I
ESSAY
What Is Sex? What Is Gender?
JOAN ROUGHGARDEN
F ASKED, most people would probably say that the
purpose of sex is to reproduce. This is not correct.
Many species reproduce without sex. For example,
there are geckoes in South Pacific islands who repro-
duce asexually. Many other species do too, including
whiptail lizards from the American southwest. In these
species, the eggs carry the full complement of genes to begin
with. (They are formed through mitosis, not meiosis.) Hence,
the eggs in such species do not need to be fertilized to develop
into embryos. Because the eggs do not need fertilization, males
do not exist in these species, which consist solely of females.
Thus reproduction is perfectly possible without sex, and the
purpose of sex is to provide the offspring with a mixture of
genes from two parents. Gene mixing is the purpose of sex, not
reproduction per se.
What about the role of male and female in this equa- in many other species. For starters, there are three types of her-
tion, and must it be a binary system? When asked, most people
define male and female on the basis of external appearance. A
male lion has his mane, a rooster his cockle-doodle-do, and so
forth. A closer inspection reveals different genitalsin mam-
mals, males with a penis, and females with a vagina. A look in-
side the body reveals males with testes and females with
ovaries. In mammals, a still deeper look into the cells reveals an
XY set of sex chromosomes in males and an XX set in females.
You may be surprised to learn, then, that these traits do not
define male and female for biologists. These traits vary too
much among individuals within a species and especially across
species to serve as defining traits. Instead, to have a definition
of male and female that
applies to all living things,
from seaweed to sea lions,
from mice to men, biolo-
gists define a male as an
individual who produces
only sperm and a female
as one who produces only
eggs throughout their
lives, while a hermaphro-
dite is an individual who
produces both sperm and
eggs over the life cycle.
The question then be-
comes, what is a sperm
Joan Roughgarden, biology professor emerita at Stanford, is the author
of Evolutions Rainbow; Diversity, Gender, and Sexuality in Nature and
People (2004). This piece is excerpted and adapted from an article, The
Gender Binary in Nature, across Human Cultures, and in the Bible,
which appears (in English) in Transsexualitt in Theologie und Neu-
rowissenschaften, Gerhard Schreiber, ed. (Walter De Gruyter Inc., 2016).
20
and what is an egg? Each gamete is a cell that contains half of
the genetic material of an adult. Two gametes fuse to produce
the cell that multiplies to become an embryo. The gametes are
of very unequal size, and there are only two sizes per species.
This is the only binary that exists. The individuals in virtually
all sexually reproducing species produce only two sizes of ga-
metes, one large and one small. There are almost no species
whose members produce sizes varying continuously from small
to large. Moreover, if the individuals of a species produce ga-
metes of only one size, such as some fungi, then male and fe-
male are not defined for that species. So, practically speaking,
the only universal sex binary in biology is the dichotomy be-
tween egg and sperm.
The prevalence of hermaphroditic species among both
plants and animals can shed light on male and female sex roles
maphrodite. A simultaneous hermaphrodite is an individual who
produces both sperm and eggs at the same time. A sequential
hermaphrodite produces eggs and sperm at different times dur-
ing life, and these come in two varieties: protandrous (male first,
then female) and protogynous (the reverse). Sequential her-
maphrodites change sex during life, shifting from making sperm
to making eggs, or vice versa. A sequential hermaphrodite also
changes in incidental characteristics, such as sex-specific body
colors, shapes, and behavior.
Most plants are hermaphroditic. For example, a flower typ-
ically contains anthers that disperse pollen and a stigma with a
sticky surface on which pollen grains are deposited by a polli-
nator or by the wind. The anthers are the male part
of the plant, the pollen grains are the sperm, and
the stigma and the ovule at the base of the stigma
that becomes the seed comprise the female part.
Hermaphrodites are the rule, not the exception,
throughout the plant kingdom. Only about six per-
cent of plant species have separate sexes.
Animals, too, present an enormous number
of hermaphroditic species, including many ma-
rine invertebrates, such as sea urchins, snails, bar-
nacles, and corals. Even among vertebrates, many
species are hermaphroditic. If one goes snorkel-
ing on a coral reef, about fifty percent of the fish
one sees belong to hermaphroditic species: parrot
fish, wrasses, groupers, etc. Across both marine
and terrestrial environments, about six percent of all animal
species are hermaphroditic. However, if insects are not counted,
the percentage rises to about 33 percent of animal species. Al-
though hermaphroditic animal species are common in the ma-
rine environment, they are relatively rare in terrestrial
environments, an unexplained fact that has invited fun but un-
proven conjectures over the years.
The Gay & Lesbian Review / WORLDWIDE
 SEX ROLES OSCAR WILDE TOURS & HE TRAVEL PRESENT
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of fertilizing many females with an almost infinite supply of
sperm, while the female is limited to producing a finite number
of large eggs. Hence, females are in control; they make the
choice of a mate lest they waste their expensive eggs on inferior
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JanuaryFebruary 2017 21
 GENDER
Gender, as distinct from sex, is normally thought of as per-
taining uniquely to humans, something thats constructed by
culture and finding highly divergent expression in different
cultures. What could gender mean when applied to animals?
For animals, I take gender to mean the appearance, behavior,
and life history of a sexed body, which is a body classified ac-
cording to the size of the gametes produced. Thus, gender is
appearance plus actionhow an organism uses its morphol-
ogy (color and shape) and behavior to carry out a reproduc-
tive role.
As we have seen, sex at the whole organism level is not a bi-
nary, even though at the gamete level the egg/sperm size dis-
tinction does constitute a binary. In fact, gender is even less
binary than sex. Even in species with exactly two sexes classi-
fied as male and female, the number of genders can be greater
than two. There may be, say, two genders of males, as when
those with the smaller gamete come in two distinct types based
on their appearance, behavior, and life trajectory.
Sandpipers, for example, are a species with three male gen-
ders and one female gender. Ruffs are sandpipers, shorebirds
that breed in summer in northern Europe. Ruffs owe their name
to a ring of feathers that males have around their necks. Shown
here is the male gender with a dark ruff, a white ruff, and no
ruff. Below is the female, also with no ruff. These genders are
genetic, and about twenty percent of the males with a ruff have
the white variety and eighty percent the black variety.
Ruffs often mate in leks, which are in effect male red-light
districts where males congregate to attract females. The black-
ruffed males defend small courts within the lek and within
these each displays to the visiting
females. The white-ruffed males
do not defend courts within the
lek and instead keep company
with the females as they feed.
Whats remarkable is the interac-
tion between the black- and
white-ruffed males. When a
white-ruffed male is nearby and a
black-ruffed male is alone on a
court, the black-ruffed male does
a little dance that invites the
white-ruffed male to join him on
the court he has been defending.
Females who then arrive at the lek
to mate prefer a black-white team
of males over a lone black-ruffed
male. Both males jointly court
and then mate with the female. A
black-ruffed male obtains more
matings when a white-ruffed male
is present than when by himself,
even though the matings are
shared. Its not clear why a female
finds a team more attractive than a
single black-ruffed male. Most in-
vestigators seem to assume that a
female automatically finds two
22
males better than onethe more masculinity the better. If more
total masculinity is so important, then two black-ruffed males
could simply team up with each other. Why two genders?
My hunch has been that a white-ruffed male builds rela-
tionships with females while he is with them away from the lek.
While the black-ruffed male is defending a court against other
black-ruffed males, the white-ruffed male is flying with females
in the field and presumably getting to know them. Perhaps the
white-ruffed male can, so to speak, make introductions when
the females arrive at the lek. He can act as a go-between, a mar-
riage broker, who can introduce a black-ruffed male to a female
and vouch for his safety. Whatever the explanation, this exam-
ple serves to illustrate the phenomenon of gender multiplicity in
animals. Two sexes does not imply only two genders.
Finally, a black-ruffed male has also been observed mating
with a ruff-less male. This is an example in the wild of a ho-
mosexual matingi.e., between two individuals who produce
the same size of gamete. However, this example is also hetero-
genderal because the two individuals belong to different male
genders.
Even in species that have only two sexes and two genders,
the gender categories are not absolute, and transgender indi-
viduals occur. The best studied example of this phenomenon
occurs in a set of sun angel hummingbird species from the
Andes. Male sun angel hummingbirds have colorful feathers on
their throats called a gorget. A female with a gorget is referred
to here as a masculine female. She also has a comparatively
shorter bill. Conversely, feminine males also exist with spe-
cial female traits such as a longer bill.
Of 42 hummingbird species surveyed throughout the
Andes, seven had both masculine females and feminine males,
nine had masculine females and no femi-
nine males, two had feminine males and
no masculine females, and 24 had neither
masculine females nor feminine males.
Pooling the species with either masculine
females, feminine males, or both, revealed
that 52 percent of the females were mas-
culine and two percent of the males were
feminine.
Males use their gorgets in territorial de-
fense of the common short flowers that fit
their shorter bills. Masculine females can,
like the males, defend a territory of short
flowers to feed in. Conversely, the femi-
nine males have longer bills than mascu-
line males, even longer than feminine
females. Hence, feminine males use dif-
ferent flowers from the masculine males,
namely relatively rare long tubular flowers that do not
need to be defended in a territory. Thus, the masculine
females occupy slightly different niches, i.e., they have
slightly different occupations, from the feminine fe-
males. Conversely feminine males also occupy some-
what different niches from the masculine males. We
see then that gender expression in birds in part reflects
a gendered difference in occupation, and transgender
birds are those whose occupation crosses over into the
occupation typical of the other gender.
The Gay & Lesbian Review / WORLDWIDE
 HOMOSEXUALITY
By homosexuality, I refer to same-sex mounting and to mu-
tual touching of genitals. Today, the reality of extensive homo-
sexuality among animals is generally accepted among
biologists. The 300 or so known examples of homosexuality in
animals, surely an underestimate, reveal much diversity. In
some species only males are homosexual, in some, only fe-
males, and in others, both sexes. In almost all, the homosexual-
ity is mixed in with heterosexuality by the same individual. In
some species, a small fraction participates in homosexuality; in
others, like our closest relative the bonobo chimpanzee, every
animal does. But rather than wondering why homosexuality ex-
ists in some animals, we may well ask why homosexual behav-
ior is not dominant, perhaps mixed in with some minimal
amount of heterosexuality to ensure reproduction?
Homosexuality is widely distributed across many higher
taxa and is especially well-documented in birds, mammals,
and insects. This wide distribution implies that homosexuality
has originated many times in the animal kingdom. Concerning
primates, their family tree indicates a pattern. From its base
near the ground, the primate tree trunk splits first into the
prosimians and the anthropoids. The prosimian branch, in-
cluding bush babies, lemurs, and tarsiers, appear to have only
incidental same-sex mounting and no same-sex courtship. The
anthropoid branch splits into two sub-branches, New World
and Old World primates. The former, which includes mar-
mosets, tamarins, and monkeys with prehensile tails, such as
the spider monkey, display some homosexual behavior. In the
Old World primates, though, homosexual courtship becomes
prominent. These primates, including macaques, baboons, gib-
bons, orangutans, gorillas, chimpanzees, bonobos, and hu-
mans, have the most advanced primate societies and the most
extensive homosexuality. In these societies, individuals form
complex relationships, fostered by both same-sex and other-
sex sexual behavior.
When confronted with evidence of homosexuality in nature,
most people immediately wonder why two males would waste
their time mating with one another rather than mating with fe-
males. The answer is that somehow they are not wasting time
that in their present circumstances, the benefits of the male
bonding outweighs the benefits of pursuing a heterosexual
courtship.
DARWIN AND SEXUAL SELECTION
The diversity of gender and sexuality in nature poses founda-
tional difficulties for biology. Biologists conceptualize gen-
der and sexuality according to Darwins The Descent of Man,
and Selection in Relation to Sex, which presents the theory of
sexual selection. Darwins theory of natural selection is Wyoming/Dakotas who lived as a woman. At right is an 1890
about evolutionary change in general, whereas the theory of
sexual selection is solely about traits pertaining to courtship
and mating. The classic example is the peacocks tail. Darwin
assumed that the peahen, the female of the species, was in a
position to choose the male she would mate with, and shes
most impressed by male with the showiest plumage. Over
many generations, males evolved the showy traits that females
prefer.
The issue we face is not whether this story ever occurs, but
JanuaryFebruary 2017
whether it is representative of how gender and sexuality are ex-
pressed throughout nature. That is to say, the preceding exam-
ples of animal species that depart from a simple sex binary, that
exhibit diverse sex roles, and that express natural homosexual-
ity all refute any claim to generality on the part of sexual-se-
lection theory. The picture of sex and gender expression
expressed in Darwins theory of sexual selection is not only far
from universal, but is perhaps exceedingly rare.
GENDER ACROSS CULTURES
The comparative anthropology of gender expression across the
world and through history is poorly documented relative to that
concerning sexuality, though the 1994 classic by Gilbert Herdt,
Third Sex, Third Gender: Beyond Sexual Dimorphism in Culture
and History, remains timely even today. That said, the available
information establishes that transgender-like and multiple gen-
der expressions are vastly more common that we in the West
have been led to believe. As recently as fifteen years ago the
medical-industrial complex was still floating the figure of
roughly one in 10,000 people as transsexual. The latest figures
show that around one in 300 is transgender in the U.S.
And as we look beyond the contemporary West, we en-
counter a huge variety of cultural forms and institutions inhab-
ited by transgender and third gender people. Among the best
studied transgender-like cultural forms are the so-called two-
spirit people of many Native American nations and tribes. The
name suggests that they are people who possess a combination
of feminine and masculine characteristics.
These people include male-bodied individuals living as
women and female-bodied individuals living as men. Here, at
left, is a 1900 photograph of Osh-Tish, a well-known male-
bodied two-spirit person from the Crow Nation of present-day
photograph of a female-bodied two-spirit person from the
Quechuan area of Northwestern South America who lived as a
man, specifically as a warrior. This warrior, dressed in male
clothing, presents in a confident, almost jaunty manner, with no
attempt to cover or bind the breasts. The middle panel is a paint-
ing of another female-bodied warrior from the Plains Indians.
Notice the bare breasts. Two-spirited people assume cross-gen-
der occupations. They do not try to hide or modify their
anatomy, because in these cultures occupation rather than gen-
23
ital morphology is the marker of gender identity.
Transgender and gay people are often accused of recruiting
young boys into their way of life. Invariably, the truth is that
the young people themselves seek out older transgender and gay
people, whom they see as kindred spirits. Anthropologist Wal-
ter Williams, in his 1986 book The Spirit and the Flesh: Sexual
Diversity in American Indian Culture, describes the ceremony
by which young boys from several Indian tribes announce their
gender and are welcomed by the tribe.
Among the Mohave tribes along the Colorado River in the
American Southwest, Williams writes about how parents deal
with a boy who has a predisposition to be two-spirited. When
the child is about ten years old, his relatives begin discussing his
predispositions. The relatives then prepare for a ceremony with-
out letting the boy know about it. The ceremony is meant to take
him by surprise, and to serve as both an initiation and a test of
his true inclinations. People from various settlements attend.
The boy is led into a circle. If he remains there, he accepts that
he will go through with the ceremony. A singer sings songs. If
the songs move him, he will dance as women do, with inten-
sity. If the boy dances as a woman for four songs, his status as
a two-spirit is confirmed. Then he is taken by the women,
bathed, receives a skirt, returns to the crowd, and announces his
new feminine name.
Among the Tohono Oodham Indians of the Sonoran Desert
of the American Southwest, the ceremony also involves a pub-
lic test of inclination. At a social gathering of family and friends,
a small brush enclosure is built. A mans bow and arrows and a
womans basket are placed inside. The boy is then brought to the
enclosure. As the adults watch, he is told to enter the enclosure.
The adults set fire to the enclosure. The boy has time to take
only one of the two items. If he takes the basket and leaves the
bow and arrows, he is confirmed as a two-spirit.
As Williams notes, in all these practices the role of a two-
spirit is not forced on the boy by others. While adults might
have their suspicions about the boys inclinations, it is only
when the boy makes the proper move that he is considered
two-spirited. By doing womans dancing, preparing a meal, or
taking the womans basket, the boy is making an important sym-
bolic gesture. Indians do not see the assumption of two-spirit
status as a free choice by the boy but rather as a revelation of his
underlying character.
Trans and gay people are too common to represent a pathol-
ogy. The existence of transgender people throughout the world
confirms that trans people comprise a significant proportion of
the human species, now and in the past. The reality, extent, and
naturalness of trans people do not need to be vouched for by
science. Unfortunately, the Western perspective on transgender
people relies on a medical construction of transgender experi-
ence and identity. That construction presumes a heterosexual
gender binary as the standard and seeks to explain variation
from that binary as a pathology.
But this medical construction is a fallacy, a scientific mis-
take that reflects an ignorance of elementary population ge-
netics. For a trait to be considered a pathology, it must be
deleterious. It has been known in population genetics for
nearly 100 years that theres a connection between how rare
a pathological trait is and how deleterious it is: the more dele-
terious, the less frequent its occurrence. For example, almost
24
lethal traits like Huntingtons disease are present at five per
100,000 births; while hemophilia A, also rare, occurs at one
birth per 8,500.
Gay and transgender people are nowhere close to being
this rare. According to the most recent 2011 demographic in-
formation from the Williams Institute at UCLA, in the U.S.
some 3.5 percent of adults identify as lesbian, gay, or bisexual,
and an estimated 0.3 percent of adults identify as transgender.
Thus around nine million Americans identify as LGBT, a fig-
ure roughly equivalent to the population of the state of New
Jersey.
The degree of rarity for pathology is set by a balance be-
tween the rate at which the pathology arises by mutation and
the rate at which it is eliminated by natural selection. This bal-
ance point is called a mutation-selection equilibrium. This table
shows the balance point between rarity and deleteriousness, as-
Frequency of Genetic Disease When Mutation
Balances Natural Selection, Relative to Severity
PERCENT REDUCTION
BIRTHS IN DARWINIAN FITNESS
1 in 10 0.001%
1 in 100 0.01%
1 in 1,000 0.1%
1 in 10,000 1%
1 in 50,000 5%
1 in 100,000 10%
1 in 1,000,000 100%
suming a standard mutation rate of healthy to pathology of one
in one million. Rarity is measured in terms of births. Deleteri-
ousness is measured in terms of the percentage loss of survival
and/or fecundity caused by the pathology (called Darwinian
fitness in population-genetic jargon).
If the pathology is lethal (bottom line of table), then the trait
is exceedingly rare, i.e., one in a million representing a fresh
mutation in each instance. For pathologies that are only slightly
deleterious, the pathology become much more common, as in-
dicated in the lines toward the top of the table. Deleteriousness
of less than, say, one percent is negligible in its impact on evo-
lution. The chance of being killed or failing to reproduce from
some random event in life is far higher than that. A small pathol-
ogy is indistinguishable from the ordinary risks in life.
So, compare the rarity of gay and transgender people with
what the table says their deleteriousness would be if they had
a pathology. For gays, their rarity lies between the top two
lines of the table. For trans people, their rarity lies between
second and third lines of the table. For both, the deleterious-
ness is effectively nonexistent, indistinguishable from the or-
dinary risks of life, given how common gay and trans people
actually are.
Consequently, the scientific validity of the medical con-
struction of how to view people who differ from the hetero-
sexual gender binary must be rejected once and for all.
Instead, a future project can be envisioned that seeks to un-
cover what the positive benefits are to the human variation in
gender and sexuality now being mistakenly pathologized in
Western culture.
The Gay & Lesbian Review / WORLDWIDE