How Humans Evolved

Robert Boyd . Joan B. Silk

Uníversity of California, Los Angeles

E
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For Sam and Ruby About the Authors

ROBERT BOYD has written widely on evo-

W. W. Norton & Company has been independent since its founding in 1923, when William Warder
Norton and Mary D. Herter Norton first published lectures delivered at the People's lnstitute, the lutionary theory, focusing especially on the
adult education division of New York City's Cooper Union. The firm soon expanded its program evolution ofcooperation and the role ofculture
beyond the lnstitute, publishing books by celebrated academics from America and abroad. By
mid-century, the two major pillars of Norton's publishing program-trade books and college texts-
in human evolution. His book Cubure and.the
were firmly established. ln the 1950s, the Norton family transferred control of their company to its Evolutionary Process received the J. I. Staley Prize.
employees, and today-with a staff of four hundred and a comparable number of trade, college,
He has also published numerous articles in
and professional titles published each year-W. W. Norton & Company stands as the largest and
oldest publishing house owned wholly by its employees. scientific journals and has edited volumes. He
is currently Professor of Anthropology at the
University of California, Los Angeles.
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Printed in the United States of America JOAN B. SltK has conducted extensive re-
Sixth Edition
search on the social lives of monkeys and apes,

Manufacturing by Courier, Kendallville including extended fieldwork on chimpanzees at

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Project editor: lVlelissa Atkin to human behavior. She has published numerous
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She is currently Professor ofAnthropology and
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Library of Congress Cataloging-in-Publication Data

Boyd, Robert, Ph. D. I
How humans evolved/Robert Boyd, Joan B. Silk.-6th ed.
p. cm.
lncludes b¡bliograph¡cal references and index.
lsBN 978-0-393-9L227 -2 (pbk.)
1. Human evolution. l. Silk, Joan B. ll. Title.
GN281.866 2011
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 of diversity in the behavior and morphology of organisms in relation to their environments,
we can see how evolution shapes adaptation in response to different selective pressures.
This approach is called "reasoning by analogy."

Primates Are Our Closest Relatives

and other primates share many characteristics, other
I Because humans
I primates provide valuable insi$hts about early humans.

We humans are more closely related to nonhuman primates than we are to âny other ani
mal species. The anatomical similarities among monkeys, aPes, and humans led the Swedish
naturalist Carolus Linnaeus to place us in the order Primates in the first scientific taxonomy,
Systema Naturøe, published in 1735. Later, naturalists such as Georges Cuvier andJohann
Blumenbach placed us in our own order because of our distinctive mental capacities and
upright posrnre. ln The Descent' of Man, hgwever, Charles Darwin firmly advocated rein-
staring humans in the order Primates; he cited the biologist Thomas Henry Huxley's essay
enumerating the many anatomical similarities between us and apes, and he suggested that
"if man had not been his own classifier, he would never have thought of founding a separate
order for his own reception." Modern systematics unambiguously confirms that humans
are more closely related to other primates than to any other living creatules.
Because \Me âre closely related to other primates, we share with them many aspects of
morphology, physiology, and development. For example, like other primates, we have well-
developed vision and grasping hands and feet. We share features of our life history with
other primates as well, including an extended period ofjuvenile development and larger
brains in relation to body size than the members of other taxonomic groups. Homologies
between humans and other primates also extend to behavior, since the physiological ahd
cognitive structures that underlie human behavior are niore similar to those of other pri
mates than to members of other taxonomic groups. The existence of this extensive array of
homologous traits, the product of the common evolutionary history of the primates, means
that nonhuman primates provide useful models for understanding the evolutionary roots

Primate Diversity and Ecology of human morphology and for unraveling the origins of human nature.

Primates Are a Diverse Order

Two Reasons to Study Diversity within the primate order helps us to understand how natural
Primates Two Reasons to Study Primates I
I selection shapes behavior.
Primates Are Our Closest
Relatives The chapters in Part T\r¡o focus on the behavior of living nonhuman pri- During the last 30 years, hundreds of researchers from a variety of academic discipìines
Primates Are a Diverse have spent thousands of hours observing many different species of nonhuman primates in
mates. Studies of nonhuman primates help us understand human evo-
Order the wild, in captive colonies, and in laboratories. All primate species have evolved adapta-
Features That Define the lution for two complementary but distinct reasons. First, closely related tions that enable them ro meet the basic challenges of life, such as finding food, avoiding
Primates predators, obtaining mates, rearing young, and coping with competitors' At the same time,
I
species tend to be similar morphologically, because, as we saw in Chapter
Primat€ Biogeography there is great morphological, ecological, and behavioral diversity among species within
4, they share traits acquired through descent from a common ancestor. the p r i m a're o rde r r aîå
i;å:iil:J*;
Fo e x
A Taxonomy of
Primates
Living
For example, viviparity (bearing live young) and lactation are traits that :iäîi:ì J ä:fi il ï1:î:îl::iåi3.
ies live in dense tropical forests; others are at home in open woodlands
Primate Diversity all placental and marsupial mammals share, and these traits distinguish subsist almost entirely on leaves; others rely on an omnivorous diet
The Strepsirrhines of fruits, leaves, flowers, seeds, gum, nectar, insects, and small animal prey. Some species
The Haplorrhines mammals from other taxa, such as reptiles. The existence of such simi-
are solitary, and others are highly gregarious. Some are active at night (nocturnal); others
Primate Ecology larities means that studies of living primates often give us more insight are acrive during daylight hours (diurnal). One primate, the fat-tailed dwarf lemur, enters
The Distribution of Food a torpid state and sleeps for six months each year. Some species actively defend territories
Activity Patterns into the behavior of our ancestors than do studies of other organisms. from incursions by other members of their own species (conspecifics); others do not. In
Ranging Behavior
This approach is called "reasoning by homology." The second reason some species, only females provide care of their young; in others, males participate actively
Predation
in this process.
Primate Sociality we study primates is based on the idea that natural selection leads to wever, evidence of diversity among closely
Primate Conservation similar organisms in similar envir ments. By assessing the patterns fferent ecological and social conditions also

TWO REASONS TO STUDY PR]MATES 103

LO2
helps researchers to understand how evolution shapes behavior. Animals that are ciosely
related to one another phylogenetically tend to be very similar in morphology, physiology,
life history, and behavior. Thus, differences observed among closely related species are
likeiy to represenr adaptive responses to specific ecological conditions. At the same time,
similarities among more distantly related creatures living under similar ecological condi
tions are likely to be the product ofconvergence.
This approach, sometimes called the "comparative method," has become an important
form of analysis as researchers attempt to explain the patterns of variation in morphology
ame principles have been borrowe
members of the human lineage. B
he comparative method Provides
means of testing hypotheses about the lives of our hominin ancestors. For exam-
ple, the observation that there are substantial differences in male and female body size, a
phenomenon called sexual dimorphism, in species that form non-pair-bonded groups sug-
gesrs thar highly dimorphic hominins in the past were not pair-bonded. In Part Three, we
will see how the data and theories about behavior produced by primatologists have played
an important role in reshaping our ideas about human origins.

Features That Define the Primates

I Members of the primate order are characterized by a number of shared,
I derivecl characters, but not all primates share all of these traits.

The animals pictured in Figure 5.1 are all members of the primate order. These animals
are similar in many ways: they are covered with a thick coat of hair, they have four limbs,
and they have five fingers on each hand. They give birth to live young, and mothers suckle
their offspring. However, they share these ancestral features with all mammals. Beyond

TABLE 5.1

1. The big toe on the foot is opposable, and hands are ptehensile. This means that
primates can use their feet and hands for grasping. The opposable big toe has been
lost in humans. F¡gure 5.1 All of these animals
2. There are flat nails on the hands and feet in most species, instead of claws, and are primates: (a) aye-aye, (b) ring-
there are sens¡tive tactile pads with "fingerprints" on f¡ngers and toes. tailed lemur, (c) langur, (d) howler,
(e) gelada baboon. Primates are a
3. Locomotion is hind-limb-dominated, meaning that the hind limbs do most of the diverse order and do not Possess
work, and the cénter of Sravity is nearer the hind limbs than the forelimbs' a suite of traits that unambigu-
4. There is an unspecialized olfactory (smelling) apparatus that is reduced in diurnal ously d¡stinguish them from other
primates. animals.

5. The visual sense is highly developed. The eyes are large and moved forward in the (d) (e)
head, providing stereoscopic v¡sion.
6. Females have small l¡tters, and gestation and juvenile periods are longer than in
these ancesrral features, it is hard to see what the members of this group of animals have
other mammals of similar size.
in common that makes them distinct from other mammals. What distinguishes a ring-
7. The brain is large compared wlth the brains of similarly sized mammals, and it has a tailed lemur from a mongoose or a raccoon? What features link the langur and the aye-aye?
number of unique anatomical features.
In fact, primates are a rather nondescript mammalian order that cannot be unambigu-
8. The molars are relatively unspecialized, and there is a maximum of two incisors, one ously characterizedby a single derived feature shared by all members. In his extensive
canine, three premolars, and three molars on each half of the upper and lower jaw. treatise on primate evolution, however, the biologist Robert Martin of the Field Museum
g. There are a number of other subtle anatomical characteristics that are useful to of Natural History in Chicago defines the primate order in terms of the derived features
systematists but are hard to interpret functionally. listed in Table 5.1.
The first three traits in Table 5.1 are related to the flexible movement of hands and feet.
Definition of the primate order. See the text for more complete descriptions of the Primates can grasp with their hands and feet (Figure 5.2a), andmost monkeys and apes can
above features. oppose their thumb and forefinger in a precision grip (Figure 5.2b). The flat nails, distinct
from the claws of many animals, and the tactile pads on the tips of primate fingers and toes

FEATURES THAT DEF]NE THE PRIMATES

CHAPTER 5 Prímate Diversíty and Ecology

Fi(ure 5.2 (a) Primates have
grasping feet, which they use
to climb, cling to branches,
hold food, and scratch them-
selves. (b) Primates can
oppose the thumb and fore-
f¡nger in a precision grip-a
feature that enables them to
hold food in one hand while
they are feeding, to pick small
ticks and bits of debris from
their hair while grooming, and
(in some species) to use tools.
(c) Most primates have flat
nails on their hands and sensi-
tive tactile pads on the tips of
their fingers.
Figure 5.3 ln most primates, the eyes are moved forward in
the head. The field of vision of the two eyes overlaps, creating
binocular, stereoscopic vision. (Photograph courtesy of Carola
Borries.)
106
(a) (c)
further enhance their dexterity (Figure 5.2c). These traits enable primates to use their hands
and feet differently from the ways most other animals do. Primates are able to grasp fruit,
squirming insects, and other small items in their hands and feet, and they can grip branches
with their fingers and toes. During grooming sessions, they delicately part their partner's
hair and use their thumb and forefinger to remove small bits of debris from the skin.
Traits 4 and 5 in Table 5.1 are related to a shift in emphasis among the sense organs,
Most primates are characterizedby a greater reliance on visual stimuli and less reliance
on olfactory stimuli than other mammals. Many primate species can perceive color, and
their eyes are set forward in the head, providing them with bin-
ocular, stereoscopic vision (Figure 5.3). Binocular vision means
that the fields ofvision ofthe two eyes overlap so that both eyes
perceive the same image. Stereoscopic vision means that each
eye sends a signal of the visual image to both hemispheres in
the brain to create an image with depth. These trends are not
uniformly expressed within the primate order; for example,
olfactory cues play a more important role in the lives of strep-
sirrhine primates than in the lives of haplorrhine primates. As
we will explain shortly, the strepsirrhine primates include the
lorises and lemurs, and the haplorrhine primates include tarsi
els, monkeys, and apes.
Features 6 andT in Table 5.1 result from the distinctive life
history of primates. As a group, primates have longer pregnan-
cies, mature at later ages, live longer, and have larger brains
than other animals of similar body size. These features reflect
a progressive trend toward increased dependence on complex
behavior, learning, and behavioral flexibility within the pri-
mate order. As the noted primatologist Alison Jolly points out,
CHAPTER 5 Primate Diversity and Ecology
"lf there is an essence t is the progressive
evolution o we will see in the
chapters that fo ese traits have a profound impact on mat-
ing and parenting strategies and the patterns ofsocial interaction
within primate groups.
The eighth feature in Table 5.1 concerns primate dentition.
Teeth play a very important role in the lives of primates and
in our understanding of their evolution. The utility of teeth to
primates themselves is straightforward: Teeth are necessary for
processing food and are also used as weapons in conflicts with
other animals. Teeth are also useful features for those who study
living and fossil primates. Primatologists sometimes rely on
tooth wear to gauge the age of individuals, and they use features
Fi¡lure 5.4 A high degree of intelligence characterizes some
of the teeth to assess the phylogenetic relationships among spe-
animals besides primates. Dolphins, for example, have very
cies. As we will see, paleontologists often rely on teeth, which
large brains in relation to their body size, and their behavlor is
are hard and preserve well, to identify the phylogenetic relation- quite complex.
ships of extinct creatures and to make inferences about their
developmental patterns, their dietary preferences, and their social structure. Closer Look 5.1
describes primate dentition in greater detail.
Although these traits are generally characteristic of primates, you should keep two
points in mind. First, none of them makes primates unique. Dolphins, for example, have
large brains and extended periods ofjuvenile dependence, and their social behavior may
be just as complicâted and flexible as that of any nonhuman primate (Figure 5.4). Second,
not every primate possesses all of these traits. Humans have lost the grasping big toe that
characterizes other primates, some strepsirrhine primates have claws on some of their fin-
gers and toes, and not all monkeys have color vision.
Primate Biogeography
N Prirnates are nrainly restrictecl to tropicarl tegions of the world.
The continents of Asia, Africa, and South America and the islands that iie near their coasts
are home to most of the world's primates (Figure 5.5). A few species remain in Mexico and
Central America. Primates \¡r'ere once found in southern Europe, but no natural population
survives there now. There are no natural populations of primates in Australia or Antarctica,
and none occupied these continents in the past.
Fi¡lure 5.5 The distribution
of living and fossil primates.
Primates are now found in Cen-
tral America, South America,
Africa, and Asia. They are found
mainly in tropical regions of the
world. Primates were formerly
found in southern Europe and
northern Africa. There have
never been indigenous popula-
tions of primates in Australla or
! Fossil only Anta rctica.
PR]MATE BIOGEOGRAPHY lo7

or various reasons, biological
anthropologists spend a lot of
time thinking about teeth. Teeth
are useful markers for taxonomic iden-
tity, since various kinds of primates
have different numbers ofteeth. Teeth
are also useful because they tell us
things about what kinds of food pri
mates eat. If we can detect a relation-
ship between dental morphology and
diet, we can apply these insights to the
fossil record. This is particularly handy
because teeth are the most commonly
preserved parts of the body. Finally,
teeth and gut morphology provide
examples of how natural selection has
created adaptations that enable animals
to cope with their environments more
effectively.
Dental Formula
To appreciate the basic features ofpri-
mate dentition, you can consult Figure
5.6, or you can simply look in a mirror,
since your teeth are much like those
of other primates. Teeth are rooted in
the jaw. The jaw holds four different
kinds ofteeth: in order, they are, first,
the incisors at the front; then come the
canines, premolars, and the molars in
the rear. All primates have the same
kinds ofteeth, but species vary in how
many of each kind of tooth they have.
For convenience, these combinations
are expressed in a standard format
called the dental formula, which is
commonly written in the following
form:
2.t.3.3
2.1.3.3
Reading from left to right, the numer-
als tell us how many incisors, canines,
premolars, and molars a particular spe-
cies has (or had) on one side of its jaw.
The top line of numbers represents
the teeth on one side ofthe upperjaw
r08
(maxilla), and the bottom line repre-
sents the teeth on the corresponding
side of the lower jaw (mandible). Usu-
all¡ but not always, the formula is the
same for both upper and lower jaws.
Like most other parts of the body, our
dentition is bilaterally symmetrical,
which means that the left side is identi
cal to the right side. The ancesual pat-
tern shown here has been modified in
various primate taxa, as the total num-
ber ofteeth has been reduced.
The dental formulas among living
primates vary (Table 5.2). The lorises,
pottos, galagos, and a number of lemu-
rids have retained the primitive dental
formula, but other strepsirrhine taxa
have lost incisors, canines, or premo-
lars. Tarsiers have lost one incisor on
the mandible but have retained two on
the maxilla. All of the New World mon-
keys, except the marmosets and tama-
rins, have retained the primitive dental
TABLE 5.2
Primate taxa
Strepsirrhines Lorises, pottos, galagos, dwarf lemurs,
mouse lemurs, true lemurs
lndris
Aye-ayes
Haplorrhines Tarsiers
New World monkeys (most species)
Marmosets, tamarins
Old World monkeys, apes, humans
Primates vary ¡n the numbers of each type oftooth that they have. The dental
formulas given here give the number of incisors, canines, premolars, and
molars on each side of the upper jaw (maxilla) and lower jaw (mandible).
formula; the marmosets and tamarins
have lost one molar. The Old World
monkeys, apes, and humans have only
two premolars.
Dental Morphology
Primates who rely heavily on gum
tend to have large and prominent inci
sors, which they use to gouge holes in
the bark of trees (Figure 5.7). In some
strepsirrhine species, the incisors and
canines are projected forward in the
jaw and are used to scrape hardened
gum off the surface of branches and
tree trunks. Dietary specializations are
also reflected in the size and shape of
the molars. Primates who feed mainly
on insects and leaves have molars with
well-developed shearing crests that per-
mit them to cut their food into small
pieces when they chew. Insectivores
tend to have higher and more pointed
cusps on their molars, which are useful
for puncturing and crushing the bod-
ies of their prey. The molars of frugivo-
res tend to have flatter, more rounded
cusps, with broad and flat areas used to
Dental Formula
2.7.3.3
2.1.3.3
2.1.2.3
2.0.3.3
1.0.1.3
1.0.0.3
2.7.3.3
1.1.3.3
2.1.3.3
2.7.3.3
2.1.3.2
2.7.3.2
2.1,2.3
2.!.2.3
Flture 5.6 The upperjaw (left) and
lower jaw (right) are shown here for a
male colobine (a) and a male gorilla (b).
Canine
ln Old World monkeys, the prominent Canine
anter¡or and posterior cusps of the
lower molars form two parallel ridges. ln
apes, the five cusps of the lower molar
eremotars eremotars
form a Y-shaped pattern.
{ {
crush their food. Primates who rely on
1cm 1cm
hard seeds and nuts have molars with
very thick enamel that can withstand (a) (b)
the heavy chewing forces needed to
process these types offood.
Guts
Well-developed
Primates who feed principally on Broad molar shearing v
insects or animal prey have relatively lnctsors crests ,l
simple digestive systems that are spe- Low rounded molar cusps Propithecus
cialized for absorption. They generally
have a simple small stomach, a small
Cerocebus
cecum (a pouch located at the upper
end of the large intestine), and a small
Large cecum Complex stomach
colon relative to the rest of the small
intestine. Frugivores also tend to have
simple digestive systems, but frugivo-
rous species with large bodies have
capacious stomachs to hold large quan-
tities of the leaves they consume along
with the fruit in their diet. Folivores
have the most specialized digestive
systems because they must deal with Lepilemur Colobus
large quantities ofcellulose and second-
ary plant compounds. Because primates
cannot digest cellulose or other struc-
Enlarged
tural carbohydrates directly, folivores large intesin-e
maintain colonies of microorganisms
Callithrix
in their digestive systems that break
down these substances. In some spe- Stout
cies, these colonies of microorganisms rncrsors Leaf eaters
are housed in an enlarged cecum; in
other species, the colon is enlarged for
this purpose. Colobines, for example,
have an enlarged and complex stomach
divided into a number of different sec-
tions where microorganisms help pro-
cess cellulose.
Flglure 5.7 The dentition and digestive
tracts of fruit-eâting (frugivorous), leaf-
Long cecum
eating (folivorous), gum-eating (gummiv-
Euoticus
orous), and insect-eating (insectivorous)
primates typically differ. lnsect eaters
109
 Primates are found mainly in tropical regions, where the fluctuations in temperature
from day to night greatly exceed fluctuations in temperature over the course ofthe year.
In the tropics, the distribution of resources that primates rely on for subsistence is affected
more strongly by seasonal changes in rainfall than by seasonal changes in temperature.
Some primate species extend their ranges into temperate areas of Africa and Asia, whele
they manage to cope with substantial seasonai fluctuations in envrronmental conditions.
Within their ranges, plimares occupy an extremeiy diverse set of habitats that includes
all types of tropical forests, savallna woodlands, mal1gl'ove s\Mamps, grasslands, high-
altitude plateaus, and deser
ir.rcluding a well-developed sense of smell, large eyes, and independently movable ears. By
contrast, monkeys, apes, and humans, which make up the suborder Haplorrhini, evoÌved
adaptations more suited to a diurnal lifestyle early in their evolutionary history. In the
Haplorrhini, traits lelated to increased complexity of behavior, inciuding large brains and
lor.rger life sparls, are most fully developed. Haplorrhine monkeys are generally larger rhan
strepsirrhines, are active during the day, are more fully dependent on vision than smell,
and live in bigger and more complex social groups.
The classification of the primates that we have adopted here reflects the pattern of
descent within the order. Tarsiers are included in the haplorrhines because genetic and
morphological data indicate that they are more closely related to monkeys and apes than
ro the strepsirrhines. However, like many of the strepsirrhines, they are smali-bodied and
nocturnal. A cladistic classification places tarsiers within the haplorrhini, but ân evolution-

A Taxonomy of Living Primates a1'y taxonomy would group tarsiers with strepsirrhines because of their overall similarity
in morphology, genetics, and behavior'.
Scientists classify primates into two suborde
Many of the primates included in the suborde\' noctumal, and, like some
of the earliest primates that lived 50 mya, they have many adaptations to living in darkness,
Primate Diversity
f . ; .:

TABLE 5.3
ii,. ti!i) riì.i'r)lr:itfi.iii!l(: iì!-ititil!r)r; ;¡til rirVl¡ir:ri il¡rü ì-wri ì¡lír;rt.,irit:r:;: i.¡:tr¡l¡rii,-¡¡¡ilr:l'
ri ;r:l{i I {lf!1;ll{iiiriirll
SUBORDER INFRAORDER SUPERFAMILY FAMITY SUBFAMITY EXAMPLES
The infraorder Lemuriformes includes lemurs, which are found only on Madagascar and
Strepsirrhini Lemuriformes Lemuroidea Cheirogaleìdae Dwarf lemurs, mouse
the Comoro Islands, offthe southeastern coast of Africa. These islands have been sepa-
lemurs
rated from Africa for 120 million years. The primitive primates that reached Madagascar
Daubentoniidae Aye-ayes evolved in total isolation from primates elsewhere in the world, as well as from many of
lndriidae lndris, sifakas the predators and competitors that primates confront in,other places. Faced with a diverse
Lemuridae Lemurs set of available ecological niches, the lemurs underwent a spectacular adaptive radiation.
When humans first colonized Madagascar about 2,000 years ago, there were approximately
Lepilemuridae
44 species of lemurs, some as small as mouse lemurs and others as big as gorillas. In the
Lorisiformes Lorisoidea Galagidae Galaginae Galagos next few centuries, all of the larger lemur species became extinct, probably the victims of
Lorisidae Lorlsinae Lorises human hunters or habitat loss. The extant lemurs are mainly smal1 or medium-sized arbo-
Perodicticinae Pottos real residents offorested areas (Figure 5.8a); they travel quadrupedalÌy orbyjumpingin an

Haplorrhini Tarsiiformes Tarsiodea Tarsiidae Tarsinae Tarsiers

Platyrrhini Ceboidea Aotidae Aoti nae Owl monkeys
(New World monkeys)
Atelidae Alouattti nae Howler monkeys
Atelinae Spider monkeys
Callitrichidae Callitrichinae Marmosets, tamarins
Cebidae Cebinae Ca puch i ns

Saimirinae Squirrel monkeys

Pitheciidae Callicebinae Titi monkeys
Pithecinae Sakis
(b)
Catarrhini Cercopithecoidea Cercopithecidae Cercopithecinae Mangabeys, macaques,
(Old World monkeys) baboons
Figlure 5.8 (a) Ring-tailed lemurs, with their distinctive striped
Colobinae Langurs, colobus monkeys tails, live in social groups and are active during daylight hours.
Hominoidea Hylobatidae Gibbons, siamang ln a number of lemur species, females are dominant over
(Apes, humans) males. (b) Sifakas use their powerful legs to jump in an upright
posture, a form of locomotion known as vertical clinging and
Hominidae Ponginae Orangutans
leaping. (c) Galagos are small, arboreal, nocturnal animals who
Homininae Gorillas, chimpanzees, can leap great distances, They are mainly solitary, though resi-
humans dents of neighboiing territories sometimes rest together during
the day.

110 CHAPTER 5 Primate Díversity and Ecology PRIMATE DIVERSITY

 Marmosets and tamarins are also notable
upright posture from one tree to another, a form of locomotion known as vertical clinging birth to twins and sometimes triplets (Figure 5'10).
and leaping (Figure 5.8b). Activity patterns of lemurs are quite variable: about half are pri domestic arrangements: in most groups there is a single breeding paír, and other
for their
marily diurnal, others are nocturnal, and some are active during both day and night. one øroup members help the parents rear the offspring'
of the most interesting aspects of lemur behavior is that females routinely dominate males. '- irr. orher New world monkeys are generally larger than the marmosets and tamarins,
In most lemur species, females are able to supplant males from desirable feeding sires; and
in some lemur species, females regularly defeat males in aggressive encounters. Although
such behavior may seem unremarkable in our own liberated times, female dominance is
very rare in other primate species.
The infraorder Lorisiformes is comprised of small, nocturnal, arboreal residents of the
forests of Africa and Asia. These animals include two subfamilies with different types of
locomotion and activity patterns. Galagos are active and agile, leaping through the trees
and running quickly along branches (Figure 5.8c). The lorises move with ponderous delib-
eration, and their wrists and ankles have a specialized network of blood vessels that allows
them to remain immobile for long periods of time. These traits may be adaptations that
heip them avoid detection by predators. Traveling aione, the lorisiformes generally feed on
fruit, gum, and insect prey. The lorisiformes leave their dependent offspring in nests buiit
in the hollows of trees or hidden in masses of tangled vegetation. During the day, females
sleep, nurse their young, and groom, sometimes in the company of mature offspring or
familiar neighbors. Fiture 5.1O Marmosets are
small-bod¡ed South American
monkeys who form Pair-bonded
The Haplorrhines are the only nocturnal haplorhine primates.
or polyandrous social grouPS.
Males and older offspring Par-
'lhe subot'der Haplorrhini conta¡ns
| tlrree infraorders: Tarsiiformes, ticipate activelY in the care of
I Platyrrhirìi, and Catarrhini. i nfa nts.

The infraorder Tarsiiformes includes tarsiers, which are enigmatic primates thât live in the
rain forests of Borneo, sulawesi, and the Philippines (Figure 5.9). Like many of the strep-
sirrhine primates, tarsiers are small, nocturnal, and arboreal, and they move by vertical
clinging a

ir own weight; mothers leave their bulky infants behind in safe hiding places when
they forage for insects. Tarsiers are unique among primates because they are the only pri-
mates that rely exclusively on tter, feeding on insects and small vertebrate prey. FiÉure 5.11 Portraits of
The two infraorder commonly referred to as the New some cebid monkeYs. (a)
world monkeys and the old world mon Muriquis, or woollY sPider mon-
, respecriveiy, because platyrrhine
keys, are large-bodied and
arboreal. TheY are extremely
peaceful creatures, rarely fight-
ing or competing over access
The infraorder Platyrrhini (New World monkeys) is divided to resources. (b) SPidertnon-
into five separate families: Aotidae, Atelidae, Callitrichidae, Ceb- keys rely heavilY on riPe fru¡t
idae, and Pitheciidae. and travel in small Partìes.
Although the New World monkeys encompass considerable They have prehensile tails that
diversity in size, diet, and social organization, they do share they can use much like an
some basic features. All but those in one genus (Aotus) are diur- extra hand or foot. (c) CaPuchin
nal, all live in forested areas, and all are mainly arboreal. Most monkeys have larger brains
New World monkeys are quadrupedal, moving along the tops in relation to their bodY sizes
of branches and jumping between adjacent rrees. Some species than any of the other nonhu-
man primates. (d) Squirrel
in the family Atelidae can suspend rhemselves by their hands,
monkeys form large multimale,
feet, or tail and can move by swinging by their arms beneath
multifemale grouPs. ln the mat-
branches. ing season, males gain weight
The family Callitrichidae is composed of rhe marmosers and and become "fatted," and then
tamarins. These species share several morphological features compete actively for access
that distinguish them from other anrhropoid primare species: to receptive females' [Photo-
they are extremely small, rhe largest weighing less than tkge.z graphs courtesY of Sue Boinski
Flture 5.9 Tarsiers are small, insectivorous primates who lb); they have claws instead of nails; rhey have only two molars, (a), Susan Perry (c), and Carlão
live in Asia. Some tarsiers form pair bonds. while all other monkeys have three; and they frequently give Limeira (d).1

PRIMATE DIVERSITY 113

CHAPTER 5 Primate Diversity and Ecology
 I The itlfraorder Catarrhini contains the monkeys arrrl ape-s of tire Olcl Worlcl
I an,i humans.
As a group, the catarrhine primates share a number of anatomical and behavioral fea-
tures that distinguish them from the New world primates. For example, most old world
monkeys and apes have narrow nostrils that face downward, while New World monkeys
have round nostrils. old World monkeys have two premolars on each side of the upper and
lowerjaws; New World monkeys have three. Most Old World primates are larger than most
New world species, and old world monkeys and apes occupy a wider range of habitats
than New World species do.
The catarrhine primates are divided into two superfamilies: Cercopithecoidea (Old
World monkeys) and Hominoidea (apes and humans). Cercopithecoidea contains one exrant
(still living) family, which is further divided into two subfamilies of monkeys: Cercopith-
ecinae and Colobinae.

The superfan-ìily Cerco¡rithecoidea cncorrpasrìes grc¡rt rliversity in sr¡cial

I
I org¿ìr'ìlzatiorr, ecolo¡jical s¡recializations, ancl biogeography.

Colobine monkeys are found in the forests of Africa and Asia and may be the most
elegant of the primates (Figure 5.12). They have slender bodies, long legs, long tails, and (a) (b) (c)
often beautifully colored coats. The black-and-white colobus, for example, has a white ring
Fi(ure 5.13 Some representative cercopithecines: (a) Bonnet macaques are one of several
around its black face, a striking white cape on its black back, and a bushy white tail that flies
species of macaques that are found throughout Asia and North Africa. Like other macaques,
out behind as it leaps from tree to tree. Colobines are mainly leaf and seed earers, and most
bonnet macaques form multimale, multifemale groups, and females spend their entire lives ¡n
species spend the majority of their time in trees, They have complex stomachs, aimost like their natal (birth) groups. (b) Vervet monkeys are found throughout Africa. Like macaques and
the chambered stomachs of cows, which allow them to maintain bacterial colonies that baboons, females live among their mothers, daughters, and other maternal kin. Males transfer
facilitate the digestion of cellulose. Colobines are most often found in groups composed to nonnatal groups when they reach maturity. Vervets defend their ranges against incursions
of one adult male and a number of adult femaies. As in many other vertebrate taxa, the by members of other groups. (c) Blue monkeys live in one-male, multifemale groups. During the
replacement of resident males in one-male groups is often accompanied by lethal attacks mating season, however, one or more unfamiliar males may join bisexual groups and mate with
on infants by new males. Infanticide under such circumstances is believed to be favored by females. [Photographs courtesy of Kathy West (a) and Marlna Cords (c).]
selection because it improves the relative reproductive success of infanticidal males. This
issue is discussed more fully in Chapter 7.
Most cercopithecine monkeys are found in Africa, though one particularly adaptable
genus (Macacø) is widely distributed through Asia and part of Northern Africa (Figure 5.13).
The cercopithecines are more variable in size and diet than the colobines are. The social
behavior, reproductive behavior, life history, and ecology of a number of cercopithecine
species (particularly baboons, macaques, and vervets) have been studied extensively and
will figure prominently in the discussions of mating strategies and social behavior in the
nexr fe\M chapters. Cercopithecines typically live in medium or large bisexual (multimale,
multifemale) groups. Females typically remain in their natal groups (the groups into \¡/hich
they are born) throughout their lives and establish close and enduring relationships with
Fiture 5.12 (a) African colo-
theír maternal kin; males leave their natal groups and join new groups when they reach
bines, like this black-and-white
sexual maturity.
colobus monkey, are arboreal
and feed mainly on leaves.
These animals are sometimes I Thc s¡l)crf(ìnìily Horninoicle;r incluclr:s two f¿rnrilir.ts of aprl s: [lylolratidae
hunted for the¡r spectacular I (¡.Íibf,n''tr) arrrl lloltrirrirl¡rc (or¿ìngr.rt;lrs, gotillirs, cllinlllír!lzccs ¿rlld hu!'ì'larìÐ"
coats. (b) Gray langurs are
The hominoids are different from the cercopithecoids in a number of ways. The most
native to lndia and have been
readíly observed difference between apes and monkeys is that apes lacktails. But there are
the subject of extensive study
many orher more subtle differences between apes and monkeys. For example, the apes share
during the last three decades.
ln some areas, gray langurs some derived traits, including broader noses, broader palates, and larger brains; and they
form one-male, multifemale retain some primitive traits, such as relatively unspecialized molars. In Old World mon-
groups, and males engage in keys the prominenr anterior and posterior cusps are arranged to form two parallel ridges'
fierce fights over membership In apes, the five cusps on the lower molars are arranged to form a side-turned Y-shaped
in bisexual groups. ln these pattern ofridges (Figure 5.6).
groups, infanticide often fol- The family Hylobatidae includes lesser apes (gibbons and siamangs), and its living
lows when a new male takes members are now found in Asia. The family Hominidae includes the larger-bodied great
over the group. (b, Photograph
apes (orangutans, gorillas, bonobos, chimpanzees, and humans). Humans and other great
courtesy of Carola Borries.)
apes are somerimes placed in different subfamilies; in this classification, the other apes are
(a) (b)

Lt4 CHAPTER 5 Primate Diversity and Ecology PRIMATE D]VERSITY


il fl
;ü
(b)
Figlure 5.14 Gibbons (a)
and siamangs (b) live ¡n pair-
bonded groups and actively
defend their territories against
intruders. They have extremely
long arms, which they use to
propel themselves from one
branch to another as they
swing hand over hand through
the canopy, a form of loco-
motion called "brachiation."
Siamangs and gibbons are
confined to the tropical forests
of Asia. Like other residents
of trop¡cal forests, their sur-
vival is threatened by the rapid
destruction of tropical forests.
(Photographs courtesy of John
Mitani.)
Figure 5.15 (a) Orangutans
are large, ponderous, and
mostly solitary creatures. Male
orangutans often descend to
the ground to travel; lighter
females often move through
the tree canopy. (b) Today,
orangutans are found only
on the islands of Borneo and
Sumatra, in tropical forests like
this one.
116
placed in the subfamily Pongidae, while humans are placed in the Hominidae. Orangurans
are found in Asia, while chimpanzees, bonobos, and gorillas are resrricted to Africa.
The lesser apes are slightly built creatures \Mith extremely long arms in relation to their
body size (Figure 5.14). Gibbons and siamangs are strictly arboreal, and they use rheir long
arms to perform spectacular acrobatic feats, moving through the canopy with grace, speed,
and agility. Gibbons and siamangs are the only true brachiators among the primates, which
means they propel themselves by their arms alone and are in free flight between handholds.
(To picture this, think about swinging on monkey bars in your elementary school play-
ground.) Al1 of the lesser apes live in pair-bonded family groups; vigorously defend their
home ranges (the areas they occupy); and feed on fruit, leaves, flowers, and insects. Siamang
males play an active role in caring for young, frequently carrying them during the day; maie
gibbons are less attentive fathers. In territorial displays, mated pairs of siamangs perform
coordinated vocal duets that can be heard over long distances.
Orangutans, now found only on the Southeast Asian islands of Sumatra and Borneo, are
among the largest and most solitary species of primates (Figure 5.15). Orangutans have been
studied extensively by Biruté Galdikas in Tanjung Puting, Borneo, for more than 20 years.
Long-term studies oforangutans have also been conducted at Cabang Panti in Borneo, and
F¡gure 5.16 (a) Gorillas are the largest of the primates. lvlountain gorillas usually live in one-
at Ketambe and Suaq Balimbing in Sumatra. Orangutans feed primarily on fruit, but they
male, multifemale groups, but some groups contain more than one adult male. (b) Most behav-
also eat some leaves and bark. Adult females associate mainly with their own infants and
ioral information about gorillas comes from observations of mountain gorillas who live in the
immature offspring and do not often meet or interact with other orangutans. Adult males Virunga Mountains of central Africa, pictured here. The harsh montane habitat may influence
spend the majority of their time alone. A single adult male may defend a home range that the nature of soc¡al organization and social behavior in these animals, and the behavior of
encompasses the home ranges of several adult females; other males wander over larger areas gorillas living at lower elevations may differ. (Photographs courtesy of John lvl¡tani.)
and mate opportunisticaliy with receptive females. When resident males encounrer these
nomads, fierce and noisy encounters may take place.
Gorillas, the largest of the apes, existed in splendid isolation from Western science populations become available, however, we are revising some eiements of this view of
until the middle of the nineteenrh cenrury (Figure 5.16a, b). Today, our knowledge of gorilla social organization. For example, lowland gorillas seem to eat substantial amounts
the behavior and ecology of gorillas is based mainly on detailed long-term srudies of one of fruit, spend more of their time in trees, and form iarger and less cohesive social gtoups
subspecies, the mountain gorilla, at the Karisoke Research center in Rwanda, which was than mountain gorillas do.
founded by the late Dian Fossey. Mountain gorillas live in small groups that contain one As humankind's closest living relatives, chimpanzees (Figure 5.17a) have played a
or two adult males and a number of adult females and their young. Each day, mountain uniquely imporranr role in the study of human evolution. Whether reasoning by homology
gorillas ingest great quantities of various herbs, vines, shrubs, and bamboo. They eat little
fruit because fruiting plants are scarce in their mountainous habitat. Adult male mountain
gorillas, called silverbacks because the hair on their backs and shoulders turns a striking
silver-gray when they mature, play a central role in the structure and cohesion of their
social groups. Males sometimes remain in their natal groups to breed, but most males leave
their natal groups and acquire females by drawing them away from other males during
intergroup encounters. The silverback largely determines the timing of group activity
and the direction of travel. As data from newly established field studies of lowland gorilla
(a) (b) (c)
Figure 5.17 (a) Chimpanzees live in multimale, multifemale social groups. ln this spe-
cies, males form the core of the social group and remain in their natal groups for life. Many
researchers believe that ch¡mpanzees are our closest llving relatives. (b) Like other apes'
chimpanzees are found mainly in forests like this area on the shores of Lake Tanganyika in
Tanzania. However, chimpanzees sometimes range into more open areas as well. (c) Bono-
bos are members of the same genus as chimpanzees and are similar in many ways. Bonobos
are sometimes called "pygmy chimpanzees," but this is a misnomer because bonobos and
chlmpanzees are about the same size. This infant bonobo is sitting in a patch of terrestial
herbaceous vegetatlon, one of the staples of the bonobo's diet. (b, c, Photographs iourtesy
of John Mitani.)
(a) (b)
CHAPTER 5 Primate Diversíty and Ecology PRIMATE DIVERSITY
 or by analogy, researchers have found observations about chimpanzees to be important t. Basølmetabolism. Basal metabolic rate is the rate at which an animal
large
bases for hypotheses about the behavior of eally hominins.
How-
Detailed knowledge of chimpanzee behavior and ecology comes from a number of
long-term studies conducted at sites across Africa. In the 1960s, Jane Goodall began her c)

well-known study of chimpanzees at the Gombe Stream National Park on the shores of 2 î::'" (ú

Lake Tanganyika in Tanzania (Figure 5.12b). About the same time, a second study was ini- above baseline levels, The number of additional calories required depends .o
õ
tiated by the Ìate Toshisada Nishida at a site in the Mahale Mountains nor far from Gombe. on how much enelgy the animal expends' The amount ofenergy expended' _o
(ú
ó)
These studies are now moving into their sixth decade. Other important study sites have inturn,dependsonthesizeoftheanimalandhowfastitmoves.Ingen- E
eral, to sustain a normal range of actívities, an ave|age-sized primate
like
been established at Boussou, Guinea; in the'Iai Forest of Ivory Coast; and ar rwo sites in õU)
a baboon or macaque requires enough energy per day to
maintain a rate (ú
the Kibale Forest of Uganda: Kanyawara and Ngogo. c0
Bonobos (Figure 5.17c), another member of rhe genus Pan,ltve in inaccessible places and about twice its basal metabolic rate.
are much less well studied than common chimpanzees. Important field studies on bonobos 3. Growthrate. Growth imposes further energetic demands on organisms'
have been conducted at two sites in the Democratic Republic of the Congo (formerly Zaire): Infants and juveniies, that are gaining weight and growing in stature'
Wamba and Lomako. Field studies of bonobos have been disrupted by civil conflicts thar require more energy than would be expected on the basis of their body
have ravaged central Africa over the last decades. BodY weight
4. Primates the energetic costs
of
Chimpanzees and bonobos form large multimale, multifemale communiries. These
commnr-tities differ from the sociaÌ groups formed by most other species of primares in two latteI stages of their pregnancies, Figure 5.19 Average basal metabolism is
wt25o/o more calories than usual; affected by body size. The dashed line rep-
important ways. First, female chimpanzees usually disperse from their natal groups when
resents a direct linear relationship between
they reach sexual maturity, while males remain in their natal groups throughout their lives. and during lactation, about 50% more calories than usual
body weight and basal metabolic rate. The
Second, the members of chimpanzee communities are rarely found together in a unified
sol¡d line represents the actual relationsh¡p
group. Instead, they split up into smallel parties that vary in size and composition from between body weight and basal metabolic
day to day. In chimpanzees, the strongest social bonds among adults are formed among rate. The fact that the curve "bends" means
males, while bonobo females form stronger bonds with one another and with their adult that larger animals use relatively less energy
sons than males do. Chimpanzees modify natural objects for use as tools in the wild. At per unit of body weight.
several sites, chimpanzees strip twigs and poke them into termite mounds and ant nests to The food that primates eat provides them with energy and essential nutrients,
Pro-
extract insects, a much-prized delicacy. In the Taï Forest, chimpanzees crack hard-shelled such as amino acids and minerals, that they cannot synthesize themselves.
reins are essential for virtually every aspect ofgrowth and reproduction
and fol the regula-
nuts using one stone as a hammer and a heavy, flat stone or a protruding root as an anvil. At
of long chains
Gombe, chimpanzees wad leaves in their mouths and then dip these "sponges" into crevices tion of many body functions. As we saw in chapter 2, f roteins are composed
molecules, so in order
to soak up water. New data also reveal tool use by wild orangutans, but chimpanzee tool of amino acids. Primates cannot synthesize amino acids from símpler
sufûcient amounts of a
use is more dive¡se and better studied. to build many essential proteins, they must ingest foods that contain
for animals and pro-
¡umber of amino acids. Fats and oils are impo¡tant sources of energy
miner-
vide about twice as much energy as equivalent volumes of carbohydrates' Vitamins'
many of the
ais, and rrace amount, of .".tni.r eiements play an essential role in regulating
Primate Ecology body's metabolic functions. Although specific vitamíns, minerals, and trace
,r."á"d in only small amounts, deficiencies of these nutrients can cause significant
elements are
impair-
Much of the day-to-day life ofprimates is driven by two concerns: getting enough to eat and ment of normal body function. For example, trace amounts of the elements
iron and copper
avoiding being eaten. Food is essentiai for growth, survival, and reproduction, and it should are important in the synthesis of hemoglobin, vitamin D is essential
for the metabolism of
of body flu-
not be surprising that primates spend much of every day finding, processing, consuming, calcium and phosphorus, and soclium regulates the quantity and distribution
them from the
and digesting a wide variety of foods (Figure 5.18). At the same time, primates musr always ids. primates cannot synthesize any of these compounds and must acquire
most plants'
foods they eat. Watef is the major constituent of the bodies of all animals
guard against predators like 1ions, pythons, and eagles that hunt them by day, and leopards and
moderate
For survival, most animals must balance their watel intake with their water
that stalk them by night. As we will see in the chapters that follow, both the distribution of loss;
food and the threat ofpredation influence the extent of sociality amongprimates and shape dehydration can be debilitaring, and significant dehydration can be fatal.
must
the patterning of social interactions within and between primare groups. At the same time that primates attempt to obtain nourishment from food, they
harmful to them'
Below, we describe the basic features of primate ecology. Later we will draw on this also take care to avoid toxins, substances in the environment that are
information to explore the relationships among ecological factors, social organization, and produce toxins called secondary compounds to protect themselves from
Many plants
primate behavior'. It is important to understand the nature of these relationships because the b"i,ri.nt"n. Thorr.",-rd. of these secondary compounds have been identified: caffeine and
same ecological factors are likely to have influenced the social organization and behavior compounds most familiar to us some secondary com-
-o.f,hi.r" are among the secondary pass through the stomach
of our earliest ancestors. porrrrdr, such as alkaloids, are toxic to consumers because they
into ,ra.ious types of ceils, where they disrupt normal metabolic functions. common
when you eat recl
alkaloids inciude capsicum (the compound that brings tears to yoLrr eyes
'li i'ial | ¡i.;;ii'rårlliì;lr¡ r¡i J.:'i:¡c-¡rJ peppers) and chocolate. other secondary compounds, such as tannins
(the bitter-tasting
in consumer's gut to reduce the digestibility of plant material'
.o-po.rr]d in tea), acr rhe
I jr,¡¡iì i,i::\/jrii". r.iri;ìli)/ii¡;¡i ir, ls;:'r¡til i;ìi itli ¡1rr:rw{h, t,¡r¡vii¿i,¡l iiIil rc¡rtotirtci.irttt, and are often
S".o.rdury compounds are particularly common among tropical plant species
concentrated in mature leãves ar-rd seeds. Young leaves, fruit, and
flowers tend to have
Fiture 5.18 A female baboon Like ail other animals, primates need energy to maintain normal metabolic processes; to
more palatable to
feeds on corms in Amboseli, legulate essential body functions; and to sustain growth, development, and reproduction. lower concentrations of secondary compounds, making them relatively
Kenya. The total amount of energy that an animal requires depends on four components: primates.

PR]MATE ECOLOGY 119

118 CHAPTER 5 Primate Diversity and Ecoloqy
 water are particularly important for arboreal &q
o
animals that do not descend from the canopy o^
o-J
Protein Carbohydrates Fats and Oils Vitamins Minerals Water of tree branches and for terrestrial animals
oz
during times of the year when surface water o-
Animals (x) X -ol
is scarce. Vitamins, minerals, and trace ele- E
Fru¡t X x ments are obtained in small quantities from f
z kg 10 kg 100 kg
Seeds many different sources.
lnsects and gum
Although primates display considerable
Flowers X .8q
diversity in their diet, some generalizations o
o^
Young leaves X X o-J
are possible: Ø
Mature leaves (x) o¿
t. All primates rely on at least one type of o-
stems x
Woody
food that is high in protein and another -ol
E
Sapxxx that is high in carbohydrates. Strepsir- f
z 1 kg '10 kg 100 kg
Gum x (x) x rhines generally obtain protein from
Fruit and some insects
insects and carbohydrates from gum
Underground .Eo
parts x and fruit. Monkeys and apes usually '
x x 8s
o-,
obtain protein from insects or young ø4
sources of nutrients for primates. (x) indicates that the nutrient content ¡s generally leaves and carbohydrates from fruit. b3
@¿
2. Most primates rely more heavily on
accessible only to an¡mals that have specific digestive adaptations.
some types of foods than on others. fr
l
z
Chimpanzees, for example, feed mainly 100 kg
on ripe fruit throughout their range
from Tanzania to Ivory Coast. Scien-. .Eo
I Primates obtain llutrients fronr rnany different sources. tists use the terms frugivore, folivore, oc
o,
ø4
Primates obtain energy and essential nurients from a variety of sources (Table 5.4). insectivore, and gummivore to refer
OJ
carbohydrates are obtained mainly from the simple sugars in fruit, but animal prey, such to primates who rely most heavily on õ2
as insects, also provide a good source offats and oils. Gum, a substance that plants produce fruit, leaves, insects, and plant gum,
respectively. Closer Look 5.1 examines
fr
z
in response to physical injury, is an important source of carbohydrates for some primates, 1 kg 10 kg 100 kg
particularly galagos, marmosets, and tamarins. Primates get most of their protein from some of the adaptations in morphology
Leaves, seeds, and herbs
insect prey or from young leaves. Some species have special adaptations that facilitate the among primates with different diets.
3. In general, insectivores are smaller than Fi(ure 5.21 Body size and diet are related among pr¡mates. The smallest spe-
breakdown of cellulose, enabling them to digest more of the protein conrained in the cells
frugivores, and frugivores are smaller cies eat mainly insects and gum; the largest species eat leaves, seeds, and
of mature leaves. Although seeds provide a good source of vitamins, fats, and oils, many herbs. Fruit-eating species fall in between.
plants package their seeds in husks or pods that shield their contenrs from seed predators. than folivores (Figure 5.21). These differ-
Many primates drink daily from streams, water holes, springs, or puddles of rainwater ences in size are related to differences in
(Figure 5.20). Primates can also obtain'À/ater from fruit, flowers, young leaves, animal pre¡ energy requirements; small animals have relatively higher energy requirements than
and the underground storage parts (roots and tubers) ofvarious plants. These sources of larger animals do, and they require relatively small amounts of high-quality foods
rhat can be processed quickly. Larger animals are less constrained by the quality of
their food than by the quantity because they can afford to process lower-quality foods
more slowly.

The nature of dietary specializations and the challenge of foraging in tropical

I
I forests influence ranging patterns.
Nonhuman primates do not have the luxury of shopping in supermarkets, where abun-
dant supplies offood are concentrated in a single location and are constantly replenished.
Instead, the availability of their preferred foods varies widely in space and time, making
their food sources patchy and often unpredictable. Most primate species live in tropical for-
ests. Although such forests, with their dense greenery, seem to provide abundant supplies
offood for primates, appearances can be deceiving. Tropical forests contain a very large
number of tree species, and individual trees of any particular species are few in number.
Primares with different dietary specializations confront different foraging challenges
(Figure 5.22). Plants generally produce more leaves than flowers or fruit, and they bear
Fiture 5.2O These savanna leaves for a longer period during the year than they bear flowers and fruit. As a result,
baboons are drinking from a foliage is normally more abundant than fruit or flowers at a given time during the year,
pool of rainwater. lvlost pri- and mature leaves are more abundant than young leaves. Insects and other suitable prey
mates must drink every day. animals occur at even lower densities than piants. This means that folivores can generally

t20 CHAPTER 5 Primate Díversity and Ecology PRIMATE ECOLOGY

 Brown
capuchin

White-fronted
capuchin

Squirrel
monkey

Emperor
tamarin

Saddleback
tamarin
F¡guro 5,23 The amount of
Dusky titi time that animals devote to var-
monkey ious types of activities is called
a "time budget." Time budgets
of different species vary con-
Percent time
siderably. These six monkey

r [".'åï"' E [:io "' Rest species all live in a tropical raln

forest in Manú National Park
lTravel Other feeding Miscellaneous in Peru.

(e) (r)
Fi$ure 5.22 (a) Some primates feed mainly on leaves, though many leaves contain toxic
secondary plant compounds. The monkeys shown here are red colobus monkeys in the
Kibale Forest of Uganda. (b) Some primates include a variety of insects and other animal
prey in their diet. This capuchin monkey in costa Rica is feeding on a wasp nest. (c) Moun-
tain gorillas are mainly vegetarians. They consume vast quantities of plant material, like this
fibrous stem. (d) This vervet monkey is feeding on grass stems. (e) Although many primates
feed mainly on one type of food, such as leaves or fruit, no primate rel¡es exclusively on one
type of food. For example, the main bulk of the muriqui diet comes from fruit, but muriquis
also eat leaves, as shown here. (f) Langurs are folivores. Here, hanuman langurs in Ramna-
gar, Nepal, forage for water plants. fPhotographs courtesy of Lynne lsbell (a), Susan perry (b),
John Mitani (c), Carlão Limeira (e), and Carola Borries (f).1
Activity Patterns
I Primate activity patterns show re$ularity in seasonal and daily cycles.
Primates spend the majority of their time feeding, mgving around their home ranges, and
resting (Figure 5.23). Relatively small portions of each day are spent grooming, playing,
fighting, or maring (Figure 5 ,24). The proportion of time devoted to various activities is
influenced to some exrent by ecological conditions. For primates living in seasonal habi-
tats, for example, the dry season is often a time of scarce resources, and it is harder to
find adequate amounts of appropriate types of food. In some cases, this means that the
proportion of time spent feeding and traveling increases during the dry season, while
the proportion of time spent resting decreases.
Primate acrivity also shows regular patterns over the course of the day. When primates
wake up, their stomachs are empty, so the first order of the day is to visit a feeding site.
Much of the morning is spent eating and moving between feeding sites. As the sun moves
directly overhead and the temperature rises, most species settle down in a shady spot to
rest, socialize, and digest their morning meals. Later in the afternoon they resume feed-
ing. Before dusk they move to the night's sleeping site; some species sleep in the same trees
every night; others have multiple sleeping sites within their ranges.
Ranginq Behavior
All prirnatcs lrave hotne rarlges) btlt only sonre s¡:ecies are territorial-
clefenclirrg theil hotne r¿ìnge aga¡11st ittcursiolrs by other lnetnbers of their
specles.

find more food in a given area than frugivores or insectivores can. However, the high con-
centration of toxic secondary compounds in mature leaves complicates the foraging strate-
gies of folivores. Some leaves must be avoided altogether, and others can be earen only in
small quantities. Nonetheless, the food supplies of folivorous species are generally more
uniform and predictable in space and time than the food supplies of frugivores or insecti-
vores. Thus, it is not surprising to find that folivores generally have smaller home ranges
than frugivores or insectivores.
In all primate species, groups range over a relativeìy fixed area, and members of a given
group can be consistently found in a particular area over time. These areas are called home
ranges, and they contain all of the resources that group members exploit in feeding, rest-
ing, and sleeping. However, the extent of overlap among adjacent home ranges and the
nature of interactions with members of neighboring groups or strangers vary consider-
abìy among species. Some primate species, like gibbons, maintain exclusive access to fixed
areas, called territories. Territory residents regularly advertise their presence by vocalizing,
and they aggressively protect the boundaries of their territories from encroachment by

CI{APTER 5 Primate Diversity and Ecology PRIMATE ECOLOGY

 resources from conspecifics. Costs and benefrts are measured in terms of the impact on
the individual's ability to survive and reproduce successfully. Territoriality is beneficial
because it prevents outsiders from exploiting the limited resources within a territory. At the
same time, however, territoriality is costly because the residents must be constantly vigi-
lant against intruders, regularly advertise their presence, and be prepared to defend their
ranges against encroachment. Territoriality is expected to occur only when the benefits of
il maintaining exclusive access to a particular piece of land outweigh the costs of protecting
these benefits.
When will the benefirs of territoriality exceed the costs? The answer to this question
""*tü depends in part on the kinds ofresources individuals need to survive and reproduce suc-
cessfully, and in part on the \May these resources are distributed spatially and seasonally'
For reasons we will discuss more fully in Chapter 6, the reproductive strategies of mam-
ü
a malian males and females generally differ. In most cases, female reproductive success
depends mainly on getting enough to eat for themselves and their dependent offspring,
and males' reproductive success depends mainly on their ability to mate with females.
As a consequence, females are more conøerned about access to food, and males are more
interested in access to females. Thus, territoriality has two different functions. Some-
times females defend food resources, or males defend food resources on their behalf.
Other times, males defend groups of females against incursions by other males. In pri-
mates, both resource defense and mate defense seem to have influenced the evolution
of territoriality.

Fiture 5.24 (a) All diurnal primates, like this capuchin

monkey, spend some part of each day resting. (b) lmmature
monkeys spend much of their "free" time playing. These
patas monkeys are play-wrestling. (c) Gorillas often rest in
close proximity to other group members and socialize dur-
ing a midday rest period. [Photographs courtesy of Susan
Perry (c) and Lynne lsbell (b).1
(c)

outsiders (Figure 5.25). Although some territorial birds defend only their nest sites, primate
territories contain all ofthe sites atwhich the residents feed, rest, and sleep and the areas
in which they travel. Thus, among territorial primates, the boundaries for the rerrirory are
essentially the same as for their home range, and territories do
not overlap.
Nonterritorial species, like squirrel monkeys and long-tailed
macaques, establish home ranges that overlap considerably with
those of neighboring groups (Figure 5.26). When members of
neighboring nonterrirorial groups meer, rhey may fight, avoid
one another, or mingle peacefully together. This last oprion
Fiture 5.26 Home range
overlap of caPuchin monkey
is unusual, but in some species, adult females sexually solicit groups on Barro Colorado
males from other groups, males attempt to mate with females lsland, Panama. Sites of inter-
from other groups, andjuveniles from neighboring groups play group encounters are marked
together when their groups are in proximity. with a dot. (Figure 3 in Crofoot,
M.C., Gilby, 1., Wikelski, M.C.,
The two main functions suggested for territoriality are & Kays, R.W., et al. 2008.)
II resource defense and mate defense. lnteraction location outweighs
the competitive advantage of
To understand why some primate species defend their home numerical superiority in Cebus
Fl¡lure 5.25 Gibbons perform complex vocal duets as part of ranges from intruders and others do not, we need to think capucinus intergroup contests.
territorial defense. about the possible costs and benefits associared with defending (PNAS 105(2): 577-581-.)

124 CTIAPTER 5 Primate Diversity and Ecology PRIMATE ECOLOGY L25

 (a) (c)

Filure 5.28 ln some cases, researchers are able to confirm predation. Here, an adult
female baboon in the Okavango Delta, Botswana, was killed by a leopard. You can see (a)
the depression in the sand that was made when the leopard dragged the female's body out
of the sleeping tree and across a small sandy clearing, (b) the leopard's footprints beside
the drag marks, and (c) the remains of the female the following morning-her jaw, bits of her
skull, and clumps of hair.

and chimpanzees in the Tai Forest (Figure 5.29). In general, terrestrial species are more
Fi(ure 5.27 Primates are preyed upon by a
variety of predators, including the python (a), vulnerable than arboreal species, and species that live in small groups are more vulnerable
lion (b), leopard (c), crowned hawk eagle (d), than animals that live in large groups. Thus, arboreal monkeys that live in large groups face
and crocodile (e). the lowest risks. Schultz and her colieagues
(d) suggest that these results may explain some
aspects of the disribution of terrestrial pri- o.25
mates in Africa, Asia, and the neotropics. In
Africa, with crowned hawk eagles and at least
Predation two large predatory felids, terrestial primates o.20
Predation is believed to be a significant source of mortality among primates, are large-bodied or live in large groups. In
I
I but direct evidence of predation is difficult to obtain. Asia, where there are no large forest raptors
and few large felids, there are several semi- o
Primates are hunted by a wide range of predators, including pythons, raptors, crocodiles, 0.'15
E
terrestrial macaque species. And in the neo- c
leopards, lions, tigers, and humans (Figure 5.27). In Madagascar, large lemurs are preyed o
tropics, where there are several species of .F
upon by fossas, pumalike carnivores. Primates are also preyed on by other primates. Chim- (ú
rco
large felids and forest raptors, there are no
panzees, for example, hunt red colobus monkeys, and baboons prey on vervet monkeys. 0.10
terrestrial monkeys at all. fL
The estimated rates of predation vary from less than l% of the population per year ro
more than l5o/o.The available data suggest that small-bodied primates are more vulnerable
to predation than larger ones and that immature primates are generally more susceptible to
I Primates have evolved an arraY o1

predation than adults. These data are not very solid, however, because systematic informa-
I d"funr", against predators. 0.5

tion about predation is quite hard to come by since most predators avoid close contact with Many primates give alarm calls when
humans, and some predators, like leopards, generally hunt at night, when most researchers they sight potential predators, and some spe-

.e*" .
are asleep. Usually predation is inferred when a healthy animal that is unlikely to have left cies have specific vocalizations for particular
the group abruptly vanishes without a trace (Figure 5.28). Such inferences are, ofcourse, predators. Vervet monkeys, for example, give
aUo
subject to error. different calls when they are alerted to the ,Ø

Another approach is to study the predators, not their prey. Crowned hawk eagles are the presence ofleopards, small carnivores, eagles, o\\: a*c""".".'%Ì:-"
only large raptors that live in the tropical rain forests of Africa. They are formidable preda- snakes, baboons, and unfamiliar humans. In
tors; although they weigh only 3 to akg(6.6 ro S.8 lb), they have powerful legs and large many species, the most common resPonse to .^ó*tst' ñro$ô'
talons and can take prey that weigh up to 20 kg (44 lb). Crowned hawk eagles carry prey predators is to flee or take cover. Small pri- ,uøq
o'óo-
back to their nests and discard the bones. By sorting through the remains under crowned mates sometimes try to conceal themselves _$ø-
hawk eagle nests, researchers can figure out what they eat. Analyses of nest remains in the from predators; larger ones may confront Fi(ure 5.29 The rate of predation by leopards (red), eagles (tan), and chimpan-
Kibale Forest of Uganda and the Taï Forest in Ivory Coast indicate that crowned hawk eagles potential predators. When slow-moving pot- zees (blue) in the Taï Forest on different primate species is shown here. Note that
prey on all of the primates in these forests except chimpanzees. Monkeys make up 60Vo to tos encounter snakes, for example, they fall the preferred prey of chimpanzees are red colobus monkeys, and chimpanzees'
80% of the crowned hawk eagles' diets at these sites, and the eagles kill a sizable fraction to the ground, move a short distance, and only predators are leopards. (From S. Schultz, R. Noê, W. S. MacGraw, and R. l.
(zYo to 16%) of the total populations of various primate species in these forests each year. freeze. At some sites, adult red colobus mon- M. Dunbar, 2OO4, "A Commun¡ty-Level Evaluation of the lmpact of Prey Behav-
Susanne Schultz of the University of Liverpool and her colleagues have com- keys aggressively attack chimpanzees who ioural and Ecological Characteristics on Predator Diet Composition," Proceedíngs
pared the characteristics of mammalian prey taken by crowned hawk eagles, leopards, stalk their infants. of the Royal Society of London, Section B 27!:725-732.)

CHAPTER 5 Pr¿mate Díversity and Ecology PRIMATE ECOLOGY 127

 Another antipredator strategy that primates adopt is to associate with members of other
primate species. In the Taï Forest of Ivory Coast, a number of monkey species share the
canopy and form regular associations with one another. For example, groups ofred colobus
monkeys spend approximately half their time with groups of Diana monkeys. Interspecific
associations may enhance predator detection if each species occupies a different portion
of the canopy and is oriented toward different predators. In addition, by associating with ost primâtes live in groups.
members of different species, monkeys may increase group size without increasing levels A group is a social unit that
of competition from conspecifics who have similar dietary preferences. is composed of animals
that share a common home range or
territory and interact more with one
another than with other members of lew platyrrhine monkeys and a few Multiple males, multiple females:
Primate Sociality their species. Groups can vary in their strepsirrhines. In some pairJiving Groups are composed of a number
SociirlitV tlir:; evolvctl in prirni.rtr:s irl ro:;¡lotrsc tu <:t;olr^r¿jicerl ¡rtr-.:;l;rtrr,;1;. size, age-sex composition, and degree species, males and females remain of adult females, a number of adult
I close together, but in others, they males, and immatures. This form
I lifc lr¿¡s both r:ogtr; arrrl Irorrcfii.q.
l-:ioci;rl of cohesiveness. We use the term social
organization to describe variation may travel independently within of social organization is character-
Nearly all primates live in social groups of one kind or another. Sociality has evolved in their territories much of the time. istic of macaques, baboons, caPu-
along these dimensions, Thàre are five
primates because thele are important benefits associated with living in groups. Primates chin monkeys, squirrel monkeys,
basic types of social systems among pri-
that live in groups may be better able to acquire and control resources. Animais that live Multiple males, one female: One
mates (Figure 5.30): and some colobines. Some sPecies,
in groups can chase away lone individuaÌs from feeding trees and can prorect their own adult female shares a territory or
such as chimpanzees and spider
access to food and other resources against smaller numbers of intruders. As we saw earlie r, Solitary: Females maintain sepa- home range with more than one
monkeys, that live in these kinds of
grouping also provides safety from predators. This is because ofthe three Ds: detection, rate home ranges or territories and adult male and offspring. This form
groups often divide up into smaller
deterrence, and dilution. Animals in groups are more likely to detect predators because associate mainly with their depen- of social organízation is only found
temporary parties (fission-fusion
there are more pairs of eyes on the lookout for predators. Animals in groups are also more dent offspring. Males establish their in marmosets and tamarins.
groups).
effective in deterring predarors by actively mobbing or chasing rhem away. Finally, the own territories or home ranges, One male, multiple females:
threat of predation to any single individual is diluted when predators srrike at random. If which may encompass the ranges Primates also vary in their mating
Groups are composed of a number
there are two animals in a group, and a predator strikes, each animal has a 50% chance of of one or more adult females. All of systems, the pattern of mating activ-
of adult females, one resident adult
being eaten. Ifthere are 10 individuals, the individual risk is decreased to 102o. the solitary primates are strePsir- ity and reproductive outcomes. There
male, and immature offspring. is a close, but not perfect, relationship
Although there are important benefits associated with sociality, there are equally impor- rhines, except for orangutans.
Males compete vigorously over res-
tant costs. Animals that live in groups may encounter more competition over access to food between social organization and mat-
Pairs: Groups âre composed of idence in these kinds ofgroups, and
ing systems. There are four main forms
and mates, become more vulnerable to disease, and face various hazards from conspecifics
one adult female, one adult female, males may band together to oust
(such as cannibalism, cuckoldry, inbreeding, or infanticide). of mating systems in primates.
and immature offspring. Species established residents. This form of
The size and composition of the groups that we see in nature are expected to reflect a Monogamy/pair-bonding: In a
that live in pairs usually defend social organization is characteristic
compromise between the costs and benefits of sociality for individuals. The magnitude of of howler monkeys, some langurs, strictly monogamous mating sYs-
the boundaries of their territories.
these costs and benefits is influenced by both social and ecological factors. tem, each male and female mates
Gibbons live in pairs, along with a and gelada baboons.

! f 'rirrt;rtrrlo¡t,i';tr; r¡t'r: riirridr:rl 0vr:f ttrhr.)tlrcr '[uor.iin¡1 ccltt-i¡lil1.ilirtn ot ìtroclaiirjtì ¡s

fl tl',, ¡ri'inritry l,ri ir¡; l;trru¡'it'rl.l r;{l(:i;Ìlii}/ ,..ìtrì(}n¡,j l.¡tinli.li(r:.j.
It is not entirely clear whether resource competition or predation was the primary factor
favoring the evolution of sociality in primates. However, many primatologists are convinced
I
that the nature of resource competition affects the behavioral strategies of primates, particu-
larly females, and influences the composition of primate groups (Closer Look 5.2). Females
come first in this scenario because their fitness depends mainly on their nutritional status: Solitary Polygyny:
one-male
well-nourished females grow faster, mature earlier, and have higher fertility rares than do
poorly nourished females. In contrast, males' fitness depends primarily on their ability to
obtain access to fertile females, not on their nutritional status. Thus, ecological pressures infln-
ence the distribution of females, and males distribute themselves to maximize their access to
( I 9ç
females. (We will discuss male and female reproductive strategies more fully in Chaprer 6.) gt 9o Figure 5.30 The major types of social
groups that primates form. When males

Pair-bonded (( 9o ç ?
and females share their home ranges,
their home ranges are drawn here in
Primate Conservation brown. When the ranges of the two
sexes differ, male home ranges are
Polygyny:
i lVllrtry:,Ì){l{)i(:r, lrf ¡ilitr-r;.rir::,;rtr: iii ir:;il rl;tt¡/ir:t ri'[l,xi!i¡r:ií¡.¡¡t itr iil¡'willl. multimale
drawn in blue and female home ranges
are drawn in red. The sizes of the male
Sadly, no introduction to rhe primate order would be complete without noting that and female symbols reflect the degree
the prospects for the continued survival of many primate species are grim. According of sexual dimorphism among males
to the International Union for the Conservation of Nature (IUCN), which assesses the Polyandry and females.

124 CHAPTER 5 Prímate Diversity and Ecology 129

with only one member of the oppo-
site sex. Most primates that live in
pairs mate mainly with each other,
but there are reports of extra-pair
matings in a number of pairJiving
primate species, and extra-pair
paternity has been confirmed in
at least one pair-living primate,
the fork-marked lemur. Thus, the
term pair-bonding may be a more
accurate description of the mating
system of most pair-living primates
than monogamy.
males in the group, but the limited species that live in multimale, multi-
available genetic data suggest that female groups and might also char-
not all males are equally successful acterize some solitary species. In
in fathering offspring. most species that form multimale,
multifemale groups males compete
Polygyny: Males mate with multi over access to mating females, and
ple females, but each female mates there is considerable skew in male
with a single male. This mating I Extinct Nearthreat
reproductive success. Fi¡Jure 5.31 Conservation
system characterizes most of the
species that live in one-male, multi- These classifications of social orga-
r 3;[:iJ:,."6 I Least concern
status of primate populations
female groups. This mating sysrem nizatio¡ and mating systems represent I rndangered f_l No data in the wild. Based on data from
the IUCN Red List of Threat-
generates considerable skew in ídealized descriptions of residence and
mating patterns. The reality is inevita-
I Vulnerable ened Species,2OIO.
male reproductive success, as resi-

Polyandry: Females mate with

dent males largely control access bly more complicated. Not all groups of É'e ,"f.d Þà"
to receptive females. However, in a particular species may have the same
multiple males, but each of the some of these species, including social organization or mating system. S..i""

males mates with only one female. blue monkeys, males from outside For example, some groups of tarsiers
Polyandry is an uncommon mating the group sometimes enter groups are composed of a single mated pair,
system among mammals but may while others include additional females. f'hc nl¿rin tl llc;¡i:; lo rlrilrr¡lic:; irr thc wiltl ;rrt: (:1,) lr¿li¡iti¡t tlr.:strttctiott,
and mate with females. I
characteúze some of the marmo- Hamadryas and gelada baboons form ü (2) lriilltirì,1Í" (J) (li:;()¿rsìo. (4) atrd livt: t;it¡ltitto lur tr¿rclt:.ìlì(l cxl)ort.
sets and tamarins. In these species, Poþynandry (promiscuity): Both one-male, multifemale units, but sev-
As arboreal resiclents of the tropics, most primate populations are directly affected by
one female usually monopolizes males and females mate with more eral of these units collectively belong
the rapid and widespread destruction of the world's forests. Colin Chapman of the Uni-
reproduction. The breeding female than one partner. This mating sys- to larger aggregations.
versity of Florida and Carlos Peres of the University'of East Anglia, recentiy reviewed the
may matê with all of the unrelated tem is generally associated with
conservation status of the world's primate populations. Their analysis is quite sobering.
Between 1980 and 1995, approximately l\Vo of the forests in Africa and Latin America were
lost, and 6Vo of the forests of Asia disappeared (Figure 5.33).
The destruction of tropical forests is the product of economic and demographic pres-
sures acting on governments and local residents. Many developing countries have huge
foreign debts that must be repaid, The need to raise funds to pay offthese debts generates
conservation status of plant and animal species around the globe, nearly half of all primate intense pressure for timber harvesting and more intensive agricultural activity. Each year,
species are nolv threatened in the wild (Figure 5.31). In Asia,54o/o of all primate species are 5 million to 6 million hectares of forest are logged, seriously disrupting the lives of the ani-
at risk of extinction. On the island of Madgascar, the conservation status of many species is mals that live in them. (A hectare is a square measuring 100 m on a side, or about 2.5 acres.)
not known, but ofthose that can be assessed, halfare endangered or critically endangered Forests are also cleared for agricultural activities. Rapid increases in the population
and one has already become extinct. A smaller proportion of species in Africa and Central of underdeveloped counries in the tropics have created intense demand for additional
and South America are in threatened categories, but the prospects are not encouraging. agricultural land. In West Africa, Asia, and South America, for example, vast expanses of
All around the world, the populations of nearly all primate species are decreasing, some forests have been cleared to accommodate the demands of subsistence farmers trying to
very rapidly. feed their families, as well as the needs of large-scale agricultural projects. In Central and
All of the great apes are now endangered. In West Africa, chimpanzee populations have South America, massive areas have been cleared for large cattie ranches.
been decimated over the last two decades. In 1990, Ivory coast was home to 8,000-12,000 In the last few decades, a new threat to the forests of the world has emerged: wildfire.
chimpanzees , By 2OO7, that number had declined by 9\o/o, according to a census conducted Major fires destroyed massive tracts of forest in Southeast Asia and South America. Ecolo-
by Christophe Boesch andhis colleagues. The decline is attributed toas}yoincrease in the gists believe that natural fires in tropical forests are relatively rare and that these devastat-
size of the human population, which created more poaching and habitat destruction and ing fires are the product of human activity. In Indonesia, massive fires in the late 1990s left
was exacerbated by civil unrest within the country. There are only 6,600 orangutans left in thousands of orangutans dead, reducing their numbers by nearly a third.
Sumatra, where forests are being logged or converted to oil palm plantations. In many areas around the world, parricularly South America and Africa, primates are
It is particularly disturbing that some of the most endangered primare species ale ones also hunted for meat. Although systematic information about the impact of hunting on
that we know the least about. For example, in 2005 researchers encountered a previously wild primate populations is scant, some case studies reveal troubling trends. In one forest
unknown type of monkey in the highlands of Tanzania(Figure 5.32). Generic data indicare in Kenya, for example, 1,200 blue monkeys and nearly 700 baboons were killed by subsis- Fiture 5.32 The kapunji mon-
that these monkeys are sufficiently different from other species to be placed in their own rence hunters in one year. In the Brazilian Amazon, one family of rubber tappers killed ZOO key was first discovered by sci-
genus, Rangwecebus, and are most closely related to baboons. This species is now restricted woolly monkeys, 100 spider monkeys, and 80 howler monkeys during an 18-month span. entists in 2005. (Photo by Tim
to two small areas of evergreen forest, 350 km (230 miles) apart, and the total population is In addition to subsistence hunting, there is also an active market for "bushmeat': in many Davenport, Wildlife Conserva-
estimated to be only 1,100 individuals. urban areas. tion Society.)

CHAPTER 5 Pr¡mate Diversity and Ecology PRlMATE CONSERVATION T3T


Fiture5.33 Deforestation
in Borneo. Tropical forests
are disappearing all over the
world, threatening the continu-
ing survival of primates and
other animals that live in them.
ln Borneo, deforestation has
greatly reduced the availabilitY
of su¡table hab¡tats for orang-
utans and other primates.
(Source: www.rainforestsos
.or g / ab o uT- r ainf o re
whatshappen ing-to-them)
sts/
In equatorial Africa, primate populations have also been decimated by outbreaks of epi-
demic disease. Twenty-six percent of one habituated group in the TaI Forest of Ivory Coast
died of the Ebola virus during â one-month period. Later, anthrax killed more members of
the same community. Several hundred gorilias, belonging to more than 100 groups, regu-
larly foraged in a swampy clearing in Odzala-Kokoua National Park in the Congo. Over the
course ofa two-year period, 957o ofthese gorillas died from Ebola.
The capture and trade of live primates has been greatly reduced since the Convention
on International Trade in Endangered Species ofWild Fauna and Flora (CITES) was drafted
in 1973. The parties to CITES, which now number 167 countries, ban commercial trade of
all endangered species and monitor the trade of those that are at risk of becoming endan-
gered. CITES has been an effective weapon in protecting primate populations around the
world. The United States imported more than 100,000 primates each year before ratifying
CITES but had reduced this number to approximately 13,000 a decade after signing the
international agreement.
Although CITES has made a major impact, some problems persist. Live capture for
trade remains a major threat to certain species, particularly the great apes, whose high com-
mercial value creates strong incentives for illegal commerce. In many communities, young
primates are kept as pets. For each animal taken into captivity, many other animals are put
at risk because hunters cannot obtain young primates without capturing their mothers,
who are usually killed in the process. In addition, many prospective pets die after capture
from injuries suffered during capture and transport or from poor housing conditions and
inappropriate diets while in captivity.
I Efforts to save endangered primate populations have met with some success.
Although much remains to be done, conservation efforts have significantly improved
the survival prospects of a number of primate species. These efforts have helped preserve
muriquis and golden lion tamarins inBraziland golden bamboo lemurs in Madagascar. But
there is no room for complacency. Promising efforts to save orângutans in Indonesia and
mountain gorillas in Rwanda have been severely impeded by regional political struggles and
armed conflict, putting the apes' habitats and their lives in serious jeopardy. A number of
different strategies to conserve forest habitats and preserve animal populations are on the
table. These include land-for-debt swaps in which foreign debts are forgiven in exchange for
commitments to conserve natural habitats, to develop ecotourism projects, and to promote
sustainable development offorest resources. But as conservationists study these solutions
and try to implement them, the problems facing the world's primates become more pressing.
More and more forests disappear each year, and many primates are lost, perhaps forever.
CFIAPTER 5 Primate Dtverstty and Ecology
IÚEEI' HELP STUDTITTG?
wwnorton.com/studyspace
Visit StudySpace to access free review materials such as:
r Vocabulary Flashcards
r Diagnostic Review Quizzes
r Study Outlines
Further Reading
Boinski, S., and P. A. Garber, eds. 2000. On the Mot¡e: How and
W'lty Animals Travel in Groups. Chicago: University of Chi-
cago Press.
Campell, C.J., A.Fuentes, K. Cl MacKinngn, M. Panger, and
S. K. Bearder, eds. 2007. Primates in Perspective. New York:
Oxford University Press.
Cowlishaw G., and R. I. M. Dunbar. 2000. Primtte Consetvø-
tionBiology. Chicago: University of Chicago Press.
Hohmann, G., M. M. Robbins, and C. Boesch, eds. 2006.
FeedingEcology in Apes and Othet Ptimøtes Ecological, Plrysi-
cal, øndBehøvioral Aspect.t. New York: Cambridge Univer-
sity Press.
IUCN 2010. IUCNRed List ofThreatened Species. Version 201'0.3.
www.iucnredlist.org.
Study Questions
1.What is the difference between homology and analogy?
What evolutionary processes correspond to these terms?
2. Suppose that a group ofextraterrestrial scientists lands
on Earth and enlists your help in identifying animals.
How do you help them recognize members of the pri-
mate order?
3. What kinds of habitats do most primates occupy? What
are the features of this kind of environmentì
4. Large primates often subsist on low-quality food such as
leaves; small primates specialize in high-quality foods
such as fruit and insects. Why is body size associated
with dietary quality in this waY?
5. For folivores, tropical forests seem to provide an abun-
dant and constant supply of food. Why is this not an
accurate assessment?
6. Territorial primates do not have to share access to food,
sleeping sites, mates, and other resources with members
ofother groups, Given that territoriality reduces the
extent of competition over resources, why are not all pri
mates territorial?
Kappeler, P. M., and M. Pereira, eds. 2003. Primate Life Histo-
ries and Socioecology. Chicago: University ofChicago Press.
Kramer, R., C. van Schaik, and.¡. Johnson, eds. 1997. Last
Stand: Prctected Areøs and the Defense of Tropical Biodbersity.
New York: Oxford UniversitY Press.
Lee, P. C., eà. lggg. Comparative Primate Socioecology' New
York: Cambridge University Press.
Martin, R. D.1990. Primøte Origins andEvolution: APhylo'
genetic Reconstruction. Princeton: Princeton University
Press.
Mittermeier, R. et al. 2009. Primates in peril: the world's 25
most endangered primates 2008-2010)' Prímate ConservL-
tion 25tl-57.
Strier, K. B. 2007. Primate Behøviotal Ecology, 3rd ed. Boston:
Allyn & Bacon.
7. Territoriality is often linked to group size, day range, and
diet. What is the nature of the association, and why does
the association occur?
8. Most primates specialize in one type of food, such as
fruit,leaves, or insects. What benefits might such spe-
cializations have? What costs might be associated with
specialization?
9. Nocturnal primates are smaller, more solitary, and rñore
arboreal than diurnal primates. What might be the
reason(s) for this pattern?
10, Sociality is a relatively uncommon feature in nature.
What are the potential advantages and disadvantages of
living in social groups? Why are (virtually all) primates
social?
11. The future of primates, and other occupants of tropical
forests, is precarious. What are the major hazards that
primâtes face?
12. How can we balance the needs and rights of people living
in the developing nations of the tropics with the needs of
the animals who live in tropical forests?
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