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William H. Calvin
Portraits of the
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Books by William H. Calvin
A Brief History of the Mind
A Brain for All Seasons
Lingua ex Machina*
The Cerebral Code
How Brains Think
Conversations with Neil’s Brain**
How the Shaman Stole the Moon
The Ascent of Mind
The Cerebral Symphony
The River That Flows Uphill
The Throwing Madonna
Inside the Brain**
*with Derek Bickerton
**with George A. Ojemann
Portraits of the Apes
WILLIAM H. CALVIN
Copyright © 2005 by William H. Calvin
All rights reserved
Version 0.9 (preview release) published in November 2005 by
WilliamCalvin.com. Printed by booksurge.com, a subsidiary of amazon.com.
William H. Calvin (2005). Almost us: portraits of the apes.
WilliamCalvin.com/booksurge.com. 200 pp. Supplementary material at
Supplementary information for this book and relevant web links,
including photographs for computer wallpaper and posters, may be found
at apes.WilliamCalvin.com. The low resolution color images on the
website may also be downloaded and utilized for education and nonprofit
purposes without requesting permission.
Cataloguing-in-publication data is on file with the Library of Congress.
WILLIAM H. CALVIN, Ph.D., is a professor at the University of Washington
in Seattle, affiliate faculty at the Living Links Center of Emory
University, and a science advisor to the Great Ape Trust of Iowa. His
research interests include how brains work, how humans evolved, how
language ability came about, and how evolution speeds up when the
earth’s climate abruptly flips. He is the author of a dozen books for
general readers and won the Phi Beta Kappa book award for science.
1. Play and the play face 9
2. A look full of meaning 25
3. The executive stare 39
4. The junior executive practices 45
5. Halfway into sharing things 55
6. The great apes 61
The omnivorous Chimpanzee 63
Bonobo, the “left-bank chimp” 81
The orange oriental Orangutan 111
The vegetarian Gorilla 129
7. The smaller apes 149
The acrobatic Siamang 151
Gibbon, the smallest ape 163
8. Not apes at all 169
9. The ape hidden in plain sight 177
Reading up on the apes 187
The view from the other side.
The bonobo, Makasi, and the toddler are about the same age.
Each time I visit the apes at my local zoo, I encounter some familiar
faces on my side of the glass windows. The apes have informal fan clubs,
people who show up regularly and stay for hours. Some bring toys and
picture books, which they prop up to show the apes. With great patience,
they try to direct the ape’s attention, just as they would do with a
young child. I just bring my camera.
While at the San Diego Zoo, a researcher told me that most of their five
ape species seemed to have its own set of followers. But, she added,
there weren’t any regulars for the monkey species at the zoo – except
for the undergraduates that she sends to study primate behavior.
Monkeys just don’t “grab” the visitors in quite the same way as apes.
It’s easy to see why. The apes look and act far more like humans than
other animals. Apes are personable, even charming. I’m not the first to
suggest that apes don’t fit either of the lay human-or-animal
categories. They are in a category all to themselves, almost us.
Yet apes are not well known to most people – most zoo visitors will be
heard referring to them as “monkeys.” Apes are super monkeys (and humans
are super apes). Portraits are how we “capture” human personalities,
however incompletely and imperfectly, and here I use portraits to show
you some glimpses into the expressions of our closest cousins.
W. H. C.
PLAYING & THE PLAY FACE
You are familiar with the play face from dogs offering to romp. The eyes
are wide open and the mouth is hanging loose, with the lips relaxed or
rounded inward to partly cover the teeth. The expression signals the
other that “This isn’t really aggression, it’s only pretending.” In some
apes, the play face is accompanied by a rhythmic vocalization resembling
(but not identical to) human laughter.
More orangutan play at England’s Twycross Zoo. Tiga, a 4-year-old male,
gets a play bite from Miti, 13-month-old female.
Here we see a grinning bonobo looking down at another’s play face. You
can also see why bonobos are said to have four hands. They were formerly
called “pygmy chimps“ but, since they are as tall as chimps, bonobo is
now their common name.
An emphatic bonobo grin from Bokela, 2. She is playing with her male pal
Bokela even grins while running.
Miti, 13 months old, explores with a stick at England’s Twycross Zoo.
Orangutans use their mouth like a third hand.
This is the bonobo Makasi, 16 months old.
It’s not often that two ape species are housed together, but the
siamangs and orangutans get along quite well together at the San Diego
Zoo. Here the adult male orangutan Satu, 10, has a play face while
play-biting siamang Hitam Lucu.. That’s her father Unkie in the
The next few pictures feature Hitam Lucu when she was 15 months old.
Here Hitam Lucu is “tagging” an orangutan.
A brief pause.
Bonobo play at the San Diego Zoo:Makasi (16 months old) and Kesi (12
There is an important lesson back there in Kesi’s expression sequence.
Kesi may have been unenthusiastic, but she certainly romped with Makasi
A micromoment from Makasi.
Fleeting “micro expressions” in humans can be seen on slow-motion video
replays. They are usually too brief to notice in real time, but a few
observers are good at it. Using videotaped interviews of people telling
lies on occasion, Paul Ekman discovered one experienced detective who
was especially good at detecting the lie (it proved a difficult task:
most experienced interrogators did little better than chance). It turned
out that this detective could make use of those brief micro expressions.
A LOOK FULL OF MEANING
Towan, 36, male orangutan in Seattle.
Belawan, 24, female orangutan in Seattle.
Female chimpanzee at Twycross.
Goya, 11, female chimpanzee at Burger’s Zooin Arnhem, the Netherlands.
Josephine, 45, female orangutan in San Diego.
Jamani, 5, female gorilla at San Diego.
Goya, an 11 year old Arnhem chimpanzee, is grooming while complaining.
Below, still disgruntled, with her mother.
This may remind you of an in-your-face drill sergeant. “When I say
‘jump,’ soldier, I want to hear you say ‘How high, sir?’”
Actually, this is social grooming. Kebara, 9, is using her lips to
“comb” the facial hair of Jamani, 5. The apes seem to derive much
pleasure from combing another’s hair – and inspecting it for insects
(which they eat).
Panbanisha, a language-reared female bonobo then seven years old, was
once fascinated with me. She kept asking for permission to touch the
visitor, day after day. Finally, she got the okay and promptly climbed
into my lap, flipped on her back, and diligently inspected my beard. She
was soon disappointed, climbed down, and lost all interest in me.
This is Lana, a 25-year-old bonobo in San Diego, holding her daughter
Kesi, who was then only two months old.
A bonobo family scene. Yenge, Kesi’s father, is starring into the
distance. Lana is grooming his hand and inspecting a find. Kesi is
hidden momentarily in Lana’s lap.
Female gorilla at San Diego.
Female chimpanzee at Twycross.
Mchumba, 5, is being “Big Sister” tothe “orphan” Makasi, a 18-month-old
THE EXECUTIVE STARE
Pete, 37, reddish-brown male gorilla in Seattle.
Alberta, 29, female gorilla in San Diego.
Violet, 28, female orangutan in San Diego.
Female gorilla in San Diego.
The sound of one foot clapping.
Gorilla in Seattle.
THE JUNIOR EXECUTIVE practices her trademark stare
Jamani, 5, female gorilla at San Diego Zoo‘s Wild Animal Park is
featured in this section.
But if those are the Junior Executive’s “business portraits,” what
follows might be called the “after hours portraits.”
HALFWAY INTO SHARING THINGS
An ape sometimes “shares food” by allowing others to take a few leaves,
while firmly holding on to the branch itself.
Possession seems to confer “ownership“ which, at least among adults, is
generally respected. That’s probably why you see so many apes carrying
branches around, draped over their shoulders. Leave it somewhere, and
someone else will take possession and own it.
It’s a long way from going around and offering food to others, one at a
time, but “tolerated scrounging“ is a start.
The 570-pound gorilla. Winston’s four-year-old grandson, Ajari, hopes
that he will share some bamboo leaves. He didn’t.
Dejection. But, just in case, staying in plain sight.
Except from mother to infant, food sharing is not common among primates.
We see it in a few species when males have meat and share with their
consort of the day. Within same-sex pairs, it is found only in some
highly tolerant capuchin monkeys and great apes: chimps, bonobos, and
–the primatologist CAREL VAN SCHAIK
THE GREAT APES
Orangutan (Asia) Gorilla (Africa)
Chimpanzee (Africa) Bonobo (Africa)
The ancestral great ape probably attained its large body size (and long
life span, crucial to maintaining culture through learning) in Asia
before rediscovering Africa and diversifying.
[After a fight between chimpanzees, each will sit around waiting for
their] adversary to make the first move [to reconcile]. The uneasiness
between them is obvious from the way they look in all directions—the
sky, the grass, their own body—while scrupulously avoiding eye contact.
Such a deadlock may last for half an hour or more, but can be broken by
a third party.
A female will approach one of the males, and after having groomed him
for a while, will slowly walk toward the other. If the first male
follows, he does so right behind her without ever looking at the other
male. Sometimes the female looks around to check and may return to tug
the arm of a reluctant male in order to make him follow. When the female
sits down close to the second male, both males will groom her, one on
each side, until she simply walks off and leaves them to groom each
other…. Such go-between behavior, called “mediation,” allows male rivals
to approach each other without taking any initiative, without making eye
contact, and perhaps without losing face.
Mediation promotes peace in the community by bringing the disputants
together. Interestingly, it’s only female chimps who mediate and only
the oldest and highest ranking among them.
– the primatologist FRANS DE WAAL
THE OMNIVOROUS CHIMPANZEE
Mother and big sister cope with Saphira, 3, at Arnhem.
Compared to the gorilla and its vegetarian diet, chimpanzees have a
number of behaviors for acquiring food. They are more versatile. Over
the course of one year, they eat over 300 food types. They will crack
open nuts. They probe for termites with a delicacy that far exceeds
dipping for syrup. Moving to block escape routes, a half-dozen chimps
will surround and catch a small monkey or bush pig.
Fresh meat may be less than three percent of their diet, but it is
prized. It has to be fresh, however. They are either cautious about, or
uninterested in, a dead monkey left on a trail for them to find. It’s as
if they didn’t know what to do with it, without the excitement of the
chase to prime them for ripping the carcass to pieces. They may kill
adult monkeys but they often leave much of the carcass uneaten, clearly
preferring young monkeys. Beggars gather around the possessor and some
meat is shared out.
Chimps are tool users in the wild but many practices are seen only in a
few places, suggesting that they are passed on by observation and
learning. I’ll scatter some tool examples among the pictures and also
include examples of communication. In general, they are things only seen
in the “three chimpanzees” (chimp, bonobo, and us), not in other wild
Chimps can hammer open tough nuts, seeking out flat stones to use as an
Some will even stabilize the anvil by wedging another stone underneath.
Chimps can mop up insects usinga leaf and then eat the insects.
A chimp will even make a thin stick into a toolby stripping off leaves
and protrusions, then punching throughthe ground into an underground
termite nest, eating the defendingtermites that cling to the stick when
it is withdrawn.
Chimpanzee Politics: subordinate males will form coalitionsto control
the power of a dominant chimp.
NOT SEEN: A chimp coalition playing against another coalition,as in the
ad hoc teams of humans for various games.
Three-year-old Saphira, playing to the crowd,apparently needs a hint as
to the right spot to groom.
Joery, 10, is the up-and-coming male at Arnhem, competition for Giambo,
15. His youthful light-colored face is beginning to darken into the
typical adult’s black face.
In the Arnhem chimpanzee colony, Luit once injured his hand
in a fight. He began supporting himself on a bent wrist, hobbling about
in an odd manner. Soon thereafter, Frans de Waal notes, all juveniles in
the colony began to walk in the same way. They kept this game up for
months, long after Luit’s injuries had healed.
NOT SEEN: The unconscious “echoing” of postures and speech,seen in a
pair of humans in conversation. If one crosses his legs or rubs his
face, the other may do so soon thereafter. A waitress who mimics a
customer’s tone of voice and gestures tends to get bigger tips.
Imitation often isn’t seen when you’d expect it. Michael Tomasello
trainedtwo young chimps to make a gesture in order to receive a favorite
treat. He then returned them to a play group of many young chimps.When
the keeper would come around with treats, both trained chimps would make
the gesture and get a treat. Despite intense interest,
none of the naïve chimps ever imitated the gesture to get a treat.
However, Andrew Whiten and coworkers trained a high-ranking female to
use a probe that dislodged treats. Soon, most of the other chimpanzees
in her social group got the idea. Prestige matters.
Honey is six months old. Her mother, Mwekundu, is a “chocolate” colored
variant seen at Twycross Zoo.
Adolescent male chimpanzee at Twycross.
This is the type of picture that appears on the cover of many picture
books about apes – except that the art editor selects the image from a
half-second before, when the eyes are still open and looking at the
Thus, a yawning ape is made to appear threatening in a misleading
Patting, the reassuring touch, and the arm around the shoulder.
Kissing and embracing.
Tickling oneself with an object.
Giambo, 15, alpha male at Arnhem.(He has been gray from an early age.)
[Gombe’s five-on-one gang warfare] involved chimps who
actually knew each other. Over the years, one community split into a
northern and southern faction, eventually becoming separate communities.
These chimpanzees had played and groomed together, reconciled after
squabbles, shared meat, and lived in harmony. But the factions began to
fight nonetheless. Shocked researchers watched as former friends now
drank each other’s blood. Not even the oldest community members were
left alone. An extremely frail-looking male, Goliath, was pummeled for
twenty minutes and dragged about. Any association with the enemy was
grounds for attack…. So us-versus-them among chimpanzees is a socially
constructed distinction in which even well-known individuals can become
enemies if they happen to hang out with the wrong crowd or live in the
–the primatologist FRANS DE WAAL
Only chimps and humans exhibit both the cooperative hunting
of game and the gang warfare that often kills a lone neighbor.
In both, the males act as a bonded “band of brothers.”
Alpha male chimpanzee at Twycross.
Tushi, 12, female chimpanzee at Arnhem.
Some chimps will use a stick to comb tangles out of their hair.
The royal wrist: adults may extend the back of their hand to be kissed
by an infant, which reassures it.
Raimee, 5, female chimpanzee at Arnhem.
When fruit is out of reach, some chimps (and orangutans)will use a long
stick to hook and pull down a branch.
Raimee, 5, female chimpanzee at Arnhem.
Zarno, 7, male chimpanzee at Arnhem.
Chimpanzees in the lab can match up, by family resemblance in portraits
taken elsewhere, a mother with a picture of her son.
Roos, 25. She was a youngster when Frans de Waal studied chimpanzees at
Burger’s Zoo in Arnhem, The Netherlands. Her early upbringing is
described in his book, Our Inner Ape.
Chimps can wad up a leaf and use itto mop up water from a deep hole.
Elderly chimpanzee at Twycross.
Chimps seldom die of the types of cancer and cardiovascular disease that
are the big killers of human adults, the pathologist Ajit Varki
discovered in talking with the ape veterinarians. Nor do they come down
with asthma. When infected with HIV, Hepatitis B or C, or the most
lethal form of malaria, chimps do not get particularly ill. That
suggests to me that some of our major diseases may be due to unfortunate
side effects of the gene alterations during human evolution, as rapid
evolution tends to leave behind a lot of unfixed “bugs.”
The detailed comparison of ape and human genes holds the promise of
clarifying how the apes successfully avoid our major diseases. That, we
hope, will suggest gene therapies to prevent and treat such diseases.
Chimpanzees are still to be found in western Africa, though as scattered
islands. There are several small, but famous, chimp populations in east
Africa, just east of the Rift Valley lakes in Uganda and Tanzania. Most
chimps, however, are in central Africa. Gorillas overlap with chimps in
both the west and the east. Bonobos are on the left bank of the Congo
River, chimps and gorillas on the right.
Not only does logging destroy their habitat, it creates roads that make
it easy for hunters to supply the bush meat trade. The bonobo
populations have been hard hit by soldiers scattered throughout the
Congo, who intimidate villagers into guiding them to the remaining
bonobos. Another big problem for the western gorilla and chimp
populations has been viral disease such as Ebola, which has greatly
reduced their numbers.
Apes in the wild are vanishing everywhere they still remain. No wild
populations of any size are stable. Unless this trend is reversed, there
will be no apes left in the wild.
The bonobo is overthrowing established notions about where we came from
and what our behavior potential is.... Even though the bonobo is not
our ancestor, but perhaps a rather specialized relative, its
female-centered, nonbelligerent society is putting question marks all
over the hypothesized evolutionary map of our species.
Who could have imagined a close relative of ours in which female
alliances intimidate males, sexual behavior is as rich as ours,
different groups do not fight but mingle, mothers take on a central
role, and the greatest intellectual achievement is not tool use but
sensitivity to others?
– the primatologist FRANS DE WAAL
THE “LEFT BANK CHIMP”
A little smooching, perhaps? Here we have, after all, father Yenge,
mother Lana, and ever-helpful daughter Kesi. This might be more
cautiously described as a bout of mutual grooming, lips used on the
other’s face. But bonobos do kiss, and perhaps it evolved out of
nibbling-style grooming. They even have reddish lips, unlike the other
You can tell a bonobo from a chimp because the bonobo has hair that
parts down the middle with occasional hair tufts. Bonobos are born with
side whiskers that chimps lack – but the chimps develop white beards.
Compared to bonobos, chimps have deep-set eyes that are closer together.
Bonobos have narrower shoulders, longer legs, and a slimmer appearance.
Their voices are more high pitched than those of chimps.
Bonobos spend more time upright – but also spend more time in the trees.
A bonobo can walk a tightrope with ease, fully upright. If a group of
chimps up in the trees is startled by an unfamiliar human, they will
jump to the ground and run away on all fours. Bonobos scramble swiftly
through the trees until a safe distance away, even though they too
customarily travel on the ground.
Chimps and bonobos were confused with one another for centuries until
bonobos were identified by Ernst Schwarz in 1929 from a Belgian
collection of ape skulls. The bonobo was originally called the “pygmy
chimp” and treated as a subspecies. But they have proved to be just as
tall as the “common chimps.” Chimps are more burly, looking as if they
work out on weight machines every day. Bonobos look as lean as if they
commuted to the office every day on a bicycle.
To avoid the ambiguity of the term “chimpanzee,“ scientists can always
use the Latin genus name Pan when referring to both species. The species
names are Pan paniscus for the bonobo and Pan troglodytes for the common
chimpanzee. Bonobos and “trogs” are sometimes heard in informal
conversation among scientists. (You will note that when I say “chimp,” I
am referring to the trogs.)
Of course, troglodytes is a misnomer as it means cave dweller. Chimps
never live in caves – but they were indeed a 19th-century candidate for
the ancestral “cave man.” Paniscus means diminutive – also wrong. The
Latin names, once given, seem impossible to change, no matter how
Makasi when 15 months old.
Yenge, Kesi (at 2 months), and Lana.
A bonobo may play a version of blindman’s buff, holding a hand over his
eyes or draping a blanket over his head – and then show off how well he
can balance on a rope or bounce off the walls without looking where he
The bonobo lives only in the equatorial forests of the central Congo,
south of the big loop of the Congo River. The phrase “The Left-bank
Chimps” refers to bonobos living on the left bank of the Congo River –
but also to frequent sexual intercourse. Indeed, the various
nonreproductive sexual behaviors occasionally seen in chimpanzees,
gorillas, and humans are seen in almost every bonobo.
It is said that chimpanzees resolve social conflicts with power while
bonobos resolve them with sex.
Yenge uses an arm across his face in an attempt to nap.
Mchumba, 5, is getting set to jump at a wall. She isn’t trying to escape
but to do a back flip off the wall. She did it over and over.
Watching her, you would have said she was “practicing“ – but maybe she
just liked the sensation and was repeating it for pleasure, the way our
children will spin themselves to become dizzy. I’ve seen Mchumba spin
several dozen times. (Maybe that’s how real practicing got started.)
This is Diatou, 28, the most experienced bonobo mother at Twycross.
Junior, 9, goes dipping for syrup, much like wild chimpanzees fish for
Bokela, 2, is entertaining herself at Twycross.
Makasi at 16 months.
Makasi at 16 months old. He was nursery reared while his mother,
Loretta, was ill. But then, because she wasn’t nursing, Loretta got
pregnant again, complicating Makasi’s re-introduction to the group. So
Makasi is getting his milk in the nursery from the keepers and spending
his days with a different bonobo group. Various bonobos are taking care
of the “orphan.” Mchumba, 4, has become “big sister,” carrying him
around and introducing him to somersaults.
This is in contrast with the younger Kesi, who gets a lot of reassuring
body contact from Lana.
Kesi at 12 months.
Lana, 25, calling.
Makasi and Kesi have found a cave in the roots of an overturned tree.
A male at the San Diego Zoo, shown here with the dipping stick for
syrup. Lana would have nothing to do with him.
Here is his replacement, Yenge, 22, who proved more acceptable to Lana.
(He is now the father of Kesi.) Zoos trade animals to minimize
inbreeding, but also to resolve such conflicts.
Banya, 15, with her daughter Bokela, 2, at Twycross.
Bokela at half-grin.
Kichele, 16, female bonobo at Twycross.
Kanzi, 25 and language-reared from an early age, now at the Great Ape
Trust of Iowa. It was a hot summer day and he just cooled off in a
If you tell him, “Kanzi, go to the office and bring back the red ball,”
a sentence he has never heard before, he will ignore all the nearby
balls and go into the office (provisioned temporarily with many balls,
though usually having none), find a red one, and carry it back. He makes
about as many errors on such tests of understanding long sentences as a
child does when two and a half years old. However, the sentences Kanzi
initiates via a keyboard labeled with words are usually no more than two
A human parent and a year-old child will both pay attention to an object
and communicate about it, but that kind of shared attention is usually
lacking in the wild great apes.
The bonobo language lesson at the Great Ape Trust of Iowa. That’s Nyota,
7, Sue Savage-Rumbaugh, and Kanzi, 25. In a conversation with Bill
Fields, outside, Sue is pointing to me – and Kanzi joins in, unprompted,
though pointing to someone else.
Then too, I was never Kanzi’s favorite visitor. Back when he was 13,
Kanzi would screech at me, over and over, each time I appeared. Sue
suggested that I needed to adopt a more humble posture that minimized my
height. I tried but finally found some earplugs. It took a week before
he stopped complaining and merely ignored me.
Captive bonobos can [express] vocabularies of several hundred words… and
are believed to be able to comprehend several thousand spoken words.
Bonobos combine symbols without being taught to do so and use a simple
grammar that is partially of their own construction…. Bonobos engage in
symbolic dialogs that may run for 20 or 30 minutes and span several
topics, each topic in turn giving way to another, and may return to the
dialog at a later point in time. They have no difficulty in leaving one
topic, moving to another, and then picking up the former topic, without
any need to recreate the former conversations that led up to it.
– the primatologist SUE SAVAGE-RUMBAUGH
Kanzi screams with pleasure upon getting a hose shower on a hot summer
day at the Great Ape Trust of Iowa. He is sprawled atop the remains of
the language lesson.
Jasongo, 15, male bonobo at Twycross with long mutton-chop whiskers.
Lana, 25, provides another example of Bonobo Hairstyle. And it’s not
even expensive, if you have the genes to make it grow in the right
directions – and the right grooming partners to compulsively maintain
Perhaps we will eventually see a spiky-hair variant called Bonobo Style
where the hairdressers send a percentage to one of the ape protection
Yenge, 23, is the big male at San Diego Zoo. Here he is sticking out his
chest to be groomed by Ikela, a 14-year-old female.
Yenge, Makasi, Lana, and Kesi.
All orangutans, indeed all great apes, build nests: simple ones during
the day, more elaborate ones for the night… The most common recipe is as
follows. Look for a tree that grows nicely horizontal branches; break
two that come out next to each other and fold them back to cross over
each other to form a roughly triangular platform. Add additional
branches, some simply bent into place and some gathered from nearby, and
keep them in place by latching them behind some protuberance. Then add
soft, leafy twigs, as many as needed to make the bed soft and comfy.
[Sometimes when it is raining, they build another nest just above, to
serve as a roof.]
– the primatologist CAREL VAN SCHAIK
If you happen to forget a screwdriver in the gorilla cage, the animals
will hesitantly approach it, briefly sniff it, and subsequently ignore
Leave it in a chimp cage, and it will be used in vigorous display,
thrown about, and forgotten.
But if you leave it in the orangutan cage, one of the animals will
unobtrusively pick it up, hide it, and use it to let itself out when
you’ve left for the day.
– the primatologist BENJAMIN BECK
[Given a mirror,] most apes open their mouth to inspect the inside,
touching their teeth with their tongue or picking at them with their
fingers guided by the mirror. Sometimes, they go so far as to
“embellish” themselves. Presented with a mirror, Suma, an orangutan at a
German zoo, gathered salad and cabbage leaves from her cage, placed them
on top of each other, then put the whole pile on her head. Staring in
the mirror, Suma carefully rearranged her vegetable hat.
– the primatologist FRANS DE WAAL
THE ORANGE ORIENTAL
Towan, 36, is an orange male with fat-filled cheek flanges that stand
out, typical of the big males in Borneo but not in Sumatra. Belawan, 23,
is a dark russet female. They are in Seattle.
You’d never know it from the sociable orangutans at the zoo, but adults
in the wild are often loners and can go a long time without seeing
another of their species. In Borneo, it takes a lot of territory to
provide enough food for each orangutan, so they spread out. They eat
mostly fruit, especially figs.
Their cleverness and sociability in captivity, however, suggested that
this solitary life wasn’t what they were once like, back before the food
supply diminished. Sure enough, some researchers in Sumatra discovered
that the orangutans at Suaq have enough food to be able to live in
social groups and develop culture. The orangutans elsewhere are simply
With this paradox resolved, orangutans are becoming our model for the
most basic of the great apes. In italics, I will note some of the things
they do in the wild that are not seen in gorillas, the smaller apes, or
An orangutan will use a bunch of leaves as a glove when handling fruit
with sharp spines.
An orangutan will break off a branch and use itas a hook to pull distant
branches within reach.
Clyde, 27, is the patriarch of the San Diego orangutans.
This is Indah, 28, a female Sumatran orangutan in San Diego.
Josephine, 45, the matriarch.
Miti is 13 months old. Here she is playing on old fire hose at Twycross
Orangutans have very agile lips, and can clench tools like cigars
between their teeth. The bigger ones can turn tools front-to-end without
ever using their hands.
Scientists who were “hand training” an orangutan to chose between two
pictures found that, instead of reaching with a hand, their orangutan
would get lazy and just point with her mouth to the correct choice. It
took a bit more persuasion before she would consent to use a button push
that the computer could also understand.
Another view of Josephine.
Miti at 13 months old.
An orangutan will use a stick to scratch its back.
A little mutual mouth grooming between orangutans in San Diego is about
to be interrupted by siamang Hitam Lucu (whose name, in one of the
languages spoken in Borneo, means “Black Mischief”).
There’s syrup in that San Diego Zoo “termite mound.” (They love honey or
a mixture of mustard and barbeque sauce.)
Violet, above, is holding the stick with a “power grip,” what’s
typically used when grabbing a branch overhead.
Karen, the orangutan to the right, instead dips with a smaller stick,
holding it between thumb and forefinger. Such a “precision grip“ is what
we humans use with most tools – and with proper tea cups.
To get across a wide gap, a female orangutan bit through a thick vine
and then used it to swing across, Tarzan style.
In zoo settings, orangutans are much faster to learn tool use than
chimpanzees and are also more creative.
Orangutans can grab a sleeping slow loris and kill it with a bite to the
NOT SEEN: group hunting.
Satu and Violet.
NOT SEEN: Orangutan Politics for establishing alliances that prevent a
big male from dominating..
The orangutan is found in both northern and southern Borneo, as well as
in the tsunami-prone western end of Sumatra‘s Aceh province. But their
island isolation is only temporary. During ice ages, when sea level is
lower (by the height of a forty-story building), all of the large
islands are connected to the Malay peninsula. This exposure of the Sunda
Shelf happens at least every 100,000 years and likely more frequently,
potentially allowing the different orangutans to mix.
The gibbons and siamangs are found throughout southeast Asia. Siamang
territories can overlap with those of gibbons because the siamangs are
largely leaf eaters while gibbons largely eat fruit.
Logging and fires have been greatly reducing all of the populations of
the Asian apes.
In present-day Sumatra, [orangutans] thrive only where the local
inhabitants do not hunt them because of religious prohibitions. From
north to south along the West Coast, the orangutans begin to disappear
where the crescents and sickles of mosques begin to give way to the
crosses of churches.
– the primatologist CAREL VAN SCHAIK
Once… swarms of termites passed through the study area. I expected that
the gorillas would, chimpanzee style, improvise twigs to extract the
termites from the decayed tree stumps. However, they totally ignored the
termites and waded their way past the infested areas to feed on
– the gorilla researcher DIAN FOSSEY
THE VEGETARIAN GORILLA
Kebara, 9, has found something she likes in the grass at the San Diego
Zoo’s Wild Animal Park. Gorillas are vegetarians except for the
occasional termite or egg.
Gorillas at the San Diego Zoo.
In the wild, a gorilla wading in a pond has been observed
to use a stick to probe the bottom for a safe path. It’s an extension of
using the stick as a cane
when moving around a swamp.
Another gorilla was observed to place a plank to walk on.
But a walking stick is only marginally tool use (using an object to
manipulate another object), and certainly not the more challenging
toolmaking studied in wild chimpanzees. And it’s not common. It took 40
years of field studies before such gorilla tool use was finally observed
in 2005 at a swampy forest clearing north of the Congo river. In the
same year, a juvenile gorilla alone in a sanctuary (a confiscated pet)
became proficient at cracking nuts.
In contrast to tool use, toolmaking requires innovation and, most
importantly, more of a tendency for others to imitate the actions (else
the innovation will die with the discoverer).
Winston, silverback male, 38, and Alberta, 29, female at the San Diego
Wild Animal Park.
Sharing food with friends, and not just family,
has not been seen in wild gorillas.
A broccoli of one’s own.
Like monkeys, nearly all gorillas fail to recognize themselves in a
The other great apes, on the first time they are given a mirror,
instantly recognize themselves. Usually, they open their mouth and move
their tongue. Next, they use the mirror to examine their backside.
The built-in baseball hat.This is Vila, 47.
Female gorilla at San Diego.
This is Ajari, 4, Winston’s grandson.
The apes invented toothpicks long ago.
Alberta, 29.The beat begins. The little drummer girl, Akenji, age 4.
This whole sequence takes only about one second from start to finish.
That’s like most of the facial expressions: Blink, and you might miss
Vila, 47. In the wild, gorillas need to eat (and excrete) about 50
pounds a day (70 for big males) of leaves and such. To extract enough
calories from such low quality food, the gorillas have evolved a longer
Modern vegetarians can manage without the bulge only because of modern
food preparation techniques.
Winston, 37, carries some carrots and broccoli back to his favorite spot
at the San Diego Wild Animal Park.
No “Gorilla Politics”: Males do not form coalitions
to limit the power of the dominant silverback male.
Gorillas have evolved into two specializations that have, I suspect,
limited their abilities to further develop their intelligence. Evolution
is unfortunately full of dead ends, usually because a species
over-specializes, doing one thing so efficiently that it can no longer
back up into a greater versatility. Trapped, by relying on doing one
thing quite well.
A single silverback male can dominate a group, chasing out other adult
males to create a harem. There are gorilla groups with multiple adult
males, but it is nothing like the mixed groups seen in the chimps and
bonobos. There you see lots of social strategies, not just brute force,
deciding who gets to mate with whom and who gets what food. There
coalitions and cooperation start to become important, the beginnings of
Gorillas also seem to have over-specialized into the vegetarian niche,
what with those big guts. Such a monotonous diet has not given gorillas
the incentive that the chimps and bonobos have had, what with needing to
develop the skills to acquire many different foods. They developed more
versatile brains and more versatile guts.
Gorillas can live only where there are suitable leaves to eat, available
in great quantity. As the African climate got drier over the last few
million years, the mountain gorillas have been pushed higher up the
mountain. Mountain tops get rain when lower elevations do not. The
danger, of course, is a drought pushing the mountain gorillas onto the
very top of the mountain – and beyond.
All primates love fruit, of course. When it is available, even big
gorillas will climb a tree, though their weight limits them to the
Surprisingly, you can occasionally see gorillas and chimps up in the
same fruit tree, eating away on nearby branches. Dian Fossey didn’t
report chimps and gorillas together but her study site is at a higher
elevation that lacks fruit trees. Just north of there in Uganda and at a
lower elevation, Craig Stanford and his coworkers have recently seen
this gorilla-chimpanzee coexistence.
With some exceptions, their observed interactions are more peaceful than
I would have expected. But, then, maybe there was enough fruit for
THE SMALLER APES
The “lesser” apes are much smaller than the four great ape species. They
are also more distantly related to us.
The earliest apes evolved from the Old World Monkeys about 23 million
years ago in Africa. They lost the monkey‘s tail, doubled brain size,
and acquired a far more versatile shoulder joint – which we can use to
attach ear rings (a monkey might find it difficult to reach behind its
head and around to the opposite ear). The gibbon-siamang branch dates
back to 15 million years ago, after apes had spread into Asia.
Siamangs and gibbons last shared a common ancestor only about 2 million
years ago, about the same as chimps and bonobos branched off from one
another. Some would consider the siamangs to simply be the most
distinctive of the subspecies of gibbon but differences in size, diet,
and female coloration suggest a new species, not a subspecies.
Unkie swings through the air with the greatest of ease – much better
than that man on the flying trapeze. Unkie, after all, is a siamang with
25 years of experience.
This is Sutera, 13, the female siamang at Seattle‘s Woodland Park Zoo.
Sutera and Simon are seen here howling a duet in a Seattle treetop.
Those inflatable throat pouches are what opera singers need for better
Hitam’s father, Unkie.
Upon seeing this portrait, my barber exclaimed, “Cool! Who does his
There was once a high-maintenance hair style something like this. It
should have been called “Siamang Style.”
Eloise and Unkie at San Diego Zoo. In the wild, siamangs pair-bond for
life. They take their time in choosing a mate, and seldom take another
mate if the first one dies.
Eloise with Hitam Lucu when she was only five months old. From Eloise’s
habitual expression, I have always imagined that she came to the San
Diego Zoo from an exclusive French restaurant.
This is Hitam Lucu at 15 months of age, expressing the siamang‘s
Simon’s version of tip-of-the-tongue.The gibbons do this as well., but
no oneseems to know what it “means.”
Hitam’s big sister, Hidiah, 5.
Siamangs usually get around by swinging through the trees. If they need
to walk on the ground, they walk upright.
The great apes can walk upright, especially when carrying something, but
their hands tend to get roughened from frequent walking on all fours,
thus reducing their abilities to do the finer movements with their
Karen and a bipedal Hitam.
The smallest ape
Tina, 23, is a Gabriella’s Crested Gibbon at the San Diego Zoo‘s Wild
Gibbons are half the size of siamangs. Unlike the black siamang females,
the female gibbon is usually golden brown. Male gibbons are mostly
black, while male siamangs are definitely black except for some facial
This is a male Pileated Gibbon at Twycross Zoo.
This is Jane, a female Lar Gibbon at Twycross..
Another male Pileated Gibbon.
This is Smiler, a male Muller’s Gibbon at Twycross. This subspecies
comes from Borneo.
A male White Cheeked Gibbon grooms its mate.
At emotionally meaningful moments, apes can put themselves into
another’s shoes. Few animals have this capacity…. Monkeys fail to
provide reassurance even if their own offspring has been bitten. They do
protect them, but show none of the cuddling and stroking with which an
ape mother calms down an upset youngster.
– the primatologist FRANS DE WAAL, 2005
NOT APES AT ALL
MONKEYS AND PROSIMIANS
That tail is a dead giveaway that this is not an ape. And don’t be
fooled by some local names for monkeys such as “Gibraltar ape.” The term
“monkey“ was only invented about 1700 and came into general use only in
the 19th century.
That distinctive moustache tells you that this is a Patas monkey. The
live in the grasslands south of the Sahara.
While some monkeys have interesting default expressions in the manner of
a mask, their faces are not versatile in the manner of the smiles,
grins, and frowns of the great apes.
Male mandrill at the Denver Zoo.
Capuchins are quite different from apes. They are little brown monkeys.
about the size of a cat, with long tails that serve as marvelous
grasping organs…. They are about the smartest monkeys I know. Sometimes
called New World chimpanzees, their brain, relative to their body, is as
large as that of an ape. Capuchins use tools, have complex male
politics, have lethal encounters between groups, and most important,
share food. This makes them ideal for studies of reciprocity and
– the primatologist FRANS DE WAAL
A red-faced Japanese macaque diligently grooms the boss at the Milwaukee
No, not an ape.Not even a monkey.(The giveaway is that pointed muzzle.)
It’s one of the more primitive prosimians. Lemurs live only on
Madagascar. This is a Red-Ruffed Lemur at Twycross Zoo in England.
These Golden Crowned Lemurs were balled up, napping together, but woke
up to stare back.
[W]e human beings, unlike all other species on the planet, are knowers.
We are the only ones who have figured out what we are, and where we are,
in this great universe. And we’re even beginning to figure out how we
– the philosopher DANIEL C. DENNETT1
The study of man showed that, in spite of his descent, he is indeed
unique among all organisms. Human intelligence is unmatched by that of
any other creature. Humans are the only animals with true language,
including grammar and syntax. Only humanity, as Darwin emphasized, has
developed genuine ethical systems. In addition, through high
intelligence, language and long parental care, humans are the only
creatures to have created a rich culture. And by these means, humanity
has attained, for better or worse, an unprecedented dominance over the
– the evolutionary biologist ERNST MAYR2
THE HIDDEN APE
Homo sapiens sapiens
You might have guessed by now that humans are the most recent of the
great apes. We’re just hidden in plain sight.
Suppose that I shuffled the order of the preceding species chapters and
then asked you to rank the ape species for their similarity to humans:
GIBBON CHIMP ORANGUTAN SIAMANG BONOBO GORILLA
Don’t turn the page until you try.
Tools and mirrors might be one way to rank them, as proxies for
intelligence. Some people, knowing about the chimpanzees’ gang warfare
directed at neighbors, would put the chimps closest to humans. Others,
impressed by the varied sexual behaviors of the bonobo, would place them
next to humans instead. (Actually, chimps and bonobos are equally close
in terms of the DNA clock, having split into the two lineages about 1.5
million years ago.)
The appropriate metaphor is, of course, a tree, not a ladder or chapter
sequence. You can see that I did not present the great apes in a time
sequence, since I left the gorillas until last (perhaps because I
consider them an overspecialized oddity, as I explained earlier).
That “myr” means millions of years.
By 1871, Charles Darwin had concluded that our closest relatives were
likely among the great apes, especially the African ones. He may have
been around the world once, but his first-hand experience with apes was
largely limited to observing the occasional ape at the London Zoo. But
in the 1860s, the anatomists – such as his friend Thomas Huxley – who
compared bones of monkeys, apes, and humans had been looking carefully
into the similarities and differences. And their bone-based conclusions
have held up pretty well. Here is a rough tree, with only a few of the
dead ends sketched in, that summarizes our current understanding of how
we relate to our ancestors and closer cousins of the last 8 million
A century after Darwin, the geneticists finally had techniques to
estimate that the protein differences between chimps and humans were
only a few percent, closer than with any other group of animals. This
conclusion from the proteins has been strengthened by DNA snippets that
have been analyzed more recently.
Then the geneticists obtained the entire genome of a chimpanzee named
Clint, enabling much more detailed comparisons with the human genome.3
Chimps and bonobos are our closest surviving relatives, followed by
gorillas, orangutans, and the lesser apes, then the Old World monkeys.
Darwin‘s tentative conclusion was right on the money.
There used to be many more ape species but only four great ape species
remain today. The “hominid“ branch, the human ancestral lineage,
starting at about 6 million years ago, also has a lot of dead-end
branches before you get to us humans. There were three surviving species
only 30,000 years ago but Neanderthals went extinct by 28,000 years ago
and Homo erectus somewhat more recently (if the hobbit-sized hominid is
really erectus – I’d bet on a pygmy-sized Homo sapiens). Now we’re
The earlier stages of great ape evolution likely happened in Asia but by
8 million years ago, it was the climate changes in Africa that created
new opportunities by reducing shade. Hominids came to specialize in the
open woodlands where there is lots of grass but still quite a few trees
to nest in at night, just as in the ancestral forest.
Grazing animals come to visit the woodland’s grass and the lions follow
them. So our ancestors had new threats to cope with, but there were new
resources as well, such as meat to scavenge and underground storage
organs like edible roots and tubers that are more common in woodland.
Early on, back near the last ancestor we shared with the chimp and
bonobo (“Pan prior”), two anatomical changes were evident. The big ape
canine teeth, so handy for threats and fighting, were much reduced in
size. Upright posture developed early as well. But about four million
years passed until the next phase began, of enlarging and reorganizing
Then they started eating a lot more meat and advanced out into the
savanna where the trees are few and far between. The grazing herds are
enormous and the lions make a good living. That our ancestors could now
make a living out there, away from the safety of trees, marks the path
to bigger brains. It takes a lot more fat in the diet to build a bigger
brain and there was a lot of animal fat on the hoof. Brain size, which
had remained at about the size seen in chimps and bonobos for four
million years, finally began to increase.
This suggests improving hunting and scavenging skills (either way, our
ancestors had to compete with those lions and hyenas), improving food
preparation, and far more sharing and cooperation. By about 200,000
years ago, we were finally Homo sapiens. We had the big brain, as did
Neanderthals. But it wasn’t until about 50,000 years ago that we also
became creative, able to innovate in the modern manner. By then, we were
likely speaking long, complicated sentences and were able to think
complicated thoughts in a way that we weren’t earlier (the subject of my
book, A Brief History of the Mind).
That gave us a new aspect of consciousness, an ability to routinely
contemplate situations in a way that, judging from the results, apes
usually cannot. We could finally speculate in complex ways, judging a
half-dozen elements for how well they hang together. And, when the
collection was as incoherent as our nighttime dreams, improving it until
the selection was finally good enough. You need a way to improve quality
offline and to judge when it’s finally good enough to act on. We do this
effortlessly whenever we speak a sentence that we’ve never spoken
before, which we do many times a day.
While we share a great deal of our mental life with the great apes,
especially the emotional judgments and the more elementary cooperation
strategies, we can safely innovate in ways they cannot. We can structure
our mental life, and keep different concepts from blending together like
a summer drink. Syntax makes long complex thoughts possible. We
structure contingent plans for the weekend. Chains of logic make our
rationality possible. We create structured music, the left hand and the
right hand playing different, interwoven melodies. Games where your next
move has to be checked against arbitrary rules are much like logic.
Structuring with quality control is so easy for us that we have
developed a fascination with discovering hidden order amid seeming
chaos. (We love those “eureka moments” enough to do crossword puzzles –
and sometimes even science research.) We puzzle out patterns from the
past and speculate about the future. Without such structuring and
quality control, the consciousness of other animals – even the great
apes – must be far less rich.
Such “higher intellectual functions“ lift us into a new realm of mental
abilities, but they are built on the foundation of what we see in the
remaining great apes. Intellect is only the icing on the basic cake, and
we still need to thoroughly understand that underlying setup to
appreciate how intellect works in our brains.
Alas, the wild populations of the great apes are crashing everywhere,
and there aren’t that many in zoos and research centers (for example,
only about 130 bonobos, worldwide). We could be left with only the bones
and the DNA, able to study real behavioral abilities only in some inbred
captive populations who have never known life in the environment that
evolution prepared them for.
I am indebted to a number of people for arranging behind-the-scenes
access at various zoos and research institutions. Even so, most of the
portraits were shot through the usual plate glass windows, terrible for
reflections and handprints. (That is, at least, better than shooting
through chain-link fence.) The other major obstacle is bright sunshine,
which always shadows the eyes; I eventually learned to visit zoos on
cloudy days. My best pictures were taken with a hand-held SLR digital
camera with a 100-400 mm image-stabilized telephoto lens, sometimes
shooting at five per second rates to catch the action.
If you want to try for yourself, you will discover that reflections from
the eye’s surface are a major challenge; studio portrait photographers
arrange the room lighting to eliminate them. Unless you can see the
eyes, it is seldom a good portrait. In real life, we get a number of
fixes on another’s direction of gaze and seldom notice the reflections.
What we “see” is really a reconstruction from a number of glimpses, not
a proper snapshot. Given only a single photographic image, we cannot
judge direction of gaze unless the reflections are softened so that the
pupil and iris can be sufficiently seen.
In the 1950s, I did a lot of semi-pro photography, mostly sports action
and candid portraits, both excellent preparation for this project. I had
the good fortune to work one summer for Eliot Elisofon, the LIFE
magazine photographic essayist, gourmet, and art collector. So my ape
photos have been digitally retouched using all the traditional darkroom
tricks that, in effect, rearrange the lighting. Brightness and contrast
have been locally manipulated to bring out dark details, distracting
backgrounds have been fogged, and skin folds highlighted. But there is
otherwise no distortion and nothing added that cannot be seen on close
inspection of the originals.
In order to allow for several rounds of revision as additional portraits
become available, I am publishing this preview edition via a
print-on-demand service of amazon.com.
In bypassing the usual one-year publication delays at a traditional
publisher, I don’t get the publicity machine that drives major book
reviews and bookstore orders. So I am relying on word-of-mouth much more
than usual. If you think the book suitable for a zoo or museum bookstore
(or book review), do write them and perhaps include a photocopy of the
book’s cover. The easy way to tell friends by email is to first go to my
ape website apes.WilliamCalvin.com and then send the URL with a covering
note. Ape portraits for computer wallpaper and posters are free
downloads from the website. Thanks.
READING UP ON APES
JULIAN CALDECOTT, LERA MILES (editors), World Atlas of Great Apes and
their Conservation (UNEP World Conservation Monitoring Centre;
University of California Press 2005).
WILLIAM H. CALVIN, A Brief History of the Mind: From Ape to Intellect
and Beyond (Oxford University Press 2004).
FRANS DE WAAL, Chimpanzee Politics: Sex and Power Among the Apes (1982,
revised edition, Johns Hopkins University Press 1998).
FRANS DE WAAL, Peacemaking Among Primates (Harvard University Press,
FRANS DE WAAL, Good Natured: The Origins of Right and Wrong (Harvard
University Press 1996).
FRANS DE WAAL, Bonobo: The Forgotten Ape (University of California
Press, 1997). With excellent photographs by Frans Lanting.
FRANS DE WAAL, The Ape and the Sushi Master (Basic Books/Perseus, 2001)
FRANS DE WAAL, Our Inner Ape (Harvard University Press, 2005).
MARC D. HAUSER, Wild Minds (Holt 2000).
SARAH BLAFFER HRDY, Mother Nature : A History of Mothers, Infants, and
Natural Selection (Pantheon Books 1999).
WILLIAM C. MCGREW, The Cultured Chimpanzee (Cambridge University Press
RONALD M. NOWAK, Walker’s Primates of the World (Johns Hopkins
University Press, 1999).
E. SUE SAVAGE-RUMBAUGH, STUART SHANKER, Apes, Language, and the Human
Mind (Oxford University Press 1998).
PAR SEGERDAHL, WILLIAM FIELDS, SUE SAVAGE-RUMBAUGH, Kanzi’s Primal
Language : The Cultural Initiation of Primates into Language (Palgrave
CRAIG B. STANFORD, The Hunting Apes (Princeton University Press 1998).
CAREL VAN SCHAIK, Among Orangutans. (Harvard University Press 2004).
RICHARD WRANGHAM, DALE PETERSON, Demonic Males: Apes and the Origins of
Human Violence (Houghton Mifflin 1996).
For evolution and cognition more generally:
WILLIAM H. CALVIN, A Brain for All Seasons: Human Evolution and Abrupt
Climate Change (University of Chicago Press 2002).
RICHARD DAWKINS, Climbing Mount Improbable (W. W. Norton 1996).
DANIEL C. DENNETT, Darwin‘s Dangerous Idea (Simon & Schuster 1995).
DANIEL C. DENNETT, Freedom Evolves (Viking 2003).
MERLIN DONALD, A Mind So Rare: The Evolution of Human Consciousness
MARK JOHNSON, Moral Imagination : Implications of Cognitive Science for
Ethics (University of Chicago Press 1993).
STEVEN PINKER, The Blank Slate: The Modern Denial of Human Nature
ELLIOTT SOBER, DAVID SLOAN WILSON, Unto Others: The Evolution and
Psychology of Unselfish Behavior (Harvard University Press 1998).
MICHAEL TOMASELLO, The Cultural Origins of Human Cognition (Harvard
University Press 2000).
Web resources. http://www.unep.org/grasp/ GRASP is the UN’s Great Ape
Survival Project. See also http://pin.primate.wisc.edu/factsheets.
Visiting the Apes. The best zoo in the United States for apes is the San
Diego Zoo and Wild Animal Park (together, they have five of the six ape
species -- but not chimpanzees). I also can recommend Seattle’s Woodland
Park Zoo (only orangutans, gorillas, and siamangs but in excellent
exhibits) and the Milwaukee Zoo (which has an excellent bonobo exhibit,
except for its window reflections ruining all of my photographs). There
are many U.S. zoos with gorillas and orangutans but no other apes.
In the UK, the Chester Zoo has a large chimpanzee exhibit and the
Twycross Zoo has all six ape species. Twycross is between Birmingham and
Leicester in the countryside north of Coventry. It can be seen as a day
trip from London by train and taxi (see my website for directions).
Arnhem, near the Dutch-German border on the Rhine, has a large
chimpanzee colony and a good gorilla exhibit. The Cologne Zoo upriver
has a number of bonobos. A more extensive zoo list, plus updates and
chapter notes for this book, is on the web at apes.WilliamCalvin.com.
alliance, 80, 125
Arnhem, 28, 31, 63, 66, 69, 72, 73, 74, 75, 76, 189
Asia, 61, 126, 149, 180
attention, 7, 102
Big Sister, 37, 92
blindman’s buff, 84
body size, 61
bonobo, 6, 11, 12, 15, 20, 32, 33, 34, 58, 64, 79, 80, 81, 82, 84, 85,
87, 92, 100, 103, 104, 105, 147, 150, 178, 180, 181, 182, 189
Borneo, 111, 121, 126, 166
brain, 2, 181, 188, 200
brain size, 149
CALVIN, 2, 3, 4, 187, 188
capuchin, 58, 171
cardiovascular disease, 77
chimpanzee, 5, 27, 28, 31, 36, 62, 63, 64, 65, 66, 67, 68, 70, 71, 72,
73, 74, 75, 76, 77, 79, 81, 82, 85, 88, 110, 123, 128, 131, 148, 171,
178, 180, 189, 203
chimps, 11, 58, 62, 63, 65, 67, 70, 71, 72, 73, 79, 81, 82, 147, 148,
150, 178, 179, 181
coalition, 65, 147
Congo, 79, 85, 131
consciousness, 181, 182, 195, 199
cooperation, 147, 181
creative, 123, 181, 198
DARWIN, 176, 179, 180, 188
DE WAAL, 62, 66, 70, 76, 80, 110, 168, 171, 187
DENNETT, 176, 188
Denver Zoo, 170
diet, 63, 147, 150, 181
evolution, 4, 77, 147, 180, 182, 188, 189
eye contact, 62
family resemblance, 75
fan clubs, 7
FIELDS, 103, 188
food, 55, 58, 63, 111, 112, 132, 143, 147, 171, 181
FOSSEY, 128, 148
fruit, 73, 111, 112, 126, 148
gang, 70, 71, 178
genes, 77, 106
gibbons, 126, 150, 158, 163
gorilla, 30, 35, 39, 40, 45, 56, 63, 79, 110, 131, 136, 147, 148, 189
grammar, 104, 176
Great Ape Survival Project, 189
Great Ape Trust of Iowa, 4, 101, 103, 104
great apes, 5, 58, 102, 109, 112, 131, 134, 160, 170, 177-182, 203
grin, 12, 99
grooming, 31, 32, 34, 62, 65, 81, 106, 121
hair, 32, 72, 81, 106, 154, 163
hands, 11, 118, 160
higher intellectual functions, 182
Homo erectus, 180
Homo sapiens, 177, 180, 181
human, 4, 7, 9, 23, 65, 67, 71, 77, 82, 85, 102, 122, 176, 177, 178,
179, 180, 195, 197, 198, 200
hunting, 71, 124, 181
ice ages, 126
imitation, 67, 131
intellect, 182, 187
intelligence, 147, 176
Kanzi, 101, 103, 104, 188
language, 4, 32, 101, 103, 104, 176, 198, 199
lemurs, 173, 174
life span, 61
Malay peninsula, 126
mental life, 181, 195
Milwaukee Zoo, 172
mirror, 110, 134
monkey, 7, 63, 64, 149, 169, 170, 171, 173, 179, 180
mother, 31, 58, 67, 75, 81, 87, 168
mouth, 9, 14, 110, 119, 121, 186
Neanderthals, 180, 181
orangutan, 10, 14, 16, 17, 25, 26, 29, 41, 58, 73, 109, 110, 111, 112,
113, 114, 118, 119, 120, 121, 122, 123, 124, 126, 127, 161, 180, 188,
Patas monkey, 169
play bite, 10
play face, 5, 9, 11
playing, 12, 65, 117, 182
posture, 103, 180
precision grip, 122
quality, 143, 181, 182, 195, 198
religious prohibitions, 127
San Diego Wild Animal Park, 132, 146
San Diego Zoo, 7, 16, 20, 45, 97, 122, 130, 155, 156, 163, 189
SAVAGE-RUMBAUGH, 103, 104, 188
Seattle, 4, 25, 26, 39, 111, 152, 153
sentences, 101, 181
sexual behavior, 80
shared attention, 102
sharing, 5, 55, 56, 58, 132, 171, 181
siamang, 16, 121, 126, 149, 150, 151, 152, 155, 157, 163, 189
STANFORD, 148, 188
stare, 5, 34, 45, 174
Sumatra, 111, 112, 126, 127
syntax, 176, 182, 198
syrup, 63, 88, 97, 122
tail, 149, 169
termite, 64, 122, 129
termite mound, 122
termites, 63, 64, 88, 128
thought, 181, 182
TOMASELLO, 67, 189
tool, 64, 80, 118, 122, 123, 131, 171
tool use, 80, 123, 131
Twycross Zoo, 10, 14, 27, 36, 67, 68, 71, 77, 87, 90, 98, 100, 105, 117,
164, 165, 166, 173, 189
understanding, 101, 179
UNEP World Conservation Monitoring Centre, 187
upright, 82, 160
VAN SCHAIK, 58, 109, 127, 188
vegetarian, 5, 63 129, 143, 147
Wild Animal Park, 45, 163
Woodland Park Zoo, 152, 189
words, 101, 104
The great apes 61
Begun, D.R. (2003) Planet of the apes. Scientific American 289: 74–83.
The omnivorous Chimpanzee 63
Family resemblance: UNEP, p.64.
Toshisada Nishida, Lack of "Group Play" in Wild Chimpanzees. Pan Africa
News 11:1 (June 2004). At http://mahale.web.infoseek.co.jp/PAN/11_1/11(1)_02.html
Cawthon KA. 2005 June 28. Primate Factsheets: Chimpanzee (Pan
troglodytes) Taxonomy, Morphology, & Ecology. http://pin.primate.wisc.edu/factsheets/entry/chimpanzee.
Ajit Varki,. Genome Research 10:1065-1070, 2000.
Maynard Olson and Ajit Varki, Nature Reviews Genetics., 4: 20-28, 2003
Bonobo, the “left-bank chimp” 81
K. Zamma & S. Fujita, Genito-genital rubbing among the chimpanzees of
Mahale and Bossou. Pan Africa News 11,issue 2 (2004)
Sue Savage-Rumbaugh: UNEP World Atlas (2005), p.91
Bonobo tool use: G. Hohmann, B. Fruth (2003) Culture in bonobos?
Between-species and within-species variation in behaviour. Current
Anthropology 44: 563–571.
van Schaik (2004), p.117.
The orange oriental Orangutan 111
Borneo food supply: http://animaldiversity.ummz.umich.edu
Orangutan speed of learning in zoo: UNEP (2005) p.168.
The vegetarian Gorilla 129
Gorilla diet: http://animaldiversity.ummz.umich.edu
Tool use: T. Breuer, M. Ndoundou-Hockemba, V. Fishlock (2005). First
Observation of Tool Use in Wild Gorillas. PLoS Biology 3(11): e380.
Mirrors: Mark Hauser (2000), p.101.
The acrobatic Siamang 151
Siamang facts: http://animaldiversity.ummz.umich.edu
The ape hidden in plain sight 177
Dennett 2003, p.2
Ernst Mayr 1999
Our mental illnesses also suggest that human evolution was too rapid to
get the bugs out. See my 2004 book, A Brief History of the Mind (Oxford
Photographs and text by
William H. Calvin, a professor at the University of
and the author of a dozen popular books on
who won the 2002 Phi Beta Kappa book
prize for science as literature.