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 Contents

■ BOX 3.3 Lucky Lemurs 94

Implications for Primate Behavioral Ecology 96

4 Evolution and Social Behavior 101

Natural Selection 103
Sources of Genetic Variation 104
Variation among Individuals 104
Mutations 104
Mechanisms of Inheritance 105
Sexual Reproduction and Mating Patterns 107
■ BOX 4.1 MHC Genes and Mate Choice 108

Variation within and between Populations 110

Random Genetic Drift 110
Gene Flow and Dispersal 112
Genetic versus Environmental Influences 112
Testing Evolutionary Hypotheses for Behavior 115
Problems with Comparing Fitness among Primates 118
Identifying Optimal Traits 119
Generational Time Lags and Changing Environments 119
■ BOX 4.2 Menopause 120

Evaluating the Role of Ecological Pressures 121

Kin Selection and Reciprocal Altruism 123
Altruism and the Challenge of Group Selection 123
Selfish Benefits of Helping Kin 124
■ BOX 4.3 Multilevel Selection 125

Hamilton’s Rule 126

Evidence for Kin Selection among Primates 128
Evaluating the Evidence for Kin Selection 130
Benefits of Helping Nonkin 131
Conditions for the Evolution of Reciprocity 131
Evidence for Reciprocity among Primates 131
Evaluating the Evidence for Reciprocity 134
Individual Strategies and Social Organizations 137
Conflict and Cooperation among Same-Sexed Individuals 138
Conflicts between the Sexes 140

5 Evolution and Sex 143

Sexual Selection 146
Sexual Dimorphism 149
Phylogenetic Constraints 149
Ecological Constraints 149
vii
Contents

Mating Patterns 150

Mating Patterns When Females Are Solitary 151
The Case of Monogamy 151
■ BOX 5.1 Gibbon Games and Tarsier Tactics 152

An Extreme Form of Polygyny 155

Polyandry 157
Ambivalent Polygamy 158
Mating Patterns When Females Live in Groups 160
Single-Male Female Groups 162
Multi-Male Female Groups 164
Extra-Group Copulations 165
Seasonal versus Aseasonal Breeders 166
The Influence of Males on Females 169
Female Mating Strategies 171
Sperm and Fertilization 171
Food and Safety from Predators 171
Allies against Aggression 172
Parental Investment 172
Good Genes 173
Sexual Signals 174
Female Choice and the Unpredictability of Ovulation 174
Sexual Swellings and the Female Dilemma 175
Male Rank and Reproductive Success 177

6 Food, Foraging, and Females 183

Food Quality and Nutrient Balancing 187
Energy and Nutrients 188
Digestibility and Edibility 190
Physical Deterrents 190
Chemical Deterrents 191
Body Size Energetics and Turnover Rates 192
■ BOX 6.1 Forest Pharmacy 193

Reproductive Energetics 195

The Spatial Distribution of Food 199
Patch Size and Defensibility 200
Effects of Patch Density on Ranging Patterns 202
The Temporal Availability of Food Resources 203
Behavioral Adjustments to Food Seasonality 204
Reproductive Seasonality 207
■ BOX 6.2 The Power of Food 208

viii
 Contents

Interpreting Diets and Their Behavioral Correlates 211

Evaluating “Critical Functions” 211
Effects of Altered Habitats 212

7 Female Strategies 215

Ecology of Female Relationships 217
Types of Relationships 217
Within- and Between-Group Competition 218
Social Dynamics in Female Groups 221
Matrilocal Societies 221
■ BOX 7.1 Mysterious Matrilines and Market Theory 222

Rank Inheritance 225

Relations among Females 227
Relations with Males 229
Age-Related Rank 230
Relations among Females 231
Relations with Males 232
Life without Kin 233
Gaining Group Membership 235
Autonomous Interests 237
Aggregations around Males 237
Avoidance of Males 238
Managing Males 239
Population Consequences of Female Strategies 241
Habitat Fragmentation and Saturation 241
Effects on Philopatric Females 242
Effects on Groups of Unrelated Females 243
Reproductive Implications 244
Manipulating Sex Ratios 245

8 Male Strategies 249

Ecology of Male Relationships 250
Types of Relationships 252
■ BOX 8.1 Using and Misusing Infants 252

Within- and Between-Group Competition 254

Social Dynamics among Males 258
Patrilocal Societies 259
Rank Acquisition and Coalitions 262
Relationships among Males 263
Relationships with Females 263

ix
Contents

Maternal Rank Inheritance 263

Ranks in Age-Graded Groups 263
Relationships among Males 265
Relationships with Females 266
When Males Disperse 267
Hierarchical Relationships 267
Gaining Group Membership 269
■ BOX 8.2 Beyond the Group 271

Unattached Males 272

Males in Pair-Bonded Societies 274
Population Dynamics 275
Genetic and Demographic Correlates 275
Male Life Histories 277

9 Developmental Stages through the Life Span 279

Fertilization to Birth 280
Infancy 281
Maternal Care 284
Paternal Care 285
Alloparental Care 286
■ BOX 9.1 Parental Prolactin 288

Weaning Conflict 291

Juvenile Challenges 294
Staying Alive 294
Age, Size, and Sex 294
Experience 296
Social Skills 297
Long-Term Bonds 298
Sex-Biased Dispersal 299
Puberty 300
Adulthood and Aging 301
■ BOX 9.2 The Legacy of Life Histories 303

Population Consequences of Life Histories 306

Life History Flexibility and Constraints 306
Demography and Conservation 307

10 Communication and Cognition 311

Components of Communication Systems 314
Modes of Primate Communication 316
Tactile Communication 316
Visual Communication 318
x
 Contents

Olfactory and Gustatory Communication 320

Vocal Communication 323
Species Recognition 323
Within- and Between-Group Distinctions 324
Long-Distance Calls 325
Close-Range Calls 326
Ontogeny 327
Intentional or Involuntary Information Sharing 327
Cognition 329
Learning and Imitation 330
Ecological Intelligence 332
Spatial Memory 332
Tool Use 333
Social Intelligence 338
Alliances 338
Tactical Deception 339
Social Traditions 340
Implications for the Ethical Treatment of Captive and Wild Primates 341
■ BOX 10.1 Rehabilitation, Reintroduction, and Sanctuary 342

11 Community Ecology 347

Primate Communities 348
■ BOX 11.1 Ethnoprimatology 348

Niche Divergence 350

Polyspecific Associations 351
Foraging Benefits 355
Predator Protection 356
Predator–Prey Interactions 358
■ BOX 11.2 Predatory Perspectives 358

Primates as Predators 360

Primates as Prey 361
Primates, Parasites, and Microbiota 362
Social Transmission 363
Environmental Disruption 364
Primate–Plant Interactions 364
Pollination 365
Seed Dispersal and Seed Predation 366
Conservation of Communities 369
Specialists, Generalists, and Social Responses 370
Preserving Diversity 370

xi
Contents

12 Conservation 373
Threats to Primates 374
Habitat Disturbances 375
Hunting Pressures 379
Disease 382
Conservation Policies 383
Economic Incentives 383
Increasing Public Awareness 385
Nongovernmental Organizations 386
■ BOX 12.1 The Primates’ People 386

Noninvasive Research 388

Diet and Habitat Change 390
Reproductive Biology and Stress 391
From Paternity to Population Genetics 393
■ BOX 12.2 From Methods to Practice 395

The Next Millennium 396

Appendix 399
Primate Names 399
Prosimians 400
New World Monkeys 405
Old World Monkeys 411
Apes 419
Bibliography 421
Glossary 519
Name Index 533
Subject Index 547

xii
Figures

1.1 Bipedal bonobos (Pan paniscus) at the San Diego Zoo. 2

1.2 Yellow baboons (Papio cynocephalus) at Amboseli National Park,
Kenya. 3
1.3 Two northern muriquis (Brachyteles hypoxanthus) at the Reserva
Particular do Patrimônio Natural (RPPN) Feliciano Miguel Abdala,
Brazil. 6
1.4 Yellow baboon (Papio cynocephalus) mother at Amboseli National
Park, Kenya. 7
1.5 Chimpanzee (Pan troglodytes) mother with infant fishes for termites. 9
1.6 Masudi, an adult male chimpanzee (Pan troglodytes) at Mahale,
Tanzania, and primatologist John Mitani. 10
1.7 Male mantled howler monkey (Alouatta palliata). 11
1.8 Japanese macaque (Macaca fuscata) matriline. 12
1.9 Geographical distribution of baboons. 13
1.10 Distribution of major vegetation zones within the range of
nonhuman primates. 14
1.11 Malabar sacred langurs (Semnopithecus hypoleucos). 14
1.12 A male mountain gorilla (Gorilla berengei berengei). 15
1.13 A subadult male orangutan (Pongo pygmaeus). 15
1.14 Vervet monkeys (Chlorocebus pygerythrus). 16
1.15 Gray-cheeked mangabey (Lophocebus albigena). 18
1.16 Black and white colobus monkeys (Colobus guereza) by the Nile
River in Murchison Falls National Park, Uganda. 19
1.17 Model for optimal primate group sizes based on the compromise
between costs of feeding competition and enhanced safety from
predators. 20
B1.1 Breeding male cotton-top tamarin (Saguinus oedipus) carrying an
infant. 25
1.18 Female-focus view of social evolution. 27
2.1 Allometric relationships between brain and body weights for 309
extant placental mammalian species. 41
2.2 Food characteristics expected from the total and relative nutrient
requirements of large and small animals. 42

xiii
Figures

2.3 Patas monkeys (Erythrocebus patas). 44

2.4 Southern red-necked night monkey, or black-headed or Peruvian
night monkey (Aotus nigriceps). 45
2.5 Northern muriqui (Brachyteles hypoxanthus) mother with a new infant. 46
2.6 Charlie, an adult male chimpanzee (Pan troglodytes) at Gombe. 47
2.7 Young chimpanzee (Pan troglodytes), Freud, grooms the ear of his
mother, Fifi. 48
2.8 Aye-aye (Daubentonia madagascariensis). 48
2.9 Comparison of platyrrhine (represented by the genus Cebus) and
catarrhine (represented by the genus Macaca) teeth. 50
2.10 Abbreviated taxonomy of the Primate Order. 52
2.11 Alternative classifications at the top and bottom of the Primate Order. 53
2.12 Contemporary distribution of nonhuman primate families. 54
2.13 An adult female slender loris (Loris lydekkerianus) at Ayyalur, Tamil
Nadu, South India. 55
2.14 Adult male Verreaux’s sifaka (Propithecus verreauxi) at Beza Mahafoly. 56
2.15 A spectral tarsier (Tarsius spectrum). 57
2.16 Two northern muriquis (Brachyteles hypoxanthus). 58
2.17 Mexican mantled howler monkey (Alouatta palliata) eating. 59
2.18 Adult female white-faced capuchin (Cebus imitator [revised from C.
capucinus]). 59
2.19 Adult female Central American squirrel monkey (Saimiri oerstedii). 60
2.20 Characteristic anatomical features of colobines and cercopithecines. 63
2.21 The Mona monkey (Cercopithecus mona). 64
2.22 Subadult female bonnet macaque (Macaca radiata). 64
2.23 Capped langur, or leaf monkeys (Trachypithecus pileatus). 65
2.24 White-handed (or lar) gibbons (Hylobates lar) suspended. 65
2.25 An adult female orangutan (Pongo pygmaeus). 66
2.26 An adult male orangutan (Pongo pygmaeus) in Wehea Forest, East
Kalimantan, Borneo. 67
2.27 Gorillas and chimpanzees, similar to the bonobo (Pan paniscus)
shown here, walk on their knuckles. 67
2.28 Location of the major chewing muscles. 69
2.29 Relationship between primate home range size (in km2) and
population group weights (kg) for different genera. 70
2.30 Distribution of dispersal regimes among 22 primate species. 71
3.1 The geographical distribution of extant nonhuman primates and
extinct primate species. 76
3.2 Alternative routes for ancestral platyrrhines. 79
B3.1 Two male olive baboons (Papio anubis). 81
3.3 Two gelada baboon (Theropithecus gelada) one-male units in the
Simen Mountains, Ethiopia. 82
3.4 A juvenile Neblina black-headed uacari (Cacajao hosomi). 83
3.5 The number of primate species on islands and continents is a
function of the area of tropical forest. 84

xiv
 Figures

3.6 Vegetation zones from the equator to the Tropic of Cancer. 86

3.7 The relationship between climatic factors and the phylogenetic
structure of mammal, carnivore, primate, and ungulate
communities based on the Nearest Taxon Index. 87
3.8 Japanese macaques (Macaca fuscata). 88
3.9 Adult female and male chimpanzees (Pan troglodytes) at Mahale
National Park, Tanzania. 90
B3.2 Hamadryas (Papio hamadryas) and olive baboon (P. anubis) hybrid. 93
B3.3 The elusive golden bamboo lemur (Hapalemur aureus). 95
3.10 The woolly lemur (Avahi laniger). 97
3.11 Orangutan tool use. 98
3.12 Bearded capuchin monkey (Sapajus libidinosus) at Fazenda Boa
Vista, Piaui state, Brazil. 99
4.1 Yellow baboons (Papio cynocephalus) at Amboseli National Park. 102
4.2 Mendel’s first law of segregation. 106
4.3 Sex ratios also affect extinction risks. 113
4.4a Barbary macaques (Macaca sylvanus). 114
4.4b Celebes or crested black macaque (Macaca nigra). 114
4.4c Lion-tailed macaque (Macaca silenus). 115
4.5 Cotton-top tamarins (Saguinus oedipus) at the University of
Wisconsin–Madison. 116
4.6 Langur (Semnopithecus entellus) male band incursion into a troop
from which the resident male had vanished. 117
4.7 Adult female black and white colobus (Colobus guereza) with her
10-week-old infant at Kanyawara, Kibale Forest, Uganda. 118
4.8 Male chimpanzee (Pan troglodytes) at Gombe National Park eating
red colobus (Procolobus badius) meat obtained by predation. 122
4.9 Percentage of time spent grooming among dyads as function of
matrilineal relatedness in a group of Japanese macaques. 128
4.10 Vervet monkey responses to recruitment vocalizations of kin and
nonkin in the presence and absence of prior grooming by the
vocalizers. 129
4.11 Chimpanzee (Pan troglodytes) coalition at Arnhem Zoo,
Netherlands. 133
4.12 Male yellow baboons (Papio cynocephalus) at Amboseli National
Park, Kenya. 134
4.13 The Prisoner’s Dilemma game. 136
4.14 Vervet monkeys (Chlorocebus pygerythrus) at the Samara Private
Game Reserve, South Africa. 137
4.15 White-faced capuchin monkey (Cebus imitator) alpha- and
beta-ranking females threaten the alpha male. 139
4.16 A fatally wounded male chimpanzee (Pan troglodytes) from the
Kahama community at Gombe. 140
4.17 Adult female northern muriqui (Brachyteles hypoxanthus) nursing her
infant. 141

xv
Figures

5.1 Olive baboon (Papio anubis) male displaying his canines. 144
5.2 Male mandrills (Mandrillus sphinx) are larger and their faces much
more vibrantly marked than females. 145
5.3 An adult red-tailed guenon (Cercopithecus ascanius) male weighs
about 2 pounds more than an adult female. 145
5.4 An adult female long-tailed macaque (Macaca fascicularis) with her
first infant eating a Ficus fruit. 147
5.5 Bolivian white-eared titi monkeys (Plecturocebus donacophilus
[revised from Callicebus donacophilus, per Byrne et al., 2016]) are
one of the few monogamously pair-bonded primates. 152
B5.1 White-handed (lar) gibbons (Hylobates lar) resting in a fig tree
above the Lam Takhong River in Central Mo Singto, Khao Yai
National Park, Thailand. 153
5.6 A fully adult male orangutan (Pongo pygmaeus). 156
5.7 An adult Zanzibar galago (Galagoides zanzibaricus) from Gedi,
Kenya, poised in a tree near a trap. 157
5.8 Body size dimorphism, relative canine size, and relative testes size
for primate genera belonging to different breeding systems. 159
5.9 Northern muriqui (Brachyteles hypoxanthus) males in an embrace
huddle. 160
5.10 Japanese macaques (Macaca fuscata) huddling. 161
5.11 Adult male Yunnan snub-nosed monkey (Rhinopithecus bieti)
walking on the branch of a Chinese white pine. 162
5.12 A juvenile male langur (Semnopithecus entellus) grooms a serious
abdominal wound he received during a fight with a competing
male band. 163
5.13 Mountain gorilla (Gorilla berengei) group. 164
5.14 Patas monkeys (Erythrocebus patas) drinking at a water hole. 166
5.15 Ring-tailed lemurs (Lemur catta). 167
5.16 Adult male Central American squirrel monkey (Saimiri oerstedii)
in a “fattened” state feeding on fruit while a six-month-old
youngster looks on. 168
5.17 Seasonality continuum in which the timing of matings and
conceptions entirely overlap either seasonally or year-round, or
only partially. 169
5.18 The polygyny threshold in birds. 172
5.19 An alpha male black-horned capuchin (Sapajus nigritus), shown in
proximity to an infant in his group. 173
5.20 Red-tailed guenon (Cercopithecus ascanius) female solicits
grooming from an adult male, who complies. 175
5.21 Female yellow baboon (Papio cynocephalus) with a sexual swelling
is being groomed by her male consort. 176
5.22 A 15-year-old alpha male Barbary macaque (Macaca sylvanus) in
contact with a younger adult male and an infant at Affenberg
Salem, a semi-free-ranging monkey sanctuary in Germany. 179

xvi
 Figures

5.23 This old male langur (Semnopithecus entellus) has a facial scar and
nearly absent incisors. 180
5.24 A consorting pair of chacma baboons (Papio ursinus) at De Hoop
Nature Reserve. 181
6.1 Northern muriqui monkeys (Brachyteles hypoxanthus). 184
6.2 Blue monkey (Cercopithecus mitis) at Kibale National Park, Uganda. 185
6.3 Vervet monkey (Chlorocebus pygerythrus) mother carrying her
nursing infant while she climbs into a feeding tree. 186
6.4 Red-tailed monkey (Cercopithecus ascanius) feeding on fig fruit
(Ficus natalensis) at Kibale National Park, Uganda. 188
6.5 Morphological adaptations to diet. 191
6.6 Selection of pods from the Bignoniaceae family. 192
6.7 Juvenile red colobus (Procolobus kirkii) eating charcoal to detoxify
secondary plant compounds on Zanzibar. 195
6.8 Range of body sizes in the major dietary categories. 196
6.9 Relationship between body mass and digestive transit times. 197
6.10a Costa Rican white-faced capuchin (Cebus imitator) adult male and
large juvenile male descending a tree to nab a coati pup (Nasua
narica). 198
6.10b Adult male grabs the coati pup. 198
6.10c Adult male returns to the safety of the trees with his prey in hand. 198
6.11 Patas monkey (Erythrocebus patas) feeding on gum from Acacia
drepanalobium tree. 199
6.12 A female red-bellied lemur (Eulemur rubriventer) feeding on a
berry from the Psychotria family. 200
6.13 Adult female diademed sifaka (Propithecus diadema) feeding on
new leaves at Vatoharanana study site, Ranomafana National Park,
Madagascar. 201
6.14 The influence of the spatial distribution of resources on the ability
of individuals to monopolize them. 201
6.15 Japanese macaques (Macaca fuscata). 204
6.16 Northern muriquis (Brachyteles hypoxanthus). 205
6.17 The advantage of cooperating at a food patch containing a limited
number of feeding sites. 207
B6.2 Wright’s (1999) “energy frugality” model, in which poor soils and
island climate result in scarce and unpredictable food resources for
the lemurs of Madagascar. 209
6.18 Lowland gorillas (Gorilla gorilla) in the Central African Republic
consume a higher proportion of fruit than their mountain gorilla
cousins. 212
6.19 Lowland gorilla group (Gorilla gorilla) sitting near forest edge in
Mbeli Bai, Nouabalé-Ndoki National Park, Republic of Congo. 213
7.1 Mother olive baboon (Papio anubis) secures her infant while nursing. 216
7.2 Matrilineal gelada baboons (Theropithecus gelada) live in one-male
groups, or bands, that are part of a larger clan. 218

xvii
Figures

7.3 The dynamic relationship between group size and group growth
rate in social primates. 219
7.4 Members of a wild patas monkey (Erythrocebus patas) matrilineal
group survey the field. 220
7.5 Stump-tailed macaques (Macaca arctoides) engage in a grooming
session. 221
B7.1 The late Irven DeVore (1934–2014) watching olive baboons (Papio
anubis) during his original field study in Nairobi National Park. 223
7.6 Vervet monkeys (Chlorocebus pygerythrus) in Amboseli National
Park, Kenya. 225
7.7 Amboseli yellow baboons (Papio cynocephalus) in a triadic interaction. 226
7.8 An adult female yellow baboon (Papio cynocephalus) handles the
infant of another female at Amboseli National Park, Kenya. 228
7.9 Adult male Barbary macaque with infant (Macaca sylvanus). 230
7.10 A troop of black and white colobus monkeys (Colobus guereza). 231
7.11 A male red-tailed guenon (Cercopithecus ascanius). 232
7.12 Ring-tailed lemurs (Lemur catta). 233
7.13 A troop of brown howler monkeys (Alouatta guariba). 234
7.14 A rare northern muriqui (Brachyteles hypoxanthus) matriline. 235
7.15 An anubis-like female baboon (Papio anubis) grooms the hybrid
male of her unit. 238
7.16 Influences on mating systems. 239
7.17 Wild female bonobos (Pan paniscus) engage in genital–genital, or
g–g, rubbing. 240
7.18 Although female bonobos (Pan paniscus) typically disperse from
their natal communities, g–g rubbing helps them to establish social
bonds and reinforce their relationships. 240
7.19 Female patas monkeys (Erythrocebus patas) in Laikipia, Kenya. 242
7.20 Barbary macaques (Macaca sylvanus) engaged in a mutual
grooming bout reinforce social bonds. 243
7.21 Increase in the Birth Sex Ratio of a group of northern muriquis
(Brachyteles hypoxanthus). 247
8.1 Survival curves for seven wild primate populations. 251
B8.1 Adult male gelada baboon (Theropithecus gelada) carries a
yearling on his back after being threatened by another male. 253
8.2 Male long-tailed macaques (Macaca fascicularis) on a river bank. 255
8.3 Variation in male reproductive skew. 256
8.4 Rhesus macaque (Macaca mulatta) consortship. 257
8.5 Rhesus macaques (Macaca mulatta) mating in a secluded treetop. 258
8.6 Male northern muriquis (Brachyteles hypoxanthus) relaxing
together. 259
8.7 Northern muriquis (Brachyteles hypoxanthus) mate in full view of
other group members. 261
8.8 Possible patterns of age-related change of dominance status of
male bonobos (Pan paniscus). 264

xviii
 Figures

8.9 Young male mountain gorillas (Gorilla berengei) at play. 265

8.10 Possible decision rules for emigration of hamadryas male baboons
(Papio hamadryas). 266
8.11 A coalition of two males of one faction in a male band of langurs
(Semnopithecus entellus) harass a member of the other faction,
who is retreating to one of his allies. 268
8.12 Male mountain gorilla (Gorilla berengei) life history patterns in
one-male and multi-male groups. 269
8.13 Juvenile white-faced capuchins (Cebus imitator) fight near the
alpha male of their group. 270
8.14 An adolescent male langur (Semnopithecus entellus) extends his
hand toward the ankle of an unfamiliar adult male who he had
been threatening while the adult male was copulating. 273
8.15 Two male Barbary macaques (Macaca sylvanus) play with an infant
on the grass. 274
8.16 A pair of nonmatrilineal white-handed (lar) gibbons (Hylobates
lar). The darker adult male is grooming a juvenile. 275
8.17 Male and female contributions to grooming exchanges differ across
species. 276
8.18 A chacma baboon male (Papio ursinus) at De Hoop Nature
Reserve surrounded by six females with infants vulnerable to
infanticide. 278
9.1 Life history model for generating predictions about the role of
production. 282
9.2 Relationship between neonatal weight and maternal weight. 283
9.3 A male infant slender loris (Loris lydekkerianus nordicus) at Giritele,
Sri Lanka. 283
9.4 Adult male chacma baboon (Papio ursinus) holds an infant during a
fight with another male at Okavango Crater. 286
9.5 Adult male buffy-headed marmoset (Callithrix flaviceps) shares
gum with an infant. 287
9.6 Juvenile female Barbary macaque (Macaca sylvanus) allomothering
an infant. 287
9.7 An adolescent female yellow baboon (Papio cynocephalus) carries
an infant. 290
9.8 The benefit, cost, and half the cost of an act of parental investment
toward an offspring. 291
9.9 Northern muriqui (Brachyteles hypoxanthus) flings herself across a
gap in the canopy. 292
9.10 An infant yellow baboon (Papio cyncocephalus) rides jockey-style
while her mother forages. 293
9.11 Primate species differ in their patterns of somatic growth. 295
9.12 An infant langur (Semnopithecus entellus) inspects the food that
her mother is eating. 296
9.13 Vervet monkeys (Chlorocebus pygerythrus) playing. 297

xix
Figures

9.14 Two adolescent male langurs (Semnopithecus entellus) play-fight in

a tree. 297
9.15 Juvenile northern muriquis (Brachyteles hypoxanthus) hanging out
together. 298
9.16 Primates have long life spans and late maturity. 302
B9.2 Maternally related yellow baboon (Papio cynocephalus) mothers
and their infants socializing in Amboseli National Park, Kenya. 303
9.17 Estimated population size from 1983 to 2010 and projected
population sizes to 2030 for northern muriquis at the Reserva
Particular do Patrimônio Natural Feliciano Miguel Abdala, Brazil. 307
9.18 Analyses of growth rates of mountain gorilla groups under
different management regimes. 308
9.19 Young bearded capuchin monkey (Sapajus libidinous). 309
10.1 An adult male Hanuman (or Bengal sacred) langur (Semnopithecus
entellus) “whoops,” a display or intergroup call. 312
10.2 An adult male yellow baboon (Papio cynocephalus) flashes his
eyelids in an aggressive display. 313
10.3 Begging hand gesture and screaming by a juvenile chimpanzee
(Pan troglodytes) from whom food was just taken away by a
dominant. 313
10.4 Immature yellow baboon (Papio cynocephalus) cowers and moves
aside as an adult male approaches. 315
10.5 Adult male Yakushima Japanese macaque (Macaca fuscata yakui)
grooms another adult male. 316
10.6a Stage one of a courtship sequence in bonnet macaques (Macaca
radiata). 317
10.6b Male grimaces. 317
10.6c The male extends his arm; the female appears to be threatening
him. 317
10.6d The pair copulate. 317
10.7 Ventro-ventral copulation between bonobos (Pan paniscus) at the
San Diego Zoo. 318
10.8 Frederico, a subadult male black-horned capuchin monkey (Sapajus
nigritus), displays his canines. 319
10.9 Female rhesus macaque (Macaca mulatta) looks at her mating partner. 319
10.10 Topography of the known cutaneous glands in prosimians. 321
10.11 A female Geoffroy’s spider monkey (Ateles geoffroyi) climbing a
tree trunk. 322
10.12 Representative audio spectrograms of calls produced by bonobos,
chimpanzees, and gorillas. 323
10.13 Male mantled howler monkey (Alouatta palliata) in forward threat. 325
10.14 Black-fronted titi monkeys (Callicebus nigrifrons) in the Brazilian
Atlantic forest. 329
10.15 Female chimpanzee (Pan troglodytes) fishes for termites while her
offspring observes at the Goualougo Chimpanzee Project. 330

xx
 Figures

10.16a A two-year old chimpanzee (Pan troglodytes) in Bossou, New

Guinea. 334
10.16b A 15-year-old adult male chimpanzee (Pan troglodytes) in Bossou,
New Guinea, using a stone hammer and anvil to crack open oil
palm nuts. 334
10.17 Young bearded capuchin monkeys (Sapajus libidinosus) learn their
mothers’ tool-using tradition. 335
10.18 Wild bonobo (Pan paniscus) dragging a branch. 336
10.19 A Japanese macaque (Macaca fuscata) at Arashiyama stone rubbing. 337
10.20a Nettles (Laportea). 337
10.20b A juvenile gorilla processes a nettle stem. 337
10.21 The “relational model” for resolving conflicts, described by de Waal. 338
10.22 Hypothetical series of ancestor species, whose cognitive
adaptations led to the human mind, in relation to their modern
descendants. 340
10.23 Vervet monkeys (Chlorocebus aethiops sabaeus) on the Caribbean
Island of St. Kitts. 345
B11.1a Agustin Fuentes during his work in Singapore in 2012, holding
a GPS collar that dropped off from an adult male long-tailed
macaque (Macaca fascicularis). 349
B11.1b Anthropologist and primatologist Erin Riley (center) conducting
an interview on macaque crop raiding with local farmers in South
Sulawesi, Indonesia. 350
11.1 A Suriname rainforest showing forest levels with different kinds of
substrates and the primates that occupy them. 352
11.2 A subadult female Nilgiri langur (Semnopithecus johnii) sitting next
to a juvenile Hanuman (or Bengal sacred) langur (Semnopithecus
entellus). 353
11.3 Blue monkey (Cercopithecus mitis) and red-tailed monkey
(C. ascanius) play chase. 353
11.4 Factors that influence the mixed-species associations of East African
guenons. 354
11.5 Red colobus (Piliocolobus badius) and Diana monkey (Cercopithecus
diana) association in the Taï National Forest, Ivory Coast. 357
B11.2 Adult male chimpanzee (Pan troglodytes) at Kibale National Park,
Uganda, with a red colobus monkey he caught. 359
11.6 Two subadult male white-faced capuchins (Cebus imitator)
dismembering a magpie jay hen. 361
11.7 Boa constrictor eating a female red howler monkey (Alouatta
puruensis) in the western Brazilian Amazon. 362
11.8 Male northern muriqui (Brachyteles hypoxanthus) feeding on
nectar from Mabea fistulifera, a member of the Euphorbiaceae
family. 365
11.9 Open ripe fruit of Micrandra spruceana (Euphorbiaceae), the most
important fruit in the diet of black-headed uacaris. 367

xxi
Figures

11.10 Hypothetical illustration of the seed shadow beneath a parent

tree, the probability of seed survival, and the effects of different
mechanisms of dispersal. 368
11.11a Flanged adult male orangutan (Pongo pygmaeus morio) moving
terrestrially in palm oil plantation in East Kalimantan, Indonesia, in
2012. 371
11.11b Unflanged adult male orangutan consuming young planted (under
one year) oil palms in palm oil plantation in East Kalimantan,
Indonesia, in 2012. 372
12.1 Annual rates of forest cover change in the buffer zone and within
Gunung Palung National Park, Kalimantan, from a 1988 baseline. 375
12.2 A fire, originally set to burn trash, spread into adjacent forest. 376
12.3 The lumber from a large tropical hardwood tree. 376
12.4 Rhesus macaques (Macaca mulatta). 377
12.5 Remaining forest fragments of southeastern Brazil, many of which
are too small and disturbed to support primates anymore. 377
12.6 An endangered Sclater’s guenon (Cercopithecus sclateri), victim of
a hunter. 380
12.7 A Yanomami Indian holding a smoked howler monkey. 381
12.8 Guenons (red-eared monkey, Cercopithecus erythrotis) and Preuss’s
monkey (Allochrocebus preussi) in the bushmeat market in Malabo,
Bioko Island. 382
12.9 Bengal sacred langurs, previously Hanuman langurs (Semnopithecus
entellus). 384
B12.1 Russell Mittermeier and the author, June 2013, in the forest at the
Private Natural Heritage Reserve, Brazil. 387
12.10 Northern muriqui (Brachyteles hypoxanthus) female with infant
standing bipedally while eating on the ground. 389
12.11 Hormonal profile from a wild female northern muriqui (previously
Brachyteles arachnoides; now B. hypoxanthus). 391
12.12 An example of dung from a wild northern muriqui (Brachyteles
hypoxanthus) collected for subsequent fecal steroid analyses. 392
12.13 Primatologist Jim Moore climbs into an abandoned nest made by a
chimpanzee (Pan troglodytes) to collect hair samples. 394
B12.2 Former Brazilian students have worked for more than a decade
with northern muriquis in Brazil. 395
12.14 Sclater’s guenons (Cercopithecus sclateri), among the most
endangered of Africa’s guenons. 397

xxii
Tables

1.1 Classification of some familiar living genera in the Primate Order 4

1.2 Northern muriqui society compared to the societies of other
well-known primates, including savanna baboons 8
1.3 Characteristics of primate groups 32
2.1 Ape life history traits 43
3.1 Major events in primate evolutionary history 77
3.2 Categories for threatened taxa 85
4.1 Processes that affect genetic variation within and between
populations 110
4.2 Types of interactions classified by their effects on fitness 127
6.1 General nutritional characteristics of broad categories of foods
consumed by primates 189
7.1 Competitive regimes and categories of female social relationships 219
8.1 Characteristics of female bonobo (Pan paniscus) and chimpanzee
(Pan troglodytes) cycles 264

xxiii
Another random document with
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 “Yes,” he said, looking intently. “The other. Mrs. Inigo. What’s
happened?”
“Pain’s got your crew,” Tucket answered. “He’s leaving you to the
Spaniards.”
Dully, as they passed about the rum-flask, they saw the boats
draw up to Pain’s ship. They saw sail made upon her, the sprit-sail
for casting, the topgallant-sails in the buntlines. Soon she was under
way, lying over to the breeze, sailing a point or two free, bound past
the Mestizos to the southward. Her men, gathered on her poop to
hoist the mizen, cursed the Broken Heart from the taffrail, firing a
volley of pistols at her in farewell. Two of the three sloops sailed at
the same time. As they passed away, crowding all sail, foaming at
the bows with their eight-knot rush, Margaret heard the chanties at
the halliards, a broken music, coming in the gusts of the wind.
Tucket took one of the oars, and sat at the thwart wearily,
searching for a helper among the sternsheets. Captain Margaret, the
Indian, could not row. The other five were exhausted. “Boys,” he
said, “if we don’t get a gait on us, we’ll be sunstruck. We’re losing
way, too. The breeze is setting us ashore.”
Margaret rose up wearily, like a man in a dream, and sat down to
row at the thwart. They pulled a ragged stroke together. They were
too tired to do much. They pulled a few strokes, and paused, to look
round. Then pulled again painfully and again paused. One of the
men took up a pistol and fired into the air. “We’ll never make it,” he
said. He fired again and again, till some one in the ship caught sight
of them and fired in reply. A boat manned by half a dozen men put
off to them, veering out line astern, so that those aboard might heave
them in at the capstan. Cammock and Perrin were two of the rowers,
the only two who could pull an oar; the negro steward steered. The
rest of them rowed like marines on pay-day going through the
platoon. The wounded and weary smiled to see them; they had
never seen such rowing. Slowly they drew up. Margaret saw Perrin
and Cammock glancing over their shoulders at him. They cheered
and waved when they saw him. He waved his unhurt arm to them.
The canoa swung round and backed alongside. Cammock, laying in
his oar, shifted his towing-line to the bow, and bent to it the painter of
Tucket’s canoa, so that the boats might tow together. “Let your oars
swing fore and aft, boys,” he said. He lifted his voice, and yelled to
the ship to heave in. “Well, sir,” he said, coming aft alongside
Margaret. “And how are you? I’m glad to see you alive. The men
said you were killed.”
“I’m all right,” he said. “But we had. We had enough, over there.
What has happened in the ship?”
“Well, sir,” said Cammock. “Pain’s gone off with most of what were
left to us. That’s what happened.”
“Tell me,” Margaret said. “How was it?”
“I was on deck with Mr. Perrin till the last canoas were putting off,”
Cammock answered. “We’d been working the guns. We saw there
was rather a mess. Then the doctor sends for us two to help him
take off Jowett’s leg. So we went into the wardroom, where he was
working. We heard the boats come alongside, and Pain lashing out
like a chocalatto-north; so when we got the arteries fast, and Jowett
in a cot, we ran on deck to see if you were safe and to see what the
row was. They’d been making enough row to caulk a flagship. That
was too late, then. Pain had been saying that you were dead, that
you’d spoiled the show and got a hundred of ’em killed. God knows
what he hadn’t said. More’n half of them were drunk. Our lot had
been up all night. So by the time we got on deck they were bundling
their bags into the canoas. There was that Fraser fellow going down
the gangway with one of our guns. I knocked him back where he
belonged. Then they began to fire. I’ll show you my hat when we get
aboard. Then they tried to rush the gangway. So I cut the fall, and
spilled ’em off it. We’d a lively fight for a few minutes. Pain and
Ackett’s lots (Ackett’s lot weren’t so bad) went off then. It was them
really kept Pain back a bit. They said they’d play hell with Springer’s
Key for me. Ackett’s lot said I’d better stand in with them. They said
they’d seen you corpsed on the beach.”
“He wasn’t far off it,” Tucket said. “But for this capitano here.”
“Mrs. Inigo went off with Iles,” said Perrin. “They’ve gone with Pain
and the rest.”
“The rest. The rest. What rest? We must have lost. Oh, my God,”
said Margaret weakly. “We must have lost half. Was any roll called?”
“No,” said Perrin. “Twelve of your men went to your sloop, Captain
Tucket, after bringing off some wounded. They said there was hell to
pay ashore, so they stayed here. That was before the attack. And
you’ve five more here; seventeen that makes, besides your ship-
keepers.”
“We’ve got eleven left,” Cammock said, “besides nineteen
wounded and the doctor. Not counting them. Tucket, we got thirty-
nine able-bodied between us. Twenty-four of them yours.”
“By the way, I’ve got your maps, Cammock,” Margaret said. “Here
they are. I’m sorry they’re so wet. I fell in the sea coming off.”
Cammock took the maps with a groan. “Thank you very much, sir,”
he said. “But I don’t much value the maps, though, when I think of
what’s happened.”
“It must have been hell,” said Perrin.
“Yes,” Margaret answered. “It was hell.”
“Why didn’t Pain take the ship?” Perrin asked. “That’s what
puzzles me.”
“Afraid you’d government friends, sir. Besides. He thought if he
took your men the Dagoes’d get you.”
Very slowly the men at the capstan walked the canoas to the side.
Margaret stood on his own deck again, asking himself how many
years had gone since he last stood there. The guns were cast loose.
Round shot, fallen from the garlands, rolled to and fro as the ship
rolled. The decks were littered with wads. Smears, as of lamp-black,
showed where the gun-sponges had been dropped. The hammock-
nettings were fire-pocked and filthy. There were marks of blood on
the coamings, where wounded men had lain, waiting to be carried
below.
“Charles,” said Perrin, “you come to the cabin and lie down. I’ll
fetch Olivia to you. She’s with the wounded. What about Stukeley?”
“He’s dead, Edward. He died of yellow fever this morning. I buried
him.”
“My God. Dead?”
“Yes. He was married there.”
“He deserted then, that time?”
“Yes.”
“And so he’s dead. The dead are.”
“The dead are our only links with God, I think,” Margaret said
gravely. “I’ve been in hell to-day, Edward. In hell. In hell.”
 He lay down on the window-seat in the cabin, where Stukeley had
so often lain. The breeze had swung the ship head to sea. He had
only to turn his head to see the fire of Tolu, burning below its pillar of
cloud. The sea-wall spurted with smoke at intervals. The flashes
were very white and bright, not like the smoke of firelocks. Cammock
came in to him with a mess of cold poultice for his head.
“We’re getting under way, sir,” he said. “The Dagoes are blowing
the spikes out of their guns. They’ll be firing soon. We’re cutting our
cable, sir. We haven’t strength to weigh. Tucket the same. Mrs.
Stukeley is coming to you, sir. I told her you were hurt. She wants
you not to get up, sir.”
He then went away. By and by Olivia entered.
“Don’t get up, Charles,” she said. “Oh. Don’t get up.”
He raised himself to greet her, looking at her sadly. She came up
to him and took his hand, and sat at his side.
“Olivia,” he said gently. “Olivia, I bring another sorrow for you. Your
husband is dead, Olivia. He died this morning.”
“Yes,” she answered quietly. “I knew he would be dead.”
He tried to read her thought; but his head was stupid with pain; he
could not. He saw only the calm, pale face with that quality of
mystery upon it which is upon all beauty.
“I feel for your sorrow,” he stammered.
“Yes, Charles,” she answered. “You feel for my sorrow, I know.”
“Olivia,” he said, “would it pain you too much to hear. To hear
about it?”
“You ought not to be talking with your wound, Charles.”
“I must tell you, Olivia. If you can bear it. It may be harder to-
morrow.”
“Tell me then,” she said. “If you feel. If you wish. I am quite calm,
Charles. Tell me everything.”
“He died of yellow fever, Olivia. In the Governor’s house there. The
tall house above the bastion there.”
“I can’t look, Charles. Don’t tell me where. I’m. Yes?”
“He was ill for four days,” Margaret continued. “They thought he
was getting better yesterday.”
“They, Charles? Who were ‘they’?”
“The Governor and a priest, Olivia.”
 “He was in the Governor’s house,” she said. “As a friend?”
“A secretary there.” His mouth had grown very dry; it was hard to
answer these questions.
“Then he stole the book of maps, Charles?”
“Yes.”
“Did you find it?”
“Yes. I found it.”
“Was there fighting? Fighting in the town? Was he alone when he
died?”
“He died before the fighting began. An hour or two before.”
“Who was with him, Charles?”
“A priest was with him.”
“A Roman Catholic priest?”
“Yes. I found the priest in the room with him. The priest had
attended him. At the end.”
“He had become a Roman Catholic?”
“Yes, Olivia.”
“I want to picture it, Charles. Was there fighting in the town when
you. When you found him?”
“There was a little firing. But the men were at breakfast. I heard a
few shots. Distant shots. So. So I left the men. And West and I made
a grave. We buried him there, Olivia. The priest. There was a
service.”
“And then. What happened then, Charles?”
“I picked these rosebuds, and this pebble, Olivia. From the grave.
For you.”
The rosebuds were crushed and dropping. She took them in the
hollow of her hand. There was a grace in all that she did. The
holding out of her delicate palm was beautiful; the movement
showed curiously her exquisite refinement; it was as though she had
said some beautiful thing; her mind showed in it. She took the relics,
looking not at them, but at Charles, her eyes swimming with tears,
her dear mind wild with tears.
“Thank you,” she said, on a sob. “You planted the roses, Charles?
You thought of it.”
He did not answer. He turned his head, to look out over Tolu. The
ship was moving slowly, heading out over the gulf; the fiery town was
dwindling; the wake whirled pale bubbles about the rudder.
“I think no other man would have thought of it,” she said quietly.
“Ah,” he said, sighing.
“Charles,” she said, “tell me now, will you. That’s over. That part.
What happened in the town?”
“The men got drunk, Olivia, and set the town on fire. They sacked
the place. When I came back to the Plaza. Oh, I can’t. I can’t.”
 “You sent off the wounded, then, did you not?”
“Yes.”
“I helped them, down in the ward-room. Oh, poor fellows. I asked
them about the battle. I think I’ve. I think, Charles, I’ve come very
near to the world in these last days. They told me a little. Then I
heard firing. We were busy below, with a sheet over the ports to
keep out the sun. Some of them are dreadfully hurt. I couldn’t see
what was happening. But I knew you were attacked. Then the
cannon overhead began. The ship shook. Then a wounded man
came down. He said you were dead, Charles. It was like the end of
everything.”
“It was, Olivia,” he said.
“I know now, Charles, what war is.”
“You didn’t see that water,” he answered. “Oh, not that water. My
act and deed. All of it. All of it by me. And there was a girl in the
Plaza. I’m disgraced. Oh, I shall never. It’s all over. All over after to-
day.”
“No, Charles. Don’t say that. You. At Springer’s Key. You can
begin again there. With better men.”
“Springer’s Key will be a failure, too,” he said bitterly. “Oh, Olivia,”
he added, going off into an hysterical laugh, “that water gets on my
nerves. There were two upset boats, and they can’t right them.”
“Charles,” she said, frightened. “You’re hurt. You’re hit again.”
“I think my old wound has broken out,” he said.
“Lie down,” she answered, rising swiftly, to lay the relics on the
table. “I’ll see to it at once. Why didn’t I think?”
“I’ll get Perrin,” he said. “Don’t you. You mustn’t. Oh, Olivia, you’ve
enough without me.”
She ripped the coat of soiled linen with the scissors in her
chatelaine. The old negro, coming in with chocolate, brought hot
water for her. Together they dressed the wound with balsam and
pitoma leaf, binding it with Indian cotton. The steward brought in fruit
and bread. They ate and drank together, mechanically, not as though
they wanted food.
“Olivia,” Margaret said, “you are in great sorrow. Some of it, Olivia,
perhaps all of it, is due to me. I want you. I want you to feel that I feel
for you. Feel deeply. Oh, my God. I’m sorry for you. You poor
woman.”
“Charles,” she said, “you mustn’t think. You’ve no right. You
mustn’t think that. That what has happened was due to you. Don’t,
Charles. You won’t, I know. I see too clearly what happened. I see
your mind, Charles, all along. I understand.” She knelt very swiftly
and kissed his hand. “There,” she said, very white. “I understand.”
Margaret closed his eyes, then looked at a gleam of flame far
distant, and at the blue band on the bows of Tucket’s sloop, plunging
the sea into milk within hail of him.
“You were right, Olivia,” he said, in a shaking voice. “I shall never
reap my plantations yonder.”
“No,” she answered. “Nor I.”
There was silence between them for a little while.
“Charles,” she added, “we both had Darien schemes.”
“Yes, Olivia.”
“They came to nothing. Because. There. We were too wild to see
what, what we were building with.”
“Yes, Olivia. And you reap sorrow. And I dishonour.”
“Not that, Charles. We reap the world.”
“Ah no, Olivia. This is the end of everything. For me it is.”
“No, Charles,” she said. “We were living in one little corner of our
hearts, you and I. In fool’s paradises. We were prisoners. This is not
the end. We only begin here.”
He sighed, thinking of the shame of the morning’s work in Tolu.
“My father’s sword,” he thought. “In that cause.”
“Charles,” she asked, “haven’t you. Something more to tell me?”
He thought for a moment, dully, wondering in his blurred brain if
she wished to hear more of the battle.
“No,” he said. “No. That is the end.”
She stood up, facing him, her great eyes looking down on him.
“I must go now to the wounded,” she said softly. “Edward and the
doctor will be wanting me. God bless you, Charles.”
“God bless you, Olivia. And comfort you.”
Before she left the cabin she turned and spoke again.
“He was married, Charles,” she said. “You never told me that.”
“Yes, Olivia, he was married.”
 “I knew that, Charles. I saw him so clearly. With a woman with a
cruel face. Oh, I knew it. It was generous of you not to tell me. But I
knew all the time.”
Late that night, in the darkened cabin, Olivia leaned upon the port-
sill, looking out over the rudder eddies, as they spun away in fire
rings, brightening and dying. She propped the cushions at her back,
so that she might rest her head. The nightmare of the past was
ashes to her. That evil fire had burned out, as Tolu had burned out.
The past and Tolu lay smouldering together somewhere, beyond
Fuerte there, beyond the Mestizos. The embers lay red there,
crusted in ash.
She had come to see clearly in the pain of her sorrow. She saw
her life laid bare and judged. She saw the moral values of things.
Great emotions are our high tides. They brim our natures, as a tide
brims the flood-marks, bringing strangeness out of the sea, wild birds
and amber. She had relics in her hand; rosebuds and a pebble,
“feathers and dust.” The rosebuds had spilled their petals. She
looked at them there, turning them over with her fingers, holding
them to the open port to see them. There was no moon; but the
great stars gave the night a kind of glimmering clearness. The sea
heaved silvery with star-tracks. Fish broke the water to flame. The
scutter of a settling sea-bird made a path of bright scales a few yards
from her. She laid the relics on the port-sill, near the open window.
Very gently she pushed the pebble into the water, leaning forward to
see the gleam of its fall. One by one she pushed the rosebuds over,
till they were all gone too. She watched the petals float away into the
wake, chased by the sea-birds. They were out of sight in a moment,
but the gulls mewed as they quarrelled over them, voices in the
darkness, crying in the air aloft. Olivia leaned there, looking after
them, for many minutes. Then she drew close the window and
covered her eyes with her hands.
It took them five days to win back to the Samballoes. They entered
Springer’s Drive a little before noon, eagerly looking out towards the
anchorage. No ships lay there, no guns greeted them from the fort.
Margaret and Cammock, walking the poop together, knew that Pain
had been before them. A thin expanse of smoke wavered and drifted
in films among the trees. When it drove down into the palms, after
rising above their level, it scattered the macaws, making them cry
out. The flagstaff lay prone, like a painted finger, pointing down the
spit to the sea. Tucket’s sloop was fifty yards ahead of the ship,
plunging in a smother. There was a cockling sea that morning, the
reefs were running white, they gleamed milky for fifty yards about
them. There was no other sign of life about the island. The smoke
was so thin that it was like mist. The beach, which had so lately been
thronged, was busy now with crabs, which scuttled and sidled,
tearing at the manchineel trees. A wounded man limped down the
sand and waved to them. Margaret, going in in his boat, saw that it
was the seaman West.
“I been here two days, sir,” he said, “waiting for you. I been living
on sapadillies. There been awful times, sir.”
“What has happened, man?”
“I came from the town, Tolu town, in one of the sloops, sir,” the
man answered. “The Lively, as they called her. When we come here,
Captain Pain got all the men you left, all the guard like, to sign on
with him. Then he set the town on fire, and scoffed all your gear, the
guns and powder and that. So I got away and hid in the wood. I was
afraid they want me to join ’em, or put a knife into me. Then I saw
’em flog that Don Toro, and two other Indians. He flog ’em on the
beach, and sent ’em back to the Main, sir. He said that would put all
them Indians off giving you a hand in the future.”
“What a devil the man must be,” Margaret said.
“So I stayed hidden, sir, ever since, hoping no Indians would come
over and find me.”
“So that’s the end,” said Margaret to himself. He would not go
ashore there. He could see the ruins of his city, a mass of fallen
earth, a heap of ashes, a sprouting crop blasted. He would never set
foot there again. That dream had ended like the other, in savagery, in
waste, in cruelty. He would let it end. The fallen gabions of the fort
would soon be tangled with grasses. In three months there would be
shrubs on the city site. The key would be jungle again, the Indians
would be savage again, the privateers would be plundering
vagabonds again. The dream was over. All that he could do now was
to proceed to Jamaica, to sell his goods there, before sailing for
England, a beaten man, threatened by the law.
 Tucket’s men helped his crew to fill fresh water. Tucket offered to
take seven of the slightly wounded men in his ship in exchange for
five unhurt men from his own crew. As the men were willing to
exchange, this brought the Broken Heart’s complement to twenty
men; enough, at a strain, for the passage to Jamaica, if no enemy
threatened and no storm arose. When the water had been filled, and
the manger stacked with wood, the men gathered stores of fruit.
They were ready to sail then. Margaret gave the Indian Robin
enough goods to make him a chief in his own land. He made gifts to
all of Tucket’s crew. To Tucket himself he gave a pair of pistols,
choice weapons, made by the best artist in Paris. Tucket asked for
his address, in writing.
“I shall come and look you up, one day,” he said, as he put the
paper in his pocket. “I shall be coming home to set up dyer. We’ll
have a great yarn, that day.”
“I shall expect you,” Margaret said. “You shall dye for me. But
won’t you come home now, captain? With me?”
“No, sir,” he answered. “I want to get that green the Indians get.
Then I’ll come home.”
“I’m sorry,” Margaret said. “Good-bye, then, Captain Tucket. I
wonder if we shall ever meet again.”
“Well. We met. Haven’t we? We neither of us expected to.”
“Good-bye, then. Let me hear from you, if a ship is going home
from here?”
“I will, indeed, sir. So long. So long, Lion.”
“So long,” said Cammock.
They were under way again, close-hauled to the breeze, going out
of Springer’s Drive to the east of Caobos. All the Holandès keys
were roaring with surf. The palms were bending. The smoke from the
key astern trailed in a faint streamer towards the Grullos. That was
the last picture which Margaret formed of the keys. The sun bright,
the palms lashing, the noise of the surf like a battle, the welter of the
surf like milk on the reefs. Tucket was in his sloop now, with all
hands gathered on deck, their faces turned to him. The men of the
Broken Heart were gathered at the hammock nettings. Margaret
thought of the sadness of parting. Two men had shaken hands only
a moment before. Now there was this gulf of sea between them. To-
morrow they would be many miles apart; and who knew whether
they would ever meet again, for all their wandering.
The bells of the ships rang out together, a furious peal. Cammock,
standing on a gun, took off his hat, and called for three cheers for the
sloop. The sloop’s men cheered the ship. The men of the Broken
Heart answered with a single cheer. The bells rang out again, the
colours dipped, the guns thundered, startling the pelicans. Tucket
had turned away now, to help to secure his guns. His helmsman let
the sloop go off three points. She was slipping fast away now, bound
towards Zambo-Gandi. Now the figures of the men could no longer
be recognized. She was hidden behind the palms of Puyadas.
Tucket was gone. Margaret never saw him again.
“That breaks the neck of that,” said Cammock. “All gone, main-
topgallant yard?”
“All gone, main-topgallant yard, sir.”
“Then hoist away.”
Under all sail the Broken Heart swayed seawards, treading down
the rollers, creaming a track across the sea, dark now in its
blueness, with crinkling wind-ruffles. When the night fell, shutting out
the Main, and the stars climbed out, solemn and golden, she was in
the strength of the trade, rolling to the northward, circled by the
gleams of dolphins, hurrying in sudden fires.
After dark that night Margaret sat on the locker-top, looking at the
wake, as it shone below him about the rudder. He was thinking over
his manifold failure, feeling disgraced and stained, a defeated,
broken man. Olivia entered quietly from the alleyway. He only felt her
enter. There was no light in the cabin. The steward was busy with
the wounded.
“Is that you, Olivia?” he asked, knowing that it was she. He felt in
his heart the gladness which her presence always gave to him. Life
could always be noble, he thought, with that beautiful woman in the
world.
“Yes, Charles,” she answered. “I’ve been with the wounded.
They’re better. How are your wounds?”
“Better, thanks. They’re always better at night.”
She drew up a chair and sat down beside him.
 “Charles,” she said, “I want you not to brood. Not to grieve. That’s
all over, Charles.”
“Not the dishonour,” he said. “That will never be over.”
“There is no dishonour, Charles. You failed. The only glory is
failure. All artists fail. But one sees what they saw. You see that in
their failure.”
“Ah. Sometimes.”
“I see that in yours, Charles.”
“Thank you, Olivia.”
“I shall always see that, Charles, when I see you.”
“My city.”
“Yes. Your city, Charles.”
“It was for you, Olivia. My city.”
“I am proud,” she said humbly.
“It failed, Olivia. It’s in ruins. Yonder.”
“Charles,” she said, kneeling, taking both his hands, hiding her
face against him, “it is in my heart. That city. Our city.”
She trembled against him, drawing her breath. He held her with
his unhurt arm, waiting till the dear face would lift, pale in that
darkness, to the laying of the gold foundation.

THE END
 TRANSCRIBER’S NOTES
The Grant Richards edition (London, 1908) was consulted for most
of the changes listed below.
Minor spelling inconsistencies (e.g. alleyway/alley-way, gold
dust/gold-dust, etc.) have been preserved.

Alterations to the text:

Formatting: abandon the use of drop-caps.
Punctuation: quotation mark pairings/nestings and missing
commas.
[Chapter III]
Change “Hands up anchor, bosum.” to bosun.
“looking aloft at the boy on the maintopgallantyard” to
maintopgallant yard.
“Whereever that vague perfume lingers, something of the old” to
Wherever.
[Chapter V]
“We’ll, go on with this at Accomac, Stukeley.” delete first comma.
“That man up on the yard there was once a slave in Virgina” to
Virginia.
[Chapter VI]
(“It’ll annoy old Brandy-face,” he thought.) to Brandyface.
“The long chase-guns were trained athwart-ships” to athwartships.
“The table was rimmed with a patten to keep the plates from
falling” to batten.
[Chapter IX]
“and so they lose, either way, But you know, Olivia, you are”
change second comma to a period.
“so that the dicussion came to an end before it had well begun” to
discussion.
[Chapter XI]
 “Margaret knocked the lance aside with difficultly” to difficulty.
[Chapter XII]
(said another rower, “this oar’l go in the slings.) to oar’ll.
“As they pased away, crowding all sail, foaming at the bows” to
passed.

[End of text]
*** END OF THE PROJECT GUTENBERG EBOOK CAPTAIN
MARGARET ***

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