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The Philosophical Quarterly Vol. 0,No.02022
ISSN 0031-8094 https://doi.org/10.1093/pq/pqac047
HOW GENE–CULTURE COEVOLUTION CAN—BUT
PROBABLY DID NOT—TRACK MIND-INDEPENDENT
MORAL TRUTH
ByNathan Cofnas
I argue that our general disposition to make moral judgments and our core moral intuitions are likely the
product of social selection—a kind of gene–culture coevolution driven by the enforcement of collectively
agreed-upon rules. Social selection could potentially track mind-independent moral truth by a process
that I term realist social selection: our ancestors could have acquired moral knowledge via reason and
enforced rules based on that knowledge, thereby creating selection pressures that drove the evolution
of our moral psychology. Given anthropological evidence that early humans designed rules with the
conscious aim of preserving individual autonomy and advancing their collective interests, the theory of
realist social selection appears to be attractive for moral realists. The goal of evolutionary debunking
arguments should be to show not that our moral beliefs are the product of natural selection, but that
realist social selection did not occur.
Keywords: evolutionary debunking arguments, moral realism, natural selection,
social selection, cultural evolution, gene–culture coevolution.
I. INTRODUCTION
Moral realists believe that there are moral truths that are objective, non-
relative, and—according to most realists—knowable (Shafer-Landau 2003;
Huemer 2005; Tersman 2006). As Shafer-Landau (2003:15) puts it, moral
truths are ‘stance-independent’: ‘the moral standards that fix the moral facts
are not made true by virtue of their ratification from within any given actual or
hypothetical perspective’. Non-naturalist realists add that moral facts cannot
be reduced to natural ones (ibid.: ch. 3).
Sceptics about moral realism frequently employ ‘debunking arguments’.
Debunking arguments present a story about the causal origins of a belief as an
undermining defeater (Kahane 2011:106). They take the following form:
Causal premise:Ccaused S’s belief that p.
Epistemic premise:Cdoes not produce reliable beliefs about p.
C
The Author(s) 2022. Published by Oxford University Press on behalf of The Scots Philosophical Association and the University of St
Andrews. This is an Open Access article distributed under the terms of the Creative Commons Attribution License
(https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium,
provided the original work is properly cited.
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2NATHAN COFNAS
Conclusion: In light of the knowledge that Ccaused the belief that p,Sis
unjustified in believing that p. (Kahane 2011:106;Nichols2014:731; Sauer
2018:30; Egeland 2022:4).
Debunkers disagree about what makes a causal process epistemically defec-
tive in the way required for a debunking argument to succeed (Sauer 2018:31).
The idea that debunking arguments show the targeted belief to be insensitive (S
would have believed that pvia Ceven if not-p)orunsafe (Scould easily have
been wrong about p) is disputed by those who reject sensitivity or safety as
necessary for justification (Egeland 2022:7–9). Another possibility is that a
C-caused belief that pis debunked by showing that Cdoes not presuppose
the truth of p(e.g., Harman 1977; Ruse and Wilson 1986;Joyce2006:220).
This, too, runs into trouble. Nichols (2015:98) observes that, for example, the
explanation for his (presumably justified) belief that Hume will be studied at
universities a century from now does not presuppose that the belief is true.
If Ccaused S’s belief that p, we cannot undermine S’s justification for the
belief that pmerely by showing that C‘does not bear on the truth of p’
(Vavova 2018:136). Such ‘irrelevant influences’ are pervasive, but many are
epistemically unproblematic (ibid.). The fact that S’s parents met each other is
part of the causal story for all of S’s beliefs. Maybe Scame to believe that ponly
after being gifted an authoritative book about the subject for their birthday. But
(in most cases) the discovery of influences like these—your parents met each
other, you received a book on your birthday—is not debunking. In order to be
debunking, the discovery that Ccaused S’s belief that phas to provide some
reason to think that Sis likely to be mistaken (White 2010:604;Vavova2018).
There are many ways in which the causal origins of a belief can be epistemically
defective. Egeland (2022:13) suggests—quite plausibly—that ‘there most likely
does not exist any special debunking principle that unifies all valid debunking
arguments’.
Evolutionary debunking arguments (EDAs) in ethics purport to debunk
moral beliefs (realistically construed) by attributing them to natural selection.
Natural selection is said to be epistemically defective with respect to mind-
independent moral truth. Street (2006:125,143) argues that it would be an
‘incredible’, ‘implausible coincidence’ if natural selection, which aims at fitness,
endowed us with ‘evaluative tendencies’ that happen to correspond with mind-
independent morality. Although evolutionary debunkers often refer to their
target as ‘moral realism’, there are probably versions of naturalistic realism that
are not undermined by EDAs (e.g., Sterelny and Fraser 2017). The terminology
is used somewhat inconsistently. For example, Street (2006) describes herself as
a critic of ‘realism’, but she advocates constructivism, which is often classified
as a kind of (naturalistic) realism. Without attempting to sort this out, I will use
the term ‘mind-independent morality’, ‘objective morality’, or simply ‘moral
realism’ to refer to the versions of realism that are susceptible to EDAs.
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GENE–CULTURE COEVOLUTION AND MORAL TRUTH 3
Some critics of EDAs counter that even if natural selection aims at fitness
rather than moral truth, beliefs might be both adaptive and moral (Enoch
2010; Wielenberg 2010;Skarsaune2011). For example, if survival is objectively
good, natural selection will endow us with the objectively correct moral belief
that survival is good even though it is not tracking moral truth per se (Enoch
2010). Various other challenges have been raised to EDAs (see Vavova 2015).
Having acknowledged the controversy, I will not attempt to defend evolution-
ary debunking in general. The EDA that I will propose does not raise any
novel epistemic questions. For the purposes of this paper, I assume for the sake
of argument that natural selection is epistemically defective with respect to
objective moral truth in the way required to run a debunking argument.
EDAs in ethics can be ‘selective’ or ‘global’ (Sauer 2018:ch.3). Selective
EDAs attribute some but not all of our moral beliefs to natural selection. For
example, Greene (2008:66–76) and Singer (2005:347–51) argue that we should
reject our deontological moral intuitions as the tainted product of natural
selection, while our utilitarian intuitions remain undebunked and therefore
potentially reliable. Global EDAs target moral realism itself, and come in two
main varieties, which are most associated with Street (2006)andJoyce(2006),
respectively.
According to Street (2006), natural selection (indirectly) shaped the content
of our core moral beliefs. In order to prompt adaptive behaviour—protecting
one’s life, caring for one’s children, returning favours (when this cultivates ben-
eficial cooperative relationships), punishing those who have inflicted deliberate
harm on oneself (when this will discourage them or others from inflicting fur-
ther harm), and so on—natural selection endowed us and some other animals
with ‘basic evaluative tendencies’ to judge that such a behaviour is ‘called for’
(ibid.: 117). In other animals, these evaluative tendencies exist only in a ‘proto’
form (ibid.: 119). Our close relatives such as chimpanzees, who have been
subject to similar selection pressures, have proto evaluative tendencies that are
analogous—or homologous—to ours. Humans have the capacity to articulate
our intuitions about what is ‘called for’ with language in order to generate
‘full-fledged’ evaluative judgments such as it is good to return favours,criminals
should be punished,orwhat George did was wrong (ibid.: 118). But these full-fledged
judgments are rooted in evaluative tendencies—which on Street’s account
seem to be equivalent to our innate moral intuitions—that were implanted in
us by natural selection simply because they increased inclusive fitness in the
ancestral environment. She concludes that ‘one enormous factor in shaping
the content of human values has been the forces of natural selection, such that
our system of evaluative judgements is thoroughly saturated with evolutionary
influence’ (ibid.: 114). In light of this discovery, our moral beliefs (realistically
construed) are all unjustified. (Street’s theory about the origin of our moral
beliefs is controversial; see, e.g., Machery and Mallon 2010;Parfit2011:535–8;
Deem 2016; Isserow 2019;Cofnas2020a;LevyandLevy2020.)
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4NATHAN COFNAS
In contrast to Street (2006), Joyce (2006) does not emphasize the claim that
natural selection (directly or indirectly) influenced the content of our moral
beliefs. Instead, he argues that the psychological disposition to acquire (re-
alist) moral beliefs in general is the product of natural selection. We would
be disposed to acquire beliefs about right and wrong even if there were no
mind-independent truth of the matter (ibid.: 220). The discovery of this fact
undermines the justification for all of our moral beliefs (realistically construed).
I argue that the evolutionary explanations of our moral beliefs that feature in
both Street- and Joyce-style EDAs may be mistaken in ways that undermine the
arguments. Section II outlines the concept of gene–culture coevolution, and
concludes that some popular gene–culture coevolutionary theories of morality
can easily be incorporated into standard EDAs. Section III argues that our
moral psychology is the product of a special kind of gene–culture coevolution
called social selection. Social selection occurs when members of a species uncon-
sciously direct their own evolution toward a mentally represented endpoint.
Anthropological evidence suggests that early humans drove the evolution of
our moral psychology by collectively establishing rules to preserve individual
autonomy and promote group interests, and imposing fitness-reducing pun-
ishments on rule violators. This raises the possibility of realist social selection—the
theory that social selection led us to evolve an innate tendency to make moral
judgments that line up with mind-independent moral truth. Our ancestors
could have established rules to protect autonomy and promote their collective
interests because they recognized the mind-independent moral imperative of
these rules via reason. Section IV argues that, if our moral beliefs are in fact the
product of social selection, the goal of EDAs should be to show that realist so-
cial selection did not occur. I offer a debunking account of the social-selection
process that shaped our moral faculty.
II. GENE–CULTURE COEVOLUTION AND EVOLUTIONARY
DEBUNKING ARGUMENTS
Gene–culture coevolution refers to the phenomenon whereby cultural evolu-
tion affects genetic evolution and vice versa (Durham 1991; Richerson and
Boyd 2005; Lewens 2015; Henrich 2016). Some scientists have defended gene–
culture coevolutionary accounts of the evolution of morality, which appear
strikingly different from the theories that feature in popular EDAs, partic-
ularly Street’s (2006). Do these theories pose a challenge for the debunking
project? Before considering this question, we should be clear on how gene–
culture coevolution relates to natural selection as traditionally understood.
Following Lewontin (1970), Sober (2000:36) writes that ‘[n]atural selection
occurs when there is heritable variation in fitness’. The heritability requirement
means that there must be a mechanism of inheritance that makes offspring
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GENE–CULTURE COEVOLUTION AND MORAL TRUTH 5
resemble their parent(s). We now know that—at least in the sorts of cases
that concerned Darwin—the mechanism of inheritance is DNA. But natural
selection does not require DNA specifically. It requires that offspring resemble
their parent(s) regardless of how this resemblance is brought about (Lewontin
1970:1; Sober and Wilson 1998:107).
As a matter of fact, DNA is not the only way in which organisms inherit traits
from their parents. Humans and (to a very limited extent) some other animals
can also inherit traits via cultural transmission. Culturally transmitted traits, or
‘cultural variants’—ideas, skills, beliefs, attitudes, and values (Richerson and
Boyd 2005:63)—can be passed from parents to children by learning,which
involves imitation, emulation, or guided discovery (Sterelny 2012). Since these
heritable cultural variants can both vary and affect fitness, adaptive variants
can spread due to natural selection acting on individuals or groups (Sober and
Wilson 1998:149–54; Richerson and Boyd 2005:13–4,68–79). As Richerson
and Boyd (2005:76) say, ‘[t]o the extent that people acquire beliefs from their
parents, natural selection acts on culture in almost exactly the same way as it
does on genes’.
DNA transmission is (in animals) only vertical (parents to offspring). In
contrast, cultural transmission can also occur horizontally (organism to other
members of its generation) and obliquely (adults to non-offspring children)
(Cavalli-Sforza and Feldman 1981). A consequence of this is that the variants
themselves can undergo Darwinian evolution in the cultural realm (Richerson
and Boyd 2005:76;cf.Dawkins1976/2016:ch.11). That is, variants can be
more or less fit in the sense that they are more or less likely to be copied by
other people and transmitted down the generations.
This brings us back to gene–culture coevolution. Culture—the panoply of
cultural variants—is a fitness-relevant aspect of the enculturated organism’s
environment. As such, it can impose selection pressures that favour certain ge-
netic variants. The selection pressures acting on individuals depend in complex
ways on the cultural variants of others in their society, and on the variants that
they themselves adopt. Genetic adaptations make culture possible, and culture
in turn creates conditions that favour new genetic adaptations, which affect
culture, and so on.
Whereas we cannot choose our DNA, we have some liberty to choose
among competing cultural variants. This creates selection pressure for the
tendency to identify and adopt the most fitness-promoting cultural variants
in our environment. While the variants themselves are undergoing their own
Darwinian selection, the individuals who participate in the culture are subject
to Darwinian selection for the tendency to choose those that are best from
their own fitness perspective. According to cultural evolutionary theorists,
this led us to evolve certain adaptive, innate learning biases. Richerson and
Boyd (2005:Table3.1) have, based on a combination of modelling work and
empirical evidence, identified three categories of learning biases under the
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6NATHAN COFNAS
headings ‘Content-based (or direct) bias’, ‘Frequency-based bias’, and ‘Model-based
bias’. These refer, respectively, to our preference for cultural variants that
are (a) intrinsically appealing (due to their perceived benefit, or because ‘the
structure of cognition makes some variants easier to learn or remember’), (b)
common or rare (we tend to conform to the majority), or (c) exhibited by
people with certain characteristics (we tend to copy others who are successful,
prestigious, or similar to ourselves).
Now, we can consider whether the phenomenon of cultural evolution or
gene–culture coevolution has implications for EDAs in ethics.
Standard EDAs typically claim that natural selection directly or indirectly
influenced our moral beliefs by acting on genetically transmitted traits. One
might suppose that EDAs lose their force if the traits in question are culturally
transmitted. Parfit appears to defend this view in the following passage:
When Street and others claim that our normative beliefs were mostly produced by evo-
lutionary forces, these writers are in part referring to cultural evolution. Some normative
beliefs became more widely spread when and because communities of people with these
beliefs were more likely to be successful. It is much less clear how we should assess the
claim that certain normative beliefs were in this way, not reproductively,butsocially or cul-
turally advantageous. It is less clear, for example, whether and how such explanations of
our normative beliefs should be assumed to debunk or undermine these beliefs. When the
acceptance of certain normative beliefs made some community or culture more likely
to survive and flourish, this fact does not as such cast doubt on the truth or plausibility
of these beliefs. Such explanations of our normative beliefs do not obviously, in Street’s
phrase, contaminate these beliefs. (Parfit 2011:537)
If, when Parfit refers to moral beliefs that proliferate because they make a
community ‘likely to survive and flourish’, he has in mind something like
natural selection, then his reasoning seems to be based on a straightforward
mistake. If explaining a moral belief in terms of natural selection debunks it,
then, it does not matter what the underlying mechanism of transmission is—
whether it is genetic, epigenetic, cultural, or anything else we could imagine.
As discussed above, natural selection is not defined by genes or DNA. Parfit
says that ‘[w]hen the acceptance of certain normative beliefs made some
community or culture more likely to survive and flourish, this fact does not
as such cast doubt on the truth or plausibility of these beliefs’. This statement
is correct in a limited sense, but it misses the point of the EDA. If the causal
explanation of our moral belief that pis that societies in which people believed
that pwere more likely to survive and flourish (i.e., were favoured by natural
selection), then our belief is just as ‘contaminated’ as any other moral belief
explained by natural selection.
Parfit’s claim that when Street and other debunkers refer to ‘evolutionary
forces’ they are ‘in part referring to cultural evolution’ is probably wrong, and
is certainly wrong with respect to Street. Street (2006:118) makes it clear that
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GENE–CULTURE COEVOLUTION AND MORAL TRUTH 7
she is talking about natural selection acting on traits that ‘must be genetically
heritable’. But that is not important. The possibility that natural selection
acts on culturally as well as genetically transmitted moral beliefs is easily
accommodated by standard EDAs.
Does the fact that gene frequencies change in response to cultural change
and vice versa pose a challenge for EDAs? Cultural evolutionists in the Boyd-
and-Richerson tradition have developed mathematical models that suggest
that morality—or at least certain core elements of it—evolved in a different
way than Street- or Joyce-style debunkers propose.
Cultural evolutionary models assume that cultural variants are distributed
in a population, and individuals with the learning biases mentioned above
choose whom to imitate. Modelling work shows that, in populations of social
learners, stable behavioural patterns emerge as a by-product of the learning
biases, particularly when the conformist bias is combined with punishment
of deviants (Boyd and Richerson 1992; Chudek et al. 2013:442). A key as-
sumption of these models is that stable behavioural patterns, or norms, are not the
product of conscious design. Groups vary in their initial distributions of norms, and
individuals choose whom to imitate based on their learning biases, not based
on an envisioned endpoint. Cultural evolutionary theorists in this tradition
emphasize their view that people are simply unable to anticipate the conse-
quences of different norms, so any attempt to deliberately design a system
of group-beneficial norms is futile (e.g., Henrich 2006). Many norm equilib-
ria are possible, and the vast majority are detrimental, not beneficial, to group
fitness (Boyd and Richerson 1992; Henrich and Boyd 2001:86; Chudek and
Henrich 2011:222). So how do group-beneficial norms ever evolve? According
to the modellers, a small number of groups will randomly stumble on group-
beneficial equilibria—namely, prosocial norms enforced by punishment—and
these practices will be favoured by cultural group selection (Boyd and Rich-
erson 2002). After group-beneficial prosocial norms spread by cultural group
selection, in a process of gene–culture coevolution people become genetically
adapted to be innately receptive to prosocial practices and values (Henrich
and Boyd 2001; Chudek and Henrich 2011; Chudek et al. 2013; see Cofnas
2018:311).
According to the story given above, our innate moral intuitions are the
product of three stages of Darwinian selection. First, there is selection among
competing cultural variants, namely, different norms. Secondly, there is cultural
group selection, which mostly involves natural selection acting on groups.
Thirdly, natural selection acts on individuals via selection pressures created by
the enforcement of prosocial norms.
Street-style EDAs assume that our moral beliefs are the product of natural
selection acting on traits that (in Street’s words, as quoted above) ‘must be
genetically heritable’. The Boyd-and-Richerson account of morality suggests
that moral evolution was driven in large part by natural selection acting on
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8NATHAN COFNAS
culturally transmitted beliefs, which only later become genetically incorpo-
rated. This might seem to pose a challenge for EDAs, but in fact it does
not. The epistemic premise of EDAs is that natural selection does not track
mind-independent moral truth. All three stages of the evolution of morality
that feature in Boyd and Richerson’s theory involve natural selection. It would
be an ‘incredible coincidence’ (to use Street’s [2006:125] expression), if the
process they describe produced moral beliefs that happened to align with
mind-independent moral truth.
III. THE SOCIAL-SELECTION ACCOUNT OF MORALITY
In the previous section I argued that gene–culture coevolutionary theories of
morality do not necessarily pose a threat to standard EDAs. Here, I describe
Boehm’s (1999,2012) account of the evolution of morality and argue that it
does pose a genuine threat. According to Boehm, morality evolved by a kind
of gene–culture coevolution called ‘social selection’: our ancestors consciously
designed blueprints of desirable societies, enforced behaviour demanded by
the blueprint, and in this way artificially selected people to have (realist) moral
intuitions. I will argue that social selection, whether it is conceived as different
from, or as a special case of, natural selection, could potentially track mind-
independent moral truth. Moral beliefs produced by social selection cannot
be undermined—at least not directly—by standard EDAs.
Regarding the evolutionary psychological theories upon which her EDA
is based, Street (2006:113) writes: ‘while I am skeptical of the details of the
evolutionary picture I offer, I think its outlines are certain enough to make
it well worth exploring the philosophical implications’. For the purposes of
the present paper, we can take the same attitude toward the social-selection
theory of morality. The details are uncertain, but the key claim is highly plau-
sible. Human societies have been organized around deliberately engineered
social codes for an evolutionarily significant amount of time. For thousands
of generations, individuals’ fitness was to some degree tied to their ability to
conform to these codes. This created selection pressures favouring individu-
als who internalized the norms of their society—that is, who had a conscience
(Boehm 2012)—and were perhaps also disposed to acquire certain specific
moral beliefs (Cofnas 2018:315–6). Contra Boyd and Richerson, these norms
were intentionally designed by agents to promote their collective goals. I will not
try to adjudicate between the various accounts of morality offered by evolu-
tionary psychologists and cultural evolutionary theorists. Rather, I will explore
what the philosophical implications would be if we accept the social-selection
account.
Boehm’s (1999,2012) theory goes as follows. Until around 250,000 years
ago, human society resembled that of chimpanzees (Pan troglodytes), with each
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GENE–CULTURE COEVOLUTION AND MORAL TRUTH 9
group being dominated by an alpha male and his coalition partner(s). Like
chimps, subordinate people sometimes formed coalitions to challenge and even
overthrow the alpha. But although one alpha might be dethroned, a new one
would always take his place. The social system itself remained stable. Alphas
enforced rules designed almost entirely to benefit themselves. They were often
resented, but their subordinates lacked the ability and inspiration to change
the system. At some point, however, there was a revolution. Our ancestors
evolved the requisite levels of intelligence, foresight, and ability to communicate
and cooperate, which allowed them to conceive and collectively execute a
plan to redesign their political system. Subordinate males banded together
to overthrow their alphas and establish a ‘reverse dominance hierarchy’ with
themselves occupying the leadership position. Instead of an alpha ruling for
his own benefit, the new coalitions of the majority of men ruled for their
collective benefit. They designed an explicit ‘blueprint’ (Boehm 1999:12,193–4)
for a desirable society, which called for political egalitarianism (at least among
adult men), measures to protect individual autonomy, and various prosocial
behaviours. They punished those who failed to live up to the demands of the
blueprint, and rewarded those who did, in ways that affected fitness.
‘[S]ocial control by groups’ was ‘initially nonmoralistic’ (Boehm 2012:15),
but it created unique selection pressures in the human lineage that led to the
evolution of a conscience, which Boehm defines as the tendency to ‘personally
[identify] with community values, which means internalizing your group’s rules’1
(ibid.: 113). When rules are made and enforced by an alpha (as among chimps),
they tend to be followed only when the alpha is watching or otherwise likely
to discover deviance. When rules are made to promote collective well-being,
however, almost all group members have a stake in enforcing them, and vio-
lations witnessed by one person can be made public to everyone by means of
language. Many generations of highly effective law enforcement favoured indi-
viduals who were particularly inclined to follow group rules. The evolutionary
function of the conscience is to compel us to follow such rules—or at least
to think very carefully before transgressing (ibid.: 114–5). In Boehm’s words:
‘prehistorically humans began to make use of social control so intensively that
individuals who were better at inhibiting their own antisocial tendencies, either
through fear of punishment or through absorbing and identifying with their
group’s rules, gained superior fitness’ (ibid.: 17). As people evolved stronger
consciences, ‘group punishment increasingly became driven by moral out-
rage’ (ibid.: 88), and consequently more intense and effective. In addition to
1Under certain conditions people’s consciences can lead them to reject their group’s rules. But
the evolutionary function of the conscience is to prompt conformity. For most of our evolutionary
history (including to some extent today), seriously opposing the values of one’s community would
have elicited swift, fitness-reducing punishments.
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10 NATHAN COFNAS
negative social selection (i.e., punishment), rule conformists could also have
been favoured as marriage or exchange partners.
I cannot do justice to all of the evidence from multiple sources that Boehm
provides to support his theory. But his main empirical evidence comes from
studies of 20th-century nomadic foragers who lived more or less as our ances-
tors in the Pleistocene. These societies are marked by striking commonalities.
Without exception,2from Australia to South America to Africa, they all en-
force strict political egalitarianism among men. Some nomadic foragers have
nominal leaders, but neither the leader nor any other adult male is allowed
to issue direct orders to another adult male. ‘These foragers very predictably
share a core of moral beliefs with an egalitarian emphasis on every hunter’s
being a political equal, while the political positions of women as non-hunters
are much more subject to diversity’ (ibid.: 80). They also have rules requiring
that food, particularly meat from large game, be shared among all group mem-
bers. They are consciously aware that their rules have the effect of preserving
individual autonomy and protecting people from bad luck (e.g., an unsuccessful
hunt). Nomadic foragers all over the world employ the death penalty against
recalcitrant rule breakers—almost always men—and especially against men
exhibiting alpha-like behaviour (ibid.: Table 1). Approximately half of docu-
mented death penalty cases across these societies involve males who were
intimidating their groups...by greedily or maliciously using supernatural power to seri-
ously threaten the welfare or lives of others; by being far too ready to kill, repeatedly, out
of greed or anger; by otherwise managing to seriously dominate others; or (much more
rarely) by being aggressively insane. (ibid.: 85)
Other crimes inviting capital punishment include theft, cheating in the context
of meat sharing, taboo violations, and engaging in proscribed sexual behaviour.
However, the death penalty is a rare last resort. Most instances of rule violation
are dealt with by less serious (but escalating) punishments ranging from teasing
to shaming to ostracism to expulsion from the group (which can be a de facto
death sentence).
2Some critics of Boehm have pointed to examples of hunter–gatherer societies that are
not egalitarian. However, these hunter–gatherers are sedentary and/or combine hunting and
gathering with agriculture or trade with agriculturalists, and are therefore not pure nomadic
foragers. Wengrow and Graeber (2015:600) do not deny that nomadic foragers were egalitarian,
but they ‘propose a relationship between seasonality and the conscious reversal of political
structures’. According to their theory, some Upper Pleistocene hunter–gatherers alternated
seasonally between nomadic and sedentary social systems. During the nomadic phase they
were egalitarian, and during the sedentary phase they were (weakly) hierarchical. But even if
some Upper Palaeolithic hunter–gatherers sometimes deviated from nomadic foraging and strict
egalitarianism (see also Singh and Glowacki 2022), this would not undermine the claim that
social selection was a decisive factor in the evolution of morality, particularly in the beginning.
When our moral faculty initially evolved—and for a substantial period of time afterwards—our
ancestors presumably were nomadic foragers.
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GENE–CULTURE COEVOLUTION AND MORAL TRUTH 11
III.1 Social selection could track mind-independent moral truth
In principle, social selection could track mind-independent moral truth very
simply. The selecting agents merely have to enforce moral behaviour in re-
sponse to their recognition of the truth. Realists typically claim that we grasp
truths in domains such as mathematics and ethics as a by-product of our gen-
eral reasoning ability, which was favoured by natural selection to solve adaptive
problems in the ancestral environment (e.g., Huemer 2005:215–6;Parfit2011:
494–7; de Lazari-Radek and Singer 2012:16–8; Cuneo and Shafer-Landau
2014:427). Suppose, our ancestors acquired moral knowledge via reason and
imposed fitness-reducing punishments on those who refused to behave morally
or espouse moral views. In that case, we would have evolved a tendency to
make moral judgments and to have specific moral intuitions that dispose us
to acquire objectively true moral beliefs. We can call this the theory of realist
social selection.
As noted, both Street and Joyce claim that scientific explanations of our
moral beliefs do not assume the existence of mind-independent moral truth.
Although Street (2006) describes her challenge to moral realism as the ‘Dar-
winian Dilemma’, she argues that the dilemma does not have to be specifically
Darwinian. ‘Ultimately,’ she says, ‘the fact that there are any good scientific
explanations of our evaluative judgements is a problem for the realist about
value. . . .The best causal accounts of our evaluative judgements, whether Dar-
winian or otherwise, make no reference to the realist’s independent evaluative
truths’ (ibid.: 155). Alluding to our (realist) moral beliefs, Joyce (2006:220)con-
tends that any belief is rendered unjustified by ‘the discovery of an empirically
supported theory that provides a complete explanation of why we have that
belief while nowhere presupposing its truth’. But realist social selection is a
scientific theory of the evolutionary origins of our moral faculty—and a theory
that seems to have prima facie support—which does assume the existence of
mind-independent moral truth.
To spell out the theory of realist social selection more precisely:
1. Humans evolved a general reasoning ability for solving adaptive problems
in the ancestral environment, which (as an accidental by-product) allowed
them to grasp mind-independent moral truth.
2.Around250,000 thousand years ago our ancestors instituted rules to pro-
mote autonomy and collective well-being in response to their recognition
that autonomy and well-being are objectively good.
3. The enforcement of rules designed to promote autonomy and collective
well-being created selection pressures for a disposition to internalize group
norms and perceive them as true in a realist sense. It also led us to evolve
moral intuitions—for example, concerning equality—that dispose us (albeit
not inevitably) to acquire some specific, objectively true moral beliefs.
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12 NATHAN COFNAS
4. Under normal circumstances, our socially selected conscience and moral
intuitions are some of the main forces that shape our moral beliefs (real-
istically construed). We easily acquire the belief that, for example, people
are moral equals because we were socially selected to identify with norms
designed to ensure political equality.
5. Joyce-style evolutionary debunkers are right that we evolved a disposition
to acquire realist moral beliefs. Street-style debunkers are right that evolved
evaluative tendencies are an ‘enormous factor in shaping the content of
human values’ (Street 2006:114). Both are wrong to think that the evolu-
tionary process that produced our moral faculty was off track with respect
to mind-independent truth.
The theory of realist social selection assumes that (at least many of) the
norms that were enforced by early humans were objectively correct, or were
based on objectively correct values. This means that the theory will be at-
tractive mainly to realists who believe that the particular norms that were in
fact enforced among nomadic foragers were the correct ones. If we accept
the anthropological claim that nomadic foragers designed norms to promote
autonomy, equality, and collective survival, those who endorse the realist-
social-selection account must assume that these things are objectively valuable.
It is true that the moral codes of nomadic foragers did not impartially rec-
ognize the moral equality of all people. They varied greatly in their treatment
of women, and frequently deemed outgroup members to have a much lesser
moral status or even none at all. But this is not a serious problem for the realist
who regards such inegalitarianism as objectively immoral. There is no reason
to expect nomadic foragers to get everything right. Realist social selection only
requires that early humans grasped some core principles of morality (perhaps
imperfectly) and enforced rules in light of this knowledge.
If the theory of realist social selection is correct, one profound implication
wouldbethatwehavetwo ways of gaining moral knowledge. First, like our
Middle Palaeolithic ancestors, we can grasp moral truth via reason. The theory
assumes that the early humans who initiated the process of social selection
obtained knowledge of moral truth via reason, so this must be possible for us,
too. Secondly, we can rely (to some extent) on our evolved moral intuitions. The
realist-social-selection theory does not imply that moral knowledge necessarily
comes to us automatically or effortlessly via intuition. But, it does imply that
our moral intuitions (properly evaluated or interpreted) can be a reliable source
of moral knowledge. The reason is that our intuitions are the product of an
evolutionary process that tracked mind-independent moral truth.
IV. A NEW EVOLUTIONARY DEBUNKING ARGUMENT
To reiterate, the social-selection theory of morality says that early humans
possessed high levels of general intelligence and foresight as well as the ability
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GENE–CULTURE COEVOLUTION AND MORAL TRUTH 13
to make sophisticated calculations regarding their own and others’ perceived
interests. Using these cognitive abilities, they designed and enforced rules to
preserve individual autonomy and equality and to advance their collective
well-being (as they understood it). From the perspective of the realist, it may
seem like early humans grasped some key elements of mind-independent
moral truth and enforced rules in accordance with that truth, which caused us
to evolve a conscience and innate moral intuitions.
If evolutionary debunking is to succeed, a new EDA needs to be formulated
to target beliefs generated by a moral faculty that evolved due to social selection.
It needs to show that some causal factor(s) that had a decisive influence over
the social-selection process was not truth tracking, and therefore, in light of this
discovery, the beliefs generated by our evolved moral faculty are unjustified.
This can be done by pushing standard EDAs back a step. Instead of tar-
geting our beliefs by attributing them (or the underlying intuitions) to natural
selection, evolutionary debunkers can target the motivation of the social selec-
tors. The source of epistemic contamination was not natural selection acting
directly on our moral faculty as in Street- and Joyce-style EDAs. Rather, nat-
ural selection shaped the desires and dispositions of the social selectors, and
thus drove the evolution of our moral faculty indirectly.
Like other animals, humans were endowed by natural selection with drives
to perform (or avoid) certain actions and to bring about (or prevent) certain
states of affairs that tended to promote (or harm) fitness in the ancestral
environment. For example, natural selection gave us a drive to eat sugary
food, which led us to eat fruit when it was available. Natural selection gave us
a drive to keep our body within a narrow temperature range that is optimal
for survival. Because the individual fitness of primates is tied to their place
in a dominance hierarchy, natural selection endowed them with a dislike of
being dominated in order to motivate them to ascend the hierarchy, if possible
(Eibl-Eibesfeldt 1989:297; Boehm 1999:170,237–9). As primates, we inherited
this disposition.
Consider the process that led to the evolution of our moral faculty (ac-
cording to the theory of social selection). It began when people who were
being exploited by an alpha banded together to depose him. The motivations
of these rebels—chief among them being a desire to secure more personal
autonomy—are readily explained by natural selection aimed at increasing in-
clusive fitness. Some non-human primates, such as chimps, can express their
resentment toward alphas by forming coalitions of a few individuals to attack
him, or ganging up on him as a larger group (de Waal 1982/2007; Goodall
1986). Unlike chimps, our ancestors evolved the necessary intellectual and
psychological capacities to do something permanent about alpha tyranny.
When coalitions of the majority of men overthrew their alphas, each man
realized that he was unlikely to become an alpha himself, and even if he did
he would likely meet the same fate as the previous one. It served the perceived
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14 NATHAN COFNAS
interests of each individual in the coalition to rule as a society of equals—to
agree that no one would be subordinate to anyone. In this way, they were using
their cognitive abilities not to acquire knowledge of mind-independent moral
truth, but to satisfy their naturally selected primate desires to avoid domination.
The reigning coalitions also imposed rules demanding that certain foods
(particularly meat) should be shared more or less equally. This served as both
an insurance policy and a means to prevent individual hoarding that could
lead to significant social inequalities and alpha-like behaviour. Everyone knew
that he himself would sometimes come up empty in a hunting or gathering
expedition. In the long run everyone would benefit from a system that ensured
that the unlucky would not be left to starve. Our ancestors who instituted
food-sharing rules were acting on impulses—primarily a desire to eat—that
evolved by natural selection to promote individual fitness.
Other rules included prohibitions against thievery, lying, and cheating. Once
again, the rule designers were acting on naturally selected desires. Boehm
(2012:52) describes nomadic foragers as ‘intuitive applied sociologists who are
purposefully trying to shape their society in ways that will help themselves
because everyone’s life is helped by better cooperation’. In this context, help-
ing themselves meant promoting outcomes that they were naturally selected to
regard as favourable. Practices like thievery, lying, and cheating undermine
the kind of cooperation that is necessary for obtaining food, for fending off
physical threats, and so on.
According to the debunking account I am proposing, our ancestors who
presided over the social-selection process were motivated by biological drives
that were in turn the product of natural selection. These drives were designed
by natural selection to track the same thing that natural selection tracks,
namely, fitness. It would be an implausible coincidence, if naturally selected
drives (which, again, track fitness) led us to impose rules that just so happened
to correspond with mind-independent moral truth.
Consider the debunking argument in outline form, which can be contrasted
with the realist-social-selection account in the previous section:
1. Natural selection endowed our ancestors with drives to perform (or avoid)
certain actions and to bring about (or prevent) certain states of affairs that
tended to promote (or harm) fitness in the ancestral environment. Among
these were a drive to eat food, to avoid domination, and so on. We also
evolved a general reasoning ability for solving adaptive problems.
2. Because our ancestors evolved (via natural selection) to regard autonomy,
eating, and survival as desirable, around 250,000 thousand years ago they
used their general reasoning ability to design and institute rules to maintain
political equality, ensure access to food, and promote cooperation.
3. The enforcement of rules designed to promote autonomy and collective
well-being created selection pressures for a disposition to internalize group
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GENE–CULTURE COEVOLUTION AND MORAL TRUTH 15
norms and perceive them as true in a realist sense. We also evolved specific
moral intuitions—for example, concerning equality—that make us partic-
ularly receptive to norms that resemble those that were enforced among
nomadic foragers.
4. Under normal circumstances, our socially selected conscience and moral
intuitions are some of the main forces that shape our moral beliefs (real-
istically construed). We easily acquire the belief that, for example, people
are moral equals because we were socially selected to identify with norms
designed to ensure political equality.
5. Natural selection tracks fitness, not mind-independent moral truth. It en-
dows us with drives to perform (or avoid) certain actions and to bring about
(or prevent) certain states of affairs that tend to promote (or harm) fitness.
In other words, like natural selection itself, naturally selected drives track fitness,
not moral truth. The process that drove the evolution of our moral faculty
(i.e., social selectors acting on naturally selected drives) was thus off track
with respect to mind-independent moral truth.
6. Joyce-style evolutionary debunkers are right that we evolved a disposition
to acquire realist moral beliefs. Street-style debunkers are right that evolved
evaluative tendencies are an ‘enormous factor in shaping the content of
human values’ (Street 2006:114). Both are wrong that natural selection
operated directly on our moral faculty.
A similar strategy of pushing EDAs back a step is employed by Kahane
(2014), and his EDA can supplement the one proposed here. He is responding
to de Lazari-Radek and Singer’s (2012) defence of utilitarianism. De Lazari-
Radek and Singer argue that belief in the principle of universal benevolence—a
core component of utilitarianism—is uniquely immune to evolutionary de-
bunking. They acknowledge, however, that the principle of universal benev-
olence requires a theory of well-being in order to tell us what benevolence
consists in—without such a theory, the principle is ‘empty of content’ (ibid.:
23). Kahane (2014) counters that our ideas about well-being appear to be
strongly influenced by natural selection. If utilitarians accept the epistemic
premise that natural selection does not track mind-independent moral truth,
they have to acknowledge that our beliefs about well-being are unjustified,
which (for all practical purposes) renders an ethical system based on the prin-
ciple of universal benevolence contentless. In a parallel way, the theory of
realist social selection assumes that nomadic foragers were acting on an ob-
jectively correct theory of well-being when they established rules to promote
their collective interests. But their views about well-being—namely, that it
consists in individual autonomy, protection from attack, freedom from being
hungry, and so on—are best explained as the result of natural selection acting
to increase inclusive fitness.
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16 NATHAN COFNAS
IV.1 Cultural change
On the social-selection model, moral change is driven largely by changes in the
balance of power among competing interest groups in society. Among nomadic
foragers, coalitions of the majority of males gained the upper hand vis-`
a-vis
would-be alphas, and they enforced egalitarian and socialist values that served
their perceived interests. Ten thousand years ago, when some groups made
the transition to sedentary living and agriculture, populations became too
large to function effectively without a political hierarchy (Boehm 1999:143–4;
Turchin and Gavrilets 2009). Hierarchy re-emerged, and the balance of power
switched back to alphas. Kings and warlords promoted values such as respect
for authority, which served their interests often at the expense of the majority.
The largest, most militaristic agricultural societies were favoured in a process of
cultural group selection (Turchin 2009,2010), which led antiegalitarian values
to spread further.
Huemer (2016) argues that the recent cross-cultural trend toward liberalism
is best explained as a consequence of people independently coming to recog-
nize that liberalism is the objectively correct moral theory. The social-selection
model provides a debunking explanation for the phenomenon of liberal con-
vergence (see Cofnas 2020b; cf. Hopster 2020). People tend to fight for moral
values that serve their perceived interests, which, in turn, are explained by natural
selection. Liberal values—the ‘moral equality of persons’, ‘respect for the dig-
nity of the individual’, and opposition to ‘gratuitous coercion and violence’
(Huemer 2016:1987)—serve the perceived interests of most people. Over the
past few centuries—and especially the last several decades—coalitions of the
majority have simply been reasserting their interests vis-`
a-vis modern-day al-
phas. The return of egalitarianism is probably facilitated by the fact that our
innate moral intuitions evolved (via social selection) to make us particularly
receptive to the egalitarian values that prevailed among nomadic foragers.
Street (2006:113–4) suggests that the content of our moral beliefs has been
shaped in part by ‘many...forces [that] were not evolutionary at all, but
rather social, cultural, historical’, and she highlights the ‘sui generis influence of
rational reflection’. I would point out that social, cultural, and historical forces
sometimes are Darwinian processes (as discussed in Section II), but often they
are not. Non-Darwinian historical accidents, whims of powerful individuals,
changes in the physical environment, and the like can also influence culturally
transmitted values. Forces like these, however, are not likely to be regarded
by the realist as truth tracking. For the realist, rational reflection does seem like
a potentially on-track process that could counteract the distorting (direct or
indirect) influence of natural selection. But I think Street is right that rational
reflection does not provide a way to step outside our evaluations and assess
them from a neutral point of view. Rather, rational reflection elaborates on a
‘starting fund’ of evaluative judgments, drawing out the implications of initial
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GENE–CULTURE COEVOLUTION AND MORAL TRUTH 17
judgments and considering some in the light of others (ibid.: 124; cf. Kahane
2011:119). I suggest that the evidence supports the idea that, insofar as reason
played a role in the evolution of our moral faculty, it has been employed in the
service naturally selected drives.
IV.2 Why favour the debunking account?
FitzPatrick (2015:893) argues that (standard) EDAs are ‘question-begging’. He
says that debunkers cannot refute realism ‘simply by proposing a story that,
if true, would cause problems for realism, and then claim that simply because
of greater parsimony we should accept it as true’. A version of FitzPatrick’s
objection could be applied to the EDA proposed here. We have two mutu-
ally exclusive theories that fit the facts, viz., the realist-social-selection account
and the debunking account. Both theories say that, having evolved the nec-
essary intellectual and psychological capacities, early humans overthrew their
alpha tyrants, established egalitarian political systems to preserve individual
autonomy, and instituted rules to promote collective well-being. Why should we
prefer the debunking explanation? Why assume that our ancestors were acting
on their naturally selected desires rather than knowledge of mind-independent
morality?
I contend that FitzPatrick’s objection does not have the same force against
the social-selection-based EDA that it has against standard EDAs. FitzPatrick’s
original point was that the evolutionary explanations upon which standard
EDAs are predicated are highly speculative. Only prior metaethical commit-
ment to antirealism would justify favouring the speculative debunking theory
over equally speculative realist accounts of the aetiology of our moral beliefs.
The situation is different when we choose between the realist-social-selection
account and the debunking-social-selection account. Realists who reject stan-
dard EDAs like Street’s can deny the entire evolutionary story upon which they
are based. In contrast, advocates of the realist-social-selection theory ipso facto
agree with debunkers that our moral faculty evolved due to social selection.
They differ only in what motivations they attribute to the social selectors:
realists say the social selectors were acting in response to their knowledge of
moral truth—debunkers, that they were acting on naturally selected desires
shared with other primates. It seems difficult for the advocate of the realist-
social-selection account to avoid positing a big coincidence. Social selection
appears to have unfolded as if it had been guided by naturally selected desires
shared with other primates (the desire to avoid being dominated, etc.). The
debunking account offers a straightforward explanation for this, namely, it re-
ally was guided by those desires. But for advocates of the realist-social-selection
account, this would be a huge coincidence. They must posit not only extra
ontology (mind-independent moral truth) but also an inexplicable coincidence.
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18 NATHAN COFNAS
There is another option available to the realist. As noted in the Introduction,
some critics of standard EDAs argue that moral beliefs can be both adaptive
and objectively correct. Therefore, natural selection could have favoured a
tendency to acquire true moral beliefs. Essentially the same argument can be
employed against a social-selection-based debunking argument. The realist
can say that natural selection gave us drives that led us to support norms that
are also objectively morally correct. For example, natural selection gave us a
love of autonomy, which made us establish an egalitarian political system, and
this made us internalize (via social selection) the moral truth that people are
equals. But, as Vavova (2015:109) notes, arguments of this type ‘presuppose
exactly what is in question: the truth of our moral beliefs. In the context of
the debunker’s attack, this is question-begging.’ The debunker who invokes
the social-selection account of morality purports to give a reason to think that
our moral beliefs are unjustified. One cannot respond by assuming the beliefs
are correct.3
V. DISCUSSION
I have considered what follows metaethically from the premise that our moral
faculty is the product of social selection as described by Boehm (1999,2012).
There are several influential competing accounts of moral evolution—some of
which have been discussed in this paper—and scientists are far from reaching
a consensus on which is best (Bloom 2019). Arguably, however, evidence is
accumulating in support of the social-selection account. Although they do
not discuss the evolution of morality, Levine et al. (2018) provide evidence
that moral reasoning (at least in some contexts) follows a contractualist logic,
which fits with the predictions of the theory of social selection. Curry et
al. (2019) argue persuasively that categories of behaviour that are relevant
to cooperation are universally moralized, which is what we would expect if
morality was designed to advance collective group interests. Wrangham (2019:
204) notes that ‘[u]ntutored children’ exhibit prosociality and sensitivity to
norm violations ‘that are not explained by kin selection or mutualism’, but
which can readily be explained by the social-selection model.
If the theory of social selection is correct, metaethics potentially has some-
thing substantial to contribute to the science of the evolution of morality. To
explain the evolution of our moral psychology, we must give an account of
the motivations of our ancestors who established and enforced social norms.
3Vav o v a ( 2015:112–3) argues that evolutionary explanations of our moral beliefs may not
actually give us a reason to think we are mistaken, but I leave this controversy aside. As noted,
in this paper I assume for the sake of argument that attributing moral beliefs to natural selection
debunks them.
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GENE–CULTURE COEVOLUTION AND MORAL TRUTH 19
One possibility is that they were motivated by their appreciation of mind-
independent moral truth acquired via reason—a process I termed realist social
selection. The alternative is that they were acting on naturally selected motiva-
tions shared with other primates—desires to avoid being dominated, to secure
food, and so on. Although I have argued in favour of a debunking account,
the theory of realist social selection must be acknowledged as a competing
hypothesis. Scientists investigating the biological evolution of morality cannot
remain metaethically neutral.4
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