness of a trait in the
environment where it appears is not among the causes of its appearance. The source of
variation is thus more properly contingent with respect to fitness. An interesting line of
reasoning and evidence points out that “evolutionary divergence is sometimes due to
differences in the order of appearance of chance variations, and not to differences in the
direction of selection” (Beatty 2010: 39).
Genetic drift is the predictable change of the frequencies of traits that are not under
selection. The absence of selection makes the dynamic depend only on the reproduction
mechanism, and although the fate of individual traits is not predictable, the overall
landscape of frequencies is. Genetic drift has long been known in formal models, studied
in the field, and used for evolutionary reconstruction: it is not only necessary, but also
causally relevant to evolution. Yet, a lively philosophical debate exists on the ontology of
drift and selection (Millstein 2002; Walsh et al. 2002). A major disagreement concerns
the epistemic status of mathematical models: granted that drift is a necessary feature of
mathematical models, what can be legitimately inferred about the existence of a process
in the world to be called drift? A statistical interpretation sees both drift and selection as
mathematical features of aggregates of individuals (Walsh 2007, Matthen 2009, 2010).
Another point of view considers them as causal physical processes (Millstein 2006,
Millstein and Skipper 2009). The notion of evidence as a way of choosing between
alternative explanatory hypotheses, selection versus drift, for example, is another object
of philosophical study. Some philosophers think scientists are more qualified to evaluate
and weigh evidence (Hull 1969: 169). Others point out that it is up to philosophy to
probe the explanatory limits—current and constitutional—of biology (Rosenberg and
McShea 2008: 2). One fortunate approach to such a task is the technical account of
evidence based on Bayesianism (Sober 2008). Bayes’ theorem belongs to the
mathematics of probability theory. It is based on prior probability, the probability of a
particular statement before the observation, and posterior probability, the probability of
the statement in the light of the observation. Bayes’ theorem is used by some schools of
philosophers of biology in explicating various issues connected with evidence and
confirmation.
Homology explanations explain the properties or the variability of a biological character,
the form of a wing or the range of different wing shapes across different groups of
organisms, for example, by citing the ancestral character or organ, and the modification
factors that affect the descendants of that ancestral organ. Like many modes of
explanation in biology, homology explanations are historical (see also 3.a). Philosopher
Ereshefsky (2012) compares homology explanations with analogy explanations, which
instead explain a character by citing the contribution of that character to a function.
Ereshefsky points out that homology explanations are more detailed and offer a better
account of observed differences. Homology explanations can be also turned into strong
historical explanations, as opposed to weak ones that only cite the ancestral, initial
condition. This happens, for example, when detailed molecular studies of the
development of the target character enlighten precise events, such as gene duplications,
that must have been crucial along the historical path. The study of genetic and
developmental pathways also gives access to hierarchical disconnect which, for
Ereshefsky, relates homology explanations to classical topics of philosophy of
science: multiple realizability and reductionism. Hierarchical disconnect happens when “a
homologue at one level of biological organization is caused by non-homologous
developmental factors at lower levels of organization” (p. 385). Along the historical path,
for example, a morphological structure can remain stable while its underlying
developmental modules change (Griffiths and Brigandt 2007). For Ereshefsky, this is a
biological example of multiple realizability, that is the fact that one level of organization
cannot be reduced to a kind at a lower level. This in turn, for Ereshefsky, counters ideas
by Alex Rosenberg about reductionism in biology (Rosenberg 2006). For Rosenberg—
Ereshefsky says—the history of homologous characters should be reducible to the
history of their physical substrata, but Ereshefsky says hierarchical disconnect shows
decoupled histories and multiple realizability.
b. A General Picture of Biology
In characterizing ways of thinking and kinds of explanations and evidence, philosophy
of biology formulates generalizations about biology. These generalizations become
particularly encompassing when philosophy of biology tries to characterize biology
explicitly and comprehensively as distinct from other sciences. Biology is generally
considered a ‘special science’—a term inherited from logical positivist philosophy of
science that doesn’t preclude, in the long run, a reduction to physics (Fodor 1974,
Rosenberg and McShea 2008). Among the most noticed and studied features of biology
there is the apparent absence of scientific laws, whose blueprint, so to speak, are physical
laws (but see Waters 1998). Many have been the endeavors of describing biology as a
science without relying on laws, but also without relegating it as a mere collection and
description of singular events where exceptions are the law.
For Ernst Mayr (1982, 2004), biology is based on concepts or principles, which are more
flexible than laws: biology is a unique science by virtue of concepts that allow for
biological explanation, including inheritance, program, population, variation, emergence,
organism, individual, species, selection, fitness, and so on. Biology is also characterized
by population thinking, introduced by Darwin, which differentiates biology from
mechanics or chemistry whose thinking is, for Mayr, essentialist or typological (for more
on this see 4.a.i).
For paleontologist Niles Eldredge biology is based on patterns. Patterns are law-like
regularities, consisting of repeated schemes of events. This notion characterizes biology
as a historical science, while reducing the gap that, in other views, separates biology
from other natural sciences like physics. The pattern of inclusive hierarchical similarity
in the biological world was seen by Linnaeus and captured in his binomial
nomenclature. Darwin saw more patterns, for example in the geographical distribution
of species and varieties. He then discovered a subset of the grand complex of repeated
events, or regular processes, that give rise to biodiversity on Earth—the pattern of
evolution. Mendel caught some patterns as well, in his observations of inheritance.
Patterns have a double nature, ontological and epistemological: “Patterns in the natural
world are extremely important.… They pose both the questions and the answers that
scientists formulate as they seek to describe the world…. Science is a search for
resonance between mind and natural pattern as we try to answer these questions”
(Eldredge 1999: 4-5).
Other scholars think of biology as a science of mechanisms. A growing philosophical
movement called “the New Mechanism” (Machamer et al. 2000) use the concept to
revise classic ideas like causation, discovery, and explanation. In this view, biologists
aim to discover and represent mechanisms with their schemas, sketches, and theoretical
models. The characterization of natural selection as a mechanism, for instance, has been
proposed, but is yet to be resolved (Skipper and Millstein 2005).
Model-based accounts have acquired particular importance in philosophy of biology
(Schaffner 1993, 1998). The semantic view of scientific theories in the 1980s (see 2.c)
was a first ambitious endeavor to characterize biology as a model-based science. Downes
(1992) pointed out the limits of the semantic view in its original formulation, but
proposed that philosophers of biology keep some of its central claims, among which are
the centrality of various kinds of models in biology and their promise of accounting
for all scientific theorizing.
At a lower degree of generality, philosophy of biology offers a proliferation of ideas
about how schools of scientists, fields, or approaches perform fundamental activities of
science like explaining, describing, understanding, or predicting. Some included the
aforementioned and conceptually challenging explanations like natural selection, drift
and historical homology explanations. Others include the practices of ecological
modeling and model organisms, (6.b) particular inferential patterns such as
adaptationism (4.a.ii), and the difference between geneticists and developmental
biologists (7). These are all typicalities, or modest generalizations, of sub-parts of
biology. Population genetics is a thoroughly studied field from this point of view. By
studying population genetics, philosophers have raised wide-ranging topics of reflection,
such as the degree of idealization of models and experiments (Plutynski 2005, Morrison
2006). The concept of the possible has played a role in accounting for how biological
idealizations can be explanatory: if biological models cannot predict or demonstrate
necessity, they can at least restrict the field of what is possible, yielding so-called “how
possibly” answers or explanations. For Rosenberg and McShea (2008) it is a “usual
scientific” strategy that “scientists try to characterize a range of possible causes of
evolution, and then to determine which of these possibilities actually obtained. The
actual is first understood by first embedding it in the possible” (p. 13).
Even the tightest case studies usually produce generalizations to a certain degree. Grote
and O’Malley (2011), for example, claim that their historical reconstruction of research
on microbial rhodopsins

offers a novel perspective on the history of the molecular life sciences…, sheds light on
the dynamic connections between basic and applied science, and hypothesis-driven and
data-driven approaches [and] provides a rich example of how science works over longer
time periods, especially with regard to the transfer of materials, methods
and concepts between different research fields. (Grote and O’Malley 2011, p. 1082)
Sometimes generalizations come in negative form. Case studies are counted as evidence
that science may not have features that are commonly thought as necessary
requirements. Grote and O’Malley (cit.) propose their case against those philosophical
descriptions that interpret scientific advancement in terms of general, overarching, and
unifying theories. They observe that “productive interactions between different fields of
science occur not only through the adaptation of theories, concepts, or models, but very
much at the level of materials or experimental systems” (p. 1094, emphasis added).
c. A General Picture of Science
What is science in and of itself? How does it differ from other forms of knowledge?
These two sides of the coin were a major motivating question for philosophy of science
since its very beginning. Philosophers of biology sometimes take the methodological
lessons they learn from biology and tentatively postulate their broader applicability to
science (see Griesemer in section 7). However, a general view of science is often far from
the explicit goals of philosophy of biology. There are historical reasons for this. In a
paper on general philosophy of science, Psillos (2012) traces the problem back to
Aristotle, through landmark thinkers like Immanuel Kant and Pierre Duhem, to the
Modern era. The problem of demarcation was surely central in logical empiricism, but in
the 1970s the idea was spread that “individual sciences are not similar enough to be
lumped together under the mould of a grand unified scheme of how science works”
(Psillos, cit.: 100). After 1950s—and certainly in the early 1980s, when philosophy of
biology became an academic field—there was a consensus on the basic fact that science
employs a variety of methods. Indeed, biology, together with other sciences like the
human and the social sciences, had shaken the Modern idea of the scientific method.
Nevertheless, some important works have been moved by the need of finding new
descriptions of science that would fit biology better than received ones, in order to
legitimately include biology among the natural sciences. Political and intellectual
movements, such as Intelligent Design, contribute to urging philosophy of biology to
tackle the demarcation of science: their strategy includes casting doubts on the scientific
status of official biological sciences and presenting alternative, religiously inspired views
as scientific theories (Sober 2007, Boudry et al. 2010). The creationist movement is
present in the philosophers’ mental map worldwide since at least 1981, when
philosopher of biology Michael Ruse was a key witness for the plaintiff in the trial
McLean vs. Arkansas, a challenge to the state law permitting the teaching of creation
science in the Arkansas school system. The federal judge ruled that the state law was
unconstitutional. A significant part of the written opinion (see
http://www.talkorigins.org/faqs/mclean-v-arkansas.html) was devoted to philosophical
considerations on the epistemological characteristics of science (Claramonte Sanz 2011).
Intelligent Design is a new form of scientific creationism, characterized by the wedge
strategy, or public insistence on supposed flaws in the established biological sciences
combined with supposed scientific demonstrations of an Intelligent Designer of the
Universe. While there are different positions about whether philosophers and scientists
should directly appear in public debates with creationists, Intelligent Design
undoubtedly engages philosophers of biology in reflecting upon possible distinctions
within science (between hypotheses, theories, and models, for example), between
science and pseudo-science, and between science and religious beliefs. With their
competence, philosophers can help scientists in what sociologist Thomas F. Gieryn
termed “boundary work” (1999). The need of working on demarcation can also stimulate
critical revisions of the patterns of scientific explanation in biology. In a book on
evidence, Elliott Sober (2008) stated in a provocatively:
If the only thing that evolutionary biologists do is go around saying ‘that’s due to natural
selection’ when they examine the complex and useful traits that organisms have, they
are engaged in the same sterile game that creationists play when they declare ‘that’s due
to intelligent design’. Assumptions about natural selection of course can be invented
that allow the hypothesis of natural selection to fit what we observe. But that is not good
enough: the question is whether there is independent evidence for those auxiliary
propositions. (Sober 2008, p. 189)

The semantic view of scientific theories is an example of a general account of science that

was put to work in justifying evolutionary biology as a model-based science, in face of
the inveterate poor fit of the received view based on syntactic theories and universal
laws (Lloyd 1983, 1984, 1988). Under the semantic view, models constitute the core of
scientific work; theories are to be seen as combinations of families of models plus
hypotheses about the empirical scope of those models. This framework contrasts with
the received view which described theories as sets of sentences—including laws—about
the world. In the semantic view, laws are conceived in a new way: from universal
empirical claims about the world, they become specifications of models, embedded in
them. An important argument for philosophers of biology in the 1980s was that the
semantic view was not developed ad hoc for evolutionary theory: it had already been
successful in describing Newtonian mechanics, equilibrium thermodynamics, and
quantum mechanics. However, it was also said, these sciences fitted the received view as
well. The semantic view was very demanding in terms of formalization, so it worked only
for a very limited area of biology Not all scientific work consists in model building, so
the semantic view is hardly considered an exhaustive view of science, and things work so
differently for different kinds of models (for example, mathematical vs. non-
mathematical) that the existence of a unitary view was questioned (Downes 1992). But
new versions of the semantic view continue to be the topic of papers and debates in
philosophy of biology.
d. Generalization as a Possible Distinctive Feature of
Philosophy with Respect to Biology
As we have seen, philosophy of biology formulates various degrees of generalizations
about biology. These generalizations may concern biology as a whole or some subfield of
biology. Even in close-distance case studies, where generalization seems far from the
goals, it can be shown that generalizations are made (2.b). Philosophy is a generalizing
activity. This feature of philosophy may be a way to approach a tricky issue, namely the
distinction between biology and philosophy of biology.

Intuitively, philosophy is different from science. The methods of philosophy are based,

for example, on logical differences and implications (Hull 1969: 162) or on thought
experiments (Rosenberg and McShea 2008: 6), while the methods of science are based
on empirical phenomena. But this distinction is not as clear-cut as it might seem.
Scientists also use thought experiments. Einstein famously used them for developing
both special relativity and general relativity, and Darwin worked intensely with thought
experiments—to mention only two representative cases. Conceptual, foundational, and
linguistic analyses are an integral part of scientific research too. This means that
scientists autonomously do philosophy in their day-to-day work. Indeed, professional
biologists, as thinkers, could and should shift from their own main activity to more
philosophical ones whenever needed. The other way around, philosophers of biology
usually want to contribute to the advancement of science. In a sense, they want to be
part of science. Furthermore, the methods of philosophy are constantly evolving,
stimulated by advancements in biology, so that it is not rare now to find philosophical
studies that include mathematical analysis, computer simulations, or even empirical
research. The science-philosophy distinction is thus very blurred. Given this situation, is
there any possible demarcation between philosophy and biology?
David Hull (2002) characterized philosophy of biology as meta-science:

Knowing some science can be of great assistance to philosophers of science, but

philosophers of science as philosophers of science do not do science…. For example, the
equation F = ma is part of science, in particular physics. The claim that F = ma is a law of
nature is part of the domain of philosophy of science. To the extent that science and
philosophy can be distinguished in practice, scientists tell us what the world is like, and
philosophers of science tell us what science is like. (Hull 2002, p. 117)
One interpretation of philosophy of science as meta-science, which seems faithful to
Hull’s idea, is that philosophy of biology seeks generalizations about biology.

The idea of philosophy of biology as generalization about biology has descriptive

advantages, but it would be contested by some philosophers. In fact, no idea in the
literature about the specificity of philosophy remains uncontroversial. When it comes to
ambiguous science-philosophy cases, philosophers debate the right way of doing
philosophy. They are much clearer in saying what it means to be a scientist than they are
in saying what it means to be a philosopher.

In a view of philosophy as meta-science, the mark of philosophy is the leaning towards

generalizations about science. This is not a yes/no requirement; it comes in degrees.
Sometimes, for instance, works in biology make generalizations about biology (see Mayr
and Gould’s examples below). These works sometimes become philosophical while other
times they remain scientific because they generalize only to a degree which is functional
to the scientific activity. This continuity accounts for the demarcation uncertainties that
are going to persist in the field.
3. Philosophy Flanking Biology
 As is demonstrated in other parts of this article, philosophy can take a theme and
develop it with great autonomy from science or adopt a critical focus on science and on
the complex relationship between science and society. According to many authors,
however, there is also potential for philosophy to actively and directly contribute to the
advancement of life sciences. While it is commonly accepted that biologists can
undertake philosophy of biology, it is more controversial whether philosophers can do
biology in a proper sense. But biologists have generally been a receptive community. In
2002, David Hull had written:
Philosophers are attempting to join with biologists to improve our understanding of…
biological phenomena. As such, they run the risk of being considered by biologists to be
‘intruders’. In point of fact, biologists have been amazingly receptive to philosophers
who have turned their hand to philosophy of biology with a significant emphasis on
“biology.” (Hull 2002: 124)

When philosophers are close to scientific practice and current problems, they see how
they can help science in advancing towards its aims, either by criticizing existing ideas
and practices or by proposing revised ones. In Hull’s line of thought, philosophers are
encouraged to devote time and effort to work with biologists, or to formulate problems in
ways that are interesting and understandable to biologists. Philosophical treatments
that are too extensive, stereotypical, or posed in a way that is not relevant to biologists,
or those with an excess of philosophical formalization that prevents access to scientists,
should be avoided: “Formalization may be an excellent way of working out problems in
the philosophy of science. It is not a very good way of communicating the results” (Hull
1969: 178).
Philosophers try to help biologists to better frame their questions, for example by
“uncovering presuppositions and making them explicit” (Sober 1984), by analyzing
conceptual foundations (Pigliucci and Kaplan 2006), or by pursuing the detection,
analysis, and sometimes solution of theoretical and methodological problems. A great
deal of the philosophy of biology consists in working on scientific language to clarify the
meaning of concepts such as life, purpose, progress, complexity, genetic program,
adaptation, and so on (Rosenberg and McShea 2008: 4). Since these concepts frame the
scientific questions, philosophy of biology can “clarify, broaden or narrow the domain of
theories, uncovering ‘pseudo-questions’” (Rosenberg and McShea, cit.: 6). Critical
analysis and conceptual clarification are particularly valued tasks, and conceptual clarity
is seen as a necessary virtue of philosophy of biology. One mode of work consists in
looking, together with biologists, at concepts or mathematical models and their
interpretations. This line of work is exemplified hereafter (3.a, 3.b): two traditional
areas where philosophy has contributed by casting some light are the connections
among biological enterprises like taxonomy, classification, and systematics (3.a), and
the abstract descriptions of natural selection (3.b).
a. Clarifying Taxonomy, Classification, Systematics,
Phylogeny, Homology
In biology,  taxonomy consists in the recognition of natural groups, that is
the taxa  (singular: taxon). Classification deals with the categories and ranks to be
assigned to taxa (for example, species, genus, family). Systematics is, by definition, a
systematic study of the living world in search for order, or, in other words, the search for
the relationships among taxa. And phylogeny is the reconstruction of the temporal
scheme of common descent and relatedness among taxa. In fact, despite these gross
characterizations, the four activities are intertwined and depend on each other. Their
distinctions, definitions, and relationships are a traditional matter of reflection for
philosophy of biology (Wilkins and Ebach 2011).
In the 1960s, philosophers, perhaps thanks to the heritage of the Western philosophical
tradition, proved to be particularly ready and equipped for helping scientists understand
their own various ways of finding order in the living world. According to Hull (1969),
one of the important early contributions of philosophy of biology to logical clarity was
the taxon-category distinction, the distinction between “individuals, classes, and classes
of classes” (p. 171). Philosophers were able to contribute, for Hull, once they accepted to
arrange their formalisms to communicate with biologists. Hull himself (1976) long
argued for species having the ontological status of evolutionary individuals unlike taxa
in other categories. Hull relied on the Biological Species Concept (BSC) that defines
species as reproductive communities. In accordance with the BSC, whereas a taxon is
determined by a set of shared characteristics, the species-rank of the taxon—its
membership in the species category—is determined by actual evolutionary relationships,
in particular by interbreeding habits. With a similar line of reasoning, Hull defended the
idea of species as historical individuals, as opposed to classes or types. Species, perhaps,
are leaving the limelight of philosophical reflection as a consequence of the body of
discoveries about the fluidity of their boundaries, the heterogeneity of their
phenomenology, and the rarity of canonic biological species across the biological world.
But the debate on individuality was complex and lively, and is still partly open today
(Wilkins 2011).
In early years, philosophy of biology demonstrated its value also by analyzing the
entanglement among biological hypotheses—explicit and implicit—in different domains.
For example, philosophers refuted the idea of taxonomy as a theory-free activity, prior
to functional attributions and to evolutionary hypotheses. Character recognition does
bring into play theoretical considerations: the definition of “kidney,” for instance,
presupposes physiological knowledge of kidney function, and/or hypotheses on the
evolutionary derivation of kidneys. The acceptance of evolutionary theory in all fields of
biology, completed during the first half of the 20th century, triggered hot debates on its
relevance to taxonomy, systematics, and classification (Hull 1970): does taxonomy have
to reflect evolution? And in what sense? Could adaptation by natural selection be a
criterion for systematics, or is pure common descent the candidate? Philosophers took
part in trying to clarify these issues.
The network of assumptions and hypotheses of biology is an enduring object of study for
philosophy. There is a certain consensus on the fact that phylogeny should constrain
systematics; that is, systematics should reflect phylogeny as much as possible. Many
authors even equate the two tasks altogether: “Practitioners of systematics study the
historical pattern of evolution among groups of living things, i.e., phylogeny” (Haber
2008). Yet, taxonomy, classification, systematics, and phylogeny are unequally
performed by different professionals upon significantly uneven living and fossil taxa,
and there are many conflicting needs and goals. Species concepts in paleontology are by
force very different from those that fit neontology (Wilkins 2011), an issue that yields
differently organized classifications (Wilkins & Ebach 2013). Even among biologists who
study currently living organisms, the ideas on how to integrate taxonomy and
classification with systematics and phylogeny vary much across specialists at all degrees
of specialization—for example,entomologists, botanists, and mammalogists.
Furthermore, a classification of domestic animals or plants is likely to incorporate much
morphological, physiological, and ecological information beyond phylogenetic
relationships if it is to be of any practical use to the knowledge communities that are
involved in rearing. And when it comes to decide what information is relevant to identify
endangered species (or other units) that should be the object of conservation biology,
different ideas —and ethical stances are on the table (Casetta and Marques da Silva
2015).

Philosophy of biology can make or support concrete proposals about how to integrate
and refine the various ways of ordering the living world. Philosopher Marc Ereshefsky
(1997), for example, has been arguing for systems of nomenclature that are alternative
to the Linnean hierarchy (species-genera-Orders etc.). The reason for this position is
that all scientific changes that have been happening after Linnaeus’ century—Darwinism,
neo-Darwinism, cladistics and new methods in systematics—have turned the hierarchy
into an obstacle rather than an aid to taxonomical work.
A specific challenge for philosophy of biology isphylogenetic trees, which are ever
revisable hypotheses corroborated to varying degrees. Epistemological and
methodological discussions concern the ways of building, interpreting, testing, and
revising trees, as well as of relating them to other domains of knowledge such as
taxonomy or adaptation. “So what can biologists meaningfully say about phylogeny?”
philosopher Matt Haber asks (2008).
Broadly, two different issues have been at the center of recent systematics debates: given
epistemic limitations, whether any inference of phylogeny may justifiably be drawn; and
given an affirmative answer, what methods ought biologists use to justifiably infer
phylogenies, and what are the limits of these inferences? (Haber 2008, p. 231)
Competing and sometimes conflicting methods have been developed to make
phylogenetic inference more exact, manageable, or informative. In this domain,
methodological differences raised heated conflicts among scientists. The parsimony
principle was questioned in its legitimacy and importance. The principle states that the
most economic hypothesis has to be more true (Sober 1988). More generally, trees have
been examined for their theory-ladenness, that is, their sensitivity to background
theoretical assumptions. Other contentious matters were the acceptability of different
kinds of data ( genetic vs. morphological, for example), and the limits of the domain of
phylogenetic inference. Some workers maintain that methods in phylogenetics should
be able to detect homologies (see also 2.a) with certainty, discerning them from
analogies. Many others argue under different conceptions that homology detection
should rely on large amounts of data, mathematical models of evolution, and
probability. Supporters of cladistics reject probability and likelihood for being an
unstable ground on which to draw evolutionary trees. They put more confidence in the
cladist practictioner’s ability to recognize derivation between characters (Hull 1970,
Haber 2008). Meanwhile, the great majority of phylogenetic trees are built by relying on
huge sets of genetic sequences and a few morphological characters by means of more
and more cost-effective computer programs (this is an example of novel computer-based
scientific techniques posing new philosophical problems, see more below). “Total
evidence” methods (Sober 2008) address the challenge of building phylogenetic trees by
integrating all the available evidence—morphological, geological, ecological, and fossil.
The question of homology is a last example of philosophical enquiry into entangled
theoretical backgrounds and hypotheses: “when are two instances of a character to be
considered instances of the same character and in what sense?” (Hull 1969: 174).
Griffiths and Brigandt (2007) recognize different concepts of homology.
The taxic approach to homology—the best known in systematics and philosophy—uses
points of resemblances between organisms (shared character states) to diagnose their
evolutionary relationships. The transformational approach focuses on the different states
in which the same character can exist and be transformed by evolution. A third
approach to homology has emerged in conjunction with findings of evolutionary
developmental biology (EvoDevo): homology at the phenotypic level is potentially
decoupled from homology of developmental processes and homology at the level of the
genome (a phenomenon called hierarchical disconnect, see also 2.a). The level of depth
thus becomes a necessary specification for homology. Griffiths and Brigandt (cit.) see
the different concepts of homology as not only compatible but strongly complementary,
and trigger a series of reflections on their relationships.
b. Formulating Natural Selection
Charles Darwin (1859) conceived natural selection as the mechanism of change,
splitting, and divergence of lineages. Natural selection is thus the most essential notion
of evolutionary theory. Darwin’s formulation of natural selection, substantiated by many
empirical studies, was essentially verbal. After Darwin, definitions of natural selection
underwent diversification as well as progressive refinement. Mathematical models of
natural selection, created in the 20th century, made the process more precise, as
population genetics introduced technical terms such as fitness and selection
pressures (Haldane 1924), and established different formulas to quantify natural
selection and to measure its intensity and effects. But population genetics and
mathematics didn’t exhaust the task of formulating natural selection in a theoretical and
general way, and philosophers of biology eventually became very actively involved. A
classical abstraction of natural selection was elaborated by Richard Lewontin (1970). It
was based on variation, fitness, and heritability. Today philosophers acknowledge the
value of that account, but they also criticize it as, at once, too simple and too demanding
(Godfrey-Smith 2009), ascribing it to a category of recipe descriptions that list the
supposedly few and simple ingredients of natural selection. David Hull and Richard
Dawkins, for example, independently introduced the original distinction
between replicator and interactor, or vehicle. A replicator is anything that passes its
structure on largely intact, while an interactor is a cohesive unit whose action in the
environment makes a difference in the replication of the replicators it carries. Richard
Dawkins (1976), interpreting the work of William Hamilton and George C. Williams
(1966), famously took the basic idea of kin selection and developed it into the gene’s eye
view, a general view of evolution where selection in the long run is seen as operating
basically on genes and organisms are seen as their vehicles. Kin selection (Maynard-
Smith 1964) is based on shared inheritance among relatives. Social donor traits are
expected to spread in the population if they increase the fitness of the donor’s close
relatives, which are likely bearers of the same traits. Inclusive fitness (Hamilton 1963,
1964) is the fitness of a trait deriving from the bearer’s survival and reproduction plus
survival and reproduction of relatives (proportionally to the amount of genetic sharing).
Dawkins’s “selfish gene” metaphor, based on these mathematical findings, was
welcomed by many biologists as a clarifying device in their day-to-day work.  Its effects
in the public perception of science are a different story that will be addressed later in the
article. Some philosophers of biology got involved in the metaphor, either to develop it
(for example, Dennett 1995) or, more often, to criticize it (see Oyama 1998). Several
philosophers tried to find other ways of characterizing natural selection. We have
already mentioned Sober’s (1984) work on this problem (2.a). Sober (1983) also
described evolutionary theory as a theory of forces and population genetics as a theory
of “equilibrium models.” Abstract accounts of natural selection and its enabling
conditions flourished again with the turn of the 21st century. For Okasha (2006), a
replicators-interactors account is too demanding and narrow, imposing unnecessary
requirements for natural selection to occur. For Godfrey-Smith (2009), the concept of a
population is the crucial one: some features of a population render it “paradigmatically
Darwinian”, making natural selection happen. One motivating idea of such descriptions
is their potential use for evaluating the operation of natural selection among units at
different levels and in different domains (Rosenberg and McShea 2008). We will see
below the example of the domain of culture: what kind of selection, if any, is plausible in
the cultural domain?
Darwin thought natural selection happened chiefly among individuals: the individual
organism was the unit of selection. But late Darwin introduced “group selection” for
explaining some traits of humans and social insects (Darwin 1971). Groups were
foreshadowed as a larger unit of selection, and group selection seemed to be the
explanation for traits that jeopardize individual interest, such as cooperation and
abnegation. In the 1960s, evolutionary modeling showed that group selection required
repeated isolation, mixture, and re-isolation, namely conditions too narrow to be found
in nature with any significant frequency (Maynard-Smith 1964). Group selection was
disavowed; traits would not evolve simply because they are good for a group, they have
to be selectively advantageous in inter-individual competition from their inception. In
the 1970s, some philosophers of biology participated in a movement along with
evolutionary biologists and social scientists to try and develop group selection into a
scientifically respectable concept (Wilson 1975). In the meantime, kin selection had
emerged as a potential alternative explanation for group-beneficial, unselfish traits,
leading many scientists to conclude that group selection may be apparent and re-
described as kin selection if group members are relatives. Philosophers got directly
involved in the debate, sometimes working directly with biologists. In 1984, Elliott
Sober, citing Williams (1966) among others, talked about the “mirage” of group fitness,
seen as a mere statistical summation of individual fitnesses: “Selection works for the
good of the organism; a consequence may be that some groups are better than others.
However, it does not follow that selection works for the good of the group” (Sober 1984:
2). In many cases, the individual-based hypothesis is simpler than the group-level
characterization, so the principle of parsimony would recommend not adding explanatory
mechanisms. However, more in general, “Group, kin, and individual selection need to be
disentangled, their difference made clear” (Sober 1984: 4). In fact, conceptual
difficulties and fallacies in the units of selection framework were the attractor for
philosophers to this debate. Sober changed his mind about group selection through a
more careful conceptual analysis and by joining the work of biologist D.S. Wilson (Sober
and Wilson 1998). Most philosophers now think that unselfish traits may be explained
or made plausible by some combination of trait-level selection, organismal selection,
and group selection in a weak sense, together with a multiplication of hierarchies of
evolutionary entities and an “extended taxonomy of fitness” that contemplates co-opted
by-products and functional shifts (Pievani 2011).
Beyond gene, individual, and group selection, some authors have also attempted to
recognize higher levels, such as family selection, species selection, and clade selection,
although authoritative biologists such as Ernst Mayr contested this idea: “in no case are
these entities as such the object of selection. Selection in these cases always takes place
at the level of individuals” (Mayr 1997). Today, for many biologists, the question of what
unit is the “true” fundamental unit of selection has been satisfactorily settled—there are
several—but there are now new theoretical and empirical questions. Given that multiple
levels of vehicles exist, how does natural selection affect selection at lower or higher
levels, and how are higher-level vehicles created by lower-level selection (Keller 1999)?
The explanatory scope of multi-level selection, as philosophers have often emphasized,
is challenged by the major evolutionary transitions in the history of life (Maynard Smith
and Szathmáry 1995). Philosophers tend to describe a major transition in evolution as a
phase of emergence of a new level—with new units—of selection. The new level contrasts
or suppresses selection at the lower level, a process baptized “de-Darwinization” by
Godfrey-Smith (2009).
4. Who Can Do Philosophy of Biology?
As David Hull observed in 2002, philosophers “are attempting to join with biologists to
improve our understanding of…biological phenomena” (p. 124). As we have seen (2.d), a
demarcation between the two profiles—the philosopher and the biologist—is possible
but labile, and the distinction is perfectly compatible with any one scholar doing both,
even simultaneously (cf. Pradeu 2009). Consider now how Hull’s quote goes on: “But
sometimes the tables are turned. Biologists take up traditional philosophical topics and
attempt to treat them even if they are not professional philosophers” (Ibidem). Biologists
can turn to philosophy in two different ways: by reflecting philosophically on their own
work or by naturalizing philosophical problems. In the first case, biologists get
interested in issues of epistemology or methodology, and, from the ground of their work,
they extrapolate ways of thinking or modes of inference. Naturalization happens when
science becomes capable to say something significant and constraining about a
traditional topic of philosophy, exemplified here by the origin of morality.
a. Philosophical Biologists
It is not infrequent that biologists undertake philosophical reflections on their own
work. This is eased by the fact that some tasks of philosophy, such as conceptual
analysis or linguistic clarification, are integral parts of scientific work (see also 2.d).
Indeed, one might say that biologists, being experts about their own theories, are
sometimes more qualified than philosophers to reflect on their own work. But what
about the tendency to generality that characterizes philosophy of biology (section2)?
Well, the generalizing route, too, can be followed by working scientists. A biologist’s
philosophical effort may certainly vary in depth, richness, reach, and influence,
depending on many factors such as the range of his or her interests in terms of both
philosophical background and aims. With a good philosophical background, a biologist
can match more effectively with debates in philosophical areas. Their aims may go
beyond strict functionality for their own research and reach a genuine desire for
capturing, defending, or criticizing something deep about their own science. Two
examples—among many others—of very influential, philosophically-oriented biologists
are ornithologist Ernst Mayr and paleontologist Stephen Jay Gould.
i. Mayr and Population Thinking
Ernst Mayr (1904-2005) was one of the greatest evolutionary biologists of the20th
century, but he also increasingly worked in the history and philosophy of biology. He
was “a crucial link between professional philosophers of science and professional
biologists” (O’Malley 2010b: 530-1). Some of his areas of reflection were the distinction
between proximate and ultimate causes, the nature of the neo-Darwinian synthesis, and
the centrality of speciation and species defined by his Biological Species Concept (see
3.a). This article will focus on Mayr’s idea of population thinking as an example of how
“one scientist uses ‘scientific concepts’ to forge a conceptual tool with a wider range of
historical and philosophical applicability” (Chung 2003: 278).

The population notion was already central in Mayr’s work in 1942. His main concern
was the methodology of systematics. Mayr had promoted a new systematics, looking for
variation in large samples as opposed to few type specimens, and considering
geography, genetics, and other sources as opposed to morphology only. As Carl Chung
reconstructs, Mayr was first to formulate the distinction between typological and
population thinking in 1955. A few years later, Mayr (1959) started pushing population
thinking as the major innovation introduced by Charles Darwin and developed by
biology as a natural historical science, different and autonomous from other sciences.
According to population thinking, “no two individuals or biological events are exactly
the same and processes in biology can be understood only by a study of variation” (Mayr
1955, cit. in Chung 2003: 288). In opposition, Mayr described typological thinking as
having its deep roots in the Western cultural tradition, particularly in Platonic
philosophy: “Implicit in this concept is that variation as such is unimportant since it
represents only the ‘shadows’ of the eidos” (Mayr 1955: 485), the essences or ideas that
lie behind diversity. As Chung points out, the main battlefield for this opposition was the
concept of species. For Mayr, the incarnation of typological thinking in biology is
the morphological species concept, according to which individuals belong to the same
species by virtue of sharing their morphological characteristics. By
contrast, biological species concepts are population concepts. The latter take into
account variation, and in particular either reproductive barriers (two populations are
different species if they coexist with no fertile mating) and/or geographical variation
with reproductive flow between populations.
What is interesting here is the philosophical reach of Mayr’s ideas: he consciously
worked them up from his systematics field work, used them for a philosophical
interpretation of biology, and tied them to broader philosophical themes. In Chung’s
words, the enterprise was “an attempt to liberate certain key ideas of the ‘students of
diversity’ from their disciplinary constraints, and to render them more generally
applicable by repackaging them into a broader historical and philosophical distinction
that pertains to all of biology” (Chung 2003: 294-5). For sure, one goal was

to legitimize the natural historical sciences, including systematics, taxonomy, and

evolutionary biology, against the criticisms of ‘the new biology’—molecular,
reductionistic, and drawing explicit inspiration from the physical sciences…
philosophically he could argue for the ‘in principle’ need for an evolutionary (population
thinking) approach in order to offer adequate and complete explanations of biological
phenomena. (Chung: 295)
Population thinking, once devised, shaped Mayr’s own interpretations of the history of
biology, and provided him solutions to philosophical problems such as the method and
the autonomy of biology. It was generally taken up by philosophers and philosophically-
minded scientists such as Michael T. Ghiselin—notice the philosophical title of his 1997
book Metaphysics and the Origin of Species. There would be many other examples from
Mayr’s work in which he developed philosophical ideas out of science with precise
polemic aims and great influence on all scholars of biology. Statements like the
distinction between ultimate and proximate causes became common currency for
scientists. This does not mean that Mayr’s ideas weren’t criticized, as they increasingly
are (Ariew 2003, Laland et al. 2011, 2013).
ii. Gould and Adaptationism
Scientists may end up doing philosophy if they get interested in the inferential structure of
their own field. Inference, that is reasoning and its rules,is a classic topic in philosophy
of science. Induction, deduction, and inference to the best explanation (IBE) are basic,
well known types of inference, but, in scientific practice, they multiply and get combined
and put to work in different ways, producing interesting conceptual and philosophical
problems. Stephen Jay Gould (1942-2002) was a prolific writer (see more in section
5.d). Among his many favorite targets were a few inferential patterns employed by
evolutionary biologists. The adaptationist inference was definitely one of the main ones
since the famous Spandrels paper with Richard Lewontin (1979). Other biologists, like
G.C. Williams (1966), had advanced proposals for revising adaptationist inferences on
different grounds. Gould’s campaign can be seen as an expression of him being a
paleontologist with great interest in inferential patterns and with a view of evolution
directly inspired to Darwin’s works.
Adaptationism consists in explaining biological phenomena by claiming that they are
adaptations. In the Spandrels paper, Gould and Lewontin used the metaphor of the San
Marco cathedral in Venice to argue that even structures that exploit
fundamental functions can nonetheless result, originally, as structural byproducts of a
whole architecture. They wanted to criticize their colleagues’ habit to tolerate lazy tests
of adaptive hypotheses that “consisted in little more than a decent qualitative fit
between observed behaviour or form, and a set of posited adaptive pressures and
constraints” (Lewens 2008: 180). As Gould elaborated after the Spandrels paper,
between structures and functions there is no one-to-one strict correspondence, but
rather, redundancy. Functions are distributed over several parts of organisms, and
conversely any part we may call a trait or structure is involved in several mechanisms,
functions, and processes in the organism’s life. In the appreciation of trade-offs between
structural internal constraints and selected functions, Gould saw a revival of Charles
Darwin’s original attention to “contrivances” (1877). Adaptive explanations will rarely
suffice. In any case, they will need to be made testable and tested (Pievani and Serrelli
2011). Since 1982, together with Elisabeth Vrba, Gould proposed and promoted the
neologism “exaptation” to address what he saw as two evolutionary mechanisms,
distinct from adaptation, involving nonetheless natural selection and primary
functions: functional shift of a structure with previously different purposes, a process
already identified by Darwin; and functional cooptation of a trait whose origin is non-
adaptive— for example, a side effect due to a developmental constraint or a random
insertion.
Gould’s ideas about adaptationism and other aspects of biological inference triggered
cascades of philosophical reflections, for example on the multiplicity of meanings of
adaptation: by adaptation we may mean either something ensuring or increasing fitness
or something that seems designed for the performance in a particular environment, in a
range of environments, or for a particular function. We may intend something which is
being positively selected, or something whose existence is due to natural selection in the
past (Godfrey-Smith 2001, Lewens 2008). Biologists with philosophers, or
philosophically-minded biologists, reflected on adaptationism often reacting, in one
sense or the other, to Gould’s school of thought. Some scholars elaborated on the
dubious ontology and instrumental nature of adaptations, while many others
interpreted the challenge as the necessity of making adaptationist hypotheses testable
(Pievani and Serrelli, cit.). Recently, plant biologist Mark E. Olson (2012) acknowledged
the relevance of the “post-Spandrels consensus” on the importance of constraints in
evolution among biologists. But he also pointed out post-Spandrels proliferation of
contradictory “selection vs. constraints” and “externalist vs. internalist” explanations for
the same data. These contradictions were due, for Olson, to Gould’s vagueness in
defining “constraint,” as well as to the lack of experimental techniques for exploring the
accessibility of unobserved forms. But today’s embryological, manipulative, and
comparative empirical strategies allow for experimental exploration of morphologies
that are not observed in nature. By combining these techniques, biologists can turn
“internalism” and “externalism” from a-priori positions into case-by-case, testable
hypotheses. Also, selection and constraint are more properly seen as complementary
and not in mutual contrast as, for Olson, the Spandrels paper tended to suggest. For
Olson, thus, a developmental “renaissance of adaptationism” is under way.

As customary, philosophical issues raised by biologists have been taken up and

elaborated further by philosophers. Commenting on the philosophical literature on
adaptationism, Godfrey-Smith (2001) distinguished three different issues on which
adaptationist, anti-adaptationist and moderate positions can be taken up.
The empirical issue is whether or not natural selection is a powerful and ubiquitous force
in the natural world, with few constraints coming from biological variation, and with no
comparable, competing causal factor. The explanatory issue is whether the most
important questions in biology are about the fitting of organisms and environments,
given that natural selection is the only answer to such big problems (other processes and
explanations are good for less important questions). The methodological issue is whether
or not starting with adaptive hypotheses—and holding to them—is good scientific
practice. In 2009, a special issue of Biology & Philosophy was published as both a
celebration and a critical appraisal of the influence of the Spandrels paper. Therein,
Lewens (2008) elaborated on Godfrey-Smith’s taxonomy, recognizing seven types of
adaptationism, and arguing for the importance of asking a prerequisite question: “what
is a trait?”.
b. Philosophical Issues Naturalized
The relevance and implications of biology for humanity became a thought-provoking
and heartfelt issue as soon as intellectuals and laypeople reacted to Darwin’s works
(1859, 1871). More than a century later, E.O. Wilson (1975) proposed a “new synthesis”
as a project of explaining the most diverse human behavioral and psychological traits by
means of evolutionary hypotheses. Wilson’s proposal led to sociobiology and more
recently to evolutionary psychology (Barkow et al. 1992). Human behavior, mind,
morality, and systems of beliefs constitute the most interesting targets of possible, and
controversial, naturalization. Naturalization is what happens when matters that are
traditionally philosophical become empirically accessible by some scientific approach or
method. David Hull wrote provocatively that “Philosophy lost physics, then biology,
then psychology. Geometry, logic and mathematics became separate disciplines with no
necessary ties to philosophy,” therefore philosophers hold very tenaciously to
“epistemology, metaphysics, ethics and aesthetics” (Hull 2002: 124), because many of their
other objects have been taken by science. But Hull also optimistically observed that one
of the strengths of philosophy of biology “is that philosophers and biologists have
ignored this distinction, working with each other on both sides of the divide” (Ivi: 117).
In fact, naturalization is critically analyzed by philosophy, and there are many different
philosophical positions on naturalism. So, naturalization is a fruitful object of
study for philosophy of biology rather than a topic thief, and philosophy has a warranted
place that is not going to evaporate by naturalization.
Philosophers’ reactions to Wilson’s “new synthesis” were, for Hull, a virtuous example of
interactivity. After an initial wholesale opposition, many of them validated the challenge
of naturalizing humans. Our species has to be seen as a proper part of the biological
world, not as separated by any ontological divide. At the same time, philosophers
highlighted inferential errors in biological explanations of human behaviors,
epistemological limits in reconstructing the past, and ethical risks. They pursued
theoretical refinements of the project, by improving multi-level selection models, for
example. Many criticized the logical-deductive architecture of sociobiology
and evolutionary psychology, frequently built on ad hoc hypotheses and adaptationist
just-so stories. The debate is ongoing, and many arguments have been developed. For
example, if many human psychological mechanisms are evolutionary novelties due to
the interaction of ancestral genes and new environments, then many of these
mechanisms are not adaptations and adaptive thinking in evolutionary psychology will
fail to identify or explain them. More cautious, problematized, and integrated endeavors
of biological explanation applied to humans are emerging, also under the stimulation of
philosophy of biology (Sterelny 2003).
i. An Example: The Biology of Morality
It was for explaining human morality that Darwin hinted at ‘group selection’ (see 3.b),
and the endeavor wasn’t finished with him. Evolutionary ethics, the association of
morality with natural selection and evolution, provides good ground to Rosenberg and
McShea’s remark that biology “…is the only scientific discipline that anyone has ever
supposed might be able to answer the questions of moral and political philosophy”
(2008: 3). Fast growing fields like neurobiology or cognitive neurosciences seem to be
making biology more and more capable of addressing topics such as the origins of
morality. Philosopher Patricia Churchland (2011), for example, studied the scientific
literature and hypothesized a particular pattern of neural activation that would
constitute a “neurobiological platform” for morality. Churchland highlighted the role
played in this platform by molecules such as oxytocin, an ancient and simple peptide,
found in all vertebrates. In mammals, oxytocin “is at the hub of the intricate network of
mammalian adaptations for caring for others” (Churchland 2011: 14). In fact, morality
would share its neurobiological platform with other familiar phenomena of human life—
attachment and bonding. More generally, “the palette of neurochemicals affecting
neurons and muscles is substantially the same across vertebrates and invertebrates”
(Churchland 2011: 45). Among mammals, then, there is a wide “range of social
patterns…, but underlying them are probably different arrangements of receptors for
oxytocin and other hormones and neurochemicals” (p. 32). The striking thing is, for
Churchland, that modest modifications in existing neural structures can lead to new
outcomes. Morality and other phenomena would result from a not-so-exceptional
modification of a pre-existing platform involved in mammalian parental cares.
Churchland’s picture of evolution is again a familiar Gouldian one (4.a.ii):
Biological evolution does not achieve adaptations by designing a whole new mechanism
from scratch, but modifies what is already in place, little bit by little bit. Social emotions,
values, and behavior are not the result of a wholly new engineering plan, but rather an
adaptation of existing arrangements and mechanisms that are intimately linked with the
self-preserving circuitry for fighting, freezing, and flight, on the one hand, and for rest
and digest, on the other. (Churchland 2011: 46)

Does the evolutionary continuity from mammalian parental care to morality constrain
ethics and traditional philosophical theories of morality? Churchland’s view, while being
only an example, is interesting in its intermediate position between strict biological
determinism and cultural determinism. While biology can provide information and
explanation on the platform for morality, the complexity of cultures provides scaffolding
for moral development and definition so that moral decision remains a practical,
dialogic, and social problem. However, the more general question is whether and how
should moral philosophy—a large and highly technical field—take into any account what
the sciences are discovering. The answers to this question are expected to come from
philosophical studies of naturalization (Dupré 2001, De Caro and Macarthur 2004). In
any case, most philosophers of biology recognize a naturalistic fallacy in the idea that
knowing more about the natural world would suffice for making moral, political, and
social decisions.
ii. Philosophy Versus Naturalization?
To some philosophers, the naturalization of philosophical problems is rather
uncontroversial, to the point that “the difference between philosophy and theoretical
science is not a matter of kind but of degree,” and the domain of philosophy is partly
“the sum of all the questions to which science cannot (yet) answer” (Rosenberg and
McShea 2008: 5). For many others, defining philosophy as some kind of
underdeveloped science is an expression of scientism and a category mistake regarding
fields of knowledge. Many philosophers point out that the biological explanation of social
actions, behaviors, and culture, may imply a Darwinian dimension without boiling
down to it. Naturalism is related to, but different from, other very general issues,
like determinism or reductionism. Some philosophers draw a distinction between a strong
“scientistic” naturalism and a pluralistic, or “liberalized,” naturalism (De Caro and
Macarthur 2004). Scientistic naturalism considers philosophy as a branch of the natural
sciences. Liberalized naturalism includes different epistemic levels of analysis of human
nature—from natural sciences to humanities—that share the exclusion of non-natural
causes or principles.
Emphasizing the impact of biology on human capacities, social institutions, and ethical
values is also a way to justify philosophy of biology as useful or even indispensable to
philosophy and, more generally, to the humanities. Some presentations of philosophy of
biology tend to justify the field by its particularity of “concerning human affairs”
(Rosenberg and McShea 2008: 8) and its being ultimately oriented to them. Some
authors (Pradeu 2009) dislike this anthropocentric strategy in philosophy of biology
and think the field could be otherwise justified. Under these overarching debates,
human organisms and the human species are understandably a hotspot of problems for
philosophy of biology, and biologists and philosophers must confront the growing
biological knowledge of humans.

The neurobiology of morality (4.b.i) is not automatically a subtraction of morality as a

philosophical problem. Indeed, many reflections from a scientifically-informed
philosophy are of primary importance to maintain vigilance and scrutiny: what are the
aims and uses of these biological studies of humans? Could they be used, for example,
for a classification of people with consequences on the distribution of rights? Would this
be justified? Would the consequent choices and decisions be acceptable? How are
scientific results co-opted in clinical practice and health care? How are communication
issues with patients handled? How influential are social and cultural biases in the
construction of the object of research? Is this acceptable and justifiable? What ideas of
morality underlie the studies? Scientists working in these difficult grounds will
constantly exercise philosophical thought (see also 5.b). For them, the rich tradition of
moral philosophy might constitute a precious aid.
Meanwhile, philosophy of biology can inform philosophy about new and obsolete
approaches in the scientific explanation of morality. Evolutionary ethics is not
necessarily tied to adaptationism (4.a.ii) or genetic determinism. The evolution of
morality seems to resemble more bricolage than engineering. Against stereotypical and
simplified views of evolution, morality cannot be identified with a set of genes, even
though genes do not necessarily lose their importance, and the question about heritable
patterns remains crucial in order to define what morality is in evolution.
Philosophy of biology can fight easy deterministic conclusions or identifications between
naturalism and determinism while promoting useful definitions of the concepts
involved: what is to be intended for morality in different contexts, and why finding an
underlying arrangement of receptors is not going to replace the need for ethical
reasoning and moral philosophy. Science is a form of knowledge and therefore subject to
epistemology, conceptual analysis and change, and ethical reasoning. On the other
hand, biology constantly brings new fuel to philosophical inquiry, even, perhaps
especially, when philosophical issues get naturalized.

5. Philosophy Bringing the Life Sciences out of

Their Research Context
There are multiple senses in which philosophy of biology brings the life sciences out of
their research contexts. First, philosophy of biology can study and sometimes aid
interactions among the life sciences or between them and other sciences. Second,
sometimes philosophy is seen as capable of developing messages, ways of thinking, and
their consequences and implications to elaborate a “philosophy of nature” and an overall
vision of the living world (Godfrey-Smith 2009, p. 15). Third, philosophy can reflect on
the roles and meanings of science and on the interactions between science and society
with an approach different from that of the social sciences. In this way, it can assume
critical points of view towards biology and reflect on how scientific claims are, could be,
and should be received and elaborated by the public.
a. Philosophy of Biology at Intersections
What happens when different scientific fields or points of view come into intimate
contact? Philosophy of biology has always been happily and effectively involved in this
matter. A classic example (seen above, 3.a) is the analysis of coexistence, interaction,
and implicit reciprocal dependencies between “the morphological, the physiological, and
the genetical” viewpoints in taxonomy (Hull 1969: 176-7). Philosophers of biology often
notice attractions and tensions and call for integrations. They attend to emergent
relationships among life sciences, as in the topical cases of micro- and macroevolution
(Serrelli and Gontier 2015b) and evo-devo (see below), or between biology and other
sciences, as in cultural evolution studies. Classic debates in philosophy of science, for
example on reductionism, provide conceptual coordinates for thinking about these
connections.

A major development in biology began in the 1980s, when technical and theoretical
advancements enabled the molecular study of development, opening the possibility of
relating development to evolution in different ways (Gilbert et al. 1996, Minelli 2010,
Olson 2012). Evo-devo, evolutionary developmental biology, was born. The contact was
all the more significant because embryology, a discipline with an ancient tradition, had
long been seen as far removed from evolutionary biology. Many evo-devo protagonists
and observers framed evo-devo within the insufficiency of the mathematical study of the
intergenerational transmission of genes. This hegemonic field, population genetics,
considered development as a “black box” and assumed a linear relation between
genotype and phenotype (Laubichler 2010). In doing so, it would be blind to
fundamental kinds of evolutionary innovation, incapable of addressing
macroevolutionary change (Minelli 2010), and, more deeply, non-mechanistic, based on
“conceptual abstractions” (Laubichler, cit.). Yet, many scholars are pursuing
a pluralistic integration. They point out that
To deny the internal consistency and the explanatory power of [the research program
developed from population genetics] would be obviously foolish…. The objection is that
there can (and should) be more to evolutionary biology than a research program
restricted to the concepts and tools of population genetics (Minelli, cit.: 216).

Some scientists and philosophers focus on a broader polemic target: the Modern
Synthesis, that is the foundation of evolutionary biology as it is practiced today, which
happened between the 1910s and 1940s and was further canonized mainly by Ernst
Mayr in the subsequent decades. Critics analyze how the Modern Synthesis excluded
lines of research as non-legitimate or irrelevant to evolution, while successfully pulling
together Darwin’s natural selection, Mendel’s theory of inheritance, mathematical
models of population genetics, and the work of the most disparate fields in biology
(Gilbert et al. 1996, Serrelli 2015). Some philosophers and scientists are therefore
proposing the idea of an Extended Evolutionary Synthesis (Pigliucci and Müller 2010).
These movements are very interesting to philosophers of biology. They provide new
access to traditional philosophical issues about scientific change, including the unity and
disunity of science (Fodor 1974, Callebaut 2010). They also invoke a necessary
collaboration between history and philosophy of science, as shown by works that revised
the received views on the nature of the divorce of embryology and evolution in the 1930s
(Love 2003, Griesemer 2007 cf. section 7).
Philosophy of biology’s remarkable interest in cultural evolution studies is an example
involving relationships between biology and other fields. The idea of similarities across
biological and cultural evolution was already suggested by Darwin and his
contemporaries, and several approaches were formalized in the second half of the 20th
century (Cavalli Sforza and Feldman 1981, Boyd and Richerson 1985). Cultural change
and stability are understood in terms of variation, selection, and
inheritance/transmission of cultural traits, only requiring some correlation in cultural
transmission from cultural parents (models from whom a cultural trait is acquired) to
offspring (individuals acquiring the cultural trait). These evolution-inspired approaches
to culture allow for a variety of unique mechanisms for cultural transmission, and
incorporate processes like drift and multi-level selection. Capitalizing on such
approaches, some social scientists are proposing that methods, findings, and theories be
systematically exchanged between the biological and cultural sciences, particularly
between disciplines that lie at the same level on a micro-to-macro scale (Mesoudi et al.
2006). This trans-disciplinarity is sometimes presented as the beginning of a late and
needed evolutionary synthesis in the social sciences, similar to the Modern Synthesis in
biology, but some philosophers think this is an overstatement while others think the
claim is based on a naïve epistemological conception (Serrelli 2016a).
Similar proposals call philosophers of biology into question. The abstract formulations
and philosophical issues of natural selection, multi-level selection, and drift (3.b) can
help to evaluate the appropriateness of transfers of methods and theories (Godfrey-
Smith 2009). General topics like population thinking (4.a.i) and adaptationism (4.a.ii),
or reductionism will emerge in cultural evolution too. Progressionism is another very
basic issue here. The misleading idea of progress was in fact roughly applied in
anthropology by evolutionists in the past with potentially discriminatory effects and
reductionist justifications of essential diversities within the human species. Now,
philosophy can assist cultural anthropology in updating old stereotyped worries about
progressionism, overcoming a derived prejudice against naturalism as such. A coherent
and analytical criticism of any form of teleological and progressive evolutionism could
thus be a way to reconstruct the broken bridge between cultural anthropology and
evolutionary studies (Panebianco and Serrelli 2016a).  Cultural evolution also raises
deep epistemological problems, not only about definitions of terms like ‘culture’, but
also about the knowledge processes that are ongoing in this intensification of contact
between the biological and social sciences (Serrelli 2016b, Panebianco and Serrelli
2016b).
b. Biology’s Critical Friend
As we have seen, philosophy of biology is expected to help science and to work hard to
keep up with scientific research. But many authors point out that philosophy must not
forget its critical role towards science. One perspective on philosophy of science sees it as
an enterprise that aims “to understand the life of science, that is, to understand the
development of science as a Wittgensteinian family of ongoing human epistemic
practices” (Reydon 2005: 252, cf. Grene and Depew 2004). Science can be seen as a part
of culture and as a sub-system of society. This critical perspective on science, with its
complex relationship with society, is always available to philosophers. Authors like
Linda Van Speybroeck expressed the concern that philosophy of science might become
“…a servant of science, leaving science undisturbed and calling only for a ‘justification’
of the philosophical practice against scientific standards” (Van Speybroeck 2007: 54).
Stephen Jay Gould’s critique of progressionism in paleontology and paleoanthropology,
and his opposition to human racial classification and measurements of intelligence, are
examples of constructive criticisms of the scientific enterprise. Current knowledge
overwhelmingly shows that human evolution is a bushy tree of coexistent and
sometimes interacting hominin species (compare Ruse 2012). But cultural and
psychological biases have shaped science in some periods. Human evolution was
expected to be, and represented as, a ladder of progress, a sequence of progressively
more evolved hominid species, substituting one another, and approaching Homo sapiens,
the climax species, often represented as a European male (Eldredge and Tattersall
1982). Stephen Jay Gould, in books such as Wonderful Life (1989), with a peculiar
method that could be called an archaeology of scientific ideas (Pievani 2012a), coined
expressions like the “iconography of hope” and tied them to social history, to some deep
teleological preferences, and to our habit of using the present as a key for understanding
the past. For many years, Gould fought for human evolution to be separated from
human hopes, satisfying tales, and “great narratives.” He concentrated on the jargon,
distinguishing evolution and progress, trend and finality, and stressing the ambiguous
and appealing fashion for terms like “missing link.” Gould’s critical metaphors put
paleoanthropology in contact with the larger cultural context and deeper psychological
roots, helping to reinforce the theoretical normalization of human evolution into a
branching model of diversification of species, typical of the broader phylogenetic tree of
primates.
Gould’s way of reasoning was particularly apt in showing that science is a human
activity. Scientists work on the ground of cultural and social biases; they are not naïve
collectors of neutral facts. Although Gould was not a sociological relativist, he showed in
many cases the importance of history in shaping science, where ideas may also be
dismissed and then taken up again. This attitude is also found in Eldredge and Gould’s
(1972) famous “punctuated equilibria” paper, where the two paleontologists revealed the
deep theory-laden nature  of fossil interpretation. The ubiquitous pattern of
evolutionary stasis over geological time periods was neglected since Darwin (1859), who
considered fossils a constitutionally incomplete documentation. But the pattern of stasis
and punctuation was, as Eldredge and Gould pointed out, data, not lack of data.
“Phyletic gradualism” was a consolidated assumption that had been blinding even
paleontologists towards their own data, while perpetuating the subordinate position of
paleontologists with respect to theoretically important fields. In fact, Eldredge and
Gould formulated a theory that explained punctuations as speciation events, and stasis
by other processes. In doing so, they posed several problems, among which was the
legitimation of paleontology as a theoretically relevant field.
In The Mismeasure of Man (1981) Gould argued against the general scientific attitude of
attaching “universal essences” to human disparities by measuring what is not
measurable, like intelligence quotient, an artificial construct. He also exposed
unconscious manipulations of the anthropometric measurements and ranking of skulls
in Samuel G. Morton’s work (Crania Americana, 1839): Morton believed in “races” and in
their polygenic origin, and he believed human intelligence was a unitary and inheritable
object. Gould found his measurements as biased by his self-confirming preconceptions.
For Gould, any scientist is an unconscious victim of his or her preconceptions. Recent
studies of Morton’s material (Lewis et al., 2011) rehabilitated Morton’s original
measurements. The authors hypothesized that Gould’s severe analysis was biased by his
own aprioristic egalitarian and liberal cultural beliefs. “In a paradoxical way Steve had
proved his own point”, Ian Tattersall observed (2013). No scientist can be immune to
this kind influences, against which, for Gould, “vigilance and scrutiny” are the
necessary, although insufficient, palliations. Along these lines, philosophy of biology is
invited to take on its critical constructive role towards science.
c. Developing Messages from Biology
Many topics in philosophy of biology are evidently relevant in the relationship between
biology and the public. Deciding the meaning of concepts, like natural selection, fitness,
or function, requires an understanding of the theories and their domains (Rosenberg
and McShea 2008). Philosophy of biology clarifies, for example, that “‘natural selection’
is not an entirely apt name for the process [it identifies], as it misleadingly suggests the
notions of choice, desire, and belief built into the theological account of adaptations” (p.
17) and corrects popular descriptions of evolution that make it look tautological and
unscientific. The notoriously slow-changing world of science education and public
understanding is under pressure by the impressive rate of life sciences growth. In the
popularization of science, hominid evolution is still depicted in a linear sequence of
species from simple to complex, from inferior to superior, from archaic to modern. It
exhibits intuitive power, not to neglect the prospective endurance of other influential
pictures, like the tree of life, that are being questioned and made complex. And any time
a naturalistic study of humans is carried out, misunderstandings and dangerous “genes-
for-morality”-kind interpretations are just around the corner, both in the form of quick
reductionistic and deterministic claims and in the form of a-priori anti-
naturalistic positions. Even important scientific advances, like evo-devo and other fields
that are pushing for an Extended Evolutionary Synthesis, can be negligently interpreted
as breaking-offs, invalidating all the previous knowledge. Opponents of science can use
this to covertly reintroduce non-naturalist and non-scientific explanations. Instead,
philosophy of biology could help citizens become more scientific, and more able to
exploit the directional role they have towards science (cf. Haarsma et al. 2014-2015).
According to a principle called “scientific citizenship,” science should be properly
understood by everyone in society. A fundamental pillar is a shared awareness of “the
nature of science” which is the wealth of philosophical reflections about what is science,
what is biology, how is our scientific knowledge best acquired, and how much can we be
confident in it (Matthews 1994). What are the ways of thinking used in biology that can
help people to get a grasp of it? Are they similar to everyday ways of reasoning? What
are the possible conceptual traps and pitfalls? What kind of knowledge can we expect
from ecological simulations? What are model organisms, and why is it important to
establish and fund them? What kinds of predictions can really be made, for example, in
medicine or ecology? Understanding the probabilistic nature of predictions would
produce not only a more educated picture of nature, but also, for example, an
“awakening of public opinion to environmental problems, hydro-geological instability,
and maintenance of territory” (Pievani 2012b: 352).

Philosophy of biology’s endeavor can sometimes go far away from day-to-day biology,
and work out general pictures, messages, and worldviews. A classic and pervasive
example is “Universal Darwinism,” developed by Richard Dawkins, Daniel Dennett and
others (Dawkins 1983) along a reasoning line from kin selection theory to a “gene’s eye
view” to a “replicator view”. According to the “selfish gene” part of the argument
(Dawkins 1976), reliably replicating molecules (precursors of genes) would be at the
origin of life, and organisms including humans would be late-comer vehicles built up in
all their minute details by genes that survived the competition. The selfish gene has been
particularly appealing and controversial for its philosophical implications. We are
unaware machines for our genes, whose interests furthermore sometimes conflict with
(and win over) ours. The selfish gene view even came to be considered the official
version of Darwinism, being the one defended and advocated on many public stages
against non-scientific views. Universal Darwinism is a philosophical view, according to
which natural selection, intended as the selective retention and accumulation of blind
variations that prove to be stable and fit, is the fundamental mechanism in the Universe.
Many philosophers of biology fought these views and all their philosophical
implications, seeing them as a hardly justified reification of some methodologically
operationalized idealizations by population genetics (Oyama et al. 2001). Other lines of
attack were the incredible polisemy and theoretical complexity of “gene” and the
ongoing revision of their causal power on the organism (Griffths and Stotz 2013). Critics
were all the more motivated by their discontent with the picture of evolutionary biology
that was being conveyed to the public and to philosophy by the selfish gene view. Many
philosophers pointed out that natural selection is arbitrarily chosen as a process to be
universalized into a general philosophical view (Godfrey-Smith 2009), and some
outlined provocative alternative views like Universal Symbiogenesis (Gontier 2007).
Other philosophical views were elaborated from the importance of chance, randomness,
and, more comprehensively, contingency in evolution. Philosophers point out studies
demonstrating that chance variation can influence evolutionary outcomes without being
constrained or directed: “Evolutionary divergence is sometimes due to differences in the
order of appearance of chance variations, and not to differences in the direction of
selection” (Beatty 2010: 39). Since the 1980s, the neutral theory of genetic evolution
promoted by Motoo Kimura (Ohta & Kimura 1971), now developed in weak neutralism
in the scientific community (Hartl & Clark 2007), exposed the huge proportion of
neutral variation all around. Population genetics models show that important events like
speciation can well happen in conditions of fitness neutrality (Gavrilets 2004). Much
earlier, population geneticists demonstrated the importance of drift (see above, 2.a). But
is evolution random or contingent at any spatio-temporal scale? Are there law-like
tendencies in large-scale evolution? If so, do these concern adaptedness, complexity, or
other features? Since environments change over time, “what is adaptive” changes
constantly through evolutionary time, so there is no strict commitment in the Darwinian
theory to long-term adaptive progress or trends (Serrelli and Gontier 2015a).

Are trends towards greater complexity a better candidate? Some philosophers think so
(McShea and Brandon 2010). Others think that nothing in the current understanding of
evolution predicts a drive towards increased complexity. Another main disagreement
concerns how to define complexity— is it through the integrated organization of
interrelated parts in a whole or just through the number and diversity of parts? Some
philosophers see evolution as a texture of contingent histories, the most ordinary—but
most relevant to us—being the story of our species and their relatives. The human tree is
just like that of other mammals. This emerging view is very different from the one
inspired by Universal Darwinism:

Evolution is a process that abounds in redundancies and imperfections, and adaptation

could be a collateral effect rather than a direct optimisation. Biology is a field of
potentialities, and not determinations…. Complex organisms exist thanks to
imperfections, to multiplicity of use and redundancy. (Pievani 2012a: 142)

For authors like Stephen Jay Gould, the preeminence of ecological contingencies and
macro-evolutionary patterns, like mass-extinctions, in natural history seems to dismiss
any idea of progress in evolution:

We are the offspring of the material and contingent relationships between localised
populations and ever-changing environments. The massive contingency of human
evolution means that particular events, or apparently meaningless details, were able to
shape irreversibly the course of natural history. (Ivi: 139)

Some theorists identify the source of contingency in the complex interplay between
ecological systems and genealogical entities at multiple and very diverse scales
(Eldredge et al. 2016).
From the disruption of the idea of a great progressive tendency in evolution, in
particular human evolution, some philosophers develop general
implications. Evolutionary humility results from a naturalistic way of seeing Homo
sapiens as a part of a contingent process and not as its culmination. From another point
of view, the discovery of the determinant role of contingent ecological events like floods
and earthquakes gives a new vision of nature. Nature is neither a harmonic Eden nor a
wicked nemesis. “The expressions of violence and unpredictability of natural
phenomena that shock our societies so much today are the normal ecological niches
where we were born. We would not be here at this moment without them” (Pievani
2012b: 352).
6. Scientifically Up-to-Date Philosophy
In a famous paper titled “What the philosophy of biology is not,” David Hull wrote that
any work in philosophy of biology should not skip “all the intricacy of evolutionary
relationships, the difficulties with various mechanisms, the recalcitrant data, the wealth
of supporting evidence” (Hull 1969: 162). Along the same line, philosophy of biology
tends to be grounded on deep knowledge and understanding of current biology by
maintaining a non-episodic familiarity with many fields that are outside philosophical
specialization.

It is important to keep in mind that the life sciences and their objects change and grow. A
basic example is the very definition of “life” (bios). Answers, as well as philosophical
problems, for such a topic come, for example, from scientific research on the origin of
life (Penny 2005) or the search for extra-terrestrial life. The origin of life has been
considered as a backwards extension of the roots of the tree of common descent. It
regresses from some universal “ancestral organisms” (LUCA, Last Universal Common
Ancestor) to simpler, minimally living entities that might be referred to as “protoliving
systems,” and it further dissolves into non-living matter along several dimensions
(Malaterre 2010). In general, philosophers work together with biologists on interpreting
the history of life, for example, on trying to make sense of major evolutionary transitions
(Maynard Smith and Szathmáry 1995). The symbiogenesis of eukaryotes and the advent
of multicellularity are examples of major transitions. Those few moments in the history
of life may be seen as points of emergence of a new level of organization. Today the very
idea of a tree of life is being challenged by evidence of massive transfers and blurred
boundaries among its branches (O’Malley 2010a). The history of biology is a history of
changing views of life and of its history, and philosophy of biology participates into this
process.
In the 50 years of existence of philosophy of biology as an academic field, the expansion
of life sciences has been explosive. Huge global problems like climate change,
biodiversity loss, new forms of diseases, and needs for resource management in our
societies have contributed to that growth. Life sciences, including biomedical, ecological,
and microbiological fields, have faced challenges related to the Modern world, not only
by being called into question or by actively catching opportunities to develop big
research projects and programs but also by contributing to the very discovery and
perception of those challenges. In parallel, life sciences have seen incredible technical
advancements: cheaper and faster technologies for DNA sequencing and molecular
analysis; computational methods allowing analysis of billions nucleotides in search for
the most likely phylogeny and simulations of evolution of proteins or complex
phenotypes; huge shared information databases like the genome projects (such as
genomics, proteomics, metabolomics), particularly the Human Genome and the various
“–omics” projects monitoring whole complexes of functional molecules; finally, the
applications of these technical advancements in creating new kinds of organisms or in
disease detection and therapy. A parallel phenomenon of the decades leading into the
21st century has been the explosion and availability of scientific literature and access.
What are the consequences for philosophy of biology? The discipline is supposed to have
a role not only in understanding, describing, and communicating science but also in
aiding the development of scientific programs. Given the explosive historical dynamics
outlined above, almost every subfield of biology requires much of a philosopher to delve
into its peculiar concepts, methods, objects, and conceptual issues. Hence, several
presentations of philosophy of biology follow a field-by-field criterion, enumerating
items like “philosophy of ecology” or “philosophy of molecular biology” (Griffiths 2011).
Such multi-field presentations reflect the dynamic and lively development of biology.
Meanwhile, periodical shifts of focus and emergence of new fields and techniques in the
scientific literature, attract the curiosity and calling for the contribution of philosophers.
But the identities of supposed sub-fields like “philosophy of ecology” or “philosophy of
microbiology” are not crystallized, and rarely will a philosopher of biology self-limits to
one sub-field. Therefore, the field-by-field approach is not followed here.
This first section provides a few examples of how philosophers of biology can chase the
developments of some particular field of life sciences. In certain moments, this pursuit
can lead to extensions of philosophy of biology itself to embrace not only new
scientific knowledge, but also newborn ways of doing science. Furthermore, deep revisions
of philosophical approaches themselves may be necessary to address new aspects
of science, namely facets of scientific practice. The examples concern molecular studies
of gene exchange that started in microbial evolution, advances in ecological modeling,
and the construction and management of model organisms.
a. Questioning Influential Ideas
In recent years, several philosophers have become interested in the growing evidence for
a variety of gene exchange mechanisms widespread in fungi, plants, and animals, not
only in prokaryotes (unicellular organisms that have long been known to wildly
transform, conjugate and acquire DNA by transduction). Following biologists such as W.
Ford Doolittle, philosophers contrasted this evidence with the idea of a universal tree of
life as a tree of sexually reproducing, genetically isolated species that multiply by genetic
isolation. For O’Malley (2010b), the idea of a universal tree of life has come to be an
unacceptable reach given the abundance of reticulation and lateral gene transfer (LGT)
in all kinds of organisms, including animals. O’Malley proposed to give up
an animal-centric philosophy of evolution. Its key tenets are that the BSC (Biological
Species Concept) is ‘universal’ to species-forming organisms, that bifurcating lines of
descent are all that matter in ancestry reconstruction, and that the rest of life (non-
animal) is simpler, less diverse, and less ‘true’ evolutionarily. The consequences of [such
an] animal-centric philosophy of evolution are that it can include at best a severely
truncated history of evolutionary events. (p. 544)
The animal-based, constraining idea of a universal tree of life was consolidated around
the mid-20th century through an attempt of “excluding the messy”, such as prokaryotes
and bacteria. For O’Malley, such exclusion was mainly due to the influence of
ornithologist and philosopher Ernst Mayr.

As this example illustrates, philosophers, by following frontier developments of

particular fields, can sense the need for a revision of overarching theoretical choices and
frames of biology. At the same time, they can wriggle out of canonical problems and
concepts and expand their philosophical interests: “It is definitely the case – for
O’Malley – that philosophy of biology has undergone a rapid radiation of topics in the
last decade and this has meant going beyond Mayr’s focus” (544). The resilience of
issues like the species concept, molded on animal breeding, is pushed by new concepts
amenable to philosophical analysis. Further, the very task and range of philosophy of
biology happen to be questioned: “Mayr’s vision of philosophy of biology as the
clarification of evolutionary concepts has…been challenged” (O’Malley 2010b: 531).
b. Understanding New Scientific Practices
According to philosopher Thomas Pradeu (2009), ecology has made “a conspicuous and
very welcome entry” in philosophy of biology, and several philosophers advocate aspects
of ecology as targets of philosophical attention. Ecology has its
own epistemological issues, for example, the weakness of ecological laws, the debate
around the idea of balance of nature, the complex problem of the predictive value of
ecological models, and the involvement in environmental decision making (Cooper
2003, Mikkelson 2007, Plutynski 2008). Other problems concern the individuation of
ecological units, scale-dependence, and generalizability of ecological models. Some
philosophers got interested in new modeling methodologies of ecological inquiry, the
area of the following example.
Ecologist Steven Peck (2008) contributed to the debate from the standpoint of an
author of complex computer simulations. He pointed out the many differences between
simulations and “analytic models” which can be written as mathematical equations. In
simulations, many of the entities, “dispositions,” rules, and relationships, are not
captured by any equation, rather they are directly written in computer code:
“conditional if-then statements, looping structures, and calls to procedures” (Peck
2008: 390). Moreover, simulations are not at all complicated versions of analytic
models because, as Peck explains,

The complex computer representation is an ecological system. One that you have
complete control over, but which provides insights and allows complex behavior to
bubble up from lower level processes and allows one to capture the emergent behavior
often seen in ecological systems. (p. 387)
For Peck, models of this kind are provocative for philosophical issues, such as what are
models? What are their aims? How do they work? A classic and influential framework
by Richard Levins (1966) identifies three constraints among which a model has to trade-
off—generality, precision, and realism. But building a simulation, for Peck, does not
consist in adding more variables and parameters in order to capture more parts and
processes of some targeted biological system. It is a creative effort yielding something
autonomous with interesting but complex relationships with other models and with the
world. Some notions in the philosophy of modeling can make a step in the direction of
simulations, like the idea of “indirect representation” in which the model descriptions
themselves are examined as opposed to “direct representation” in which representation
is used to describe a real-world system (Godfrey-Smith 2006). But this distinction for
Peck is not enough to capture the fact that simulations become experimental systems in
themselves. Playing with the model means a “bracketing out of nature to explore the
model itself [yielding] important insights to the model – separate from what it is
supposed to represent” (Peck 2008: 395).

Complex computer simulations push philosophy of biology to reflect on new accounts of

model building and interpretation, and also to deal with new ways in which scientific
communities structure themselves.
c. Rethinking the Philosophical Approach from New
Ways of Doing Science
Chasing the state of the art of life sciences—even of one or few fields at once—is very
demanding. By doing it, however, philosophers of biology can continually fuel their
thought. We have seen, for example, that philosophers, relying on molecular discoveries
about gene exchange, can revise their agenda, downplaying traditionally-framed
problems (for example, what is a species?) and even questioning great background
pictures against which biology is thought (the tree of life, for example). By striving to
understand scientific methodologies, including completely new ones such as computer
simulations in ecology, philosophers are brought to probe their accounts of science to
work out new ones, and to get a better hold on them and their connections. Related to
all these efforts there is another tension in philosophy of science, which has been made
explicit in the last few years: the philosophical orientation toward scientific
practice (Boumans and Leonelli 2013). A recent trend, represented, for example, by the
Society for Philosophy of Science in Practice (SPSP), is pointing out the limits of an
exclusive use of conceptual analysis, proposing instead “a philosophy of scientific practice
based on an analytic framework that takes into consideration theory, practice and the
world simultaneously” (Boumans et al. 2011). Practice is defined as an ensemble of
“organized or regulated activities aimed at the achievement of certain goals” and is an
object of philosophical reflection.
Steven Peck’s analysis of ecological simulation, mentioned earlier, can also be seen as an
example of a call to scientific practice. For Peck, simulations are more of experimental
systems than representations. A simulation is useful because it helps “thinking more
deeply and creatively into the nature of the problem” (Peck 2008: 396), but it can be
appreciated only by considering the community in which the authors of the simulation are
present and maintain the simulation (otherwise, the simulation dies because it cannot
be understood or replicated by others). Crucial aspects are the authors’ willingness to
expose the multiple perspectives that are encoded in the simulation, to confront them
with other modelers and with “those who study the ecology of natural systems directly”
(p. 399), and the authors’ engagement in discussing, modifying, and exploring the
simulation further. In this new kind of engagement, Peck sees a “hermeneutic circle,” a
“back and forth conversation among modelers, their models, and those who study the
ecology of natural systems directly” characterized by each actor’s attempt to
“understand her own perspective in light of others’ perspectives.” The hermeneutic
circle is, for Peck, what “opens the door to deeper understanding of what the simulation
model is showing us about the world” (p. 399). Scientific community practices are
crucial for understanding what’s going on. Logical analysis, either of the model alone or
of its relationships with some natural ecological system, does not seem to bring
philosophy a long way.
Another example of a rising scientific practice is constituted by model organisms, a term
introduced in the late 1990s and becoming more and more used. Official lists of model
organisms include species such as the mouse, zebrafish, fruit fly, nematode worm, thale
cress. Mice and other animals are extremely important in biomedical research due to the
extrapolations to Homo sapiens that are considered possible with some conditions
(Piotrowska 2013). In philosophy of biology, there has been interest in understanding
what “model organisms” are and in demarcating them against the larger set of
experimental organisms. Ankeny and Leonelli (2011) define model organisms as:
Non-human species that are extensively studied in order to understand a range of
biological phenomena, with the hope that data and theories generated through use of
the model will be applicable to other organisms, particularly those that are in some way
more complex than the original model. (p. 313)

The two philosophers work out several concepts embedded in this definition in order to
capture the specificity of model organisms. For the issue at hand, the most important
aspect is the new kind of structured scientific communities that maintain a model
organism stable in space and time. Examples of model organisms are the fly Drosophila
melanogaster that has been studied since the dawn of genetics, knockout mice Mus
musculus, and the plant Arabidopsis thaliana. The research community of a model
organism performs intensive research with “a strong ethos of sharing resource
materials, techniques, and data” (p. 317). While initially the organism can be chosen for
experimental advantages (being easy to breed, for example), the cumulative
establishment of techniques, practices, and results—for example, through databases and
stock centers—leads to self-reinforcing standardization, comparability, and stability: “…
the more the model system is studied, and the greater the number of perspectives from
which it is understood, the more it becomes established as a model system” (Creager et
al. 2007: 6).
To summarize, a recent stream in philosophy of biology associates the need for first-
hand, recent, and deep scientific information with a need to consider newly emerged
scientific practices that often involve innovative scientific communities and that are
capable to generate new questions. It is no surprise that philosopher Leonelli, in a paper
on different modes of abstraction that are performed by different communities on the
model organism Arabidopsis thaliana, writes: “Focusing primarily on modelling practices,
rather than on models thus produced, might prove a useful way to gain insight on some
long-standing debates within the philosophy of scientific modelling and representation”
(Leonelli 2007: 510). By following biology, philosophy rethinks its own methods and
foundations.
7. History and Philosophy of Biology
Good philosophy of biology may be done non-historically, by working in a purely
conceptual way. On the other hand, philosophy is sometimes specifically interested in
grand historical processes. The inception of empirical and theoretical novelties and the
birth of whole new fields provide the opportunity to probe classic accounts of scientific
change such as Popper’s falsificationism (Popper 1935, 1963), Kuhn’s paradigm change
(Kuhn 1962), Lakatos’s methodology of scientific programmes (Lakatos 1970), or
Kitcher’s theoretical unification (Kitcher 1981). Philosophical categories such as
reductionism can be tested in their capability to account for the historical relationships
between biological fields. At the birth of molecular genetics, for example, a philosophical
question was whether the older Mendelian genetics was being reduced to it. As Griffiths
(2010) remarks, philosophers of biology achieved more adequate models of theory by
debating whether or not the molecular revolution in biology was a case of successful
scientific reduction.
At all scales, philosophy of biology has a constant need to refer to history, while some
authors complain that philosophy of biology pays too little attention “to the question
how and why things in the field have become the way they are today” (Reydon 2005:
149-150). The rediscovered interdependence between history and philosophy of biology
may be seen in light of the more general problem of the “history and philosophy of
science” (HPS) studies, which has been viewed in a more integrated way. The recursive
and expansive dynamics between history and philosophy of biology proves to be a
generator of dense and complex elaborations in all the involved fields.

James Griesemer (2007) constitutes an example of how philosophical views serve

historical analysis. The topic is the molecular study of development at the heart of evo-
devo (see also 5.a). Griesemer suggests a revision of conventional narratives that
describe evo-devo as a union between genetics and development, the study of which was
supposedly abandoned since the 1930s (see Gilbert et al. 1996). For Griesemer this
separation between fields is artificial: embryology and genetics have always been “like
the segments of a centipede: moving together with limited autonomy” (p. 376).
Griesemer’s long, reframing argument goes through the philosophical characterization
of scientists as process followers: “there is no doubt that scientists do follow processes,
that this is an important and central activity in their work, and that they achieve causal
understanding as a result of doing it” (p. 377). Research styles, are, for Griesemer,
“commitments to follow processes in a certain way.” Griesemer constructs the idea of
genetics and embryology as nothing but research styles that “package commitments to
follow processes according to particular sorts of marking interactions and tracking
conventions together with commitments to represent processes in particular ways” (p.
381). In this view, the separation between genes and development ceases to be
considered as an ontological divide: “It does not follow from the divergence of research
styles and representational practices in genetics and embryology that nature is divided
into separate processes of heredity and development” (p. 414). An important ingredient
of Griesemer’s view is the use of representations with their dual course. Before being
pressed into service as tools for communicating results and interpretations,
representations are “working objects, developed as bench or field tools for tracking
phenomena and following processes” (p. 388). Scientists, to be able to follow processes,
produce representations that, in turn, “provide reinforcing feedback that organizes
attention into foreground and background concerns” (p. 380). This dynamic becomes
important when representations survive the experimental work in which they are
produced and get used by other scientists. In scientific communities, process-marking
and process-following strongly reinforce each other in an attention-guiding feedback,
operated by representations. Griesemer analyzes Gregor Mendel’s experiments and
describes Mendel—universally considered as the founding father of genetics—as a
process follower and as a developmentalist. The different and successive notations
appearing in Mendel’s writings (for example, A + 2Aa + a; then A + A + a + a) are the
representations sequentially devised by Mendel in order to follow his enduring interest,
that is, the developmental process of hybrids. Therefore, in this account, Mendel was a
developmentalist who offered lasting representations that, in turn, helped some of his
followers to focus on patterns of intergenerational transmission, backgrounding
development. A particular power is given, in this account, to representations like
Mendel’s notation: “In Mendel’s demonstration of notational equivalence, the possibility
of reinterpreting his work on the development of hybrid characters in terms of a factor
theory of hereditary transmission was so strong that it took very careful analysis by
historians to show that Mendel probably did not hold the factor theory with which he is
credited” (Griesemer 2007: 404). The consolidation of research styles directly concealed
the awareness of their unity: “theories of heredity entail methodologies of development,
and conversely” (p. 414); “genetics entails an idealized and abstracted account of
development and development entails an idealized and abstract account of heredity” (p.
417); and “the gene theory not only had an embryological origin, it never really left
embryology at all” (p. 414). An implication for the present and future of evo-devo is that
putting heredity and development “back together again” can be thought of, in part, as a
problem of conceptual reorientation, change in theoretical perspective, and
rehabilitation of research styles that were out of the limelight rather than extinct due to
failure. The commonly suggested linear progress from embryology to genetics to evo-
devo is a historical artifact: there is no “historical progression of fields or lines of work
that take the scientific limelight in turn” (p. 417).
Whether the described implications are historical or philosophical is difficult to discern,
particularly in such cases as evo-devo, which are in current philosophical focus. It is
more productive to say that Griesemer’s analysis shows the interplay between
philosophy and history of biology. The view of scientists as “process-followers” can
indeed be seen as a methodological and a philosophical proposal.

Philosophically, the view of scientists as process-followers is pretty well elaborated and

detailed with the related notions of marking (mental and manipulative), foregrounding
and backgrounding, research styles, and so on. Griesemer explicitly confronts his
proposed view with more widespread approaches. For example:

The notion of following a process unifies [commonly separated] descriptions of science

as theoretical representation, as systematic observation, and as technological
intervention, [and] cuts across many analytical distinctions commonly used to describe
science (e.g., theory vs. observation, theory vs. experiment, hypothesis-testing vs.
measurement, active manipulation vs. passive observation, scientific methods vs.
scientific goals). (p. 375)

The marking activity, in particular, “is not easily assigned to either of the traditional
categories of passive observation and active experiment. It is nonmanipulative yet active
work” (p. 379), and since such work allows to identify causality, theory, and
methodology are intimately related. This local metatheorizing can be taken up by other
philosophers and elaborated further.

Historians, too, are invited to test the view in describing other cases in the history of
biology. To historians, however, there is also a reflexive message: just as scientists mark
and follow processes in the systems they study, so do historians with the social and
historical processes they follow. History and philosophy of science, in a word, would
consist in a “following of scientists while they follow nature” (p. 376). This means that
history and philosophy of science itself may be subject to research styles, and may need
theoretical reorientations to gain a new understanding. In general, for example, since
narratives tend to follow historical developments within scientific styles, history often
gives a false impression of scientific disciplines (embryology and genetics, for example)
that are separate because their theories describe different processes, like development
and heredity. So historians and philosophers are invited to abandon “limelight
narratives” that, while making “instructive drama,” confound our understanding of
social processes. In the specific case of study:
Embryology is not well tracked in narratives that foreground the success of genetics after
the split…. We must instead follow the ‘bushy’ divergences and reticulations of several
sciences as they spawn new lines of work if we are to understand, and follow, science as
a process. (p. 376, emphasis added)
8. Conclusion
We have seen that philosophy of biology can be actively involved in some scientific
activities. When this is not the case, philosophy of biology is largely an interpretive
description or re-description of instances of biology. Generalities about science in terms of
general descriptions of science, biology, or biology’s sub-fields, as well as in terms of
general philosophical problems about science, are usually the goals, sources, and
backgrounds of philosophy of biology. At the same time, tentative generalities about
science—such as the just seen view of “scientists as followers of processes”—can serve
historical reconstructions and translate into methodological proposals for historians,
too. However, the service is reciprocal, since present or past historical cases substantiate
—if they don’t suggest—new philosophical views of science, which is a specific tendency
of philosophy. The conclusion is that not only philosophy and science, but also history,
form an entangled, integrated whole in current research, thought, and practice.
9. References and Further Reading
a. Cited Examples
 Ankeny, Rachel, Sabina Leonelli. 2011. “What’s so special about model organisms?” Studies In
History and Philosophy of Science Part A 42(2): 313-323.
 Ariew, André. 2003. “Ernst Mayr’s ‘ultimate/proximate’ distinction reconsidered and
reconstructed.” Biology and Philosophy 18: 553–565.
 Eds. Barkow, Jerome H., Leda Cosmides , and John Tooby . 1992. The Adapted Mind: Evolutionary
Psychology and the Generation of Culture. New York, NY: Oxford University Press.
 Beatty, John., 2010. “Reconsidering the importance of chance variation.” Evolution – The Extended
Synthesis. Eds. Pigliucci Massimo and Gerd B. Müller. Cambridge-London: MIT Press, pp. 21-44.
 Boudry, Maarten, Stefaan Blancke, and Johan Braeckman. 2010. “How not to attack Intelligent
Design creationism: philosophical misconceptions about methodological naturalism.” Foundations
of Science 15(3): 227-244.
 Eds. Boumans, Marcel., Hasok Chang, and Rachel Ankeny. 2011. “Philosophy of Science in
Practice.” European Journal for Philosophy of Science 1(3).
 Boumans, Marcel, and Sabina Leonelli. 2013. “Introduction: on the Philosophy of Science in
Practice.” Journal for General Philosophy of Science 44(2): 259-261.
 Boyd, Robert, and Peter J. Richerson. 1985. Culture and the Evolutionary Process. Chicago:
University of Chicago Press.
 Callebaut, Werner. 2010. “The dialectics of dis/unity in the evolutionary synthesis and its
extensions.” Evolution – The Extended Synthesis. Eds. Pigliucci, Massimo, and Gerd B. Müller.
Cambridge-London: MIT Press, pp. 443-481.
 Casetta, Elena, and Jorge Marques da Silva. 2015. “Facing the Big Sixth: from prioritizing species to
conserving biodiversity.” Macroevolution – Explanation, Interpretation, Evidence. Eds. Serrelli,
Emanuelle, and Nathalie Gontier. Berlin: Springer, pp. 377-403.
 Cavalli-Sforza, Luigi L., and Marcus W. Feldman. 1981. Cultural Transmission and Evolution: A
Quantitative Approach. Princeton, NJ: Princeton University Press.
 Chung, Carl. 2003. “On the origin of the typological/population distinction in Ernst Mayr’s changing
views of species, 1942–1959.” Studies in History and Philosophy of Biological and Biomedical
Sciences 34(2): 277-296.
 Churchland, Patricia S. 2011. Braintrust. What Neuroscience Tells Us about Morality. Princeton, NJ:
Princeton University Press.
 Claramonte Sanz, Vincente. 2011. Diseño Inteligente: La Pseudociencia del Siglo XXI: Aspectos
Filosóficos, Sociológicos y Políticos de la Nueva Ideología Antievolucionista, Editorial Académica
Española.
 Cooper, Gregory. 2003. The Science of the Struggle for Existence: On the Foundations of Ecology.
Cambridge University Press.
 Eds. Creager, Angela N. H., Elizabeth Lunbeck, and M. Norton Wise. 2007. Science without Laws:
Model Systems, Cases, Exemplary Narratives. Durham and London: Duke University Press.
 Darwin, Charles R. 1859. On The Origin of Species. John Murray, London, sixth ed. quoted, 1872
edition.
 Darwin, Charles R. 1871. The Descent of Man, and Selection in Relation to Sex. John Murray,
London.
 Darwin, Charles R. 1877. The Various Contrivances by Which Orchids are Fertilized by Insects. John
Murray, London.
 Dawkins, Richard. 1976. The Selfish Gene. Oxford: Oxford University Press.
 Dawkins, Richard. 1983. Universal Darwinism. Evolution from Molecules to Man. Ed. Bendall D. S.
Cambridge: Cambridge University Press.
 Eds. De Caro, Maria, and David Macarthur. 2004. Naturalism in Question. Cambridge (MA): Harvard
University Press.
 Dennett, Daniel C. 1995. Darwin’s Dangerous Idea: Evolution and the Meanings of Life.  Simon &
Schuster, New York.
 Downes, Stephen M. 1992. “The importance of models in theorizing: a deflationary semantic
view.” Proc. of the Phil. of Science Ass. (PSA), volume 1. The University of Chicago Press, pp. 142-
153.
 Dupré, John. 2001. Human Nature and the Limits of Science. Oxford: Oxford University Press.
 Eldredge, Niles. 1999. The Pattern of Evolution. New York: Freeman and Co.
 Eldredge, Niles, and Stephen J. Gould. 1972. “Punctuated equilibria: an alternative to phyletic
gradual- ism.”  Models in palaeobiology. Ed. Shopf, Thomas J. M. San Francisco CA: Freeman.
 Eldredge, Niles., Telmo Pievani, Emanuele Serrelli, and, Ilya Temkin. 2016. Evolutionary Theory: A
Hierarchical Perspective. University of Chicago Press.
 Eldredge, Niles, and Ian Tattersall. 1982. The Myths of Human Evolution. New York: Columbia
University Press.
 Ereshefsky, Marc. 1997. “The evolution of the linnaean hierarchy.” Biology and Philosophy 12(4):
493–519.
 Ereshefsky, Marc. 2012. “Homology thinking.” Biology and Philosophy 27(3): 381-400.
 Fodor, Jerry. 1974. “Special sciences and the disunity of science as a working
hypothesis.” Synthese 28: 97-115.
 Gavrilets, Sergey. 2004. Fitness Landscapes and the Origin of Species. Princeton: Princeton
University Press.
 Ghiselin, Michael T. 1997. Metaphysics and the Origin of Species. Albany, NY: State University
Press of New York.
 Gilbert Scott F., John M. Opitz, and Rudolf A Raff. 1996. “Resynthesizing evolutionary and
developmental biology.” Developmental Biology 173:357-372.
 Godfrey-Smith, Peter. 2001. “Three kinds of adaptationism.” Adaptationism and Optimality. Eds.
Orzack, Steve H., and Elliott Sober. Cambridge University Press, pp. 335-357.
 Godfrey-Smith, Peter. 2006. “The strategy of model-based science.” Biology and Philosophy 21(5):
725-740.
 Godfrey-Smith, Peter. 2009. Darwinian Populations and Natural Selection.  Oxford: Oxford
University Press.
 Gontier, Nathalie. 2007. “Universal symbiogenesis: an alternative to universal selectionist accounts of
evolution.” Symbiosis 44: 167-181.
 Gould, Stephen. J. 1981. The Mismeasure of Man. New York: W.W. Norton.
 Gould, Stephen J. 1989. Wonderful Life. The Burgess Shale and the Nature of History. New York: W.
W. Norton.
 Gould, Stephen J., and Richard C. Lewontin. 1979. “The spandrels of San Marco and the Panglossian
paradigm: a critique of the adaptationist programme.” Proceedings of the Royal Society of London
Part B: Biological Sciences 205: 581-59.
 Gould, Stephen J., and Elisabeth S. Vrba. 1982. “Exaptation – a missing term in the science of
form.” Paleobiology 8: 4-15.
 Griesemer, James R. 2007. “Tracking organic processes: representations and research styles in
classical embryology and genetics.” From Embryology to Evo-Devo: A History of Developmental
Evolution. Eds. Laubichler, Manfred D., and Jane Mannenschein. Cambridge, MA: MIT Press, pp.
375-433.
 Eds. Griffiths, Paul E., and Ingo Brigandt. 2007. “The Importance of Homology for Biology and
Philosophy.” Biology and Philosophy 22(5).
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 Oyama, Susan, Paul E. Griffiths, and Russell D. Gray eds. 2001. Cycles of Contingency. Cambridge
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 Peck, Steven L. 2008. “The hermeneutics of ecological simulation.” Biology and Philosophy 23(3):
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 Penny, David. 2005. “An interpretive review of the origin of life research.” Biology and
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 Pievani, Telmo. 2011. “Born to cooperate? Altruism as exaptation, and the evolution of human
sociality.” Origins of Cooperation and Altruism. Eds. Sussman, Robert W. and C. Robert Cloninger.
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 Pievani, Telmo. 2012. “Many ways of being human, the Stephen J. Gould’s legacy to Palaeo-
Anthropology (2002-2012).” Journal of Anthropological Sciences 90: 33-49.
 Pievani, Telmo. 2012. “Geoethics and philosophy of earth sciences: the role of geophysical factors in
human evolution.” Annals of Geophysics 55(3): 349-353.
 Pievani, Telmo, and Emanuele Serrelli. 2011. “Exaptation in human evolution: how to test adaptive vs
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 Piotrowska, Monika. 2013. “From humanized mice to human disease: Guiding extrapolation from
model to target.” Biology and Philosophy 28(3): 439-455.
 Plutynski, Anya. “Explanatory unification and the early synthesis.” The British Journal for the
Philosophy of Science 56(3): 595-609.
 Plutynski, Anya. 2008. “Ecology and the environment.”  The Oxford Handbook of Philosophy of
Biology. Ed. Ruse, Michael. New York: Oxford University Press, pp. 504-524.
 Popper, Karl R. 1935. Logik der Forschung, Vienna: Julius Springer Verlag. Eng. Tr. The Logic of
Scientific Discovery. London: Hutchinson, 1959.
 Popper, Karl R. 1963. Conjectures and Refutations: The Growth of Scientific Knowledge. London:
Routledge.
 Psillos, Stathis. 2012. “What is General Philosophy of Science?” Journal for General Philosophy of
Science 43(1): 93-103.
 Ramsey, Grant. 2013. “Can fitness differences be a cause of evolution?” Philosophy and Theory in
Biology 5: e401.
 Reydon, Thomas A. C. E. 2005. “Bridging the gap between history and philosophy of
biology.” Metascience 14(2): 249-253.
 Rosenberg, Alexander. 2006. Darwinian Reductionism. Or, How to Stop Worrying and Love
Molecular Biology. Chicago: The University of Chicago Press.
 Ruse, Michael. 2012. The Philosophy of Human Evolution. Cambridge & New York: Cambridge
University Press.
 Schaffner, Kenneth F. 1993. Discovery and Explanation in Biology and Medicine. Chicago:
University of Chicago Press.
 Schaffner, Kenneth F. 1998. “Model organisms and behavioral genetics: a rejoinder.” Philosophy of
Science 65:276-288.
 Serrelli, Emanuele. 2015. “Visualizing macroevolution: from adaptive landscapes to compositions of
multiple spaces.” Macroevolution: explanation, interpretation and evidence. Eds. Serrelli,
Emanuele, and Niles Gontier. Interdisciplinary Evolution Research series, Springer, pp. 113-162.
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 Serrelli, Emanuele. 2016. “Evolutionary genetics and cultural traits in a ‘body of theory’
perspective.” Understanding Cultural Traits. A Multidisciplinary Perspective on Cultural
Diversity.  Eds. Panebianco, Fabrizio, and Emanuele Serrelli. Springer, Switzerland.
 Serrelli, Emanuele. 2016. “Removing barriers in scientific research: concepts, synthesis and
catalysis.” Understanding Cultural Traits. A Multidisciplinary Perspective on Cultural
Diversity.  Eds. Panebianco, Fabrizio, and Emanuele Serrelli. Springer, Switzerland.
 Serrelli, Emanuele, and Niles Gontier, eds. 2015. Macroevolution: Explanation, Interpretation and
Evidence. Springer. [DOI 10.1007/978-3-319-15045-1]
 Serrelli Emanuele, and Niles Gontier. 2015. “Macroevolutionary issues and approaches in
evolutionary biology.” Macroevolution: explanation, interpretation and evidence. Eds. Serrelli,
Emanuele, and Niles Gontier. Interdisciplinary Evolution Research series, Springer, pp. 1-25. [DOI
10.1007/978-3-319-15045-1_1]
 Skipper, Robert A., and Roberta L. Millstein. 2005. “Thinking about evolutionary mechanisms:
natural selection.” Studies in the History and Philosophy of Biological and Biomedical Sciences 36:
327-347.
 Sober, Elliott. 2007. “What is wrong with Intelligent Design?” The Quarterly Review of
Biology 82(1): 3-8.
 Sober, Elliott, and David S. Wilson. 1998. Unto Others: The Evolution and Psychology of Unselfish
Behavior. Cambridge, MA: Harvard University Press.
 Sterelny, Kim. 2003. Thought in a Hostile World: The Evolution of Human Cognition. Oxford:
Blackwell.
 Tattersall, Ian. 2013. “Stephen J. Gould’s intellectual legacy to anthropology.” Stephen J. Gould’s
Legacy. Nature, History, Society. Eds. Daniele, Antonio G., Allesandro Minelli, and Telmo Pievani.
New York: Springer-Verlag.
 Van Speybroeck, Linda. 2007. “Philosophy of biology: about the fossilization of disciplines and other
embryonic thought.” Acta Biotheoretica 55(1): 47-71.
 Walsh, Denis M. 2007. “The pomp of superfluous causes: the interpretation of evolutionary
theory.” Philosophy of Science 74(3): 281-303.
 Walsh, Denis M., Tim Lewens, and André Ariew. 2002. “The trials of life: natural selection and
random drift.” Philosophy of Science 69(3): 452-473.
  Waters, C. Kenneth. 1998. “Causal regularities in the biological world of contingent
distributions.” Biology and Philosophy 13(1): 5-36.
 Wilkins, John S. 2011. Species: A History of the Idea. Berkeley, CA: University of California Press.
 Wilkins, John S., and Malte C. Ebach. 2013. The Nature of Classification: Relationships and Kinds in
the Natural Sciences. Palgrave Macmillan.
 Williams, George C. 1966. Adaptation and Natural Selection: A Critique of Some Current
Evolutionary Thought. Princeton University Press.
 Wilson, David S. 1975. “A theory of group selection.” Proceedings of the National Academy of
Sciences USA 72(1): 143-146.
 Wilson, Edward O. 1975. Sociobiology: The New Synthesis. Cambridge (MA): Harvard University
Press.
b. Classics
i. First Generation
 Beckner, Morton. 1959. The Biological Way of Thought. New York: Columbia University Press.
 Hull, David L. 1964. “The metaphysics of evolution.” British Journal for the History of Science 3:
309-337.
 Hull, David L. 1969. “What philosophy of biology is not.” Synthese 20: 157-84.
 Hull, David L. 1970. “Contemporary systematic philosophies.” Annual Review of Ecology and
Systematics 1(1): 19-54.
 Hull, David L. 1974. Philosophy of Biological Science. Englewood Cliffs, NJ: Prentice-Hall.
 Hull, David L. 1976. “Are species really individuals?” Systematic Biology 25(2): 174-191.
 Grene, Marjorie G. 1959. “Two evolutionary theories, I–II.” British Journal for the Philosophy of
Science 9: 110–27; 185–93.
 Grene, Marjorie G., and Everett Mendelsohn, eds. 1976. Topics in the Philosophy of Biology.
Dordrecht: D. Reidel.
 Schaffner, Kenneth F. 1967. “Approaches to reduction.” Philosophy of Science 34: 137-147.
 Schaffner, Kenneth F. 1967. “Antireductionism and molecular biology.” Science 157: 644-647.
 Ruse, Michael. 1973. The Philosophy of Biology. London:Hutchinson.
 Wimsatt, William C. 1972. “Teleology and the logical structure of function statements.” Studies in
History and Philosophy of Science 3: 1-80.
ii. Second Generation
 Amundson, Ron. 1988. “Logical adaptationism.” Behavioral and Brain Sciences 11: 505-506.
 Amundson, Ron. 1989. “The trials and tribulations of selectionist explanations.” Issues in
Evolutionary  Epistemology. Eds. Hahlweg, Kai, and Clifford A. Hooker. State University of New
York Press.
 Beatty, John. 1980. “Optimal-design models and the strategy of model building in evolutionary
biology.” Philosophy of Science 47: 532-561.
 Beatty, John. 1982. “Classes and cladists.” Systematic Zoology 31: 25-34.
 Brandon, Robert. 1990. Adaptation and Environment. Cambridge: MIT Press.
 Burian, Richard M. 1988. “Challenges to the evolutionary synthesis.” Evolutionary Biology 23: 247-
259.
 Darden, Lindley. 1977. “William Bateson and the promise of Mendelism.” Journal of the History of
Biology 10: 87-106.
 Darden, Lindley. 1976. “Reasoning in scientific change: Charles Darwin, Hugo de Vries, and the
discovery of segregation.” Studies in History and Philosophy of Science 7: 127-169.
 Depew, David, and Bruce H. Weber. 1985. Evolution at a Crossroads: The New Biology and the New
Philosophy of Science. Cambridge MA: Bradford Books/MIT Press.
 Depew, David, and Bruce H. Weber. 1995. Darwinism Evolving: Systems Dynamics and the
Genealogy of Natural Selection. Cambridge, MA: Bradford Books/MIT Press.
  Dupré, John A. 1987. “Human kinds.”. The Latest on the Best. Ed. Dupré, John A. Bradford
Books/MIT Press, pp. 327-348.
 Gieryn, Thomas F. 1999. Cultural Boundaries of Science: Credibility on the Line. Chicago: University
of Chicago Press.
 Griesemer, James R. 1988. “Genes, memes and demes.” Biology and Philosophy 3:179-184.
 Griesemer, James R., and Michael J. Wade. 1988. “Laboratory models, causal explanation and group
selection.” Biology and Philosophy 3: 67-96.
 Kitcher, Philip. 1981. “Explanatory unification.” Philosophy of Science 48: 507-531.
 Kitcher, Philip. 1982. “Genes.” British Journal for the Philosophy of Science 33: 337-359.
 Kitcher, Philip. 1984. “Species.” Philosophy of Science 51: 308-333.
 Lloyd, Elisabeth A. 1983. “The nature of Darwin’s support for the theory of natural
selection.” Philosophy of Science 50: 112-129.
 Lloyd, Elisabeth A. 1984. “A Semantic approach to the structure of population genetics.” Philosophy
of Science 51: 242-264.
 Lloyd, Elisabeth A. 1988. The Structure and Confirmation of Evolutionary Theory. Greenwood Press.
 Mills, Susan K., and John Beatty. 1979. “The propensity interpretation of fitness.” Philosophy of
Science 46: 263-286.
 Rosenberg, Alexander. 1978. “The supervenience of biological concepts.” Philosophy of Science 45:
368-386.
 Rosenberg, Alexander. 1982. “On the propensity definition of fitness.” Philosophy of Science 49: 268-
273.
 Sober, Elliott. 1983. “Parsimony in systematics: philosophical issues.” Annual Review of Ecology and
Systematics 14(1): 335-357.
 Sober, Elliott. 1983. “Equilibrium explanation.” Philosophical Studies 43: 201- 210.
 Sober, Elliott. 1984. The Nature of Selection. Evolutionary Theory in Philosophical Focus. Chicago:
University of Chicago Press.
 Sober, Elliott. 1988. Reconstructing the Past: Parsimony, Evolution, and Inference. Cambridge:
Cambridge University Press.
 Weber, Bruce H., James Smith, and David Depew. 1988. Entropy, Information, and Evolution: New
Perspectives on Physical and Biological Evolution. Cambridge, MA: Bradford Books/MIT Press.
c. Contemporary
i. Reviews
 Byron, Jason M. 2007. “Whence philosophy of biology?” The British Journal for the Philosophy of
Science 58(3): 409-422.
 Callebaut, Werner. 2005. “Again, what the philosophy of biology is not.” Acta Biotheoretica 23: 93-
122.
 Griffiths, Paul E. 2011. “Philosophy of biology.” The Stanford Encyclopedia of Philosophy (Summer
2011 Edition). Ed. Zalta, Edward N.. URL =
<http://plato.stanford.edu/archives/sum2011/entries/biology-philosophy/>.
 Hull, David L. 2002. “Recent philosophy of biology: a review.” Acta Biotheoretica 50: 117-128.
 Gers, Matt. 2009. “The long reach of philosophy of biology.” Biology and Philosophy 26(3): 439-447.
 Müller-Wille, Staffan. 2007. “Philosophy of biology beyond evolution.” Biological Theory 2(1): 111-
112.
 O’Malley, Maureen A., and John A. Dupré. 2007. “Size doesn’t matter: towards a more inclusive
philosophy of biology.” Biology and Philosophy 22(2): 155-191.
 Pradeu, Thomas. 2009. “What philosophy of biology should be.” Biology and Philosophy 26(1): 119-
127.
ii. Some Monographs
 Pigliucci, Massimo, and Jonathan Kaplan. 2006. Making Sense of Evolution. Chicago: University of
Chicago Press.
  Sober, Elliott. 2008. Evidence and Evolution: The Logic Behind the Science. Cambridge University
Press.
 Sober, Elliott, and David S. Wilson. 1998. Unto Others: the Evolution and Psychology of Unselfish
Behavior. Cambridge (MA): Harvard University Press.
d. Anthologies and Textbooks
 Ayala, Francisco, and Robert Arp, eds. 2009. Contemporary Debates in Philosophy of Biology. Wiley-
Blackwell.
 Callebaut, Werner, ed. 1993. Taking the Naturalistic Turn: Or, How Real Philosophy of Science is
Done. Chicago:University of Chicago Press.
 Grene, Marjorie G., and David Depew. 2004. The Philosophy of Biology: An Episodic History.
Cambridge: Cambridge University Press.
 Hull, David L., and Michael Ruse, eds. 1998. The Philosophy of Biology. Oxford: Oxford University
Press.
 Hull, David L., and Michael Ruse, eds. 2007. The Cambridge Companion to the Philosophy of
Biology. Cambridge: Cambridge University Press.
 Kampourakis, Kostas, ed. 2013. The Philosophy of Biology: A Companion for Educators. Springer
Science & Business.
 Linquist, Stefan, ed. 2010. Philosophy of Evolutionary Biology. Vol. 1. Berlington, VT: Ashgate.
 Rosenberg, Alexander, and Robert Arp, eds. 2009. Philosophy of Biology: An Anthology. Wiley-
Blackwell.
 Rosenberg, Alexander, and Daniel W. McShea. 2008. Philosophy of Biology: A Contemporary
Introduction. Routledge.
 Ruse, Michael, ed. 1989. What the Philosophy of Biology is.  Dordrecht, The Netherlands: Kluwer.
 Ruse, Michael, ed. 2008. The Oxford Handbook of Philosophy of Biology. New York: Oxford
University Press.
 Sarkar, Sahotra, and Anya Plutynski. 2008. A Companion to the Philosophy of Biology. Blackwell
Publishing.
 Sober, Elliott. 1993. Philosophy of Biology. Westview Press/Oxford University Press.
 Sober, Elliott. 2006. Conceptual Issues in Evolutionary Biology. Cambridge, MA: MIT Press.
 Sterelny, Kim, and Paul E. Griffiths. 1999. Sex and Death: An Introduction to the Philosophy of
Biology. Chicago: The University of Chicago Press.
e. Journals
i. Dedicated
 Acta Biotheoretica (since 1935)
 History and Philosophy of the Life Sciences (since 1979)
 Biology and Philosophy
 Journal of the History of Biology
 Studies in History and Philosophy of Biological and Biomedical Sciences Journal of the History of
Biology
 Biological Theory
 Studies In History and Philosophy of Science Part A
 Philosophical Transactions of the Royal Society B
 Philosophy and Theory in Biology (since 2009)
ii. Generalist
 Philosophy of Science
 The British Journal for the Philosophy of Science
 Foundations of Science
 Metascience
 Studies in History and Philosophy of Science
f. Organizations
 International Society for the History, Philosophy, and Social Studies of Biology
(http://www.ishpssb.org/)
 Philosophy of Science Association (PSA) (http://www.philsci.org/)
 Society for Philosophy of Science in Practice (SPSP) (http://www.philosophy-science-practice.org/)
 Inter-Divisional Teaching Commission (IDTC) of the International Union for the History and
Philosophy of Science (IUHPS): http://www.idtc-iuhps.com/
g. Online Resources
 T. Lewens, “The philosophy of biology: a selection of readings and resources” page at the University
of Cambridge: http://www.hps.cam.ac.uk/research/philofbio.html

Author Information
Emanuele Serrelli
Email: emanuele.serrelli@epistemologia.eu
University of Milano-Bicocca
Italy