A MODERN SYNTHESIS OF PHILOSOPHY AND BIOLOGY

Marion Godman

Forthcoming in Kelly Becker & Ian Thomson (eds.), History of Philosophy, 1945 to
2015. Cambridge: Cambridge University Press.

Biology was not of much concern to the logical empiricists, who settled for physics as
their model of science and scientific explanation. In the course of the period of this
volume, the situation has changed drastically. Philosophy of biology is now a large
and respected academic specialisation in its own right.

There is much to be said for the birth of philosophy of biology coming out of the
Modern (Evolutionary) Synthesis (MS) during the 1930s, 40s the 1950s. While
biologists, such as Theodosius Dobzhansky, Julian Huxley, George Simpson, and
Ernst Mayr launched the MS, philosophers as interpreters and critics played a
significant part in determining its faith, as we shall see. Moreover, as noted by
Marjorie Grene and David Depew, “The work of defending, expanding, challenging,
and, perhaps, replacing the Modern Synthesis has tended to bring out the philosopher
in many evolutionary biologists” (2004, 248).

So wherein lied the intellectual potency of the MS? One of the central points of the
MS was to unify disciplines such as Mendelian genetics, palaeontology, systematics,
biometrics and ecology.1 In particular, the MS aimed to improve on the theoretical
framework of Darwin’s evolution by fusing natural selection with Mendelian
population genetics.

Population genetics allows for studies of evolutionary change and changes in fitness,
by studying changes in gene frequencies within populations. This change is thought to
occur according to the degree of genetic variation within a population coupled with
the degree of heritability of different genes (where heritability is a measure of
resemblance between parents and offspring). The idea is then that natural selection is
demonstrated by the relative speed and accuracy by which genes replicate. Success in
reproduction is supposed to indicate the adaptive superiority of organisms with those
genes leaving more offspring than others (see e.g. Brandon 1978).

I will focus on three themes in the philosophy of biology where from the MS and
onward there have been particularly high levels of cross-fertilization amongst
philosophers and biologists. It is however worth mentioning some of what this focus
omits. The domains of ecology and embryology, although envisioned as part of the
MS by at least Huxley and Dobzhansky, have always had a bit of a strained
relationship with evolutionary theorising, and the philosophy of ecology has also been
rather autonomous in philosophy of biology (Levins 1968). Nowadays topics in
immunology and synthetic biology, which would have been seen as peripheral earlier,
are also receiving a great deal of philosophical scrutiny. Moreover, many

1
The MS preceded molecular genetics and the discovery of the molecular mechanism of inheritance
empirically that was borne out by the uncovering the double-stranded DNA molecule using x-ray
crystallography. Most now think molecular genetics enhances rather than replaces an evolutionary
perspective.
philosophers of biology, rather than turning to the theoretical framework of biology,
are more concerned with the (distinctive) scientific practice and methods of biology.

This is how I plan to proceed. First, I’ll look at two debates at the core of the MS,
about the nature of species and the dominance of genetic selection, where
philosophical engagement has been particularly noticeable and relevant to the
development of the MS. Finally I will highlight a particular synthesis of philosophy
and biology in the study of culture that also casts a new light on the traditional
questions of the MS.

1. Sorting out species

The quest for the origin and justification for classification of organisms into species
lied at the heart of MS just as it did for Darwin. What explains the diversity of
different life forms that is clustered in different kinds of species? Or one can turn the
question around as Daniel Dennett (1995) does and ask why there is not continuous
variation, but “gaps in the design space”? One of Dobzhansky’s central themes was to
connect the origin of species with the question of the origin of genetic variation. He
showed how species are formed as a result of both geographical and genetic
mechanisms for isolating populations. He used population genetics to model the
variations and changes in chromosomes he had noted in the microscope when
studying different species and sub-species of fruit flies (Drosphila) (Grene & Depew,
253. f.). Thus, although there are many salient characters, which are similar amongst
members of a species, if the right mechanisms for isolation are in place, natural
selection will exploit the phenotypic consequences of genetic diversity within a
natural population.

Debates soon ensued about what this meant for the nature of species. Ernst Mayr
building on the work of Dobzhansky claimed that the upshot of the MS is that species
are constituted by their members’ capacity to generate gene flow. What has become
known as Mayr’s Biological Species Concept (BSC) thus states that species are
interbreeding natural groups which are reproductively isolated from other such groups
(1969). Gaylord Simpson on the other hand considered selection and maintenance of
adaptations within phyletic lineages to be the central moral of the MS (1961). It is
unclear whether he explicitly disagreed with Mayr, but he at least seemed to have
found Mayr’s “non-dimensional” view of species insufficient. For him individuals
with long stretches of historical time between them and which may lack the
propensity to interbreed with one another should still be able to qualify as members of
the same species if they belong to the same lineage. Simpson’s Evolutionary lineage
concept hence explicitly proposed that species are historical and not just populations
at a particular time-slice.2 Simpson’s view arguably also gained in currency against
Mayr when it was translated into the notable research programmes of phylogenetics
and cladistics where species were classified in terms of their historical branching
relationships (Mischler & Brandon 1987).3

2
The other main concern raised against the BSC is how it deals with organisms that reproduce
asexually.
3
At the same time it is unclear if this view is compatible with punctuated equilibriums – Gould and
Eldrige’s view that the gaps in the fossil record reflect the fact that long states of equilibrium for
species are punctuated by period of rapid change.
Several other debates have arisen in response to these disagreements – or, perhaps,
different points of emphasis. A central one issue for philosophers has been to
determine how much of a departure from traditional typological or natural kind
thinking this post-Darwinian view of species really is. One way to read the success of
the population-thinking in MS is to take it as a vindication of Darwin’s “natural
system of classification” (1859, p. 485). In particular, it can be seen as a vindication
of species as classes. If we are concerned with the direction of gene flow and of
natural selection in natural populations, it won’t do to study random assortments of
individuals. Indeed all the to-do about species in the MS suggests there is something
special about species as a natural populations.

But does this suffice to render species natural kinds? Of course everyone agrees that
for species to evolve they cannot be eternal, unchangeable, entities. Indeed,
evolutionary biology has helped demonstrate that this is a rather unattractive view of
natural kinds in science in general! However, further push back has been offered
against the claim that species are types or natural kinds. Since variation between
individuals is the central condition for natural selection, the MS seemed committed to
thinking that any average within a species would be a mere statistical abstraction –
not anything real belonging to the population or kind. Thus, it seems an Aristotelean
natural state model must fall by the wayside in modern biology (Sober 1980).

As Phillip Honenberger (2014) has highlighted, the real starting point for this
discussion was Marjorie Grene’s “Two evolutionary theories” (1958) and David
Hull’s “The Effect of Essentialism on Taxonomy--Two Thousand Years of Stasis”
(1965) – two of the first philosophy of biology articles published in the British
Journal of the Philosophy of Science.4 In her article, Grene focuses on a possible
inconsistency she detects amongst the authors of the MS (focusing on Simpson):
insofar as the legitimacy of type or kind concepts is denied, one cannot make
reference to notions like “adaptive types” in the accounts of species. Grene elaborates:

When evolutionists talk about slight variations, we may ask, variations of

what? Of bristle number, or length of limb, or skin pattern or pigmentation, or
what you will, but of some trait or other. Variants must differ from their
neighbors, however minutely, in some character or characters. But characters
are not and cannot be particulars. They are sortals, predicates that sort out
different kinds. (1990, 239)

While Grene therefore thinks kind and sortal concepts are both justified and
indispensable to biology, Hull is of the opposite view. He believes that any non-
instrumental endorsement of natural kinds is doomed to fail. The problem as he sees it
is the connection between kinds, classes and essentialism, and affirming natural kinds
would force us to postulate some necessary intrinsic properties which all and only all
members of a kind share in common. In his view, the MS just shows that there are no
such properties (1965). His negative thesis also develops into a positive thesis in the
case of species. He suggests species are particular individuals rather than classes
supporting lawlike generalisations (1976). Species on this view do not have parts or

4
Honeberger also nominates Hull and Grene as “two of the most influential philosophers of biology”
and notes that two out of the three biennial prizes offered by one of the most important organizations in
the field, the International Society for History, Philosophy, and Social Studies of Biology (ISHPSSB)
are named after Grene and Hull respectively (2014, 13).
members; but are spatial-temporal slices of the genealogical nexus. The view that
species are individuals also seems to fit well with Simpson’s view that species are
lineages enabled by Mayr’s interbreeding and reproductive isolation.

In the 1980s and onward, we see philosophers of biology give roughly three different
responses to this debate about whether species are kinds: The first follows Sober and
Hull in their rejection of species as natural kinds, but also embraces the diversity of
views on species as indicative of an ontological pluralism where different epistemic
perspectives are allowed to carve up species and the tree of life differently dependent
on their interests (e.g. Kitcher 1984; Dupré 1993). The second and third strategy
follows Grene in maintaining the usefulness of natural kind thinking for species and
biological kinds in general. Despite variation on the genetic and phenotypic levels,
these authors suggest the cluster of traits belonging to each species is something
which permits inductive generalizations, although they are not exceptionless. The
second strategy also suggests that one can maintain this view without essentialist
commitments; species and higher taxa are rather groups of individuals that share a
cluster of properties maintained by causal homeostasis (e.g. Boyd 1999). Finally the
third strategy can be traced back to a further reply which Grene herself gave to Hull
which is that there is no inconsistency in taking species as both kinds with individual
members and as individuals with temporal parts (1989). This thought has later been
developed by philosophers that suggest species are in fact kinds with historical
essences (Griffiths 1999; Okasha 2002). Interestingly all these options have recently
been challenged by Michael Devitt (2008) who has garnered much attention by
arguing that biological intrinsic essentialism should be resurrected!

II. How dominant is genetic selection and inheritance?

Stephen Jay Gould has compellingly shown how Dobzhansky and Simpson gradually
“hardened” their views about the synthesis (1983). The role of natural selection and
hence of adaptations at the expense of genetic drift became all the more dominant in
their writing, moreover, selection at the level of molecular genetics also came to
dominate over all other possible levels of selection.

Toward the end of the 1970s a clear divide between hard adaptationism and its critics
emerged in evolutionary biology as well as within philosophy of biology. We can
situate the divide in two key publications at the time. First, ethologist Richard
Dawkins’ publication of The Selfish Gene gave a forceful expression of the hard
adaptationist programme by offering an unreservedly gene-centred approach to
evolution (1976). Dawkins argued that there is good reason to privilege genetic
selection over other levels due to the faithfulness by which genes can be copied. In
virtue of this genes are genuine replicators. Such faithfulness in replication, Dawkins
argued, matters if there are to be adaptations and cumulative selection where small
improvements can be retained over time. Of course genes can only replicate by
forming collectives or interactors – what are usually called organisms. However,
organisms and phenotypes are ephemeral as they themselves are not capable of
genuine replication. The simple and clear logic of the gene’s eye view of evolution
also attracted more philosophers to use this framework for explaining, for example,
sexual selection (Cronin 1991), human morality (Dennett 1995), and even the
scientific process itself (Hull 1988),
The second crucial publication was instead one of the most forceful and influential
rejoinders to adaptationism: “The spandrels of San Marco and the Panglossian
Paradigm” by Gould and Richard Lewontin (1979). Some of the points raised were
mainly ontological (how likely are adaptations?) and others, mainly epistemological
(how can we be sure that a putative trait is an adaptation?). We cannot assume, they
argued, that a trait’s current functional role is a guide to the past, nor that it is the
reason for the trait’s existence; things that are currently adaptive (e.g. human literacy)
may not be adaptations, and, conversely, adaptations may not be currently adaptive
(e.g. human taste for sugar). Moreover one should be wary of adaptationism since for
an organism in a given environment there are often competing demands and
“optimal” solutions (or “optimal” genes) might not be retained. Finally, many traits
may be precisely like the “spandrels”– biproducts of adaptations propre. Hence, they
concluded, adaptationist explanations should not be a matter of so-called Panglossian
faith5; alternative hypotheses, such as drift and biproducts also need to be entertained.
As a compelling application, philosopher Elisabeth Lloyd has argued that
adaptationist bias pervades thinking of the female orgasm when evidence rather
indicates that it is a by-product of shared developmental pathways among males and
females (2005).

Not only were many philosophers heavily influenced by the adaptationist critique of
Gould and Lewontin, both biologists both taught and actively collaborated with
several individuals who became leading philosophers of biology, such as Robert
Brandon, Richard Levins, Elisabeth Lloyd, Elliott Sober and Will Wimsatt. A
common theme amongst these writers was to soften, rather than harden the MS by
postulating different traits and different levels of selection than those that a gene’s-
eye view would predict.

Lloyd and Gould (1993) built on the core element of population thinking that we saw
was stressed already by Dobzhansky; namely that greater variation of a trait within a
natural population – rather than between populations – is typically a condition for
natural selection. Based on the paleontological record they observed that this may also
lead to a kind of species selection. They argued that those species that are divided into
sub-populations where members can explore many more options in different
environments, will also have the best chance to respond to environmental change. In
that sense they predict that evolution would select for phylogenetic plasticity – a trait
that is selected amongst different species – where each species is a population – and
which is not necessarily adaptive at the genetic or individual level.

This latter example would be a special case of group selection that more generally
was making a return to philosophy of biology especially during the 1990s. Previously
many had been persuaded by William Hamilton’s explanations of altruistic behaviour
in terms of inclusive fitness. Hamilton (1964) had argued that seemingly altruistic
behaviour should not be explained by group selection, i.e. by the “group of altruists”
benefiting. Like many others, he thought that ultimately such selection would be
undermined by the possibility of individual defection. Instead he argued for a version
of the gene’s-eye view in explaining altruistic behaviour toward kin and those prone

5
A reference to Voltaire’s Candide where the character, Dr. Pangloss consistently asserts his
confidence in the world we live in being the best of all possible worlds.
to reciprocation via the benefits of the chance of reciprocation or the chance of genes
shared amongst kin propagating.

The tide turned somewhat when biologist David Sloan Wilson and philosopher Elliott
Sober developed a framework for multi-level selection, culminating in their
influential defence of group selection (1998). In brief, say we have different groups of
bonobos (Pan paniscus) that vary according to their trait of food sharing. In their
mathematical models, Wilson and Sober showed how those groups with more
pronounced food sharing traits will “reproduce” at greater rate than those who share
food less or not at all. Crucially this remains true in their model even if it is more
beneficial to the individual bonobo to keep the food to herself. The upshot of their
model is hence to offer a more general explanation of altruistic behaviour than what
the gene-centred view can offer. Slightly simplified, their appeal is to count groups
according to traits in virtue of which they share a common fate.6

You may have noticed that neither of these models of group selection depart from
adaptationism per se, in the sense that they still seek legitimate adaptations, albeit at
different levels. However, during the 1990s philosophers, psychologists and
developmental biologists teamed up to develop a more thoroughgoing alternative to
the MS. Its first guise was developmental systems theory (DST) (Griffiths & Gray
1994). Central to DST was to extend the notion of inheritance in a similar way as
group selection had extended the notion of genetic selection. Both MS and the gene’s
eye view can rightly be said to ignore what happens between genotype and
phenotype. DST in contrast embraced a wide range of heritable developmental
resources that contain information allowing the organism to reconstruct a life cycle
(Oyama et al 2001, 3-4). Two key assumptions of DST are made here. One is that a
developmental system has the capacity to carry information about what it has been
selected to represent – much like many think genes do (Maynard Smith 2000). The
other assumption confronts adaptationism itself. According to DST the organisms do
not merely passively respond to problems posed in their environment; for example,
many organisms build nests and provide a “nursery” for their offspring, thereby
directing or at least biasing the selection pressures in important ways. The phrase,
niche construction, was coined to describe this phenomenon, but whether it is a
genuine evolutionary force in its own right remains controversial (for a discussion see
Laland & Sterelny 2006).

Have the collaborations of DST been successful in overturning the hardening of the
Modern Synthesis or the popularity of Dawkins’ gene’s-eye view of evolution? I
would say it depends on whom you ask. First, much of the work of DST is now
subsumed by the broader paradigm of the Extended (evolutionary) Synthesis that like
DST embraces niche construction, epigenetic inheritance, group and multilevel
selection – or simply non-genetic inheritance and non-genetic selection (Mameli
2004). But while I think it is fair to say that this extended MS and partly reformed
perspective on Darwinian evolution is rather mainstream amongst philosophers these
days, it has yet to become biological orthodoxy. Indeed this is probably what
continues to fuel philosophical engagement with the issues.

6 Admittedly, this is a rather less intuitive way of counting populations than counting organisms or
species.
III. Culture annexed?
The last section naturally leads us to the question of how far the MS can be extended.
Can human culture also be Darwinized and synthesized along with the rest?
Interestingly, although advocates of the MS hardened their stance about genetic
selection, they were not averse to including culture in the framework of evolutionary
thinking. Culture, after all, is the natural continuation of the central questions of MS
that I’ve described here. If the questions about the nature of species are directed to our
own species and its nature, our capacity for creativity and culture is, if not unique, at
least one of our most conspicuous capacities. Surely our systems of inheritance and
evolutionary forces must have something to say about this.

At the same time, evolutionary thinking about culture and the diversity of human life
has had to confront the most significant and often harmful aberrations of genetic
explanations, first among which is eugenics. This was evident in the resounding
response from biologists to the eugenic programme that came both during and after
the 1959 centennial celebration of the publication of Darwin’s Origin of Species at the
University of Chicago, where the diversity of cultures was heralded as an important
achievement in human evolution (Grene & Depew 2004, 331 ff). Conrad H.
Waddington, a developmental biologist who also was an inspiration to the DST,
suggested that there might also be a separate system of cultural transmission, and
Dobzhansky, who disagreed with Waddington on much, published a book arguing
that our genes and culture evolve “hand in hand” (1962, 75).

This mid-century consensus amongst biologists was nevertheless broken by E.O

Wilson’s publication of Sociobiology: The New Synthesis and the follow-up On
Human Nature. In the latter publication, Wilson appears to flirt with eugenic notions
like using knowledge of human genetics and molecular engineering to possibly
change our nature to one of higher intelligence and creativity (1978, 208). In fact, it
was E.O. Wilson who was the chief antagonist of anti-adaptationist critiques in the
1980s and onward. In the background there were also strong ideological
disagreements. Wilson was an avid anti-Marxist who viewed culture with suspicion,
as a space where fanaticism could run unleashed. On the other side, there were people
like Waddington, Lewontin and Levine who were committed Marxists. However, the
ideas of Wilson were far from uniformly rejected, especially amongst some rebellious
anthropologists and psychologists that set out a programme of evolutionary
psychology. As the name suggests, they focused on adaptationist explanations of
universal psychological traits or “cognitive programmes” that in their view tell us,
say, what to be fearful of and how to react to a partner infidelity. In fact, as part of
their methodology, they suggested we should look for indications of how traits may
not fit or with our current modern environment as indicators that the trait would have
evolved in the Pleistocene (Tooby & Cosmides 1990). Evolutionary psychologists on
the other hand paid little attention to the creation of that modern environment itself
and we might think that this was the main crux with this programme. If we are so
unfit for our modern culture, how can it possibly have evolved?

In parallel with evolutionary psychology, other Darwinian research programmes have

tried to address precisely this issue by picking up the proposals of Dobzhansky and
Waddington’s: what if culture can constitute a separate system of inheritance from
genetics. Dawkins (1976) and Dennett (1995) for example proposed to do this by
treating cultural entities such as melodies, religious and moral ideas as analogous to
genes – that is, replicator memes – which have effective strategies for spreading
amongst the minds of humans. Many however felt that memetics would leave too
much of human agency and action out of the picture. Accordingly, more recently it
has been argued that the notion of replication should be replaced by the broader
notion of reproduction. Because reproduction need not be high fidelity it can
encompass a broader spectrum of social learning where cultural traits are transmitted
by observing or imitating habits (Godfrey Smith 2009; Heyes 2012). The proposal
dovetails with one of the ideas we reviewed about species; namely that cultural kinds
(e.g. religions, ideologies and folk wisdom), because their instances are united by
cultural reproduction, could be kinds with historical essences (Millikan 1999;
Godman 2015).

If cultural traits are also to be able to undergo (cultural) selection, however, several
additional conditions to reproducibility arguably have to be met. One is to show that
reproduction can suffice for the gradual improvements necessary for cumulative
evolution (Sterelny 2006). Another is to demonstrate how genetic selection interacts
with, rather than counteracts, the effects of culture. So far mathematical models and
simulations for gene-culture co-evolution have been important parts of making this
argument (e.g. Richerson & Boyd 2005). Tim Lewens suggests that using these
population models to study cultural change can be understood as a kinetic view of
culture (2015). Just as the MS relied on changes in gene frequencies within
population as a means of detecting processes of natural selection, the kinetic view of
culture relies on changes in learning rules within a population as guide to when a
cultural trait will evolve. Brian Skyrms has for instance investigated how a rule like
“imitate your best neighbour” would allow for a population to change quite rapidly
(2003). However, as is frequently complained about model-based science, it is still an
open question how often, if ever, these conditions are met in the real world. But it is
easy to see how research on cultural evolution is engaging—somehow or another,
human culture must have evolved.

IV. A return to a philosophy of nature?

The fascinating ideas involved in the Modern Synthesis have permeated not only
philosophy of biology, but also many other areas in philosophy, like philosophy of
language, metaethics and epistemology, which I have not been able to cover in this
short essay. In fact, although I have spoken of the exchange, inspiration and
collaborations between philosophers and biologists, I am struck that this might
slightly have mis-characterised the enterprise. Instead of philosophy of biology
becoming a new specialisation uniting philosophers and biologists, what we may have
witnessed is instead a modern reinvention of the Aristotelean tradition of a natural
philosophy or a philosophy of nature, where an essential part of any study of the
living world is its philosophical interpretation (see also Godfrey Smith 2009, p. 3).
And perhaps, to put things pointedly, it may be that for progress on understanding
evolution, its key concepts, and their legitimate extension, the increasing institutional
demands on specialisation, found both within biology and philosophy, is often an
obstacle.

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