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Why bother with history? The simple answer is that to fully under-
stand the present, one must look to the past. Science does not exist in a
vacuum, and a history of science without social context is lifeless. As I
will discuss, various social, economic, religious, and political notions
and values beginning prior to the nineteenth century and leading up to
the present have shaped evolutionary theory. As well, opposition to
evolutionary theory has resurfaced at regular intervals. We are currently
experiencing a period of strong opposition to evolutionary theory. Not
since the 1920s and 1930s has opposition to evolution from Christian
evangelicals been so pronounced, with confrontational, hostile attacks
designed to raise doubt about evolution and promote creationism. Not
surprisingly, opponents of evolution have also objected to the study and
teaching of human sexuality (sexology), often linking permissive sexu-
ality and homosexuality with the teaching of evolution. In addition,
some secular political groups and scientists have assailed evolutionary
theory for its presumed assumptions and implications about human na-
ture. It is my hope that an appreciation of the history of evolutionary
theory will help the reader anticipate, understand, and evaluate
opposition to it. For when the subject is the evolution of human
sexuality, critics often come from several directions.
Coincidentally, the birth of evolutionary theory just preceded the
birth of modern human sexuality: that is, the invention of the “hetero-
sexual” and “homosexual.” While evolution and human sexuality
crossed paths briefly in the early days, enthusiasm for an evolutionary
theory of human sexual attraction and behavior took nearly 100 years to
develop (Miller, 2000). New interest in the evolution of human sexual-
ity arose about the time that the first homosexual rights groups began
to form (Bullough, 1994; Terry, 1999) and preceded by a few years a
radical shift in American culture toward more open sexuality. This
cultural shift was fueled by feminism, the birth control pill, and in-
creased recreational drug use. Renewed scholarly interest in the study
of human sexuality can be traced to the enormous publicity (and noto-
riety) received by Alfred Kinsey’s two volumes on human sexual be-
havior in 1948 and 1953. In America for the last 50 years, both
evolution and sexology have been favorite targets of far-right secular
and religious conservatives in the so-called “Culture Wars” (Hunter,
1992).
Michael R. Kauth 25
DARWIN’S IDEA:
FROM NATURAL SELECTION
TO EVOLUTIONARY PSYCHOLOGY
In the distant future I see open fields for far more important re-
searches. Psychology will be based on a new foundation, that of
the necessary acquirement of each mental power and capacity by
gradation. Light will be thrown on the origin of man and his his-
tory.
Darwin did not publish his ideas for another 20 years, and then only
after learning that Alfred Russel Wallace had made a similar discovery.
In 1858, as co-discoverers, Darwin and Wallace presented their ideas
about natural selection to the Linnaean Society,1 and one year later Dar-
win published On the Origin of Species By Means of Natural Selection,
eloquently describing the vast diversity of replicating species as a func-
tion of a natural physical process. He argued that over generations traits
are favored that enhance the organism’s survival and reproduction in a
particular environment better than alternatives. Small beneficial
changes accumulate and over time can produce large effects. He re-
ferred to traits that enhance an organism’s survival and reproduction as
adaptations (Darwin, 1859/1958). Darwin proposed two basic explana-
tions for the origin and diversity of species: natural selection and sexual
selection.
In natural selection, individuals who are best suited to the ecology
survive and transmit their heritable characteristics to their offspring.
Thus, evolutionary success means producing viable offspring within a
particular ecology. Darwin (1958) also realized that individuals in a
species vary in their expression of traits and that these variations are at
least partially inherited. Thus, trait differences among individuals allow
for differential reproductive success (leaving more viable offspring
than rivals). On the average, individuals with robust adaptational traits
will out-reproduce individuals with weaker versions of the traits, or
none at all. As individuals with the beneficial traits out-reproduce their
competitors, the traits spread throughout the population over many
generations.
Of course, organisms have many traits. Reproductive trade-offs en-
sure that variability of a given trait is maintained within the population.
Darwin reasoned that trait mutations spontaneously occur in the popula-
tion. Although most mutations are not beneficial and in fact impair the
organism’s survival and disappear with the individual (Badcock, 2000),
rare mutations may benefit survival and reproductive success. Over
time, mutations may significantly alter or split a species.
The environment (e.g., climate, food sources and availability, preda-
tors, parasites) “selects” the trait (Cosmides, Tooby, & Barkow, 1992).
This aspect of natural selection is often overlooked by critics who ac-
cuse evolutionists of genetic or biological determinism (see Rose &
Rose, 2000; criticism of EP is discussed below). Simply, particular
ecologies favor particular traits for survival. Those organisms that pos-
28 HANDBOOK OF THE EVOLUTION OF HUMAN SEXUALITY
sess environmentally favored traits (or can develop them) flourish in the
ecology, while other organisms fail. If the trait is heritable, it can be
transmitted to offspring who go on to produce a more resilient, prolific
lineage. In short, natural selection is about reproductive success in a
given environment. Survival is only the means to reproductive success.
Darwin’s theory of natural selection gained steady followers and, by
the turn-of-the-century, it was the predominant explanation for evolu-
tion among biologists (Larson, 2004). The theory of sexual selection
had a different history. Darwin (1871/1981) expanded on his earlier
ideas about sexual selection in The Descent of Man and Selection in Re-
lation to Sex, viewing it as a major force in evolution, although he was
puzzled by why it worked (Miller, 2000). He proposed that sexual se-
lection explains how costly traits and subgroup differences (varieties)
persist in the population, such as sex and racial differences in humans.
Darwin saw that some highly favored traits, particularly in males, exact
a high cost to the organism by shortening survival through health conse-
quences or by increasing risk of predation. Sexually selected traits
broadly include sexual ornamentation (e.g., bright feathers, large ap-
pendages, large physical size) and ritual courtship displays (e.g., strut-
ting, erection of peacock’s tail) (Miller, 2000; Ridley, 1993). If
particular traits are preferred by one sex more than the other, dramatic
subpopulation differences within a species can result, such as bright
coloration in males but drab females, hulking males but smaller fe-
males. Darwin reasoned that these traits persist in the population be-
cause they confer a reproductive advantage to individuals. He
identified two independent forms of sexual selection: competition for
mates (intrasexual selection) and mate choice (intersexual selection
or sometimes, epigamic selection). For reasons discussed below, males
primarily compete for mates across species, although female-female
competition for mates does occur. Likewise, female mate choice is pre-
dominant across species, but males also make mate choices. In short,
what distinguishes sexual selection from natural selection is that sexual
selection is all about sex.
Naturalists in Victorian England understood that males attract fe-
males by ornamentation and courtship displays and compete with each
other for mates, but the idea of female choice conflicted with their no-
tion of passive females (Larson, 2004; Miller, 2000). Darwin (1981)
had compiled a great deal of evidence for female mate choice. However,
he was unsure why females choose and why males usually do not.
Miller (2000) has argued that sexual prudery as well as male bias among
scientists contributed to the marginalization of sexual selection for
Michael R. Kauth 29
nearly 100 years. But, times and thinking changed and, for the past 30
years, sexual selection theory has generated a number of insights into
human sexuality. Sociobiologists and evolutionary psychologists have
employed sexual selection theory to explain why individuals are at-
tracted to certain features in a mate and to explain mating patterns, mate
monitoring, infidelity, and jealousy.
Intrasexual Selection
males may not mate for a season, or ever. However, many females enjoy
an average reproductive success.
Intrasexual selection, or male-male competition for mates, is likely to
produce sexual weapons (e.g., horns, spurs, large claws, increased size)
to intimidate or fight rivals, as well as sexual ornaments (e.g., elaborate
tails, bright coloration, eliciting odors, behavioral displays) to attract
females. The sexual weapon is selected if it contributes to dominance
over rival males, since dominant males are more likely to mate and
produce abundant offspring than defeated males who may not mate
at all (Miller, 2000; Wilson, 2000). Over time a sexual weapon can
become more pronounced as the trait is favored in male-male compe-
tition and preferred by females as an honest indicator of fitness, con-
sistent with Fisher’s runaway selection (Fisher, 1930/1958). (For
more discussion on sexual selection theory and runaway selection, see
Sefcek, Brumbach, Vasquez, & Miller, this volume.)
Male-male competition for mates spawns a social hierarchy and fuels
intra-individual conflict in an effort to gain status. Individual and coali-
tion-based aggression is common among traditional human hunting and
foraging and agricultural societies, and coalition-based aggression is of-
ten associated with personal gain, including more resources, more
wives, and greater reproductive success (Geary, 1998). Even so, many
men are targets of aggression and are dominated by a few men. To re-
duce intrapersonal aggression and increase social status, men may ac-
quiesce when confronted by more dominant rivals, form alliances to
gain resources, defend against threats, and provision food with the ulti-
mate goal of gaining access to quality mates (de Waal, 2002; Symons,
1979; Tiger, 1969/1984).
The evolutionary history of male-male competitive aggression ac-
counts for men’s preferences for dominance-submission dynamics, in-
cluding a preference for social hierarchies and signs of status, power,
and control (Symons, 1979; Tiger, 1969/1984). As a result, men in gen-
eral should be sensitive to social status and symbols of rank; this may
account for the long existence of patriarchy, social classes, insignias as
status markers, and bragging among men. In addition, men should be
drawn to strong leaders and should enjoy male-male play-fighting (e.g.,
sports) (Buss, 1994/2003). Male-male competition may also explain the
formation of male alliances or coalitions, a preference for male compan-
ionship, and the desire for loyalty among buddies. Effective long-term
alliances are grounded in mutual obligation, trust, and probably some
degree of affection. Even so, men may form alliances opportunistically
and terminate quickly in pursuit of dominance (Symons, 1979). Finally,
Michael R. Kauth 31
Intersexual Selection
disease, especially in less fit males (Zuk, Johnsen, & Maclarty, 1995).
Parasite infestation lowers the quality of secondary sex characteristics
and lowers mating opportunities (female choice) (Folstad & Karter,
1992; Hamilton & Zuk, 1982; Zuk, Thornhill, & Ligon, 1990). Further,
high levels of testosterone are associated with behavioral and health
risks and a shortened life span (Geary, 2005). Thus, a male with robust
secondary sex characteristics–despite the parasite load in the environ-
ment and despite behavioral and health costs of high hormone levels–is
broadcasting his “good genes” and phenotypic condition (Zahavi,
1975).
In human males, sexually selected fitness indicators include mascu-
line, symmetrical features, such as broad shoulders, large chest, narrow
waist, large hands, taller than average height, and vigorous physical ac-
tivity (Buss, 2003; Miller, 2000; Symons, 1979). (For a more discussion
on fitness indicators and mate choice, see Sefcek et al., this volume.) In-
deed, women generally report a preference for handsome, masculine
men who are somewhat taller than average, athletic, and physically fit
(Barber, 1995; Thornhill & Gangestad, 1993; Thornhill, Gangestad, &
Comer, 1995).
Some preferred male traits do not directly benefit female reproduc-
tion. However, by intimidating or defeating rivals or by attracting
mates, the traits indirectly advantage reproductive fitness. Because sex-
ual selection is not a conscious process, the preferred traits in a mate are
simply perceived as “attractive” (Symons, 1979). If the “attractive”
male traits are heritable, then the female can count on her sons having
similar “sexy” traits. In a polygynous society (one male mates with
many females), a “sexy” son is advantaged over less attractive males, as
long as females continue to favor the trait. Further, the daughters can in-
herit their mother’s attraction to those traits, producing a positive feed-
back loop. Runaway selection will lead to the traits and to the
preference for those traits becoming more pronounced over time in the
population (Fisher, 1958).
Other preferred male traits may have no known benefit to female re-
productive success. Even so, if the trait is consistently preferred,
intersexual selection will drive it to become more pronounced but vari-
able in men. An example is the human penis (Miller, 2000). The flaccid
human penis is much larger and more visible than the penises of goril-
las, chimpanzees, orangutans, and bonobos; flaccid nonhuman primate
penises are typically thin and hidden. When erect, the silverback go-
rilla’s penis is less than two inches long, and the erect chimpanzee’s pe-
nis averages about three inches. However, the erect human penis
Michael R. Kauth 33
averages between four and eight inches (Men’s Health, 2006). Miller
(2000) has argued that the human penis was selected by hominin fe-
males for its tactile properties during copulation. In other words, during
sexual activity, size mattered to women (compared to a thin, small pe-
nis, like other primates). If this is indeed the case, Miller has suggested
that clitoral orgasm may be how ancestral women identified truly fit,
attentive, energetic male lovers.
Clearly, the human penis was not selected for visual appearance, or it
would be brightly colored or exquisitely ornate, rather than plain and
unadorned. “Sperm competition” is often cited as an explanation for the
larger human penis, but this is unlikely (Miller, 2000; Taylor, 1996).
Sperm competition refers to the amount of semen produced in order to
wash out or block a competitor’s semen (Baker & Bellis, 1995). Testicle
size is better correlated with sperm competition than penis size (Miller,
2000). For instance, silverback gorillas are vigilant guardians of their
harem and have very small testicles because sperm competition is low,
while chimpanzees are polygynous and males have comparatively large
testicles. Human males have testicles that are intermediate between go-
rillas and chimpanzees, suggesting that early men engaged in mild-to-
moderate polygynous mating and sperm competition. There is sugges-
tive but inconclusive evidence that the human penis evolved via
intrasexual selection as a threat or dominance display to other men, sim-
ilar to some nonhuman primates (Etcoff, 1999; Miller, 2000). When ap-
proached by an unknown animal, male vervet monkeys get an erection,
displaying their red and blue genitals, and make a threatening face. Al-
though I know of no human societies where men publicly display their
penis to threaten other men, it is possible. The pervasive phallic symbol-
ism in most human societies and male insecurity about their penis size
(at least among contemporary Western men), especially in situations
where comparison is likely (e.g., in the locker room), lend support for
the idea that a large human penis signaled dominance.
In addition to selecting for genetic quality and health (excluding
non-fitness traits such as penis size), women prefer men who have re-
sources to invest in them and their children and who are willing to in-
vest. (See Sefcek et al., this volume.) Older age, high social status,
social dominance, and alliances indicate resource potential (Kauth,
2000). Across 37 cultures, Buss and colleagues (Buss, 1989, 1998,
2005; Buss, Larsen, & Westen, 1996; Buss & Schmitt, 1993) have
found that in general women prefer men who are older, resourceful,
masculine, and emotionally committed to invest in a relationship and
family. In this context, emotional jealousy can be seen as a woman’s
34 HANDBOOK OF THE EVOLUTION OF HUMAN SEXUALITY
even when producing milk, are large human female breasts a sexually
selected trait due to male choice? Probably so. It is likely that large,
symmetrical human breasts evolved as fitness indicators of fertility and
good health (Miller, 2000). As noted earlier, breast symmetry is associ-
ated with fertility (Grammer, Fink, Juette, Ronzal, & Thornhill, 2001;
Manning, Scutt, Whitehouse, & Leinster, 1997), which is easier to
judge when breasts are large and more visible. As well, fertility is asso-
ciated with youth, and a young woman’s breasts are high and firm,
while older women have lower, droopier breasts, features that are ac-
centuated if the breasts are large (Miller, 2000). Thus, it is likely that
larger (but variable) human breasts evolved via men’s choice for this
trait as an indicator of fitness and fertility. If breast size and shape were
not fitness indicators, these features would be more consistent in
women.
coined this term and championed the idea that (Lamarckian) evolution
(toward superiority) could be sped up by encouraging attractive men
and women of good “fitness” to marry each other and produce children.
Galton’s so-called positive eugenics (“more children from the fit”)
stood in contrast to negative eugenics (“less children from the unfit”),
which was strongly influenced by social contagion and degeneracy
theories. In the early twentieth century, for example, birth control was
promoted in the United States as a means to reduce the number of chil-
dren from degenerate (re: nonwhite) races (Terry, 1999). Negative
eugenicists called for eliminating aid to the poor and mandatory sexual
sterilization of individuals who suffered from feeble-mindedness or
other hereditary disorders. By 1935, more than 60,000 persons had
been sexually sterilized in the United States, mostly in California
(Larson, 2004). In Germany, the idea of limiting the spread of heredi-
tary disease in the population by isolating or sterilizing affected indi-
viduals was particularly popular. Between 1933 and 1939, 300,000
Germans were sexually sterilized. After 1939, euthanasia programs re-
placed the sterilization programs. Contagion and degeneracy theories
and a peculiar version of social evolution supported Nazi ideology and
their extermination of millions of Jews, Gypsies, and gay men and lesbi-
ans (Larson, 2004; Oosterhuis, 1991). [In the 1990s, Serbian President
Slobodan Milosevic cited a similar eugenicist rationale for the “ethnic
cleansing” of more than 200,000 Bosnian Muslims (Milosevic to face,
2001).] The atrocities committed by Nazi eugenicists have been used
repeatedly to cast suspicion on the motives of evolutionists.
effort to maximize their own investment. Children will resist their par-
ents’ attempts to control and shape them (e.g., cry, resist, rebel), while
demanding more parental resources (e.g., by complaining, manipulat-
ing, throwing tantrums). Further, children compete with their siblings
for scarce parental resources by coercing, cheating, and fighting or by
manipulating a parent’s affection (Badcock, 2000; Pinker, 2002).
Trivers (1971, 1985) also developed the idea of reciprocal altruism
or reciprocity to explain cooperation between unrelated individuals.
Anthropological studies have long demonstrated that cooperative re-
lationships are common in human societies and facilitate survival,
even though altruism should bear a high cost for the helper. Trivers ar-
gued that if helping others entailed a social obligation to assist the
helper when needy, cooperation among unrelated individuals would be
favored. The relatively long period of parental care and long human
lifespan presents hundreds of thousands of opportunities to aide others
and be aided in times of need. Thus, reciprocity helps to account for
friendship and alliances, as well as sympathy, gratitude, and guilt.
Being reproductively selfish, some individuals will take advantage of
others’ generosity, without reciprocating. In fact, some cheating is
adaptive (Trivers, 1971, 1985). Given that cheating is a constant threat,
Trivers proposed that individuals evolved psychological mechanisms to
regulate their degree of cooperation and to detect cheating or deceit in
others. As cheating became more sophisticated to avoid detection,
cheater detecting became more pervasive. Indeed, very effective cheat-
ers might disguise their true motives even from themselves to mask
their awareness of intent. Thus, Trivers proposed that self-deception,
including biased perceptions, biased memories, and biased reasoning,
are major components of an evolved human psychology.
Assumptions
Evolutionary Theory
Evolutionary Psychology
first, that the details of present and past social arrangements are the in-
evitable manifestations of the specific actions of genes” (p. 236).
Dawkins (1985), in his review of their book, rejected this charge as “a
simple lie.” He added, “The myth of the ‘inevitability’ of genetic effects
has nothing whatever to do with sociobiology, and has everything to
do with Rose et al.’s paranoic and demonological theology of science”
(p. 59). S/EP have emphasized evolved traits and universal features be-
cause that is what they study. Sociobiologists and evolutionary psychol-
ogists know that they are discussing whole organisms with extensive
developmental histories and varied social relationships (e.g., Buss,
2003; Pinker & Bloom, 1992; Symons, 1979; Wilson, 2000).
Gould (1981) has complained that sociobiologists’ emphasis on hy-
pothetical genes “for” a trait reveals their genetic determinism, lament-
ing: “All biologists know that there is no gene ‘for’ aggression, any
more than for your lower-left wisdom teeth” (1981, p. 359). Gould
seems to imply that genes and environment are independent, when in
actuality, behavior is not simply cultural, and wisdom teeth are biologi-
cally determined (Ridley, 1993). The Roses (2000) have also fulmi-
nated against “biology-as-destiny” ideologies, taking pains to trace
their lineage from Aristotle’s patriarchal concept of human nature
through Nazi eugenics to S/EP’s presumed determinism. However,
sociobiologists and evolutionary psychologists view development as an
ongoing interaction between the organism’s genes and everything else
that influences them (the environments). Change the gene or the envi-
ronment, and the outcome changes (Tooby & Cosmides, 1992). Gould
(1981), for example, seems to suggest that biological, evolved mecha-
nisms are immutable, although environmental processes permit change
and choice (Pinker, 2002; Segerstråle, 2000).
Tooby and Cosmides (1992) have acknowledged that the form-to-
function approach–explaining why already known features came to be
as they are–runs the risk of post hoc explanation but noted that physi-
cists and geologists accept such risk when they investigate known phe-
nomena. “In science,” they pointed out, “this is usually called
‘explanation’” (p. 77). Further, an explanation is not post hoc if the the-
ory predicts a fact that was unknown. Evolutionary analyses have pre-
dicted thousands of discoveries that were later confirmed. Adaptational
hypotheses are testable, although sufficient evidence is often lacking to
know which explanation from a set of alternative explanations is correct
(Pinker & Bloom, 1992). Gould and Lewontin (1979) have argued that
many supposed adaptations identified by sociobiologists are really
non-adaptive by-products. Despite their shrill complaints, Pinker
48 HANDBOOK OF THE EVOLUTION OF HUMAN SEXUALITY
(2002) has kindly interpreted the pair’s arguments as simply a call for
restraint and rigor in attributing adaptation.
Finally, charges that S/EP is “bad” science are largely related to dis-
agreements between political and philosophical ideologies (Gould,
1999; Levins & Lewontin, 1985; Lewontin, Rose & Kamin, 1984; Rose
& Rose, 2000). In particular, Gould, Lewontin, and the Roses have op-
posed description of natural processes that refute their worldview, de-
manding that science and nature conform to their political values.
Dawkins has stated that natural selection does not advocate anything,
declaring that Nature is a poor model for deriving values (cited in
Segerstråle, 2000).
Valid Criticisms
Bisexuality?
Intelligent Design
CONCLUSION
This brief history has described the social and political context of
evolutionary theory, particularly with regard to human sexuality. Many
different disciplines are now engaged in evolutionary analyses, each
with their own assumptions, concepts, methodologies, and language.
Sociobiology’s attempt to encompass other disciplines erupted into
hostile attacks that largely failed to incite a scholarly discussion about
knowledge, human nature, moral truth, and the role of science in soci-
ety. However, EP has proven to be a robust approach for explaining
evolved sexual psychologies.
FURTHER READINGS
Buss, D. M. (2005). Handbook of Evolutionary Psychology. Hoboken, NJ: Wiley.
Darwin, C. (1871/1981). The Descent of Man and Selection in Relation to Sex. Prince-
ton, NJ: Princeton University Press.
Geary, D. C. (1998). Male, Female: The Evolution of Human Sex Differences. Wash-
ington, DC: American Psychological Association.
Michael R. Kauth 61
NOTES
1. Darwin formulated his ideas about natural selection and sexual selection during
and immediately after his voyage on the Beagle but kept his ideas largely a secret for
about two decades while refining his argument (Larson, 2004). He worried about the
effect his theory would have on the religious beliefs of others, particularly his Christian
wife. In 1858, fellow naturalist Wallace asked Darwin to review his manuscript, de-
scribing the principles of natural selection (Badcock, 2000; Larson, 2004). Wallace
had independently discovered natural selection. While Wallace emphasized the pres-
sure of ecological forces, Darwin stressed individual competition as a driving force for
evolution.
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