se x is a spe ct ru m

Sex Is a Spectrum
t h e biol ogic a l l i m i ts
of t h e bi na ry

agust í n f u e n t e s

pr i nce­t on u n i v e r sit y pr e ss
pr i nce­t on & ox for d
Copyright © 2025 by Agustín Fuentes
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 c on t e n t s

Acknowl­edgments vii

Introduction: Sex Biology Is In­ter­est­ing 1

1 The Evolution of Sex 5

2 Animal Sex Biology: Mixing It Up 22

3 ­Humans Are Messy 38

4 ­Humans Then 48

5 ­Humans Now 74

6 No Biological ­Battle of the Sexes 109

7 Why the Binary View Is a Prob­lem 125

Notes 151
Index 193
 Ac k now l­e d g m e n t s

Good science and the best scholarship are collaborative. This

book is pos­si­ble due to the amazing work of incredible schol-
ars and their willingness to share insights, discoveries, methods,
and challenges. I am especially indebted to Anne Fausto-­Sterling
and Sarah Richardson for their monumental contributions and
for facilitating my and the scholarship of so many ­others on the
topic of sex. Other scholars who are central to my learning,
thinking, and working in this area include Sari Van Anders,
John Archer, Samantha Archer, Bruce Bagehemil, Rick Bri-
beiscas, Kate Clancy, Catherine Clune-­Taylor, Lucy Cooke,
Amanda Cortez, Kimberly Crenshaw, Molly Crockett, Simone
de Beauvoir, Andrea DiGiorgio, Phyllis Dolhinow, L. Zachary
DuBois, Holly Dunsworth, Lise Eliot, Cordelia Fine, Richard
Frum, Lee Gettler, Patricia Gowaty, Scott Gilbert, Matthew
Gutmann, Donna Haraway, bell hooks, Sarah Hrdy, Daphna
Joel, Rebecca Jordan-­Young, Katrina Karkazis, Craig K ­ irkpatrick,
Alexandra Kralick, Sarah Lacy, Roger Lancaster, Judith Lorber,
Katherine C. MacKinnon, Donna Maney, Jon Marks, Jim Mc­
Kenna, Stephanie Meredith, Mary Midgley, Mia Miyagi, ­Monique
Borgerhoff Mulder, Serena Nanda, Robin Nelson, Cara Ocob-
ock, Gina Rippon, Joan Roughgarden, Julienne Rutherford,
Heather Shattuck-­Heirdorn, Rick Smith, Zuleyma Tang-­
Martinez, Caroline VanSickle, Cara Wall-­Scheffler, Polly Wiess-
ner, Mary Jane West-­Eberhard, Adrienne Zihlman, Marlene

vii
 viii A c k n o w l­e d g m e n t s

Zuk, the GENDERSCI lab at Harvard, and many ­others. I

thank my amazing colleagues in the Department of Anthropol-
ogy at Prince­ton for their support and intellectual community
and all the students at UC Berkeley, Central Washington Uni-
versity, The University of Notre Dame, and Prince­ton Univer-
sity who have inspired me to think and learn.
I effusively thank Rebecca Jordan-Young and Donna Maney
for their generous, constructive, insightful, and comprehensive
reviews on ­earlier versions of the manuscript and an anonymous
reviewer for their strong critique and extensive commentary.
This book would not have been pos­si­ble without the amazing
Alison Kalett, Hallie Schaeffer, and the entire team at PUP, and
the wonderful support of Melissa Flashman at Janklow & Nesbit
Associates. Hank Southgate did a terrific copyedit. My ­family,
especially my m ­ other and ­sisters, have s­ haped my engagement
with ­these topics, and Shelley the Wonderdog offered support
throughout the writing p­ rocess. Fi­nally, and most impor-
tantly, this book and my capacity to write it results in large part
from the more than thirty years of partnership, discussions,
debates, and collaborations with Devi Snively.
Se x Is a Spe ct ru m
 Introduction
s e x biol o g y i s i n­t e r ­e s t­i ng

imagine you are a fish called the bluehead wrasse, living off
the coast of Florida. As you grow up, you, just like all to the
other bluehead wrasse your age and size, develop one set of
reproductive organs. You are what we’d call female, so you pro-
duce eggs. Th
­ ere is only one very large member of your group,
and they are the group male, so produce sperm. But over the
next few weeks you grow r­ eally fast, becoming the second-­
largest fish on your reef. Then the male gets eaten. Almost im-
mediately your body starts to change, your reproductive organs
mold, shift, and alter their form. You become the group’s sperm
producer. As a bluehead wrasse, you can have one body and one
set of DNA, but multiple forms of reproductive biology across
your lifetime.
Bluehead wrasse reproductive biology is not the most com-
mon pattern in the animal kingdom, but it’s also not that weird.
When most ­people think of the biology of reproduction, they
typically envision two fixed kinds in each species: female and
male. This is (mostly) right when it comes to the reproductive
organs themselves, but not accurate for entire bodies and lives.

1
 2 I n t r o du c t i o n

Most species do have two types of reproductive organs, and

they are often found in two slightly dif­fer­ent forms of that spe-
cies’ body plan. But not always. As with the wrasse, many fish
start out with one set of reproductive organs, and once they
grow to a certain size, they redo their anatomy and develop a
new set of reproductive organs. Each earthworm’s body has
both types of its species’ reproductive organs. Bees have two
kinds of reproductive organs across three kinds of bodies. All
mammal ­mothers lactate, but in some species of bat, ­fathers do
too. And, as in the two dif­fer­ent types of orangutan male, one
with big face flanges and the other without, t­ here can also be
quite a bit of variation in bodies and be­hav­ior even among
­those individuals within the same species that have the same
reproductive organs.
There is an explosion in research on the biology of repro-
duction—what ­we’ll call sex biology—in the animal kingdom.
While we continue to find that ­there are impor­tant differences
in reproductive biology producing female, male, and some-
times intersex bodies in any given species, ­there is also a lot of
variation, and overlap, in the a­ ctual biology and be­hav­ior that
make up ­these categories. The variation we observe across the
animal kingdom d­ oesn’t represent unusual exceptions to some
kind of rule of sex; rather, this spectrum of variation tells us that
females and males are not two dif­fer­ent kinds of ­thing. Sex biol-
ogy is not about two distinct kinds, a binary; instead, it’s about
patterns of variation in bodies, be­hav­ior, and lives that differ,
overlap, and intertwine. Sex biology, as it turns out, makes life
quite in­ter­est­ing.
The explosion in research is not l­ imited to how other animals
“do sex.” Th
­ ere is also enormous investigation into h­ uman bod-
ies, reproductive pro­cesses and patterns, health, hormones,
genitals, ge­ne­t ics, be­hav­ior, and other related topics. For
 I n t r o du c t i o n 3

example, we now know that h­ uman brains d­ on’t come in “male”

and “female” versions. Also, unlike some other mammals, all
­human caretakers (regardless of their reproductive organs) can
undergo changes in their brains, bodies, and be­hav­ior when
they take care of babies. Fascinatingly, ­human sexual be­hav­ior,
including the targets of attraction and arousal, is not necessarily
linked to what kind of reproductive organs one has. And, most
importantly, ­human sex is never just about biology; we have
gender too.
In short, t­ here is a lot g­ oing on in science regarding sex and
gender in h­ umans. Unfortunately, ­there is also a ton of misun-
derstanding in society about what biology, especially sex biol-
ogy, tells us and what it ­doesn’t tell us. And t­ here is lack of
awareness of just how diverse and variable h­ umans are. To bet-
ter understand biology and sex in ­humans, we need to learn
about our bodies, histories, cultures, and be­hav­ior. We have to
understand what it means that every­thing about h­ umans is a
supercomplicated blend of biology and culture. We need to
combine our knowledge of biology, sex, and the h­ uman experi-
ence into a new narrative. My goal in this book is to put forward
this new narrative and show how the biology of sex actually
works, what it does and does not tell us, and how we might in-
corporate this knowledge into our education, lives, and laws.
To do so, I ­w ill first summarize what is currently known
about the biology of sex in animals and how, and why, that re-
lates to ­humans. This is impor­tant ­because understanding ani-
mal biology is at the heart of understanding ­human biology (we
are, ­after all, animals). From ­there, I ­will illustrate what we know
about sex, in biology and be­hav­ior, in the ­human past and pre­
sent, across the last two million years of our lineage’s existence
and among the eight billion ­humans living ­today. What this
knowledge from the animal world, the h­ uman past, and the
 4 I n t r o du c t i o n

h­ uman pre­sent shows us is that biology as it relates to sex is not

binary, meaning that it does not come in two distinct kinds:
male and female. This is not to say that females and males are
the same. They a­ ren’t. Nor is it that biological variation related
to sex does not ­matter. It does. It’s just that not all ­humans fit
neatly into the categories of female or male, and biological
­measures of ­human bodies rarely segregate into two non-­
overlapping categories. Neither “female” nor “male” describes
a uniform or distinct biological type.
I ­w ill conclude by discussing why a binary view is a detri-
mental way to think, and talk, about sex biology and the h­ uman
experience. Reproductive biology is an impor­tant structuring
part of h­ uman lives; however, producing ova or sperm, having
XX or XY chromosomes, or having a clitoris or a penis, does
not tell us nearly as much biologically as many believe. Nor
does it consistently or accurately inform us about an individu-
al’s childcare capacity, homemaking tendencies, interest in lit­
er­a­ture, engineering and math capabilities, or tendencies
­toward gossip, vio­lence, compassion, or a love of sports. By
contrast, placing reproductive biology in the context of the rest
of the body, and in relation to be­hav­ior, history, society, and
experience, we are much better prepared to ask, and answer,
questions about health, habits, proclivities, happiness, and the
many ways to successfully be ­human.
However, at its core, biology is about evolution, and evolu-
tionarily speaking ­there is a lot of variation in sex biology and
be­hav­ior, both across and within species. So, to r­ eally under-
stand how biology and sex work in h­ umans, we need to start not
with us ­today but right back near the start of life on earth, with
the evolution of sex.
 1

The Evolution of Sex

in the beginning . . . ​­There was no sex. The earliest life forms

on earth reproduced asexually, internally copying their key bio-
logical material and then dividing into two versions of them-
selves.1 ­These organisms ­were, and remain, the prokaryotes, that
is, microscopic single-­celled organisms most ­people call bacteria
(which scientists divide into archaea and bacteria)2 and blue-­
green algae (cyanobacteria). Then, somewhere around two bil-
lion years ago, evolutionary changes resulted in what we call
“protoeukaryotes” that evolved into the common ancestor of
all eukaryotic organisms, including animals and plants. Eukary-
otes have DNA in the form of chromosomes contained within a
nucleus. Eukaryotes mostly reproduce via a ­process of combin-
ing ge­ne­tic information from two dif­fer­ent individuals to create
a new individual, or, as that ­process is commonly called: sex.
The p­ rocess by which sex evolved is complex but basically
involved restructuring of the ways in which DNA is packaged
and copied, the internal structuring of the cell’s physiology, and
the ways in which cells divide and fuse. A main challenge for sex
is the ability to create a copy of one’s ge­ne­tic material and pack-
age it so that it can meet up and fuse with another of your same
kind of organism’s ge­ne­tic material and create a new organism.

5
 6 Chapter 1

Prokaryotes could already copy their ge­ne­tic material and di-

vide. The trick was to keep the system of copying ge­ne­tic mate-
rial, but not divide into two new organisms, and create a ­little
package of that material that could be combined with other
such packages (called gametes) from other individuals.
Over hundreds of millions of years, single-­celled eukaryotes
developed the innovation of “mating types”—­dif­fer­ent forms of
the same organism that could produce slightly dif­fer­ent gametes.
Once two dif­fer­ent mating types got their gametes to fuse to-
gether, they created a new organism with two copies of all ge­ne­tic
material. Each organism produced via the fusion of gametes from
two mating types is a ­little genet­ically dif­fer­ent from each of the
parents due to the mixing and matching of ge­ne­tic sequences that
happens during the gamete production and fusion pro­cesses.
From the very start of sex, variation was the name of the
game. Even in their earliest appearances, the number of mating
types (often called “sexes”) per kind of species has been vari-
able, ranging from two and sometimes three in most animals,
to as many as seven in single-­celled organisms, thirteen in slime
molds, and thousands in some fungi. Sex across its evolutionary
history is quite variable.3

Why Sex?
On the face of it, sex is a paradox. W
­ ouldn’t life be easier if or-
ganisms just kept copying themselves to reproduce? Why add
extra challenges and potential prob­lems? In 1975, a classic book
on sex and evolution began with the comment, “This book is
written from a conviction that the prevalence of sexual repro-
duction in higher plants and animals is inconsistent with evo-
lutionary theory.”4 If sexual reproduction is more difficult and
complicated than asexual reproduction, why did it evolve?
 T h e E volu t ion of Se x 7

The favored hypothesis is that sex is a response to dealing

with changing environments. Sex is a biological way to generate
more variation for organisms such that they have better chances
of meeting challenges the world throws at them. But sex a risky
venture.5
­Here is a simplified “sex is better than no sex” scenario . . .
Imagine a single-­celled organism living in a pond, an amoeba-­
like t­ hing that filters ­water for food. It might do just fine asexually
copying itself if the ­water temperature stays more or less con-
stant. But what if the ­water temperature shifts? The organism
might not be able to deal with the new temperature. However,
­there could be a multitude of t­ hese organisms in the pond, each
slightly dif­fer­ent in its ability to adjust to fluctuating tempera-
tures. Blending ge­ne­tic material with another similar but distinct
member of one’s species could be a good option ­because it can
offer greater flexibility and thereby a higher chance of survival for
the offspring. But not all new ge­ne­tic combinations created by
sex do better. In fact, some do worse. Such is the risk of sex. It’s
the overall payoff that m ­ atters: if some sexually produced off-
spring do better, on average, compared to t­ hose of asexual repro-
ducers, the system (sex) has a chance of catching on. And lucky
for us it generally does. Other­wise, all life on earth would be
asexual, and the world would be a lot less in­ter­est­ing.

Dif­f er­ent Gametes = Biological Sex Differences?

Once sexual reproduction (the fusion of gametes) became pos­
si­ble, t­ here was a quick transition from isogamy (making same-­
sized gametes by all individuals in a species) to anisogamy
(making different-­sized gametes by dif­fer­ent individuals in a
species) in many (but not all) kinds of life. The exact details of
the emergence of anisogamy are not fully known or agreed on,6
 8 Chapter 1

but it’s coincident with the evolution of more complex forms

of multicellular life over the last billion years. ­Today, having two
dif­fer­ent types of gametes is the norm among most sexually
reproducing beings.7
Anisogamy usually manifests as two types of gametes: one
large and one small. The larger gametes are called ova (colloqui-
ally known as “eggs”) and the smaller gametes are sperm. Both
types are haploid cells, meaning they each carry one copy of each
parental chromosome, or 50 ­percent of the full complement of
an organism’s DNA (remember: all organisms produced by sex
have two copies of the ge­ne­tic material, one from each parent).
However, ova also have the full range of cellular machinery and
large assemblage of proteins and enzymes, while sperm only
have a few additional proteins and enzymes accompanying the
DNA. On average, each individual ovum is more energetically
costly to produce than each individual sperm, due to the size and
contents of each. However, it is not accurate to compare one
ovum to one sperm in the sense of costs of production b­ ecause
in most sexual reproductive systems (especially mammals),
many, many, many more sperm need to be produced than ova.
For example, in h­ umans, the correct comparison is 1 ovum = ​
~15 million sperm, as that’s the number of sperm per ejaculate nec-
essary for effective chances of a successful sperm-­ovum fusion.
This difference in size and contents of gametes became a cen-
tral focus in the nineteenth and twentieth centuries, when re-
searchers began to hypothesize downstream differences in the
mating types, or sexes, that produced them. One might think
that it was the study of gametes that led the researchers to de-
velop the hypotheses that gametes are the key to sex difference,
but it was the other way around: belief that ­there had to be a key
difference between the sexes—­evidence of a biological binary—
led to the focus on gametes.
 T h e E volu t ion of Se x 9

The idea of biologically distinct sexes only became common

in the eigh­teenth ­century. From the early Greek ideas about sex
of Aristotle and Hippocrates and the Roman anatomical ideas
of Galen through the ­Renaissance, and into the start of the eigh­
teenth ­century, science and the medical world did not consider
females and males as two separate kinds of biologies or beings.
Rather, they w ­ ere seen as hierarchically ranked (male above
female) versions of the ­human form. This was termed the “one
sex model.”8 But from the eigh­teenth into the nineteenth
­century, the belief in a “two sex” model, with males and females
reflecting dif­fer­ent biologies, emerged, but it had no specific
definitional focus. It is from this newer two-­sex worldview that
the hypothesis of differences in gamete size as the key to female
and male distinction emerged.9 The general assumption, which
became central for many biologists in the twentieth ­century,
was that the two gamete-­producing versions (sexes) of a given
species have substantive biological differences in the p­ roduction
of their gametes and in the physiology and be­hav­ior associated
with reproduction. Given t­ hese assumed biological differences,
it stood to reason that each gamete-­producing version (each
sex) would be ­under distinct evolutionary pressures ­because of
biological differences.
In 1871, Charles Darwin proposed the central evolutionary
argument for differences between the sexes—­that ­there are
evolved differences in males and females and that they are based
on dif­fer­ent investment in reproduction. This argument was
focused on anisogamy (different-­sized gametes) by the biolo-
gist Angus Bateman in the late 1940s. Darwin10 argued that
evolutionary pro­cesses, called natu­ral and sexual se­lection,
made males and females very dif­fer­ent, and Bateman11 argued
that the primary cause of this was “that females produce much
fewer gametes than males” and that anisogamy would
 10 Chapter 1

ultimately result in “an indiscriminatory eagerness in the males

and a discriminatory passivity in the females,” with “greater de-
pendence of males for their fertility on frequency of insemina-
tions.” Basically, Bateman argued that cheap sperm and costly
ova drive evolutionary pro­cesses, making males and females
very dif­f er­ent.12 He developed his arguments by looking at flies
where he thought males’ energetically inexpensive sperm w ­ ere
their only “cost” for reproduction, and thus reproducing for
them consisted of trying to get as much sperm into as many as
females as pos­si­ble. On the other hand, he saw the ova pro-
duced by females as biologically very costly, such that females
­were l­imited in how much they could reproduce and had to
carefully consider each mating b­ ecause they could only have a
­limited amount relative to males. Bateman’s argument (cheap
sperm and costly ova make males and females very dif­fer­ent
organisms) became baseline theory for evolutionary biology
around 1966 and centralized in the world of assumed “biological
fact” by another biologist, R. L. Trivers, in 1972.13 Trivers con-
nected anisogamy to parental care via a ­simple mathematical
equation. Bateman’s notions of sex differences entered near
universal “truth” in biology with the work of E. O. Wilson in
1975 and G. A. Parker in 1979.14
As it turns out, Bateman was mostly wrong.
The prob­lem came down to a counting error. Bateman made
key errors in how he counted mating success for males and mis-
represented the fruit fly mating system he was studying. More
than thirty years of reexamination of Bateman’s experiments,
continued work on his model subjects (fruit flies), and in-­depth
study of sexually reproducing species demonstrate that
­acceptance of Bateman’s paradigm as a “law” of biology led to
two key prob­lems. First, it led to a simplification of Bateman’s
data, resulting in an incomplete and biased understanding of
 T h e E volu t ion of Se x 11

his original findings. Second, ­because of the oversimplification

of how reproductive biology works, it hampered scientists’
­ability to interpret real­ity with regard to male and female sexual
be­hav­ior.15
This is a serious hitch in the science of reproduction. Bate-
man’s early studies, and the authors that subsequently en-
shrined them into biological canon, have led to widespread
confusion, mistaken assertions, and a deep-­seated belief in an-
isogamy as the core explanation of every­thing about sex. That
is not to say t­ here a­ ren’t patterns of differential stressors and
challenges across dif­fer­ent reproductive systems. ­There are.
Evolutionary histories have ­shaped sex biology into a range of
typical patterns for ova and sperm producers, and we can learn
a lot from the study of ­those patterns. However, while the gam-
etes themselves mostly come in two types (a binary), most of
the rest of the biology in organisms does not. B ­ ecause they rely
on the simplistic, and erroneous, assumptions about anisog-
amy, many of the classic assertions about why differences in sex
biology emerge, what ­those differences are, and how we should
think about sex, are incomplete and sometimes flat-­o ut
wrong.16 ­We’ll deal with this topic in greater detail in the fol-
lowing chapters. But before diving into the ­giant debate about
sex biology and h­ umans, we need to cover some basics, b­ ecause
reproductive biology is more than just gamete making.

Getting Gametes Together

The goal for any sexually reproductive system is to develop a
method for ensuring that gametes get together, fuse, and pre-
pare to develop into adult organisms. Not surprisingly, life on
earth has developed a huge range of ways to get this done. Evo-
lution and biology are anything but s­ imple or uniform. While
 12 Chapter 1

individuals in many species are large-­or small-­gamete produc-

ers for much or all of their lives, t­ here are some species of animals
where all members are si­mul­ta­neously large-­and small-­gamete
producers, and a number of species where individuals switch
back and forth between being large-­and small-­gamete producers
across their lifetimes. For most biologists, the term “female” de-
scribes an organism when it is a large-­gamete producer, the term
“male” when it is a small-­gamete producer, and the term “inter-
sex” is used for individuals that produce both types of gamete at
the same time.17 When this naming system is used across dif­fer­
ent species, some organisms would be called male or female or
intersex their ­whole lives, but for ­others, the terms would switch
depending on the life stage they are in. Be warned, however,
­because the biology and be­hav­ior of organisms sharing the terms
“male,” “female,” and “intersex” are rarely identical within each
category across dif­fer­ent species. Take, for example, the follow-
ing four female mammals: a queen mole rat, a dominant hyena,
a doe kangaroo, and a m ­ other cotton-­top tamarin monkey. Their
biology, be­hav­ior, and life experience, while sharing a few ­things
in common, are all radically dif­fer­ent from one another. In fact,
the patterns and experiences of reproductive biology are not
even necessarily the same even within individuals in a single spe-
cies with the same reproductive organs, such as between a queen
mole rat and a regular female mole rat.
­Things are ­really complicated in reproductive biology,
and the rest of the body, beyond the gametes, but an overview of
the typical patterns in systems for gamete fusion is a necessary
starting point in understanding sex biology. In the most basic
reproductive systems, individuals produce gametes and eject
them into the local environment (usually w ­ ater), ideally in
proximity to another member of their species. And they leave
it to the gametes to find each other and fuse, creating a zygote.
 T h e E volu t ion of Se x 13

This is called “spawning” and is common in aquatic species,

including many fish, jellyfish, crabs, starfish, sea cucumbers,
aquatic insects, and even some amphibians.
The next level of reproductive complexity involves one
gamete-­type producer (usually the large-­gamete producer) cre-
ating some form of gel or other gooey/sticky matrix to deposit
the ova in a cluster, which is often then attached to some object
in the environment, like a plant or a rock in a pond, river, or
ocean. Sometimes members of a given species carry the large
sticky masses of ova on their bodies. Individuals who can pro-
duce small gametes approach the cluster of ova and eject sperm
onto/into it, and thus fusion can take place, creating a mass of
zygotes in the gooey matrix. A lot of amphibians, like salaman-
ders and frogs, and many fish reproduce this way. Stepping up
to the next level of complexity (physiologically at least) are sys-
tems wherein one sex develops an internal physiology that takes
in and stores a cluster of already fused gametes (zygotes). Usu-
ally, this internal storing of the zygotes ­until they are ready to
move on their own is done by large-­gamete producers, but in a
range of species, such as sea­horses, it is the small-­gamete pro-
ducers that internally store and ­later “birth” the zygotes.
It is with internal fusion of the gametes that t­ hings get a bit
trickier, as the small gametes need to get into the body of the
large-­gamete producer, reach the ova, and then the products of
the fused gametes need to be excreted into the world at some
­later stage of development. ­There are more than a few twists to
internal fusion. The first step is to get the gametes together in
the body of the large-­gamete producer, which usually involves
some form of copulation (see below), and then t­ here are a few
variants on internal dynamics of the system. One way to deal
with internal fusion and subsequent development is the pro-
duction of a nutrient-­rich, protective, solid, organic container,
 14 Chapter 1

called an “egg.” The development of the egg is catalyzed by the

gamete fusion p­ rocess and the contributions of additional phys-
iological structures in the large-­gamete producer who then ex-
cretes it. The zygote then develops inside the organic container
but outside the body of the large-­gamete producer.18 The or-
ganic container (egg) system is found in many animals, from
insects to reptiles to birds and even a few mammals (like the
duck-­billed platypus). Eggs are fantastically in­ter­est­ing struc-
tures, and the internal physiology associated with them is var-
ied but has some core consistencies (described below).
The final type of internal gamete fusion and development
system is the more evolutionarily recent, and highly ­complicated,
mammalian innovation of gestation and lactation. Basically,
in this system the large-­gamete producer develops a specific
internal physiology (a uterus) that enables it to act as a host for
a ­process wherein a small number of large gametes are selectively
released into a section of the internal biology. Then behavioral
and physiological actions (copulating) requiring specialized
organs (genitals19) occur between small-­and large-­gamete
­producers, and small gametes are introduced into the large-­
gamete producer’s internal system. Then, if gamete fusion oc-
curs, the resultant zygote implants inside the tissues of the
large-­gamete producer’s uterus and develops into an embryo
and then a fetus.
This is a relatively long-­term developmental p­ rocess that
­involves shared physiological connections between the devel-
oping zygote/embryo/fetus and the host large-­gamete pro-
ducer, with the embryo/fetus wholly dependent on, and part
of, the host’s body. At a certain point in development, the host
excretes (births) the developed fetus (an infant), and some
form of caretaking involving high-­quality nutrition (lactation)
ensues. Frequently, the host and/or other members of the
 T h e E volu t ion of Se x 15

species help the developing young individual for an extended

period of time. This par­tic­u­lar system sets up some expanded
physiological obligations for large-­gamete producers and
throws in a w­ hole new set of biological and behavioral com-
plexities, and constraints, for the reproductive ­process.

Let’s Talk about Gonads

The basic unit of reproductive biology is the gonad.20 We all
have them. This nifty ­little organ produces gametes. Gonads
that produce large gametes (ova) are called ovaries, and ­those that
produce the small gametes (sperm) are testes. In most species
of animal, a majority of individuals usually have one of the two
gonad types (ovaries or testes) as adults. However, ­there are a
number of species in which all individuals shift from one gonad
type to the other across their lifetimes, o­ thers that have both
gonad types as adults, and t­ here are also a number of species
(largely invertebrates) that have single gonads (called ovotes-
tes) that produce both types of gamete. Th­ ere is also significant
variation within species, including ­humans, where a small
but not insignificant percentage of individuals have some mix
of the above (an untypical gonad, two gonad types in the same
body, or even ovotestes). Most gonads also have associated duct
and tube systems for the storage and excretion of the gametes,
a significant/crucial component of structuring a species’ sex
biology.
In vertebrates, gonads also produce a range of hormones
­related to the growth and functioning of reproductive tracts, ac-
cessory sex glands, and organs related to copulation. ­These hor-
mones contribute to a range of behavioral actions associated
with copulation and other facets of development and function-
ing in organisms’ bodies. Testes and ovaries emerge from the
 16 Chapter 1

same embryonic tissues, but they vary in certain aspects of their

mature physiology. All types of gonads produce the same
­hormones, but they often do so in differing levels. Therefore,
individual organisms with e­ ither testes or ovaries, individuals
having both, and t­ hose with ovotestes differ with re­spect to
levels of some hormones (but not kinds) between them.

Details of Internal Fusion Systems

For animals in which gamete fusion is internal t­ here are a vari-
ety of additional organs and physiological systems that make up
key parts of sex biology. First, t­ here are copulatory organs
whose role is to facilitate the excretion and/or absorption of
gametes. At a basic level, t­ hese are a variety of bodily modifica-
tions for clasping during copulation and ­simple openings con-
nected to gonad duct systems. At a more complex level, they
include invagination of body surfaces and cavities, external
genitals (such as a clitoris and penis), and internal structures
(such as a vagina) that interact to facilitate getting the large and
small gametes into interaction with one another.
As for the core layout of reproductive organs, most verte-
brates have a cloaca, a common opening/chamber for waste
excretion from the urinary and digestive tracts that also serves
as the exit point for sperm and/or ova. In most birds and rep-
tiles, the cloaca is divided into a urodeum for the urine, a co-
prodeum for fecal ­matter, and the proctodeum, which is the
final chamber next to the cloacal opening associated with copu-
latory organs and the excretion of sperm and eggs (not ova but
­actual eggs).
In vertebrates that lay eggs, ­there is specific differentiation in
internal sex biology between large-­and small-­gamete produc-
ers. For example, egg-­producing reproductive tracts develop
 T h e E volu t ion of Se x 17

into a system whereby the ovaries produce ova that are trans-
ported to an oviduct, a long tube with multiple cell types across
dif­fer­ent parts of it. As the ova passes through the tube, they
fuse with sperm (if pre­sent), acquire vari­ous proteins and ma-
terials that make up the yolk and white of the egg, and then stop
in the shell gland. In the shell gland, the internal materials of the
egg finalize, and the hard calcium-­rich exterior develops around
to contain them, ­after which the egg is excreted via the cloaca.
Mammals have even more complicated reproductive physi-
ology. Most have some form of external genitalia derived from
the embryonic tissues called the genital tubercle. In ­humans,
typically, this tissue mass develops into a penis or clitoris, with
a wide range of variation in the adult endpoints (see chapter 5).
Associated areas of tissue (the labioscrotal/urogenital fold) de-
velop into the scrotum or the labia majora, and other tissues of
the urogenital folds develop into the skin of the penis and the
internal labia (if pre­sent). Many mammals have internal testes,
but several of them (including ­humans) have an external scrotal
sack and testes that reside in it. All ovaries are internal.
A core difference between mammals and other animals is
that mammals gestate. Zygotes develop inside the uterus into
embryos and then fetuses.21 To accomplish this, mammals
evolved a complex system involving connection from the ova-
ries to a fallopian tube and into the uterus. In one lineage of
mammals, including us, the uterus develops a special organ
called the placenta plugging the zygote directly into the ­maternal
physiology and providing nutrients and removing waste. This
intense placental linkage between maternal and fetal bodies is
distinctive in the animal kingdom. It creates a suite of patterns
that structure the bodies of individuals with this reproductive
anatomy relative to individuals who do not have it.22 At the end
of gestation, one or more fetuses are birthed needing to be fed
 18 Chapter 1

a high-­quality nutrient (milk) u­ ntil they are ready to fend for

themselves food-­wise. To accomplish this, mammals develop
mammary glands that produce high-­quality milk during lacta-
tion. The ovary-­fallopian tube-­uterus physiology and mam-
mary glands are typical of large-­gamete-­producing mammalian
bodies, but not small-­gamete-­producing ones.

Caring for Infants ­Matters

Animal bodies are ­shaped by sex biology, and in many cases,
their be­hav­ior is too. While reproduction for many animals ends
with the excretion of gametes, ­there are ­others for whom care of
the embryos, fetuses, and postbirth young is a central part of the
reproductive cycle and shapes the animals’ ways of living.
On the simplest end of the caretaking spectrum, in many
species of fish and amphibian, adults ­w ill guard masses of
­z ygotes/eggs and carry and protect masses of embryos
(“brood” them) in their mouths, on their backs, or in invagina-
tions or pouches in their body ­until the embryos develop into
young capable of their own movement and feeding. Many rep-
tiles stay with laid eggs and protect them ­until they hatch, with
some snakes and lizards carry­ing the eggs internally, and then
give “birth” to the hatchlings. Some reptiles, like crocodilians,
even care for the young for a period a­ fter birth. Birds almost
always care extensively for eggs and thereafter the hatched
young. Bird parents provide warmth and protection to the
eggs, then care and food for the developing hatchlings, and
ultimately some level of training for adult life. And, of course,
mammals take the cake in terms of pre-­and postnatal care.
Some lineages, such as primates, elephants, cetaceans (­whales
and dolphins), and many social carnivores, intensively care for
young for years ­after birth.
 T h e E volu t ion of Se x 19

For most of the nineteenth and twentieth centuries, biolo-

gists mostly assumed that b­ ecause of the supposed implications
of anisogamy, the large-­gamete producers (“females”) evolved
as the primary caregivers and thus their bodies, and brains,
­were “designed” for parenting. In contrast, small-­gamete pro-
ducers (“males”) ­were considered to not be ­under such ­pressures
and would thus be ­shaped by evolution to maximize getting
their sperm to the ova in order to enable the fusion of gametes,
regardless of what happened postfusion. This simplistic view of
anisogamy is wrong, and so ­were many of the nineteenth-­and
twentieth-­century biologists’ assumptions both about the limi-
tations on “females”23 and about parental care and its evolution.
To be fair, from very early on, it was clear to many scientists that
­there was indeed variation in parenting and other reproductive
patterns, and sex-­related biology, across the animal kingdom,
but ­these variations ­were often seen as exceptions and abnor-
malities, and studied as such. However, as ­we’ll see throughout
this book, what ­were previously thought of as exceptions or
minority patterns are more often than not a core part of the
­actual range of biological variation in and between species, and
seeing and studying them as part of an expected range offers a
better understanding of biology and be­hav­ior.
Clearly, t­ here are some biological patterns that structure
large-­gamete-­producing bodies differently than small-­gamete-­
producing bodies, especially in t­ hose species that lay eggs or
gestate and lactate. However, ­these differences in reproductive
anatomy do not predict the entirety of the animal’s biology,
be­hav­ior, and ecol­ogy. In fact, when it comes to care, t­ here is
much variation, and the specifics of that variation are central to
structuring how any given sex-­biology system works. The as-
sumption that large-­gamete producers are uniformly destined
to be the caretakers of young is not true.24 In fish, “males” offer
 20 Chapter 1

care more often than “females.”25 Among amphibians, ­there is

a wide array of caretaking patterns across all gamete-­producing
types,26 and with birds, both gamete-­producing types are joint
primary caretakers in nest brooding and posthatching care in
81 ­percent of all species.27 When it comes to mammals, repro-
ductive biology dictates a specific postbirth role of lactation for
large-­gamete producers, but regarding all other forms of active
caretaking of infants and juveniles, t­ here is a range of variation
from small-­gamete producers who do 90 ­percent of the infant
care, to fully cooperative care by an entire group, to exclusive
maternal care. And this range of dif­fer­ent patterns generates a
diversity of mammalian sex biology and be­hav­ior between,
across, and within gamete-­producing categories.28

­There Are Many Ways to Do Sex Biology

At its core, the fusion of gametes forms a zygote, then an em-
bryo, then a fetus, and eventually an adult organism. Thus, any
biological system dealing with sex must be prepared to respond
to three core challenges. The first is to make gametes, the sec-
ond is to get ­those gametes together, and, fi­nally, the last is to
provide a physiologically and ecologically ­viable environment
for a zygote to develop and mature.
The first two challenges create specific biological characteris-
tics of organisms: gamete-­production physiology and gamete-­
placement biology and be­hav­ior. The third challenge, finding a
­v iable environment, ranges from simply finding a spot in the
­water of the right temperature and excreting gametes, to internal
gamete fusion and laying eggs, to a range of internal gestation
strategies, to complex parenting be­hav­ior. Clearly, this varies by
type of organism. For many, some form of investment and/or
care beyond gamete fusion is needed.
 T h e E volu t ion of Se x 21

When parental care is needed, a new set of challenges

emerges. Th ­ ere must be a physiology sufficient to facilitate the
development of the zygote into an embryo, and then into a
young organism. Then be­hav­ior sufficient to facilitate care for
the development of the young organism(s) is usually required,
sometimes for many years a­ fter birth. Fi­nally, animals that have
the physiology and be­hav­ior to care for young also need a
­socioecological system (a way of being socially) and set of re-
sponses to ecological challenges in which their par­t ic­u­lar
physiology and be­hav­ior can be effective. It is in this physiology
and be­hav­ior of parenting that the dynamic variation in sex bi-
ology becomes most apparent and most in­ter­est­ing for a con-
versation that is eventually about ­humans. The ways in which
organisms interact with their environments and each other are
impor­tant in understanding sex biology.29
Surveying the range of animal sex biology, a group of biolo-
gists recently summarized their findings as follows: “ ‘Sex’ com-
prises multiple traits. . . . ​Individuals may possess dif­fer­ent
combinations of chromosome type, gamete size, hormone
level, morphology, and social roles, which do not always align
in female and male specific ways or persist across an organism’s
lifespan. Reliance on strict binary categories of sex fails to
accurately capture the diverse and nuanced nature of sex.”30
In short, ­there are many, many ways to “do” sex biology ef-
fectively. The next chapters lay this out for other animals and then
for ­humans.
 2

Animal Sex Biology

m i x i ng i t u p

­e very textbook lays out how animals are supposed to “do”

sex biology. But most animals ­don’t read textbooks.
Sex biology is often misunderstood as a story of two kinds of
creatures, the male and the female. For example, many assume
male mammals are larger than female mammals. But ­there are
multiple examples of large-­gamete-­producing individuals that are
the same size as—or larger than—­small-­gamete producers.1
And in many species, including ­humans, ­there is huge variation,
and overlap, in size across populations and between individuals
within the large-­and small-­gamete-­producing categories. Most
­people also incorrectly assume that female animals are less ag-
gressive, provide most or all care of young, and are choosier in
mating than males, when counterexamples exist among species
of fish, birds, lizards, and mammals.2 The assumption of fixed
“sex” differences and “sex” uniformity e­ ither between or within
species is a major stumbling block to understanding biology
and be­hav­ior in organisms.
Science is supposed to constantly adapt to new data and
analyses. The first periodic ­table of ele­ments had 63 entries (in

22
 A n i m a l Se x Biol ogy 23

1869); ­today, we have 118 due to discoveries of new ele­ments. A

­century ago, tool use was seen as the near-­exclusive domain of
­humans, but the study of apes, monkeys, birds, elephants, and
many other animals revealed tool use across much of the animal
kingdom. The longer a system, a species, or a group of animals
is studied, the more fine-­tuned our understanding becomes.
Of course, t­ here are patterns and trends associated with re-
productive systems that are common across much of life or in
specific lineages. But ­there are also numerous variations and
modifications in the patterns of sex biology in animals. In some
lineages, and sometimes within the same gamete-­producing
categories, many previously considered “aty­pi­cal” variations are
as common as the expected patterns. In o­ thers they are not.
Identifying what is “typical” or common and assuming that this
identification represents sufficient understanding of a species
or system is a scientific error. Ignoring biological variation, even
if small, is bad science ­because the goal of biology is under-
standing how bodies are made and work: how all the parts
­relate to one another. So, ignoring aspects of some parts and the
variation of ­those parts blinds us to key data needed to do
the best science. Au­then­tic understanding of the world grows
as more of the range of what is ­really out ­there is added in.
Large-­and small-­gamete producers can and do experience
dif­fer­ent evolutionary pressures resulting in some sexual se­
lection for sex-­biology “roles.” But any one pattern, even if it’s
very common, is seldom ubiquitous, and the ­actual biological
outcomes—­what the “roles” and bodies look like and do—­vary
substantially across dif­fer­ent species and lineages.3 In mam-
mals, the reproductive physiology associated with gestation
and lactation shapes the bodies of t­ hose who gestate and lactate
relative to the bodies of t­ hose who do not. However, even in
mammals with this very specific set of physiological constraints,
 24 Chapter 2

how this system translates to the specifics of body size and

shape, hormone patterns, care of young, and sexual be­hav­ior in
large-­gamete producers alone can vary dramatically from opos-
sums to rats to hyenas to dolphins to monkeys to elephants to
­humans.4 So, while ­there are general sex-­biology patterns that
are common, even typical, it is in examining the variations
across biologies, within and between kinds of animal, that we
see ­there is not one “best” or “right” or single way to characterize
sex biology in the animal kingdom.
Let’s take a brief tour.
The Hymenoptera (bees, ants, and wasps) are haplodiploid,
meaning they have a ge­ne­tic system in which some individuals
develop from fused (diploid) zygotes and o­ thers from unfused
(haploid) sex cells. But this is not a binary “male/female” sys-
tem. In bees, the system of sex biology includes two types of
ge­ne­tic systems that produce three types of bodies: one small-­
gamete producer and two large-­gamete producing physiologies.
But one of the large-­gamete types does not actually produce
any gametes (usually) or engage in sexual activity, while the
other one can and does both.5 So instead of simply labeling
both types “female,” we call the first type “workers” and the
second type a “queen.” The small-­gamete producers are called
“drones.” The queen is much larger than the other two body
types and can control the pattern of gametic fusion in her own
reproductive tract to manage the ratios of the three body types
in subsequent generations. The behavioral, ecological, and
physiological lives of bees involve three sex-­biology types.
In the Hemiptera (the official name for about eight thousand
va­ri­e­ties of wingless, sucking insects commonly called scale in-
sects), all zygotes are diploid, but in order to develop into a small-­
gamete producer, the zygote must undergo a “turning off ” of
chromosomes or lose certain chromosomes right ­after gamete
 A n i m a l Se x Biol ogy 25

fusion. In other insect lineages, all offspring of any given individ-

ual ­will be only large-­gamete or only small-­gamete producers. Yet
in ­others, which reproductive physiology (small-­or large-­gamete
producing) ­w ill develop is entirely temperature-­dependent.
Parthenogenesis—­asexual reproduction by which large gametes
develop into zygotes without any need for small gametes—is also
found in a minority of insects. Strangely, while bodies with both
small-­and large-­gamete-­producing physiology are fairly common
in many invertebrate animals, they are extremely rare in insects.6
Small-­and large-­gamete producers’ bodies, be­hav­iors, ecologies,
and physiologies are not uniform across the insect world; t­ here is
no one universal reproductive biology, or one consistent way to
become or to be “female” or “male” in insects.
Worms, too, have fascinating sex biologies. Earthworms, for
example, are all intersex. As adults, each individual has fully
developed and functioning testes and ovaries along with a
­series of organs and glands that act to bring large and small
gametes together, e­ ither in the individual’s own body or via a
mutual gamete exchange with another earthworm. Addition-
ally, certain species are also capable of parthenogenesis (asexual
reproduction with no gamete fusion at all).7 The sex biology of
earthworms enables a variety of sex “roles” but is not character-
izable as a binary. For example, in some species, the same worm
can have “self-­sex” between the two reproductive physiologies
in their bodies, have sex with another individual involving the
mutual exchange of both large and small gametes, and undergo
parthenogenesis. That’s pretty much the ­whole range of repro-
ductive options for living ­things in one individual. Alongside
this dynamic in sex “roles,” the development of such worm sex
biology also involves pretty complex sex-­determination sys-
tems. Most worms have polyploidy,8 with three or as many as
twelve (!) sets of chromosomes related to sex biology.
 26 Chapter 2

Other types of worms have dif­fer­ent variants of sex biology.

The roundworm C. elegans has only two sex-­biology types, but
they are not the typical pattern of a small-­gamete producer and
a large-­gamete producer. In C. elegans, individuals usually ­either
produce both gamete types or just small gametes. The large-­
gamete-­only sex biology rarely shows up. However, individual
C. elegans can flexibly alter temporarily, or permanently, the
specifics of their reproductive system (that is, go from one “mode”
of gamete-­producing physiology to another) in the face of en-
vironmental challenges.9
Organisms with backbones offer yet a new set of variations
to consider. Take teleost fish, which are the most common type
of bony fish. They have a sex biology that is flexible, with major
changes in reproductive physiology within the lifespan of indi-
viduals being common. Many fish are “sequentially intersex,”
meaning individuals start out with one set of sex biology and
then change one or more times to dif­fer­ent sets of sex biology
across their lives.10 Such changes involve transformations, as
adults, of multiple biological systems, and include behavioral,
anatomical, physiological, neuroendocrine, and molecular pro­
cesses.11 Fascinatingly, the bluehead wrasse usually develops
into an adult as a large-­gamete producer and then might shift to
a small-­gamete producer if their body size grows very large. But
this usually only happens when the largest bluehead wrasse in
a small area of a reef dies. When this happens, the next largest
wrasse changes their sex biology and shifts from large-­to small-­
gamete-­producing physiology and takes over as the sole small-­
gamete producer in that specific area of the reef. However, the
system is not without its variations. Sometimes a small, youn­
ger wrasse does not mature into a large-­gamete producer but
has a body that looks just like one, even though it is producing
small gametes. ­These individuals are called “female-­mimics”
 A n i m a l Se x Biol ogy 27

and covertly mate with other small-­bodied bluehead wrasses

without the large local “male” noticing them.12 The bluehead
wrasse has three sex-­biology types, and any individual in the
species may or may not “be” one, two, or all three of them dur-
ing their lifetimes.
Other fish species switch in the opposite direction, starting
out as a small-­gamete producer and switching to a large-­gamete
producer only when they become much larger bodied. Many
anemonefish (“Nemo” from the Disney movie, for example)
live in groups of two dominant reproducing adults and a collec-
tion of nonbreeding subordinate individuals. The larger bodied
of the breeding pair produces large gametes and the smaller one
makes small gametes. But, if the larger one dies, then the smaller
member of the breeding pair undergoes a sex-­biology change
to become a large-­gamete producer, and one of the nonbreed-
ing members w ­ ill step in to take over as the small-­bodied “male”
of the breeding pair.
All California sheepshead fish change from large-­to small-­
gamete-­producing physiology once they grow large enough to
defend a territory.13 But some species change multiple times.
Okinawa pygmy gobies start out as a large-­gamete producers, and
then, if they grow to become the largest goby in the group, they
transition to small-­gamete-­producing physiology. However, if
a larger fish enters the group, the one that just switched from
large-­to small-­gamete producing switches back to large gam-
etes in the presence of the new, larger goby.14 The triggers for
sex-­biology change in fish are frequently social (a change in
makeup of the social group or local breeding population) or
directly tied to body size. In some coral gobies, change in sex
biology in one direction or the other depends on the sex-­
biology makeup of the population they are born into or migrate
into. The gobies can shift their biology in e­ ither direction
 28 Chapter 2

(­toward large-­or small-­gamete-­producing physiology) so that

any two fish can form a breeding pair. The ability to shift sex
biology flexibly by all individuals in a population reduces the
costs and risks incurred by moving between the coral clusters
where the gobies like to live.15
Internal gamete fusion is a ­whole dif­fer­ent ballgame from the
external fertilization we have just learned about in fish. Having
the gametes fuse internally is associated with a set of specific
reproductive organs in birds, reptiles, and mammals. With in-
ternal fusion in reptiles and birds, the large-­gamete-­producing
individuals’ bodies develop egg-­producing physiology and then
excrete the egg. In mammals with internal gestation (develop-
ment of the zygote into embryo and then fetus all internally),
­there is need for a very distinctive physiology, including f­ allopian
tubes, uterus, and, in most, a placenta as well. Th ­ ese specific
organs and bodily modifications in reptiles, birds, and mam-
mals create patterns of differences in development and adult
physiology between large-­and small-­gamete producers, giving
them reproductive physiologies that are less flexible than ­those
in the invertebrates and in fish. It is in ­these birds, reptiles, and
especially mammals, where one sees, on average, greater physi-
ological differences between the sex biologies of large-­and
small-­gamete producers. However, what ­those patterns are
across and within species, what is “typical” for any given repro-
ductive physiology, and how much overlap exists between the
bodies and be­hav­ior of large-­and small-­gamete producers
within a given species vary quite a lot.
Take reptiles, for example. They have a distinctive climate-­
influenced sex biology: many species have a temperature-­
dependent sex determination.16 In many species of lizard, ­turtles,
and crocodilians, the sex biology of a developing embryo is not
predetermined by a specific ge­ne­tic makeup. Rather, it’s a suite
 A n i m a l Se x Biol ogy 29

of hormonal responses driven by the local temperature around

the eggs during a critical period of development that shapes the
reproductive tracts and eventual associated adult physiology.
For many reptiles, developing large-­or small-­gamete-­producing
physiology is a context-­dependent outcome not predetermined
by the ge­ne­tic results of gamete fusion.
It’s not just how sex is determined that is in­ter­est­ing in
­reptiles; patterns of sex biology can also vary greatly across spe-
cies. ­There are species where females are larger than males (in
sea turtles and some snakes), o­ thers where males are larger than
females (in many lizards, such as Gila monsters where small-­
gamete producers have larger heads), and yet o­ thers where
­there is l­ ittle or no size difference (such as grass lizards). ­There
are a number of hypotheses for why ­these diverse patterns
emerge in dif­fer­ent reptile species, but it’s likely that both com-
petition between small-­gamete producers (likely due to sexual
se­lection) and environmental constraints on body size (for
both small-­and large-­gamete producers) are involved.17
­There is a particularly in­ter­est­ing mode of sex biology in a
cluster of lizard species (and some snakes): ­there are no small-­
gamete producers.18 In some whiptail lizard species, small-­
gamete-­producing physiologies have dis­appeared, and only
large-­gamete producers with egg-­laying reproductive tracts re-
main. ­These lizards evolved a complicated way to duplicate their
chromosomes and divide the results so that when they produce
large gametes, ­there is already a full complement of ge­ne­tic
­material in them with novel rearranging so it’s not just cloning.
­These gametes have no fusion with other gametes but are ready
to develop into an egg and eventually into a new individual.19
In t­ hese lizards, sex biology not only ­isn’t binary, it’s unitary.
Fi­nally, reptiles, regardless which sex-­biology variants they
exhibit, offer ­little in the way of care of the young. Some species
 30 Chapter 2

do guard eggs before they hatch, and other species carry the
eggs in their cloaca ­until they hatch rather than laying them, but
their care goes no further. Crocodilians are the only exceptions,
with adults ­doing some caretaking ­after the eggs hatch.20 It’s in
the animals that go all-­out for their offspring posthatching
(birds) and postbirth (mammals) that a w ­ hole new level of sex-­
biology complexity emerges.
Birds lay very large eggs, relative to adult body size, that
hatch into young who cannot fend for themselves (this type of
offspring is called “altricial”). Birds must care for their young.
In most birds, parental care is overseen by both the large-­and
small-­gamete producers. Such care likely emerged in protobirds
(better known as dinosaurs) from some forms of early care of
young that increased and facilitated the development of very
large eggs and highly altricial young.21 While most birds have
biparental care of young, the details of that caregiving vary
widely. How large-­and small-­gamete producers contribute to
such care is related to differences in social systems, mating
­systems, environmental context, and bodies.22 Parental care is
central to bird existence, but across—­and occasionally within—
species, ­there is not a consistent one-­to-­one correlation be-
tween a bird’s reproductive physiology (being a large-­or small-­
gamete producer) and how it participates in care for young
once the eggs are laid and ­after the young hatch.
In many bird species, ­there is sexual dimorphism (differ-
ences in the shapes, sizes, and/or colors of bodies) between
large-­and small-­gamete producers, but what form that takes is
not consistent or in a single pattern across all species. Body-­size
differences between large-­and small-­gamete producers often
relate to aspects of social and mating systems, to the patterns of
parental care, and to aspects of the specific ecologies of that
species. Like so many other animals, the patterns of how sex
 A n i m a l Se x Biol ogy 31

biology varies with bird species is dynamic and multifactorial:

­there are not simply two uniform sex types.23 For example, in
many species, small-­gamete producers are larger bodied. But
in some, especially birds of prey (like hawks and ­eagles), large-­
gamete producers are larger bodied. In birds where small-­
gamete producers do most of the competing for mates, they are
usually more colorful or have the most complex songs, but not
always. Also, sometimes t­ here is intense competition between
small-­gamete producers and almost no sexual dimorphism.
And in some species (like kingfishers and some parrots), large-­
gamete producers are more brightly colored than small-gamete
producers. In songbirds, small-­gamete producers sing most. But
in some s­ pecies, large-­gamete producers also sing, and in t­ hose
species, they are often more colorful.24 ­There are even some
bird species with multiple “sex” categories. The white-­throated
sparrow has some changes to its chromosomes that effectively
produce four chromosomal types that have dif­fer­ent plumages
and a mating system wherein certain types are not compatible
with o­ thers. So, while t­ here are only two gamete-­producing
physiologies in the species, ­there are functionally four sexes in
the real­ity of the ­actual mating system.25
The takeaway is that while t­ here are typical trends in bird
morphology and be­hav­ior that differ between large-­ and
­small-­gamete producers, especially in that only large-­gamete
producers lay eggs, t­ here is no one-­to-­one universal correlation
between a bird’s reproductive physiology and its body size,
plumage color or brightness, mating or parenting be­hav­ior,
song patterns or the specifics of its social system. “Sexes” are not
uniform ­things in birds.
As varied as birds are, mammals, including h­ umans, do sex
biology differently from every­one ­else. The bodily pro­cesses
associated with mammalian internal gametic fusion and the
 32 Chapter 2

potential for, and actuality of, gestation result in some very spe-
cific physiological patterns. Mammals also produce and provide
high-­quality nutrition to infant(s) during the earliest stages of
life. Large-­gamete-­producing mammals typically can gestate and
lactate. Small-­gamete-­producing mammals cannot gestate
(with some variation in intersex individuals, including in
­humans) and rarely lactate.26 Mammals also exhibit high levels
and a wide variety of postbirth care of their young, including
care by one adult, care by two adults, care by many adults,
and care by entire groups. It is common, but not at all ubiqui-
tous or uniform, that large-­gamete producers are the primary
caretakers relative to other group members.
While sexually dimorphic patterns associated with reproduc-
tion are common in mammals, they are neither ubiquitously
pre­sent nor consistent across all species. ­There is variation in
body shape and size, physiological pro­cesses, hormone pat-
terns, genitals, caretaking, and a range of be­hav­ior. While one
can certainly assert that t­ here are typical patterns of reproduc-
tive physiology and be­hav­ior in mammals, ­there is si­mul­ta­
neously a wide range of variation rejecting the assertion of a
single “way to be” within and between mammalian species.27
­There are many successful ways to be reproductively effective
mammals. Hyenas, “monogamous” primates, and naked mole
rats offer fascinating examples.
Unlike the devious characters in the Lion King or the
­scoundrels or scavengers often represented in books and nature
documentaries, spotted hyenas are active and successful hunt-
ers, form strong and deeply bonded social groups, and mess
with most ­peoples’ assumptions about mammalian patterns of
sex biology.28 Spotted hyena large-­gamete producers are larger,
heavier, and more aggressive than small-­gamete producers, and
socially dominant over them. The external genitalia of ­these
 A n i m a l Se x Biol ogy 33

hyenas include labia that develop into what looks like a scrotum
and a clitoris that develops into what is called a “pseudopenis”
that is fully erectile and through which mating and birth take
place.29 One might think that such differences simply mean that
hyenas with ovaries are more “masculine” than ­those with tes-
tes. That is to say they get more androgen hormone exposure
during development and thus mimic “typical male mammal”
development. But that’s not the case. The patterns of androgen
concentrations (­those sets of hormones often associated with
development of “typical” sex biology for mammals with testes)
in spotted hyenas are complex and vary by and are dependent
on age, w­ hether animals reside in the group they ­were born into
or not, their social status within the group, and on reproductive
state.30 Fetal large-­gamete-­producing hyenas are not “masculin-
ized” by hormones in the womb. Case in point, the develop-
ment of the genitals in hyenas with ovaries is not, as researchers
once thought, controlled by testosterone or other androgen
exposure but is, in fact, mediated by a very complex interplay
between a range of multiple hormones. This distinct physiology
is impor­tant b­ ecause hyenas are not “intersexed” in any manner
or form, or by any biological definition. They are neither
“­hermaphroditic” or “masculinized females.” The hyena system
reflects typical ge­ne­tic and internal reproductive physiology
patterns for large-­gamete-­producing mammals but demon-
strates a nontypical way to be a successful large-­gamete-­
producing mammal. Hyenas force a reevaluation of how one
can, or should, define “masculine” or “feminine” in the context
of sexual biology and motivate reflection about the value of
­simple models of “male” and “female” in mammals.31
Similarly, ­there is a w
­ hole group of primates that challenges
the expected roles of large-­and small-­gamete producers. Most
primates live in big groups of several males and females and
 34 Chapter 2

young, and o­ thers live in groups of one male and many females
and young. But ­there is also a range of species that live in small
groups of two adults and their offspring. Th ­ ese so-­called “mo-
nogamous primates” include the nighttime dwelling owl mon-
keys, the titi monkeys, the marmoset and tamarin monkeys of
South Amer­i­ca, and the gibbons (a small ape) of Southeast
Asia. All are assumed to have a par­tic­u­lar pattern of sex biology
and be­hav­ior that differentiates them from the assumed mam-
malian pattern of large males and smaller, infant-­focused
­females. The general argument is that t­ hese primates have a
“monogamy package” of bodies and be­hav­ior consisting of
specific deviations from the mammalian typical pattern. The
variations on the typical mammalian pattern include large-­and
small-­gamete producers of the same size and shape (mono-
morphic), small-­gamete producers who do much or sometimes
all of the infant caretaking, and no dominance differences
across females and males. This “monogamy package,” while
deviating from the typical expected pattern for mammals, is
still based on the assumptions of anisogamy, and thus still fits
into the typical assumptions about mammalian sex biology.
The prob­lem is that this monogamy package does not exist.32
Basically, the only ­thing that all of the “monogamous” primates
share in common is that they primarily live in groups of two
adults and some young. In some species that live in pairs, sex
is only between the two adults (sexual monogamy). But in
­others, sex can happen with individuals outside the two-­adult
group (social monogamy). In some pair-­living species, small-­
gamete producers are larger than large-­gamete ones and do
­little to no infant care, as in many other nonmonogamous pri-
mates. In ­others, both types of gamete producers are the same
size, or large-­gamete producers might even be a bit bigger, and
small-­gamete producers do at least half if not 100 ­percent of the
 A n i m a l Se x Biol ogy 35

carry­ing and social care of infants. In pair-­living primate spe-

cies, the two adults are sometimes codominant, and other
times the large-­gamete producer is dominant, and t­ here are
also, occasionally, more than two adults in the group (“monoga-
mous” species are sometimes neither monogamous nor pair
living). Fi­nally, the hormonal profiles across large-­and small-­
gamete producers in the wide range of “monogamous” primates
vary depending on their reproductive condition/status, w ­ hether
they care for infants, the demographic makeup of the group,
age, and dominance status. Even within what was assumed to
be a straightforward “monogamy exception” to the typical
mammalian system of sex biology and be­hav­ior, t­ here is a huge
amount of variation in the biology and be­hav­ior of ­these pri-
mates living in small groups.
Naked mole rats are particularly strange mammals. They
have ­little hair, spend almost all their time in extensive burrows,
­don’t regulate their temperature well, and live in large colonies
with one reproducing, often tyrannical, “queen,” and a few re-
producing small-­gamete producers, with all other members
functioning as nonreproducing “workers.”33 The naked mole rat
social system is superficially like that of some social insects
(such as bees). However, ­there are no differences in sex biology
regarding chromosomes or internal reproductive physiology
between the two “types” of large-­gametes producers (“queen”
vs. non-­queen). ­There are no biologically distinct “workers” as
in bees. Nor is ­there pheromonal control of “workers” or any
lasting biological changes that alter their internal physiology.
The mole rat “queen” (sometimes ­there are two in a colony) has
a longer body than other mole rats and behaves differently.
While ­there are some hormonal differences in naked mole rats
relative to other species, they do not show any of the massive
sex-­biology differences between sex-­biology types that we see
 36 Chapter 2

in insects. In other words, naked mole rats have developed an

insect-­like social and behavioral system without major modifi-
cation to the standard set of mammalian sex biology. Naked
mole rats demonstrate that expectations of a consistent pattern
of social ­organization and be­hav­ior based on standard mam-
malian sex biology are not accurate.
Hyenas, naked mole rats, and “monogamous” primates are
only a few examples of variation relative to typical mammalian
patterns, but they illustrate that sex biology and its relation to
bodies and lives is not a clear-­cut ­process. Internal gestation,
lactation, and care of young does not limit mammals to one type
of social system, one strict set of be­hav­iors, or one single way
to successfully be “female” or “male.”34

Is ­There a Sex Binary in Other Animals?

Chapter 11 of one of my favorite books35 is titled “Beyond the
Binary: Evolution’s Rainbow,” and it opens with a quote attrib-
uted to the twentieth-­century evolutionary biologist J. B. S.
Haldane: “The universe is not only queerer than we suppose, it is
queerer than we can suppose.” ­Here “queer” means strange, fasci-
nating, dif­fer­ent, confounding of expectations. This quote is
fitting for the real­ity for sex biology. Gametes (sperm and ova)
in most animals can be described as binary, one large and one
small. But bodies, physiology, and be­hav­ior are not so easily
classified, and are queer indeed.
The distribution of large-­and small-­gamete-­producing
physiologies, what forms they take, ­whether they are found in
the same body, dif­fer­ent bodies, one body at dif­fer­ent times,
or even consistently within a given animal lineage varies. What
“female” or “male” means physiologically, behaviorally, and
ecologically can be variable across and within species, and a
 A n i m a l Se x Biol ogy 37

s­ imple take is often insufficient to categorize, or understand,

sex biology. Therefore, to effectively understand sex biology
and its relation to bodies and be­hav­iors, one must examine
closely the evolutionary and ecological history, as well as the
current biology and be­hav­ior, of a species or lineage. And
­nowhere is this more evident, impor­tant, and convoluted than
in ­humans.
 3

­Humans Are Messy

almost five thousand years ago, a group of ­people in the

eastern Iberian Peninsula (near where Valencia, Spain, is now)
laid to rest one of their kin and buried ivory, weapons, crystals,
and ostrich egg shells alongside the bones. When archeologists
discovered and analyzed the grave site, they asserted that the
shape of the pelvic bones was typical of a male and the ­treasures
indicated the individual was a revered leader. Further, ­because
of the bone shape and the conclusion that the individual was a
leader, they concluded the individual must be a man.
Fifteen years a­ fter the initial discovery, researchers analyzed
the teeth using a new technique that detects proteins specific to
X and Y chromosomes. Lo and behold, the revered leader’s pro-
teins reflected no Y chromosomes. The individual was reclassi-
fied as female.1 Oops.
What happened h­ ere? The initial researchers assumed men’s
bodies and lives are one way and ­women’s another. In this in-
stance, they believed that the specific grave goods ­were signs of
high social status and reflected a specific set of “men’s” and
“­women’s” social roles. And, in combination with the pelvic
bones looking “typically male,” they made the assessment to
classify both the biology and gender and call the individual a

38
 Hum a ns A r e M essy 39

man. But we d­ on’t know if this individual was classified as a

“man,” a “­woman,” another category, or even if t­ hose categories
­were the ones used in that society five thousand years ago. What
we can say is that the individual had elite grave goods and a
body the reflected a variation on what is often assumed to be
“typical” sex biology. Placing con­temporary assumptions about
gender roles and a binary sex biology classification on past
­humans, or on pre­sent ­humans, is almost aways shoddy science.
The moral of this story? ­Human sex and gender are compli-
cated. ­Every ­human body, and life, is a blend of biological and
cultural pro­cesses, and nowhere is that more evident than sex
biology and the h­ uman experience. And that blend is what this
book is ultimately about.

­Human Sex Biology Is Complicated

In the big picture of h­ uman history, and in the biological
­sciences, the binary definition of sex biology is a recent inven-
tion. It emerged out of par­tic­u­lar cultural and philosophical com-
mitments centered in the ­European enlightenment and infused
the biological sciences across the eigh­teenth to the twentieth
centuries.2 For at least the last ­century and a half, biologists gen-
erally use the terms “female” and “male” when referring to what
type of gametes (ova or sperm) a sexually reproducing organism
produces. But gametes ­don’t magically appear out of thin air.
They are created, managed, and maintained by complex physi-
ological pro­cesses and organ systems. Biologically, “sex” involves
all the pro­cesses of sexual reproduction—­not just gametes.
Late in the twentieth c­ entury, scholars challenged the abso-
lutism of the binary view of sex biology and proposed defini-
tions of sex as bimodal (having two broad ranges that overlap
extensively). Now, in the twenty-­first ­century, many researchers
 40 Chapter 3

see the bimodal model as still too much of an oversimplification.

What is referred to as “sex” biologically comprises multiple traits
and pro­cesses with variable distributions and patterns. The best
current data and analyses produce a con­temporary understand-
ing that a “reliance on strict binary categories of sex fails to ac-
curately capture the diverse and nuanced nature of sex.”3
­There are few universally consistent connections between
the ability to make sperm, ova, both or neither, and the majority
of bodily functions. Even in mammals, with internal gestation
and lactation, ­there are variable connections between gamete-­
production physiology and bodily characteristics like body-­fat,
muscle density, height, blood pressure, metabolism, brain func-
tion, and a range of other aspects of biological function both
across and sometimes within species. ­There are even fewer uni-
form connections between gamete production and specific be-
havioral patterns, aside from the specific be­hav­ior associated
with giving birth and lactating. And even in the arena of birthing,
the postbirth be­hav­ior and the investment by adults into raising
offspring is not consistently associated with a specific gamete
type. While one can describe gametes as binary (of two distinct
kinds), the systems, be­hav­iors, and bodies that produce them are
not. And in the case of h­ umans, it’s even more complicated.

­Humans Are Biocultural

The material structures of ­human bodies, as with all animals,
are the product of complex interactions of biological, develop-
mental, and environmental systems. Th ­ ings like hair color and
type, nose, feet, and head shape, the length of legs and arms,
and the shades of our skin are generally described as our “biol-
ogy.” However, ­these structural parts of the body exist within
and are part of the ­human cultural context. ­Humans shave, cut,
 Hum a ns A r e M essy 41

or color their hair, bind their feet in shoes altering their shape,
change their noses surgically, with ornaments, and via life ex-
perience, eat a range of foods, and participate in activities that
affect growth patterns and limb morphology; they even alter
the shades of their skin by activity, clothing, chemistry, and tat-
toos. The h­ uman foot may be a collection of tissues, ligaments,
bones, and muscles arising from the interactions of ge­ne­tics
and development, but for e­ very ­human, their foot—­including
its form and function—is as much ­shaped by their cultural
selves as it is part of their biological structure.
How ­humans taste food, whom they are attracted to, the
sports they play, and how well they play them, are always af-
fected by biology and culture. Taste bud reactivity, muscles and
coordination, and the targets of sexual desire all emerge from
the mutual, and interactive, development of bodies and lived
experiences. Adult height and weight, one’s ability to score well
on standardized exams, views on raising ­children, ­resistance (or
lack thereof) to disease-­causing bacteria, and even what any
given person thinks of as natu­ral be­hav­ior for a man or a w
­ oman,
are a product of dynamic relationships that interweave the bio-
logical and the cultural, as well as the historical, into a single
result: us.4 Although we are biological organisms, the totality of
the h­ uman experience cannot be reduced to ­either specific in-
nate (biological) or external (environmental/cultural) influ-
ences. It is a synthesis of both: ­humans are biocultural.
The ­human brain is about 40 ­percent of its adult size at
birth, relatively smaller than any other mammal. Most ­human
brain growth happens out in the world, not in the womb. The
brain and the entire ­nervous system develop, create, and alter
connections and pathways and flows of biochemical signals in
constant exchange with bodies, senses, the social and physical
environments each ­human inhabits, including their own
 42 Chapter 3

thoughts, perceptions, and experiences of all ­these ­things.5 This

is the embodiment of culture: h­ uman bodies are literally cul-
tural organisms. Cultural experiences are integrated into skin,
muscles, nerve fibers, neurons, and other physiological systems,
and the entire p­ rocess is central to the development, makeup,
and function of the ­human animal.6
­Humans are born into a world of social and physical ecolo-
gies, patterns, institutions, and ideologies that become inextri-
cably entangled with our biology, even before we leave the
womb.7 Even basic perceptions such as smell, color, sound, and
taste are mutually s­ haped by physiology and cultural experi-
ence. Think about many of the ­things associated with sex
­biology in ­humans: femininity, masculinity, sexuality, sports
abilities, child-­care, health, body shape, how one speaks, how
one walks, and so on. ­Every single one of ­these ele­ments reflects
a deep interweaving of biological, cultural, and social ­factors,
and more. ­Every aspect of the ­human experience of sex biology
is deeply biocultural. And gender is central in that experience.

An Ova Is Not a ­Woman

and a Sperm Is Not a Man
When most h­ umans talk about sex, they are not r­ eally referring
to biology. More often, they are talking about gender. Gender
is the social and structural experiences framed by socie­ties and
cultures that includes norms and expectations for be­hav­ior
and appearance associated with aspects of sex biology.8 Gender
is femininity, masculinity, and other frames for social roles. As
such, the terms “man” and “­woman” are primarily about gender,
not biology. Simone de Beauvoir lays this out in her discussion
of “­women” in the patriarchal society of her time stating, “One
 Hum a ns A r e M essy 43

is not born, but rather becomes, a ­woman. No biological, psy-

chic, or economic destiny defines the figure that the ­human
female takes on in society; it is civilization as a w ­ hole that elab-
orates this intermediary product between the male and the
eunuch that is called feminine.”9
Gender is a crucial ­factor in ­human lives ­because it shapes
and constrains be­hav­ior, with groups and institutions reward-
ing/recognizing ­those who meet gender expectations and often
punishing/dismissing ­those who violate or contravene them.10
Gender is all around us.11 Gender roles are a power­ful influence
when thinking about sex biology b­ ecause they affect e­ very as-
pect of daily life for individuals. Gender can shape biology it-
self. Case in point: a recent study of over a thousand brain scans
from twenty-­nine countries demonstrated that greater gender
­inequality in a society seems to affect the neurobiological re-
sponses of ­women differently than in the contexts of socie­ties
with reduced ­inequality.12
Gender norms consciously and subconsciously assert how
we believe certain bodies and body parts “should” be and
how the h­ umans with t­ hose bodily p­ resentations “should” be-
have: for example, who can and should wear a dress, a sari, a
­miniskirt, tight shorts, or a kilt, and what reactions the “wrong”
bodies wearing them elicit. At the same time, society’s gender
expectations dictate who holds a door open for whom, how one
sneezes or belches, and even how one sits or stands. Nearly
­every aspect of daily life is influenced by the gender-­space/role
one occupies. Look at upper classes in certain E ­ uropean nations
across the fifteenth to eigh­teenth centuries, for instance. Men
wore makeup, long wigs, high-­heeled shoes, and tights to convey
masculinity. ­Today, in many of t­ hose same regions, t­ hese items
are associated with ­women and femininity. Or consider the real­
ity that cultural roles for and social ­acceptance of transgendered
 44 Chapter 3

individuals was a typical aspect of many socie­ties’ gender sys-

tems historically.13 Such traditions became repressed or banned
with the colonial expansion of Chris­tian­ity and par­tic­ul­ar
­political systems over the past five centuries but are now be-
coming more common in some con­temporary cultures again.14
Gender, as a cultural experience, is not consistent or uni-
form. Overviews of global gender dynamics demonstrate sub-
stantial cross-­cultural variation in how gender plays out. Using
a ­simple woman/man dichotomy or feminine/masculine spec-
trum does not cover the entirety of gender roles across h­ uman
socie­ties.15 This is not to say ­there are not core commonalities
in con­temporary gender systems, especially in the context of
childcare, l­ abor, economic and religious roles, kinship, p­ olitical
structures, and expressed sexuality.16
However, ­there is rarely a perfect one-­to-­one correlation be-
tween patterns of biological variation and ­peoples’ lived experi-
ences of their bodies through gender. Being classified as, or
feeling like, a “boy” or a “girl” or “masculine” or “feminine” does
not necessarily correlate to what one’s genitals look like, what
ge­ne­tic sequences reside on their twenty-­third chromosome, or
­whether or not one can make ova or sperm. For example, a ma-
jority of, but not all, ­humans with a penis and testes associate
with the gendered categories of man or masculinity. But do all
­those individuals who identify as such have the same bodies?
Do they have the same desires and attractions in sexual part-
ners? Of course not. Even in a cluster of individuals identifying
as man and masculine ­there are substantive variations in e­ very
one of the key bodily and psychological aspects of that gendered
role. Such variation does not make individuals better or worse
at the gender role but demonstrates that such roles should not
be thought of as uniform, or in binaries, ­because that is not how
­people experience them. Also, for many h­ umans, aspects of
 Hum a ns A r e M essy 45

their sex biology do not “match” their assigned or personally

identified gendered categories, and ­there are ­those who sense
themselves as not corresponding to any of the available catego-
ries at all. Gender and biology have a complex relationship—
not a 1:1 correlation.
The assumption that gendered differences are primarily or
“best” explained by specific differences in sex biology is both
oversimplified and incorrect.17 To assume that every­one who
has a vagina, clitoris, and labia, or a penis and scrotum is a uni-
form kind of ­human relative to the “other” kind is insufficient
and ill-­informed. One’s sex biology is impor­tant and plays a
major role in how any given individual engages with gender and
other cultural dynamics of their society. But it is neither ­simple,
nor uniform, nor binary. It’s biocultural.

Two Terms ­We’ll Use

Given all that w ­ e’re covering in this book, we need to set some
terms to assist with clarity and precision in the remaining
­chapters. Any effective examination of the ­human experience
in relation to sex biology is always contingent on two ­things:
the sex-­biology variation in play, and the specifics of the cultur-
ally structured gendered system. It’s the dynamic interweaving
of ­these two traits that distinguishes h­ umans from many other
animals. For this reason, the best con­temporary scientific ap-
proach to this discussion is to frame it in terms of “gender/sex.”
Gender/sex is the notation used to reflect the intertwined
biocultural real­ity of bodies and experiences more accurately.18
Think of it as an extension of the ongoing rejection of the nature
vs. nurture frame in the study of h­ umans. By adopting a biocul-
tural framing, research on ­human bodies and lives is no longer
forced to arbitrarily, and often erroneously, separate the material
 46 Chapter 3

form from the lived experience (nature from nurture, or—­

better put—­biology from culture). This has proven e­ xtremely
impor­tant in health research, cognition, neuroscience, and a
myriad of other arenas.19 ­Today, gender/sex is the term increas-
ingly used by researchers who study sex, biology, gender, sexu-
ality, and be­hav­ior in ­humans to acknowledge the real­ity that
the biological and the cultural are always entangled and often
inseparable. So for the rest of the book, ­we’ll use the terms “gen-
der/sex” and “biocultural” in many locations both for clarity as
to what we are talking about and to reflect the con­temporary
state of science regarding ­humans.20
One final set of terms is necessary. Most research on ­human
sex biology uses “sex at birth” as the key category for dividing
up subjects by “sex” in the study. However, “sex at birth” usually
means a designation based on a classification of the genitals of
the newborn. Th ­ ere are usually no tests for chromosomal com-
plements, gonad types, or anything ­else along ­those lines. As
such, the “sex at birth” category is a ­little sloppy, and sometimes
misses the mark. What “sex at birth” is hoping to ­measure
is ­really what is called the 3G category of sex.21 The three Gs are
genes, gonads, and genitals. A “3G female” is a ­human who has
XX twenty-­third chromosomes, ovaries, and a clitoris/vagina/
labia. A “3G male” is a h­ uman who is XY and has testes and a
penis/scrotum. The assumption is that a look at the genitals
gives you a solid inference of the other 2 Gs and thus that this
works as the classification system and the working definition of
“sex.”22 The rationale for using this categorization is that the 3
Gs are highly but not absolutely correlated with one another.
Thus, while 3G categories reflect a set of typically patterned
variation related to reproductive biology, they are not even
close to 100 ­percent accurate at capturing the a­ ctual range of
­human biological variation, even in relation to the 3 Gs.23
 Hum a ns A r e M essy 47

3G categorization does not reflect the biological real­ity for at

least 1 ­percent of all h­ umans,24 if not more. This represents
at least ~80,000,000 h­ umans living right now. 3G female and
3G male are not absolutes, nor are they the only definition of
“sex” biologically or of the terms “male” and “female.” 3G cate-
gories do not always or necessarily map to the terms “man” and
“­woman,” nor are they necessarily correlated with the gender
dynamics of masculine and feminine in only one direction or
manner. But they are biologically definable categories (if at
times incomplete), which is better than “sex at birth,” or “man”
or “­woman,” and unfortunately most of the data w ­ e’ll be work-
ing with in the following chapters ­either assumedly or explic­itly
uses 3G categories. So when discussing t­ hese data, we ­will be
using ­these categories too.
Now, with a solid grasp of basic animal sex biology, the
­biocultural real­ity of being ­human, and the framing of ­human
gender/sex, we are ready to tackle sex biology and the ­human
experience past and pre­sent.
 4

­Humans Then

if you line up a monkey, an ape, and a ­human, you’ll see a lot

that looks similar, and some key differences. Then, if you watch
their be­hav­ior, ­there’ll be some similarities, but many more dif-
ferences. This is called the comparative approach: comparing
one kind of organism with its closest relatives to see what in
their bodies and be­hav­ior likely comes from a shared past. We
can also study an organism’s past by looking at the fossil rec­ord,
the history of its ancestors. Both approaches help us under-
stand h­ uman sex biology. H ­ umans are primates, a par­tic­u­lar
type of primate called hominoids (apes), and a very specific
lineage of hominoids called hominins (figure 1). The bodies we
walk around in and the socie­ties we live in are s­ haped by our
primate, hominoid, and hominin evolutionary histories.
Each node of the h­ uman past (primate, hominoid, hominin)
offers evolutionary clues to current h­ uman sex biology and how
it relates to bodies and minds.

Primates Are Us
Primates are mammals1 and thus have a reproductive physiol-
ogy that involves shaping large-­gamete-­producing bodies for
potential gestation and lactation. Being mammals also indicates

48
 Hum a ns Then 49

Other mammals Other primates Hominoids Humans

Hominins

Primates

Mammals

figure 1. The ­human evolutionary tree.

the presence of an extended system of care of the young ­after

birth.2 Primates have large, complicated brains and lives centered
around social relationships. A key part of ­these social relations
involves very needy infants who are often the center of atten-
tion.3 In many primate species, the ­mother is the main caretaker,
but from shortly ­after birth infants are moved from arm to arm
of other group members, spending time being held by, cared for,
and playing with siblings, kin and other members of the group.4
Some primate species have f­athers as the primary or only
­caretakers, and in o­ thers t­ here are no single primary roles, and
dif­fer­ent aspects of care are spread across multiple group mem-
bers. Primates as a lineage evolved a diversity of modes of infant
care and social life and a varied physiology that relates to both.
­Because primates evolved a focus on social relationships as
their main way to make it in the world, sexual interaction out-
side of reproduction is a big part of many species’ lives.5 Pri-
mate sex biology and be­hav­ior are not just about to getting
gametes together. Indeed, many primate species have socially
 50 Chapter 4

learned greetings, vocal calls, tool use, diverse ways of grooming

and developing and maintaining friendships in both sexual and
nonsexual contexts. ­These “cultural”6 pro­cesses play impor­tant
roles in primates’ daily lives. Primates’ social be­hav­ior and rela-
tionships play central structuring roles in how sex biology is
­shaped and experienced.
Before we explore primates further, a quick note: in keeping
with the typical terminology in primate studies, in the rest of
this section w ­ e’ll use the term “male” for small-­gamete produc-
ers and “female” for large-­gamete producers. This shorthand is
meant to reflect the biology associated with 3G females and 3G
males, not anything about specific behavioral roles or patterns.
­There is ­little known about the physiology of intersex nonhu-
man primates, except that the frequency of such variation is
prob­ably similar to that in h­ umans. However, ­there are few, if
any, studies that have looked specifically at variation in 3G cat-
egories in primates, but we do know that t­ here is substantial
biological and behavioral variation within the categories of
“male” and “female” across and within many primate species.
­There are impor­tant differences across primate species related
to sex biology. In roughly a third to a half of primate species,
males are a l­ittle larger bodied than females (10–20 ­percent). In
a smaller percentage of species, t­ here is even greater variation
in body size, with male bodies 25 ­percent to 100 ­percent larger
than females. In the species where males are substantially larger,
they can have huge canine teeth.7 In such species, males often
use their body size as a social tool to dominate and control the
group (but not always successfully8) and/or to fight with one
another over resources.9 This extreme variation in body and ca-
nine size in some primate males is argued to emerge from sexual
se­lection. As Darwin originally proposed, sexual se­lection results
from competition between or within large-­and small-­gamete
 Hum a ns Then 51

producers and is related to achieving successful reproduction. In

species with very large-­bodied males, females may be shaping
males’ evolution by preferring to mate with larger males, or
males may evolve larger bodies and bigger canine teeth via the
pressures of male-­male competition (or both).10
Usually, sexual se­lection in primates is seen as being facili-
tated by the basic differences in sex biology between males and
females, with males having low investment in reproduction
(beyond copulation) and females having very high investment.
Sound familiar? As ­we’ve already discussed, this assumption
based on gamete size and related investment is not wholly sup-
ported, and the original model of what anisogamy implies is not
universally accurate. ­There are many permutations and much
diversity in reproductive dynamics across lineages and species.
But ­there are some patterns in primates. In most species with
large size variation, females (­mothers and ­others) do the major-
ity of infant care, and males do fight among themselves (some-
times a lot). But t­ here are exceptions. In gorillas, where males
can be more than twice the size of females, adult males are very
tolerant of young and participate in their care.11 In macaque
and baboon monkeys, where males are up to 25–50 ­percent
larger than females, males typically d­ on’t do much caretaking,
but it is not totally uncommon for males to adopt and care for
infants whose m ­ others died, and in many of t­ hese species, some
males spend as much if not more time with infants and young
than they do with other males.12 The bottom line is that in
highly sexually dimorphic species of primates, reproductive
physiologies usually correlate with some patterned differences
in bodies and be­hav­ior, and that sexual se­lection likely had a
role in structuring some of ­these patterns of sex biology.
But the same is not true for species with l­ ittle or no body-­size
differences. In ­these less dimorphic primate species, behavioral
 52 Chapter 4

differences are small, absent, or, in some cases, in the reverse

direction of what they are in the species where males are much
larger. In chapter 2, we reviewed the pair-­living primates where
­there is often care of infants e­ ither by all group members or pri-
marily by an adult male. Most primates that live in small or pair-­
bonded groups are relatively monomorphic (same-­sized), and
in the ones where ­there is a lot of male care of young, ­there are
substantial commonalities in hormone activity across adults
who care for infants regardless of their sex biology.13 ­There are
also non-­pair-­living primates that are monomorphic, and in
­those species, we d­ on’t see the pattern of male dominance, nor
its biological correlates, common in primates with large size
variation between sexes.14
While primates come in all shapes and sizes, from the tiny
mouse lemur that can fit into the palm of a ­human hand to
the mountain gorillas who weigh three times what most
­human adults do, they are relatively consistent in one critical
way: primate infants are large, slow growing, and fully depen-
dent on care for a longer period that than most other mam-
mals. For primates, the patterns of care for young in a given
species often exert substantive influence on bodies and
be­hav­ior.

­Humans Are Hominoids (Apes)

­ ecause we are evolutionarily a part of the hominoid lineage,
B
that group of primates (the living apes) provides a glimpse of
what is shared between us and them, offering insight into what
is deeply ancestral to h­ uman sex biology and the h­ uman experi-
ence. Thus, it should not be surprising that apes demonstrate a
lot of variation in how sex biology plays out in bodies and be­
hav­ior, within and between species.
 Hum a ns Then 53

Gibbons are the smallest apes and are ­humans’ most distant
ape relatives. Gibbon females and males are mostly the same
size with similar-­sized canine teeth. However, in a few species,
males are slightly bigger, especially in the siamang, and in many
gibbon species adult males and females have different-­colored
fur.15 The most striking difference between many male and fe-
male gibbons are their vocalizations. All gibbon species give
long melodious vocalizations, called songs, but generally males
and females give dif­fer­ent versions, often as complementary
duets. Gibbons primarily live in small groups, often pair-­
bonded, with just two adults and some young. Female and male
be­hav­ior overlaps extensively except that females do most care
of the young, with the exception of siamangs, where males who
do a bit more care of the young than other gibbon males.16
Orangutans shared a common ancestor with the African
apes and h­ umans more than ten million years ago. Orangutans
are highly sexually dimorphic with adult males up to twice the
size of adult females. But t­ here are two dif­fer­ent types of “male”
orangutan, both basically 3G, but at least one (or two) of the Gs
acts differently, and other aspects of their bodies and be­hav­ior
also differ from one another. Most adult males develop a set of
distinctive fatty cheek pads called “flanges” on their face and a
large throat sack that enables them to give deep and power­ful
long calls. However, some adult males ­don’t develop the flanges
or do long calls. This “two kinds of 3G male” is an orangutan-­
distinctive sex biology.17 The males that do not develop the
flanges and other aspects of large adult male appearance seem
to not have the circulating levels of a set of hormones (LH,
testosterone, and DHT) necessary for development of the
flanges and a full throat sack. However, t­ hese unflanged males
do have sufficient hormone levels and patterns (specifically tes-
ticular ­steroids and FSH) to develop full sexual function and
 54 Chapter 4

sperm production.18 Their gonadal and other endocrine (hor-

mone) function is not aty­pi­cal, but it is dif­fer­ent from that of
the flanged males. So, unlike most primates, orangutans have
three body types related to sex biology: small females, flanged
males, and unflanged males, and t­ hese types behave differently.
Flanged males spend the majority of their adult lives alone,
coming together with females for a few days now and then to
mate. Unflanged males also spend most of their time alone, but
sometimes hang out for a while with another unflanged male,
especially when they are young. Females spend the majority of
their time with their young offspring and on occasion with one
or a few other females. Orangutan social lives and sex biology
are distinctively dif­fer­ent from ­those of the other ­great apes.19
Gorillas, as noted above, have enormous size variation.
Males can be more than twice the size of females. Gorillas often
live in groups with one adult male and a number of females and
young. But as many as 40 ­percent of gorilla groups have more
than one adult male. It is not uncommon for many gorilla
groups to come together in large multigroup assemblages from
time to time with many interactions, including homosexual
sexual be­hav­ior, between the members of the dif­f er­ent groups.20
While gorilla m­ others are the primary caretakers of infants, go-
rilla males are highly tolerant and offer both protection of the
young and often spend a good amount of time playing and in-
teracting with them.21 Gorillas do map to the general pattern of
male dominance in largely dimorphic primates, but they differ
from many such species in the fact that males are attentive to,
and engage with, the young regularly.
Chimpanzees and bonobos, both members of the genus Pan,
are h­ umans’ closest primate relatives. Th­ ese two dif­fer­ent ape
species illustrate a range of behavioral diversity but share a
more or less identical basic sex biology. Chimpanzees (Pan
 Hum a ns Then 55

troglodytes) have slightly more sex-­related variation in body size

than ­humans and have much larger canine teeth. They live in
communities that break up into smaller subgroups for much of
the time, occasionally coming together as the full group. Adult
males in most chimpanzee communities spend the majority of
time with other adult males, and females mostly spend time
with their young and a few other females. Males are often so-
cially dominant to females and use aggression regularly. Sexual
interactions occur heterosexually and homosexually as part of
the complex social lives of chimpanzees. M ­ others do most
of the infant care, but older siblings, still with the ­mother, often
chip in. The patterns of relationships across the range of chim-
panzee populations (and subspecies) vary. For example, the
Western chimpanzee subspecies has smaller groups, less effec-
tive male aggression ­toward females, and more participation in
group aggression and more hunting by females (in fact, they are
the main hunters who use wooden sticks as spears).22
Bonobos (Pan paniscus) have approximately the same level
of dimorphism as chimpanzees and pretty much the same sex
biology, but their social system and behavioral profiles are dif­
fer­ent. In bonobos, females tend to be the dominant individu-
als, and w ­ hole communities are more often together than in
chimpanzees. Unlike chimpanzees, where dif­fer­ent communi-
ties avoid one another or get into potentially lethal fights,
bonobo communities often get together in a calm and friendly
social manner (including having lots of sex). While infant care
is still largely by the m
­ other, interactions with the young are
more widespread and undertaken by a broad range of members
in the bonobo community. Sexual interactions are a central
facet of bonobo daily life, and nonreproductive sexual be­hav­
ior is common across all ages and sexes, homosexually and
heterosexually.23
 56 Chapter 4

The striking ­thing about chimpanzees and bonobos is that

they have pretty dif­fer­ent be­hav­ior and social systems, and
­patterns of sexual be­hav­ior, despite having the same sex biology.
And this is not only between the two species: dif­fer­ent popula-
tions and groups of chimpanzees can also vary quite a bit in
their social relations and sex-­related patterns. And both species
of Pan are quite dif­fer­ent behaviorally from gorillas and orang-
utans. The diversity of be­hav­iors and bodies in the hominoids,
who all have a very similar sex biology, demonstrates that for
­these closest ­human relatives a given pattern of sex biology does
not specify one par­tic­u­lar set of behavioral and social lives. Sex
biology in the ape experience is variable and is not universally
tied to one kind of relationship between bodies, be­hav­ior, what
type of gametes are produced, or which Gs one has: this is the
key part of our inheritance from our history as hominoids.
Evolution is as much about discontinuity as it is about con-
tinuity. Understanding that h­ umans and other hominoids share
an evolutionary history, a continuity, is to acknowledge that this
shared past shapes ­humans’ bodies and lives. But that’s only
part of the story. It’s the history of our more recent evolution
since our split with the other apes, a discontinuity, where we
find deeper and more specific insights into h­ uman sex biology
and experience ­today.24

­Humans as Hominins
The hominins separated from other hominoids sometime be-
tween seven and ten million years ago and diversified into a
cluster of lineages of ape-­like beings who walked upright on two
legs. B
­ ecause we are talking about organisms that lived in the
deep past (and all we have are fossils), we ­can’t truly assess any
of their Gs. When talking about fossils, we use the terms “male”
 Hum a ns Then 57

and “female” as proxies for 3G categories based on typical skel-

etal patterns for 3G ­humans ­today. Admittedly, ­these classifica-
tions are wrong at least some of the time, even in ­those with
bodies just like ours. Remember the “revered leader” case from
the opening of chapter 3.
The earliest hominins are not very well known, but by about
four million years ago, the hominin genus called Australopithecus
emerged, and in its early members are the likely roots of the
­human lineage. The Australopithecines ­were a bit smaller than
most h­ umans t­ oday and likely had a good deal of body-­size di-
morphism, with males larger than females. However, unlike many
other size-­dimorphic primates, they did not have significant
canine-­teeth dimorphism, and it’s unclear if the social be­hav­ior
and social systems of the early Australopithecines w ­ ere more
ape-­, other-­primate-­like, or more human-­like.25 The Australopith-
ecines lived in small groups with multiple adults and young, and
likely had very strong social bonds between group members.26
Between three and four million years ago, the antecedents of
the h­ uman niche (the way we h­ umans live in the world) ­were
clearly developing. By at least 3.3 million years ago, Australo-
pithecines ­were making and using stone tools and beginning to
interact with, and modify, the world around them in what ap-
pear to be precursors to human-­like lives.27 At least some of the
Australopithecines evolved a human-­like reproductive pattern
of giving birth to newborns that need much care and attention
from multiple group members in addition to the m ­ other.28
While this mix of very needy infants with l­ ittle motor develop-
ment and lots of neurobiological growth outside the womb is
found in other primates and some other mammals, the ­human
pattern of cooperative care of extremely helpless infants is rare
and creates a distinctive set of biological and behavioral patterns
for ­human bodies that have much to do with the emergence
 58 Chapter 4

h­ uman gender/sex (see below). Between two and three million

years ago, the first members of the genus Homo (­humans)
showed up and began the specifically ­human story.

­Humans as Genus Homo

As noted ­earlier, h­ uman babies are born with about 40 ­percent
of their total brain growth completed. Having more than half of
brain development occurring outside the womb in the world is
a feature not found in any other mammal or primate. ­Because
of the h­ uman infant’s slowly developing neuroanatomy, coor-
dination, movement capacities, and information-­processing
skills, they are almost completely dependent on ­others for the
first few years of life.29 The closest comparison would be a
chimpanzee, whose brain is about 60 ­percent grown when
born, but who can cling to her m ­ other’s fur and climb around
on the ­mother’s body almost immediately ­after birth. ­Humans
have no fur, and our infants cannot coordinate their limbs or
even hold their head up on their own when born. The ability to
simply move themselves along the ground, let alone climb on a
caretaker, ­doesn’t emerge for months. H ­ uman infants must be
carried, tended to, protected, fed, and nurtured more than
­those of any other species. But this helplessness is a necessary
step to developing the physiological, psychological, and social
pathways to adulthood, and to becoming the neurobiologically,
socially, and technologically complex beings we are. The p­ hysical
and perceptual development of touch, sounds, smells, tastes,
looks, likes, movement, and every­thing ­else is entangled in, and
­shaped by, the infant’s cultural landscape and all the actors in it.
To develop effectively, ­human brains and bodies, and thus
minds, are completely soaked in their cultural environment.
This is where the ­human biocultural experience begins; this
pattern evolved in our ancestors.
 Hum a ns Then 59

The high post-­birth-­brain-­growth and bioculture-­acquiring

infant pattern began in the genus Homo at least a million years
ago. But one cannot evolve giving birth to helpless infants first
and then figure out how to respond to it. The infants would die.
For such a system of infant development to evolve, ­there must
have been a serious capacity for cooperative collaboration and
contribution to infant well-­being already in place. But how did
such a caretaking system evolve, and what does that story that
tell us about our sex biology?
From its earliest appearance more than two million years
ago, the genus Homo shows ­little sexual dimorphism relative to
­earlier hominins and to most other apes. Homo has small body-­
size variation related to sex biology (~10–15 ­percent) and no
differences in canines. Aside from the typical skeletally identifi-
able correlates of birthing and slight body-­size variation, the
fossil and paleoarcheological rec­ord hold ­little evidence of
physical demarcations, and even less of behavioral and social
variation, between 3G categories in the genus Homo.30
Compared to other hominins (like the Australopithecines),
the genus Homo underwent specific morphological changes in
the pelvic girdle, legs, feet, arms, hands, skulls, teeth, and f­ aces,
alongside less easily ­measurable, but significant, behavioral and
cognitive shifts across the last two million years or so.31 During
this time period, much of what we take for granted as “­human”
emerged. ­Things like hypercooperation, complex social interac-
tions, and increasingly complex material technologies such as
tools and fire.32 Members of the genus Homo worked together,
assisting one another day in and day out, expanding the intensity
of mutual care and the depth of social bonds and reliance on
each other.33 Via this intensive cooperation and coordination,
Homo developed more intricate and diverse foraging and hunt-
ing patterns involving not only the use of stone, wood, and bone
tools, but also increasingly complex collaborative be­hav­ior and
 60 Chapter 4

communication. Around this time, the brains of the genus Homo

started getting more neurobiologically complex, necessitating a
longer growth and development period: Homo childhood ex-
tended to be the longest developmental p­ rocess for any primate
(or mammal). By at least about eight hundred thousand to a
million years ago, our ancestors began to experiment with fire,
eventually working together to use it to reshape bone, wood,
and stone, and to cook a plethora of dif­fer­ent foods. By three
hundred to four hundred thousand years ago, ­there is strong
evidence for social connections across dif­fer­ent groups and
places, the emergence of long-­distance exchange networks,
and increasingly dynamic intergroup relations. Then, in the past
few hundred thousand years, we see the emergence of engrav-
ings, carvings, and eventually art. And all this was accompanied by
increasingly complex communication and information sharing,
eventually resulting in that amazing system called language.34
Emerging from this Homo evolutionary history are three key
patterns that offer insight into how ­human sex biology and be­
hav­ior, our experience of gender/sex, evolved. The first is an
intense cooperation and coordination around childcare. The
second is increased diversity in social and sexual relationships.
And the third is the development of categories of culturally
­mediated perceptions and expectations of and for dif­fer­ent
members of the group related to aspects of sex biology, what we
call gender and gendered roles.

It Takes a Group to Raise a Child

In most primate species, ­mothers do the majority of infant care,
but some amount of allocare (care of young by individuals
other than the ­mother) is common across the primates.35
­Usually, the allocare is done by females related to the m
­ other,
 Hum a ns Then 61

and/or siblings, and/or the ­father, but in ­those species that

exhibit intensive allocare, the entire range of group members
assist in the raising of the young. Allocare be­hav­ior by other
group members alters the evolutionary pressures on ­mothers
and affects both group members’ and ­mothers’ bodies. Hor-
mone dynamics and physiologies shift in the caretakers of
heavy allocare species, making them somewhat dif­fer­ent from
other species with the same basic reproductive anatomy but
mother-­only caretaking systems.36 ­Humans are the only pri-
mates that combine intensive allocare be­hav­ior with superex-
tended childhood: we have a deeply cooperative and collabora-
tive multicaretaker childcare system, and ­human infants are
born in more need of more intense and long-­term care and as-
sistance than any other primate. H ­ uman infant caretaking
evolved as a group effort that reshaped ­human bodies.
Recent overviews of the available evidence for the evolution
of the distinctive ­human infant caretaking system demonstrate
that ­humans not only evolved a suite of biological pro­cesses to
deal with time-­and energy-­intensive infants, but that we also
take care of ­these infants through a variety of cultural adapta-
tions, including material culture, provisioning of food, and
shared child care. ­Humans evolved to be biocultural coopera-
tive caretakers in ­every sense of the word.37
By between a million and five hundred thousand years
ago, the genus Homo displayed remarkable levels of cooperation
and collaboration, more so than any other primate or hominin,
and they used this capacity to develop a par­tic­u­lar type of care-
taking.38 Caring for infants shifted from a primary reliance on
the ­m other to a system wherein h­ umans w ­ ere raised by
“­Mothers and ­Others.”39 The ­human infant is a distinctive evo-
lutionary twist on primate babies in that the level of care, atten-
tion, and engagement that it requires to develop could only
 62 Chapter 4

emerge in a system where multiple potential caretakers are al-

ready pre­sent. The way our lineage enabled such helpless infants
to emerge, paving the way to our incredible cognitive and neu-
robiological biocultural evolution, was the co-­option of an al-
ready strongly developed pattern of cooperation and applying
that to childrearing. Multiple researchers over the past few
­decades have demonstrated via fossil, archeological, and com-
parative studies on con­temporary ­human bodies, that caretaking
in the genus Homo evolved to involve more than the m ­ other;
siblings and other members of the group ­were crucial to the ef-
fective raising of a child. Carrying of Homo infants, a large cost,
was spread across group members.40 Older individuals and sib-
lings ­were and are key caretakers, and caretaking by a range of
older individuals might even have resulted in the evolution of
longer ­human life-­spans.41 All h­ uman bodies (not just 3G-­female
ones) evolved strong physiological/hormonal responses for
caretaking be­hav­ior. For example, unlike many mammals, ­human
adult 3G-­male bodies change physiologically in the presence of
infants, most strongly when they are engaged in caretaking of
them.42 The extent and structure of cooperative parenting and
broadscale caretaking in the genus Homo created a distinctive
context for ­human evolution and ­shaped ­human sex biology.43
The genus Homo evolved a system where the ­mother is never
alone, so the pressures of successful reproduction are not solely
on ­mothers’ bodies. The evolutionary implication of this deeply
cooperative and complex caretaking is a rejection of a s­ imple
calculus of extreme evolutionary differences for bodies and
­be­hav­ior based on gamete producing and individual costs of
gestation and lactation.44 The data from the fossil rec­ord and
archeological evidence related to reproduction and energetic
investments in the genus Homo indicate that we cannot model
an understanding of h­ uman sex biology simply by separating
small-­and large-­gamete producers assuming a fixed and massive
 Hum a ns Then 63

difference of investment in reproductive effort. Nor can we lean

solely on the expected constraints of having a uterus and ovaries,
or testes, to understand ­human sex biology’s relation to the rest
of our bodies and be­hav­ior. Indeed, a ­decade ago, a cluster of
biologists45 specifically reviewing Bateman’s princi­ples (the
­asserted universal impacts of anisogamy) and con­temporary
sexual se­lection theory demonstrated that “­human mating strat-
egies are unlikely to conform to a single universal pattern.”46
­There are a few other aspects of h­ uman mating that differ from
­those of many other mammals: almost ­every adult produces off-
spring, and very few produce way more than o­ thers (­humans have
a very low “reproductive skew”). H ­ umans have the lowest varia-
tion between individuals in number of offspring produced and
the strongest similarity between females and males in number of
offspring produced of almost any mammal. In short, h­ umans
evolved a distinctive relationship between sex biology, mating,
reproduction, and parenting be­hav­ior.47 Therefore, explanations
for ­human bodies and be­hav­ior that rest on the assumption of
major evolved differences between 3G females and 3G males re-
lated to reproduction and parenting are inconsistent with the ex-
isting data on ­human evolution, and ­human reproduction, and
thus offer insufficient explanatory power.48 The ­human lineage
evolved a particularly intense, social, and communal mode of re-
production and childcare that shapes how our sex biology works
and what it means for h­ uman bodies and be­hav­ior. But sex biol-
ogy is not just about making and caring for babies.

Sex Is More Than Reproduction

Genital-­genital contact, genital manipulation via hands, mouth,
and feet, genital-­anal contact, and other forms of sexual activity
are found outside of reproductive possibilities in many mon-
keys, and in all apes and h­ umans. In some primates, social sex
 64 Chapter 4

is as frequent as reproductive sex (if not more so).49 In such

species, sex is a social tool, and sex biology must be understood
in the context of both its role in reproduction and its use in
social relations.50
Both members of the ape genus Pan (our closest primate
relatives) excel at social sex. Bonobos are famous for frequent
social sex, and chimpanzees51 also participate in sexual be­hav­
ior outside of reproductive possibilities.52 Bonobos engage in
sexual activity in dominance relations, feeding, group cohesion,
play, bonding, fighting and reconciling, and pretty much every­
thing ­else (including reproduction). Being so sexually active
has hormonal and other physiological impacts.53 Chimpanzees
also use sexual be­hav­ior as a social tool, but not quite as much
as bonobos. And in chimpanzees it may be more common be-
tween males than females.54 In gorillas55 and orangutans,56 sex
outside of reproduction is less common, but it is pre­sent, most
often in homosexual contexts. And ­humans outdo all our pri-
mate cousins with an enormous range of nonreproductive
sexual activity. In fact, one could argue that most sexual activity
in our species occurs outside of reproductive possibilities.57
The pervasiveness of sexual activity as a social tool in apes
and in con­temporary ­humans suggests that such be­hav­ior is an
ancestral hominoid state for apes and ­humans. Across homi-
noid and hominin evolutionary history, sexual interactions are
often detached from reproductive function but are attached to
other social and physiological functions. So, any explanation for
the evolutionary impact, functions, and constraints or benefits
of sex biology cannot only be explained by its relation to repro-
duction. Given that ­humans are much more socially sexual than
any related species (except maybe bonobos), the most likely
explanation is that across hominin history, and especially in the
last two million years of the evolution of the genus Homo, the
 Hum a ns Then 65

role of socio-­sexual be­hav­ior has been amplified as a major as-

pect of ­human sex biology. Sex biology is not just for making
babies, not in the apes, not in the hominins, and most certainly
not in ­humans.58 But, ­humans do make babies, have lots of
social-­sexual relations, and build amazingly strong social bonds.
All of ­these processes—­making babies, having sex, and making
bonds—­are interconnected.

Social and Sexual Pair Bonds Are Complicated

In the animal be­hav­ior lit­er­a­ture, a pair bond is a special, pre-
dictable relationship between two adults, often a reproductive
relationship. ­Humans frequently have pair bonds. This fact has
been used to argue that the presence of ­human pair bonds
stems from patterns of difference in 3G-­male and 3G-­female sex
biology. Some researchers suggest that ­humans evolved hetero-
sexual pair bonds and the nuclear f­ amily ­because of the specif-
ics of our sex biology. The story goes something like this: given
basic differences in mammalian reproductive biology and the
costs of ­human infants, ­humans who give birth and gestate
(largely 3G females) are ­under extreme pressures, so bonding
with a high-­quality partner (a 3G male in this proposal) im-
proves her chances of successfully having and raising an in-
fant.59 This same argument assumes that 3G males would prefer
to mate with many 3G females and not invest highly in just one,
but the costs of the ­human infant and the competition for ac-
cess to the best-­quality m ­ others make some degree of pair
bonding a good option (especially if the 3G males can keep the
option to “cheat” on occasion open). This story is used to
­explain the “­human nature” of the nuclear ­family, 3G females as
primary caregivers, and 3G males’ tendency to seeking more sex
partners than females. But is it accurate?
 66 Chapter 4

In ­human evolution, the pair bond has been characterized as

a “special and predictable relationship between a male and a fe-
male that involves tight social connections and a sexual relation-
ship, and usually includes mating and the raising of young.”60
Many scholars have argued that this “pair bond” set-up is mo-
nogamous, is the basis of ­human society, and evolved in our
hominin ancestors due to the assumed core costs and structures
of mammalian large-­gamete-­and small-­gamete-­producers’ bod-
ies.61 For example, some primatologists argue that the hetero-
sexual pair bond (and monogamous mating) precedes and gives
rise to the nuclear f­ amily structure in h­ uman evolution, and that
its appearance is a key point in the evolution of h­ uman society.62
But pair bonds are not necessarily linked to a nuclear ­family
structure in h­ umans or in other animals. In fact, pair bonds are
not necessarily linked to reproduction at all.63 Rather, pair bonds
are an effective way to enhance and expand the social networks
and cooperative possibilities, and they are not always related to
reproduction.64 And, as far as reproduction is concerned, we
know that, over approximately the last two million years, our
lineage evolved a complex system for raising and caring for
young that involves more than one m ­ other and one child (and
one sperm producer). We are not a species that evolved a system
of reproduction and social groups based on heterosexual pairs.
More than twenty-­five years ago, I began reviewing all the
available data for pair bonds in primates and h­ umans.65 I dis-
covered that primates (including h­ umans) are not more mo-
nogamous or pair-­bonded than other mammals,66 and that
while ­there are many primate species that live in small groups
consisting of a male-­female pair plus offspring, only some of
­those exhibit pair bonds. Work since then demonstrates that
pair bonds come in a number of dif­fer­ent types and are rarely
associated with ­actual reproductive monogamy.67
 Hum a ns Then 67

It turns out that t­ here are two types of pair bond: social and
sexual. The social pair bond is a “strong biological and psycho-
logical relationship between two individuals that is measurably
dif­fer­ent in physiological and emotional terms from general
friendships or other acquaintance relationships.” The sexual
pair bond is a “pair bond that has a sexual attraction component
such that the members of the sexual pair bond prefer to mate
with one another over other mating options.”68 In mammals,
including h­ umans, pair bonds combine social relations with a
range of biological activity.69
­Humans have social and sexual pair bonds. H ­ uman social
pair bonding occurs across the entire range of gender/sex and
age categories, with relatives and with unrelated individuals.70
­Humans have sexual pair bonds both heterosexually and homo-
sexually. And ­human sexual pair bonding is not necessarily re-
lated to reproduction. Therefore, the pair bond, sexual and/or
social, while critically impor­tant in h­ umans, is not something
that emerged directly out of our sex biology or something that
is constrained by it.

Gender?
Gender did not appear out of thin air. H ­ umans assign cultural
meaning to almost every­thing in the world, and our ancestors
began ­doing that a long time ago.71 Cultural meaning is ­assigned
not just to the material world around us but also to dif­fer­ent
aspects of the h­ uman experience, like age and life-­stage, body
type, hair patterns, genitals, demeanor, voice tenor, w ­ hether
one gives birth or lactates or not, who one has sex with, caring
for infants and o­ thers, and much more. The social dynamic that
we call “gender” t­ oday emerged from complex biocultural evo-
lution in the ­human lineage. H ­ uman gender concepts and
 68 Chapter 4

experiences have been ­shaped over evolutionary time by the

mix of cooperative caretaking, complex social sexuality, increas-
ingly complex social lives, and the specifics of sex biology and
other aspects of ­human bodies.
Gender is a set of expectations, perceptions, and be­hav­ior
that a social group believes about how bodies and be­hav­ior
should be in relation to aspects of sex biology. Obviously, bio­
logy plays a role in gender, but the specific history of a society,
its ecol­ogy, its demographic makeup, and a w ­ hole range of
other p­ olitical and economic f­ actors contribute the specific
­details of what gender-­particular roles emerge in any given
group. ­Humans’ deep shared evolutionary history and complex
dynamic variation in cultural diversity are why we have some
common patterns of gender across socie­ties and so much varia-
tion within and between them.72 However, while we can study
gender dynamics in ­humans ­today and in the very recent past,
it is almost impossible to see gender roles in the fossil and ar-
cheological rec­ord. That leaves us with the very unsatisfactory
answer that the capacity for gender clearly evolved, but what
gender roles looked like in the past is not at all clear. Interest-
ingly, what ­little evidence we do see suggests that if ­there was
something like gendered roles ­earlier in ­human evolution, they
­were prob­ably not identical to ­those of ­today.73
3G-­male Homo ­were, on average, about 10–15 ­percent larger
than 3G females, and we can assume that ­earlier Homo 3G males
had, on average, slightly higher muscle mass and density and
greater upper-­body strength (as they do ­today). We also know
that the nutritional and caloric needs of ­human 3G females with
a uterus go up at the very end of pregnancy and during lacta-
tion. This increase in energetic costs for female Homo likely
started around 1–1.5 million years ago and was part of the evo-
lutionary development of cooperative parenting.74
 Hum a ns Then 69

­ ese differences in body size and the fact of costly infants

Th
are often offered as clues to gender roles in h­ uman ancestors.
For example, it is common in recreations of h­ uman ancestors to
envision a man (an assumed 3G male) standing with a medium-­
sized dead animal slung over his back (the hunter), a w ­ oman (an
assumed 3G female) sitting or kneeling holding an infant (the
caretaker), and maybe another w ­ oman tending a fire with a tod-
dler at her side (the meal preparer), and then another man, usu-
ally a bit older, sitting making the stone tools (the tool maker).75
­These images are con­temporary gendered imaginings, not data
from the past. ­There is almost nothing in the fossil or archeo-
logical rec­ord that suggests a specific 3G-­sex biology or age
makeup or gender role for who did what or that ­there ­were “jobs”
like this in the past.76 Rather, for most of the history of the
genus Homo, ­there is no evidence for specific roles of “tool spe-
cialists,” “hunt specialists,” “meal preparers,” or “childcare spe-
cialists.”77 The gendered assumptions of “man the toolmaker,”
“man the hunter,” and “­women the caregiver” are recent cultural
inventions.78
In the comparative sense, we know that ape females use and
make tools slightly more than males and that young apes learn
to use tools primarily by watching their ­mothers. In-­depth anal-
yses of the reconstructed pro­cesses of stone-­tool creation and
use by Homo do not distinguish clear or consistent sex-­biology
differences in capacities or patterns in tool making and use.79
Analyses of recent, especially agricultural, tool use and manu-
facture (over approximately the last five to ten thousand years)
reveal some gendered role differentiation in use of tools, but
­these appear to be largely based on variation related to upper-­
body size/strength and the requisites of certain types of l­ abor.80
However, in most Homo populations over the last million years,
but before the last five to ten thousand years or so, ­there is
 70 Chapter 4

evidence of more overall upper-­body strength in both t­ hose

assumed to be 3G males and 3G females compared to most con­
temporary ­humans of ­either category.81 So, using con­temporary
­humans as the basis to understand ancient Homo strength and
capacity is underplaying the relative muscular/skeletal strength
of both Homo 3G males and 3G females in the past and over-
playing the relevance of sex-­biology-­related strength dimor-
phism for a majority of activities in ­human prehistory.82
In some con­temporary h­ uman socie­ties, men (the gender)
do hunt large game more than ­women (the gender), and in
many socie­ties, ­mothers, grand­mothers, and s­ isters are the
main individuals charged with childcare. However, assump-
tions about such role differentiation and its patterns in the
­human past, especially in the context of hunting and caretaking,
are challenged via a range of archeological, fossil, and recent
ethnographic analyses.83 This is not to say that vari­ous gender
roles did not exist. ­There is some evidence that while members
of Homo groups likely hunted together, they may have differed
in their roles in the hunt and in posthunt pro­cessing of foods
and hides. For example, recent evidence for hunting in the
­h uman past demonstrates not only a high likelihood of
­3G-­female participation in hunting (and other physically de-
manding activities) but show that they prob­ably excelled at
endurance aspects of hunting endeavors.84 In another example,
Neanderthal teeth from three dif­fer­ent sites show cultural wear
patterns, likely due to pro­cessing of meat and hides associated
with hunting, but ­there is some indication of slightly dif­fer­ent
wear and tooth-­chipping in teeth classified as 3G male and 3G
female. This suggests that ­there may have been slightly dif­fer­ent
“jobs” regarding the use of teeth in t­ hese Neanderthals. What
­these differences could be is hard to determine, but the fact
that the patterns are ­there in the fossil teeth suggests that some
 Hum a ns Then 71

form of gendered differences may have existed. But they w ­ eren’t

necessarily the same gender roles we see ­today.85 In the case of
caregiving, w ­ e’ve already seen the evidence for h­ uman coopera-
tive parenting, but at a broader level, t­ here is also robust evi-
dence that care for the injured, sick, and el­derly by all group
members emerged early on in the genus Homo, and that coor-
dinated and extensive compassion was a major ele­ment in the
success of our genus.86
Art is another area of frequent assertions about paleo-­gender.
­There is very l­ittle evidence of any gendered patterns in any
form of art before the last four to five thousand years, except a
recent survey of one kind of cave art: the outline of hands. Ar-
cheologists examined thirty-­two of ­these hand images from
eight dif­fer­ent cave art sites and calculated ­whether the hands
­were most likely male or female, based on the assumptions
about the ­human hand’s second digit to fourth digit (2D:4D)
ratio, and ­whether they w ­ ere adult or youth (based on size).87
The initial conclusion was that about 75 ­percent of the hand
stencils w­ ere likely done by females (smaller 2D:4D ratios). A
sex difference! Or not. It turns out that while the 2D:4D ratio
has been argued to be a good m ­ easure of patterned variation
between 3G males and 3G females (in mostly UK, ­European,
and US test subjects) and assumed to reflect differences in
­androgen hormone levels during fetal development (higher in
3G males than in 3G females), it d­ oesn’t. 2D:4D variation be-
tween 3G males and 3G females varies by population, is some-
times absent, and most likely reflects gendered activities rather
than a result of universal sex-­biology differences in prenatal
androgen exposure.88 2D:4D variation does not work as a sex-­
or gender-­classification system across all living ­humans, or for
­human ancestors. Interestingly, at least five of the hand prints
appeared to be t­ hose of teen­agers, and in other studies t­ hese
 72 Chapter 4

same kinds of hand stencils often show that many of the artists’
fin­gers ­were cut at the knuckles or w
­ ere missing.89 It is not at all
clear if this age pattern and the mutilation of hands tells us any-
thing about gender, but it does show that the making of this
kind of art was certainly a p­ rocess that resulted from the inter-
weaving of biological and cultural dynamics.
Many scholars note that heightened social and material com-
plexity (more complicated technology, p­ olitical systems, econ-
omies) and i­nequality, all usually associated with patterns of
con­temporary gender roles, do start showing up in many, but
not all, socie­ties, in the most recent phase of h­ uman evolution,
which is the last five to eight thousand years or so.90 This sug-
gests that the current sex biology of the genus Homo, which has
been around for much more than the last eight thousand years,
did not create con­temporary gender roles.91
Clearly, body size, strength, and reproductive physiology all
play substantive roles in h­ uman society and its structuring of
gender roles, but not always in consistent manners nor neces-
sarily in the same manners across time and culture. In the last
five to eight thousand years, con­temporary gender roles became
widespread across our species, initially seen in the archeological
evidence of burial patterns and grave goods.92 But the patterns
of inequity and gender roles that emerged and became common
in certain places and times do not always map to the majority
patterns of the pre­sent day—­remember the Iberian revered
leader from chapter 3 and the many recent archeological finds
of ancient “warriors” that w ­ ere assumed to be 3G male but have
turned out to be 3G female, at least in one or more of the Gs.93
Over the last six thousand years, researchers find increased evi-
dence of difference in the bone and tooth chemistry between
3G-­male and 3G-­female bodies, suggesting social differences
causing differences in nutritional status. Muscle scars and wear
 Hum a ns Then 73

marks on bones at archeological sites suggest slightly dif­fer­ent

lifestyles or work patterns between ­people, with some of t­ hese
patterned differences emerging between 3G males and 3G fe-
males (as best we can assess 3G sex from the remains). ­There is
also an uptick in the birth rates and a reduction in the time be-
tween births in the past five to eight thousand years, suggesting
that in many groups 3G females who can give birth ­were more
frequently pregnant and occupied in lactation than at e­ arlier
periods in ­human evolution. ­These biological markers indicate
the presence of be­hav­iors and lifeways that seem to reflect spe-
cific culturally s­ haped social dynamics and perceptions that
mark the emergence of some con­temporary gender roles.

­Humans Then S­ haped ­Humans Now

The cumulative evidence from the ­human evolutionary past
demonstrates that anisogamy and sexual se­lection hypotheses
based on less than solid assumptions about large-­and small-­
gamete-­producing biology do not offer adequate insight for
understanding the evolution of h­ uman bodies or be­hav­ior.
­Human evolutionary history is much more than gamete pro-
duction, mating competition, and simplistic assumptions about
sex biology and gender roles. ­Humans have evolved diverse
modes of ­family and community and an extremely dynamic and
very social sexuality. While recent gendered roles are broadly
distributed ­today, they are just that: recent. This brief journey
through the evolution of ­human bodies and be­hav­ior sets the
stage for the next necessary set of information: the details of sex
biology and its patterns of variation in ­humans ­today.
 5

­Humans Now

put together a random set of two hundred ­people and invite

fifteen 3G males and fifteen 3G females from that group to step
forward. Line them up by height, tallest to shortest. Looking over
the lineup, you w­ on’t see all 3G males on one side and all 3G fe-
males on the other. Instead, ­there’d most likely be more 3G males
in the taller half and more 3G females in the shorter half, but
individuals of both categories would be interspersed across the
entire lineup. That 3G males are on average 10 to 15 ­percent larger
than 3G females in our species does not mean that ­every male is
larger than ­every female. It just means that the averages in height
between 3G females and 3G males are separated by that percent-
age.1 Height in ­humans is not a sex binary and not a true dimor-
phism. Height is one morph (a m ­ easurable shape) with a range
of variation that can be divided into overlapping clusters com-
posed of 3G males and 3G females. But it does not automatically
have to be divided that way. Height distribution can also be
sorted by age, by ­people from dif­fer­ent latitudes and dietary prac-
tices, by athletes versus non-­athletes, and by a range of other vari-
ables depending on what questions you are asking. ­There are not
two forms of ­human, a tall and a short version; rather, t­ here is a
range of variation with some patterns in that variation.

74
 H u m a ns Now 75

median woman = 5 ft. 4 in. median man = 5 ft. 9½ in.

6
Number of adults
(in hundreds)

5
4
3
2
1

2 3 4 5 6 7 8 9
US men’s and women’s height
(in feet)

figure 2. US variation in height distribution in “men” and “­women.”

From A. Fausto-­Sterling. 2014. “The Science of Difference.” HuffPost,
May 23, updated December 6, 2017. https:/​/­www​.­huffpost​.­com​/­entry​
/­the​-­science​-­of​-­difference​-­lets​-­do​-­it​-­right​_­b​_­5372859​. Data from the
CDC: https:​/­/­www​.­cdc​.­gov​/­nchs​/­fastats​/­body​-­measurements​.htm.

Take, for example, figure 2, which shows the usual graph for
the range of heights in the United States when comparing 3G-­
male and 3G-­female bodies.2 One can focus on the median or
on the shaded clusters, but what is most significant, and often
overlooked, is that if one takes away the dif­fer­ent shading by 3G
category and the median markers, one is left with a distribution
of height wherein ~78 ­percent of individuals overlap and are
not sortable, by height, into a specific 3G category. So, despite
an average difference in height between 3G male and 3G fe-
males in the United States, height is an extremely unreliable
way to identify any individual’s 3G category.
The majority of ­human sex biology works in the same way as
height does: it is morphology with a range of expression across
­human bodies. This range of variation can often be broken
 76 Chapter 5

down into overlapping clusters via placing the bodies being

­measured into the categories of 3G females, 3G males, and
­people who do not fall neatly into 3G categories. But d­ oing so
automatically, before one has a specific question about the as-
pect of biology being ­measured, can obscure impor­tant pat-
terns and mislead one into thinking that 3G categories are the
main categories of key relevance. For example, while it is not
apparent in the black and white image, figure 2 (the original)
uses gendered (cultural) shading patterns (blue and pink) to
identify the clusters of 3G individuals and ignores non-3G in-
dividuals completely. Even though the graph is of a m ­ easurable
piece of ­human biology (height), it is presented in a heavi­ly
cultural fashion (as overlapping gender-­colored categories).
What if we w ­ ere interested in the effect of childhood diet on
height? Or about the length of the femur (our largest leg bone)
relative to the overall height? If we break the sample pool into
male and female before we take our m ­ easurements, we might
miss some very impor­tant variation that is not related to sex, or
we might see the variation as sex-­related when it might not be.
The immediate breaking of a sample pool into male and female
is a pattern of data p­ resentation, analyses, and discussion that
sometimes makes effective and accurate conversation about,
and understanding of, ­human variation difficult.
Most work on sex biology is based on specific assumptions
about “sex differences” and thus is framed as a female vs. male
comparison. Researchers rarely ask the initial question about
the overall patterns of variation of the specific trait or variable
being m­ easured (height, muscle strength, hormone levels, e­ tc.)
for the study subject pool. Instead, the subject pool is usually
divided into “male” and “female,” without any concrete defini-
tion of t­ hose terms beyond self-­reporting of assigned sex at
birth (based on genitals), before the data collection starts. The
 H u m a ns Now 77

prob­lem is that if one assumes t­ here is a difference and sets up

the research proj­ect to look for that difference, the likelihood
of finding difference is pretty high.3
­Because of the widespread pattern of assumed difference in
“males” and “females,” and the assumption that anyone not fitting
into ­those categories can be ignored, the methods (and theory)
in the vast majority of research into sex biology are conceived,
conducted, and presented as the study of “sex differences” and not
as the study of variation in sex biology. In this book, we are inter-
ested in the ­actual variation in and the patterns of sex biology. So,
while we review and discuss a range of studies on con­temporary
­human sex biology in this chapter, we are stuck with the ways in
which the studies ­were designed and the terminology, and meth-
ods, based on assumptions of difference they used. This makes
our task, trying to see what the ­actual ­human variation pattern is
before assuming that “sex” is the best way to divide up that varia-
tion, a ­little harder. But we have to use the data and analyses that
are available. I’ll try to point out the issues with certain studies
and some of the more egregious biases as we work through the
data, but I would suggest keeping ­these potential biases in
the forefront of your mind as you navigate this chapter.
­Humans vary in many aspects of our bodies, inside and out.
Much of this variation comes in par­tic­u­lar patterns. Some of it
is directly related to sex biology. But if we focus primarily on
averages and see every­thing from a core assumption of differ-
ence, we often obscure the biological real­ity of the trait in
­question. When thinking about bodies, we should consider the
forest of variation and not focus only on a few average trees,
certain tree clusters, or predetermined ways of looking at the
forest. This chapter’s brief survey of ­human bodies, hormones,
reproductive physiology, brains, and DNA reveals a lot about
­human variation, sex biology, and the ­human experience.
 78 Chapter 5

­Human Bodies Vary a Lot

All h­ umans share the same physical structures. ­Human tissues
(tendons, ligaments, bones, blood, skin, ­etc.) are all made of the
same materials. But tissues and the bodies they compose can
form a variety of shapes. In the h­ uman species, body mass (the
relation of weight to height) varies by as much as 40 to
50 ­percent across populations, and in sheer weight, the heaviest
­humans are as much as four times as heavy as the lightest. The
width of the h­ uman body at the pelvis (the hip) varies by about
25 ­percent across our species, and average body heights range
from about just ­under five feet to about six feet (from about 150
to 185 cm). Over the last 20,000 years, our species, on average,
has decreased in overall body mass, but recently in many areas
of the planet, as health care and nutritional patterns have
shifted, ­people have shown dramatic increases in height, adi-
pose accumulation, and body mass.4 Some of this bodily varia-
tion patterns within and between 3G categories.5
The USA National Health and Nutrition Examination Sur-
vey (NHANES) dataset illustrates some of this variation.
NHANES is a collection of physical and social data drawn from
a nationally representative sample of about 5,000 persons each
year since the 1960s, and individuals are identified by 3G cate-
gory before data are collected.6 In the 2015–2018 USA
NHANES7 dataset, non-­p regnant 3G-­female adults over
20 years old weighed between 110 and 263 lbs. and 3G males
over 20 between 136 and 287 lbs. In this same dataset, the 3G
females range from 4′11″ to 5′8″ and 3G males 5′4″ to 6′2″: so,
­there’s clear differentiation at the extremes, but the distribu-
tions overlap extensively. If one takes off the 3G labels and just
describes the variation, the majority of individuals fall into the
space between 136 and 263 lbs. regardless of 3G category, but
 H u m a ns Now 79

t­ here is more differentiation in 3G categories at the low and

high extremes. Th ­ ese data represent the 5th though the 95th per-
centiles, so 10 ­percent of all individuals in the dataset group fell
above and below the core numbers in height and weight, adding
a bit more complexity—­some individuals in both 3G categories
are even bigger and smaller than ­these numbers reveal. Keep in
mind that the weight data could be as much a 50 ­percent dif­fer­
ent and the height data as much as 15 to 20 ­percent dif­fer­ent if
­measured in other parts of the world. ­Human bodies vary a lot.
­There are also patterns of variation in skele­tons. While all
­humans have the same bones in pretty much the same shapes
(with the size variation as noted above), t­ here are a few skeletal
ele­ments that vary, on average, across and within 3G categories.
The shape of the pelvic girdle is, on average, more “flared” in 3G
females with a larger central space than in 3G males. The com-
mon argument for this difference is that a large-­headed h­ uman
infant requires a bigger pelvic opening in females than males.
But recent work challenges this explanation, demonstrating
that larger pelvic openings in females are common in many pri-
mates and some other mammals with smaller infant heads. Pel-
vic girdle variation is not just about birth requirements. A wide
array of patterns in organ size and positioning, plus hormone
and developmental pro­cesses, influence adult pelvic shape and
size, and ­there is a very large degree of overlap between 3G
­categories.8 Remember the “revered leader” skeleton? One of
the reason scholars labeled the skeleton a “male” was due to the
assessment of the “male-­like” pelvic girdle. And they w ­ ere
wrong. The pelvic girdle fell into a range where ­there is a ­little less
overlap between 3G categories, but t­ here was overlap nonethe-
less. Ignoring the possibilities of variation in f­ avor of an a­ ssumption
of fixed difference can lead to prob­lems when assessing skele­
tons, an issue front and center in the forensics world.9
 80 Chapter 5

In addition to the patterned pelvic variation, ­there are some

patterns in skulls. 3G females tend to have more vertical fore-
heads, smaller browridges above the eyes, fewer bony buildups
around muscle attachments, and smaller mastoids (the bony
protrusion just b­ ehind and below the ears). Also, the a­ ngle of
the mandibular ramus (the back of the jawbone) tends to be
larger in 3G females. ­These patterns are common, but ­there is
huge overlap between individuals in the two 3G categories in
­every one of ­these features. For example, I am a 3G male and
my mandibular ramus ­angle is large. Also, the relative appear-
ance and differences in many of ­these areas of the skulls are af-
fected by muscle action during life. So, the mastoids and neck
muscle attachments of 3G females who spend their lives carry­
ing weight on their heads ­will be larger than ­those who did not
grow up with this practice (and possibly larger than ­those of 3G
males as well). The shape of skele­tons differs across many vari-
ables, including some patterns between 3G categories, but t­ here
are not two distinct sex morphs of the ­human body.

Muscles Shape Bodies

As muscles act, they pull at the surface of the bones, causing
alterations to ­those bones, affecting the shape of the body. 3G
males, on average, have greater muscle mass than 3G females of
the same body size, and on average muscle strength in the upper
body is greater in 3G males than in 3G females. Th­ ese variations
in musculature, combined with variation in height, are a major
reason why, on average, bodies can look dif­fer­ent across 3G
categories.
A recent review of strength training, compiling data from
over five hundred published sources, helps contextualize some
of this variation.10 Muscle strength is directly related to muscle
 H u m a ns Now 81

architecture, the neural drive to the muscle (called voluntary

activation), and muscle fiber composition. The patterns of
­variation in muscle strength appear to be explained by both
overall body-­size dimensions (larger bodies have more muscle)
and differences in muscle mass. But training, lifetime activity,
and age also affect muscle ­performance and mass. 3G males on
average have more muscle mass than 3G females in absolute
terms (as a proportion of total body mass) and carry a greater
proportion of muscle mass in the upper body. This pattern
emerges around fifteen years of age, and by adulthood patterns
of differentiation in strength, on average, between 3G categories
are more pronounced in upper-­body than lower-­body muscles.11
When ­measured via vari­ous ­resistance, strength training, and
controlled exertion methods (pushing, pulling, and lifting
weights), adult 3G-­female upper-­body muscle and trunk strength
averages around 60 ­percent of 3G-­male upper-­body strength. On
the same ­measures, 3G-­female lower-­body muscle strength is
usually about 70 ­percent of 3G-­male lower-­body strength. ­These
figures all come from studies where the 3G categories are pre-­
identified and the tests are explic­itly comparing ­those catego-
ries (usually not also looking at within-­category variation or
patterns of overlap of the variation). Th ­ ese data are also com-
posites of many studies where overall size differences between
bodies and differences in muscle mass and density are not al-
ways controlled for. So, it is not totally clear what role size dif-
ference as a category is playing relative to 3G sex as category of
comparison. This is relevant as ­actual strength m­ easures of what
dif­fer­ent muscles do in real life (as opposed to in the gym or lab
lifting weights) are often smaller in between category compari-
sons with more overlap. But some patterns of between 3G cat-
egory ­measured strength still exist, to some degree, when com-
parisons are made between same height and weight 3G-­female
 82 Chapter 5

and 3G-­male individuals. This may be due to on average varia-

tion in patterns of muscle fiber in 3G males and 3G females, but
it is not clear if ­those differences are due to differential gendered
experiences and/or training, developmental sex-­biology differ-
ences, or a combination of the two.12
Much research controlling for the same muscle thickness
(and matched muscle size) between 3G males and 3G females
demonstrates some level of greater strength p­ erformance in
males in 76 to 88 ­percent of the comparisons (so 12 to 24 ­percent
of the comparisons showed no such difference between 3G cat-
egories). But other research comparing muscle strength in simi-
lar manners does not reveal the same patterns.13 In power-­lifting
competitions, 3G males can usually outperform even heavier-­
bodied 3G females.14 This suggests that for at least some mus-
cles (mostly in the upper body), this variation in strength is not
due simply to size but to the a­ ctual structure and functioning
of the muscles. However, we also know that a­ ctual structure and
functioning of the muscles is not a reflection exclusively of one’s
biology, ­because gendered experiences profoundly affect that
biology.
In all ­these assessments of muscle strength comparing 3G
males and 3G females, ­there is significant overlap. Unfortu-
nately, the vast majority of such muscle-­strength comparisons
do not control for lifetime activity or kinds of training and
­dif­fer­ent effects of societal gender norms and expectations for
activity patterns of bodies in the dif­fer­ent 3G categories: for ex-
ample, what kind of play/exercise boys and girls are expected,
or allowed, to undertake and how ­these affect muscle and
strength development in growing bodies. While all h­ umans can
increase muscle size and strength with strength training, in-
creases in upper-­body, but not lower-­body, strength are more
substantial for young adult 3G females than for young adult 3G
 H u m a ns Now 83

males (maybe due to the starting-­point differences), and older

adult 3G females increase relative lower-­body strength more
than older adult 3G males via strength training. Fi­nally, a pre-
ponderance of evidence suggests that patterns of variation in
strength between 3G-­male and 3G-­female bodies are not due to
differences in voluntary activation (the brain-­muscle connec-
tion), but rather to on average variation in muscle mass, size,
and density. That is, ­there does not appear to be any sex-­based
difference in the neural functioning of the muscular system, so
it remains remain unclear if t­ here is a set of dif­fer­ent gene activ-
ity related to muscle strength differences, or if the differences
are primarily due to muscle structure and development.15
Previous and ongoing physical activity, training, and lifetime
be­hav­ior make a major difference in adult muscle p­ erformance.16
For example, the speed of the ball in the fastest-­kicked goal in
the w ­ omen’s World Cup in 2023 was greater than that of the
fastest goal kicked in the men’s 2022–23 Premier League sea-
son.17 This is likely due to the increased quality in training and
a lifetime of sports engagement by t­ oday’s ­women’s football
(soccer) players. So some, but not all, of the variation in 3G
sexes’ muscle ­performance and the overlap between them may
be reduced, or increased, via gendered norms and be­hav­iors
favoring or restraining physical exercise/activity. For example,
many elite female athletes are often coached and trained in a
way that minimizes, rather than maximizes, bodily bulking up,
as the importance of appearing feminine is weighed against
­performance goals, both ­because of broad cultural norms and
­because of the importance of looking feminine to get endorse-
ment deals. Th ­ ere are also culturally mediated gendered differ-
ences in body image dissatisfaction across genders in a range of
sports and athletic practices that also impact or restrict how
dif­fer­ent individuals train and shape their bodies.18
 84 Chapter 5

In much of the con­temporary world, t­ here is an increased

focus on, and participation in, activities related to muscle de-
velopment and training in boys and men (who are most often
3G males) relative to girls and ­women (who are most often 3G
females). Across dif­fer­ent cultural gendered landscapes, one
sees a range of dif­fer­ent demands on the musculo-­skeletal sys-
tems in development from infancy to adulthood, in gender/sex
dynamics, and with age. ­Little of this cross-­cultural variation
has been m ­ easured, and few studies focus on non-­athlete or
non-­trained individuals. Most ­humans are neither athletes nor
strength trainers. Also, while the relative m ­ easures of muscles
mass and strength patterns represent a large number of samples,
they are mostly from specific cultural exercise and training
­contexts and in WEIRD19 nations (primarily ­European ­Union,
United Kingdom, Australia, United States, and Canada), so
­there may be some impor­tant differences in such patterns across
a more global comparison of ­human bodies.
While some overlapping but patterned variation appears in
body composition and in muscles, it’s in the genitals and repro-
ductive tract that well-­known sex-­biology variation, and lesser-­
known sex-­biology similarities, are found.

Genitals, Gonads, and Uteri, Oh My!

Despite the ­popular use of genitals to classify someone as male
or female, genitals ­don’t come in just two distinct forms. In the
nineteenth ­century, it was clear to scientists studying ­human
biology that a range of external genital morphology was pos­si­
ble, and t­ here was no way to sort it into two totally distinct
categories. ­Because genitals did not work as a distinction, by
the late 1800s researchers looking for markers of a binary sex
biology moved to the gonads, identifying ­those with testes as
 H u m a ns Now 85

male, and ­those with ovaries as female. However, very quickly

thereafter, improvements in microscopy and surgical t­ echniques
(especially biopsies) quickly revealed that some individuals had
both ovarian and testicular tissue in the same body, or even the
same gonad, and some had gonadal tissue that was neither.20
Both genitals and gonads have par­tic­u­lar patterns of variation
resulting in a set of typical morphology and function, but they
are not binary.
­Human genitals emerge ­after the first six weeks of develop-
ment from masses of embryonic tissue called the urogenital
folds.21 One ele­ment in the urogenital fold, the genital tubercle,
develops into ­either penile or clitoral forms. Another part forms
the labia or scrotum. This p­ rocess is not directly based on what
genes one has but rather by the actions of hormone and tissue
interactions—­although t­ here is a strong but not 100 ­percent
concordance of having certain ge­ne­tic sequences and the out-
comes of this p­ rocess.22 Another area, the gonadal ridge, re-
sponding to gene activity from multiple chromosomes, creates
a suite of hormone-­tissue interactions central in gonad forma-
tion.23 The end points of gonad formation are usually testes or
ovaries, which become involved in cholesterol-­derived s­ teroid
hormone production in interaction with hypothalamo-­pituitary
axis (HPA), a major neuroendocrine system, and gametogen-
esis (the production of ova or sperm). While it is not common,
gonadal development does produce ovotestes and/or both
­ovaries and testes in the same body as a recurring component
in the range of ­human sex-­biology variation.
In most h­ umans who are XX for their twenty-­third chromo-
somes and have no testes, or t­ hose who are not XX but who ex-
perience early life ge­ne­tic activation and developmental pro­cesses
similar to the pattern in t­ hose having two X chromosomes,24 a
uterus develops via the fusion of a set of tissues (the Mullerian
 86 Chapter 5

ducts) around and in the urogenital folds between the meso-

nephric ducts and the gonadal ridges. The vagina emerges from
interactions of the Mullerian ducts, the urogenital folds, the
Wolffian ducts, and other tissues, but the exact details are not
well understood.25 This internal set of developments (vagina,
uterus) seems facilitated by a range of pro­cesses centrally con-
nected to actions of a cluster of gene activation and inactivation
combined with estrogens and related hormones between the
tenth and twentieth weeks of embryonic development.26 ­We’ll
loop back to uteruses below.
The amazing t­ hing about the genitals and gonads is that,
­despite what seem like pretty dif­fer­ent endpoints in most
­people, the vari­ous versions are all made of the same stuff. Adult
external genitals are usually observably dif­fer­ent and categoriz-
able as such (for example, a penis and scrotum or a clitoris and
labia), but ­there is substantial commonality across a range of
shape and size. This commonality is clearly seen in sexual re-
sponse physiology: in the clitoris and penis, erectile tissues
function the same way. Understanding this developmental real­
ity helps us contextualize the large range of variation in ­human
genitals. Penis size and shape vary. Clitoris and labia vary. And
it is not always evident ­whether an infant has a clitoris or penis
at birth.27 Most importantly, most of this variation is irrelevant
to adult reproductive function; size and shape d­ on’t m­ atter that
much at all.
Genitals do not equal “biological sex,” or “male” or “female,”
but their adult forms do reflect critical sex-­biology variation
and patterns related to reproduction and other physiological
pro­cesses. And speaking of physiological pro­cesses, the very
vis­i­ble and often misunderstood dynamics of fat (adipose tis-
sue) plays a central role in how we both see and understand sex
biology in bodies.
 H u m a ns Now 87

Fun Facts about Fat (Adipose Tissues)

Fat ­matters. It shapes bodies, interacts with hormones, pro-
vides energy, regulates physiology, and has cultural meaning.28
Plus, adipose tissues are part of our sex biology. ­Humans de-
posit fat more or less the same way that most mammals do, but
we have very chunky infants (a high fat-­to-­bodyweight p­ ercentage
at birth) and “wear” our fat a l­ ittle differently. Biologically, “fat”
is adipose tissue and comes in two types, white and brown.
White adipose is mostly laid down u­ nder the skin in the ab-
dominal, lower back, and gluteofemoral regions (butt and
upper legs) and around the organs. Brown fat is mostly depos-
ited around the shoulders and neck. While ­there are typical
human-­wide patterns of deposition, specific patterns of where
and how fat gets laid down vary by geography, populations, 3G
sex, age, familial groups, and individuals.29 The pattern of fat
deposition is the same for 3G sexes u­ ntil puberty, when 3G fe-
males usually begin to increase total fat mass, eventually devel-
oping about 10 ­percent more total fat, on average, than a 3G
male of the same height and weight.30 Deposition location also
varies, with adult 3G females depositing more fat around the
chest area and gluteofemoral region and 3G males more around
the abdomen and the internal organs (usually). Th ­ ese patterns
are on average, and ­there is a huge range of variation from indi-
vidual to individual, and across the entire species.31 For exam-
ple, breasts in h­ umans with testes and no ovaries or uterus are
not uncommon. Approximately 30 to 60 ­percent of 3G males
experience a degree of adipose (and glandular tissue, see
below) deposition in their chest leading to the developments
of breasts at some point during their lifetimes (called ­either
gynecomastia or pseudogynecomastia or both).32 In ­humans
who menstruate and become pregnant, pregnancy and
 88 Chapter 5

menopause are both associated with changes in fat mass and its
patterns of deposition, but the specifics of how this pattern
plays out vary dramatically across geographic and cultural
­contexts. Th­ ere is mounting evidence that multiple X and Y
chromosome genes (plus many other genes), epige­ne­tic pro­
cesses, sociocultural stressors, and trauma influence specific
adipose deposition and metabolism. And t­ here are even mosaic
effects (where a person has two or more genet­ically dif­fer­ent
sets of cells in their body) on dif­fer­ent adipose locations within
the same individual.33 Adipose deposition, fat, while showing
some key average patterns, is quite variable across and within
3G-­sex categories in ­humans.
It’s worth noting that h­ uman breasts and butts are distinctive
largely b­ ecause of fat and bipedalism. And breasts and butts
often play culturally salient roles in our gender/sexed lives. The
basic mammalian pattern of adipose deposition, the fact that
­humans walk upright on two legs, and that we only have two
nipples (most mammals have multiple sets), combined with the
pull of gravity creates a distinctive look for ­humans. This “look”
involves breasts in most 3G females postpuberty (and in some
3G males) and pronounced buttocks for all ­humans, with po-
tentially relatively larger buttocks in t­ hose with ovaries and
uteruses. This gives ­human bodies a dif­fer­ent look than other
primates. The more frequent and greater development of breast
tissue in 3G females, the slightly greater adipose deposition
rates postpuberty, and average body-­size differences (smaller)
accentuate the appearance of breasts and butts on their bodies.
This small pattern of variation in fat deposition frequently
forms a basis for perceptions of 3G bodies as being more dif­fer­
ent than they are. Another vis­i­ble part of our bodies that plays
an outsized role in our interpretation of gender/sex and as-
sumptions about differences is hair.
 H u m a ns Now 89

­Human Hair
­ here are two key types of hair in ­humans. Vellus hair—­
T
nonpigmented, soft, and small—­covers much of the body (and
is hard to see). Terminal hairs are longer, more rigid, more pig-
mented, penetrate further into the dermis than vellus hairs, and
are what are what most think of as “hair.” Growth and thyroid
hormones are the main actors shaping hair patterns, but in cer-
tain areas of the body (face, genital region, upper torso, thighs)
androgens (see “Hormones Are Complicated” below) also play
a major role.
­Humans are weird when it comes to hair. We have relatively
scarce amounts on our body core, yet dense patches on our
heads, armpits, and around the genitals. 3G males have, on aver-
age, more and denser terminal hair than 3G females, but t­ here is
a lot of overlap between individuals within a group and some-
times extreme differences between populations of h­ umans
around the planet.34 ­There are groups of ­humans where both 3G
males and 3G females have very ­little body terminal hair and
other groups where all members of a group have substantial ter-
minal hair. In some groups, 3G males have higher densities and
distribution of terminal hair on the face than in other groups,
and it is not uncommon for 3G females in t­ hose groups to also
have greater facial hair. Hirsutism, defined as the presence of
“excess” body or facial terminal hair on 3G females, is m ­ easured
relative to a “standard expectation” for hair growth. This is a cul-
tural, not a biological, m ­ easure. Using this m ­ easure, 10 to
15 ­percent of ­those who identify as ­women get labeled hirsute
across the ­human species, with some ­human groups having
~35 ­percent of 3G females incorrectly labeled as having “male-­
like” hair patterns. This is incorrect ­because “hirsute” patterns of
hair growth are well within the typical range of variation for our
 90 Chapter 5

species, so labeling them “male-­like” is a gendered (cultural)

classification, not a biological one. ­There are vastly dif­fer­ent
levels of terminal hair growth on nonscalp bodily areas across
and within dif­fer­ent groups of p­ eople in our species, and thus
many 3G females in some populations ­will have higher levels of
terminal hair than 3G males in other populations and vice
versa.35 We know that terminal hair patterns are connected to
androgens, estrogens, growth, and thyroid hormone dynamics
and population developmental differences: how much and
what type of hair one has does not simply boil down to a “sex
difference,” as the patterns and variation are related to a variety
of physiological, geographic, ge­ne­tic, developmental, and sex-­
biology ­parameters.36 ­Human hair does not come in a binary,
but rather a spectrum with some hormone-­related effects that
often, but not always, pattern with re­spect to 3G categories.
So far, ­we’ve focused primarily on the vis­i­ble differences:
bodies, muscles, genitals, fat, hair. But what about our insides?

Organs Are Biocultural

Most ­human organs ­don’t seem related to sex biology. Kidneys,
livers, hearts, pancreases, gall bladders, lungs, and so on are not
discernably “male” or “female” except that, on average, the or-
gans in 3G-­male bodies are larger than t­ hose in 3G-­female bod-
ies. Basic organ function does not vary “by sex,” and 3G sex is not
a determining ­factor in decisions regarding transplants. While
­human organs are not biologically “sexed,” some are gender/
sexed: they are s­ haped in slightly differing manners as they
develop inside the gendered body. For example, on average,
3G-­male kidneys are larger, but they also, on average, have more
and/or larger nephrons (the part of the kidney that filters your
blood) than ­those in 3G females, which might affect function.
 H u m a ns Now 91

­ ere is also evidence from rodents, and a few ­human studies,

Th
that long-­term exposure to dif­fer­ent hormonal levels and dy-
namics (as happens in ­human bodies via age, 3G sex, and other
physiological and life-­experience variables) affects the be­hav­ior
of endocrine receptors on certain organs and potentially some
of their physiological characteristics. Higher-­c irculating
­testosterone, or more dramatic cycling of progesterone and es-
tradiol, might reshape some of the ways in which a given organ
functions. Also, inflammation, which can be heavi­ly impacted
by pregnancy, physiological/social stress, and a range of cultural
and nutritional dynamics, can also affect organs. A lifetime of
being in a par­tic­u­lar body in a par­tic­u­lar culture can shape
­organs in specific ways via patterns of endocrine exposures,
pregnancy (or not), alcohol and drug use, social, economic, nu-
tritional, and psychological differences in lived experiences of
gender/sex, age, race, socioeconomic class, caste, and so on.37

Hormones Are Complicated

The endocrine system, our hormones, is implicated in almost
every­thing we do. When considering sex biology, researchers
and the public often turn to hormones as a (or the) core dif-
ferentiation between females and males. But hormones d­ on’t
create bodies and d­ on’t make “males” or “females.” Rather, en-
docrine systems are impor­tant but not exclusive components
in the development of bodily phenotypes (what bodies look
like) and how they function.38 Hormones via the endocrine
system are how the body gets cells, tissues, organs, and other
parts to communicate and interact for critical pro­cesses, like
growth, metabolism, and reproduction.39 The endocrine s­ ystem
consists of the pituitary gland, the hypothalamus, the thyroid
and parathyroid glands, the adrenal glands on the kidneys, parts
 92 Chapter 5

of the pancreas, the gonads (testes and ovaries) and the hor-
mones that each of t­ hose glands/organs produce. Hormones
come from, and mediate, ge­ne­tic action and changes in be­hav­
ior and physiology in response to varying aspects of physical,
social, and developmental environments and experiences. And
they do so not by themselves, but via an intricate system of
varying hormone receptors and a network of feedback loops
within the endocrine system and between the endocrine sys-
tem and vari­ous organs and other physiological components of
bodies. Thinking that ­measuring individual hormone types or
levels is sufficient information for understanding how this sys-
tem works is incorrect and unscientific.40 Plus, all h­ umans have
the same hormones.41 ­There are no “male” or “female” hormones.
Rather, levels and actions of specific hormones, hormone re-
ceptors, and their outcomes vary with age, gonad physiology,
ge­ne­tic variation, life experiences, social contexts, nutritional
state, reproductive state, and much, much more. Endocrine sys-
tem activity is deeply biocultural.42
The main hormones secreted by the gonads are estrogens,
progestogens, and androgens.43 Both ovaries and testes pro-
duce all three (and a few o­ thers), but the patterns of production
vary dramatically, with ovaries capable of producing higher
­levels of estrogens and progestogens, and testes capable of pro-
ducing higher levels of androgens (specifically testosterone).
The gonadal production of hormones is directly connected to
and mediated by the hypothalamus and the pituitary. Ovaries
and testes are variants on the same theme, not dif­fer­ent kinds
of organ.44 However, gonadal actions and their outcomes vary
in patterned manners.45 Usually, t­ here is a concordance be-
tween the presence of ovaries or testes and specific other Gs
(ge­ne­tics and genitals). However, t­ here is a nonnegligible range
of variation in the specifies of gene action/inaction, gonadal
 H u m a ns Now 93

development, and other endocrine and physiological dynamics

such that the circulating levels of gonadal hormones, and their
effects, across individuals fall outside of what are seen as typical
3G patterns with regularity. Thus, 3G patterns are not always
accurate predictors of endocrine status, as t­ here is much varia-
tion in the system.46
Hormones also vary across the life cycle. Some initial varia-
tion in the circulating levels of testosterone and estrogen in
bodies emerges in the late embryo stage. Just ­after birth, the
hypothalamic-­pituitary-­gonadal (HPG) axis is activated, and
­humans undergo a “mini puberty” ­until about 3–6 months of
age. Th
­ ere are peaks in estradiol and testosterone that act to
facilitate the developmental trajectory of the genitals and other
reproductive anatomy (glandular development around the
nipples and ovarian and testicular gamete-­production areas).47
This early p­ rocess prob­ably sets the stage for l­ater functioning
of estrogen and androgen receptors and aspects of their varia-
tion and action in dif­fer­ent bodily areas. Differences in fluctuat-
ing gonadal hormone levels dis­appear in most bodies from
~1.5 years of age u­ ntil between ~8–10 years of age. At about
6–8 years of age, h­ umans start adrenarche, the adrenal glands
and the pituitary slowly increase the production of the hor-
mones DHEA, DHEAS, and ACTH, setting the stage for ­actual
puberty.48 Typically starting between 10 and 14 years of age,
puberty includes the full maturation of the HPG axis and the
genitals (called gonadarche) and the development of patterned
changes in hair growth, muscle mass, fat deposition, breast tis-
sue, an acceleration in height growth, and, in t­ hose with ovaries,
the occurrence of menarche.49
Menarche is the onset of menstruation and the specific
ramping up of the hypothalamic-­pituitary-­ovarian (HPO) axis.
From puberty forward, patterned variation in the levels of
 94 Chapter 5

circulating gonadal hormones becomes common between in-

dividuals with ovaries and ­those with testes. L ­ ater in life (about
fifty to fifty-­f ive years of age), ­those with ovaries undergo
­menopause wherein changes in the ovaries result in lowered
production of estradiol and inhibin (a protein), which alters the
production of the two main gonadotropins, follicle-­stimulating
hormone (FSH) and luteinizing hormone (LH), leading to the
cessation of reproductive cycling.50 Menopause has a variable
range of physiological effects in addition to the changes in
the hormone levels and is distinctive to h­ umans (but possibly
occurs in a few other animals as well). Individuals with testes
undergo an “andropause” slightly ­later than menopause, with a
decrease in the level of production of testosterone accompanied
by a suite of physiological and physical changes.51
Patterned variation in hormone activity is an impor­tant as-
pect of sex biology. And, while ­there is much variation between
individuals, ­there are species-­wide typical patterns. Estradiol (a
main form of estrogen) generally rises in individuals with ova-
ries during “mini-­puberty” and then drops to levels similar to
­those in individuals with testes from ages two to eight. During
­actual puberty, estradiol rises in all ­humans but is usually two
to three times higher in t­ hose with ovaries. As adults, t­ hose with
ovaries, and who menstruate, typically have cyclical increases
in estradiol during the follicular stages of the menstrual cycle.
Progesterone spikes in all h­ umans at birth and then drops to
low levels u­ ntil the age of seven or eight, when it rises moder-
ately in t­ hose with ovaries and remains stable in t­ hose with tes-
tes. In adulthood, levels of progesterone are about the same in
all h­ umans, except during the luteal phase of ­those who men-
struate, where it is up to twenty times higher, and during preg-
nancy, when it also increases. Testosterone levels are usually
about three to ten times higher in infants with testes than in
 H u m a ns Now 95

t­ hose with ovaries, but drop ­after mini-­puberty, and all h­ umans
have about the same testosterone levels u­ ntil true puberty. At
puberty, all ­humans’ testosterone rises, but ­those with testes
typically undergo a larger spike. As adults, most ­humans with
testes typically have circulating testosterone levels on average
about ten to e­ ighteen times higher than t­ hose with ovaries.
However, t­ here are a number of variants on this theme, both in
­humans with testes who produce very low amounts of testos-
terone and in variations such as PCOS (Polycystic ovary syn-
drome), which occurs in five to twenty ­percent of ­human adults
with two X chromosomes and often results in high levels of
androgens (especially testosterone) produced.52 As with all
hormones, the varying levels between bodies at dif­fer­ent points
in development and life history can have differing effects on
vari­ous aspects of the developing body, physiology, and be­hav­
ior. The patterns of hormone variation ­matter for ­human bod-
ies, but they are not binary in nature or form.
One ­can’t do a review of hormones without talking a bit
more about testosterone. “T” holds an almost mythical role in
the public view, and many think T is what “makes” a “man.” But
is that correct? T is prob­ably the most scientifically researched,
discussed, and debated, and certainly the most publicly misun-
derstood of the gonadal hormones. On the one hand, ­there are
researchers who believe that testosterone’s main job is to sup-
port the anatomy, physiology, and be­hav­ior that increases a
male’s reproductive output—­T helps them [males] reproduce
and direct energy to be used in ways that support competition
for mates, in short, that testosterone is the biological and
evolved “difference” between males and females.53 ­Others argue
that “T” is best understood in the context of a dynamic and
malleable set of relationships. ­These scholars suggest that T is
neither the creator of masculinity nor the essence of maleness,
 96 Chapter 5

but is involved in growth, muscle-­mass development, and re-

productive dynamics that produce key patterns of variation in
and between 3G bodies, and thus T is intricately related to the
gender/sex experience.54 ­These scholars also argue that T is
often responding to, not necessarily driving, be­hav­ior and
­physiology. The preponderance of research demonstrates that
testosterone action and effect is interlaced in and with biological,
social, and cultural contexts. A full understanding of this (and
other) hormone’s role and impacts is not simply volumetric
(how much is produced or is circulating), but has to do with the
dynamics of receptors, feedback loops, patterns and variations
of development, and a myriad of other physiological and social
­factors requiring an integrative, and nonbinary, approach.55
As for function, the entire current corpus of work on testos-
terone does offer some basic agreement: T is involved in as-
pects of muscle mass, aggression, and sexual behavior/desire in
many animals, including h­ umans—­and not just in t­ hose with
testes—­but, then again, so is estrogen.56 The distinctive evolu-
tionary histories of dif­fer­ent animal linages, like hyenas, naked
mole rats, and titi monkeys, shape the specific patterns and dy-
namics of T and other hormones related to their sex biology in
specific manners. Clearly, the distinctive aspects of h­ uman evo-
lution altered some of the dynamics of T in ­humans. While the
variance between testosterone levels, and some of its interac-
tions/effects, can be, on average, substantial between t­ hose
with testes and t­ hose with ovaries, within-3G sex correlation
between testosterone and muscles, sex and aggression are small
and often inconsistent. And, the patterns of T in relation to the
wide array of p­ eople who do not fit into 3G categories are l­ ittle
studied or understood. Moreover, the interactions of T and par-
enting be­hav­ior in ­humans are pronounced, which is not sur-
prising given the distinctive ­human evolutionary history
 H u m a ns Now 97

around the caretaking of young.57 At the end of the day, while

testosterone is an impor­tant hormone with substantive sex-­
biology-­related variation, it is not the marker of distinction be-
tween 3G categories or the “maker” of males or men.58 The idea
that testosterone—or any single hormone—is the biological
basis of a sex binary is not supported by the research and real­ity
of the complexities of the endocrine system.59
However, ­there is a suite of pro­cesses closely mediated by a
diversity of hormones, sex biology, and reproduction, which
occurs only in some ­human bodies and not in ­others.

Menstruation, Gestation, and Lactation:

Now H­ ere’s a Difference
Unlike all ­we’ve covered so far in this chapter, ­there are three
core pro­cesses that do mark clear, and distinct, biological dif-
ferences between h­ umans who have them and t­ hose who d­ on’t.
­These pro­cesses are central to mammalian reproduction and, as
usual, the ­human lineage has done some weird ­things with
them relative to other mammals.
Menstruating ­people and p­ eople who could menstruate
make up half of humanity and thus a core aspect of sex biol-
ogy.60 Ovulation in ­humans typically happens on a cyclical
basis in ­those with ovaries across a certain portion of the lifes-
pan, between menarche and menopause. The menstrual cycle
itself involves a range of physiological pro­cesses related to the
preparation of the bodies of ­those with ovaries and uteruses for
potential gamete fusion, pos­si­ble pregnancy, and subsequent
interactions with a zygote, embryo, and fetus. ­These pro­cesses
are regulated by the hypothalamic-­pituitary-­ovarian (HPO)
axis, specifically interactions between follicle-­stimulating
 98 Chapter 5

hormone and luteinizing hormone, progesterone, estradiol, and

cortisol (plus a few o­ thers). Unlike much common (and unfor-
tunately medical) belief, the menstrual cycle is not a closed
system isolated from other aspects of one’s body and life that is
more or less the same across every­one who menstruates. In-
stead, it is a p­ rocess that is impacted by and responds to many
­factors, including development, lived experience, age, nutri-
tional state, activity, and so on.61 The timing of first and last
menstrual periods, the concentrations of hormones, and the
thickness of the endometrium (the uterine lining) all vary with
and are particularly responsive to stressors from the social and
physical environment. As with so many key pro­cesses and traits
in ­human sex biology, ­there is not one “normal” menstrual cycle
or one “best” way for bodies to menstruate. And while only
about half of h­ umans can potentially menstruate, menstruation
is a biological p­ rocess, a cultural experience, and an impor­tant
part of the entire ­human gender/sex landscape.62 Even in an
aspect of biology that only exists in a half of h­ uman bodies,
an ­actual dimorphism, the dynamics of the ­process reject a sim-
plistic description of the ­human experience of sex biology.
If one has ovaries plus the associated fallopian tubes and
uterus, one can usually undergo pregnancy, which is the major
physiological ­process involving the complex of gestation and
birthing. ­Humans without a uterus are biologically unable per-
form this physiological p­ rocess. Along with menstruation, preg-
nancy is prob­ably the most substantive patterned difference
between many in the 3G category of female and all 3G males.
However, it is vital to note that not all ­humans categorized as
3G females can (or do) become pregnant, for a variety of rea-
sons, and some h­ umans who do not fit into the 3G categories
can, and do, experience pregnancy.63 ­There is also a range of
physiological variation, including liver, heart, endocrine, and
 H u m a ns Now 99

neurobiological changes, that can emerge between ­those indi-

viduals who have gestated and lactated and t­ hose who have
not.64 Given ­these patterns of variation, even in the experience
of pregnancy ­humans do not map to a ­simple binary.
Pregnancy often results in an infant, and lactational feeding
of infants is a significant aspect of ­human reproduction. While
breast tissue itself emerges in most h­ umans to some extent, lac-
tation is pos­si­ble due a specific ­process, postpuberty, typical in
3G-­female individuals. In the p­ rocess, estrogen, progesterone,
and a few other hormones help facilitate the development of
ducts, stroma, and glandular tissue with milk-­p roducing
­globules that mix with the adipose tissue on the upper chest
congregated around the nipples. In most h­ umans with lactation-­
capable breasts who become pregnant, changes in prolactin,
estrogen, and progesterone levels at the terminal end of
­pregnancy and ­after birth facilitate the maturity and activity of
the milk-­producing globules in the breast glandular tissue, en-
abling the production and expression of h­ uman milk. As noted
above, breasts appear in as much as 30 to 60 ­percent of 3G
males, but they are usually not capable of lactation.65 ­There are
rare cases of h­ uman 3G males exposed to high levels of prolac-
tin expressing milk-­like substances (galactorrhea—­not true
lactation). In some intersex individuals, lactation occurs, and it
has been successfully induced in some individuals with XY
twenty-­third chromosomes and some transwomen who have
under­gone progesterone/progestin hormone therapies.66 In-
triguingly, when one looks across the mammalian world, t­ here
are two species of fruit bat where 3G males do regularly lac-
tate67 (via mammary glands with milk-­producing globules) and
feed offspring with their milk. While ­there are clear and typical
patterns of variation for lactation (it’s usually only in t­ hose
­humans who give birth), the biological variation across our and
 100 Chapter 5

other species demonstrates that even in the key reproductive

dynamic of lactation ­there is no ­simple binary.
­There are two final sets of “internal” biology that deserve our
attention, places that are the basis for deep and power­ful as-
sumptions about sex biology: the brain and DNA.

Brains/Neurobiology
Minds are a core to how we perceive, interpret, and respond to
the world. And at the heart of the h­ uman mind is the h­ uman
brain. ­There are some patterns of variation between the genders
of men and ­women across ­human cultures in be­hav­ior, psycho-
logical illness, and the manners in which one perceives sights,
sounds, and other signals from the world. This is often assumed
to be due to biological “differences” in the brains of 3G males
and 3G females. And, as usual, t­ hose h­ umans who do not fit into
the 3G categories are largely ignored in this area of research.
Are ­there male and female brains? A group of researchers
reviewed hundreds of studies (conducted over more than thirty
years) of ­human brain imaging and physical postmortem brain
analyses. They discovered few patterned variants between 3G
categories despite the strong history of repeated claims of “sex
difference” in the h­ uman brain.68 As with much in h­ uman bod-
ies, 3G-­male brains are on average larger than 3G-­female brains
from birth and stabilize at about 11 ­percent larger as adults. This
size difference is often assumed to be distinct from the other
commonly reported, but largely inaccurate, “differences” in
brains: a higher white/gray m ­ atter ratio, more intra-­versus in-
terhemispheric connectivity, and higher regional cortical and
subcortical volumes in 3G males. ­There is also some reported
3G-­related variation in the connectome (a comprehensive map
of neural connections in the brain), and ­there are relatively
 H u m a ns Now 101

successful computer-­based predictions of 3G sex of any given

brain based on assessment of shapes and volumes of areas.
However, all t­ hese patterns are primarily correlated with size
differences, and they vary greatly across populations and indi-
viduals. Brain-­imaging studies fail to identify consistent repro-
ducible differences in how brains react to learning challenges
(called activation differences) between 3G categories (and be-
tween genders) in verbal, spatial, or emotion pro­cessing. The
largest recent overview of brain studies found only two small
areas that are pretty consistent in their pattern of variation not
due to overall brain size: subcortical structures called the amyg-
dala and putamen. Also, the INAH-3 nucleus in the anterior
hypothalamus appears to be relatively larger, on average, in 3G
males. Overall, when the totality of findings on structural varia-
tion in ­human brains are assessed, the 3G-­sex category explains
only about 1 ­p ercent of total variation. This means that
99 ­percent of the structural variation in brains across h­ umans is
explained by ­factors not connected to 3G categories. Being
“male” and “female” tells us very ­little about brain structure.
The conclusion of many researchers is that differences
emerging from 3G-­m ale/3G-­female structural brain
­comparisons are largely trivial and often population-­specific:
the ­structure of the ­human brain is not sexually dimorphic.69
This perspective and the overall data on brain structure and 3G
sex are best summarized by a group of fourteen neuroscientists
who state, “Although ­there are gender/sex differences in brain
and be­hav­ior, ­humans and ­human brains are comprised of
unique ‘mosaics’ of features, some more common in females
compared with males, some more common in males compared
with females, and some common in both females and males.
Our results demonstrate that regardless of the cause of ob-
served gender/sex differences in brain and be­hav­ior, h­ uman
 102 Chapter 5

brains cannot be categorized into two distinct classes: male

brain/female brain.”70
­There are researchers in the neurobiological arena who dis-
agree that brains cannot be classified into female and male ver-
sions, arguing that t­ here are 3G-­sex variants in brain structure
that are not due to average size differences. Shortly a­ fter the
publication of the large overview noted above, a group of other
neuroscientists published an analy­sis of age, 3G sex, and brain
allometry (relative brain size)71 using the UK Biobank dataset
(of about forty thousand brains of primarily [95 ­percent] white
British nationals) and a specific type of analy­sis software to ret-
rofit the brain images for analy­sis.72 This group found age differ-
ences and 3G-­sex differences and age-­by-­sex differences in the
volumes and shapes in older individuals (forty to sixty-­five years
of age).73 While all their m­ easures overlapped extensively across
3G categories, they did find some form of 3G-­sex differences in
67 ­percent of the relative size ­measurements, with 37 ­percent of
regions larger in 3G males and 30 ­percent in 3G females. They
also found that 49 ­percent of all brain volumes generally
­decreased with age, and 28 ­percent increased, although aging
effects varied across all categories of analyses. Importantly, the
graphs of the data from the two 3G categories provided in
the study show dramatically more overlap than the height graph
at the start of this chapter, demonstrating that variation between
all individuals in the dataset vastly outweighs the variation
­between 3G categories. Also, the main research group and sub-
sequent commentaries on this topic highlight the fact that ­there
are, to date, no known corollaries of be­hav­ior or function connected
with reported relative volume/shape differences. It is not clear
that any of ­these patterns do, or mean, anything at all.
In a commentary article published just a­ fter the UK Biobank
study, another group of neuroscientists agree that t­ here is key
 H u m a ns Now 103

variation but argue that the patterns represented are not dimor-
phic, but are a continuum, with 3G categories exhibiting mean
or variability differences (as with height, but much more over-
lap).74 In other words, even for researchers who think ­these
results demonstrate significant patterned variation, t­ here are
not two types of brain (male and female), but rather t­ here is
one range of brain function and 3G individuals are distributed
along it with extensive overlap and some small differentiation.
Interestingly, a study published a few years before on the same
UK Biobank dataset, but a smaller sample and dif­fer­ent meth-
odology, also found some patterns of variation by 3G category,
but they w ­ ere dif­fer­ent ones, and many w
­ ere greatly reduced or
dis­appeared with size corrections.75 Fi­nally, the researchers
from the large overview presented above redid their key com-
parisons to include the UK Biobank study and five o­ thers stud-
ies looking at the same dataset or using similar methods and
discovered that the results of all six varied in locations of varia-
tion, direction of the variation, and w ­ hether or not variation
occurred in a given location in the brain.76 The is no single, or
same, pattern of variation across all studies. The current data
and analyses demonstrate that ­there is ­little evidence for sub-
stantive structural differences across most of the brain, aside
from ­those related to overall size.
This is not to say that brains in 3G-­male and 3G-­female bod-
ies, t­ hose of p­ eople who do not fit into 3G categories, t­ hose of
men and ­women, and ­those of very masculine or very feminine
individuals, do not respond, or function, somewhat differently
in response to specific stimuli, contexts, and actions; some-
times they do.77 The fact that brain activity and the functioning
of vari­ous aspects of brains varies by life experience, nutritional
status, exposure to trauma, age, gender, degree of masculinity
and femininity, and a myriad of other sociocultural and
 104 Chapter 5

ecological f­ actors, has been long known. Brains are biocul-

tural.78 This dynamic of little-­to-no structural differences but a
wide range of activity and a diversity of ways that the brain re-
acts suggests that rather than focusing on a search for “differ-
ences” between 3G categories, an approach that examines
gender-­influenced patterns and dynamics using a nonbinary
frame is likely to be more beneficial and scientifically fruitful.79
Even t­ hose arguing for clear and consistent structural differ-
ences acknowledge that “males and females are (at the group-­
level) often exposed to systematically dif­fer­ent environments
across the lifespan” and that this gendered real­ity could have
systematic impacts on neurobiological activity and function.80
The current data are pretty clear: ­there are not two brain types
or even a continuous gradient from masculine to feminine;
rather, the h­ uman brain appears to be a multidimensional
“­mosaic” of countless brain attributes that differ in distinctive
patterns across individuals.81

DNA
I end this overview of con­temporary h­ uman sex biology with
the place most ­people want to start: DNA. Specifically, the first
“G” in 3G: the genes on the twenty-­third chromosome. This
chromosome pair comes in two forms: X and Y. Th ­ ese chromo-
somes are often called the “sex chromosomes” ­because they
contain many genes related to the development of gonads and
other ele­ments of the reproductive tract. But as the X and Y
chromosomes themselves are neither sufficient to “produce
sex” nor l­imited to directing the development of sex biology,
the very heart of the supposed binary of sex is neither a binary
nor just about sex biology.82 The X chromosome has about 155
million base pairs of DNA and about 900 genes, not all of which
 H u m a ns Now 105

are related to sex biology. The Y chromosome has about 59 mil-

lion base pairs and about 55 genes.83 Most h­ umans are e­ ither
XX or XY, but ­there are also many variations on this theme.
Usually in individuals with more than one X chromosome t­ here
is a p­ rocess of “X chromosome inactivation” where all but
one of the Xs is “shut down” in each cell (not always the same
X) so that its genes are not expressed. But this ­process is not
always 100 ­percent effective, which can lead to mosaic develop-
ment and a range of potential physiological effects.
Generally, if a h­ uman embryo has one X and one Y chromo-
some, genes on the Y chromosome “turn on” and facilitate the
development of testes and related physiology. If one has two Xs,
then the active genes facilitate the development of ovaries,
uterus, and related physiological patterns. Th
­ ese developmental
trajectories primarily target reproductive physiology, but also
have downstream effects on patterns of hormonal action, mus-
cle, and bone development.
But being XX or XY does not always correlate with the
­patterns just described.84 One can be XY and have a variation
in the activation of segments of DNA so that the specific genes
that facilitate testes development never turn on and/or their
protein products are differently made or transported. The same
is true for someone who is XX and their ovary-­related ge­ne­tic
sequences. Variations that depart from a 3G pattern of gonad
and genital development are rare, but as noted e­ arlier, they are
a consistent aspect of ­human biology. For example, the SRY
gene on the Y chromosome is assumed to be a TDF (testes
determining ­factor) and known to steer the developing embryo
­toward the formation of testes. But it can be pre­sent and not
“turn on” or sometimes can have been translocated to another
part of the genome. Th ­ ere are also a number of other genes
shared by all h­ umans, being on the X chromosome or one of
 106 Chapter 5

the twenty-­two autosomes, such as DAX1, SOX9, and Wnt4,

that all appear to have dosage effects (when ­there is too much
or too ­little of a gene product) that affect the patterns of gonad
and genital development.85 Variation in the activity of ­these
genes in embryos affects much of the variation in gonads and
genitals in 3G-­sex individuals and in ­those who do not map
perfectly into the 3G categories.
­There is a wide array of other variations on the dynamics of
the twenty-­third chromosome, such as the occurrence of XO
(no Y or second X), XXY, and XYY individuals. ­There is also a
variety of scenarios where physiological pro­cesses are impacted
by more than just gene action in the development of an indi-
vidual and facilitate a range of outcomes that challenge the 3G-­
sex category model, such as XX individuals with a penis, XY
individuals with a clitoris, vagina, and labia, and individuals
who are XX or XY but have mixed sex gonads, genitals, and
other variants. Overtly non-3G bodies make up a very small
proportion of all ­humans, but many more covert variations aris-
ing from ­these same dynamics give rise to a range of genital,
gonad, and endocrine structures and patterns that are outside
the typical 3G expectations but are still loosely classifiable into
3G categories.86 Such variation is understudied and poorly un-
derstood but is a recurrent aspect of h­ uman biological pro­
cesses and reflects the normal range of variation in reproductive
development.
Fi­nally, recent work on the ­human transcriptome (the initial
biochemical products of genes) demonstrates some m ­ easurable
variation in initial products by identical genes in dif­fer­ent tis-
sues of the bodies of 3G males and 3G females. However, the
end product or action that t­ hese genes are associated with is
almost always the same across all bodies. This is impor­tant
­because even when genes are ­d oing some t­ hings slightly
 H u m a ns Now 107

differently in 3G-­category bodies, they mostly end up with

the same outcome.87 ­There is, however, a specific subset of the
genes that act differently depending on which specific tissues
associated with 3G categories they are expressed in. For exam-
ple, t­ here are about 6,500 genes that show some 3G-­sex-­based
variants in their transcriptome action. About 6,000 of ­those
variants are related to expression in mammary tissue, and the ma-
jority of the rest are in reproductive related tissues, with a few
in adipose, skeletal, and muscle tissues—­all of the specific areas
where endocrine, growth, and structural variations affecting sex
biology are the most prominent—­and all ­measured in adults.88
This intriguing ge­ne­tic data suggest that ­there are some dy-
namic relationships between growth/development, endocrine
and reproductive physiology, gene expression, and the lived
experiences of h­ umans based on aspects of sex biology. Yet
again, even at the level of gene functions, ­there are dynamic and
variable effects and pro­cesses, but ­there is no sex binary.89

Normal?
The ­actual biological variation of any ­human trait comes with a
set of ideas about what constitutes a “normal” version of that
trait. This idea of “normal” is also almost always accompanied
by opinions about what that trait should be like. But this should
give us pause and makes us ask the question: what is “normal”
­human biology?90
The average ­measure in a range of distribution of a given trait
is called the “mean” of that range. It is not necessarily the best
or ideal form, nor is it necessarily the most functional, adaptive,
or even aesthetic. It is a statistical repre­sen­ta­tion of the sum of
the range of values in a given m­ easure divided by the total num-
ber of values in the dataset. In many of the traits related to sex
 108 Chapter 5

biology that vary across h­ uman bodies, the mean has a cultural
value, and it is thought to be “normal.” And ­people are judged
against it. Therefore, when asking about aspects of sex biology,
the range of variation and functions or lack of function as-
sociated with that range should be the focus, not necessarily the
mean or some subset of the typical distribution. Comparing
means of some traits, like height and hair, have far more cul-
tural meaning than biological significance. Other traits, such as
gonad type or specific hormone levels, can have equally salient
cultural and biological significance, but the two might not be
connected in a ­simple fashion. Still other traits, like muscle
strength, vary in their salience and importance depending on
what specific questions are being asked and what comparisons
are being made. In the real­ity of ­human sex biology, means,
ranges, and traits have both cultural and biological implications
that are almost always entangled.
Ultimately, ­there are some impor­tant patterns of variation
between 3G categories, but ­there is a much more overlap in
functions and outcomes than can be captured by the concept
of sexual dimorphism or “sex differences.”91 “Female” and
“male” do mean something biologically but are not two ­dif­f er­ent
kinds.92 X and Y chromosomes, gonads, pregnancy, circulating
hormone levels, ge­ne­tic pro­cesses, developmental dynamics,
musculature, hair growth, adipose deposition, and brain mor-
phology and function all impact the ­human experience. But
­these aspects of our biology, and their variation, are not best
understood as binary sex differences. H­ umans are biocultural
and have gender/sex. Variation is our norm, and our variation
does not reflect two kinds of ­human.
 6

No Biological ­Battle of the Sexes

in 1993, a book called Men Are from Mars, W ­ omen Are from
Venus hit the bestseller list and stayed ­there for 121 weeks. In it,
the author and relationship counselor John Gray claimed that
men, like the Roman god of war, Mars, are by nature aggressive
and violent, but protectors, and w ­ omen are emotive, coy, and
maternal like the goddess of love, Venus. He told the readers
that successful male-­female romantic relationships are based on
recognizing natu­ral (biological) differences in communication,
emotion, and be­hav­ior between men and ­women.1
Sound familiar?
Charles Darwin told us that “man is more courageous,
­pugnacious, and energetic than ­woman” with “more inventive
genius.”2 Angus Bateman claimed males are promiscuous and
females sexually passive, coy, and choosy, and E. O. Wilson told
us that the evolved, biological differences between men and
­women naturally led to universal male dominance.3 Too many
biologists and other kinds of scholars across the nineteenth and
twentieth centuries made ­these kinds of assertions. And ­these
assertions, just like t­ hose of John Gray, are wrong.4 We know
that 3G females are not necessarily sexually passive and that 3G
males are not necessarily more courageous or energetic, nor

109
 110 Chapter 6

have ­humans evolved for male dominance or two dif­fer­ent

kinds of sex personality. Yet, more than three ­decades ­later, Men
Are from Mars, ­Women Are from Venus is still selling tens of thou-
sands of copies, and some biologists still refer to sex as a binary,
imagining that a male/female split, via anisogamy, pits the sexes
against one another and explains much in the ­human experi-
ence. The commitment to the binary view and its “­battle of the
sexes” implications is deep and resilient. Given all we know, this
should not stand. So, yet again, let’s take a closer look.
It’s true that many species have sex-­biology patterns that lead
to marked distinctions in bodies and lives between sex cate-
gories. Take, for example, peafowl (peacocks and peahens).
Peacocks, the small-­gamete producers, have much larger bod-
ies, extravagant coloration, and enormous tail feathers, poten-
tially creating dif­fer­ent modes of movement, risk, and related
be­hav­ior relative to the peahen, the large-­gamete producers.
But h­ umans are nothing like peafowl. We are not a species with
dramatic, evolved patterns of sex difference.
The core premise of the ­battle of the sexes (in mammals) is
that in order to successfully reproduce, the 3G female is locked
into gestating and caring for infants and the 3G male is wired to
get as many copies of his DNA (via sperm) into as many 3G
females’ reproductive tracts as pos­si­ble. The classic anisogamy
position in biology asserts that t­ here is a vast disparity in
­energetic investment in gamete production and investment in
offspring between 3G males and 3G females that is related to
differences in reproductive organs and physiology. As a result,
­there is an evolution of distinctive anatomical, physiological,
neurobiological, and behavioral pro­cesses due to differential
evolutionary pressures on the sexes.
When it comes to h­ umans, the biology used to “demon-
strate” the validity of a ­battle of the sexes is body size and
 No Biol ogic a l Bat t l e of t h e Se x e s 111

strength (larger and stronger 3G males) and specific behavioral

and hormonal differences (especially more testosterone in 3G
males). For example, some researchers assert that “sex differ-
ences in muscle and strength are largely the result of sexual se­
lection for male-­male competitive traits, whereas females have
increased body fat (in place of muscle) due to natu­ral se­lection
for maternal investment capacity in the context of our unusually
large brains.”5 ­These researchers believe that evolution has
­shaped ­human bodies along two distinct lines of development,
producing 3G males geared to fight each other for females and
3G females ­shaped for raising young. Chapters 4 and 5 outlined
why t­ hese assumptions ­don’t map onto what we know about
­human evolution and how they oversimplify descriptions of the
existing variation in h­ uman biology. Along t­ hese same lines,
­others argue that it is specific patterns of circulating levels of
­testosterone that create massive 3G-­male and 3G-­female bodily
differences and that “T” even “pushes the psy­chol­ogy and be­hav­
ior of the sexes apart.”6 In the last chapter, we covered how this is
an oversimplified assignment of superpowers to one of the dy-
namic, and variable, aspects of the ­human endocrine system.7
So, as we see, ­there are scholars who argue that ­human 3G-­
category bodies and minds are deeply and evolutionarily dif­fer­
ent and that this “biological real­ity” places h­ uman men and
­women in direct conflict. And ­there is a large swath of the pub-
lic that believes this as well. However, scientific data and analy-
ses demonstrate that this sex-­conflict position is not true.

Of Minds, Means, and Be­hav­ior

The “females and males as very dif­fer­ent” camp often points to
cognition (“minds”) as the proof of their position. Every­one
knows men and ­women think differently, right? Over the past
 112 Chapter 6

few ­decades, massive studies called meta-­analyses8 reviewed pat-

terns in many of t­ hese cognitive variables, such as math, verbal,
and spatial-­ability skills, communication dynamics (verbal and
nonverbal), social and personality variables such as aggression,
negotiation, helping, sexuality, leadership, introversion/­
extroversion, general psychological well-­being, some motor
­be­hav­iors (throwing, balance, flexibility, ­etc.), and a few other
psychological states and be­hav­iors (moral reasoning, cheating
be­hav­ior, e­ tc.). Th
­ ese meta-­analyses involved data from more
than twenty thousand separate studies involving more than
twelve million participants.9 The results are clear: Across most
topic areas in psychological science, the difference in responses
and outcomes between males and females is small or very small.10
­Here “small” and “very small” are ­measures of how far apart
the means of massively overlapping variation are. That is, pretty
much every­thing being ­measured in ­these studies overlaps al-
most completely between 3G categories (usually based on self-­
reported genders), but the means of the distributions of the
­measured variables, when separated by 3G-­sex category or gen-
der, can be dif­fer­ent from one another. Think of the height ex-
ample from e­ arlier in the book: 78 ­percent of folks in the United
States are not identifiable to 3G sex simply by height, but the
means of the overlapping distributions of 3G-­male and 3G-­
female height are dif­fer­ent. So, on average, one can say 3G males
are taller than 3G females. But that might not tell you much at
all about any specific individual, or about height as a biological
characteristic, given the massive overlap between 3G categories.
In the “mind” studies, most of the differences between the means
are much, much smaller than in the height example. The differ-
ence between the means in ­these studies is assessed by the com-
mon statistical tool called the “cohens d” ­measure, which re-
flects how far apart the means in the overlapping distributions
 No Biol ogic a l Bat t l e of t h e Se x e s 113

of the ­measurements are in standardized units. So “small” sug-

gests that the means are very close to one another and “very
small” even closer (with almost 100 ­percent overlap). The
between-­gender differences in t­ hese huge meta-­analyses ­were
small in 46.1 ­percent of all cases and very small in 39.4 ­percent
of all cases. The largest and most recent metanalysis demon-
strated this same massive overlap, with about 84 ­percent of
mean differences being small and very small across most traits
examined.11 Again, we ­really ­aren’t so dif­fer­ent. And cognition
is certainly not binary.
While the vast majority of the traits ­measured in ­these stud-
ies overlap extensively, ­there are a few patterns worth noting.
Men (the self-­identified gender) consistently scored higher in
hand-­grip strength, sprinting, throwing velocity and throwing
distance, masturbation, views on casual sex, physical aggres-
sion, and ­mental rotation of objects. ­Women (the self-­identified
gender) scored higher on indirect aggression, agreeableness,
and smiling. Th­ ese patterns reflect the effects of variation in
muscle mass and body size, training, and patterns of encultura-
tion (gender). But none of them stand out as clear and unequiv-
ocal markers of evolved patterns of male-­female conflict.
The patterns of higher scores for casual sex and masturbation
in men often receive a lot of attention, and are used to argue for
an evolved 3G-­male reproductive strategy of “more sex, less in-
vestment.”12 However, self-­reporting on sexuality and sexual
experience is notoriously problematic (e.g., men overreport
and ­women underreport) due to the power­ful gendered expec-
tations and cultural constraints in the con­temporary world.13
Even if the reports w
­ ere mostly accurate and self-­identified men
do indeed have a greater interest in casual sex, more sex part-
ners (on average), and more frequent masturbation, it would
not signify that “man’s mind” differs from “­woman’s mind” due
 114 Chapter 6

to evolved patterns. One would need to have a comparative

sample without the con­temporary global cultural views on sex,
sexuality, and patriarchal social structures to control for such
variables. Also, from a purely biological/anatomical perspec-
tive, the wholly external location and the physical structure of
the penis and testes makes it easy to manually manipulate to
stimulate the sexual physiological response (e.g., masturba-
tion). While the clitoris is also external and highly responsive
to stimulation, across the h­ uman socie­ties where the studies
on sexuality are done, ­there are substantively dif­fer­ent beliefs,
expectations, and restrictions on sexuality, including masturba-
tion, for t­ hose with clitorises as opposed to t­ hose with penises.
This likely plays into the self-­reported differences in sexual
­be­hav­ior and self-­sex.
In mammals with hands, especially primates, one often sees
individuals with penises engage manual stimulation for its physi-
ological reward (primates masturbate a lot). The increased
­frequency of reported masturbation in ­humans with external
penises and testes is at least as likely a by-­product of the combi-
nation of primate hands and genital anatomy in the context of
specific h­ uman cultural systems, as it is a reflection of an evolved
difference in mind and body between ­those with and without
the penis/external testes combo. Similarly, the lower self-­
reported rates for h­ uman females may not be due to any biologi-
cal difference, but rather to cultural ones. In many primate spe-
cies (including ­humans), females do masturbate, and do so more
than in most other mammals. Female primates are also observed
to use objects to masturbate. In fact, research on monkeys and
apes demonstrates that “females exhibit more sophisticated
kinds of masturbatory behaviours than males.”14 Primates use
sexual be­hav­ior outside of reproduction a lot, and masturbation
is part of that. ­Humans are typical primates in this re­spect (see
 No Biol ogic a l Bat t l e of t h e Se x e s 115

below); it’s just that for us sexual be­hav­ior also has substantive
cultural connotations and consequences. And t­ hese cultural
­realties in much of the con­temporary world often restrict the
expression of sexuality by t­ hose classified as female. Thus, mea-
sures of sexual be­hav­ior need to take that into account. As the
biologist Anne Fausto-­Sterling has eloquently and repeatedly
demonstrated, “Sexuality is a somatic fact created by cultural ef-
fect.”15 In other words, sexuality is not strictly programmed by
our genes, or genitals, but rather emerges in concert with the
gender/sex experience of ­humans as they develop.
­Actual data on sexual activity are difficult to collect and
quantify, but one can look to the 2010 National Survey of Sexual
Health and Be­hav­ior, a US-­based study of 5,865 adolescents and
adults (2,936 self-­identified men [gender] and 2,929 self-­
identified w­ omen [gender] ages fourteen to ninety-­four) to get
a general idea.16 In the study, 55 ­percent of the men reported
masturbation in the past month, and 71 ­percent in the last year;
31 ­percent of the w ­ omen reported masturbation in the last
month, and 54 ­percent in the last year, except t­ hose over sev-
enty. 85 ­percent of men in their twenties and thirties reported
having vaginal intercourse in the last year, compared to
74 ­percent in their forties, 58 ­percent in their fifties, 54 ­percent
in their sixties, and 43 ­percent in their seventies. For the ­women,
81 ­percent in their twenties and thirties reported having vaginal
intercourse in the last year, compared to 70 ­percent in their for-
ties, 51 ­percent in their fifties, 42 ­percent in their sixties, and
22 ­percent in their seventies. Men and ­women of all age groups
reported engaging in oral sex and masturbation with a partner.
For both oral sex and partnered masturbation, the man/woman
pattern is almost identical. More than 20 ­percent of the men
between ages twenty-­five to twenty-­nine reported anal sex, with
youn­ger and older men reporting much lower numbers. More
 116 Chapter 6

than 40 ­percent of the men e­ ighteen to fifty-­nine years old

reported participating in anal sex during their lifetimes. W ­ omen
are almost identical with slightly higher frequencies of anal
sex over a larger age range (­e ighteen to sixty-­n ine) than
males. Across all age categories, 8–10 ­percent of the men and
5–9 ­percent of w ­ omen reported same-­sex sexual activity during
their lifetime with much higher figures (up to 17 ­percent) for
the twenty to thirty-­nine age group. The survey did not ask
about sexual orientation, so it’s unclear what percentage of
­these numbers reflect homosexually oriented individuals as op-
posed to heterosexual or bisexual persons engaging in same-­sex
sexual be­hav­ior.17
While this is just one example, a plethora of studies of ­human
sexual be­hav­ior demonstrate that much ­human sexual activity
is outside of reproductive possibilities; h­ umans have sex across
a wide range of contexts and in multiple manners and that t­ here
is a lot of variation across individuals in sexual activity, orienta-
tions, desires, and practices.18 One pattern that emerges is the
overall decline in sexual activity with age (especially over sixty).
However, ­women report a higher decline than men. This out-
come may be related to physiological changes associated with
the sex biology of menopause.19 It also might be related to pat-
terns of cultural heteronormativity and patriarchy that encour-
age older men/youn­ger ­women pairings. Given every­thing
known about h­ uman sexuality, it is most certainly biocultural.
Overall, the data show few if any consistent patterned differ-
ences between ­human men and ­women in ­actual sexual activity.
Regardless, many still argue that the real differences between
sexes are not in a­ ctual sexual activity but the patterns of interest
in sex as s­ haped by evolved patterns of bodies and lives based
on assumptions about anisogamy (­whether you produce small
gametes or large gametes, ­etc.). This pattern of differential
 No Biol ogic a l Bat t l e of t h e Se x e s 117

interest in sex is assessed via a person’s “sociosexual o­ rientation,”

which is ­measured via the Sociosexual Orientation Inventory
(SOI). The basic argument is that t­ here is a pervasive pattern of
evolved differences between males and females, and that this
can be seen in their attitudes about sex in general, sexual fantasy,
and sexual be­hav­ior.20
In general, men (the gender) tend to score higher than
­women (the gender) on the SOI. Men also tend to report
higher interest in sexual activity and in sexual fantasies, higher
numbers of preferred or ­actual sexual partners, and prefer
short-­term versus long-­term sexual opportunities (on average).
­There is also a robust degree of self-­reported variation cross-­
culturally (across genders and 3G categories) in SOI-­related
patterns.21 However, ­there are no neurobiological patterns or
aspects of reproductive physiology that indicate such patterns
come from specific bodily variation in 3G males and 3G fe-
males. Even testosterone, which plays an impor­tant role in
sexual arousal in our species, does so in all h­ umans not more or
less in one 3G sex. Th ­ ere is a set of well-­studied physiological
links between testosterone-­level fluctuation, sexual activity,
sexual bonding, and parenting in some groups of 3G males.22
Unfortunately, ­there is not nearly as much study of the specific
dynamics of T in 3G-­female sexual activity and sociosexuality
as t­ here is in 3G males, and almost no study of this at all in
­people who do not neatly fit into 3G categories, so truly scien-
tific comparison is difficult.23 Given the substantive average
variation in circulating T levels between 3G females and 3G
males, and the on-­average patterns of SOI difference, one might
think that the assertion that higher T in 3G males is related to
(but not uniquely causal of) the between-­category variation
reported in sociosexual orientation is accurate. However, it is
pos­si­ble that circulating T i­ sn’t the right ­measure to assess with
 118 Chapter 6

regards to sociosexuality. Above a low threshold, t­ here’s l­ ittle to

no increase in libido with an increase in circulating T among
most 3G males, and smaller increments of T seem to have libido
effects in 3G females. So, rather than simply circulating T as the
prime agent that underlies libido, it might be T-­receptor varia-
tion, and a range of ele­ments related to gender and cultural
­dynamics also in play.24 The main researchers who study socio-
sexuality argue that T is not the only or even the primary ­factor,
stating that “the most consistent finding was that men scored
higher than w ­ omen on sociosexuality across cultures. Several
dif­fer­ent theories w ­ ere evaluated concerning why men and
­women differ in this way. They all received at least some empiri-
cal support. As a result, we are left with the relatively unsatisfy-
ing conclusion that sociosexual sex differences are predictable
from several theoretical perspectives, none of which is con-
spicuously superior to the o­ thers. . . . ​At pre­sent, it appears that
multiple perspectives are required to more fully explain the
cultural and gender-­linked variance in sociosexuality.”25
­There is some gender/sex-­related variation in self-­reported
perspectives on sexuality, but this variation is not simply a re-
flection of an “evolved difference” stemming from gamete sizes
and reproductive tracts. ­Humans have a lot of sex (more than
most other animals), and ­there are relatively few, if any, signifi-
cant biological features or variants that constantly differ
­between 3G males and 3G females in relation to kinds and pat-
terns of ­actual sexual activity. ­Humans are biocultural creatures,
and the social and cultural context in which we develop is g­ oing
to have enormous influence on something as complex as our
self-­reported perceptions of sex and sexuality.26
Aggression is another area where patterns of variation
emerge. It is also an area that has been held up as an indicator
of evolved male-­female conflict.27 On average, 3G males have a
 No Biol ogic a l Bat t l e of t h e Se x e s 119

larger body size and greater upper-­body muscle mass and there-
fore can pose a potentially greater risk of harm in purely physi-
cal fights than do smaller 3G females (assuming no training in
hand-­to-­hand combat). Across many socie­ties, young adult
men (the gender), on average, tend to participate in some cul-
turally defined categories of physical risk-­taking be­hav­ior at
higher rates than young adult w ­ omen (the gender).28 In many
socie­ties, men engage in overall higher rates of physical aggres-
sion than w ­ omen, especially in conflicts with other men. But in
aggression between heterosexual partners t­ here is very l­ittle
difference in the use of physical aggression, with ­women being
slightly more likely to use it.29 However, in this same context
(heterosexual partnerships/pairs), physical aggression by men
often ­causes more injury than that by ­women. This is not sur-
prising given that 3G males are on average slightly larger than
the ­woman they are fighting with, and often have a greater
upper-­body strength due to both physiological and cultural pat-
terns, and are likely to have more previous experience in physi-
cal fighting. This outcome is a biocultural blend of patterns, not
a smoking gun for evolved differences in physical vio­lence be-
tween 3G categories.
­There is a par­tic­u­lar consistency in certain reported patterns
in aggression and vio­lence, especially at the level of hom­i­cide
and sexual assault/rape, when using the 3G-­male/3G-­female
comparison.30 Some have argued that this is related to the
­differences in circulating testosterone and/or the evolution of
vio­lence and aggression as a basal adaptation of 3G-­male be­hav­
ior. However, t­ here is no solid evidence from the h­ uman evolu-
tionary rec­ord or ­human physiology to support such an asser-
tion.31 While 3G males are indeed the initiators of the vast
majority of hom­i­cide and sexual assault/rape in our species, it
is also true that most 3G males are never involved in such acts.32
 120 Chapter 6

Hom­i­cide and sexual assault/rape are, in most cases, not driven

by reproductive outcomes but are acts of social vio­lence, power,
and control. Acts not reflective of specific evolved sex-­biology
patterns or traits but rather events catalyzed, or facilitated, by
cultural histories, dynamics, and structures in the context of
­human gender/sex.33
Do patterns of sexual vio­lence and hom­ic­ ide tell us some-
thing about 3G males evolving to be more aggressive than 3G
females? Not r­ eally. Men (the gender) appear to be more ag-
gressive in some contexts and ­women (the gender) in ­others.
Men’s physical aggression and men’s participation in antisocial
aggression (e.g., violent crime) are more common and often
more serious than t­ hose of w ­ omen. But the majority of all
­humans (men and ­women) do not engage in violent aggression,
or crime, with any regularity, or even at all. Overviews of pat-
terns in con­temporary ­human socie­ties, and of ­human socie­ties
of the past, and in-­depth studies of h­ uman bodies in the past
and ­today (chapters 4 and 5) strongly suggest that while ­there
are gendered patterns of be­hav­ior that ­matter, ­there is no evi-
dence for an evolved pattern of specific 3G-­male hyperaggres-
sion as a key adaption in ­humans.34
As for brains and minds, many researchers conclude35 that
male/female physical/structural brain differences are largely
trivial and often population-­specific: the structure of the h­ uman
brain is not sexually dimorphic, and while some features are
more common in 3G females than in 3G males and vice versa,
­there are no “male brains” and “female brains.”36 ­There is not
even a continuous gradient from men to ­women or masculine to
feminine; rather, the ­human brain is a multidimensional mosaic
of myriad attributes that differ in unique patterns across indi-
viduals.37 ­There is patterned variation in the ­measured means for
some ­mental states and cognitive be­hav­iors in ­humans, but ­these
 No Biol ogic a l Bat t l e of t h e Se x e s 121

are almost always small or very small and usually in the midst of
massively overlapping variation. A focus on means obscures the
likely more biologically relevant realities and dynamics of varia-
tion in h­ uman bodies and minds. If one seeks to validate as-
sumptions about difference by looking only for difference, then
that is all one sees. The patterns of variation in a forest are often
more impor­tant than the average height of certain types of trees.
The same goes for ­humans and ­human socie­ties.
When the ­great overlap (and complexity) in sexuality, ag-
gression, and cognition fails to support the ­battle of the sexes
argument, its proponents often turn to the costs of reproduc-
tion as a trump card. But as we already know, scientific evidence
about h­ uman reproduction and the evolution of h­ uman care-
taking systems rejects the ­simple “large-­gamete producers
­versus small-­gamete producers” view of the world. To summa-
rize: parental investment in ­humans is more complex than one
3G male and one 3G female investing in reproduction. ­There is
massive evidence that the genus Homo evolved complex coop-
erative caretaking of young early in the Pleistocene and that by
the mid-­Pleistocene ­human childhoods had extended substan-
tially b­ ecause of it.38 The evolutionary implication of this real­ity
rejects a s­ imple calculus based on individual 3G females’ costs
of gestation and lactation and includes a caretaking regime
where the ­mother is one of many caretakers and the range of the
specific infant caretaking energetic burdens is redistributed
across the group. This is not to say that the “costs” of reproduc-
tion are not high in the genus Homo, but rather that the costs
have, in an evolutionary response, been socially and physiologi-
cally mediated and spread across group members to enable the
complex, extended childhood associated with a range of capaci-
ties connected to substantial neurobiological and social devel-
opment ­after birth.
 122 Chapter 6

The nuclear f­ amily structure is often brought up as evidence

for some aspect of the conflict between or severely dif­fer­ent
roles for 3G males and 3G females (male the provider and fe-
male the caretaker). The assumption is that the core nature of
the nuclear f­ amily deep in h­ uman evolutionary history struc-
tures both Homo bodies and socie­ties. However, as we reviewed
in chapter 4, ­there is ­little fossil or comparative evidence of two-­
adults-­plus-­offspring as the core social structure for Homo. That
is, existing fossil and archeological data do not offer clarity re-
garding group structure aside from the high likelihood of Homo
groups consisting of multiple adult females and males and
young. ­W hether the within-­group structure consisted of mul-
tiple bonded pairs is not detectable, but having pair bonding as
a common facet of Homo social ­organization is likely given the
near-­universal ­human social be­hav­ior pattern of pair bonding
and its physiological manifestations. Pair bonding does not
equal monogamy or two-­adult exclusive mating or a specific
two-­adult caretaking pattern; it is not equal to or indicative of
a nuclear f­ amily structure.39 In other words, the current fossil,
comparative, and con­temporary data offer no support for the
nuclear ­family as the “traditional” or core evolutionary basis of
the h­ uman ­family or social group.40

Man ≠ Male and ­Woman ≠ Female:

Beyond the B­ attle of the Sexes
The “­battle of the sexes” assumes that man is interchangeable
with male, as is ­woman with female. But this is not necessarily
the case. Yes, 3G males and 3G females are typical clusters of
genes, gonads, and genitals, and they vary in some patterned ways,
but one’s 3G state, typical or other­wise, does not uniformly tell
 No Biol ogic a l Bat t l e of t h e Se x e s 123

us what one’s gender identity, one’s degree of masculinity or

femininity, one’s sexuality, or one’s sense of self in regard to re-
productive roles is or ­will be. And ­these 3G categories often tell
us even less about one’s other be­hav­iors. Also, ­there is a cluster
of humanity who are not easily classifiable into 3G categories
and who need to be included in all t­ hese analyses if we wish to
cover the full biological range of being h­ uman. As a collection
of researchers in the areas of sex and gender r­ ecently noted,
“Gender/sex is multidimensional and each component is
­dynamic and responsive to both internal forces (biological,
cognitive) and external forces (social interactions, culture). In-
dividuals show variability across the dif­fer­ent components of
gender/sex, presenting a mosaic of biological and psychologi-
cal characteristics that may not all align in a single category of
the gender binary.”41
Given every­thing ­we’ve covered so far, it’s clear that the
­human “sexes” are not just biology or just culture, they are not
“the same,” but nor are they “dif­fer­ent kinds.” Rather than envi-
sioning typical reproductive physiologies, bodies, and minds in
competition with one another (the “­battle” of the sexes), the
goal should be to move beyond assumptions about conflict be-
tween and uniformity of “sex” as categories and shift ­toward
examining the ­actual patterns and variations in Homo sapiens
and allowing the information emerging from that ­process to
structure the inquiry. Male/female, man/woman, small ­gamete/
large gamete, ovaries/testes, penis/vagina, and so on are not
necessarily default comparisons to use when thinking about
­people. Of course, such comparisons can be appropriate for
many questions, but we need to be open to letting ­actual behav-
ioral, physiological, ge­ne­tic, cultural, and historical data and
variation drive the modes of categorization for our investiga-
tions, not cultural assumptions about what sex “­really” means.
 124 Chapter 6

To be clear, ­there are patterned variations in sex biology in

­human bodies, and they contribute to the shaping of p­ eoples’
lives. But t­ hose patterns are not ubiquitous. They are seldom
truly dimorphic, almost never binary, and do not always or use-
fully match to the categories of man or w ­ oman. In fact, close
attention to the a­ ctual variation in caretaking, sexuality, gender
identity, parental investment, aggression, and other be­hav­iors
might suggest other ways to cluster ­humans as the targets of the
research focus and thus frame the question(s) differently.
Man is not more courageous, pugnacious, energetic, or inven-
tive than ­woman. Men are not biologically more promiscuous
than ­women, who are in turn passive and coy. Evolved, biologi-
cal differences between men and ­women do not naturally lead
to universal male dominance. And certainly, men are not from
Mars and ­women are not from Venus. We are all the same spe-
cies from the same planet. However, this does not mean that
­there are not conflicts between gendered expectations and ex-
periences, that ­there is no oppression and exclusion between
and within genders, especially in the context of patriarchal sys-
tems, or that reproduction, child care, and patterns of vio­lence
are equitably spread across all members of our species. What is
impor­tant to understand is that the vast majority of the patterns
and pro­cesses related to gender/sex that ­humans experience
­today are not biologically predetermined, evolutionary or-
dained, or inevitable. They are biocultural, created by the en-
tanglements of h­ uman bodies, histories, lives and cultures. And
thus they are not necessarily fixed or permanent aspects of the
­human experience.
 7

Why the Binary

View Is a Prob­lem

t­ oday, the state of biological knowledge is one that “ac-

knowledges sex is a rich, vari­ous, and diverse phenomenon that
can—­and should—be ­measured across multiple levels of bio-
logical ­organization and can be variable within an individual,
within a species, and across dif­fer­ent species. The study of sex
diversity and variability in the animal kingdom has been hin-
dered by imposing binary assumptions and limitations on what
sex is, or can be, across species. By simply acknowledging that
sex can, and does, exist outside a strict binary framework, we
can evolve and improve how we define, ­measure, and analyze
‘sex’ in our research.”1
Despite this knowledge, a binary perspective remains perva-
sive. Understanding and communicating the realities of ­human
sex biology is not just an academic or philosophical undertak-
ing; allegiance to the binary view in industry, government,
­education, and by the public fosters ignorance, harm, and suf-
fering. ­Humans are always immersed in complex, variable, and
biocultural gender/sex realities. Variation in ­human bodies,
cultures, and lives illustrates that “average,” “ideal,” “typical,”

125
 126 Chapter 7

and “binary” are seldom the most effective ways to talk or think
about h­ uman gender/sex.2 This knowledge is power­f ul, but
only if it is shared and used. In that spirit, this final chapter of-
fers brief examples of a few areas in con­temporary society
where binary views of sex biology and gender are invoked, and
it illustrates why such actions are both wrong and harmful.

Sexuality and Sexual Orientation

Back in 1991, a neuroscientist named Simon LeVay argued that
a certain brain region, called the INAH-3, was less than half the
size in gay men than in straight men, and that gay men and
straight˝ ­women had INAH-3 of about the same size.3 He be-
lieved he’d found the biological marker of being gay, in men.
LeVay worried what society would do with the knowledge that
“gayness” is located biologically. In discussing the possibility of
a “gay” gene, he said, “We are ­going to have this power over our
own natures, particularly over our ­children’s natures. What we
do with ­these choices ­will be one of the major ethical questions
of this ­century. Are parents g­ oing to have f­ ree reign, or should
society step in?”4 Luckily, ­there ­will be no manipulation of em-
bryos for sexual orientation, as subsequent research demon-
strated that t­ here is no “gay” area of the brain, nor are t­ here
“gay” genes. In fact, recent work reveals an incredibly complex
and dynamic suite of ge­ne­tic associations with sexuality and
sexual orientation and demonstrates that the neurobiological,
behavioral, and social relations to sexuality are equally com-
plex.5 The quest for a “gay brain” or a “gay gene” stems from the
belief of a “natu­ral” sex binary tied to heterosexual reproduction
such that any significant deviation from that pattern, like homo­
sexuality, must come from a biological alteration or aberration.
But real­ity is far more complicated. Of course, t­ here are biological
 W h y t h e B i n a r y V i e w I s a P r ob l e m 127

components to sexuality and sexual orientation, but ­these are

not simply tied to a specific aspect of sex biology.
­There is an enormous lit­er­a­ture examining and debating
which biological and social categories or variants m ­ atter most
for sexuality and sexual orientation, with ­little agreement aside
from the fact that it is complicated.6 The state-­of-­the-­art under-
standing is that ­there is no consistent, or effective, sole predictor
or single determinant for sexuality and sexual orientation in
­humans: not one’s gender, nor one’s genitals, nor one’s gonads
or hormones, nor one’s genes or brain structures. Many ­people
assume that heterosexuality is the “natu­ral” mode of sexual be­
hav­ior in h­ umans and that other modes of sexuality, homo­
sexuality, bisexuality, asexuality, and so on are ­either aberrations
or deviations from the typical pattern. Such an assumption is a
misleading way to discuss h­ uman sexuality. The physiological
system of arousal is almost identical across all ­humans (regard-
less of what kinds of genitals are in play7), but the psychological,
social, and emotional triggers for such arousal vary widely, are
related to gendered identities, and are not solely, or even pri-
marily, based on the specific variants of sex biology one has.
­There is variation between, and within, gender and sex cat-
egories in sexuality. The sociosexuality index (SOI), initiation
of sexual vio­lence, self-­reporting of masturbation, and a range
of other behavioral patterns vary between and within genders
and 3G sexes. For example, we noted in the last chapter that
decline in interest in sexual activity seems to be affected by
variations in sex biology related to age, relationship longevity,
gender, and sexuality. Th ­ ere is a general decline in interest in
sexual activity with age (especially over sixty years) in h­ umans,
but the decline for self-­identified ­women is larger. This trend is
especially acute in married or long-­term ­couples (both hetero-­
and homosexual), where ­women’s participation in sexual
 128 Chapter 7

activity with their partners is negatively correlated with the

length of time together.8 This difference might be influenced by
physiological changes affecting genitals and physiologies in
bodies with uteri and ovaries during menopause combined
with other social and psychological dynamics of aging. It also
may be influenced by cultural structures and patterns, such as
patterned partner-­age differences, expected sexual roles and be­
hav­ior, societal views of beauty, bodies, and sexuality, and a
range of structured gender inequities in con­temporary socie­
ties.9 As with most t­ hings, t­ here is not a uniform pattern; rather,
­there is only an “on average” one, and it is surely biocultural—­
disentangling the role of sex-­biology variation from the cultural
and gender context might not be pos­si­ble in this case (or even
in most cases of sexuality).
Sexual be­hav­ior varies substantially across many animal spe-
cies, especially social mammals. In primates, sexual be­hav­ior of
all kinds is more common in social contexts than in purely re-
productive ones. That is not to say that reproductive sexual ac-
tivity, which is by definition heterosexual in mammals and birds
but not necessarily so in other animals (remember worm sex
biology), is not common—­indeed, it is typical. But the key
point ­here is that the range of sexual activity, and pair-­bonding,
in ­humans is not necessarily constrained by (or even related to)
the necessities of reproduction. ­Humans exhibit the most di-
versity and highest frequency of sexual be­hav­ior of the primates
(except for perhaps bonobos), and all h­ uman cultures demon-
strate a wide range of sexual be­hav­iors and sexualities, with
varying patterns of cultural restrictions and constraints. So, for
­humans the accurate characterization of sexuality and sexual
be­hav­ior is that it is diverse, largely social, and often occurs
across multiple partner combinations across ­human lifetimes
in the context of a range of cultural rules, expectations, and
 W h y t h e B i n a r y V i e w I s a P r ob l e m 129

institutions. One cannot scientifically look exclusively to sex

biology or any biological specifics to fully understand any given
­human’s sexuality and sexual orientation. Representing ­human
sexuality as a binary system (men are one way and w ­ omen are
another, male/female, gay/straight, e­ tc.) sets up erroneous and
scientifically unsupported understandings and expectations
regarding sexuality and sexual orientation. The binary approach
tells us very l­ittle about h­ uman experience, and it likely ob-
structs effective and impor­tant research into sexuality.

Making a ­Family
In 2020, the Republican governor of the US state of Tennessee,
Bill Lee, signed a bill to ensure that foster care and adoption
agencies could exclude LGBTQ families and o­ thers based on
“religious beliefs.”10 By 2024, ­there ­were at least thirteen states
in the United States that had similar laws.11 The laws are based
on false beliefs about what is biologically “natu­ral,” and there-
fore “right” for ­humans. ­These laws assume that only hetero-
sexual pairings of certain types of p­ eople (cisgender) provide
appropriate and successful conditions for the rearing of
­children. This is factually incorrect, as the c­ hildren of LGBTQ
parents fare just as well as c­ hildren of non-­LGBTQ parents.12
But t­ hese beliefs and real l­ egal structures are not about the data
or facts; rather, they reflect a deep commitment to the logic of
the sex-­biology binary: if female bodies are made for repro-
duction and caretaking, and male bodies for protection and
­provisioning, then the biological (natu­ral) basis for f­ amily is a
heterosexual male and female ­couple.
The core of mammalian reproduction is gametic fusion in a
body capable of gestating and birthing. But the necessary bio-
logical details for the creation and gestating of an embryo tell
 130 Chapter 7

us only a small slice of the dynamics of a h­ uman f­ amily. The

­human f­ amily is not l­imited to or by the physiology of repro-
duction. ­Humans create and nurture families, communities of
care, wherein adults, and o­ thers, collaborate to care for each
other and young. But families come in all shapes and sizes. The
idea of the heterosexual nuclear ­family as basal to humanity is
refuted by a range of scholarship.13
­Humans with uteri play a central, biological role in the cre-
ation of offspring (gestation and lactation), and thus are often
at the heart of the ­human ­family experience. Birth ­mothers can
technically undertake much of the reproductive ­process alone,
but historical, social, and biological features of humanity indi-
cate that the ­human system of reproduction and caretaking has
evolved to be a multi-­individual ­process. ­Humans with uteri are
center stage regarding gestation and birthing, but human-­wide
physiology is structured to enable caretaking of infants and
strong bonding by individuals across age groups regardless of
which variants of sex biology one has. Successful rearing
of ­children and the formation of communities of bonding
and care (families) occur in a variety of ways across a variety of
­humans. Unfortunately, ­there are some con­temporary socie­ties
with structures, and laws, as noted above, that seek to inhibit or
limit the diversity and possibilities in ­human patterns of ­family
formation, community care, and childrearing by mandating
that only one combination of ­humans (one 3G male and one
3G female) is the correct nucleus of a f­ amily. Given what we
know about sex-­biology variation, pair-­bonding, ­family forma-
tion, and how the caretaking system in h­ umans evolved, t­ hese
laws, and beliefs, place a wide range of p­ eople, especially the
­children who benefit from the diversity and commitment of
caretakers, at substantive risk. ­Human families are not neces-
sarily composed of only biological kin, are not necessarily based
 W h y t h e B i n a r y V i e w I s a P r ob l e m 131

on the u­ nion of two cisgender dif­fer­ent 3G-­sex adults, and they

are not only about reproduction. Families are complex relations
and networks of bonding, caretaking, and collaborating be-
tween socially close individuals. They are not the constrained
product of, or necessarily related to, a sex binary.

Medicine
Historically in the medical sciences, researchers ­were often
hesitant to use female animals in laboratory studies. Male lab
animals (mostly mice) outnumber female ones by a ­factor of at
least five, especially in tests focused on obtaining species-­wide
results.14 Traditionally, medical researchers considered female
animals, including ­humans, too behaviorally and hormonally
unstable and physiologically complicated to give generalizable
results in many cases.15 In the medical binary view of sex biol-
ogy, ­there are two types in e­ very species, the male and the fe-
male. The male is the standard, the stable specimen, b­ ecause his
only contribution to reproduction is sperm. The female, with
her instability, is s­ haped by her possession of a uterus and her
reproductive cycling; thus approaches to her biology, and
health, are centered around her capacity to reproduce.16
And that capacity is thought to involve messy hormones, be­
hav­ior, and physiology, rendering the female too complicated
and a poor subject for general assessment of nonreproductive-­
related aspects of biology, health, and well-­being. ­Because of
this belief, many experiments, or rather experimenters, assume
that how something affects a 3G male (be it a mouse or h­ uman)
is the best proxy for the rest of the species. This approach as-
sumes minimal relevant variation among 3G males and too
much variation among 3G-­female bodies. This belief has kept
female animals, and animals who do not fit neatly into 3G-­male
 132 Chapter 7

categories, out of much laboratory research.17 For the last

­century, based on the binary view, most of the medical testing
world18 saw male mice (and male ­humans) as the ideal baseline,
the “normal” for understanding bodies. This commitment to a
­simple reproductive physiology binary as a key ­factor when it
comes to researching health and medicine is not supported by
current understanding of ­human biological variation and has
resulted in medical science ­doing outsized harm to ­women.19
To be clear, rejecting a binary view of sex biology in medical
research and treatment does not mean ignoring patterns of
variation across and between bodies, including ­those associated
with uteri, gonads, genes, and genitals.20 ­There are critical pat-
terns of physiological variation related to sex biology, and they
often ­matter for issues of health. However, to assume that the
most basic dividing line for h­ umans is between two typical sets
of reproductive physiology, or that the optimal approach in
medical science is to minimize variation in test subjects by
avoiding complexity, has negative outcomes.
Take drug reactions. Most reports assert that w ­ omen (gener-
ally meaning 3G females)21 are about 1.5–2 times as likely as
men (generally meaning 3G males) to experience severe side
effects when using pharmacological treatments. ­Women also
may not derive the same benefits from many drugs as do men
(on average). This is usually assumed to be ­because of differ-
ences in 3G-­male and 3G-­female biologies. But is that the case?
If one restricts comparisons to 3G categories, most of the
documented variation in effects of drugs on 3G males and 3G
females is not based solely on body size. Th ­ ere are other physi-
ological pro­cesses at play, possibly related to aspects of sex biol-
ogy. One might take this fact at face value and argue that this
reinforces a binary view. It does not. One aspect likely affecting
­these outcomes is that most of the dosages for drugs ­were, up
 W h y t h e B i n a r y V i e w I s a P r ob l e m 133

u­ ntil recently, largely based on the clinical testing done solely,

or primarily, in men.22 Another aspect is that the percentage of
­women’s adverse side effects may be overestimated ­because
­women (the gender) use prescription drugs more than men
(the gender) and the databases are not adjusted for ­actual drug-­
use rates by men and ­women (by 3G-­sex category).23 A third
explanation for pos­si­ble differences in outcomes is that “men”
and “­women” are not discrete biological categories, so testing
phar­ma­ceu­ti­cals in the categories “men” and “­women” is not
effectively assessing the range of ­human biological variation
(the ­actual p­ eople) taking the drugs. Fi­nally, h­ uman bodies and
their physiologies are ­shaped by their lived experiences, so gen-
dered cultural facets may affect how bodies and physiologies
respond to drugs.
For example, ­there is a long-­standing argument that the
sleeping aide zolpidem (Ambien) affects ­women differently
than men. The development of the original dosage data for zol-
pidem came from testing on men (assumedly 3G males), and
­because ­women (­here meaning 3G females) are, on average,
smaller in overall body and muscle mass than men, the assump-
tion was that the same dose would affect ­women more than
men (on average). Recent testing on ­women (3G females)
shows that they are not more heavi­ly affected by the drug, nor
are t­ here dif­fer­ent chemical impacts by zolpidem in w ­ omen’s
bodies. To the contrary, it appears that ­there is a slower (up to
35 ­percent) clearance time of the drug in 3G females (on aver-
age) as compared to that in 3G males. This is not due to weight
or BMI. Something ­else is ­going on. The initial binary-­based
solution was to blanketly call for reduced dosages in ­women
relative to men, which can result in the drug having less effect
and thus failing to achieve its target goal (battling insomnia) in
­women. But such an approach does not get to why the differences
 134 Chapter 7

exist. Asking about the ­actual physiological response, rather

than assuming 3G males and 3G females are dif­fer­ent kinds of
­humans, is a better approach. It is likely that zolpidem clearance
in ­humans is mediated, in part, by a system of enzymes called
Cytochrome P450 (CYP), and their activity might be related to
aspects of testosterone.24 If this is the case (which is not yet
confirmed), then the attention should be focused on the vary-
ing levels of acting testosterone in attenuating the effectiveness
of zolpidem. Testosterone is not characterizable as a male or
female hormone, but rather by variation in circulating levels
across ­humans, with 3G males usually having much higher lev-
els than 3G females.
What appears to be in play in the zolpidem case, and in the
majority of assessments of drug-­impact variation, is that
the ­sex-­binary assumption of males and females as two discrete
categories misses the substantial physiological variation in
­humans within, between, and across sex biologies, gender, and
3G categories.25 Rather than two categories (men and ­women)
as the comparison, the patterns of variation in enzymatic activ-
ity, endocrine function, muscle activity, and so on should be the
focus for understanding the specifics of drug action in h­ uman
systems. One category (males) cannot stand in for all h­ umans.
That category, even if restricted to 3G males, in and of itself does
not effectively capture all the variation in each of its own con-
stituent Gs (ge­ne­tic variants, testes variants, and genital vari-
ants). For example, t­ here is substantial variation in muscle mass
(as much as 100 ­percent) and massive variation in postpubertal
circulating testosterone (by as much as 300 ­percent) between
3G-­male individuals.26 The solution is also not simply to add an
equal number of 3G females to the testing sample (although
that is a good start). We know that ­human females vary substan-
tially even within the 3G definition, in body size,27 in genital
 W h y t h e B i n a r y V i e w I s a P r ob l e m 135

morphology,28 breast biomechanics,29 and endocrine d­ ynamics.30

The binary two-­kinds-­of-­human view restricts effective scien-
tific understanding of pharmacological impacts in ­humans.
Shifting to psychiatric conditions, many neurobehavioral
disorders differ in prevalence between the categories of men
and ­women and boys and girls. Recognized syndromes such as
autistic spectrum, ADHD, dyslexia, depression, anxiety, de-
mentia, and eating disorders all have varying frequencies across
genders, ethnicities, and nations. Such syndromes are complex
and dynamic, and are almost always biocultural. Gender/sex
clearly plays a role in many (or all) of ­these conditions. ­These
conditions represent complex interlacing of physiological, neu-
rological, social, experiential, and individual pro­cesses. Al-
though studies of brain variation situated in the sex binary often
intend to offer to a better understanding of health disparities in
psychiatric and psychological conditions, their assumptions of
a “male” and “female” brain are likely obfuscating the a­ ctual re-
lationships as the binary frame excludes the complexities of
interindividual neurobiological variation and ­human biocul-
tural experience.31

Cardiac Disease
Since 1984, more w ­ omen than men have died of ischemic
heart disease and heart failure each year, and yet more men
(­here meaning 3G males) have been diagnosed with heart disease
than ­women (­here meaning 3G females).32 Doctors ­diagnose
heart conditions by a combination of clinical ­presentation, bio-
markers, and imaging. Obstructive coronary artery disease
(CAD) remains the current focus of most assessment and ther-
apeutic strategies, but ­women have lower rates of typical CAD
than men. ­Women also have a higher rate of diverse symptoms
 136 Chapter 7

and adverse outcomes likely linked to coronary and microvas-

cular dysfunction.33 Also, understanding of antiplatelet therapy,
a primary tool of cardiovascular disease prevention, was, ­until
very recently, largely based on research with almost exclusively
3G males, and it appears that ­there may be a wider range of
variation in function of t­ hose drugs in 3G females.34 ­There is
also substantive evidence that the manifestations of cardiac dis-
ease and distress, and the effectiveness of vari­ous treatments,
vary across ­humans (a lot), and one salient aspect of that varia-
tion is in the categories “men” and “­women,” which are not sim-
ply biological. Some of this gender difference is likely linked to
variations in 3G-­sex-­related physiology, but the data are clear
that socioeconomic status, nutrition, smoking, alcohol use,
racism, sexism, stress hormones, inflammation patterns, im-
mune cell function diversity, and cellular aging all play key roles
as well.35 This pattern suggests that rather than a binary man/
woman or 3G-­male/3G-­female frame, which much of the med-
ical world currently holds,36 a dynamic gender/sex and bio-
cultural frame should be applied to the issue of cardiac disease
and its treatments.

Organ Transplants
While t­ here is no evidence that most organs originate in a bi-
nary fashion (3G-­male and 3G-­female organs) in ­humans, ­there
are on average worse outcomes for transplants from 3G-­female
donors regardless of the 3G sex of the recipient. Also, kidneys
and hearts (but not livers) transplanted from 3G females
have specifically higher failure rates in 3G-­male recipients.37
One might take a binary view and argue that this indicates some
level of “female” organs and some kind of incompatibly with
“male” immune systems. And one would likely be wrong. It
 W h y t h e B i n a r y V i e w I s a P r ob l e m 137

turns out that more 3G females (and ­women in general) donate

organs than do 3G males and men. ­Those 3G females and
­women who donate are also generally sicker and older than the
population on average and a larger percentage of the 3G males
who receive organ transplants are also generally sicker and
older than the population on average. This pattern creates a dy-
namic that is not best examined through only a sex-­biology
lens. Rather, it’s most likely that organs are responsive to effects
of gender/sex and other aspects of life. Thus, the details of the
lives of the donors and recipients and the traumas—­social and
physiological—­within them are as likely as, if not more, rele-
vant to transplant success than if donor was XX or XY genet­
ically. This situation is described in a large review entitled “How
Sex and Gender Affect Transplantation,” in which the t­ wenty-­six
authors state, “In this review, we summarize the data regarding
sex-­and gender-­based disparity in adult and pediatric kidney,
liver, lung, heart, and hematopoietic stem cell transplantation
and argue that t­ here are not only biological but also psychologi-
cal and socioeconomic issues that contribute to disparity in the
outcome, as well as an inequitable access to transplantation for
­women and girls.”38

Pregnancy and Related Physiological Changes

­ ere is a specific set of organs that, due to mammalian repro-
Th
ductive biology, establishes critical patterns of variation be-
tween most 3G-­female and 3G-­male bodies. The dynamics of
gestation and lactation, as already noted multiple times in this
book, m­ atter im­mensely and group t­ hose who can get pregnant
and do get pregnant into a cluster of relevance to medical re-
search and treatment. However, the binary approach generally
deployed by the medical world does not cover every­thing ­here.
 138 Chapter 7

Not all of ­those who get pregnant and give birth are classified or
identify as w ­ omen, or 3G females, and t­ here are also plenty of
individuals who have the physiology for gestation and lactation
but cannot or do not get pregnant and give birth. Thus, the focus
of research into pregnancy and birth should not simply be on
­women or 3G females as one of the two categories in a binary.
Rather the focus should be on the patterns, dynamics, and ex-
periences of the range of t­ hose who can and do get pregnant. For
example, in 2020 the National Academies of Science, Engineer-
ing, and Medicine, examining the data and relevant patterns of
variation changed their terms for research in this arena to “preg-
nant ­people” or “pregnant individuals” in place of “pregnant
­women.”39
While sex biology plays a core role in pregnancy, a mountain
of data supports the assertion that the individual experience of
pregnancy, like so much ­else in ­human lives, strongly inter-
twines with social, economic, p­ olitical, racialized, and related
aspects.40 Even something as specific as where one gives birth
can have dramatic impacts on bodies and lives.41 Pregnancy, like
so much of the ­human, is also biocultural. ­There is a substantial
range of physiological variation between individuals who can
and do get pregnant in terms of length of gestation (which varies
by as much as many weeks), hypertensive disorders, labor/­
delivery experience, and much more.42 Given this range of varia-
tion in biology, and the experience and context of pregnancy
and birth, rather than seeing w ­ omen, p­ eople who can get
­pregnant, and ­people who do get pregnant as a monolithic cat-
egory, it makes better scientific and social sense to examine the
patterns and pro­cesses of the variation across and among t­ hose
who do get pregnant to better facilitate healthy outcomes.
­Humans who gestate and give birth undergo a suite of physi-
ological dynamics that are not pre­sent in the same form and
 W h y t h e B i n a r y V i e w I s a P r ob l e m 139

intensity in other ­human bodies. Gestating and giving birth

play a core role in the construction of gendered perceptions
about, and expectations of, ­women. Many gendered, and
­medicalized, perspectives pre­sent pregnancy as debilitating or
as some form of malady. This is common in patriarchal systems
that argue for hyperdifferentiation between w ­ omen’s and men’s
places and capacities in society, often promoting men in the
public sphere of power and overemphasizing w ­ omen as tied to
the home and caretaking. But this use of gestation as a tool of
restrictions and oppression is not justified biologically. While
the impacts of gestation are physiologically substantive and
have distinctive effects on bodies, they are not necessarily the
restrictive, debilitating constraint many cultural and gendered
assumptions assert they are. Pregnant individuals can usually
retain their typical levels of social and physical activity and
physiological functioning across the pregnancy.43 And, ­because
the ­human reproductive system evolved as a cooperative ­system
involving many individuals beyond the ­mother, the individual
gestating and giving birth’s physiological system is “expecting”
a range of collaborative action by other individuals involved
(by their family/community). However, ­because of the mis-
guided binary-­based belief about limitation in social roles and
physical incapacitation in ­humans who gestate and give birth,
­there is often robust social, economic, and p­ olitical inequity for
all ­women.

Sex Contextualism in Medicine

Approaching sex biology as a culturally contextualized pat-
terned variation across and within categories is a framework
that helps improve medical care by rejecting a s­ imple binary and
showing that investing in a­ ctual understanding of biological
 140 Chapter 7

and social variation and patterns across bodies is key to ef-

fectively linking biological diversity to facilitating health.
Drawing on the conceptual framework of “sex contextual-
ism” can be particularly useful when it comes to medicine and
biomedical research.44 This view emphasizes that “male” and
“female” or “men” and “­women” do not mean the same ­things
in all contexts, nor are ­these the only subclasses or categories
that are applicable or useful in biomedical research.45 As
­there is often substantial variation in some aspects of biology
within the categories of “men” and “­women,” using them as
the “bins” every­one gets put into for a given biomedical as-
sessment, as opposed to focusing on the patterns of variation
in the biology related to the medical issues of interest, can
lead to missed opportunities for achieving health.46 Think of
the zolpidem example and w ­ hether or not “male” and “fe-
male” are the best categories for comparisons in that case
(they a­ ren’t), or the concern about biological variability in
mice and ­humans classified as “female” and how this has af-
fected medical research by keeping “females” and their com-
plexity out of it. The data are clear; the medical world should
think beyond a sex binary as the only way to ask questions
about health. Sex contextualism, by contrast, recognizes the
pluralism and context-­specificity of operationalizations of
“sex” across medical research and urges prac­ti­tion­ers to at-
tend to, be clear about, and be consistent with the uses and
meanings of the classification used in the design, interpreta-
tion, and communication of that research.47 This approach
in medicine may also be impor­tant for the sporting world.
Sports could also benefit from thinking more deeply about
the use, meaning, and impact of sex categories, adding nu-
ance and context to better engage ­human variation and ath-
letic ­performance.
 W h y t h e B i n a r y V i e w I s a P r ob l e m 141

Sports and the Definition of ­Woman

Caster Semenya is a South African athlete, winner of two Olym-
pic gold medals and three World Championships in the
­women’s 800-­meter race. ­After her streak of wins, Ms. Semenya
was forced to undergo “sex confirmation” testing, becoming a
center of attention in the b­ attle of sex and gender in the sporting
world. Her case, and ­others, resulted in vari­ous sets of debates,
bans, and restrictions on ­women athletes. Ms. Semenya, identi-
fied as female on her birth certificate, has genitals concordant
with that classification, grew up a girl, and is a w ­ oman. How-
ever, she is not a 3G female. Ms. Semenya has higher-­circulating
testosterone than is typical for 3G females, a Y chromosome,
and internal gonads that are the equivalent of undescended
­testes.48 Ms. Semenya is a w ­ oman whose sex biology is outside
what is typical for 3G females but well within the range of varia-
tion found in our species. The binary view, however, does not
allow space for her.
Between 2017 and 2019, the International Olympic Commit-
tee and World Athletics, the two main groups controlling track
events, barred ­women who had naturally occurring levels of
testosterone (T) above 5 nmol/L from competing in races
­between 400 meters and 1,500 meters (including hurdles). The
assertion was that such a level of circulating T was outside of
the “normal” range for “­women” and that it veered into the
range of “men,” thus offering an unfair competitive edge to
­those w­ omen. In making this decision, the groups drew on re-
search suggesting that while testosterone levels did not corre-
late with ­performance in most sporting events, t­ here ­were some
correlations found between circulating testosterone and levels
of success in the 400m, 400m hurdles, and 800m races, and the
hammer throw and pole vault for w ­ omen athletes. Researchers
 142 Chapter 7

argued that a higher testosterone level seemed to confer a 1.8–

2.8 ­percent competitive advantage, but the ­actual mechanism
of this advantage was not clear. This same study found that male
athletes in throwing events had lower than expected testoster-
one levels and that testosterone did not appear to play a key role
in longer-­distance ­running or in most other sporting events as-
sessed for men or w ­ omen.49 In a 2017 article, the main group of
researchers relied on by the International Olympic Committee
and World Athletics concluded that testosterone does not have
a biasing effect in most sporting events regardless of the gender
classification of the event and that differences in circulating tes-
tosterone have no biasing effect in any of the men’s categories,
but that ­there is, at least in a few track events, “a significant role
of endogenous androgens for athletic ­performance in ­women.”50
However, in 2021, ­after publishing a few more studies arguing
that higher testosterone benefits w ­ omen in some track events,
that same group of researchers published a little-­noticed
­correction to their ­earlier work. In it, they stated that their con-
clusions about the role of testosterone on athletic p­ erformance
in ­women elite athletes “should be interpreted based on an ex-
ploratory study setting where it was not pos­si­ble to adjust for
potential confounding ­factors.” And they concluded, “To be
explicit, t­ here is no confirmatory evidence for causality in the
observed relationships reported.”51 In short, ­there are patterns
of associations between testosterone and many other bodily
and experiential ­factors that influence, and may potentially bias,
outcomes in sporting events, but the specific relationships, es-
pecially w­ hether it is the level of testosterone that makes the
difference, are not clear.52
In some, but certainly not all, sporting competitions, upper-­
body strength, per-­unit of area muscle strength, overall body
size, and leg length and muscle p­ erformance are biological
 W h y t h e B i n a r y V i e w I s a P r ob l e m 143

variables that may offer specific benefits. They are also aspects
of ­human bodies that are affected, often substantially, by ­variation
in sex biology. ­Because ­these variants do manifest, on average,
proportionately differently in 3G females and 3G males, and
­these 3G categories are most often societally correlated with the
categories men and ­women, many sports ­today have developed
men’s and ­woman’s sports where the competitions are gender-­
uniform, such as basketball, baseball, football (soccer), tennis,
and track and field events.53 Given recent societal changes (in
some places) to accept a broader range of gendered identities,
­there are increasing situations where individuals who w ­ ere as-
signed a gender at birth and have changed that gender (are now
transgendered individuals) wish to compete in their current
gender category. At the same time, t­ here are also individuals, as
with Ms. Semenya, assigned a gender at birth, who remain in
that gender, but do not fully fit the current assumptions about
the biology “under­lying” that gender, who are being excluded
from competitions based on a highly specific (and culturally
structured) definition of “­woman” (basically, how much testos-
terone the body naturally produces). It is worth noting that
being of the right “sex” is brought up as an issue in ­women’s
sports but not in men’s.
Before making any further comments on t­ hese cases, we
must recognize the fact that ­women’s professional and amateur
sports are relatively new as ­organizations and institutions, un-
derfunded, and undersupported relative to men’s sports. ­There is
abundant evidence that societal gender structures and gender/
sex dynamics of girls’ and w ­ omen’s lives structure their bodies
and possibilities affecting their capacities, ­performances, and
risk of injury in sports differently than ­those of boys and men
who are trained for sports.54 This gendered training dynamic is
illustrated in ­running sports, where the gender differentials in
 144 Chapter 7

p­ erformance are the smallest at elite levels where the training

is at the highest caliber and closer to equity as opposed to larger
differences at lower and amateur levels where training and in-
frastructure quality and options are highly divergent in quality
(primarily favoring boys and men).55 Any discussion of ­women
athletes’ capacities and ­performance must recognize that t­ here
is an under­lying inequity resulting in w ­ omen athletes not yet
achieving the full potential relative to their physiological ca-
pacities at the degree to which men athletes do.
Coming back to the two cases, transwomen and ­women who
are defined out of being a w ­ oman by testosterone levels, one
notes that each is dif­fer­ent both in relation to the dynamics of
sex biology and to the ways in which gender/sex works and
gender is constructed, but both are united as being identified as
a “prob­lem” by the binary view. In the case of defining what a
­woman is by testosterone production, this is clearly not simply
a biological m ­ easure, as it reflects a social construct of what the
“correct” range of testosterone should be. While testosterone
and other androgens often play roles in much physiological
functioning related to physical activity, it is not clear if or how
an individual’s physical ­performance at sports, including limb
length, training regime, muscle density, oxygen-­carrying capac-
ity of the red blood cells, lung capacity, and so on, translates into
a deciding ­factor as to one’s gender and broadscale physiological
capacity. The decision to define who is a ­woman based on
­specific ranges of testosterone levels is a social and cultural one,
not a biological one, and it has a range of ethical and societal
implications.56
For the case of transwomen competing in w ­ omen’s sporting
events, it is a bit more complicated. If an individual is compet-
ing in a sport where aspects of sex biology such as upper-­body
strength, muscle density, or overall body size offer benefits,
 W h y t h e B i n a r y V i e w I s a P r ob l e m 145

then an individual may indeed have an advantage if they are at

the top end of the distribution of variation in ­those bodily areas
relative to their competitors. But the structure and patterns of
such advantage vary by sport, individual life histories, body
variation, and other variables.57 This is the case already in
gender-­uniform sports, as certain individuals have biological
variants that offer them a par­tic­u­lar set of advantages (think of
the swimmer Michael Phelps, a 3G male and a man, relative to
the majority of men he competed against58). However, in the
case of transwomen, it is argued that some of ­those benefits ­will
be substantially outsized b­ ecause of lingering effects of their
pretransition physiology. But this assumes that the individual
transitioned postpuberty from a body with testes and that they
are in the upper end of 3G-­male physiological outcomes as op-
posed to being in the large area of overlap between 3G-­male and
3G-­female bodies. Given the extremely small number of trans-
gendered athletes overall and the even smaller number of ­those
individuals who are winning an outsized percentage of their
competitions, it is difficult, problematic, and largely unscientific
to make any broad-­scale blanket generalizations of this s­ ituation.
And any rules created from such generalizations, especially if
based exclusively on natal gonads and timing of transition, ig-
nore the substantive variation in bodies and capacities (which
is unfortunately what World Athletics did59). However, one
­thing is clear: the current gender inclusion and exclusion sys-
tem for many sporting events, leagues, and competitions is in-
sufficient to engage, effectively, the sex-­biology variation and
the gender/sex real­ity of ­humans.60
This focus on sports events, athletes’ bodies, and testoster-
one is in­ter­est­ing and ­matters for the athletes and the sporting
world. But it is a poor venue to examine, and think about, varia-
tion in sex biology in h­ umans. Only a minuscule percentage of
 146 Chapter 7

h­ umans have the pattern of biological variation that confers

bodily capacity, skillset, and opportunity to be elite athletes or
even athletes in amateur and school competitive contexts.
­There are about 85,000–95,000 professional athletes in the entire
world and about 170,000 individuals worldwide who compete
for Olympic slots.61 To put this in perspective: t­ here are only
between 6,000 and 11,000 professional athletes in the entire
United States, out of a population of over 330,000,000. This
means that elite athletes make up .0012 ­percent of living h­ umans
globally and .003 ­percent of ­people in the United States. Using
one thousandth of one ­percent of ­humans, who are mostly at
the extreme ends of the h­ uman range for many biological
­factors and with a lifetime training related to their sports, as the
key group to understand human-­wide patterns of sex biology is
misleading and faulty science. The focus on athletes to legislate
issues of transgender makes even less sense when one considers
that only 0.6 ­percent of the population 13 years of age and above
in the United States identifies as transgendered, and only a tiny
percentage of them are athletes.62
In the day-­to-­day life of the vast majority of h­ umans, a slight
competitive edge in ­running an 800-­meter race is prob­ably not
relevant. Variation in circulating levels of testosterone is not the
sole determinant, or necessarily a main determinant, of any in-
dividual’s be­hav­ior, capacities, experience, or identity.63 This is
not to state that testosterone is not an impor­tant hormone for
­humans (and most animals), ­because it is, as are estrogen, pro-
gesterone, and so on. It is also not to state that T is not associ-
ated with many aspects of physiology and has some differing
impacts on bodily systems across ­humans, specifically in indi-
viduals with testes or individuals who receive external testos-
terone. Testosterone has a range of impor­tant functions and
 W h y t h e B i n a r y V i e w I s a P r ob l e m 147

impacts in the ­human body in multiple manners. However, if

we are truly interested in an accurate and comprehensive un-
derstanding of sex biology, then focusing on something as re-
cent and hyperculturally modified at professional/elite athletics
is a poor place to look.

Rest­rooms
In 2016, the US state of North Carolina passed a law prohibiting
transgender individuals from using the rest­room that corre-
sponded to their gender. This action was part of the ­political,
­legal, and public debates over ­whether transgender individuals
should be able to use public rest­rooms (including locker rooms
and changing rooms) that match their gender or if they should
be forced to use ones that match a social categorization of the
person based on which genitals they have (or had) and how
they ­were assigned at birth. Between 2021 and 2023, legislatures
in 34 US states introduced over 300 anti-­transgender bills re-
flecting a wide range of proposed restrictions on issues of
health, ­legal rights, and access for individuals who did not fit
certain beliefs about gender and sex biology.64 By early 2024,
­there w­ ere more than 450 active anti-transgender bills moving
through the legislatures in 41 states.65 In June 2023, the state of
Florida implemented H ­ ouse Bill 1521, which mandates that
transgender individuals must use public rest­rooms that corre-
spond with the sex assigned to them at birth, even if they have
legally updated their gender on their birth certificate and driv-
er’s license. If they do not comply, they can be charged with
trespassing—­punishable by up to a year in jail. In January of
2024, Utah followed with a similar law. ­These laws and their
associated outcomes are a clear, power­ful, and targeted attempt,
 148 Chapter 7

based on the binary notion of sex biology and gender, to curtail

the rights of a group of ­humans whose bodies and lives chal-
lenge the binary view.
Everyday ­people encounter the world in a myriad of dif­fer­ent
ways b­ ecause of the combinations of their distinct sex-­biology
variations, their gender/sex, and a society’s expectations of gen-
der. In some socie­ties, rest­rooms in public, in school, and in the
workplace are divided along a binary gender system with sepa-
rate facilities for men/boys and women/girls. For example, in
the United States, depending on one’s gender classification, the
structures and locations for urinating and defecating (urinals,
toilets, stalls, dividers, ­etc.) are quite dif­fer­ent as are the layout
and often the sanitary conditions of the rest­rooms. Some of the
differences are asserted to be connected to genital differences
and the capacity to urinate while standing. But most of the struc-
tures are highly gendered and reflect not so much the physical
necessities of sex-­biology variation (all ­humans can effectively
urinate and defecate when sitting) but the cultural assumptions
about what that variation means. It is also an explicit cultural
assertion that a binary division is the only acceptable option for
­human gender/sex regarding urination, defecation, and washing
up. Th
­ ose individuals with sex-­biology and gender/sex variation
fully within the range of the h­ uman pattern but not fitting this
specific set of genital-­based (and broadly 3G) binary assump-
tions must then decide which rest­room to enter and use and
risk, in many cases, harassment, vio­lence, or ­legal repercussions
if someone ­else decides ­they’ve chosen the “wrong” one.66
The rest­room example is just one of so many public misper-
ceptions rooted in the binary. ­W hether it is in the workplace,
the p­ olitical or religious sphere, school, or broader public
­venues, everyday ­people vary a lot, but most assumptions about
what that variation is and what it means are highly restrictive
 W h y t h e B i n a r y V i e w I s a P r ob l e m 149

and ignore the ­actual variation in the ­human system. Simplistic

binary assertions about how men or w ­ omen “are” and “should”
be, and assumptions that every­one with the same type of
­genitals or gonads shares the same set of ways of being, perme-
ate the world, making it difficult not only for t­ hose who do not
fit the general assumptions for the categories of men and
­women, but also ­those who do.

The Binary Is Wrong and Harmful

This book has endeavored to introduce what is actually known
about sex biology and the h­ uman experience and to explain
why the most accurate way to discuss and research it is via a
biocultural framework. The best science requires a full consid-
eration of patterns and diversity and the recognition that
­humans have gender/sex. Thinking broadly about the evolution
of sex, sex biology across the animal kingdom, and the ways in
which our biological understanding of sex has changed over
time helps us see diversity in sex biology as typical. Th­ ere are
few ­simple one-­to-­one universal “truths” about being a female
or male, and treating ­those two categories as dif­fer­ent kinds of
being is not supported by the science of sex biology. Reviewing
­human evolutionary history and the patterns of sex biology in
current ­human bodies opens us up to a better understanding of
why and how ­humans are in the world and rejects simplistic
binary assertions about men and ­women.
The take-­home message is that patterns of sex biology, while
always impor­tant, are not the exclusive determinants of the en-
tirety of or the specifics of any given individual’s gender/sex
experience or their experience of self. The biological, behav-
ioral, and social variation exhibited by h­ umans is almost always
substantively broader, and more dynamic, than culturally
 150 Chapter 7

created binary descriptions and bound­aries for gender and sex.

Moving past the binary view does not mean ignoring variation
and differences across humanity, including t­ hose related to sex
biology. Rather, the necessity to develop better understandings
of ­those differences, and that variation, is exactly why ­there is a
need to move past the sex and gender binary. Let’s do every­
thing we can to make that happen.
 no t e s

1. The Evolution of Sex

1. The earliest life prob­ably started with RNA (single-­stranded ge­ne­tic material),
and then DNA (double-­stranded ge­ne­tic material) emerged a bit ­later
2. The bacteria are divided into archaebacteria and eubacteria, but much current
work separates the two further into archaea and bacteria as distinct evolutionary
lineages
3. U. Goodenough and J. Heitman. 2014. “Origins of Eukaryotic Sexual Repro-
duction.” Cold Spring Harbor Perspectives in Biology 6 (3): a016154. https://­doi​.­org​
/­10​.­1101​/­cshperspect​.a016154; D. Bachtrog, J. E. Mank, C. L. Peichel, M. Kirkpat-
rick, S. P. Otto, T. L. Ashman, M. W. Hahn, et al. 2014. “Tree of Sex Consortium. Sex
Determination: Why So Many Ways of D ­ oing It?” PLoS Biology 12 (7): e1001899.
https://­doi​.­org​/­10​.­1371​/­journal​.­pbio​.1001899.
4. G. C. Williams. 1975. Sex and Evolution. Monographs in Population Biology 8.
Prince­ton, NJ: Prince­ton University Press, v.
5. For a very basic overview, see C. Zimmer. 2009. “On the Origin of Sexual Re-
production.” Science 324 (5932): https//­doi​.­org​/­10​.­1126​/­science​.­324​_­1254​. For two
much more in-­depth, classic overviews, see G. C. Williams. 1975. Sex and Evolution.
Monographs in Population Biology 8. Prince­ton, NJ: Prince­ton University Press;
and U. W. Goodenough. 1985. “Origins and Evolution of Eukaryotic Sex.” In Ori-
gins and Evolution of Sex, edited by H. O. Halvorson and A. Monroy, 123–40. New
York: Alan R. Liss. For more recent reviews, see U. Goodenough and J. Heitman.
2014. “Origins of Eukaryotic Sexual Reproduction.” Cold Spring Harbor Perspectives
in ­Biology 6 (3): a016154. https//­doi​.­org​/­10​.­1101​/­cshperspect​.a016154.
6. J. Lehtonen and G. A. Parker. 2014. “Gamete Competition, Gamete Limitation,
and the Evolution of the Two Sexes.” Molecular H ­ uman Reproduction 20 (12): 1161–68.
https://­doi​.­org​/­10​.­1093​/­molehr​/gau068; D. Bachtrog, J. E. Mank, C. L. Peichel,
M. Kirkpatrick, S. P. Otto, T. L. Ashman, M. W. Hahn, et al. 2014. “Tree of Sex Con-
sortium. Sex Determination: Why So Many Ways of ­Doing It?” PLoS Biology 12 (7):
e1001899. https://­doi​.­org​/­10​.­1371​/­journal​.­pbio​.1001899.

151
 152 Not e s to Ch a p t er 1

7. Since this book is ­really just focused on the animals, we ­won’t even try to get
into the sex biology of plants, which is far more complicated. If you are interested,
read D. Charlesworth. 2002. “Plant Sex Determination and Sex Chromosomes.” He-
redity 88: 94–101. https://­doi​.­org​/­10​.­1038​/­sj​.­hdy​.6800016; J. R. Pannell. 2017. “Plant
Sex Determination.” Current Biology 27 (5): R191–­R197. https://­​.­doi​.­org​/­10​.­1016​/­j​
.­cub​.­2 017​.­01​.052; and especially B. Subramaniam and M. Bartlett. 2023. “Re-­
imagining Reproduction: The Queer Possibilities of Plants.” Integrative and Com-
parative Biology 63 (4): 946–59. https://­doi​.­org​/­10​.­1093​/­icb​/icad012.
8. T. Laqueur. 1990. Making Sex: Body and Gender from the Greeks to Freud. Cam-
bridge, MA: Harvard University Press.
9. V. Sanz. 2017. “No Way Out of the Binary: A Critical History of the Scientific
Production of Sex.” Signs: Journal of ­Women in Culture and Society 43 (1): 1–27.
https://­doi​.­org​/­10​.­1086​/692517; S. Richardson. 2013. Sex Itself: The Search for Male
and Female in the ­Human Genome. Chicago: The University of Chicago Press; see also
E. Martin. 1991. “The Egg and the Sperm: How Science Has Constructed a Romance
Based on ­Stereotypical Male-­Female Roles.” Signs 16 (3): 485–501.
10. C. Darwin. 1871. The Descent of Man and Se­lection in Relation to Sex. London:
John Murry.
11. A. J. Bateman. 1948. “Intrasexual Se­lection in Drosophila.” Heredity 2:349–68.
12. Bateman’s assertions ­were that females are l­ imited in how many gametes they
can produce due to the size and energetic costs of ova, and thus their investment in
each gamete is very high. Basically, Bateman (working with fruit flies) argued that
males can produce unlimited amounts of small sperm, while females can only pro-
duce a ­limited number of large ova. So, given this difference in costs to the organisms,
males should try to mate with as many females as pos­si­ble (get as much sperm out
­there as pos­si­ble), but females, with costly, l­ imited ova, should be coy, sexually pas-
sive, and very choosy. This is what Darwin said in 1871 about animal evolution in
general (including ­humans), but in 1948, Angus Bateman put the cause of this as-
sumed real­ity squarely on the fact of anisogamy.
13. G. C. Williams. 1966. Adaptation and Natu­ral Se­lection. Prince­ton, NJ: Prince­ton
University Press; R. Trivers. 1972. “Parental Investment and Sexual Se­lection.” In Sexual
Se­lection and the Descent of Man, edited by B. Campbell, 52–95. Chicago: Aldine.
14. E. O. Wilson. 1975. Sociobiology: The New Synthesis. Cambridge, MA: Belknap
Press of Harvard University Press; G. A. Parker. 1979. “Sexual Se­lection and Sexual
Conflict.” In Sexual Se­lection and Reproductive Competition, edited by M. S. Blum and
N. A. Blum, 123–66. New York: Academic Press. Please note all of the most promi-
nent scholars making t­ hese arguments are men: Darwin, Bateman, R. L. Trivers,
E. O. Wilson, and G. A. Parker.
15. Z. Tang-­Martinez and T. B. Ryder. 2005. “The Prob­lem with Paradigms: Bate-
man’s Worldview as a Case Study.” Integrative and Comparative Biology 45: 821–30.
 Not e s to Ch a p t er 1 153

See also P. Gowaty, Y. Kim, and W. Anderson. 2012. “No Evidence of Sexual Se­lection
in a Repetition of Bateman’s Classic Study of Drosophila melanogaster.” Proceedings
of the National Acad­emy of Sciences of the United States of Amer­i­ca 109: 11740–45.
https://­doi​.­org​/­10​.­1073​/­pnas​.1207851109; H. Kokko and M. Jennions. 2003. “It
Takes Two to Tango.” Trends in Ecol­ogy and Evolution 18 (3): 103–4; C. M. Drea. 2005.
“Bateman Revisited: The Reproductive Tactics of Female Primates.” Integrative and
Comparative Biology 45 (5): 915–23. https://­d oi​ .­o rg​ /­10​ .­1093​ /­i cb​ /­4 5​ .­5​ .915;
M. Borgerhoff-­Mulder. 2004. “Are Men and ­Women ­Really So Dif­fer­ent?” Trends in
Ecol­ogy and Evolution 19 (1): 3–6.
16. M. Ah-­King. 2013. “On Anisogamy and the Evolution of ‘Sex Roles.’ ” Trends in
Ecol­ogy and Evolution 28 (1): 1–2. https://­doi​.­org​/­10​.­1016​/­j​.­tree​.­2012​.­0 4​.­0 04​. A.
Fausto-­Sterling. 2020. Sexing the Body: Gender Politics and the Construction of Sexual-
ity. London: Hachette; L. Z. DuBois and H. Shattuck-­Heidorn. 2021. “Challenging
the Binary: Gender/Sex and the Bio-­logics of Normalcy.” American Journal of H ­ uman
Biology 33 (5): e23623. https://­doi​.­org​/­10​.­1002​/­ajhb​.23623; J. F. McLaughlin,
­Kinsey M. Brock, I. Gates, P. Anisha, M. Piattoni, A. Rossi, and S. E. Lipshutz. 2023.
“Multivariate Models of Animal Sex: Breaking Binaries Leads to a Better Understand-
ing of Ecol­ogy and Evolution.” Integrative and Comparative Biology 63 (4): 891–906.
https://­doi​.­org​/­10​.­1093​/­icb​/icad027; C. T. Ross, P. L. Hooper, J. E. Smith, A. V.
Jaeggi, E. A. Smith, S. Gavrilets, F. T. Zohora, et al. 2023. “Reproductive ­Inequality in
­Humans and Other Mammals.” Proceedings of the National Acad­emy of Sciences of
the United States of Amer­i­ca 120 (22): e2220124120. https://­doi​.­org​/­10​.­1073​/­pnas​
.222012​4120.
17. Much of the biological lit­er­a­ture on nonhuman animals still uses the term
“hermaphrodite” for individuals who have both gonad types (or an ovotestis). “Her-
maphrodite” is no longer used for h­ umans, and increasingly less so for other animals,
due to historical baggage and the fact that the term “intersex” is more expansive and
encompasses multiple biological variants regarding gonads and other sex biology.
18. Except not always. ­There are a number of fish and reptiles that make ­these
structures but still store them internally u­ ntil the zygotes develop into juvenile or-
ganisms and are capable of making it on their own. Then they excrete them.
19. Many egg-­laying species also have genitals for assisting in the internal fusion
of gametes.
20. For relatively concise and pretty good overviews of animal reproductive
systems, see G. C. Kent. 2021. “Animal Reproductive System.” Encyclopedia Britan-
nica, November 19. https://­w ww​.­britannica​.­com​/­science​/­animal​-­reproductive​
-system.
21. This system is a bit dif­fer­ent in marsupial and placental mammals. In placen-
tals, all the fetal development is internal, but in marsupials the last portions of it take
place in a sack external to the reproductive tract.
 154 Not e s to Ch a p t er 1

22. Z. M. Thayer, J. Rutherford, and C. W. Kuzawa. 2020. “The Maternal Nutri-

tional Buffering Model: An Evolutionary Framework for Pregnancy Nutritional
Intervention.” Evolution, Medicine, and Public Health 1: 14–27.
23. See L. Cooke. 2022. Bitch: On the Females of the Species. New York: Basic Books.
24. See, e.g., A. Liker, R. P. Freckleton, V. Remeš, and T. Székely. 2015. “Sex Dif-
ferences in Parental Care: Gametic Investment, Sexual Se­lection, and Social Environ-
ment.” Evolution 69 (11): 2862–75; H. Kokko and M. D. Jennions. 2008. “Parental
Investment, Sexual Se­lection and Sex Ratios.” Journal of Evolutionary Biology 21 (4):
919–48; Drea, “Bateman Revisited.”
25. R. L. Goldberg, P. A. Downing, A. S. Griffin, and J. P. Green. 2020. “The Costs
and Benefits of Paternal Care in Fish: A Meta-­Analysis.” Proceedings of the Royal So-
ciety B 287 (1935): https://­doi​.­org​/­10​.­1098​/­rspb​.­2020​.1759.
26. A. I. Furness, I. Capellini. 2019. “The Evolution of Parental Care Diversity in
Amphibians.” Nature Communications 10: 4709. https://­doi​.o­ rg​/­10​.­1038​/­s41467​-­019​
-­12608​-5.
27. A. Cockburn. 2006. “Prevalence of Dif­fer­ent Modes of Parental Care in Birds.”
Proceedings of the Royal Society: Biology 273 (1592): 1375–83.
28. See J. K. Rilling and L. J. Young. 2014. “The Biology of Mammalian Parenting
and Its Effect on Offspring Social Development.” Science 345 (6198): 771–76; L. S. B.
Hrdy. 2009. ­Mother and ­Others: The Evolutionary Origins of Mutual Understanding.
New York: Belknap; L. T. Gettler. 2016. “Becoming DADS: Considering the Role of
Cultural Context and Developmental Plasticity for Paternal Socioendocrinology.”
Current Anthropology 57: S38–­S51; S. Rosenbaum and L. T. Gettler. 2018. “With a
­Little Help from Her Friends (and F ­ amily) Part I: The Ecol­ogy and Evolution of
Non-­maternal Care in Mammals.” Physiology and Be­hav­ior 193: 1–11; and S. Rosen-
baum and L. T. Gettler. 2018. “With a L ­ ittle Help from Her Friends (and ­Family) Part II:
Non-­maternal Caregiving and Physiology Caregiving in Mammals.” Physiology and
Be­hav­ior 193: 12–24.
29. This is called “niche construction,” and it is a very salient and impor­tant part
of evolutionary pro­cesses. In par­tic­u­lar, patterns of niche construction can play cen-
tral roles in expanding and restricting the roles and types of variation pos­si­ble in sex
biology. This w ­ ill be a central theme in chapters 2 and 3. For a good overview article,
see ­here: K. Laland, B. Matthews, and M. W. Feldman. 2016. “An Introduction to
Niche Construction Theory.” Evolutionary Ecol­ogy 30: 191–202. And for an online
review see ­here: K. Laland and L. Chiu. 2020. Niche Construction. https://­
nicheconstruction​.­com​/­information​/.
30. J. F. McLaughlin, K. M. Brock, I. Gates, A. Pethkar, M. Piattoni, A. Rossi, and
S. E. Lipshutz. 2023. “Multivariate Models of Animal Sex: Breaking Binaries Leads
to a Better Understanding of Ecol­ogy and Evolution.” Integrative and Comparative
Biology 63 (4): 891–906. https://­doi​.­org​/­10​.­1093​/­icb​/icad027.
 Not e s to Ch a p t er 2 155

2. Animal Sex Biology: Mixing It Up

1. An analy­sis of more than four hundred mammalian species indicates that al-
though males tend to be larger than females when dimorphism occurs, males are not
larger in most mammals, and suggests a need to revisit other assumptions in sexual
se­lection and sex-­biology research. K. J. Tombak, S. B. S. W. Hex, D. I. Rubenstein. 2023.
“New Estimates Indicate That Males Are Not Larger Than Females in Most Mammals.”
Nature Communications 15 (1): 1872. https://­doi​.­org​/­10​.­1038​/­s41467​-­024​-­45739​-5.
2. L. Cooke. 2022. Bitch: On the Females of the Species. New York, Basic Books.
3. T. Janicke, I. K. Häderer, M. J. Lajeunesse, and N. Anthes. 2016. “Darwinian Sex
Roles Confirmed across the Animal Kingdom.” Science Advances 2 (2): e1500983;
C. M. Drea. 2005. “Bateman Revisited: The Reproductive Tactics of Female Pri-
mates.” Integrative and Comparative Biology 45 (5): 915–23. https://­doi​.­org​/­10​.­1093​
/­icb​/­45​.­5​.915; and Cooke, Bitch.
4. Cooke, Bitch.
5. ­There are some species of bee where “workers” do end up laying ova and thus
reproducing some haploid offspring, contributing up to 10 ­percent of the hives’
males. D. A. Alves, V. L. Imperatriz-­Fonseca, T. M. Francoy, P. S. Santos-­Filho,
J. ­Billen, and T. Wenseleers. 2009. “The Queen Is Dead—­Long Live the Workers:
Intraspecific Parasitism by Workers in the Stingless Bee Melipona scutellaris.” Molecu-
lar Ecol­ogy 18 (19): 4102–11. https://­doi​.­org​/­10​.­1111​/­j​.­1365​-­294X​.­2009​.­0 4323​.x.
6. H. Blackmon, L. Ross, D. Bachtrog. 2017. “Sex Determination, Sex Chromo-
somes, and Karyotype Evolution in Insects.” Journal of Heredity 108 (1): 78–93.
7. D. J. Diaz Cosin, M. Novo, and R. Fernández. 2010. “Reproduction of Earth-
worms: Sexual Se­lection and Parthenogenesis.” In Biology of Earthworms, edited by
A. Karaca, 69–86. Berlin: Springer-­Verlag.
8. Multiple sets of chromosomes, not just two.
9. C. H. Chandler, G. E. Chadderdon, P. C. Phillips, I. Dworkin, and F. J. Janzen.
2011. “Experimental Evolution of the Caenorhabditis elegans Sex Determination Path-
way.” Evolution 66 (1): 82–93. https://­doi​.­org​/­10​.­1111​/­j​.­1558​-­5646​.­2011​.­01420​.x.
10. E. V. Todd, H. Liu, S. Muncaster, and N. J. Gemmell. 2016. “Bending Genders:
The Biology of Natu­ral Sex Change in Fish.” Sexual Development 10 (5–6): 223–41.
https://­doi​.­org​/­10​.­1159​/000449297.
11. Todd et al., “Bending Genders.”
12. K. Semsar and J. Godwin. 2004. “Multiple Mechanisms of Phenotype Devel-
opment in the Bluehead Wrasse.” Hormones and Be­hav­ior 45: 345–53.
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14. Todd et al., “Bending Genders.”
 156 Not e s to Ch a p t er 2

15. P. L. Munday, T. Kuwamura, and F. J. Kroon. 2010. Bi-­Directional Sex Change

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Reduced Sexual Dichromatism.” Frontiers in Ecol­ogy and Evolution 4. https://­doi​.­org​
/­10​.­3389​/­fevo​.­2016​.00022.
25. C. Arnold. 2016. “The Sparrow with Four Sexes.” Nature 539: 482–84. https://­
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26. Given the range variations in development, it is physiologically pos­si­ble that
one could have a mix of reproductive physiology that does not fit neatly into typical
patterns and be capable of gestation. However, this is likely to be quite rare. See, for
instance, B. A. H. Schultz, S. Roberts, A. Rod­gers, and K. Ataya. 2009. “Pregnancy
 Not e s to Ch a p t er 2 157

in True Hermaphrodites and All Male Offspring to Date.” Obstetric Gynecol­ogy 113:
534–36. Also, ­there are a few species of bats where small-­gamete producers lactate.
27. Cooke, Bitch; Drea, “Bateman Revisited”; S. R. Zajitschek, F. Zajitschek,
R. Bonduriansky, R. C. Brooks, W. Cornwell, D. S. Falster, M. Lagisz, et al. 2020.
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28. See C. Delle-­Amores. 2024. “Love Them or Hate Them, Hyenas Are Getting
the Last Laugh.” National Geographic Magazine, February 8. https://­w ww​
.­nationalgeographic​.­com​/­animals​/­article​/­spotted​-­hyenas​-queens.
29. S. E. Glickman, L. G. Frank, J. M. Davidson, E. R. Smith, and P. K. Siiteri. 1987.
“Androstenedione May O ­ rganize or Activate Sex-­Reversed Traits in Female Spotted
Hyenas.” Proceedings of the National Acad­emy of Sciences of the United States of Amer­i­ca
84 (10): 3444–47. https://­doi​.­org​/­10​.­1073​/­pnas​.­84​.­10​.3444.
30. A. Conley, N. J. Place, E. L. Legacki, G. L. Hammond, G. R. Cunha, C. M. Drea,
M. L. Weldele, et al. 2020. “Spotted Hyaenas and the Sexual Spectrum: R ­ eproductive
Endocrinology and Development.” Journal of Endocrinology 247 (1): R27–­R44.
https://­doi​.­org​/­10​.­1530​/­JOE​-­20​-0252.
31. Conley et al., “Spotted Hyaenas and the Sexual Spectrum.”
32. And, in fact, t­ here is a solid argument that “monogamy” is not one ­thing.
­There can be social monogamy (but having sex with more than one partner) and/
or sexual monogamy (sex with only one partner). See A. Fuentes. 1999.
“­Re-­evaluating Primate Monogamy.” American Anthropologist 100 (4): 890–907; A.
Fuentes. 2002. “Patterns and Trends in Primate Pair Bonds.” International Journal of
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Encyclopedia of Primatology, edited by A. Fuentes, 831–35. London: John Wiley &
Sons; E. Fernandez-­Duque, M. Huck, S. Van Belle, and A. Di Fiore. 2020. “The Evo-
lution of Pair-­Living, Sexual Monogamy, and Cooperative Infant Care: Insights from
Research on Wild Owl Monkeys, Titis, Sakis, and Tamarins.” American Journal of
Physical Anthropology 171 (suppl. 70): 118–73. https://­doi​.­org​/­10​.­1002​/­ajpa​.24017.
33. M. M. Holmes and B. D. Goldman. 2021. “Social Be­hav­ior in Naked Mole-­
Rats: Individual Differences in Phenotype and Proximate Mechanisms of
­Mammalian Eusociality.” Advances in Experimental Medicine and Biology 1319: 35–58.
https://­doi​.­org​/­10​.­1007​/­978​-­3​-­030​-­65943​-­1​_2.
34. S. E. Glickman, R. V. Short, and M. B. Renfree. 2005. “Sexual Differentiation
in Three Unconventional Mammals: Spotted Hyenas, Elephants and Tammar Wal-
labies.” Hormones and Be­hav­ior 48: 403–17. https://­doi​.­org​/­10​.­1016​/­j.​ ­yhbeh​.­2005​.­07​
.013; Cooke, Bitch; Drea, “Bateman Revisited”; Zajitschek et al., “Sexual Dimor-
phism in Trait Variability.”
35. Cooke, Bitch.
 158 Not e s to Ch a p t er 3

3. ­Humans Are Messy

1. M. Cintas-­Peña, M. Luciañez-­Triviño, R. Montero Artús, A. Bileck, P. Bortel,
F. Kanz, K. Rebay-­Salisbury, et al. 2023. “Amelogenin Peptide Analyses Reveal
­Female Leadership in Copper Age Iberia (c. 2900–2650 BC).” Scientific Reports 13
(1): 9594. https://­doi​.­org​/­10​.­1038​/­s41598​-­023​-­36368​-­x ​. But also see ­later in the
book (chapter 5) to recognize that XX or XY are not always accurate indicators of
biological sex.
2. T. Laqueur. 1990. Making Sex: Body and Gender from the Greeks to Freud. Cam-
bridge, MA: Harvard University Press; V. Sanz. 2017. “No Way Out of the Binary: A
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.­org​/­10​.1­ 016​/­j​.­yhbeh​.­2023​.105445.
4. A. Fuentes. 2022. Race, Monogamy and Other Lies They Told You: Busting Myths
about H ­ uman Nature. 2nd ed. Oakland: University of California Press.
5. C. Kuzawa, H. T. Chugani, L. I. Grossman, L. Lipo­v ich, O. Muzik, P. Hof,
R. Patrick, D. E.Wildman, et al. 2014. “Metabolic Costs and Evolutionary Implica-
tions of ­Human Brain Development.” Proceedings of the National Acad­emy of Sciences
of the United States of Amer­i­ca 111 (36): 13010–15; G. Downey and D. H. Lende. 2012.
“Evolution and the Brain.” In The Encultured Brain: An Introduction to Neuroanthro-
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6. S. Han and Y. Ma. 2015. “A Culture-­Behavior-­Brain Loop Model of H ­ uman
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Brain. Oxford: Oxford University Press; C. C. Sherwood and A. Gomez-­Robles.
2017. “Brain Plasticity and ­Human Evolution.” Annual Reviews of Anthropology
46: 399–419; A. Fuentes. 2019. Why We Believe: Evolution and the ­Human Way of Being.
New Haven, CT: Yale University Press/Templeton Press.
7. M. Lock. 2015. “Comprehending the Body in the Era of the Epigenome.” Cur-
rent Anthropology 56 (2): 151–77; A. Fuentes. 2023. “­Humans Are Biocultural, Science
 Not e s to Ch a p t er 3 159

Should Be Too.” Science 382: 6672. https://­doi​.­org​/­10​.­1126​/­science.​ adl151; A. Fuen-

tes and P. Wiessner. 2016. “Reintegrating Anthropology: From Inside Out.” Current
Anthropology 57 (suppl. 13): 3–12; Downey and Lende, “Evolution and the Brain”;
T. Leatherman and A. H. Goodman. 2020. “Building on the Biocultural Syntheses:
20 Years and Still Expanding.” American Journal of ­Human Biology 32 (4): e23360;
T. Ingold and G. Paalson. 2013. Biosocial Becomings: Integrating Social and Biological
­Anthropology. Cambridge: Cambridge University Press.
8. Gender also includes individual expressions of identity expectations and ex-
periences. S. Nanda. 2000. Gender Diversity: Cross-­Cultural Variations. Long Grove,
IL: Waveland Press; J. Lorber. 2022. The New Gender Paradox. Oxford: Polity Press.
9. S. de Beauvoir. [1949] 2011. The Second Sex. Translated by J. Thurman. New York:
Vintage Books, 300.
10. Lorber, New Gender Paradox; J. S. Hyde, R. S. Bigler, D. Joel, C. C. Tate, and
S. M. van Anders. 2019. “The ­Future of Sex and Gender in Psy­chol­ogy: Five Chal-
lenges to the Gender Binary.” American Psychologist 74 (2): 171–93. https://­doi​.­org​
/­10​.­1037​/amp0000307; S. M. van Anders. 2022. “Gender/Sex/ual Diversity and
Biobehavioral Research.” Psy­chol­ogy of Sexual Orientation and Gender Diversity,
November 10. Advance online publication. https://­d x​.­doi​.­org​/­10​.­1037​/sgd00​
00609.
11. K. Cheslack-­Postava and R. M. Jordan-­Young. 2012. “Autism Spectrum Disor-
ders: T ­ oward a Gendered Embodiment Model.” Social Sciences and Medicine 74:
1667–74.
12. A. Zugman, L. M. Alliende, V. Medel, R. A. I. Bethlehem, J. Seidlitz, G. Ring-
lein, C. Arango, et al. 2023. “Country-­Level Gender I­ nequality Is Associated with
Structural Differences in the Brains of ­Women and Men.” Proceedings of the National
Acad­emy of Sciences of the United States of Amer­i­ca 120 (20): e2218782120. https://­doi​
.­org​/­10​.­1073​/­pnas​.2218782120.
13. This is a recent term and was not used by the socie­ties I reference h­ ere, as t­ here
­were gendered roles beyond the binary male/female in many of them. Generally, the
term t­ oday includes t­ hose individuals whose gender identity does not correspond
with the one assigned to them at birth by their culture, usually based on the assess-
ment of genitals at birth, and who seek to live in the other gender role.
14. R. Smith. 2021. “Imperial Terroir: ­Toward a Queer Molecular Ecol­ogy of
­Colonial Masculinities.” Current Anthropology 62 (suppl. 23): S155–­S168; K. Tallbear.
2015. “Settler Love Is Breaking My Heart.” The Critical Polyamorist Blog: Polyamory,
Indigeneity, and Cultural Politics, May 29. http://­w ww​.­criticalpolyamorist​.­com​
/­homeblog​/­settler​-­love​-­is​-­breaking​-­my​-heart; Q. McLamore. 2023. “Disarming
Transphobia: ‘Rapid-­Onset Gender Dysphoria’ Is a P ­ opular Weapon in the Anti-­
Trans Arsenal. It Is Nothing but Unscientific Bunk.” Aeon, January 23. https://­aeon​
.­co​/­essays​/­the​-­real​-­reason​-­why​-­theres​-­a​-­global​-­rise​-­in​-­trans​-youth.
 160 Not e s to Ch a p t er 3

15. Nanda, Gender Diversity. See also Hyde et al., “­Future of Sex and Gender in
Psy­chol­ogy.”
16. This is especially prominent in gender system of primarily patriarchal socie­
ties. A. Saini. 2023. The Patriarchs: The Origins of I­ nequality. New York: Penguin Press.
17. B. E. deMayo, A. E. Jordan, and K. R. Olson. 2022. “Gender Development in
Gender Diverse C ­ hildren.” Annual Review of Developmental Psy­chol­ogy 4 (1): 207–29;
Fuentes, Race, Monogamy and Other Lies; A. Fausto-­Sterling. 2020. Sexing the
Body: Gender Politics and the Construction of Sexuality. London: Hachette; C. Fine.
2010. Delusions of Gender: How Our Minds, Society, and Neurosexism Create Difference.
New York: W. W Norton.
18. S. M. van Anders. 2015. “Beyond Sexual Orientation: Integrating Gender/Sex
and Diverse Sexualities in Sexual Configurations Theory.” Archives of Sexual Be­hav­ior
44: 1177–213.
19. N. Krieger. 2019. “­Measures of Racism, Sexism, Heterosexism, and Gender
Binarism for Health Equity Research: From Structural Injustice to Embodied
Harm—­An Ecosocial Analy­sis.” Annual Review of Public Health 41: 37–62; M. Yudell,
D. Roberts, R. DeSalle, and S. Tishkoff. 2016. “Taking Race Out of ­Human Ge­ne­tics.”
Science 351 (6273): 564–65. https://­doi​.­org​/­10​.­1126​/­science​.aac4951; M. Lock. 2015.
“Comprehending the Body in the Era of the Epigenome.” Current Anthropology 56
(2): 151–77; Fuentes, “­Humans Are Biocultural”; Fuentes and Wiessner, “Reintegrat-
ing Anthropology”; Downey and Lende, “Evolution and the Brain”; Leatherman and
Goodman, “Building on the Biocultural Syntheses”; Ingold and Paalson, Biosocial
Becomings.
20. H ­ ere and in the following definitions I am drawing on a range of lit­er­a­ture,
but most specifically the excellent summaries of that lit­er­a­ture and the definitions
provided in Hyde et al., “­Future of Sex and Gender”; L. Z. DuBois and H. Shattuck-­
Heidorn. 2021. “Challenging the Binary: Gender/Sex and the Bio-­logics of Nor-
malcy.” American Journal of ­Human Biology 33 (5): e23623. https://­doi​.­org​/­10​.­1002​
/­ajhb.​ 23623; van Anders, “Gender/Sex/ual Diversity and Biobehavioral Research”;
and A. Fausto-­Sterling. 2019. “Gender/Sex, Sexual Orientation, and Identity Are in
the Body: How Did They Get ­There?” The Journal of Sex Research 56 (4–5): 529–55.
https://­doi​.­org​/­10​.­1080​/­0 0224499​.­2019​.1581883; S. Richardson. 2013. Sex Itself:
The Search for Male and Female in the H ­ uman Genome. Chicago: The University of
Chicago Press.
21. D. Joel. 2012. “Genetic-­Gonadal-­Genitals Sex (3G-­Sex) and the Misconcep-
tion of Brain and Gender, or, Why 3G-­Males and 3G-­Females Have Intersex Brain
and Intersex Gender.” Biology of Sex Differences 3 (1): 27; DuBois and Shattuck-­
Heidorn, “Challenging the Binary.”
22. The study might not explic­itly state this, but it is almost always the case that
when the researcher says “male” and “female,” they mean 3G male and 3G female,
 Not e s to Ch a p t er 4 161

but in real­ity, researchers are almost always only taking the “sex at birth” assignment
­unless stated other­wise in the a­ ctual research work.
23. Most researchers assume they are using something like 3G categories, but in
real­ity, they are often using self-­reported “sex” (male/female) or “gender” (man/
woman) or “sex” assessed based on physical observation of overall appearance. This
means that ­there is usually some small error in the assignments and thus the data are
often slightly askew.
24. M. Blackless, A. Charuvastra, A. Derryck, A. Fausto-­Sterling, K. Lauzanne,
and E. Lee. 2000. “How Sexually Dimorphic Are We? Review and Synthesis.” Ameri-
can Journal of ­Human Biology 12 (2): 151–66.

4. ­Humans Then
1. ­Here I leave out the marsupials, who are very fascinating, but not as central to
this discussion as are the placental mammals (as that is what ­humans are).
2. Relative to other nonmammal animals. Birds do have extended care, but not
at the physiological and behavioral level of mammals.
3. S. Hrdy. 2009. ­Mothers and ­Others: The Evolutionary Origins of Mutual Under-
standing. Cambridge, MA: Harvard University Press; E. Fernandez-­Duque, C. R.
Valeggia, and S. P. Mendoza. 2009. “The Biology of Paternal Care in H ­ uman and
Nonhuman Primates.” Annual Review of Anthropology 38 (1): 115–30; S. Rosenbaum
and J. B. Silk. 2022. “Pathways to Paternal Care in Primates.” Evolutionary Anthropol-
ogy 31 (5): 245–62. https://­doi​.­org​/­10​.­1002​/­evan​.21942.
4. Hrdy, ­Mothers and ­Others; C. Ross and A. MacLarnon. 2000. “The Evolution
of Non-­maternal Care in Anthropoid Primates: A Test of the Hypotheses.” Folia
Primatol (Basel) 71 (1–2): 93–113. https://­doi​.­org​/­10​.­1159​/000021733; S. Rosen-
baum and L. T. Gettler. 2018. “With a ­Little Help from Her Friends (and ­Family) Part
I: The Ecol­ogy and Evolution of Non-­maternal Care in Mammals.” Physiology &
Be­hav­ior 193, Part A: 1–11. https://­doi​.­org​/­10​.­1016​/­j​.­physbeh​.­2017​.­12​.025.
5. E. Clarke, K. Bradshaw, K. Drissell, P. Kadam, N. Rutter, and S. Vaglio. 2022.
“Primate Sex and Its Role in P ­ leasure, Dominance and Communication.” Animals 12
(23): 3301. https://­doi​.­org​/­10​.­3390​/ani12233301.
6. A. Whiten, R. A. Hinde, K. N. Laland, and C. B. Stringer. 2011. “Culture
Evolves.” Philosophical Transactions of the Royal Society of London: Biological Sciences
366 (1567): 938–48. https://­doi​.­org​/­10​.­1098​/­rstb​.­2010​.0372; H. Whitehead, K. N.
Laland, L. Rendell, R. Thorogood, and A. Whiten. 2019. “The Reach of Gene–­
Culture Coevolution in Animals.” Nature Communications 10 (2405): https://­doi​.­org​
/­10​.­1038​/­s41467​-­019​-­10293​-y.
7. J. M. Plavcan. 2012. “Sexual Size Dimorphism, Canine Dimorphism, and Male-­
Male Competition in Primates.” ­Human Nature 23: 45–67; but also see A. Fuentes.
 162 Not e s to Ch a p t er 4

2021. “Searching for the ‘Roots’ of Masculinity in Primates and the ­Human Evolution-
ary Past.” Current Anthropology 62: S13–­S25. https://­doi​.­org​/­10​.­1086​/711582; and the
discussion in this and the subsequent chapter in regard to h­ umans in this context.
8. N. Smit, B. Ngoubangoye, M. J. E. Charpentier, and E. Huchard. 2022. “Dynam-
ics of Intersexual Dominance in a Highly Dimorphic Primate.” Frontiers in Ecol­ogy
and Evolution 10: 931226. https://­doi​.­org​/­10​.­3389​/­fevo​.­2022​.931226.
9. P. M. Kappeler, E. Huchard, A. Baniel, C. Canteloup, M. J. E. Charpentier,
L. Cheng, E. Davidian, et al. 2022. “Sex and Dominance: How to Assess and Interpret
Intersexual Dominance Relationships in Mammalian Socie­ties.” Frontiers in Ecol­ogy
and Evolution 10: 918773. https://­doi​.­org​/­10​.­3389​/­fevo​.­2022​.918773; R. J. Lewis.
2018. “Female Power in Primates and the Phenomenon of Female Dominance.” An-
nual Review of Anthropology 47: 533–51. https://­doi​.­org​/­10​.­1146​/­annurev​-­anthro​
-­102317​-045958; R. J. Lewis. 2020. “Female Power: A New Framework for Under-
standing ‘Female Dominance’ in Lemurs.” Folia Primatologica 91: 48–68. https://­doi​
.­org​/­10​.­1159​/000500443.
10. G. Parker. 2006. “Sexual Conflict over Mating and Fertilization: An Overview.
Philosophical Transactions of the Royal Society: Biological Sciences 361: 235–59. https://­
doi​.­org​/­10​.­1098​/­rstb​.20 05.1785.
11. C. Campbell, A. Fuentes, K. MacKinnon, S. Bearder, and R. Stumpf. 2011.
Primates in Perspective. 2nd ed. Oxford: Oxford University Press, chap. 19.
12. Campbell et al., Primates in Perspective, chaps. 14 and 15.
13. L. T. Gettler. 2016. “Becoming DADS: Considering the Role of Cultural
­Context and Developmental Plasticity for Paternal Socioendocrinology.” Current
Anthropology 57: S38–­S51; E. Fernandez-­Duque, C. R. Valeggia, and S. P. Mendoza.
2009. “The Biology of Paternal Care in ­Human and Nonhuman Primates.” Annual
Review of Anthropology 38 (1): 115–30.
14. In fact, we sometimes see female dominance (such as in many lemurs).
15. T. Q. Bartlett and L. E. O. Light. “Sexual Dichromatism.” In The International
Encyclopedia of Primatology, edited by A. Fuentes. Hoboken: John Wiley & Sons, Inc.
https://­onlinelibrary​.­wiley​.­com​/­doi​/­10​.­1002​/­9781119179313​.wbprim0427.
16. Campbell et al., Primates in Perspective, chap. 17.
17. A. Kralick. 2023. “When Ape Sex ­Isn’t ­Simple.” Anthropology News, March 14.
https://­www​.­anthropology​-­news​.­org​/­articles​/­when​-­ape​-­sex​-­isnt​-­simple​/#citation.
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Hormone Profiles in Captive Male Orangutans: Implications for Understanding
Developmental Arrest.” American Journal of Physical Anthropology 109 (1): 19–32.
https://­doi​.­org​/­10​.­1002​/­(SICI)1096​-­8644(199905)109:1<19::AID​-­A JPA3>3​.­0​
.­CO;2​-­3​. And E. M. Thompson, A. Zhou, and C. D. Knott. 2012. “Low Testosterone
Correlates with Delayed Development in Male Orangutans.” PloS One. 7 (10):
e47282. https://­doi​.­org​/­10​.­1371​/­journal​.­pone​.0047282.
 Not e s to Ch a p t er 4 163

19. Campbell et al., Primates in Perspective, chap. 18.

20. Campbell et al., Primates in Perspective, chap. 19.
21. S. Rosenbaum, L. Vigilant, C. W. Kuzawa, and T. S. Stoinski. 2018. “Caring for
Infants Is Associated with Increased Reproductive Success for Male Mountain Goril-
las.” Scientific Reports 8: 15223.
22. Campbell et al., Primates in Perspective, chap. 20.
23. Campbell et al., Primates in Perspective, chap. 20.
24. A. Fuentes. 2018. “How H ­ umans and Apes Are Dif­fer­ent, and Why It M ­ atters.”
Journal of Anthropological Research 74 (2): 151–67. https://­doi​.­org​/­10​.­1086​/697150;
J. Marks. 2012. “The Biological Myth of ­Human Evolution.” Con­temporary Social
­Science: Journal of the Acad­emy of Social Sciences 7 (2): 139–57.
25. Actually, none of the hominins have even the teeny canine dimorphism one
sees in bonobos. G. Suwa, T. Sasaki, S. Semaw, M. J. Rogers, S. W. Simpson, Y. Kuni-
matsu, M. Nakatsukasa, et al. 2021. “Canine Sexual Dimorphism in Ardipithecus rami-
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United States of Amer­i­ca 118 (49): e2116630118. https://­doi​.­org​/­10​.­1073​/­pnas​
.2116630118.
26. N. Malone, A. H. Wade, A. Fuentes, E. Riley, M. Remis, and C. Jost-­Robinson.
2014. “Ethnoprimatology: Critical Interdisciplinarity and Multispecies Approaches
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27. S. Harmand, J. Lewis, and C. Feibel, C. J. Lepre, S. Prat, A. Lenoble, X. Boës,
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2017. The Creative Spark: How Imagination Made ­Humans Exceptional. New York:
Dutton.
28. P. Frémondière, L. Thollon, F. Marchal, F. Marchal, C. Fornai, N. Webb, and
M. Haeusler. 2022. “Dynamic Finite-­Element Simulations Reveal Early Origin of
Complex ­Human Birth Pattern.” Communications Biology 5 (377): https://­doi​.­org​/­10​
.­1038​/­s42003​-­022​-­03321​-z.
29. C. Kuzawa, H. T. Chugani, L. I. Grossman, L. Lipo­v ich, O. Muzik, P. Hof,
R. Patrick, D. E. Wildman, et al. 2014. “Metabolic Costs and Evolutionary Implica-
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of the United States of Amer­ic­ a 111 (36): 13010–15; K. R. Rosenberg. 2021. “The Evolu-
tion of ­Human Infancy: Why It Helps to Be Helpless.” Annual Review of Anthropology
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30. S. Lacy and C. Ocobock. 2023. “­Woman the Hunter: The Archaeological Evi-
dence.” American Anthropologist 126 (1): 19–31. https://­doi​.­org​/­10​.­1111​/­aman​.13914.
31. R. A. Foley. 2016. “Mosaic Evolution and the Pattern of Transitions in the
Hominin Lineage.” Philosophical Transactions of the Royal Society: Biological Sciences
371: 1–14; Fuentes, Creative Spark; Fuentes, “How ­Humans and Apes Are Dif­fer­ent”;
 164 Not e s to Ch a p t er 4

J. Galway-­Witham, J. Cole, and C. B. Stringer. 2019. “Aspects of ­Human Physical and

Behavioural Evolution during the Last 1 Million Years.” Journal of Quaternary Science
34 (6): 355–78.
32. Foley, “Mosaic Evolution”; Fuentes, Creative Spark; Fuentes, “How H ­ umans
and Apes Are Dif­fer­ent”; Galway-­Witham, Cole, and Stringer, “Aspects of H ­ uman
Physical and Behavioural Evolution.”
33. P. Spikins, A. Needham, B. Wright, C. Dytham, M. Gatta, and G. Hitchens.
2018. “Living to Fight Another Day: The Ecological and Evolutionary Significance of
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.­1016​/­j​.­quascirev​.­2018​.­08​.011; P. Spikins, C. Dytham, J. French, and S. John-­Wood.
2021. “Theoretical and Methodological Approaches to Ecological Changes, Social
Behaviour and ­Human Intergroup Tolerance 300,000 to 30,000 BP.” Journal of Ar-
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-­0 9503​-5.
34. Fuentes, Creative Spark; Fuentes, “How ­Humans and Apes Are Dif­fer­ent”;
S. Mithen. 2024. The Language Puzzle: Piecing Together the Six Million Year Story of How
Words Evolved. New York: Basic Books.
35. Actually, allocare is pretty common in highly social mammals (like canids,
some cats, mongooses, ­etc.).
36. Rosenbaum and Gettler, “With a ­Little Help from Her Friends Part I.”
37. Rosenberg, “Evolution of ­Human Infancy.”
38. Fuentes, Creative Spark; Fuentes, “How H ­ umans and Apes Are Dif­fer­ent”;
Galway-­Witham, Cole, and Stringer, “Aspects of H ­ uman Physical and Behavioural
Evolution.”
39. Hrdy, ­Mothers and ­Others.
40. L. T. Gettler. 2010. “Direct Male Care and Hominin Evolution: Why Male-­
Child Interaction Is More Than a Nice Social Idea.” American Anthropologist 112:
7–21; A. M. Kubicka, R. Wragg Sykes, A. Nowell, and E. Nelson. 2022. “Sexual
­Be­hav­ior in Neanderthals.” In The Cambridge Handbook of Evolutionary Perspectives
on Sexual Psy­chol­ogy, 516–60. Cambridge: Cambridge University Press.
41. K. Hawkes and J. Coxworth. 2013. “Grand­mothers and the Evolution of
­Human Longevity: A Review of Findings and ­Future Directions.” Evolutionary An-
thropology 22: 294–302. https://­doi​.­org​/­10​.­1002​/­evan​.21382.
42. Hrdy, ­Mothers and ­Others; Kuzawa et al., “Metabolic Costs”; Rosenberg,
“Evolution of H ­ uman Infancy”; Gettler, “Direct Male Care and Hominin Evolution”;
Gettler, “Becoming DADS.”
43. P. Spikins. 2015. How Compassion Made Us ­Human: The Evolutionary Origins
of Tenderness, Trust & Morality. Barnsley: Pen & Sword Archaeology; J. M. Burkart,
S. B. Hrdy, and C. P. Van Schaik. 2009. “Cooperative Breeding and ­Human Cognitive
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 Not e s to Ch a p t er 4 165

Nuclear ­Family: An Evolutionary Perspective on Parenting.” Current Opinion in Psy­

chol­ogy 7: 98–103. https://­doi​.­org​.­10​.­1016​/­j​.­copsyc​.­2015​.­0 8​.­013​. Gettler, “Direct
Male Care and Hominin Evolution”; Gettler, “Becoming DADS.”
44. See case study overview for Neanderthals h­ ere: Kubicka et al., “Sexual Be­
hav­ior in Neanderthals.” See also P. K. Stone and A. Netzer Zimmer. 2023. “Issues of
Gender, Identity, and Agency on Paleopathology.” In The Routledge Handbook
of ­Paleopathology, edited by A. L. Grauer, 435–56. Oxfordshire: Routledge.
45. G. R. Brown, K. N. Laland, and M. B. Mulder. 2009. “Bateman’s Princi­ples
and ­Human Sex Roles.” Trends in Ecol­ogy and Evolution 24 (6): 297–304. https://­doi​
.org/ 10.1016/j.tree.2009.02.005.
46. Brown, Laland, and Mulder, “Bateman’s Princi­ples and ­Human Sex Roles,”
297.
47. C. T. Ross, P. L. Hooper, J. E. Smith, A. V. Jaeggi, E. A. Smith, S. Gavrilets,
F. T. Zohora, et al. 2023. “Reproductive ­Inequality in ­Humans and Other Mammals.”
Proceedings of the National Acad­emy of Sciences of the United States of Amer­i­ca 120 (22):
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48. H. Dunsworth. 2020. “Expanding the Evolutionary Explanations for Sex Dif-
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­Reviews 29 (3): 108–16. https://­doi​.­org​/­10​.­1002​/­evan​.21834; A. Fuentes. 2016. “The
Extended Evolutionary Synthesis, Ethnography, and the H ­ uman Niche: ­Toward an
Integrated Anthropology.” Current Anthropology 57 (suppl. 13): 13–26. https://­doi​.­org​
/­10​.­1086​/685684; Gettler, “Direct Male Care and Hominin Evolution”; Sear, “Be-
yond the Nuclear F ­ amily”; R. Sear. 2021. “The Male Breadwinner Nuclear F ­ amily Is
Not the ‘Traditional’ H ­ uman F ­ amily, and Promotion of This Myth May Have Ad-
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49. Clarke et al., “Primate Sex and Its Role in ­Pleasure.”
50. M. Rodrigues. 2021. “Same Sex Sexual Be­hav­ior in Chimpanzees Challenge
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-­our​-­gendered​-­biases​-­about​-­evolution​/.
51. ­There are four subspecies of chimpanzees, and all have been reported to en-
gage in sexual be­hav­ior outside of reproduction, but not at the levels seen in
bonobos.
52. R. Stumpf. 2010. “Chimpanzees and Bonobos: Inter-­and Intra-­species
­Diversity.” In Primates in Perspective, 2nd ed., edited by C. Campbell, A. Fuentes,
K. MacKinnon, S. Bearder, and R. Stumpf, 340–56. Oxford: Oxford University Press;
A. Sandel and R. Reddy. 2021. “Sociosexual Behaviour in Wild Chimpanzees Occurs
in Variable Contexts and Is Frequent between Same-­Sex Partners.” Behaviour 158:
1–28. https://­doi​.­org​.­10​.­1163​/­1568539X​-bja10062.
 166 Not e s to Ch a p t er 4

53. G. Hohmann and B. Fruth. 2000. “Use and Function of Genital Contacts
among Female Bonobos.” Animal Be­hav­ior 60: 107–20. https://­doi​.­org​.­10​.­1006​/­anbe​
.­2000​.1451; L. R. Moscovice, M. Surbeck, B. Fruth, G. Hohmann, A. V. Jaeggi, and
T. Deschner. 2019. “The Cooperative Sex: Sexual Interactions among Female Bono-
bos Are Linked to Increases in Oxytocin, Proximity and Co­ali­tions.” Hormones and
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54. Sandel and Reddy, “Sociosexual Behaviour in Wild Chimpanzees.”
55. Sandel and Reddy, “Sociosexual Behaviour in Wild Chimpanzees”; J. Yamag-
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57. A. Fuentes. 2022. Race, Monogamy and Other Lies They Told You: Busting Myths
about ­Human Nature. 2nd ed. Oakland: University of California Press; K. Wellings,
M. Collumbien, E. Slaymaker, S. Singh, Z. Hodges, D. Patel, and N. Bajos.
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58. Again, for a case study in Neanderthals, see Kubicka et al., “Sexual Be­hav­ior
in Neanderthals.”
59. O. Lovejoy. [1981] 2009. “Reexamining ­Human Origins in Light of Ardipithe-
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60. Fuentes, Race, Monogamy and Other Lies.
61. For recent versions of this approach, see B. Chapais. 2010. Primeval Kinship:
How Pair-­Bonding Gave Birth to H ­ uman Society. Cambridge, MA: Harvard University
Press; and Lovejoy, “Reexamining ­Human Origins”; see also Ross et al., “Reproduc-
tive ­Inequality in ­Humans and Other Mammals.”
62. B. Chapais. 2010. Primeval Kinship: How Pair-­Bonding Gave Birth to H ­ uman
Society. Cambridge, MA: Harvard University Press.
63. Fuentes, Race, Monogamy and Other Lies.
64. A. Fuentes. 2009. Evolution of ­Human Be­hav­ior. Oxford: Oxford University
Press; and A. Fuentes, M. Wyczalkowski, and K. C. MacKinnon. 2010. “Niche Con-
struction through Cooperation: A Nonlinear Dynamics Contribution to Modeling
Facets of the Evolutionary History in the Genus Homo.” Current Anthropology 51 (3):
435–44; Fuentes Creative Spark.
65. See A. Fuentes. 1999. “Re-­evaluating Primate Monogamy.” American Anthro-
pologist 100 (4): 890–907; and A. Fuentes. 2002. “Patterns and Trends in Primate Pair
Bonds.” International Journal of Primatology 23 (4): 953–78.
66. Less than 3 ­percent of all mammalian species are monogamous.
 Not e s to Ch a p t er 4 167

67. See multiple chapters in P. T. Ellison and P. B. Gray, eds. 2009. The Endocrinol-
ogy of Social Relationships, 270–93. Cambridge, MA: Harvard University Press; and
J. T. Curtis and Z. Wang. 2003. “The Neurochemistry of Pair Bonding.” Current Direc-
tions in Psychological Science 12 (2): 49–53; and S. L. Díaz-­Muñoz, and K. L. Bales.
2016. “Monogamy.” In “Primates: Variability, Trends, and Synthesis.” Introduction to
special issue on Primate Monogamy. American Journal of Primatology 78 (3): 283–87.
https://­doi​.­org​/­10​.­1002​/­ajp​.22463.
68. Fuentes, Race, Monogamy and Other Lies.
69. See multiple chapters in Ellison and Gray, Endocrinology of Social Relation-
ships, 270–93; and Curtis and Wang, “Neurochemistry of Pair Bonding.”
70. However, just being a relative does not automatically generate a pair bond.
71. Fuentes, Creative Spark; M. W. Conkey. 2010. “Images without Words: The
Construction of Prehistoric Imaginaries for Definitions of ‘Us.’ ” Journal of Visual
Culture 9: 272–83; M. Kissel and A. Fuentes. 2017. “Semiosis in the Pleistocene.”
­Cambridge Archaeological Journal 27 (3): 1–16. https://­doi​.­org​/­10​.­1017​/S0959774​
317000014.
72. S. Nanda. 2014. Gender Diversity: Cross-­Cultural Variations. 2nd ed. Long
Grove, IL: Waveland Press; W. Wood and A. H. Eagly. 2002. “A Cross-­Cultural
­Analy­sis of the Be­hav­ior of ­Women and Men: Implications for the Origins of Sex
Differences.” Psychological Bulletin 128 (5): 699–727; J. Lorber. 2022. The New Gender
Paradox. Oxford: Polity Press.
73. S. Lacy and C. Ocobock. 2023. “­Woman the Hunter: The Archaeological Evi-
dence.” American Anthropologist 126 (1): 19–31. https://­doi​.­org​/­10​.­1111​/­aman​.13914.
74. L. C. K. Aiello. 2002. “Energetic Consequences of Being a Homo erectus
­Female.” American Journal of ­Human Biology 14: 551–65; L. C. Aiello and J. C. K. Wells.
2002. “Energetics and the Evolution of the Genus Homo.” Annual Review of Anthropol-
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75. D. Haraway. 1989. Primate Visions: Gender, Race, and Nature in the World of
Modern Science. Oxfordshire: Routledge; Fuentes, Creative Spark; Conkey, “Images
without Words.”
76. Lacy and Ocobock, “­Woman the Hunter: The Archaeological Evidence.”
77. Stone and Zimmer, “Issues of Gender, Identity, and Agency on Paleopathol-
ogy”; Fuentes, Creative Spark; Lacy and Ocobock, “­Woman the Hunter: The Ar-
chaeological Evidence”; C. Ocobock and S. Lacy. 2023. “­Woman the Hunter: The
Physiological Evidence.” American Anthropologist 126 (1): 7–18. https://­doi​.­org​/­10​
.­1111​/­aman​.13915.
78. See J. M. Adovasio, O. Soffer, and J. Page. 2007. The Invisible Sex: Uncovering
the True Roles of ­Women in Prehistory. Washington, DC: Smithsonian Books; Stone
and Zimmer, “Issues of Gender, Identity, and Agency on Paleopathology”; Conkey,
“Images without Words.”
 168 Not e s to Ch a p t er 4

79. D. Stout. 2011. “Stone Toolmaking and the Evolution of ­Human Culture and
Cognition.” Philosophical Transactions of the Royal Society B 366 (1567): 1050–59;
D. Stout and N. Khreisheh. 2015. “Skill Learning and ­Human Brain Evolution: An
Experimental Approach.” Cambridge Archaeological Journal 25: 867–75. https://­doi​
.­org​/­10​.­1017​/S0959774315000359; Lacy and Ocobock, “­Woman the Hunter: The
Archaeological Evidence.”
80. K. W. Arthur. 2010. “Feminine Knowledge and Skill Reconsidered: ­Women
and Flaked Stone Tools.” American Anthropologist 112 (2): 228–43; J. Gero. 1991.
“Genderlithics: W ­ omen’s Roles in Stone-­Tool Production.” In Engendering
­Archaeology: ­Women and Prehistory, edited by J. Gero and M. W. Conkey, 163–93.
Oxford: Blackwell; K. Weedman. 2005. “Gender and Stone Tools: An Ethnographic
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81. A. Murray, R. Pinhasi, and J. Stock. 2014. “Lower Limb Skeletal Biomechan-
ics Track Long-­Term Decline in Mobility across ∼6150 Years of Agriculture in Cen-
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.­jas​.­2014​.­0 9​.­0 01​. A. A. Macintosh, R. Pinhasi, and J. T. Stock. 2017. “Prehistoric
­Women’s Manual L ­ abor Exceeded That of Athletes through the First 5500 Years of
Farming in Central E ­ urope.” Science Advances 3 (11): eaao3893. https://­doi​.­org​/­10​
.­1126​/­sciadv​.aao3893; Lacy and Ocobock, “­Woman the Hunter: The Archaeologi-
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82. Ocobock and Lacy, “­Woman the Hunter: The Physiological Evidence.”
83. E.g. Lacy and Ocobock, “­Woman the Hunter: The Archaeological Evidence”;
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Hunters of the Early Amer­i­cas.” Science Advances 6 (45): eabd0310. https://­doi​.­org​
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Do? The Division of L ­ abor among Neandertals and Modern H ­ umans in Eurasia.”
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A. A. Sigma, S. Chilczuk, K. Nelson, R. Ruther, and C. Wall-­Scheffler. 2023. “The
Myth of Man the Hunter: W ­ omen’s Contribution to the Hunt across Ethnographic
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85. A. Estalrrich and A. Rosas. 2015. “Division of ­Labor by Sex and Age in Nean-
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 Not e s to Ch a p t er 4 169

of ­Human Evolution 80: 51–63; see also Kubicka et al., “Sexual Be­hav­ior in
Neanderthals.”
86. Spikins et al., “Living to Fight Another Day”; Spikins et al., “Theoretical and
Methodological Approaches.”
87. D. Snow. 2013. “Sexual Dimorphism in E ­ uropean Upper Paleolithic Cave Art.”
American Antiquity 78: 746–61. https://­doi​.­org​/­10​.­7183​/­0 002​-­7316​.­78​.­4​.746;
­Conkey, “Images without Words”; Lacy and Ocobock, “­Woman the Hunter: The
Archaeological Evidence.”
88. P. Sorokowski and M. Kowal. 2024. “Relationship between the 2D:4D and Pre-
natal Testosterone, Adult Level Testosterone, and Testosterone Change: Meta-­analysis
of 54 Studies.” American Journal of Biological Anthropology 183 (1): 20–38. https://­doi​.­org​
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“No Sex Difference in Digit Ratios (2D:4D) in the Traditional Yali of Papua and Its
Meaning for the Previous Hypotheses on the Inter-­populational Variability in 2D:4D.”
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89. B. McCauley, D. Maxwell, and M. Collard. 2018. “A Cross-­cultural Perspective
on Upper Palaeolithic Hand Images with Missing Phalanges.” Journal of Paleolithic
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90. Fuentes, Creative Spark; A. Fuentes. 2019. Why We Believe: Evolution and the
­Human Way of Being. New Haven, CT: Yale University Press/Templeton Press;
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91. Fuentes, Creative Spark; Lacy and Ocobock, “­Woman the Hunter: The Ar-
chaeological Evidence”; Ocobock and Lacy, “­Woman the Hunter: The Physiological
Evidence.”
92. For example, M. Cintas-­Peña and L. García Sanjuán. 2019. “Gender Inequali-
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­Neolithic Near East. Oxford: Oxford University Press; M. Cintas-­Peña, M. Luciañez-­
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 170 Not e s to Ch a p t er 5

5. ­Humans Now
1. Note I am specifically stating male and female (3G), not man and ­woman . . . ​
if one used only gender categories, t­ here would be even more mixing and less con-
sistency between one of the categories and height.
2. A. Fausto-­Sterling. 2020. Sexing the Body: Gender Politics and the Construction
of Sexuality. London: Hachette.
3. See for example C. Sanchis-­Segura and R. R. Wilcox. 2024. “From Means to
Meaning in the Study of Sex/Gender Differences and Similarities.” Front Neuroen-
docrinol 73: 101133. https://­doi​.­org​/­10​.­1016​/­j​.­yfrne​.­2024​.101133.
4. C. Ruff. 2002. “Variation in ­Human Body Size and Shape.” Annual Reviews in
Anthropology 31: 211–32; A. Fuentes. 2022. Race, Monogamy and Other Lies They Told
You: Busting Myths about H ­ uman Nature. 2nd ed. Oakland: University of California
Press.
5. Fausto-­Sterling, Sexing the Body; L. Z. DuBois and H. Shattuck-­Heidorn. 2021.
“Challenging the Binary: Gender/Sex and the Bio-­logics of Normalcy.” American
Journal of ­Human Biology 33 (5): e23623; Fuentes, Race, Monogamy and Other Lies;
W. D. Lassek and S. J. C. Gaulin. 2022. “Substantial but Misunderstood ­Human
Sexual Dimorphism Results Mainly from Sexual Se­lection on Males and Natu­ral
Se­lection on Females.” Frontiers in Psy­chol­ogy 17 (13): 859931. https://­doi​.­org​/­10​.­3389​
/­fpsyg​.­2022​.859931.
6. See the National Center for Health Statistics. https://­www​.­cdc​.­gov​/­nchs​
/­nhanes​/­index​.htm.
7. C. D. Fryar, M. D. Carroll, Q. Gu, J. Afful, and C. L. Ogden. 2021. “Anthropo-
metric Reference Data for ­Children and Adults: United States, 2015–2018.” Vital
Health Statistics (36): 1–44.
8. H. M. Dunsworth. 2020. “Expanding the Evolutionary Explanations for Sex
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https://­doi​.­org​/­10​.­1002​/­evan​.21834.
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34. The variation in hair patterns across our species has been mispresented and
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Sex Differences.” Biology of Sex Differences 15:14. https://­doi​.­org​/­10​.­1186​/­s13293​-­024​
-­0 0585​-4.
77. M. Ingalhalikar, A. Smith, D. Parker, T. D. Satterthwaite, M. A. Elliott, K. Ruparel,
H. Hakonarson, et al. 2014. “Sex Differences in the Structural Connectome of the
­Human Brain.” Proceedings of the National Acad­emy of Sciences of the United States of
Amer­i­ca 111 (2): 823–28. https://­doi​.­org​/­10​.­1073​/­pnas​.1316909110; J. M. Rauch and
L. Eliot. 2022. “Breaking the Binary: Gender versus Sex Analy­sis in H ­ uman Brain
Imaging.” Neuroimage 264: 119732. https://­doi​.­org​/­10​.­1016​/­j​.­neuroimage​.­2022​
.119732; Rippon, Gendered Brain; R. Jordan-­Young. 2011. Brainstorm: The Flaws in the
Science of Brain Differences. Cambridge, MA: Harvard University Press.
78. G. Downey and D. H. Lende. 2012. “Evolution and the Brain.” In The Encul-
tured Brain: An Introduction to Neuroanthropology, edited by D. H. Lende and
G. Downey, 103–38. Cambridge, MA: MIT Press.
 178 Not e s to Ch a p t er 5

79. Rauch and Eliot, “Breaking the Binary”; Eliot, “Remembering the Null Hypoth-
esis”; Rippon, Gendered Brain; L. Eliot, A. K. Beery, E. G. Jacobs, H. F. LeBlanc, D. L.
Maney, and M. M. McCarthy. 2023. “Why and How to Account for Sex and Gender in
Brain and Behavioral Research.” Journal of Neuroscience 143 (37): 6344–56. https://­doi​
.­org​/1­ 0​.­1523​/J­ NEUROSCI​.­0 020​-­23​.2023; A. Kaiser. 2012. “Re-­Conceptualizing ‘Sex’
and ‘Gender’ in the H ­ uman Brain.” Zeitschrift für Psychologie 220: 130–36; D. L. Maney.
2015. “Just Like a Circus: The Public Consumption of Sex Differences.” Current Topics
in Behavioral Neuroscience 19: 279–96. https://­doi​.­org​/­10​.­1007​/­7854​_­2014​_339.
80. DeCasien et al., “Sex Differences in the ­Human Brain.”
81. Eliot et al., “Dump the ‘Dimorphism’ ”; Eliot, “Remembering the Null
Hypothesis.”
82. S. Richardson. 2013. Sex Itself: The Search for Male and Female in the H ­ uman
Genome. Chicago: The University of Chicago Press; Kung et al., “Prenatal Androgen
Exposure and Sex-­Typical Play Behaviour.”
83. Many of the genes on the X chromosome are not related directly or only to
reproductive biology, but most of the genes on the Y are.
84. Prum, ­Performance All the Way Down.
85. B. Croft, T. Ohnesorg, J. Hewitt, J. Bowles, A. Quinn, J. Tan, V. Corbin, et al.
2018. “­Human Sex Reversal Is Caused by Duplication or Deletion of Core Enhancers
Upstream of SOX9.” Nature Communications 9 (1): 5319. https://­doi​.­org​/­10​.­1038​
/­s41467​-­018​-­07784​-9; P. Bernard and V. R. Harley. 2007. “Wnt4 Action in Gonadal
Development and Sex Determination.” The International Journal of Biochemistry & Cell
Biology 39 (1): 31–43. https://­doi​.­org​/­10​.­1016​/­j.​ ­biocel​.­2006​.­06​.007; K. K. Niakan and
E. R. B. McCabe. 2005. “DAX1 Origin, Function, and Novel Role.” Molecular Ge­ne­tics
and Metabolism 86 (1): 70–83. https://­doi​.­org​/­10​.­1016​/­j​.­ymgme​.­2005​.­07​.019.
86. Fausto-­Sterling, Sexing the Body; Prum, ­Performance All the Way Down.
87. O. M. Muñoz-­A guirre, S. Kim-­Hellmuth, V. Wucher, A. D. H. Gewirtz,
D. J. Cotter, P. Parsana, S. Kasela, et al. 2020. “The Impact of Sex on Gene Expression
across ­Human Tissues.” Science 369 (6509): eaba3066. https://­doi​.­org​/­10​.­1126​
/­science​.aba3066.
88. M. Gershoni and S. Pietrokovski. 2017. “The Landscape of Sex-­Differential
Transcriptome and Its Consequent Se­lection in ­Human Adults.” BMC Biol 15 (7).
https://­doi​.­org​/­10​.­1186​/­s12915​-­017​-­0352​-z.
89. Richardson, Sex Itself.
90. A. Wiley. 2021. “Pearl Lecture: Biological Normalcy: A New Framework for
Biocultural Analy­sis of H ­ uman Population Variation.” American Journal of ­Human
Biology 33 (5): e23563. https://­doi​.­org​/­10​.­1002​/­ajhb​.23563.
91. See Fausto-­Sterling, Sexing the Body, for an excellent and extensive overview
of ­these patterns and their explanations and impacts. See also C. Ainsworth. 2015.
“Sex Redefined: The Idea of 2 Sexes Is Overly Simplistic.” Scientific American,
 Not e s to Ch a p t er 6 179

February 8. https://­www​.­scientificamerican​.­com​/­article​/­sex​-­redefined​-­the​-­idea​-­of​
-­2​-­sexes​-­is​-­overly​-simplistic1; and Richardson, Sex Itself.
92. Hyde et al., “The ­Future of Sex and Gender in Psy­chol­ogy”; and Fausto-­
Sterling, Sexing the Body; S. M. van Anders. 2022. “Gender/Sex/ual Diversity and
Biobehavioral Research.” Psy­chol­ogy of Sexual Orientation and Gender Diversity, No-
vember 10. Advance online publication. https://­dx​.­doi​.­org​/­10​.­1037​/sgd0000609.

6. No Biological B
­ attle of the Sexes
1. This book has sold more than fifteen million copies, been reprinted many times,
and remains a near best seller in the 2020s. John Gray’s website offers (for a fee) to
help you know who your soul mate is, increase intimacy and passion, create romance
with your partner, and enjoy a lifetime of ­great sex (Marsvenus​.­com).
2. C. Darwin. 1871. The Descent of Man, and Se­lection in Relation to Sex. London:
John Murray, 401.
3. A. J. Bateman. 1948. “Intra-­sexual Se­lection in Drosophila.” Heredity 2: 349–68;
E. O. Wilson. 1979. “Sex and ­Human Nature.” The Wilson Quarterly 3 (4): 92–105.
4. H. Dunsworth. 2021. “This View of Wife.” In A Most In­ter­est­ing Prob­lem, edited
by J. DaSilva, chap. 9. Prince­ton, NJ: Prince­ton University Press; L. Cooke. 2021.
Bitch: On the Female of the Species. New York: Basic Books; M. Landau. 1991. N ­ arrative
of ­Human Evolution. New Haven, CT: Yale University Press; A. Fausto-­Sterling. 1992.
Myths of Gender. New York: Basic Books; M. Midgley. 2003. The Myths We Live By.
Oxfordshire: Routledge; A. Saini. 2017. Inferior: How Science Got ­Women Wrong—­
and the New Research That’s Rewriting the Story. Boston: Beacon Press.
5. W. D. Lassek and S. J. C. Gaulin. 2022. “Substantial but Misunderstood ­Human
Sexual Dimorphism Results Mainly from Sexual Se­lection on Males and Natu­ral
Se­lection on Females.” Frontiers in Psy­chol­ogy 13: 859931. https://­doi​.­org​/­10​.­3389​
/­fpsyg​.­2022​.­859931​. ­These authors state specifically that (1) sexual dimorphism in
stature, fat mass, and fat distribution have been significantly s­ haped by disruptive
natural-­selection regimes operating on females and males, with some likely overlay
of subsequent sexual se­lection acting via mate choice in the case of fat distribution;
(2) sexual dimorphism in lean mass, muscle mass, and strength are largely due to
sexual se­lection arising from a long history of aggressive male mating competition,
with the some pos­si­ble influence of divergent natu­ral se­lection due to sex differences
in foraging ecol­ogy; and (3) a large lit­er­a­ture seems to have overemphasized the role
of mate choice, and underestimated the role of male competition for mates, in shap-
ing ­human sex differences. For a version of this same argument that focuses on the
female need to stay healthy to raise young, see h­ ere: J. F. Benenson, C. E. Webb, and
R. W. Wrangham. 2021. “Self-­Protection as an Adaptive Female Strategy.” Behavioral
and Brain Sciences 45: e128. https://­doi​.­org​/­10​.­1017​/S0140525X21002417.
 180 Not e s to Ch a p t er 6

6. See C. Hooven. 2021. T: The Story of Testosterone, the Hormone That Dominates
and Divides Us. New York: Henry Holt and Com­pany, 25.
7. See, for example, R. Jordan-­Young and K. Karkazis. 2019. Testosterone: An
­Unauthorized Biography. Cambridge, MA: Harvard University Press; C. Fine. 2016.
Testosterone Rex: Myths of Sex, Science and Society. New York: W. W. Norton; S. M. van
Anders. 2013. “Beyond Masculinity: Testosterone, Gender/Sex, and ­Human Social
Be­hav­ior in a Comparative Context.” Frontiers in Neuroendocrinology 34: 198–210.
https://­doi​.­org​/­10​.­1016​/­j​.­y frne​.­2 013​.­07​.001; S. M. van Anders, J. Steiger, and
K. L. Goldey. 2015. “Effects of Gendered Be­hav­ior on Testosterone in W ­ omen and
Men.” Proceedings of the National Acad­emy of Sciences of the United States of Amer­i­ca 112:
13805–10; A. Fausto-­Sterling. 2020. Sexing the Body: Gender Politics and the Construction
of Sexuality. London: Hachette; S. Richardson. 2013. Sex Itself: The Search for Male and
Female in the H­ uman Genome. Chicago: The University of Chicago Press; L. T. Gettler.
2016. “Becoming DADS: Considering the Role of Cultural Context and Developmen-
tal Plasticity for Paternal Socioendocrinology.” Current Anthropology 57: S38–­S51.
8. Meta-­analyses are systematic syntheses of many studies.
9. As usual, ­these studies are mostly on ­people in North Amer­i­ca and ­Europe.
Thus, they are prob­ably underrepresenting the broader variation in ­humans across
the planet.
10. While the authors use the terms “male” and “female,” they are actually assess-
ing gender (men/women, boys/girls) in ­these studies. J. Shibley Hyde. 2005. “The
Gender Similarities Hypothesis.” American Psychologist 60 (6): 581–92; E. Zell,
Z. Krizan, and S. R. Teeter. 2015. “Evaluating Gender Similarities and Differences Using
Metasynthesis.” American Psychologist 70: 10–20. https://d­ oi.​ ­org​/1­ 0​.­1037​/a0038208.
11. Zell, Krizan, and Teeter, “Evaluating Gender Similarities and Differences.”
12. R. W. Wrangham and M. L. Wilson. 2004. “Collective Vio­lence—­Comparisons
between Youths and Chimpanzees.” In Youth Vio­lence: Scientific Approaches to P ­ revention,
edited by J. Devine, J. Gilligan, K. A. Miczek, R. Shaikh, and D. Pfaff, 233–56. New
York: New York Acad­emy of Sciences; Hooven, T: The Story of Testosterone, 25; Be-
nenson, Webb, and Wrangham, “Self-­Protection as an Adaptive Female Strategy”;
Lassek and Gaulin, “Substantial but Misunderstood H ­ uman Sexual Dimorphism”;
D. Buss and D. P. Schmitt. 2019. “Mate Preferences and Their Behavioral Manifesta-
tions.” Annual Review of Psy­chol­ogy 70: 77–110. See also previous mention of Darwin,
E. O. Wilson, Robert Trivers, e­ tc. (chapter 3). See also R. Nelson. 2021. “The Sex in
Your Vio­lence: Patriarchy and Power in Anthropological World Building and Every-
day Life.” Current Anthropology 62: S92–­S102. https://­doi​.­org​/­10​.­1086​/711605.
13. M. G. Alexander and T. D. Fisher. 2002. “Truth and Consequences: Using the
Bogus Pipeline to Examine Sex Differences in Self-­Reported Sexuality.” Journal of
Sex Research 40 (1): 27–35; H. Donnan and F. Magowan. 2010. The Anthropology
of Sex. Oxfordshire: Routledge.
 Not e s to Ch a p t er 6 181

14. R. Thomsen. 2013. “Masturbation in Non-­Human Primates.” In The Encyclo-

pedia of ­Human Sexuality, vol. 1. Boston: Wiley-­Blackwell Press.
15. Fausto-­Sterling, Sexing the Body.
16. The statistics are summarized data presented in the categories used by the
main data article: D. Herbenick, M. ­Reece, V. Schick, S. A. Sanders, B. Dodge, and
J. D. Fortenberry. 2010. “Sexual Be­hav­ior in the United States: Results from a Na-
tional Probability Sample of Men and ­Women Ages 14–94.” Journal of Sexual Medi-
cine 7 (suppl. 5): 255–65.
17. The authors of the study make the point that their survey undersampled ho-
mosexually oriented populations.
18. C. Ryan and C. Jetha. 2010. Sex at Dawn: The Prehistoric Origins of Modern
Sexuality. New York: Harper; Donnan and Magowan, Anthropology of Sex; J. L. ­Carroll.
2009. Sexuality Now: Embracing Diversity, 3rd ed. Boston: Cenage Learning; and also
J. DeLamater and W. N. Friedrich. 2002. “­Human Sexual Development.” Journal of
Sex Research 39 (1): 10–14.
19. See S. T. Lindau, L. P. Schumm, E. O. Laumann, W. Levinson, C. A.
O’Muircheartaigh, and L. J. Waite. 2007. “A Study of Sexuality and Health among
Older Adults in the United States.” New E ­ ngland Journal of Medicine 357 (8): 762–74;
and A. Brewis and M. Meyer. 2005. “Marital Coitus across the Life Course.” Journal
of Biocultural Science 37: 499–518.
20. D. P. Schmitt. 2005. “Sociosexuality from Argentina to Zimbabwe: A
48-­Nation Study of Sex, Culture, and Strategies of ­Human Mating.” Behavioral and
Brain Sciences 28: 247–311; but see also the update on this study for more complexity
and a less clear set of assertions about mate choice patterns than in the 1993 and 2005
original studies: Buss and Schmitt, “Mate Preferences and Their Behavioral
Manifestations.”
21. K. Wellings, M. Collumbien, E. Slaymaker, S. Singh, Z. Hodges, D. Patel, and
N. Bajos. 2006. “Sexual Behaviour in Context: A Global Perspective.” Lancet 368
(9548): 1706–28. https://­doi​.­org​/­10​.­1016​/­S0140​-­6736(06)69479​-8.
22. Gettler, “Becoming DADS.”
23. M. Cappelletti and K. Wallen. 2016. “Increasing ­Women’s Sexual Desire: The
Comparative Effectiveness of Estrogens and Androgens.” Hormones and Be­hav­ior 78:
178–93. https://­doi​.­org​/­10​.­1016​/­j​.­yhbeh​.­2015​.­11​.003; S. J. Parish, J. A. Simon, S. R.
Davis, A. Giraldi, I. Goldstein, S. W. Goldstein, N. N. Kim, et al. 2021. “International
Society for the Study of ­Women’s Sexual Health Clinical Practice Guideline for the
Use of Systemic Testosterone for Hypoactive Sexual Desire Disorder in ­Women.”
Climacteric 24 (6): 533–50. https://­doi​.­org​/­10​.­1080​/­13697137​.­2021​.1891773.
24. S. M. van Anders. 2012. “Testosterone and Sexual Desire in Healthy ­Women
and Men.” Archives of Sexual Be­hav­ior 41 (6): 1471–84. https://­doi​.­org​/­10​.­1007​
/­s10508​-­012​-­9 946​-2.
 182 Not e s to Ch a p t er 6

25. Schmitt, “Sociosexuality from Argentina to Zimbabwe,” 273.

26. In fact, t­ here is good evidence that gender roles and cultural pressures affect
the way ­people respond to questions about sexuality. See Alexander and Fisher,
“Truth and Consequences.”
27. W. Wood and A. H. Eagly. 2002. “A Cross-­Cultural Analy­sis of the Be­hav­ior
of ­Women and Men: Implications for the Origins of Sex Differences.” Psychological
Bulletin 128 (5): 699–727; and J. Archer. 2009. “Does Sexual Se­lection Explain
­Human Sex Differences in Aggression?” Behavioral and Brain Sciences 32: 249–311.
28. R. Bribiescas. 2021. “Evolutionary and Life History Insights into Masculinity
and Warfare: Opportunities and Limitations.” Current Anthropology 62: S23.
29. ­There is ­little extensive data on vio­lence and homosexual partnerships.
30. J. Archer. 2019. “The Real­ity and Evolutionary Significance of Psychologi-
cal Sex Differences.” Biological Reviews 94 (4): 1381–1415. https://­doi​.­org​/­10​.­1111​
/­brv​.12507; and Zell, Krizan, and Teeter, “Evaluating Gender Similarities and
Differences.”
31. M. Gutmann. 2019. Are Men Animals? New York: Basic Books; M. Guttman,
R. G. Nelson, and A. Fuentes. 2021. “Epidemic Errors in Understanding Masculinity,
Maleness, and Vio­lence.” Current Anthropology 62: S5–­S12. https://­doi​.­org​/­10​.­1086​
/712485; A. Fuentes. 2022. Race, Monogamy and Other Lies They Told You: Busting
Myths about ­Human Nature. 2nd ed. Oakland: University of California Press.
32. ­There is a wide array of definitions and framings of sexual assault that vary by
and are related to cultural and gendered beliefs, roles, laws, and experiences. This
makes global assessments of what percentage of men perpetrate it, and how to ac-
curately ­measure global patterns complex. However, the WHO definition is a good
working frame: “any sexual act, attempt to obtain a sexual act, unwanted sexual com-
ments or advances, or acts to traffic, or other­wise directed, against a person’s sexual-
ity using coercion, by any person regardless of their relationship to the victim, in any
setting, including but not l­imited to home and work.” See P. A. Fernandez. 2011.
“Sexual Assault: An Overview and Implications for Counselling Support.” Austral-
asian Medical Journal 4 (11): 596–602. https://­doi​.­org​/­10​.­4 066​/­AMJ​.2011858; and
R. Jewkes, C. Garcia-­Moren, and P. Sen. 2002. “Sexual Vio­lence.” In World Report on
Vio­lence and Health, edited by E. G. Krug, L. L. Dahlberg, J. A. Mercy, A. B. Zwi, and
R. Lozano, 147–74. Geneva: World Health ­Organization.
33. Nelson, “The Sex in Your Vio­lence”; Fernandez, “Sexual Assault.”
34. See R. W. Sussman and C.R. Cloninger. 2011. Cooperation and Altruism. New
York: Springer; and D. Fry. 2007. Beyond War: The ­Human Potential for Peace. Oxford:
Oxford University Press; Fuentes, Race, Monogamy and Other Lies; A. Fuentes. 2021.
“Searching for the ‘Roots’ of Masculinity in Primates and the H ­ uman Evolutionary
Past.” Current Anthropology 62: S13–­S25. https://­doi​.­org​/1­ 0​.­1086/​ 711582; Guttman,
Are Men Animals?
 Not e s to Ch a p t er 6 183

35. G. Rippon. 2019. Gendered Brain: The New Neuroscience That Shatters the Myth
of the Female Brain. London: The Bodley Head Ltd.; D. Joel, Z. Berman, I. Tavor,
N. Wexler, O. Gaber, Y. Stein, N. Shefi, et al. 2015. “Sex Beyond the Genitalia: The
­Human Brain Mosaic.” Proceedings of the National Acad­emy of Sciences of the United
States of Amer­i­ca 112 (50): 15468–73; D. Joel. 2012. “Genetic-­Gonadal-­Genitals Sex
(3G-­Sex) and the Misconception of Brain and Gender, or, Why 3G-­Males and
­3G-­Females Have Intersex Brain and Intersex Gender.” Biology of Sex Differences 3
(1): 27; S. Liu, J. Seidlitz, J. D. Blumenthal, L. S. Clasen, and A. Raznahan. 2020.
“Integrative Structural, Functional, and Transcriptomic Analyses of Sex-­Biased Brain
­Organization in ­Humans.” Proceedings of the National Acad­emy of Sciences of the
United States of Amer­i­c a 117 (31): 18788–98. https://­d oi​ .­o rg​ /­10​ .­1073​ /­p nas​
.1919091117; M. Hirnstein, K. Hugdahl, and M. Hausmann. 2019. “Cognitive Sex
Differences and Hemispheric Asymmetry: A Critical Review of 40 Years of Re-
search.” Laterality: Asymmetries of Body, Brain and Cognition 24 (2): 204–52. https://­
doi​.­org​/­10​.­1080​/­1357650x​.­2018​.1497044.
36. Joel et al., “Sex Beyond the Genitalia.”
37. L. Eliot, A. Ahmed, H. Khan, and J. Patel. 2021. “Dump the ‘Dimorphism’:
Comprehensive Synthesis of ­Human Brain Studies Reveals Few Male-­Female
­Differences beyond Size.” Neuroscience and Biobehavioral Reviews 125: 667–97.
https://d­ oi.​ ­org/​ 1­ 0.​ ­1016/​ j­ .​ ­neubiorev.​ ­2021.​ ­02.​ 026; L. Eliot. 2024. “Remembering the
Null Hypothesis When Searching for Brain Sex Differences.” Biology of Sex Differ-
ences 15:14. https://­doi​.­org​/­10​.­1186​/­s13293​-­024​-­0 0585​-4.
38. L. S. B. Hrdy. 2009. ­Mother and ­Others: The Evolutionary Origins of Mutual
Understanding. New York: Belknap; C. Kuzawa, H. T. Chugani, L. I. Grossman,
L. Lipo­v ich, O. Muzik, P. Hof, R. Patrick, D. E. Wildman, et al. 2014. “Metabolic
Costs and Evolutionary Implications of ­Human Brain Development.” Proceedings of
the National Acad­emy of Sciences of the United States of Amer­ic­ a 111 (36): 13010–15;
K. R. Rosenberg. 2021. “The Evolution of ­Human Infancy: Why It Helps to Be Help-
less.” Annual Review of Anthropology 50: 423–40. https://­doi​.­org​/­10​.­1146​/­annurev​
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39. K. L. Bales, C. S. Ardekani, A. Baxter, C. L. Karaskiewicz, J. X. Kuske, A. R. Lau,
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40. See A. Fuentes. 2017. The Creative Spark: How Imagination Made ­Humans
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 184 Not e s to Ch a p t er 7

41. J. S. Hyde, R. S. Bigler, D. Joel, C. C. Tate, and S. M. van Anders. 2019. “The
­ uture of Sex and Gender in Psy­chol­ogy: Five Challenges to the Gender Binary.”
F
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7. Why the Binary View Is a Prob­lem

1. K. O. Smiley, K. M. Munley, K. Aghi, S. E. Lipshutz, T. M. Patton, D. S. Prad-
han, T. K. Solomon-­Lane, et al. 2024. “Sex Diversity in the 21st ­Century: Concepts,
Frameworks, and Approaches for the ­Future of Neuroendocrinology.” Hormones and
Be­hav­ior 157: 105445. https://­doi​.­org​/­10​.­1016​/­j​.­yhbeh​.­2023​.105445.
2. S. M. van Anders. 2013. “Beyond Masculinity: Testosterone, Gender/Sex, and
­Human Social Be­hav­ior in a Comparative Context.” Frontiers in Neuroendocrinology
34: 198–210. https://­doi​.­org​/­10​.­1016​/­j​.­yfrne​.­2013​.­07​.001; A. Fausto-­Sterling. 2020.
Sexing the Body: Gender Politics and the Construction of Sexuality. London: Hachette;
S. Richardson. 2013. Sex Itself: The Search for Male and Female in the H ­ uman Genome.
Chicago: The University of Chicago Press.
3. S. LeVay. 1991. “A Difference in Hypothalamic Structure between Heterosexual
and Homosexual Men.” Science 253 (5023): 1034–37. https://­doi​.­org​/­10​.­1126​/­science​
.1887219.
4. “Researcher Simon LeVay Talks About the Science of Sexuality.” Elmhurst
University, October 18, 2012. https://­www​.­elmhurst​.­edu​/­news​/­researcher​-­simon​
-­levay​-­talks​-­about​-­the​-­science​-­of​-­sexuality​/.
5. A. Borsa, M. Miyagi, K. Ichikawa, K. De Leon De Jesus, K. Jillson, M. ­Boulicault,
and S. S. Richardson. 2024. “The New Ge­ne­tics of Sexuality.” GLQ 30 (1): 119–40; A.
Ganna, K. J. H. Verweij, M. G. Nivard, R. Maier, R. Wedow, A. S. Busch, A. Abdellaoui,
et al. 2019. “Large-­Scale GWAS Reveals Insights into the Ge­ne­tic Architecture of
Same-­Sex Sexual Be­hav­ior.” Science 365 (6456): eaat7693. https://­doi​.­org​/­10​.­1126​
/­science​.aat7693; Fausto-­Sterling, Sexing the Body; C. E. Roselli. 2018. “Neurobiol-
ogy of Gender Identity and Sexual Orientation.” Journal of Neuroendocrinology 30
(7): e12562. https://­doi​.­org​/­10​.­1111​/­jne​.12562.
6. Fausto-­Sterling, Sexing the Body; Roselli, “Neurobiology of Gender Identity
and Sexual Orientation”; A. Fuentes. 2022. Race, Monogamy and Other Lies They Told
You: Busting Myths about H ­ uman Nature. 2nd ed. Oakland: University of California
Press; C. Hooven. 2021. T: The Story of Testosterone, the Hormone That Dominates and
Divides Us. New York: Henry Holt and Com­pany; R. Jordan-­Young and K. Karkazis.
2019. Testosterone: An Unauthorized Biography. Cambridge, MA: Harvard University
Press; C. Fine. 2016. Testosterone Rex: Myths of Sex, Science and Society. New York:
W. W. Norton; van Anders, “Beyond Masculinity”; M. Gutmann. 2019. Are Men Ani-
mals? New York: Basic Books; and many, many more. . . .
 Not e s to Ch a p t er 7 185

7. Once the initial sensory stimulus engages the limbic system, neural stimuli
(brain actions) are transmitted to much of the body via the endocrine system (using
the same hormones, testosterone, vasopressin, and oxytocin, in all ­humans). This
leads to physical and psychological excitement that includes increased blood flow
and swelling of tissues (called vasocongestion) and a tensing of the muscles (called
myotonia) throughout the body, sporadic increases in blood pressure, lubrication in
the vagina and inner labia, erection of the clitoris and erection of the penis, glandular
secretions across parts of the body, and ultimately (or potentially) a variable suite of
physiological changes associated with orgasm. Fuentes, Race, Monogamy and Other
Lies; see also R. S. Calabrò, A. Cacciola, D. Bruschetta, et al. 2010. “Neuroanatomy
and Function of H ­ uman Sexual Be­hav­ior: A Neglected or Unknown Issue?” Brain
and Be­hav­ior 9: e01389. https://­doi​.­org​/­10​.­1002​/­brb3​.1389.
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10. Associated Press. 2020. “Tennessee Governor Signs Anti-­Gay Adoption Bill.”
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11. “Child Welfare Nondiscrimination Laws.” Movement Advancement Proj­ect,
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12. “Same-­Sex Parents Are 7 Times More Likely to Raise A ­ dopted and Foster
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 186 Not e s to Ch a p t er 7

14. ­There are also many other, very serious, prob­lems with animal testing. See
H. Ferdowsian, A. Fuentes, L. Johnson, B. King, and J. Pierce. 2022. “­Toward an
­Anti-­Maleficent Research Agenda.” Cambridge Quarterly of Healthcare Ethics 31 (1):
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We Owe Lab Animals?” New York Times, January 23. https://­www​.­nytimes​.­com​
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15. This is not true of all medical researchers, especially in the past ­decade or two.
­Today, t­ here are many out­spoken advocates of including female nonhuman animals
(and h­ umans) in research. Most major government funding agencies of Canada, the
United States, and the ­European U ­ nion now require such inclusion.
16. For an excellent overview, see E. Cleghorn. 2021. Unwell ­Women: Misdiagnosis
and Myth in a Man-­Made World. New York: Dutton.
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/­female​-­mice​-­hormones​.html.
18. Which was primarily the United States and E ­ urope ­until the last fifty years or so.
19. ­There is a range of solid scholarship on this blend of racism and sexism and
how it s­ haped con­temporary medical practice in the United States; see Cleghorn,
Unwell ­Women; C. Willoughby. 2022. Masters of Health: Racial Science and Slavery in
US Medical Schools. University of North Carolina Press. See also A. K. Beery and
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.­07​.002; Ghorayshi, “Guess Which Sex Behaves More Erratically”; D. R. Levy,
N. Hunter, S. Lin, E. M. Robinson, W. Gillis, E. B. Conlin, R. Anyoha, et al. 2023.
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20. L. Greaves and S. A. Ritz. 2022. “Sex, Gender and Health: Mapping the Land-
scape of Research and Policy.” International Journal of Environmental Research and
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21. This is primarily referring to 3G females, but we know that the term “­women”
is not a biological term and not all ­women are 3G females. This, however, still sup-
ports the statement as the range of biological variation in the term “­women” (and in
“men”) is much larger than is envisioned, or engaged, by much medical research.
22. I. Zucker and B. J. Prendergast. 2020. “Sex Differences in Pharmacokinetics
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doi​.­org​/­10​.­1186​/­s13293​-­020​-­0 0308​-5.
23. If this is the case, ­there are not ­really any significant differences between male
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 Not e s to Ch a p t er 7 187

24. D. J. Greenblatt, J. S. Harmatz, and T. Roth. 2019. “Zolpidem and Gender: Are
­ omen R
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32. K. Sobhani, D. K. Nieves Castro, Q. Fu, R. A. Gottlieb, J. E. Van Eyk, and
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/­s13293​-­018​-­0201​-y.
33. L. J. Shaw, R. Bugiardini, and C. N. Merz. 2009. “­Women and Ischemic Heart
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 188 Not e s to Ch a p t er 7

35. W. M. Schultz, H. M. Kelli, J. C. Lisko, T. Varghese, J. Shen, P. Sandesara,

A. A. Quyyumi, et al. 2018. “Socioeconomic Status and Cardiovascular Outcomes:
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/­CIRCULATIONAHA ​.­117​.029652; T. M. Powell-­Wiley, Y. Baumer, F. O. Baah,
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36. See Brigham and ­Woman’s Hospital information as a good example: “Heart
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https://­give​.­brighamandwomens​.­org​/­7​-­differences​-­between​-­men​-­and​-­women​/.
37. See F. Puoti, A. Ricci, A. Nanni-­Costa, W. Ricciardi, W. Malorni, and
E. ­Ortona. 2016. “Organ Transplantation and Gender Differences: A Paradigmatic
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38. A. Melk, B. Babitsch, B. Borchert-­Mörlins, F. Claas, A. I. Dipchand, S. Eifert,
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.0000000000002655.
39. National Academies of Sciences, Engineering, and Medicine. 2020. Birth Settings
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emies Press. https://­nap​.­nationalacademies​.­org​/­resource​/­25636​/­interactive​/.
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M. McLemore, et al. 2019. “The Giving Voice to M ­ others Study: Inequity and
­Mistreatment during Pregnancy and Childbirth in the United States.” Reproductive
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­Efforts to Address Them.” KFF, November 1. https://­www​.­kff​.­org​/­racial​-­equity​-­and​
-­health​-­policy​/­issue​-­brief​/­racial​-­disparities​-­in​-­maternal​-­and​-­infant​-­health​-­current​
-­status​-­and​-­efforts​-­to​-­address​-­them​/.
41. National Academies of Sciences, Engineering, and Medicine, Birth Settings in
Amer­i­ca.
42. A. M. Jukic, D. D. Baird, C. R. Weinberg, D. R. McConnaughey, and A. J. ­Wilcox.
2013. “Length of H ­ uman Pregnancy and Contributors to Its Natu­ral V ­ ariation.”
­Human Reproduction 28 (10): 2848–55. https://­doi​.­org​/­10​.­1093​/­humrep​/det297;
A. J. Butwick, M. L. Druzin, G. M. Shaw, and N. Guo. 2020. “Evaluation of US State-­
Level Variation in Hypertensive Disorders of Pregnancy.” JAMA Network Open 3
(10): e2018741. https://­doi​.­org​/­10​.­1001​/­jamanetworkopen​.­2020​.18741.
43. Such as training and r­ unning into the third trimester and across lactations:
A. S. Tenforde, K. E. Toth, E. Langen, M. Fredericson, and K. L. Sainani. 2015.
 Not e s to Ch a p t er 7 189

“­Running Habits of Competitive Runners during Pregnancy and Breastfeeding.”

Sports Health 7 (2): 172–76. https://­doi​.­org​/­10​.­1177​/1941738114549542.
44. The term “sex contextualism” was coined by the ­philosopher and historian of
science Sarah Richardson: see S. S. Richardson. 2022. “Sex Contextualism.” Philoso-
phy, Theory, and Practice in Biology 14: 2. https://­doi​.­org​/­10​.­3998​/­ptpbio​.2096.
45. J. S. Williams, M. R. Fattori, I. R. Honeyborne, and S. A. Ritz. 2023. “Consider-
ing Hormones as Sex-­and Gender-­R elated ­Factors in Biomedical Research:
­Challenging False Dichotomies and Embracing Complexity.” Hormones and Be­hav­ior
156: 105442. https://­doi​.­org​/­10​.­1016​/­j​.­yhbeh​.­2023​.105442; M. Pape, M. Miyagi,
S. A. Ritz, M. Boulicault, S. S. Richardson, and D. L. Maney. 2024. “Sex Contextual-
ism in Laboratory Research: Enhancing Rigor and Precision in the Study of Sex-­
Related Variables.” Cell 187 (6): 1316–26. https://d­ oi.​ ­org/​ 1­ 0.​ ­1016​/j­ .​ ­cell​.­2024​.­02​.008.
46. Pape et al., “Sex Contextualism in Laboratory Research.”
47. For a ­great discussion on this topic, see interview of Sarah Richardson by
Heather Shattuck-­Heidorn and Kelsey Ichikawa on the GenderSci Lab website: “No
Sex without Context: A Q&A with Sarah Richardson on ‘Sex Contextualism.’ .”
https://­w ww​.­genderscilab​.­org​/­blog​/­q​-­and​-­a​-­sarah​-­r ichardson​-­on​-­sex​-­contex​
­tualism​. See also L. Greaves and S. A. Ritz. 2022. “Sex, Gender and Health: Mapping
the Landscape of Research and Policy.” International Journal of Environmental Re-
search and Public Health 19: 2563. https://­doi​.­org​/­10​.­3390​/ijerph19052563.
48. C. Semenya. 2023. “­Running in a Body That’s My Own.” The New York Times,
October 21. https://­w ww​.­nytimes​.­com​/­2 023​/­10​/­21​/­opinion​/­r unning​-­body​
-­semenya​.html.
49. Despite ­these data and facts, in 2023 World Athletics banned the ­women with
higher natu­ral testosterone levels from all track events u­ nless they underwent six
months of hormone-­repressing treatment. E. Pells. 2023. “Track Bans Transgender
Athletes, Tightens Rules for Semenya.” AP News, March 23. https://­apnews​.­com​
/­article​/­transgender​-­track​-­semenya​-f3499b00b932948f96838adb3b010f11.
50. S. Bermon and P. Y. Garnier. 2017. “Serum Androgen Levels and Their Rela-
tion to ­Performance in Track and Field: Mass Spectrometry Results from 2127 Ob-
servations in Male and Female Elite Athletes.” British Journal of Sports Medicine 51
(17): 1309–14. https://­doi​.­org​/­10​.­1136​/­bjsports​-­2017​-097792.
51. “Correction: Serum Androgen Levels and Their Relation to ­Performance in
Track and Field: Mass Spectrometry Results from 2127 Observations in Male and
Female Elite Athletes.” British Journal of Sports Medicine 2021 55 (17): e7. https://­doi​
.­org​/­10​.­1136​/­bjsports​-­2017​-­0 97792corr1​. Erratum for: British Journal of Sports
Medicine 2017 51 (17): 1309–14.
52. D. J. Handelsman, A. L. Hirschberg, and S. Bemon. 2018. “Circulating Testos-
terone as the Hormonal Basis of Sex Differences in Athletic P ­ erformance.” Endocrine
 190 Not e s to Ch a p t er 7

Reviews 39 (5): 803–29. https://­doi​.­org​/­10​.­1210​/­er​.­2018​-­0 0020​. See also Jordan-­

Young and Karkazis, Testosterone.
53. But see S. Bekker and S. Mumford. Forthcoming. Open Play: The Case for
Feminist Sport. London: Reaktion Books, for a much more in depth and complicated
history of why “­women’s” sports came to be.
54. Bekker and Mumford, Open Play; J. L. Parsons, S. E. Coen, and S. Bekker. 2021.
“Anterior Cruciate Ligament Injury: T ­ owards a Gendered Environmental Approach.”
British Journal of Sports Medicine 55: 984–90; L. C. Hallam and F. T. Amorim. 2022. “Ex-
panding the Gap: An Updated Look into Sex Differences in R ­ unning ­Performance.”
Frontiers in Physiology 12: 804149. https://­doi​.­org​/­10​.­3389​/­fphys​.­2021​.804149.
55. Hallam and Amorim, “Expanding the Gap.”
56. Julian Savulescu. 2019. “Ten Ethical Flaws in the Caster Semenya Decision on
Intersex in Sport.” The Conversation, May 9. https://­theconversation​.­com​/­ten​
-­ethical​-fl aws-­in-­the-­caster-­semenya-­decision-­on-­intersex-­in-­sport-116448; Seme-
nya, “­Running in a Body That’s My Own”; See also Jordan-­Young and Karkazis,
Testosterone.
57. T. A. Roberts, J. Smalley, and D. Ahrendt. 2021. “Effect of Gender Affirming
Hormones on Athletic P ­ erformance in Transwomen and Transmen: Implications for
Sporting Organisations and Legislators.” British Journal of Sports Medicine 55: 577–83.
58. V. Seibert. 2014. “Michael Phelps: The Man Who Was Built to Be a Swimmer.”
The Telegraph, April 25. https://­www​.­telegraph​.­co​.­uk​/­sport​/­olympics​/­swimming​
/­10768083​/­Michael​-­Phelps​-­The​-­man​-­who​-­was​-­built​-­to​-­be​-­a​-­swimmer​.html.
59. E. Pells. 2023. “Track Bans Transgender Athletes, Tightens Rules for Seme-
nya.” Associated Press, March 23. https://­apnews​.­com​/­article​/­transgender​-­track​
-­semenya​-f3499b00b932948f96838adb3b010f11.
60. See for example D. Lopiano. 2022. “A Fair and Inclusive Solution for Trans-
gender ­Women in Sports.” Forbes, August 4. https://­w ww​.­f orbes​ .­com​ /­s ites​
/­donnalopiano​/­2022​/­08​/­0 4​/­a-​ ­fair​-­and​-­inclusive​-­solution​-­for​-­transgender​-­women​
-­in​-­sports​/­​?­sh​=7bdd75d012ef; and S. Erikainen, B. Vincent, and A. Hopkins. 2022.
“Specific Detriment: Barriers and Opportunities for Non-­Binary Inclusive Sports in
Scotland.” Journal of Sport and Social Issues 46 (1): 75–102. https://­doi​.­org​/­10​.­1177​
/0193723520962937; and B. Hamilton, A. Brown, S. Montagner-­Moraes, C. Comeras-­
Chueca, P. G. Bush, F. M. Guppy, and Y. P. Pitsiladis. 2024. “Strength, Power and
Aerobic Capacity of Transgender Athletes: A Cross-­Sectional Study.” British Jour-
nal of Sports Medicine 58 (11): 586–97. https://­doi​.­org​/­10​.­1136​/­bjsports​-­2 023​
-108029.
61. “The Global Players’ and Athletes’ Association for Professional Sport—­UNI
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.­com​ /­topics​ /­f eatures​ /­i tem​ /­t he​ -­g lobal​ -­p layers​ -­and​ -­athletes​ -­a ssociation​ -­f or​
-­professional​-­sport​-­uni​-­world​-­athletes​-explained#.
 Not e s to Ch a p t er 7 191

62. J. L. Herman, A. R. Flores, and K. K. O’Neil. 2022. “How Many Adult and
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63. Jordan-­Young and Karkazis, Testosterone; but see Hooven, T: The Story of Tes-
tosterone, for opposite claims.
64. K. Nakajima and C. H. Jin. 2022. “Bills Targeting Trans Youth Are Growing
More Common—­and Radically Reshaping Lives.” NPR, November 28. https://­www​
.­npr​.­org​/­2022​/­11​/­28​/­1138396067​/­transgender​-­youth​-­bills​-­trans​-sports.
65. The Trans Legislation Tracker. https://­translegislation​.­com​/.
66. G. R. Murchison, M. Agénor, S. L. Reisner, R. J. Watson. 2019. “School Rest­
room and Locker Room Restrictions and Sexual Assault Risk among Transgender
Youth.” Pediatrics 143 (6): e20182902. https://­doi​.­org​/­10​.­1542​/­peds​.­2018​-2902;
L. Z. DuBois, J. A. Puckett, D. Jolly, S. Powers, T. Walker, D. A. Hope, R. Mocarski,
et al. 2024. “Gender Minority Stress and Diurnal Cortisol Profiles among Transgen-
der and Gender Diverse ­People in the United States.” Hormones and Be­hav­ior 159:
105473. https://­doi​.­org​/­10​.­1016​/­j​.­yhbeh​.­2023​.105473.
 I n de x

ACTH (adrenocorticotropic sex biology of, 3–4, 22–37, 47; study

hormone), 93 of sex and variability in, 125. See also
activation differences, 101 specific animals
ADHD (attention deficit hyperactiv- anisogamy, 9–10, 11, 63, 73, 152n12;
ity disorder), 135 assumptions about, 116; classic
adipose tissues, 87–88; deposition of, position in biology in, 110;
88. See also fat emergence of, 7–8; original model
adrenal glands, 91–92, 93 of, 51; parental care and, 10, 19;
adverse drug events (ADEs), 186n23 simplistic view of, 19
age, correlation of sexual activity anti-­Mullerian hormones (AMHs),
with, 127 172n24, 174n43
aggression, 112–13; ­battle of the sexes antiplatelet therapy, 136
argument on, 119, 121; participation ants, 24
in antisocial, 120; testosterone and, anxiety, 135
96; variations in, 118–20 apes, 52–56; sexual activity in, 64–65;
agreeableness, 113 tool use of, 69
algae, blue-­green, 5 aquatic species, 13. See also fish
allocare, 60–61, 164n35. See also Aristotle, 9
childcare arousal, physiological system of, 127
altricial offspring, 30 art, 60, 71, 72
Ambien (zolpoidem), 132–33, 140 asexual reproduction, 5, 6, 7, 25, 127
amphibians, 13, 20 Australopithecines, 57–58, 59
anal sex, 115–16 autistic spectrum, 135
androgens, 89, 92; concentrations of,
33; hormone exposure and, 33 baboons, 51
andropause, 94 bacteria, 5; division of, into archaebac-
anemonefish, 27 teria and eubacteria, 151n2
animals: evolution of, 152n12; Bateman, Angus, 9–11, 63, 109, 152n12,
prob­lems with testing of, 186n14; 152n14

193
 194 i n de x

“­battle of the sexes” concept, 110, 121, body size and strength, 110–11
122–23 bonobos, 54, 55, 56, 64, 128
Beauvoir, Simone de, 42–43 brains, 3, 100–104; size of ­human, 41
bees, 24, 35, 155n5; reproductive organs
in, 2 California sheepshead fish, 27
behavioral diversity, 54 canine teeth, 50, 53, 70; dimorphism
bimodal definition of sex, 39–40 in, 57, 163n25
binary view of sex biology, 4, 84–85, cardiac disease, binary view of, 135–36
125–50; cardiac disease in, 135–36; cetaceans (­whales and dolphins), 18
inadequacy of, 3–4, 21, 39–40, childcare: animals and, 18–20, 32, 49;
149–50; making ­family as part of, ­humans and, 55, 66, 110; role of
129–31; in medical research and group in 59–63, 68, 71
treatment, 131–35; organ transplant chimpanzees, 54–55, 56, 64; sexual
as aspect of, 136–37; pervasiveness be­hav­ior in, 64; subspecies of,
of, 125; pregnancy in, 137–39; sex 165n51
contextualism in, 139–40; sexuality chromosomes, 4, 5, 158n1; multiple
and sexual orientation in, 126–29; sets of, 25, 155n8; sex, 104;
sports and, 141–47 twenty-­third, 44, 46, 85, 99, 104,
biocultural framing, 45, 46 106; X, 38, 88, 95, 104–5, 108; XX, 85,
biocultural view: of ­humans, 40–42, 95, 105, 158n1; XY, 46, 99, 105, 106,
47, 109; of organs, 90–91 137; Y, 38, 88, 104–5
biological binary, 8 clitoris, 16, 17, 33, 45, 86, 114
biological differences in communica- cloaca, 16, 30
tion, 109 clothing, 41, 43
biological sex, 158n1; gonads and, 86 cognition: ­battle of the sexes
biology, 11, 40; animal, 3–4, 22–37, 47; argument on, 121; differences
gender and, 45; ­human, 84–86; between males and female in,
reproductive, 1–2, 3–4, 11, 12. See also 111–12; as nonbinary, 113
sex biology “Cohens d” ­measure, 112–13
biomedical research, 140 communication: biological differences
biopsies, 85 in, 109; dynamics in, 112
bipedalism, 88 comparative approach, defined, 48
birds, 16, 23; heterosexuality in, 128; connectome, 100
parental care by, 18, 19–20, 30–31, coprodeum, 16
161n2. See also par­tic­u­lar species cortisol, 98
birth ­mothers, reproductive ­process crocodilians: infant care by, 18, 30; sex
in, 130 biology in, 28–29
bisexuality, 127 cultural meaning, assignation of, 67
Bitch: On the Female of the Species cyanobacteria, 5
(Cooke), 36 Cytochrome P450 (CYP), 134
 i n de x 195

Darwin, Charles, 9, 14, 109, 152n12; on extroversion, 112

sexual se­lection, 50–51 EXX2 gene, 172n24
DAX1 (nuclear receptor protein), 106
dementia, 135 fallopian tubes, 97
depression, 135 ­family, making of, as binary view
developmental dynamics, 108 prob­lem, 129–31
DHEA (dehydroepiandrosterone), 93 fat, 86, 87–88; brown, 87; variation in
DHT (dihydrotosterone), 53 deposition, 88. See also adipose
dimorphism, 55, 155n1; sexual, 30–31, tissues
51, 53, 55, 59, 101, 108, 179n5 Fausto-­Sterling, Anne, 115
dinosaurs, 30 females, 1, 12, 42–43, 50, 108;
discontinuity, evolutionary 56 biologists’ use of term, 12, 39; body
diversity in sex biology, 149–50 size of, 134; brains in, 120; childcare
DNA, 1, 8, 104–7, 110, 151n1; in and, 19; cognition in, 111–12;
eukaryotes, 5; information revealed communication between males
by, 77 and, 109; in dominance, 162n14;
dolphins, 18 height distribution in, 74–75;
drones, 24 physical skills of, 113; relationships
drugs: effect on 3G males and 3G with males, 109; sexual activity by,
females, 132–33. See also specific 127–28; Sociosexual Orientation
names Inventory (SOI) ­measure of, 117;
duck-­billed platypus, 14 variation between males and, 109,
ducts, Mullerian and Wolffian, 124. See also 3G females; ­women
85–86, 99 female/woman distinction, 42, 122–23,
dyslexia, 135 170n1
femininity, 33, 42, 43, 83, 103, 123
­eagles, 31 fetuses, 14, 17–18, 20, 28, 97; fetal
eating disorders, 135 development, 17, 153n21
eggs, 14, 16–17, 29. See also ova fish: anemonefish, 27; bluehead
elephants, 18, 23 wrasse, 1–2, 26–27; California
embryos, development of zygotes sheepshead as, 27; caretaking in,
into, 18, 20, 21 19–20; female-­mimics, 26–27;
endocrine system, 91–92, 185n7 gobies, 27–28; sex biology change
endometrium, 98 in, 27; teleost, 26
estradiol, 94 fixed sex differences, 22
estrogen, 92, 99 flanges, 53, 54
eubacteria, 151n2 follicle-­stimulating hormone (FSH),
eukaryotes, 5–6 53, 94, 97–98
evolution, 11, 56; animal, 152n12; of sex, fossil rec­ord, 48, 56, 59, 62, 68, 69–70,
4, 5–21 122
 196 i n de x

frogs, 13 hair, 88, 89–90, 173n34

fruit flies, 10, 152n12 Haldane, J. B. S., 36
hawks, 31
galactorrhea, 99 hearts, 90, 98, 136; transplantation of,
gall bladders, 90 137
gametes, 7–8, 15, 36, 39; fusion of, height distribution in men and
14, 16, 20, 24, 28, 31–32, 153n18; ­women, 74–75, 76
getting together, 11–15; large, 50; helping, 112
large versus small producers of, hematopoietic stem cell transplanta-
23–24, 50, 121; production of, 40; tion, 135, 137
types of, 8 Hemiptera, 24–25
gametogenesis, 85 hermaphrodites, 153n17
gay gene, quest for, 126 heteronormativity, 116
gender, 42–45, 67–73; be­hav­ior heterosexuality: aggression between
patterns in, 120; binary views of, 123, partners in, 119; in mammals and
126; biology and, 3, 45; as a cultural birds, 128; as natu­ral mode of sexual
experience, 44; defined, 68; be­hav­ior in ­humans, 127; sex binary
expectations and experiences in, and, 126
159n8; global dynamics in, 44; Hippocrates, 9
multidimensionality of, 123; in hirsutism, 89
neurobehavioral disorders, 135; hom­i­cide, patterns of, 120
norms in, 43–44; relationship with hominins 48; lack of canine dimor-
biology, 45; roles in, 43, 159n13. phism in, 163n25
See also ­under sports hominoids (apes), 48, 56–58; ­humans
“gender/sex” term, 45–46 as, 52–56
ge­ne­tic pro­cesses, 108, 151n1 Homo: ­humans as genus, 58–60; social
genital-­genital contact, 63 ­organization in, 122; two-­adults-­
genitals: animal, 32–33; ­human, 16–17, plus-­offspring as core social
18, 84–86, 114 structure for, 122
genital tubercle, 85 Homo sapiens, 123
gestation, 14, 23 homo­sexuality, 116, 126, 127
gibbons, 53 hormones, 91–97; gonadal production
Gila monsters, 29 of, 92; luteinizing, 53, 54, 94, 97–98.
gonadal ridge, 85 See also specific hormones
gonadarche, 93 HOXA13 gene, 172n24
gonads, 15–16, 108; biological sex and, ­human(s), 38–47; biocultural view of,
86; formation of, 85 40–42, 108, 109, 118; biological
gorillas, 51, 52, 54, 64 variation of traits in, 107–8;
gynecomastia, 87 development of genitals in, 85;
 i n de x 197

evolutionary history of, 49, 73; as kangaroos, 12

genus Homo, 58–60; as hominoids, kidneys, 90, 136
52–56; level of sex in, 118; makeup of
families among, 130–31; as primates, labia, 33, 45, 85, 86
48; sexual be­hav­ior in, 3; shaping labioscrotal/urogenital fold, 17
of, 73; social pair bonding in, 67 lactation, 14, 18, 19, 23, 99
­human biology, genitals and, 84–86 large-­gamete production, 50
­human bodies: material structures of, leadership, 112
40–41; variation in, 78–80, 125 lemurs, 52, 162n14
­human brain, size of, 41 LeVay, Simon, 126
­human evolution, 49f, 111; hetero- LGBTQ families, prejudice against,
sexual, 66; pair bonds in, 66 129–30
­human sex biology, 48–49, 75–76; limbic system, 185n7
complications of, 39–40 LIM1 gene, 172n24
­human transcriptome, 106–7 Lion King, The, 32
hunting, 70 liver, 90, 98; transplantation of, 137
hyenas, 12, 32–33, 36, 96 lizards, 28–29
Hymenoptera (bees, ants, and lungs, 90; transplantation of, 137
wasps), 24 luteinizing hormone (LH), 53, 54, 94,
hypercooperation, 59 97–98
hypertensive disorders, 138
hypothalamic-­pituitary axis (HPA), 85 macaque monkeys, 51
hypothalamic-­pituitary-­gonadal male/man distinction, 42, 122–24,
(HPG) axis, 93 170n1
hypothalamic-­pituitary-­ovarian males, 1, 12, 50, 108; biologists’ use of
(HPO) axis, 93, 97–98 term, 12, 39; body size in dominat-
hypothalamus, 91 ing group, 50; brains in, 120;
cognition in, 111–12; communica-
INAH-3 nucleus, 101, 126 tion styles of, 109; conflict between
infants. See childcare females and, 111; flanged and
inhibin, 94, 174n43 unflanged, 54; height distribution
insects, scale, 24 in, 74–75; mammal, 155n1;
insomnia, battling, 133 one-­to-­one universal truths in, 149;
internal fusion systems, 16–18 oral sex and masturbation in, 115;
International Olympic Committee physical skills of, 113; promiscuity
and World Athletics, 141 and, 124; relationships with
intersex category, 12, 25, 153n17 females, 109; size of INAH-3 in gay
introversion, 112 versus straight, 126; Sociosexual
isogamy, 7 Orientation Inventory (SOI)
 198 i n de x

males (continued) meta-­analyses, 180n8

­measure of, 117; universal dominance minds, difference between men’s and
of, 109; variation between females ­women’s, 113
and, 109, 124. See also 3G males; men “mind” studies, 112–13
mammals, 17, 18, 31, 48–49; core mole rats, naked, 12, 35, 36, 96
premise of the ­battle of sexes and, monkeys: baboons, 51; macaques, 51;
110; gametic fusion in reproduction, tamarins, 12; titis, 96
129–30; heterosexuality in, 128; monogamy: in animals, 34–35, 36; in
internal gestation in, 28; male, 155n1; ­humans, 66, 122, 157n32, 166n66
marsupial, 153n21; masturbation in, motor be­hav­iors, 112
114–15; pair bonds in, 67; physiologi- Mullerian ducts, 85–86
cal and behavioral care of, 161n2; muscles: concentric versus eccentric
placental, 153n21; postbirth care in, contractions of, 171n11; ­factors in
32; reproductive biology in, 20; ­performance of, 83; relationship
reproductive physiology in, 23–24; between strength and architecture
sex biology for, 33; social, 164n35 of, 80–81; in shaping ­human bodies,
mammary glands, 18 80–84; strength of, 80–81; 3G
mandibular ramus, 80 assessments of, 82–83; in 3G
“man’s mind,” differences between females versus 3G males, 81;
“­woman’s mind” and, 113–14 variation in, 81–83
marsupials, 153n21, 161n1 myotonia, 185n7
masculinity, 42, 44, 95, 103, 123;
clothing and, 43 National Academies of Science,
masturbation, 113–15; self-­reported, 127 Engineering, and Medicine, 138
mating types (aka sexes), 6 National Health and Nutrition
medical research: binary view bias in, Examination Survey (NHANES),
131–35; sex contextualism in, 139–40 78–79
men: biocultural makeup of, 41; natu­ral se­lection, 9
common assertions about, 109, 124; Neanderthals, 70, 165n44; teeth in, 70
gender classification of, 42–44, 69, negotiation, 112
124; sexual activity of, 115–17; social neurobehavioral disorders, 135; gender
roles of, 38; 3G categories and, 47, in, 135
69, 136. See also males; testosterone; neurobiology, 100–104
3G males niche construction, 154n29
menarche, 93–94 nonlinear scaling relationship between
Men Are from Mars, ­Women Are from region and brain size, 177n71
Venus (Gray), 109–10, 179n1
menopause, 94; physiological changes obstructive coronary artery disease
in, 128; sex biology of, 116 (CAD), 135–36
menstruation, 97–98 one sex model, 9
 i n de x 199

orangutans, 53–54, 64; sex biology of, biology and be­hav­ior in, 49–50;
54; social lives of, 54; types of, 2 sexual be­hav­ior of, 128; sexual
organs: sex biology and, 90–91; se­lection in, 51; shapes and sizes of,
transplants of, as aspect of binary 52; social be­hav­ior and relationships
view prob­lem, 136–37 among, 33–34, 50; social sex in, 63
ova, 8, 13, 15 proctodeum, 16
ovaries, 15–16, 17, 92, 94 progesterone, 92, 94, 97
ovotestis, 16, 153n17 prokaryotes, 5, 6
ovulation, 97–98 prolactin, 99
oxytocin, 185n7 protobirds, 30
pseudogynecomastia, 87
pair bonds, 52, 53, 65–67, 122, 130, pseudopenis, 33
167n70 psychiatric conditions, 135
pancreas, 92 psychological states and be­hav­iors, 112
Pan genus, 54–55, 64 puberty, 87, 93, 94–95; mini puberty,
parathyroid glands, 91 93, 94
Parker, G. A., 10, 152n14
parthenogenesis, 12, 25 race, 91, 186n19
PAX2 gene, 172n24 religious beliefs, 44, 129
peafowl, 110 reproduction: asexual, 2, 5, 6, 7, 28, 127;
pelvic girdle, 59, 79 organs in, 1–2; patterns and trends
penis. See genitals associated with, 23; sexual, 6, 7, 8;
periodic ­table of ele­ments, 22–23 sexually dimorphic patterns
Phelps, Michael, 145 associated with, 32; types of organs
phenotypes, development of bodily, 91 in, 2
physiology, reproductive, 23, 156n26 reproductive biology, 11, 20; differ-
pituitary gland, 91, 92 ences in, 2, 4
placenta, 17 reproductive physiology, 23, 156n26
placental mammals, 153n21 reproductive skew, 63
polycystic ovary syndrome (PCOS), 95 reptiles: infant care by, 29–30; sex
polyploidy, 25 biology in, 28–30
power-­lifting competitions, 82
pregnancy, 98–100, 108; binary view salamanders, 13
of, 137–39 scrotum, 33, 85
primates, 48–52; f­ ather as caretaker in, sea ­horses, 13
49; h­ umans as, 48; masturbation in, Semenya, Caster, 141, 143
114–15; monogamy in, 34, 35, 36; sensory stimulus, 185n7
­mother as caretaker in, 49; sex: as bimodal, 39–40; evolution of,
pair-­living, 52; patterns of care in, 4, 5–21; purposes of, 6–7, 11, 63–65;
52; reproductive sex in, 64; sex self-­reported, 161n23
 200 i n de x

“sex at birth” term, 46, 47, 77 prob­lem, 126–29; gender/

sex binary, 36–37, 97; as tied to sex-­related variation in self-­
heterosexual reproduction, 126 reported perspectives, 118;
sex biology, 1–4, 5–21, 54, 65, 76–77, 127; responding to questions about,
basic animal, 22–37, 47; binary views 182n26; self-­reporting on, 113; as
of, 4, 39–40, 84, 126; climate-­ somatic fact and cultural effect,
influenced, 28–29; copulation, 13, 14; 115
as culturally contextualized pattern, sexual orientation, 116, 129
139; diversity in, 149–50; ­human sexual pair bond, 67
experience of, 39–40, 42; impor- sexual reproduction, 7, 8, 63–64;
tance of, 45; mammalian, 36; evolution of, 6
medical binary view of, 131; of sexual se­lection, 9, 50–51
menopause, 116; misunderstandings sexual vio­lence: initiation of, 127;
of, 22; niche construction in, 154n29; patterns of, 120
organs and, 90–91; patterns in, 23, sex uniformity, 22
110, 124; triggers for change in fish, siamangs, 53
27; typical, 39; variations in, 4, 18, skele­tons, patterns of variation in, 79
26–27, 29, 77, 110, 124, 130. See also skulls, 80
biology social pair bond, 67
sex chromosomes, 104 social sex, 64
sex confirmation testing, 141 sociosexuality, 118
sex contextualism, 189n44; in binary Sociosexuality index (SOI), 127
view prob­lem, 139–40 songs, 53
sex differences, 76–77 SOX9, 106
sex diversity and variability, 125 spatial-­ability skills, 112
sexism, 136, 186n19 spawning, 13
sex organs. See genitals sperm, 8, 15, 131
sex reproduction, 39 sports: binary view prob­lem and,
sexual activity: age and, 127; difficulty 141–47; gender-­uniform, 145;
in collecting data on, 115; as social impact of sex categories in, 83, 140;
tool, 64; ­women’s participation in, ­running, 143–44; testosterone and,
127–28 141, 142
sexual arousal, 117–18 SRY gene, 105
sexual assault, 182n32 strength training, 80, 82–83
sexual be­hav­ior of primates, 128
sexual dimorphism, 30–31, 51, 53, 55, 59, teeth: canine, 50, 53, 70; Neanderthal,
101, 108, 179n5 70
sexual interactions, 55 testes, 15–16, 92, 94, 114; undescended,
sexuality, 112, 115; ­battle of the sexes 141
argument on, 121; as binary view testes determining ­factor (TDF), 105
 i n de x 201

testosterone, 53, 54, 91, 92, 94–97, 119, reproductive strategy of “more
185n7; range of functions in, sex, less investment,” 113; sexual
146–47; sexual arousal and, 117–18; nature of, 109–10; testosterone
sports and, 141, 142, 144; variation in, 111; types of, 53. See also men;
in circulating levels of, 134, 141–42, males
146 thyroid glands, 91
3G categories, 46–47, 161n23 tissues, adipose, 87–88
3G females, 46–47, 50, 56–57, 186n22; transgender individuals: early
­battle of the sexes argument and, ­acceptance of, 43–44; growth in
122; biology associated with, 50; legislation against, 147; rest­room
bodies of, 62; brains of, 100; cardiac prohibitions and, 147–49; sporting
disease in, 135–36; as correct events and, 144–45
nucleus of ­family, 130; defined, 46; transplantation: heart, 137; hemato-
differences from 3G males, 62, 65; poietic stem cell, 135; liver, 137;
effect of drugs on, 132–33; height of, lung, 137
74–75, 112; ­human hair in, 89, 90; Trivers, R. L., 10, 152n14
infant care by, 110; in laboratory turtles, 28
research, 131–32; muscle strength in, two sex model, 9
80–84; nutritional and caloric
needs of, 68; organs in, 90; organ UK Biobank dataset, 102–3
transplants in, 136–37; pelvic girdle undescended testes, 141
in, 79; physical description of, 68; unflanged males (orangutan), 54
pregnancy and related physiological urodeum, 16
changes in, 137–39; sexual nature of,
109–10. See also females; ­women vaginas, 16, 45
3G males, 46–47, 56–57; ­battle of the vasocongestion, 185n7
sexes argument and, 122; biology vasopressin, 185n7
associated with, 50; bodies of, 62; vellus hair, 89
brains of, 100; cardiac disease in, vertebrates: egg laying in, 16–17;
135–36; as correct nucleus of ­family, gonads in, 15–16
130; defined, 46; differences from vocalizations, 53
3G females, 62, 65; DNA in, 110; voluntary activation, 81, 83
effect of drugs on, 132–33; height of,
74–75, 112; hyperaggression in, 120; wasps, 24
initiation of hom­i­cide and sexual WEIRD (Western, Educated,
assault by, 119–20; in laboratory Industrialized, Rich, Demo­cratic)
research, 131–32; muscle strength in, nations, 84
80–84; organs in, 90–91; organ Wilson, E. O., 10, 109, 152n14
transplants in, 136–37; pelvic girdle Wnt4, 106
in, 79; physical description of, 68; WNT gene, 172n24
 202 i n de x

­women: biocultural makeup of, 41; X and Y chromosomes. See ­under

common assertions about, 109, 124; chromosomes
gender classification of, 42–44, 69,
124, 143; sexual activity of, 115–17; zolpidem (Ambien), 132–33, 140
social roles of, 38; 3G categories zygotes, 12, 13, 14, 17, 24, 153n18;
and, 47, 136. See also females development into embryos of,
worms, sex biology of, 2, 25, 26 20, 21
 A NO T E ON T H E T Y PE

This book has been composed in Arno, an Old-style serif typeface in the
classic Venetian tradition, designed by Robert Slimbach at Adobe.