Fear of fitness indicators:
How to deal with our ideological anxieties
about the role of sexual selection
in the origins of human culture
Geoffrey F. Miller
Assistant Professor
Psychology, Logan Hall 160
University of New Mexico
Albuquerque, NM 87131-1161, USA
(505) 277-1967 (office voice/fax)
(505) 277-1394 (dept fax)
gfmiller@unm.edu
http://www.unm.edu/~psych/faculty/gmiller.html
Published as:
Miller, G. F. (2003). Fear of fitness indicators: How to deal with our ideological anxieties
about the role of sexual selection in the origins of human culture. In Being human:
Proceedings of a conference sponsored by the Royal Society of New Zealand, pp.
65-79. Wellington, NZ: Royal Society of New Zealand, Miscellaneous series 63.
Running Head: Fear of fitness indicators
Fear of fitness indicators
Geoffrey F. Miller
Introduction: Mental evolution through mate choice
The human mind evolved somehow. Yet it remains unclear what evolutionary
selection pressures favored our large brains, our creative intelligence, and our unique
capacities for language, art, music, humor, romantic love, and moral commitment. Following
Herbert Spencer's misleading phrase 'the survival of the fittest', most theorists throughout
the last 140 years have tried and failed to identify the 'survival value' of these capacities.
How did they contribute to life on the Pleistocene savanna of Africa, in which our hominid
ancestors evolved?
Most evolutionary thinking about human mental evolution has emphasized the
“production” payoffs for creative intelligence: our ancestors could invent better tools, which
allowed more effective hunting, gathering, and defense against predators and rival groups,
thereby improving survival prospects. This production orientation fit well with 20th century
social science’s post-Marxist emphasis on economics and technological progress as
determinants of human history. It was easy to project such an orientation back into human
prehistory.
However, in recent years, an increasing number of evolutionary thinkers have begun
emphasizing sexual reproduction rather than economic production as the driving force
behind the evolution of animal and human behavior. This innovation actually dates back to
Darwin’s (1871) book The descent of man, and selection in relation to sex. Darwin argued
that sexual selection (for reproduction) is distinct from natural selection (for survival), and
that traits inexplicable in terms of 'survival value' can often be explained as ornaments for
attracting sexual partners. Over time, discriminative mate choice results in the elaboration of
any traits that are noticed and appreciated by the opposite sex, such as body size and
symmetry, beautiful plumage, healthy skin and fur, energetic gait, creative song-production,
or social dominance. Thus, mate choice can explain the evolution of the peacock’s tail, the
nightingale’s song, the elk’s antlers, and, Darwin argued, human intelligence, music, and
morality. Unfortunately, Darwin’s Victorian peers rejected his evidence for mate choice, and
could not accept that animals possessed the capacities of perception, judgment, and
discrimination necessary for mate choice to play a central role in evolution (Cronin, 1991).
As a result, almost all of 20th century psychology, anthropology, neuroscience, and the
humanities developed without recognizing any significant role for mate choice in human
mental evolution.
The dramatic revival of sexual selection theory since about 1980 (Andersson, 1994)
has confirmed the utility of Darwin's distinction between natural and sexual selection. Sexual
selection often produces extravagant traits that have high costs and complexity, yet these
traits are often unique to one species, and absent in closely-related species. For example,
closely-related birds often display quite different plumage colors and patterns, and sing very
different courtship songs. By contrast, natural selection for ecological utility tends to produce
convergent evolution, where many lineages independently evolve the same, efficient, lowcost solutions to the same environmental problems – traits such as wings, eyes, teeth,
claws, hearts, and lungs. Sexual selection theory’s revival has been so dramatic that
scientific journals concerning the evolution of animal and human behavior are now
dominated by research on mate choice and its effects. Yet this revival has gone largely
unnoticed in mainstream psychology, neuroscience, and the social sciences, which still view
'survival of the fittest' as evolution's bottom line, and which therefore have trouble seeing any
evolutionary rationale for those aspects of human nature most concerned with selfornamentation, display, status, ideology, fashion, and aesthetics.
Darwin’s view of the human mind as a set of courtship adaptations can be extended
and refined in the light of modern sexual selection theory, evolutionary psychology, and
evolutionary anthropology. In my recent book The mating mind (Miller, 2000a), I argued that
many of our most distinctive mental traits evolved through mutual mate choice, in both
2
Fear of fitness indicators
Geoffrey F. Miller
sexes, to advertise intelligence, creativity, moral character, and heritable fitness. Indeed, the
human brain itself can be viewed as a sexual ornament, since it has most of the hallmarks of
sexually-selected traits: it is unique among living primates, it has high metabolic costs and
enormous complexity, and its capacities are conspicuously displayed during courtship
(especially verbal courtship).
The fact that male and female brains are fairly similar is congruent with evidence that
human mate choice is mutual, with males and females almost equally choosy about the
mental traits of long- term sexual partners. Sex differences are a distinctive outcome of
sexual selection, but sexual selection does not always produce sex differences. So far,
much of evolutionary psychology has focused on sex differences in courtship behavior and
mate choice (e.g. Buss, 1999), but sexual selection theory is equally applicable to
conspicuous, costly, ornamental traits in humans that are displayed by both sexes when they
are reproductively mature.
Human mental traits as fitness indicators
If our most distinctive mental capacities evolved through mate choice, then to
understand human nature, we have to understand how mate choice shapes traits as
courtship adaptations. In the last 15 years, there has been a scientific revolution in our
understanding of this issue, and the revolution depends on a branch of evolutionary theory
called “biological signaling theory”. Signaling theory reaches all the way from the deepest
fundamentals of evolution – mutation, genetic variation, and selection – right up to the most
complex manifestations of beauty in nature and human culture.
All species face the problem of “mutational meltdown” (Ridley, 2001): cosmic rays
and copying errors are constantly introducing mutations into the genetic code. These
mutations are almost always harmful, because a change in any gene in much more likely to
disrupt the gene’s function than to improve it. Yet, without the rare beneficial mutation,
evolution would never make any progress. Thus, any genetic code (e.g. DNA) capable of
progressive evolution is also vulnerable to progressive degeneration through mutation.
Without a way to “purge” harmful mutations from its genes, every species would eventually
lose its ability to survive and reproduce effectively in its environment – it would suffer a
mutational meltdown. In fact, many species that give up sexual reproduction in favor of nonsexual reproduction suffer precisely this fate: they almost always go extinct within a million
years (Ridley, 2001). This reasoning has led many biologists to the view that sexual
reproduction itself may have evolved as a way to purge harmful mutations (e.g. Atmar, 1991;
Kondrashov, 1988; Michod, 1995; Michod & Hasson, 1990).
How can sex get rid of mutations? Mate choice can discriminate against individuals
whose bodies and behaviors reveal a high “mutation load”. If an animal is showing poor
health, poor coordination, and poor intelligence, this may because it is carrying a higher than
average number of harmful mutations. A choosy animal would do well to avoid mating with
such an unfit individual, lest its offspring inherit a similarly high number of mutations.
Therefore, traits that happen to reveal mutation load more accurately than other traits (e.g.
through more complex development that requires the interaction of more genes) will tend to
be favored in mate choice. Under sexual selection, such traits will tend to grow ever larger,
costlier, and more complex, so they function as ever more reliable indicators of good genes.
Insofar as genetic quality implies expected evolutionary fitness, these sexually-selected traits
could be called 'fitness indicators' (Andersson, 1994; Grafen, 1990; Johnstone, 1995;
Michod & Hasson, 1990; Rowe & Houle, 1996; Zahavi & Zahavi, 1997).
Examples of fitness indicators are ubiquitous, because almost every sexually
reproducing species has one or more, in at least one sex. The classic Darwinian examples
of fitness indicators are extravagant ornaments such as the peacock’s tail (Darwin, 1871;
Cronin, 1991). Recent research has shown that the quality of many such ornaments actually
conveys reliable information about an individual’s general health, fitness, and genetic quality.
This fitness-indicating function appears to hold true not only for peacock tails (Manning &
3
Fear of fitness indicators
Geoffrey F. Miller
Hartley, 1991; Moller & Petrie, 2002; Petrie, 1994), but for a very wide range of otherwise
puzzling traits in many species, including:
 the length of eye-stalks in stalk-eyed flies (Hingle, Fowler, & Pomiankowski, 2001;
Wilkinson, Presgraves, & Crymes, 1998),
 the amount of orange-red caretenoid pigment in guppy tails (Gong & Gibson, 1996;
Kodric-Brown, 1985; Nicoletto, 1991),
 the symmetry of tail streamers in barn swallows (Moller, 1994),
 the rate of abdomen-drumming by wolf spiders (Kotiaho, 2000),
 the energetic intensity of copulation in a species of highly promiscuous beetle
(Psilothrix viridicoeruleus) (Shuker et al., 2002),
 the size of the penis bone in bats (Hosken et al., 2001)
 the length and complexity of bird song in aquatic warblers (Catchpole & Leisler,
1996),
 the length and loudness of mating songs by gibbons (Colinshaw, 1996; Geissman,
2000), and
 the precision of song repetition in humpback whales (Chu & Harcourt, 1986; Miller,
Biassoni, & Samuels, 2000; Noad, Cato, & Bryden, 2000; Payne, 2000).
Human bodies are full of reliable fitness indicators. Male human beards, penises,
and upper-body muscles, and female breasts, buttocks, and orgasmic capacities, show
evidence of having evolved through sexual selection (Dixson et al., 2003; Dixson, 1998;
Etcoff, 1999; Miller, 2000a). There are also some traits that do not show large sex
differences, such as long head hair, hairless skin, highly expressive faces, and full lips, that
function as sexual attractants in our species (Langlois et al., 2000). These fleshy fitness
indicators don't fossilize, and so have been neglected in studies of human evolution.
However, all these traits are valued in modern mate choice, are displayed during courtship,
and are mostly unique to humans. Also, the sexually differentiated traits (male beards,
penises, and muscles, and female breasts and buttocks) develop fully only during puberty, in
time for sexual attraction.
Human brains make particularly good fitness indicators because their growth
depends on about half the genes in the genome, thereby summarizing a huge amount of
information about mutation load (Miller, 2000a,b,c; Miller & Todd, 1998; also see Madden,
2001). Brains are also good indicators of nutritional state and general health, because they
have such high energetic costs, constituting only 2% of body weight, but consuming over
25% of adult metabolic energy (60% in infancy). Indeed, the brain requires so much glucose
that experimentally induced hypoglycemia can dramatically lower cognitive ability
(McCrimmon, Deary, & Frier, 1997; Sommerfield, Deary, & McAulay, 2003). Moreover, the
more important brains became during primate evolution (for whatever reason), the more
incentive mate choice would have had to focus on specific indicators of brain quality (i.e.
mental fitness as distinct from physical fitness).
Recent genetic analyses suggest that at least 1.6 harmful new mutations per
individual per generation have been arising in our lineage for the last several million years
(Eyre-Walker & Keightley, 1999; Crow, 1999). This probably exceeds the mutation rate that
natural selection could contain in the absence of sexual selection for genetic quality.
Without mate choice for fitness indicators, we would have suffered a mutational meltdown
long ago.
What is the evidence that good brains and good minds are really sexually attractive
in our species? Isn’t the putative attractiveness of creative intelligence just a conceit of the
educated classes? Cross-cultural research on human mate choice suggests not. For
example, in David Buss’s (1989) study of mate choice in 37 cultures across 5 continents
around the world, five out of the top seven traits most desired by both men and women were
psychological rather than physical traits. These psychological traits included kindness (rated
#1 by both sexes), intelligence (rated #2 by both sexes), exciting personality, adaptability,
4
Fear of fitness indicators
Geoffrey F. Miller
and creativity. The only physical traits to make the top seven were physical health and
physical attractiveness. Purely sociological or economic traits (such as social class or
wealth) did not feature prominently in self-reported mate preferences in either sex of any
culture. There is abundant evidence now for the importance in human mate choice of
creative intelligence (e.g. Kanezawa, 2000; Li et al., 2002; Lynn et al., 2002; Miller, 1999a,
2000c; Reynolds & Gifford, 2001; Zebrowitz et al., 2002; Zechner et al., 2001) and kindness
(e.g. Dessalles, 1998; Goldberg, 1995; Kelly & Dunbar, 2001; Johnson, 1996;
From fitness indicators to human culture
What do fitness indicators have to do with human culture? The biological signaling
theory that explains naturally evolved fitness indicators is remarkably similar to the theory of
conspicuous consumption developed by Thorstein Veblen (1899) to explain extravagant
culturally invented luxuries (Miller, 1999). Highly fit animals show off their genetic quality by
growing costly, precisely formed ornaments that their lower-fitness rivals cannot afford or
imitate. Likewise, wealthy humans show off their pecuniary prominence by buying costly,
precisely formed luxuries that their poorer rivals cannot afford or imitate. Indeed, the 20th
century saw a parallel but largely independent development of signaling theory in biology (to
explain costly sexual ornaments – e.g. Johnstone, 1995; Zahavi & Zahavi, 1997), in
economics (to explain costly, conspicuous advertising and marketing efforts by corporations
– e.g. Frank, 1999; Hellofs & Jacobson, 1999; Kirmani & Rao, 2000), and in sociology (to
explain costly, conspicuous badges of social status – e.g. Veblen, 1899; Packard, 1960;
Bourdieu, 1987; Illouz, 1997; Neiman, 1997; Gagnier, 2001).
How deep do the parallels go between these three strands of research – biological
signaling theory, economic signaling theory, and sociological signaling theory? A skeptic
might accept that the basic logic of costly advertisement may be the same, but argue that
the functions of human economic and social-class signaling must be quite different from the
fitness-indicator function of sexual ornaments in other animals. That is, we may display our
wealth and status not to attract sexual partners, but to do something else. This view
maintains the comfortable division between the animal world of genes, traits, and sex, and
the human world of culture, ideology, and money.
However, I think the reasonable default view is that our wealth-displays and statusdisplays serve much the same mate-attracting and mate-retaining functions as the fitnessdisplays of other species. One way to see the potential overlap of functions is to consider in
more detail the typical features that are shared by biological fitness indicators and human
cultural displays such as art, music, and story-telling.
Conspicuous cost and conspicuous precision in fitness indicators
Fitness indicators often look rather different from standard naturally selected
adaptations that solve standard survival problems. The evolved function of indicators is
basically to advertise an individual’s fitness to other individuals. The only way to do this
reliably is to grow a trait, or to emit a behavior, that a lower-fitness rival could not produce in
such a high-quality form. Thus, fitness can be advertised according to several overlapping
principles:
 Do something that a lower-fitness rival could not afford to do – in terms of time,
energy, resources, or risk.
 Do something that would make your mutations, health problems, and
psychopathologies very conspicuous
 Do something that requires special virtuosity to overcome some difficulty
 Do something that is very hard to do well, and very easy to do badly
Any fitness indicator that was easy to “fake” – such that a low-fitness individual could
produce a high-quality version of the trait – would no longer be attended to by other animals,
because it would no longer convey reliable information (Grafen, 1990; Zahavi & Zahavi,
1997). Thus, fitness indicators must be hard to fake in order to be reliable, and in order to
remain evolutionarily relevant. The relevant evolutionary slogan is not “survival of the fittest”,
5
Fear of fitness indicators
Geoffrey F. Miller
but “truth in advertising”. There seem to be two major features that can make fitness
indicators harder to fake: conspicuous cost and conspicuous precision.
Conspicuous cost. Low-fitness individuals cannot afford high-cost indicators,
because they have a smaller budget of available time, energy, or resources, and must
devote a higher proportion of their budget to basic survival needs (Johnstone, 1995; Zahavi
& Zahavi, 1997). They cannot afford the luxury of high-quality sexual ornamentation. Often,
sexual ornamentation requires high growth costs (e.g. energy and resources to produce a
peacock’s tail, an elk’s antlers, or a human’s brain), high maintenance costs (e.g. preening,
grooming, practice), and high repair costs (e.g. annual regrowth of moulted plumage,
rewiring of brain areas injured by violence or stroke).
Conspicuous precision. Low-fitness indicators cannot manage to grow their bodies
or brains with sufficient precision that they can display high-quality ornaments. Imprecise
growth results in asymmetric, irregular, and unevenly colored ornaments. For example,
animals with higher mutation loads tend to grow less symmetric bodies and ornaments,
because the mutations interfere with the precision of right versus left side body growth
(Gangestad & Thornhill, 1997; Moller & Swaddle, 1998; Thornhill, 1998). Also, humans with
higher mutation loads tend to grow brains that show various anatomical and functional
abnormalities, resulting in lower intelligence and higher rates of psychopathology (Yeo,
1999; Yeo et al., 1999).
The most common forms of conspicuous precision in sexual ornamentation seem to
be: symmetry, averageness, the precise repetition of complex motifs, and the precise
imitation of external phenomena. Human facial beauty for example involves the first two:
faces are rated as most beautiful when they are bilaterally symmetrical and when they are
close to the statistical average form for a human face (Etcoff, 1999; Langlois et al., 2000).
By contrast, much of human culture involves the precise repetition of complex motifs (e.g.
ornamental motifs in visual art, rhythm in music, movements in dance – see Gombrich, 1979;
Zahavi, 1978) and the precise imitation of external phenomena (e.g. representational
accuracy in art, accurate depiction of human nature in songs and novels, concordance
criteria of truth in mental models of the world and scientific theories – see Gombrich, 1977;
Miller, 2000a, 2001). Yet, repetition and mimesis are not uniquely human principles of
fitness-display: humpback whale show off their fitness through the precise repetition of 30minute whale songs to attract mates, and nightingales show off their fitness through the
precise imitation of sounds made by other species.
From this fitness-indicator viewpoint, virtuosity is much more important than novelty
in human cultural displays. Virtuosity is universally admired in every craft from every culture
(Boas, 1955; Gombrich, 1977, 1979; Turner, 1992). In fact, the term “art” is used as an
honorific term to identify almost any product of human behavior that is created with skill and
attention that goes beyond the pragmatically necessary, that is “made special” in some way
(Dissanayake, 1992) – i.e. that follows the logic of signaling theory. Whether in the
geometric art of M. C. Esher, the complex wood carvings of Maori peoples, or the prehistoric
cave paintings of Lascaux and Altamira, the ability to repeat complex motifs with precision,
consistency, and skill is taken as a sign of expertise, and is considered “beautiful”. Indeed,
Kohn and Mithen (1999) have even proposed that the evolution of tool-making ability in
human evolution was driven in part by sexual selection favoring capacities to create large,
symmetric, beautiful hand-axes.
The ideological problems with the fitness indicator theory of human evolution
The argument thus far has been, roughly:
 Some of our most distinctive mental traits (e.g. art, music, language, creativity, ideology)
are hard to explain as survival adaptations
 Darwin’s process of sexual selection through mate choice can explain many traits in
many species that are hard to explain as survival adaptations
 Sexual selection often produces “fitness indicators” – traits that advertise an individual’s
genetic quality through conspicuous cost and precision
6
Fear of fitness indicators
Geoffrey F. Miller

Fitness indicators follow the “truth in advertising” logic of biological signaling theory,
which is similar to theories of wealth-signaling and product-quality-signaling in
economics, and theories of social-status-signaling in sociology
 Some human mental traits and cultural forms have already been analyzed as wealthsignals and status-signals, so it is not a great reach to consider them as possible
fitness-signals as well
 Some human mental traits and cultural forms bear other hallmarks of sexual selection,
so they may serve the same biological function (mate-attraction) as fitnesssignals in other species
In response to longer versions of this argument (e.g. reviews of my book The mating
mind, and questions after my public talks), two reactions are often voiced: “So what?” and
“Oh no!”. The “So what?” reaction I deal with at much greater length in the book itself. My
focus for the rest of the paper here is on the “Oh no!” reaction. Often, this reaction arises
from concerns about the political, religious, and ideological implications of this fitnessindicator view of human mental evolution and of human culture.
Basically, the idea that the human mind evolved as a bundle of fitness indicators
does not sit comfortably with contemporary views of human nature and human society. In
fact, it violates at least six core values commonly accepted in modern society:
1. Fitness variation vs. human equality. Fitness indicators are designed by evolution
to reveal individual differences in general intelligence, health, fitness, and genetic quality.
They maximize apparent differences between people both objectively (by amplifying small
differences in genetic quality into big differences in apparent physical and psychological
attractiveness), and subjectively (by shaping our social-perception instincts to pay a great
deal of attention to fitness indicators). This leads us to a virtual obsession with the minutiae
of individual differences in our social interactions – not only in mate choice, but in our
choices of friends, allies, co-workers, mentors, and leaders. Fitness indicators make all
sorts of differences loom very large indeed, including psychological differences in
intelligence, kindness, agreeableness, aggressiveness, conscientiousness, conservatism,
neuroticism, psychoticism, extroversion, and ideology. Insofar as different ages, sexes,
ethnic groups, and races may have somewhat different averages and degrees of variation in
these traits, our obsession with differences is easily generalized from individuals to such
groups, leading to various forms of ageism, sexism, ethnocentrism, and racism. Necessarily,
this difference-obsession also tends to eclipse our awareness of what we humans have in
common – and it is precisely our common human nature that is the basis for the classical
liberal enlightenment concepts such as “human rights”, “equality before the law”, “universal
suffrage”, and “democracy”. Thus, a descriptive emphasis on the role of fitness indicators in
human evolution appears to sit very uncomfortably with a normative respect for human
equality.
2. Judging people by their indicators vs. by their character. A related problem is that
the components of genetic quality that are advertised by fitness indicators can seem quite
impersonal. If intelligence, creativity, sense of humor, and even kindness are simply
reflections of how many mutation-free genes we carry, then human dignity may seem
reducible to DNA. Equally, when we think we are judging people by unique, deeply
personal qualities, fitness indicator theory suggests we are really paying attention to speciestypical indicators that can be arrayed along dimensions of superiority vs. inferiority. This
viewpoint seems to leave little room for developing a personal identity apart from one’s
heritable indicators, or for being judged by others as anything other than an advertisement
for one’s genes.
3. Mate choice for good genes vs. romantic love. This is really a special case of the
above point, but it is an especially important, emotionally charged special case. The fitness
indicator view suggests that sexual attraction is largely a matter of integrating physical and
psychological cues of genetic quality, and finding the highest-fitness partner who will actually
reciprocate one’s affection. By identifying the major fitness indicators, the ways in which we
weigh and integrate them, and the ways in which we search for mates from the pool of
7
Fear of fitness indicators
Geoffrey F. Miller
possibilities (e.g. Miller & Todd, 1998; Todd & Miller, 1999), evolutionary psychology works
to demystify human love and romance. Many of us do not want love demystified, because
the romantic commitments entailed by true love do not seem capable of surviving such
demystification. In short, we like to pretend that there was only one predestined “true love”
out there for us, and that we made no comparative judgments concerning the relative fitness
of our true love and other potential mates. Indeed, putting one’s marriage into the arena of
comparative judgment risks undermining the emotional commitments that keep one from
wandering into infidelity and divorce. Thus, fitness indicator theory seems hostile to the
mythology of romantic love, and to the emotional demands of marriage.
4. Heritable fitness vs. parental and cultural influences on human development. A
major discovery in sexual selection research has been that many aspects of general fitness
are heritable. Indeed, the most common evolutionary rationale for mate choice may be that
it allows individual to get better genes for their offspring. Moreover, the better a fitness
indicator works as a cue of general genetic quality, the more heritable will be individual
differences in the quality of that indicator. This sounds fine in the abstract, but it becomes
problematic when we start viewing as fitness indicators many human mental traits – such as
general intelligence, language ability, sense of humor, artistic ability, musical ability,
kindness, and altruism. If intelligence is a good fitness indicator, then intelligence is
probably highly heritable – and all the behavior genetics research suggests that it is (e.g.
Plomin et al., 2001). Before the rise of fitness indicator theory, it was possible to argue that
heritable traits must not be that important, because selection would have long ago eliminated
any heritable variation in traits that really mattered in survival or reproduction (see Miller,
2000b,c). However, fitness indicator theory points out that some sexually-selected traits can
remain both extremely important in social and sexual life, and very highly heritable (Miller,
2000c; Rowe & Houle, 1996). Thus, heritability can no longer be taken as a symptom of
evolutionary irrelevance. This leaves the “nature vs. nurture” debate in a very unbalanced
state, with behavior genetics research showing “nature” (one’s genotype) looking ever more
powerful and socially important, leaving “nurture” (one’s shared family environment in
childhood) looking impotent. That is, the quality of parenting and early family life simply
doesn’t influence how an individual will turn out as an adult, except in cases of extreme
abuse or neglect. Thus, a descriptive emphasis on the role of fitness indicators in human
evolution seems to fit very poorly with a normative respect for the importance of parenting
and culture in individual human development. We want parenting to be important to justify
our efforts and attentiveness as parents, but fitness indicator theory combined with behavior
genetics findings shows that this desire for parental relevance is largely futile.
5. Narcissistic self-display vs. civilized humility. Loudly advertising one’s fitness
violates our values of humility, decorum, and tact. Fitness indicator theory views self-display
as central to human evolution, so could be construed as providing an excuse for all manner
of narcissistic showing-off – bragging, strutting, consuming conspicuous luxuries, and
derogating one’s less extroverted rivals. Yet many cultural norms have developed precisely
to restrain runaway showing-off by rivals – especially young, single, male rivals – within
society. The law itself can be viewed largely as an attempt by older, mated men to minimize
the anti-social effects of runaway status competition by young, single men. These anti-social
effects range from noise pollution (e.g. playing rock or rap music at high volume from one’s
car) to lethal violence (e.g. the tendency of accidental slights to escalate into mortal combat
among aggressive young men). If fitness indicator theory provides a clear, coherent
rationale for such showing-off (i.e. it has high reproductive payoffs), it becomes harder for
older do-gooders to argue that all such showing-off is irrational and contrary to the true
interests of the perpetrators. Indeed, fitness indicator theory appears to give the perpetrators
an ironclad excuse for all manner of loud, aggressive, self-display – fitness signaling as a
basic human right, perhaps. Thus, highlighting the evolutionary payoffs to fitness signaling
seems to conflict with our desire to justify social norms against narcissistic self-display and
showing-off.
8
Fear of fitness indicators
Geoffrey F. Miller
6. Fitness signaling vs. transcendental cultural meaning. Perhaps the hardest thing
to accept about fitness indicator theory is what it does to the content of human cultural
creations, such as works of art, music, literature, and religious inspiration. It suggests that
the content of such works is a peripheral and somewhat incidental feature of the work’s
function as a fitness indicator. For indicators, form is closely tied to function: indicators are
most informative when they display easy-to-assess “surface” qualities such as high cost,
symmetry, regularity of design, and accuracy of mimesis. To the sexual selection theorist,
the visual themes of a painting may be less important than the monetary cost of the
pigments used, the amount of time and energy invested by the painter, and the correlation
between the painter’s manifest skill level and his or her mutation load. Fitness indicator
theory puts these “superficial” features at the heart of human cultural effort, and relegates
thematic content to the periphery, as just another form of cognitive ornamentation. This
viewpoint threatens the Romantic-era ideologies of transcendence, genius, inspiration,
salvation, and social utility that remain the major popular justification for the arts, the
humanities, and religion. Thus, fitness indicator theory seems corrosive to cultural content,
symbolism, and social values. It seems nihilistic to propose that our capacities for language,
art, and music evolved to proclaim just one message that has been repeated loudly and
insistently for thousands of generations: “I am fit, my genes are good, mate with me”.
Resolving the ideological problems with fitness indicator theory
How is it possible for one biological concept to affront so many of our fundamental
values? It seems quite astounding that a scientific idea should so consistently fall on the
wrong side of the ideological fence. I think it is no coincidence. Look at it this way: our
human norms and values developed as reactions to patterns of natural human behavior that
we decided should be discouraged. If a great deal of human behavior consists of advertising
one’s fitness, and if many ways of doing that impose social costs on others, and if moral
norms develop to minimize social costs, then a lot of moral norms should be aimed directly
against the irresponsible use of fitness indicators. For example:
 We teach equality to fight arrogance and dominance, and to limit the subjective
distress caused by inequality
 We teach that everyone has a unique identity apart from his or her indicators, to
promote sympathy in our social and sexual relationships
 We teach romantic love to limit the callousness with which we judge potential
mates, to limit mate-switching and infidelity, and to stabilize families
 We teach that nurture is important, to remind parents to care about the subjective
well-being of their children
 We teach humility to limit obnoxious showing-off and fitness-displaying
 We teach that cultural works have deep, socially important meanings, to promote
the production of cultural works that really may have such meanings, rather than
the superficial signs of high fitness.
These social norms do not just fall randomly from the sky. They emerged as moral
instincts and cultural inventions to combat the excesses of sexual self-advertisement and
sexual competition. Our moral aversion to fitness indicators may tempt us to reject them as
an important part of sexual selection. But if we reject them, then it is hard to see how our
moral norms evolved in the first place. It is possible, perhaps even necessary, to admit that
much of human behavior evolved to advertise fitness, while simultaneously realizing that the
essence of wisdom and morality is not to take our fitness indicators too seriously. This is not
to say that our capacities for wisdom and morality are cultural inventions that liberate us from
the imperatives of our genes. Our moral instincts may be just another set of evolved
adaptations. It is not question of “us” over-riding our genetic predispositions, but of using
one set of predispositions to over-rule others – just as our evolved desire to preserve our
looks can over-ride our evolved tastes for fats and sugars.
9
Fear of fitness indicators
Geoffrey F. Miller
A major prediction of fitness indicator theory is that in every culture, and perhaps in
every individual, there should be a dynamic balance between our mating instincts for fitnessdisplay, and our moral instincts for restraining the social and personal costs of runaway
fitness-display and sexual competition. In many societies, certain groups of people tend to
become advocates for one side or the other. Often, it seems to me that the advocacy
groups break down as follows – on average, in most places and times:
Against runaway fitness signaling:
Women
Old folks
Conservatives
Fascists, Communists
Marxism
The bourgeoisie
Monotheists
Obsessive-compulsives, Paranoids
Group-minded do-gooders
Economists
Ecologists
Sociologists
Volkswagen-drivers
Immanuel Kant
Al Gore
Tipper Gore
For runaway fitness signaling:
Men
Young folks
Liberals
Libertarians
Consumerism
The bohemians
Pagans
Narcissists, Psychopaths
Irresponsible individualists
Marketers, Advertisers
Sexual selection theorists
Socialites
Porsches-drivers
Friedrich Nietzsche
Bill Clinton
Britney Spears
If this amateur sociology is even vaguely correct, then we need not worry that the
advocates of fitness indicator theory will cause any permanent injury to the social fabric.
There will remain a dynamic ideological balance between those in favor of runaway fitness
signaling and those opposed to it, as long as we have a demographically balanced society.
Some social benefits of the fitness indicator view
Of course, one is not likely to win over many hearts and minds by pointing out that a
scientific theory is not quite as horribly unsettling as it first appears. One must offer some
positive social benefits of adopting the new view. I believe there are several such benefits to
emphasizing the role of sexual selection in human mental evolution. These benefits seem to
fall into five categories: intellectual, educational, economic, aesthetic, and spiritual.
Intellectual benefits. To scientists, the intellectual, theoretical, and empirical benefits
of a new viewpoint are paramount – indeed, other ideological considerations are considered
irrelevant at best, and corrosive at worst. In most of my other writings, my focus has been
on the purely scientific benefits of fitness indicator theory as a framework for understanding
human nature. In this paper, my focus is on the non-scientific, “ideological” issues, so I will
just mention one major intellectual payoff offered by the fitness indicator viewpoint:
“consilience” (Wilson, 1999), which is the potential to integrate seamlessly the biological and
social sciences with the arts and humanities (Miller, 2000a). Signaling theory offers a
common language and functionalist framework that spans the realms of the biological, the
economic, and the cultural. Of course, the proof is in the pudding, and it remains to be seen
whether this potential for consilience is fulfilled over the coming decades of research.
Educational benefits. Ever since Darwin, most theories of mental evolution have
stressed the economic and technological payoffs for human creative intelligence. This can
easily lead to a sort of nerd’s eye view of public education, in which business management
and computer programming are viewed as the pinnacles of human cognitive achievement,
and in which art, music, sports, social life, and sexual education can be marginalized as
10
Fear of fitness indicators
Geoffrey F. Miller
“extra-curricular activities”. The fitness indicator view inverts this cognitive hierarchy by
pointing out that economic success and technological invention are just two among many
natural ways that individuals advertise their fitness – and they are not even considered the
most sexually attractive or romantic ways by most people. If educational policy recognized
that training individuals to be socially and sexually successful was at least as important to
their subjective well-being and quality of life as training them to be technically proficient at
some employable skill, then fitness indicator theory would offer a strong argument for putting
the arts and humanities at the core of the curriculum. At the moment, the interests of
corporate employers seem to dictate much of educational policy in many industrialized
nation, and those interests typically demand that young humans spend thousands of hours
learning frustrating, counter-intuitive, evolutionarily novel skills such as mathematics, word
processing, and accounting. The corporate world is not much interested in promoting
fitness-indicating skills that young people seems intrinsically motivated to learn and practice,
which are socially and sexually rewarding, and which tap their naturally evolved cognitive
abilities – such as art, music, story-telling, drama, natural history, humor, sports, and
socializing. At the very least, fitness indicator theory can help educational policy-makers
become less hypocritical and self-deceiving about whose interests are being served by
current educational practices.
Economic benefits. Early theories of human evolution coincided with the peak of
industrial capitalism based on heavy industry – the world of coal, oil, steel, railroads,
automobiles, armaments, and factories. In this world, the economy seemed to be based
upon the transformation of raw materials into physical tools, weapons, vehicles, and
buildings. Materialism – whether Marxist or capitalist, dialectical or techno-scientific –
seemed an appropriate way to describe how the world works and how the human brain
evolved to understand that world. However, with the rise of sexual selection theory and
fitness indicator theory, the world appears not so much a set of raw materials and physical
products, as a system of signals, advertisements, status games, and socio-sexual
transactions. This new view seems much better suited to understand the global 21st century
economy as it is – a system that delivers service, entertainment, education, experience, and
information – rather than as it was in the mid-19th century world of heavy industry. Indeed,
without the fitness indicator perspective, it is very difficult to understand modern marketing
phenomena, which are at the heart of modern business practice (Miller, 2000d,e; Saad &
Gill, 2000). All successful contemporary businesses understand that profits arise not from
commodifying their products (i.e. stripping them of brand identity, advertising associations,
and fitness-signaling potential), but from ‘adding psychological value’ to their products by
increasing their fitness-signaling, status-signaling, and identity-signaling potential (Frank,
1999; Klein, 2000; Twitchell, 1999). The fitness indicator view also helps us why status has
become so dematerialized in advanced economies – why it is based much more on one’s
experiences of education, travel, entertainment, and self-fulfillment, than on what physical
products one owns (see Brooks, 2001; Conniff, 2002; Pine & Gilmore, 1999). By adopting
the new signaling-based view of human mental evolution, it is much easier to see the
connections between prehistoric life and post-industrial life, between fitness-signaling and
status-signaling, between personal identity and brand identity, and between the social
functions of products for consumers and the profitability of products for manufacturers and
marketers.
Aesthetic benefits. Art theory and art criticism throughout the 20th century was
dominated by the novelty-obsessed doctrines of Modernism, Post-modernism, and socialconstructivism. This often led to the privileging of elite, often arcane forms of culture –
whether the “high art” of the New York/London axis or the fashionably obscure forms of “pop
culture” – over the ordinary folk arts enjoyed by most people across culture. Since folk arts
everywhere tend to value virtuosity over novelty (Boas, 1955; Gombrich, 1977, 1979), clarity
over obscurity, and pragmatic sexual signaling-power over theoretical “meaning”, this 20th
11
Fear of fitness indicators
Geoffrey F. Miller
century emphasis often left folk arts out in the cold. A major benefit of fitness indicator
theory is that it puts folk art – ordinary ornamental and representational forms of visual
display – back to the heart of aesthetic theory (Miller, 2000a, 2001). It gives art theorists and
critics a richer new language for describing the forms of art and ornamentation that have
been most popular and widely appreciated across cultures and across history. The fitness
indicator perspective makes it as easy to talk about the aesthetics of the Aeron office chair,
the BMW 540i automobile, and the suburban tract mansion as it was for Modernist theorists
to talk about Picasso, Pollock, and Pei. Also, it reconnects contemporary aesthetic theory to
an alternative, non-Modernist intellectual tradition that runs from Aristotle through a long list
of rather neglected biophilic theorists:
 Charles Darwin (1871),
 Friedrich Nietzsche (1883-1888/1968),
 Herbert Spencer (1890),
 Ernst Grosse (1897),
 Thorstein Veblen (1899, 1914),
 Yrjö Hirn (1900),
 Frans Boas (1955),
 Pearl Binder (1958),
 Desmond Morris (1962, 1985),
 Thomas Munro (1963),
 Ernst Gombrich (1977, 1979),
 Amotz Zahavi (1978),
 Richard Dawkins (1982),
 Pierre Bourdieu (1987),
 Frederick Turner (1992, 1995),
 Denis Dutton (1983), and
 George Hershey (1996, 1999).
This tradition of biophilic aesthetics overlaps perfectly with modern evolutionary psychology
research on the beauty of human faces and bodies (e.g. Etcoff, 1999; Thornhill, 1998).
Spiritual benefits. I can’t stand the word “spiritual”, and I have no idea what it really
means, but some people seem to think it’s important. Many such people seem highly
motivated to understand their place in the cosmos, and to feel the sense of oceanic oneness
with the universe described so well by William James (1902) in The varieties of religious
experience. The fitness indicator view might be useful in promoting such feelings, insofar as
it:
 views our most distinctive human abilities and aspirations as natural outgrowths of
the biological world
 connects the most abstract forms of cognition and culture (the “head”) to the
embodied functions of sexual reproduction and social interaction (the “heart”)
 reconciles our sense of personal identity and uniqueness to our function as conduits
of genetic knowledge from the distant past to the distant future
In these ways, the fitness indicator view is profoundly anti-reductionist and anti-materialist,
while still being scientifically precise, testable, empirically well-supported, and theoretically
fruitful.
References
Andersson, M. (1994). Sexual selection. Princeton, NJ: Princeton U. Press.
Atmar, W. (1991). On the role of males. Animal Behavior, 41, 195-205.
Binder, P. (1958). The peacock’s tail. London: George C. Harrap & Co.
Boas, F. (1955). Primitive art. New York: Dover.
12
Fear of fitness indicators
Geoffrey F. Miller
Bourdieu, P. (1987). Distinction: A social critique of the judgment of taste. Cambridge, MA:
Harvard U. Press.
Brooks, D. (2001). Bobos in paradise: The new upper class and how they got there. New
York: Touchstone.
Buss, D. H. (1999). Evolutionary psychology: The new science of mind. New York: Allyn &
Bacon.
Catchpole, C. K., & Leisler, B. (1996). Female aquatic warblers (Acrocephalus paludicola)
are attracted by playback of longer and more complicated songs. Behaviour, 133(1516), 1153-1164.
Colinshaw, G. (1996). Sexual selection and information content in gibbon song bouts.
Ethology, 102(4), 272-284.
Conniff, R. (2002). The natural history of the rich: A field guide. New York: W. W. Norton.
Chu, K., & Harcourt, P. (1986). Behavioral correlations with aberrant patterns in humpback
whale songs. Behavioral Ecology and Sociobiology, 19(5), 309-312.
Cronin, H. (1991). The ant and the peacock: Altruism and sexual selection from Darwin to
today. Cambridge, UK: Cambridge U. Press.
Crow, J. F. (1999). The odds of losing at genetic roulette. Nature, 397, 293-294.
Darwin, C. (1871). The descent of man, and selection in relation to sex (2 vols.). London:
John Murray. (Reprinted in 1981 by Princeton U. Press.)
Dawkins, R. (1982). The extended phenotype: The gene as the unit of selection. Oxford,
U.K.: W. H. Freeman.
Dessalles, J. L. (1998). Altruism, status and the origin of relevance. From J. R. Hurford, M.
Studdert-Kennedy, & C. Knight (Eds.), Approaches to the evolution of language, pp.
130-147. Cambridge, UK: Cambridge U. Press.
Dissanayake, E. (1992). Homo Aestheticus: Where art comes from and why. New York: The
Free Press.
Dixson, A. F. (1998). Primate sexuality: Comparative studies of the prosimians, monkeys,
apes, and human beings. Oxford, UK: Oxford U. Press.
Dixson, A. F., Halliwall, G., East, R., Wignarajah, P., & Anderson, M. J. (2003). Masculine
somatotype and hirsuteness as determinants of sexual attractiveness to women.
Archives of Sexual Behavior, 32(1), 29-39.
Dutton, D. (Ed.). (1983). The forger's art: Forgery and the philosophy of art. Berkeley:
University of California Press.
Etcoff, N. (1999). Survival of the prettiest: The science of beauty. New York: Doubleday.
Frank, R. (1999). Luxury fever: Why money fails to satisfy in an era of excess. Princeton,
NJ: Princeton U. Press.
Gagnier, R. (2001). The insatiability of human wants. Chicago, IL: U. Chicago Press.
Gangestad, S.W., and Thornhill, R. (1997) Human sexual selection and developmental
stability. In J. A. Simpson & D. T. Kenrick (Eds.), Evolutionary social psychology,
pp.169-195. Hillsdale, NJ: Erlbaum
Giessman, T. (2000). Gibbon songs and human music from an evolutionary perspective. In
N. L. Wallin & B. Merker (Eds.), The origins of music, pp. 103-123. Cambridge, MA:
MIT Press.
Goldberg, T. L. (1995). Altruism towards panhandlers: Who gives? Human Nature, 6(1), 7989.
Gombrich, E. H. (1977). Art and illusion: A study in the psychology of pictorial
representation (5th ed.). London: Phaidon Press.
Gombrich, E. H. (1979). A sense of order: A study in the psychology of decorative art.
Ithaca, NY: Cornell U. Press.
Gong, A., & Gibson, R. M. (1996). Reversal of female preferences after visual experience to
a predator in the guppy, Poecilia reticulata. Animal Behaviour, 52(5), 1007-1015.
Grafen, A. (1990). Biological signals as handicaps. J. Theoretical Biology, 144, 517-546.
Grosse, E. (1897). The beginnings of art. New York.
13
Fear of fitness indicators
Geoffrey F. Miller
Hellofs, L. L., & Jacobson, R. (1999). Market share and customer’s perceptions of quality:
When can firms grow their way to higher versus lower quality? J. Marketing,
January, 16-25.
Hershey, G. (1996). The evolution of allure: Sexual selection from the Medici Venus to the
Incredible Hulk. Cambridge, MA: MIT Press.
Hershey, G. (1999). The monumental impulse: Architecture’s biological roots. Cambridge,
MA: MIT Press.
Hingle, A., Fowler, K., & Pomiankowski, A. (2001). Size-dependent mate preferences in the
stalk-eyed fly Cyrtodiopsis dalmanni. Animal Behaviour, 61(3), 589-595.
Hirn, Y. (1900). The origins of art: A psychological and sociological inquiry. New York:
MacMillan.
Hosken, D. J., Jones, K. E., Chipperfeld, K., & Dixson, A. (2001). Is the bat os penis
sexually selected? Behavioral Ecology & Sociobiology, 50(5), 450-460.
Illouz, Eva (1997). Consuming the romantic utopia: Love and the contradictions of
capitalism. Berkeley, CA: U. California Press.
James, W. (1902/2002). The varieties of religious experience: A study of human nature.
Centenary edition. London: Routledge.
Johnson, R. C. (1996). Attributes of Carnegie Medalists performing acts of heroism and of
the recipients of these acts. Ethology and Sociobiology, 17, 355-362.
Johnstone, R. A. (1995). Sexual selection, honest advertisement, and the handicap
principle: Reviewing the evidence. Biological Reviews, 70, 1-65.
Kanazawa, S. (2000). Scientific discoveries as cultural displays: A further test of Miller’s
courtship model. Evolution and Human Behavior, 21, 317-321.
Kelly, S., & Dunbar, R. I. M. (2001). Who dares, wins. Heroism versus altruism in women’s
mate choice. Human Nature, 12(2), 89-105.
Kirmani, A., & Rao, A. R. (2000). No pain, no gain: A critical review of the literature on
signaling unobservable product quality. J. Marketing, 64, April, 66-79.
Kondrashov, A. (1988). Deleterious mutations as an evolutionary factor III. Mating
preference and some general remarks. J. Theoretical Biology, 131, 487-496.
Klein, N. (2000). No logo. London: Flamingo.
Kodric-Brown, A. (1985). Female preference and sexual selection for male coloration in the
guppy (Poecilia reticulata). Behavioral Ecology and Sociobiology, 17(3), 199-205.
Kohn, M., & Mithen, S. (1999). Handaxes: Products of sexual selection? Antiquity, 73, 518526.
Kotiaho, J. S. (2000). Testing the assumptions of conditional handicap theory: Costs and
condition dependence of a sexually selected trait. Behavioral Ecology and
Sociobiology, 48(3), 188-194.
Langlois, J. H., Kalakanis, L., Rubinstein, A. J., Larson, A., Hallam, M., & Smooth M. (2000).
Maxims or myths of beauty? A meta-analytical and theoretical review. Psychological
Bulletin, 126(3), 390-423
Li, N. P., Kenrick, D. T., Bailey, J. M., & Linsenmeier, J. A. W. (2002). The necessities and
luxuries of mate preferences: Testing the tradeoffs. J. Personality and Social
Psychology, 82(6), 947-955.
Lynn, R., Irwing, P., & Crammock, T. (2002). Sex differences in general knowledge.
Intelligence, 30, 27-39.
Michod, R. E. (1995). Eros and evolution: A natural philosophy of sex. New York: AddisonWesley.
Madden, J. (2001). Sex, bowers and brains. Proc. Royal Society of London B, 268, 833838.
Manning, J. T., & Hartley, M. A. (1991). Symmetry and ornamentation are correlated in the
peacock’s train. Animal Behaviour, 42(6), 1020-1021.
McCrimmon, R. J., Deary, I. J., Frier, B. (1997). Auditory information processing during
acute insulin-induced hypoglycemia in non-diabetic human subjects.
Neuropsychologia, 35(12), 1547-1553,
14
Fear of fitness indicators
Geoffrey F. Miller
Michod, R. E., & Hasson, O. (1990). On the evolution of reliable indicators of fitness.
American Naturalist, 135, 788-808.
Miller, G. F. (1999a). Sexual selection for cultural displays. In R. Dunbar, C. Knight, & C.
Power (Eds.), The evolution of culture, pp. 71-91. Edinburgh, UK: Edinburgh U.
Press.
Miller, G. F. (1999b). Waste is good. Prospect, Feb., pp. 18-23.
Miller, G. F. (2000a). The mating mind: How sexual choice shaped the evolution of human
nature. New York: Doubleday.
Miller, G. F. (2000b). Mental traits as fitness indicators: Expanding evolutionary
psychology’s adaptationism. In D. LeCroy & P. Moller (Eds.), Evolutionary
perspectives on human reproductive behavior (Annals of the New York Academy of
Sciences, Volume 907), pp. 62-74. New York: New York Academy of Sciences.
Miller, G. F. (2000c). Sexual selection for indicators of intelligence. In Bock, G. R., Goode,
J. A., & Webb, K. (Eds.), The nature of intelligence (pp. 260-275.). Novartis
Foundation Symposium 233. New York: Wiley.
Miller, G. F. (2000d). Marketing. In J. Brockman (Ed.), The greatest inventions of the last
2,000 years, pp. 121-126. New York: Simon & Schuster.
Miller, G. F. (2000e). Memetic evolution and human culture. Quarterly Review of Biology,
75(4), 434-436.
Miller, G. F. (2001). Aesthetic fitness: How sexual selection shaped artistic virtuosity as a
fitness indicator and aesthetic preferences as mate choice criteria. Bulletin of
Psychology and the Arts 2(1), 20-25.
Miller, G. F., & Todd, P. M. (1998). Mate choice turns cognitive. Trends in Cognitive
Sciences, 2(5), 190-198.
Miller, P. J. O., Biassoni, N., & Samuels, A. (2000). Whale songs lengthen in response to
sonar: Male humpbacks modify their sexual displays when exposed to man-made
noise. Nature, 405(6789), 903.
Moller, A. P. (1994). Sexual selection and the barn swallow. Oxford, UK: Oxford U. Press.
Moller, A. P., & Petrie, M. (2002). Condition dependence, multiple sexual signals, and
immunocompetence in peacocks. Behavioral Ecology, 13(2), 248-253.
Moller, A. P. & Swaddle, J. P. (1998). Asymmetry, developmental stability and evolution.
Oxford, UK: Oxford U. Press.
Morris, D. (1962). The biology of art. London: Methuen.
Morris, D. (1985). Bodywatching: A field-guide to the human species. London: Jonathan
Cape.
Munro, T. (1963). Evolution in the arts, and other theories of culture history. Cleveland, OH:
Cleveland Museum of Art.
Neiman, F. D. (1997). Conspicuous consumption as wasteful advertising: A Darwinian
perspective on spatial patterns in classic Maya terminal monument dates. In G. A.
Clark & C. M. (Eds.), Rediscovering Darwin: Evolutionary Theory in Archaeological
Explanation. Archaeological Papers of the American Anthropological Association, No.
7, pp. 267-290.
Nicoletto, P. F. (1991). The relationship between male ornamentation and swimming
performance in the guppy, Poecilia reticulata. Behavioral Ecology and Sociobiology,
28(5), 365-370.
Nietzsche, F. (1883-1888/1968). The will to power. New York: Vintage. (Trans. W.
Kaufmann & R. J. Hollingdale from Nietzsche’s notebooks 1883-1888).
Noad, M. J., Cato, D. H., Bryden, M. M. (2000). Cultural revolution in whale songs. Nature,
408(6812), 537.
Packard, V. (1960). The waste makers. New York: Van Rees Press
Payne, K. (2000). The progressively changing songs of humpback whales: A window on the
creative process in a wild animal. In N. L. Wallin & B. Merker (Eds.), The origins of
music, pp. 135-150. Cambridge, MA: MIT Press.
15
Fear of fitness indicators
Geoffrey F. Miller
Petrie, M. (1994). Improved growth and survival of offspring of peacocks with more
elaborate trains. Nature, 371, 598-599.
Pine, B. J., & Gilmore, J. (1999). The experience economy: Work is theater & every
business a stage. Cambridge, MA: Harvard Business School Press.
Plomin, R., DeFries, J. C., McClearn, G. E., & McGuffin, P. (2001). Behavioral genetics (4th
Ed.). New York: Worth.
Reynolds, D. Jr., & Gifford, R. (2001). The sounds and sights of intelligence: A lens model
channel analysis. Personality and Social Psychology Bulletin, 27(2), 187-200.
Ridley, M. (2001). The cooperative gene: How Mendel’s demon explains the evolution of
complex beings. New York: Free Press.
Rowe, L., and Houle, D. (1996) The lek paradox and the capture of genetic variance by
condition dependent traits. Proc. Roy. Soc. London B, 263: 1415-1421.
Saad, G., & Gill, T. (2000). Applications of evolutionary psychology in marketing.
Psychology and Marketing 17(12), 1005-1034.
Shuker, D., Bateson, N., Breitsprecher, H., O’Donovan, R., Taylor, H., Barnard, C., Behnke,
J., Collins, S., & Gilbert, F. (2002). Mating behavior, sexual selection, and copulatory
courtship in a promiscuous beetle. Journal of Insect Behavior, 15(5), 617-631.
Sommerfield, A. J., Deary, I. J., & McAulay, V. (2003). Moderate hypoglygemia impairs
multiple memory functions in healthy adults. Neuropsychology, 17(1), 125-132.
Spencer, H. (1890). The principles of psychology (3rd Ed.). London.
Thornhill, R. (1998). Darwinian aesthetics. In C. Crawford & D. Krebs (Eds.), Handbook of
evolutionary psychology, pp. 543-572. Mahwah, NJ: Erlbaum.
Todd, P.M., & Miller, G. F. (1999). From Pride and Prejudice to Persuasion: Satisficing in
mate search. In G. Gigerenzer & P. Todd. (Eds.), Simple heuristics that make us
smart, pp. 286-308. New York: Oxford U. Press.
Twitchell, James (1999). Lead us into temptation: The triumph of American materialism.
New York: Columbia U. Press
Turner, F. (1992). Beauty: The value of values. Charlottesville, VA: U. Virginia Press.
Turner, F. (1995). The culture of hope: The new birth of the classical spirit. New York: Free
Press.
Veblen, T. (1899/1994). The theory of the leisure class. Reprinted by Dover, New York.
Veblen, T. (1914/1964). The instinct of workmanship and the state of the industrial arts.
Reprinted by Transaction Publishers, New Jersey.
Wilkinson, G. S., Presgraves, D. C., & Crymes, L. (1998). Male eye span in stalk-eyed flies
indicates genetic quality by meiotic drive suppression. Nature, 391(6664), 276-279.
Wilson, E. O. (1999). Consilience: The unity of knowledge. New York: Random House.
Yeo, R. A. (1999). Asymmetry, developmental stability, and evolution. Laterality, 4, 389-394.
Yeo, R. A. , Gangestad, S. W., Edgar, C., & Thoma, R. (1999). The evolutionary-genetic
underpinnings of schizophrenia: The Developmental Instability model. Schizophrenia
Research, 39, 197-206.
Zahavi, A. (1978). Decorative patterns and the evolution of art. New Scientist, 19, 182-184
Zahavi, A., & Zahavi, A. (1997). The handicap principle: A missing piece of Darwin's puzzle.
New York: Oxford University Press.
Zebrowitz, L. A., Hall, J. A., Murphy, N. A., & Rhodes, G. (2002). Looking smart and looking
good: Facial cues to intelligence and their origins. Personality and Social Psychology
Bulletin, 28(2), 238-249.
Zechner, U., Wilda, M., Kehrer-Sawatzki, H., Vogel, W., Fundele, R., & Hameister, H.
(2001). A high density of X-linked genes for general cognitive ability: a run-away
process shaping human evolution? Trends in Genetics, 17(12), 697-701.
16
Fear of fitness indicators
Geoffrey F. Miller
17