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Thirteenth Edition

Biology
Kenneth A. Mason
University of Iowa

Jonathan B. Losos
William H. Danforth Distinguished University
Professor and Director, Living Earth Collaborative,
Washington University

Tod Duncan
University of Colorado Denver

Based on the work of

Peter H. Raven
President Emeritus, Missouri Botanical Garden;
George Engelmann Professor of Botany Emeritus,
Washington University

George B. Johnson
Professor Emeritus of Biology, Washington
University
BIOLOGY

Published by McGraw Hill LLC, 1325 Avenue of the Americas, New York, NY 10019. Copyright ©2023 by McGraw Hill LLC. All rights
reserved. Printed in the United States of America. No part of this publication may be reproduced or distributed in any form or by any means, or
stored in a database or retrieval system, without the prior written consent of McGraw Hill LLC, including, but not limited to, in any network or
other electronic storage or transmission, or broadcast for distance learning.

Some ancillaries, including electronic and print components, may not be available to customers outside the United States.

This book is printed on acid-free paper.

1 2 3 4 5 6 7 8 9 LWI 27 26 25 24 23 22

ISBN 978-1-265-12884-5
MHID 1-265-12884-7

Cover Image: Tree Frog: Shutterstock/Kurit afshen, Mushrooms: Russell Illig/Getty Images, Antibodies immunoglobulins attacking corona-
virus: Corona Borealis Studio/Shutterstock

All credits appearing on page or at the end of the book are considered to be an extension of the copyright page.

The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website does not indicate an endorsement
by the authors or McGraw Hill LLC, and McGraw Hill LLC does not guarantee the accuracy of the information presented at these sites.

mheducation.com/highered
Brief Contents

Committed to Excellence xi 27 Prokaryotes 560

Preparing Students for the Future xv 28 Protists 586
29 Seedless Plants 610

I The Molecular Basis of Life

30 Seed Plants 625
Part 1 31 Fungi 643
32 Animal Diversity and the Evolution of Body Plans 666
1 The Science of Biology 1 33 Protostomes 688
2 The Nature of Molecules and the Properties of Water 18 34 Deuterostomes 721
3 The Chemical Building Blocks of Life 35

Part II Biology of the Cell 62

Part VI Plant Form and Function 762

35 Plant Form 762

4 Cell Structure 62 36 Transport in Plants 788
5 Membranes 92 37 Plant Nutrition and Soils 807
6 Energy and Metabolism 112 38 Plant Defense Responses 825
7 How Cells Harvest Energy 127 39 Plant Sensory Systems 838
8 Photosynthesis 153 40 Plant Reproduction 866
9 Cell Communication 175

VII Animal Form and Function

10 How Cells Divide 194
Part 899
Part III Genetic and Molecular Biology 216 41
42
The Animal Body and Principles of Regulation 899
The Nervous System 923
11 Sexual Reproduction and Meiosis 216 43 Sensory Systems 953
12 Patterns of Inheritance 230 44 The Endocrine System 979
13 The Chromosomal Basis of Inheritance, and Human 45 The Musculoskeletal System 1003
Genetics 248 46 The Digestive System 1023
14 DNA: The Genetic Material 268 47 The Respiratory System 1044
15 Genes and How They Work 290 48 The Circulatory System 1063
16 Control of Gene Expression 318 49 Osmotic Regulation and the Urinary System 1085
17 Biotechnology 341 50 The Immune System 1103
18 Genomics 368 51 The Reproductive System 1133
19 Cellular Mechanisms of Development 390 52 Animal Development 1155

Part IV Evolution 416 Part VIII Ecology and Behavior 1185

20 Genes Within Populations 416 53 Behavioral Biology 1185
21 The Evidence for Evolution 443 54 Ecology of Individuals and Populations 1215
22 The Origin of Species 463 55 Community Ecology 1241
23 Systematics, Phylogenies, and Comparative Biology 484 56 Dynamics of Ecosystems 1264
24 Genome Evolution 505 57 The Biosphere and Human Impacts 1288
58 Conservation Biology 1317
Part V Diversity of Life on Earth 524
Appendix A
25 The Origin and Diversity of Life 524 Glossary G-1
26 Viruses 539 Index I-1

iii
About the Authors
Kenneth Mason maintains an association with the University of Iowa, Department of Biology after having served
as a faculty member for eight years. His academic positions, as a teacher and researcher, include the faculty
of the University of Kansas, where he designed and established the genetics lab, and taught and published
on the genetics of pigmentation in amphibians. At Purdue University, he successfully developed and grew large
introductory biology courses and collaborated with other faculty in an innovative biology, chemistry, and
physics course supported by the National Science Foundation. At the University of Iowa, where his wife served as
©Kenneth Mason president of the university, he taught introductory biology and human genetics. His honor society memberships
include Phi Sigma, Alpha Lambda Delta, and, by vote of Purdue pharmacy students, Phi Eta Sigma Freshman
Honors Society.

Jonathan Losos is the William H. Danforth Distinguished University Professor in the Department of Biology
at Washington University and Director of the Living Earth Collaborative, a partnership between the university,
the Saint Louis Zoo, and the Missouri Botanical Garden. Losos’s research has focused on studying patterns
of adaptive radiation and evolutionary diversification in lizards. He is a member of the National Academy of
Sciences, a fellow of the American Academy of Arts and Science, and the recipient of several awards, including
the Theodosius Dobzhanksy and David Starr Jordan Prizes, the Edward Osborne Wilson Naturalist Award, and
©Jonathan Losos the Daniel Giraud Elliot Medal, as well as receiving fellowships from the John Guggenheim and David and Lucile
Packard Foundations. Losos has published more than 250 scientific articles and has written two books, Lizards
in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles (University of California Press, 2009) and
Improbable Destinies: Fate, Chance, and the Future of Evolution (Penguin-Random House, 2017). He is currently
in the process of writing his next book, on scientific research on the ecology and evolution of domestic cats.

Tod Duncan is a Clinical Associate Professor at the University of Colorado Denver. He currently teaches the
first semester of introductory biology, which focuses on molecular and cellular systems; he also teaches upper-
division courses in virology and cancer biology. Tod also runs the first-semester introductory biology labs in
the course-based undergraduate research (CURE) format, in which up to 800 students each semester perform
metagenomic analysis of purebred Labrador retriever gastrointestinal microbiota. Previously, he taught general
microbiology, virology, the biology of cancer, medical microbiology, and cell biology. A bachelor’s degree in cell
©Lesley Howard biology with an emphasis on plant molecular and cellular biology from the University of East Anglia in England
led to doctoral studies in cell cycle control, and postdoctoral research on the molecular and biochemical
mechanisms of DNA alkylation damage in vitro and in Drosophila melanogaster. Currently, he is interested in
factors affecting retention and success of incoming first-year students in diverse demographics.

iv
 Contents

Committed to Excellence xi 4.4 The Endomembrane System 72

Preparing Students for the Future
Soames Summerhays/Natural Visions
xv 4.5 Mitochondria and Chloroplasts: Cellular
Generators 76
4.6 The Cytoskeleton 78
4.7 Extracellular Structures and Cell Movement 82
4.8 Cell-to-Cell Interactions 85

I The Molecular Basis

5 Membranes 92
Part 5.1 The Structure of Membranes 93
5.2 Phospholipids: The Membrane’s Foundation 96
of Life 5.3 Proteins: Multifunctional Components 98
5.4 Passive Transport Across Membranes 100
1 The Science of Biology 1 5.5 Active Transport Across Membranes 104
5.6 Bulk Transport by Endocytosis and Exocytosis 106
1.1 The Science of Life 2
1.2 The Nature of Science 4
1.3 An Example of Scientific Inquiry: Darwin and
6 Energy and Metabolism 112
Evolution 8 6.1 The Flow of Energy in Living Systems 113
1.4 Core Concepts in Biology 12 6.2 The Laws of Thermodynamics and
Free Energy 114
2 The Nature of Molecules and the 6.3 ATP: The Energy Currency of Cells 117
Properties of Water 18 6.4 Enzymes: Biological Catalysts 118
6.5 Metabolism: The Chemical Description of Cell
2.1 The Nature of Atoms 19 Function 122
2.2 Elements Found in Living Systems 23
2.3 The Nature of Chemical Bonds 24 7 How Cells Harvest Energy 127
2.4 Water: A Vital Compound 26
7.1 Overview of Cellular Respiration 128
2.5 Properties of Water 29
7.2 Glycolysis: Splitting Glucose 132
2.6 Acids and Bases 30
7.3 The Oxidation of Pyruvate Produces
Acetyl-CoA 135
3 The Chemical Building Blocks of Life 35 7.4 The Citric Acid Cycle 136
3.1 Carbon: The Framework of Biological Molecules 36 7.5 The Electron Transport Chain and
3.2 Carbohydrates: Energy Storage and Structural Chemiosmosis 139
Molecules 40 7.6 Energy Yield of Aerobic Respiration 142
3.3 Nucleic Acids: Information Molecules 43 7.7 Regulation of Aerobic Respiration 143
3.4 Proteins: Molecules with Diverse Structures and 7.8 Oxidation Without O2 144
Functions 46
7.9 Catabolism of Proteins and Fats 146
3.5 Lipids: Hydrophobic Molecules 56
7.10 Evolution of Metabolism 148
Dr. Gopal Murti/Science Source
8 Photosynthesis 153

II Biology of the Cell

8.1 Overview of Photosynthesis 154
8.2 The Discovery of Photosynthetic
Part Processes 155
8.3 Pigments 157
4 Cell Structure 62 8.4 Photosystem Organization 160
4.1 Cell Theory 63 8.5 The Light-Dependent Reactions 162
4.2 Prokaryotic Cells 66 8.6 Carbon Fixation: The Calvin Cycle 166
4.3 Eukaryotic Cells 68 8.7 Photorespiration 169

v
 9 Cell Communication 175 14.3 Basic Characteristics of DNA Replication 275
14.4 Prokaryotic Replication 278
9.1 Overview of Cell Communication 176
14.5 Eukaryotic Replication 283
9.2 Receptor Types 179
14.6 DNA Repair 285
9.3 Intracellular Receptors 180
9.4 Signal Transduction Through Receptor
Kinases 182
15 Genes and How They Work 290
9.5 Signal Transduction Through G Protein–Coupled 15.1 The Nature of Genes 291
Receptors 186 15.2 The Genetic Code 294
15.3 Prokaryotic Transcription 297
10 How Cells Divide 194 15.4 Eukaryotic Transcription 299
10.1 Bacterial Cell Division 195 15.5 Eukaryotic pre-mRNA Splicing 301
10.2 Eukaryotic Chromosomes 197 15.6 The Structure of tRNA and Ribosomes 303
10.3 Overview of the Eukaryotic Cell Cycle 200 15.7 The Process of Translation 306
10.4 Interphase: Preparation for Mitosis 201 15.8 Summarizing Gene Expression 310
10.5 M Phase: Chromosome Segregation and the Division 15.9 Mutation: Altered Genes 311
of Cytoplasmic Contents 202
10.6 Control of the Cell Cycle 206 16 Control of Gene Expression 318
10.7 Genetics of Cancer 211
16.1 Control of Gene Expression 319
Steven P. Lynch
16.2 Regulatory Proteins 320
16.3 Prokaryotic Regulation 322

III Genetic and Molecular

16.4 Eukaryotic Regulation 326
16.5 Chromatin Structure Affects Gene Expression 329
Part 16.6 Eukaryotic Posttranscriptional Regulation 331
16.7 Protein Degradation 335
Biology
17 Biotechnology 341
11 Sexual Reproduction and Meiosis 216 17.1 Recombinant DNA 342
11.1 Sexual Reproduction Requires Meiosis 217 17.2 Amplifying DNA Using the Polymerase Chain
Reaction 346
11.2 Features of Meiosis 218
17.3 Creating, Correcting, and Analyzing Genetic
11.3 The Process of Meiosis 219 Variation 349
11.4 Summing Up: Meiosis versus Mitosis 225 17.4 Constructing and Using Transgenic Organisms 351
17.5 Environmental Applications 355
12 Patterns of Inheritance 230 17.6 Medical Applications 357
12.1 The Mystery of Heredity 231 17.7 Agricultural Applications 361
12.2 Monohybrid Crosses: The Principle of
Segregation 233
18 Genomics 368
12.3 Dihybrid Crosses: The Principle of Independent
Assortment 236 18.1 Mapping Genomes 369
12.4 Probability: Predicting the Results of Crosses 238 18.2 Sequencing Genomes 372
12.5 Extensions to Mendel 239 18.3 Genome Projects 375
18.4 Genome Annotation and Databases 377
13 The Chromosomal Basis of Inheritance, 18.5 Comparative and Functional Genomics 380
and Human Genetics 248 18.6 Applications of Genomics 385

13.1 Sex Linkage and the Chromosomal Theory of

Inheritance 249 19 Cellular Mechanisms of
13.2 Exceptions to Mendelian Inheritance 251 Development 390
13.3 Genetic Mapping 253 19.1 The Process of Development 391
13.4 Human Genetics 256 19.2 Cell Division 391
13.5 Human Genetic Mapping and Association Studies 262 19.3 Cell Differentiation 393
19.4 Nuclear Reprogramming 398
14 DNA: The Genetic Material 268 19.5 Pattern Formation 402
14.1 The Nature of the Genetic Material 269 19.6 Evolution of Pattern Formation 407
14.2 DNA Structure 271 19.7 Morphogenesis 410

vi Contents
tamoncity/Shutterstock 24.4 Gene Function and Expression Patterns 516
24.5 Applying Comparative Genomics 517

Part
IV Evolution Jeff Hunter/The Image Bank/Getty Images

20 Genes Within Populations 416

20.1 Genetic Variation and Evolution 417
Part
V Diversity of Life
on Earth
20.2 Changes in Allele Frequency 418
20.3 Five Agents of Evolutionary Change 420
20.4 Quantifying Natural Selection 425
20.5 Reproductive Strategies 426 25 The Origin and Diversity of Life 524
20.6 Natural Selection’s Role in Maintaining Variation 430 25.1 Deep Time 526
20.7 Selection Acting on Traits Affected by Multiple 25.2 Origins of Life 526
Genes 432 25.3 Evidence for Early Life 529
20.8 Experimental Studies of Natural Selection 434 25.4 Earth’s Changing System 531
20.9 Interactions Among Evolutionary Forces 436 25.5 Ever-Changing Life on Earth 532
20.10 The Limits of Selection 437
26 Viruses 539
21 The Evidence for Evolution 443
26.1 The Nature of Viruses 540
21.1 The Beaks of Darwin’s Finches: Evidence of Natural
Selection 444 26.2 Viral Diversity 544
21.2 Peppered Moths and Industrial Melanism: More Evidence 26.3 Bacteriophage: Bacterial Viruses 546
of Selection 446 26.4 Viral Diseases of Humans 547
21.3 Artificial Selection: Human-Initiated Change 448 26.5 Prions and Viroids: Infectious Subviral Particles 555
21.4 Fossil Evidence of Evolution 450
21.5 Anatomical Evidence for Evolution 454 27 Prokaryotes 560
21.6 Convergent Evolution and the Biogeographical 27.1 Prokaryotic Diversity 561
Record 456
27.2 Prokaryotic Cell Structure 565
21.7 Darwin’s Critics 458
27.3 Prokaryotic Genetics 569
27.4 The Metabolic Diversity of Prokaryotes 574
22 The Origin of Species 463 27.5 Microbial Ecology 576
22.1 The Nature of Species and the Biological Species 27.6 Bacterial Diseases of Humans 578
Concept 464
22.2 Natural Selection and Reproductive Isolation 468
28 Protists 586
22.3 The Role of Genetic Drift and Natural Selection in
Speciation 470 28.1 Eukaryotic Origins and Endosymbiosis 587
22.4 The Geography of Speciation 471 28.2 Overview of Protists 589
22.5 Adaptive Radiation and Biological Diversity 473 28.3 Characteristics of the Excavata 591
22.6 The Pace of Evolution 478 28.4 Characteristics of the SAR: Stramenopila 594
22.7 Speciation and Extinction Through Time 479 28.5 Characteristics of the SAR: Alveolata 596
28.6 Characteristics of the SAR: Rhizaria 600
23 Systematics, Phylogenies, and 28.7 Characteristics of the Archaeplastida 601
Comparative Biology 484 28.8 Characteristics of the Amoebozoa 604
28.9 Characteristics of the Opisthokonta 606
23.1 Systematics 485
23.2 Cladistics 486
23.3 Systematics and Classification 490
29 Seedless Plants 610
23.4 Phylogenetics and Comparative Biology 493 29.1 Origin of Land Plants 611
23.5 Phylogenetics and Disease Evolution 499 29.2 Bryophytes Have a Dominant Gametophyte
Generation 613
24 Genome Evolution 505 29.3 Tracheophytes Have a Dominant Sporophyte
Generation 615
24.1 Comparative Genomics 506 29.4 Lycophytes Diverged from the Main Lineage
24.2 Genome Size 509 of Vascular Plants 618
24.3 Evolution Within Genomes 513 29.5 Pterophytes Are the Ferns and Their Relatives 619

Contents vii
 30 Seed Plants 625 Susan Singer/McGraw Hill

30.1 The Evolution of Seed Plants 626

30.2
30.3
30.4
Gymnosperms: Plants with “Naked Seeds” 626
Angiosperms: The Flowering Plants 630
Seeds 636
Part
VI Plant Form and
30.5 Fruits 637 Function
31 Fungi 643 35 Plant Form 762
31.1 Classification of Fungi 644
35.1 Organization of the Plant Body: An Overview 763
31.2 Fungal Forms, Nutrition, and Reproduction 645
35.2 Plant Tissues 766
31.3 Fungal Ecology 648
35.3 Roots: Anchoring and Absorption Structures 772
31.4 Fungal Parasites and Pathogens 652
35.4 Stems: Support for Above-Ground Organs 776
31.5 Basidiomycota: The Club (Basidium) Fungi 654
35.5 Leaves: Photosynthetic Organs 781
31.6 Ascomycota: The Sac (Ascus) Fungi 656
31.7 Glomeromycota: Asexual Plant Symbionts 658 36 Transport in Plants 788
31.8 Zygomycota: Zygote-Producing Fungi 658
36.1 Transport Mechanisms 789
31.9 Chytridiomycota and Relatives: Fungi with
Zoospores 660 36.2 Water and Mineral Absorption 792
31.10 Microsporidia: Unicellular Parasites 661 36.3 Xylem Transport 795
36.4 Rate of Transpiration 797
32 Animal Diversity and the Evolution 36.5 Water-Stress Responses 799
of Body Plans 666 36.6 Phloem Transport 801

32.1 Some General Features of Animals 667 37 Plant Nutrition and Soils 807
32.2 Evolution of the Animal Body Plan 668
37.1 Soils: The Substrates on Which Plants Depend 808
32.3 Animal Phylogeny 672
37.2 Plant Nutrients 811
32.4 Parazoa: Animals That Lack Specialized
Tissues 676 37.3 Special Nutritional Strategies 813
32.5 Eumetazoa: Animals with True Tissues 679 37.4 Carbon–Nitrogen Balance and Global Change 816
32.6 The Bilateria 684 37.5 Phytoremediation 819

33 Protostomes 688 38 Plant Defense Responses 825

33.1 The Clades of Protostomes 689 38.1 Physical Defenses 826
33.2 Flatworms (Platyhelminthes) 690 38.2 Chemical Defenses 828
33.3 Rotifers (Rotifera) 693 38.3 Animals That Protect Plants 831
33.4 Mollusks (Mollusca) 694 38.4 Systemic Responses to Invaders 832
33.5 Annelids (Annelida) 700
33.6 Ribbon Worms (Nemertea) 703 39 Plant Sensory Systems 838
33.7 Bryozoans (Bryozoa) and Brachiopods (Brachiopoda) 704 39.1 Responses to Light 839
33.8 Roundworms (Nematoda) 706 39.2 Responses to Gravity 843
33.9 Arthropods (Arthropoda) 708 39.3 Responses to Mechanical Stimuli 845
39.4 Responses to Water and Temperature 847
34 Deuterostomes 721 39.5 Hormones and Sensory Systems 849
34.1 Echinoderms 722
34.2 Chordates 724 40 Plant Reproduction 866
34.3 Nonvertebrate Chordates 726 40.1 Reproductive Development 867
34.4 Vertebrate Chordates 727 40.2 Making Flowers 869
34.5 Fishes 729 40.3 Structure and Evolution of Flowers 874
34.6 Amphibians 734 40.4 Pollination and Fertilization 877
34.7 Reptiles 738 40.5 Embryo Development 882
34.8 Birds 743 40.6 Germination 888
34.9 Mammals 747 40.7 Asexual Reproduction 891
34.10 Evolution of the Primates 752 40.8 Plant Life Spans 893

viii Contents
©Dr. Roger C. Wagner, Professor Emeritus of
Biological Sciences, University of Delaware
46 The Digestive System 1023
46.1 Types of Digestive Systems 1024

VII Animal Form and

46.2 The Mouth and Teeth: Food Capture and Bulk
Processing 1026
Part 46.3 The Esophagus and the Stomach: The Early Stages
of Digestion 1027
Function 46.4 The Intestines: Breakdown, Absorption, and
Elimination 1029
46.5 Accessory Organ Function 1032
41 The Animal Body and Principles 46.6 Neural and Hormonal Regulation of the Digestive
of Regulation 899 Tract 1034
46.7 Food Energy, Energy Expenditure, and Essential
41.1 Organization of Animal Bodies 900
Nutrients 1035
41.2 Epithelial Tissue 901
46.8 Variations in Vertebrate Digestive Systems 1039
41.3 Connective Tissue 904
41.4 Muscle Tissue 907 47 The Respiratory System 1044
41.5 Nerve Tissue 908
47.1 Gas Exchange Across Respiratory Surfaces 1045
41.6 Overview of Vertebrate Organ Systems 909
47.2 Gills, Cutaneous Respiration, and Tracheal
41.7 Homeostasis 912 Systems 1046
41.8 Regulating Body Temperature 914 47.3 Lungs 1049
47.4 Structures, Mechanisms, and Control of Ventilation
42 The Nervous System 923 in Mammals 1052
42.1 Nervous System Organization 924 47.5 Transport of Gases in Body Fluids 1056
42.2 The Mechanism of Nerve Impulse Transmission 927
42.3 Synapses: Where Neurons Communicate with Other 48 The Circulatory System 1063
Cells 932 48.1 Invertebrate Circulatory Systems 1064
42.4 The Central Nervous System: Brain and Spinal Cord 938 48.2 The Components of Vertebrate Blood 1065
42.5 The Peripheral Nervous System: Spinal and Cranial 48.3 Vertebrate Circulatory Systems 1068
Nerves 945
48.4 Cardiac Cycle, Electrical Conduction, ECG,
and Cardiac Output 1071
43 Sensory Systems 953 48.5 Blood Pressure and Blood Vessels 1075
43.1 Overview of Sensory Receptors 954
43.2 Touch, Pressure, and Body Position 955 49 Osmotic Regulation and the Urinary
43.3 Hearing, Vibration, and Balance 957 System 1085
43.4 Taste, Smell, and pH 963
49.1 Osmolarity and Osmotic Balance 1086
43.5 Temperature, Pain, Electric Currents, and Magnetic
Fields 965 49.2 Nitrogenous Wastes: Ammonia, Urea, and
Uric Acid 1087
43.6 Vision 967
49.3 Osmoregulatory Organs 1088
43.7 Evolution and Development of Eyes 973
49.4 Evolution of the Vertebrate Kidney 1090

44 The Endocrine System 979 49.5 The Mammalian Kidney 1092

49.6 Hormonal Control of Osmoregulatory Functions 1097
44.1 Regulation of Body Processes by Chemical Messengers 980
44.2 Overview of Hormone Action 985 50 The Immune System 1103
44.3 The Pituitary and Hypothalamus: The Body’s Control
Centers 988 50.1 Innate Immunity 1104
44.4 The Major Peripheral Endocrine Glands 993 50.2 Adaptive Immunity 1109
44.5 Other Hormones and Their Effects 997 50.3 Cell-Mediated Immunity 1114
50.4 Humoral Immunity and Antibody Production 1117
45 The Musculoskeletal System 1003 50.5 Autoimmunity and Hypersensitivity 1123
50.6 Antibodies in Medical Treatment and Diagnosis 1125
45.1 Types of Skeletal Systems 1004
50.7 Pathogens That Evade the Immune System 1127
45.2 A Closer Look at Bone 1006
45.3 Joints 1009
45.4 Muscle Contraction 1010
51 The Reproductive System 1133
45.5 Vertebrate Skeleton Evolution and Modes 51.1 Animal Reproductive Strategies 1134
of Locomotion 1017 51.2 Vertebrate Fertilization and Development 1136

Contents ix
 51.3 Structure and Function of the Human Male 54.4 Life History and the Cost of Reproduction 1224
Reproductive System 1140 54.5 Environmental Limits to Population Growth 1227
51.4 Structure and Function of the Human Female 54.6 Factors That Regulate Populations 1229
Reproductive System 1144
54.7 Human Population Growth 1232
51.5 Contraception and Infertility Treatments 1148
54.8 Pandemics and Human Health 1236

52 Animal Development 1155 55 Community Ecology 1241

52.1 Fertilization 1156
55.1 Biological Communities: Species Living Together 1242
52.2 Cleavage and the Blastula Stage 1160
55.2 The Ecological Niche Concept 1243
52.3 Gastrulation 1162
55.3 Predator–Prey Relationships 1248
52.4 Organogenesis 1166
55.4 The Many Types of Species Interactions 1252
52.5 Vertebrate Axis and Pattern Formation 1171
55.5 Ecological Succession, Disturbance, and Species
52.6 Human Development 1178 Richness 1258

K. Ammann/Bruce Coleman Inc./Photoshot

56 Dynamics of Ecosystems 1264

VIII Ecology and

56.1 Biogeochemical Cycles 1265
56.2 The Flow of Energy in Ecosystems 1271
Part 56.3 Trophic-Level Interactions 1276
56.4 Biodiversity and Ecosystem Stability 1280
Behavior 56.5 Island Biogeography 1283

53 Behavioral Biology 1185 57 The Biosphere and Human

53.1 The Natural History of Behavior 1186 Impacts 1288
53.2 Nerve Cells, Neurotransmitters, Hormones, and 57.1 Ecosystem Effects of Sun, Wind, and Water 1289
Behavior 1187 57.2 Earth’s Biomes 1293
53.3 Behavioral Genetics 1188 57.3 Freshwater Habitats 1296
53.4 Learning 1190 57.4 Marine Habitats 1299
53.5 The Development of Behavior 1191 57.5 Human Impacts on the Biosphere: Pollution and Resource
53.6 Animal Cognition 1194 Depletion 1303
53.7 Orientation and Migratory Behavior 1195 57.6 Human Impacts on the Biosphere: Climate
53.8 Animal Communication 1197 Change 1309
53.9 Behavior and Evolution 1200
53.10 Behavioral Ecology 1201
58 Conservation Biology 1317
53.11 Reproductive Strategies 1204 58.1 Overview of the Biodiversity Crisis 1318
53.12 Altruism 1206 58.2 The Value of Biodiversity 1322
53.13 The Evolution of Group Living and Animal 58.3 Factors Responsible for Extinction 1325
Societies 1210 58.4 An Evolutionary Perspective on the Biodiversity
Crisis 1335
54 Ecology of Individuals and 58.5 Approaches for Preserving Endangered Species and
Ecosystems 1338
Populations 1215
54.1 The Environmental Challenges 1216
Appendix A
54.2 Populations: Groups of a Single Species in One
Place 1218 Glossary G-1
54.3 Population Demography and Dynamics 1222 Index I-1

x Contents
Committed to Excellence
With the new 13th edition, Raven and Johnson’s Biology continues
the momentum built over the last five editions. We continue to pro­
vide an unmatched comprehensive text fully integrated with a con-
tinuously evolving, state-of-the-art digital environment. We remain
committed to our roots as the majors biology text that best inte-
grates evolution throughout the text. There is an emphasis on the
relevance of evolution throughout the ecology section, not only in
all four ecology chapters, but also in the chapters on behavioral
and conservation biology. In the animal form and function section,
we emphasize evolution in the context of physiology. We have also
moved the examples and insights from the chapter devoted to the
evolution of development, to place them into the appropriate con-
texts throughout the book. This emphasizes the importance of evo-
lution and develop­ment by continually providing examples, rather
than gathering them together in a single chapter. In the opening
molecular chapters, we have added additional examples of the
action of evolution at the molecular level.
We have also renewed our commitment to the ideas set forth
in the Vision and Change report from the AAAS, which provides a
framework for modern undergraduate biology education. This re­port
is now more than a decade old, yet still retains relevance. Perhaps the
most important idea articulated by Vision and Change was an
­emphasis on core concepts. This emphasis is important because it is
integral to how experts organize information in their brains. This
allows an expert to incorporate new information more readily than a
novice, who lacks an adequate conceptual framework.
We have designed a new Visual Outline for each chapter
opening and a more detailed Visual Summary at the end of each
chapter. These provide a visual representation of one way an expert
organizes the main concepts of a chapter, emphasizing connections
and hierarchies of concepts. These are intended not to be exhaustive
roadmaps of every chapter, but to show how important concepts for
each chapter can be organized and connected. The presentation will
appeal to students who are visual learners, and allow them to “see”
the chapter as a whole. We encourage students to create their own
visual maps of how the important concepts in each chapter are
interconnected.
One unanticipated consequence of the Vision and Change
movement was that publishers chasing new approaches began to
produce books so “feature-laden” as to be virtually unreadable by
the average student. This continues to be an issue with many text-
books on the market. We have not abandoned the idea that narra­
tive flow is important, even in a science textbook. While we
include a variety of features to improve student learning, they are
integrated into the text and are not included at the expense of the
concise, ac­cessible, and engaging writing style we are known for.
A priority of each revision is to assess each chapter for clarity and
readability. With each revision, we strive to extend ­ the clear
­emphasis on evolution and scientific inquiry that have made this a
leading textbook of choice for majors biology students.
Faculty want a textbook that emphasizes both student-­
centered ap­proaches and core concepts for the biological sciences.
As a team, we continually strive to improve the text by integrating
the latest cognitive and best practices research with methods that are
known to positively affect learning. We emphasize scientific ­inquiry,
and have increased the quantitative component in the ­Scientific
Thinking figures, as well as in the Inquiry and Data Analysis
­questions. Our text continues to be a leader with an organization that
emphasizes important biological concepts, while keeping the s­ tudent
engaged with learning outcomes that allow as­sessment of progress
in understanding these concepts. An inquiry-based approach with
robust, adaptive tools for discovery and assessment in both text and
digital resources provides the intellec­tual challenge needed to pro-
mote student critical thinking and en­sure academic success.
We continue to use our digital environment in the revision of
Biology. A major strength of both text and digital resources is assess­
ment across multiple levels of Bloom’s taxonomy that develops
critical-thinking and problem-solving skills in addition to com­
prehensive factual knowledge.
McGraw Hill Connect® offers a powerful suite of online tools
that are linked to the text and in­cludes quantitative assessment tools.
The Data and Graphing Interactive exercises have been expanded for
the 13th edition. This valuable digital tool uses data, controlled by
the students, to engage them in actively exploring quantitative
aspects of biology. Our adaptive SmartBook 2.0 learning system
helps students learn faster, study efficiently, and retain more knowl-
edge of key concepts.
The 13th edition continues to employ the aesthetically stun-
ning art program that the Raven and Johnson Biology text is known
for. Complex topics are represented clearly and suc­cinctly, helping
students build the mental models needed to understand biology.
We’re excited about the 13th edition of this quality textbook
providing a learning path for a new generation of students. All of
us have extensive experience teaching undergraduate biology, and
we’ve used this knowledge as a guide in producing a text that is up
to date, beautifully illustrated, and pedagogically sound for the
­stu­dent. We are also excited about the continually evolving digital
environment that provides a unique and engaging learning environ­
ment for modern students. We’ve worked hard to provide clear,
explicit learning outcomes that closely integrate the text with its
media support materials to provide instructors with an excellent
complement to their teaching.
Ken Mason, Jonathan Losos, Tod Duncan
xi
Cutting Edge Science
Changes to the 13th Edition
We authors started work on this new edition at the beginning of an
unprecedented pandemic that has affected the entire world. In addi-
tion to the personal and professional upheaval that all have experi-
enced, we were faced with the challenge of how to respond to the
pandemic in this new edition. Given that this pandemic is primarily
a biological phenomenon, we felt that it was important to include
accurate information, but not to let the book be overwhelmed by this
single event. So, we have included new material on the disease
COVID-19, the virus SARS-CoV-2, and the ways scientists have
analyzed and responded to this new pathogen. Rather than have a
single section on this, we have included material throughout the
book wherever it is relevant, including information about the virus
itself, diagnostic methods, vaccines, evolution, and the population
dynamics and environmental origin of epidemic diseases.
Part I: The Molecular Basis of Life
Chapter 1—Edited for clarity and readability for the student.
The Scientific Thinking figure was completely redone.
Chapter 2—Minor edits for clarity, especially regarding hydrogen
bonding and water.
Chapter 3—Minor edits for clarity, especially regarding the
structure of nucleic acids and proteins. Several figures were
corrected for inaccuracies or ambiguities. Section on the role of
trans fats was rewritten for clarity and currency.
Part II: Biology of the Cell
Chapter 4—The entire section on prokaryotic cell structure was
rewritten to take into account new data. This provides a different,
and more modern, view of prokaryotic cell structure. A new
figure comparing bacterial and archaeal flagella was added.
Chapter 5—Minor edits for clarity and readability for the student.
Chapter 6—Minor edits for clarity and readability for the
student. The speculative material on evolution of metabolism
was removed.
Chapter 7—Minor edits for clarity and readability for the student.
Chapter 8—Some editing for clarity. Also, the section on the
experimental history of photosynthesis was rewritten. This reduced
the amount of material while still providing students an historical
and experimental context for the rest of the chapter.
Chapter 9—Edited for clarity and to update some material for
currency.
Chapter 10—The section on chromosome structure was rewritten
to take into account new data on the organization of chromatin in
the nucleus. This complements the updates to the last edition, and
xii Committed to Excellence
includes a new figure showing the organization of chromosomes in
the nucleus.
Part III: Genetic and Molecular Biology
We reorganized the two chapters on transmission genetics to pro-
vide a more logical flow of concepts, and to emphasize modern
human genetics.
Chapter 11—Edited for clarity and readability for the student.
Chapter 12—This chapter was entirely rewritten to bring a more
modern focus. The history was retained to provide context for the
entire chapter. Mendel’s model is presented in a clearer, more
modern form. The extensions to Mendel were reorganized and
also given a more modern perspective. Two new figures were
added to replace older figures.
Chapter 13—This chapter was completely rewritten to form a
more cohesive whole with chapter 12, on Mendelian genetics. It
begins with an updated treatment of the chromosomal theory of
inheritance, which complements and extends chapter 12. The
remainder of the chapter is devoted to a much more modern view
of human genetics. Material on human genetics from the old
chapter 12 was moved to this chapter and substantial additions
were included to keep pace with the rapid pace of change in
human genetics.
Chapter 14—The material on DNA structure was rewritten to
make some difficult concepts more clear to students.
Chapter 15—The material on alternative splicing was updated.
The section on mutation was completely rewritten to take new
data into account, and to include current information on genetic
variation.
Chapter 16—The chapter was edited for clarity and currency.
Chapter 17—The chapter was revised to include new and relevant
technologies highlighted by the recent pandemic of COVID-19.
There is a description of PCR-based tests for pathogens, and
updated material on antibody-based tests to detect pathogens and
exposure to pathogens. There is also a section on new vaccine
technologies, including mRNA and subunit vaccines.
Chapter 18—The sections on genome projects and annotation
were both rewritten to take new data into account. These provide
as current a view of these very fast-moving areas of biology as is
possible for an introductory text.
Chapter 19—Minor edits for clarity and readability for some
difficult concepts.
Part IV: Evolution
Chapter 20—The section on genetic variation in populations was
extensively revised reflecting new information based on wide-
spread genomic investigation. Genomic variation in humans is
now discussed in great detail, quantifying the extent of variation
that exists and how that variation is apportioned within and
between populations.
Chapter 22—New information on the genes involved in the
evolution of the beaks of Darwin’s finches was included.
Chapter 23—Discussion on the evolution of the HIV virus was
revised to include new discoveries and trends. An entirely new
section was added detailing how phylogenetic analysis has helped
track the evolution and spread of the virus causing COVID-19.
The drawing of Archaeopteryx was revised to correspond to
new analyses of fossils indicating that these early birds were dark
in color.
Chapter 24—The comparative genomics section was rewritten
to emphasize the evolution of the vertebrate genome. The
material on primate genome evolution was updated to reflect
new data.
Part V: Diversity of Life on Earth
Chapter 25—The material on supergroups was updated, including
a new figure.
Chapter 26—A new section was added to provide students with
contemporary information about the coronaviruses. Particular
emphasis was placed on SARS-CoV-2, the causative agent of the
COVID-19 pandemic. A new figure was added to help explain
the infectious cycle of coronaviruses in general. The chapter was
edited for readability.
Chapter 27—This chapter was edited to improve readability.
Epidemiological data on infectious diseases was updated to
reflect changes since the 12th edition.
Chapter 28—This chapter was completely restructured and large
parts were rewritten to reflect current thinking in the taxonomy
of the protists. Additional edits were made for readability and
comprehension.
Chapter 31—Edited for clarity and readability.
Chapter 32—Discussion of relationships at the base of the
phylogeny for all animals was revised to reflect new understand-
ing and debate about relationships among sponges, ctenophores,
and other animals. Additional changes were made to reflect other
changes in understanding of phylogenetic relationships among
animal taxa, such as the position of chaetognaths. Aspects of
taxonomy and natural history were updated in line with new
findings.
Chapter 33—This chapter was revised and reorganized to reflect
new understanding of relationships among protostome taxa.
Chapter 34—Discussion of human evolution was revised in light
of new discoveries.
Part VI: Plant Form and Function
There have been no major changes in the plant form and function
chapters. There has been overall editing for readability and in response
to recommendations by reviewers and users of the 12th edition.
Part VII: Animal Form and Function
Chapter 41—Minor edits for clarity in the material on homeosta-
sis and thermoregulation.
Chapter 42—Minor edits for clarity and accuracy.
Chapter 43—This chapter was reorganized to reflect the relation-
ship among different types of receptors. The depiction of the
evolution of the inner ear was revised to illustrate the transition
from reptilian to mammalian hearing structures. Treatment of the
phylogenetic analysis of the evolution of genes related to eye
structure was modified in light of new discoveries.
Chapter 44—Minor edits for clarity and readability in material
on receptor function.
Chapter 45—Illustration of convergent evolution in skeletal
structure of birds, bats, and pterosaurs was revised.
Chapter 47—Discussion of external gills and respiration in
salamanders was revised to reflect the variation in structures
found within this order.
Chapter 50—Much of the chapter was rewritten to improve the
clarity and flow of ideas and emphasize connections between
innate and adaptive immunity. A new section on vaccination was
added to complement the new section in chapter 17 on vaccine
technology.
Chapter 51—Discussion of variation in reproductive systems
among animals was revised with regard to parthenogenesis and
reproductive mode.
Chapter 52—Edited for clarity and readability of difficult
concepts.
Part VIII: Ecology and Behavior
Chapter 53—Discussions of behavioral genetics, migration, and
other topics were revised to incorporate new findings on the
genetic basis of behavior.
Chapter 54—A new section was added on pandemics and human
health that covers the general topic, but extensively details the
population biology of the COVID-19 pandemic. Human popula-
tion trends and other timely data were updated to stay current.
Chapter 57—The section on deforestation was greatly updated
with new information and trends, and a section on the effect of
massive wildfires was added. Discussion of zoonotic diseases
was increased, with emphasis on COVID-19. New discussion of
Committed to Excellence xiii
microplastics in the environment was added. The section on El
Niño events was revised. All of the data on biosphere impacts of
humans were updated to stay current.
Chapter 58—Figures on the rate at which species are becoming
endangered were updated and a section on the new-found data on
decline in insects was added. Data on the relationship between
human population growth and biodiversity decline was updated,
as was information on the rate of establishment of invasive
species. Information on case studies throughout the chapter was
also updated to reflect the current situation.
A Note From the Authors
A revision of this scope relies on the talents and efforts of many
people working behind the scenes and we have benefited greatly
from their assistance.
Beth Bulger was the copy editor for this edition. She has
­labored many hours and always improves the clarity and consis-
tency of the text. She has made significant contributions to the
quality of the final product.
We were fortunate to work again with MPS to update the art
program and improve the layout of the pages. Our close collabora-
tion resulted in a text that is pedagogically effective as well as
more beautiful than any other biology text on the market.
In the end, the people we owe the most are the generations
of students who have used the many editions of this text. They
have taught us at least as much as we have taught them, and their
questions and suggestions continue to improve the text and sup-
plementary materials.
Finally, we need to thank instructors from across the country
who are continually sharing their knowledge and experience with
us through market feedback and symposia. The feedback we
­received shaped this edition. All of these people took time to share
their ideas and opinions to help us build a better edition of Biology
for the next generation of introductory biology students, and they
have our heartfelt thanks.
Reviewers for Biology, 13th edition
Beth Cliffel Triton College
David Cox Lincoln Land Community College
Sarah Cuccinello University of Tampa
Stella Doyungan Texas A&M University–Corpus Christi
Sherry L. Harrel Eastern Kentucky University
Christopher Ivey California State University, Chico
Lori Kayes Oregon State University
Nathan Lanning California State University, Los Angeles
Karen Neal Reynolds Community College
Bruce Stallsmith University of Alabama
Shalini Upadhyaya Reynolds Community College
Tom Warren Snead State Community College
Jennifer Whitt Texas A&M University–Corpus Christi

We have the continued support of an excellent team at

McGraw Hill Education. Ian Townsend, Portfolio Manager for
Biology, has been a steady leader during a time of change. Senior
Product Developer Liz Sievers, provided support in so many
ways it would be impossible to name them all. Kelly Hart,
Content Project Manager, and David Hash, Designer, ensured
our text was on time and elegantly designed. Kelly Brown,
Senior Marketing Manager, is always a sounding board for more
than just marketing, and many more people behind the scenes
have all contributed to the success of our text. This includes
Sarah Hosch and Elizabeth Drumm who greatly helped develop
the new Visual Outlines and Visual Summaries, and the digital
team, to whom we owe a great deal for their efforts to continue
improving our Connect assessment tools. The digital Subject
Matter Experts for this edition include: Faye Nourollahi, Austin
Dreyer, Cynthia H. Dadmun, Timothy Hadlock, Elizabeth
Drumm, and Lauri Carey.
Throughout this edition we have had the support of spouses
and families, who have seen less of us than they might have liked
because of the pressures of getting this revision completed. They
have adapted to the many hours this book draws us away from
them, and, even more than us, looked forward to its completion.

xiv Committed to Excellence

Preparing Students for the Future

Developing Critical Thinking with the Help of . . .

Scientific Thinking Figures
Key illustrations in every chapter highlight how the frontiers of
knowledge are pushed forward by a combination of hypothesis and
experimentation. These figures begin with a hypothesis, then show
how it makes explicit predictions, tests these by experiment, and
finally demonstrates what conclusions can be drawn, and where
this leads. Scientific Thinking figures provide a consistent frame-
work to guide the student in the logic of scientific inquiry. Each
illustration concludes with open-ended questions to promote sci-
entific inquiry.
Data Analysis Questions
It’s not enough that students learn concepts and memorize scientific
facts; a biologist needs to analyze data and apply that knowledge.
Data Analysis questions inserted throughout the text challenge
students to analyze data and interpret experimental results, which
shows a deeper level of understanding.
Inquiry Questions
These questions challenge students to think about and engage in
what they are reading at a more sophisticated level.

SCIENTIFIC THINKING 32

Hypothesis: The plasma membrane is fluid, not rigid. Temperature (°C) 30

Prediction: If the membrane is fluid, membrane proteins may
diffuse laterally.
Body

28
Test: Fuse mouse and human cells, then observe the distribution
of membrane proteins over time by labeling specific mouse and 26
human proteins. open habitat
24 shaded forest
Human
cell 24 26 28 30 32
Air Temperature (°C)

Figure 54.3 Behavioral adaptation. In open habitats, the

Mouse
cell Intermixed Puerto Rican crested lizard, Anolis cristatellus, maintains a relatively
Fuse membrane proteins constant temperature by seeking out and basking in patches of
cells
sunlight; as a result, it can maintain a relatively high temperature even
when the air is cool. In contrast, in shaded forests, this behavior is not
possible, and the lizard’s body temperature conforms to that of its
surroundings.
Allow time for (inset) Jonathan Losos
mixing to occur

? Inquiry question When given the opportunity, lizards

Result: Over time, hybrid cells show increasingly intermixed proteins.
Conclusion: At least some membrane proteins can diffuse laterally in
the membrane.
Further Experiments: Can you think of any other explanation for
these observations? What if newly synthesized proteins were inserted
into the membrane during the experiment? How could you use this
basic experimental design to rule out this or other possible explanations?
Figure 5.6 Test of membrane fluidity.
regulate their body temperature to maintain a temperature
optimal for physiological functioning. Would lizards in open
habitats exhibit different escape behaviors from those of
lizards in shaded forest?
Data analysis Can the slope of the line tell us something
about the behavior of the lizard?

xv
Visual Maps
There are two new but related features in Biology, 13th edition
that help students build a conceptual framework into which they
can insert new knowledge. Every chapter contains a Visual
Outline at the beginning of the chapter and a more detailed Visual
Summary at the end of the chapter. These complementary features
serve as conceptual bookends to help students place chapter topics
into a conceptual framework rather than a linear list. When topics
are placed in the context of a map, students see relationships and
connections between topics that are not apparent in the standard
chapter outline and chapter review formats.
Visual Outline
Every chapter contains a list of chapter contents but now also
contains a Visual Outline that shows topics in relation to each
other. The Visual Outlines differ from “concept maps” in that
there is a hierarchical aspect because they present a conceptual
framework for the chapter. In looking at the Visual Outline
before reading a chapter, students will be familiar with key
terminology and how key topics generally relate to each other.
Images are placed in the Visual Outline as a graphic connection
to figures in the chapter.

Fluid Cellular
mediate Transport
mosaic model membranes

3 types
Consists
of

Passive Active Bulk

transport moves moves transport transport
down up
Phospholipid
bilayer 3 types Concentration
gradient
types
Transmembrane Simple
proteins diffusion
Requires
energy Endocytosis
Cytoskeleton Osmosis

Exocytosis
Cell surface Facilitated
markers diffusion ATP

Dr. Edwin P. Ewing, Jr./CDC

Visual Summary
As an extension of the Visual Outline, a Visual Summary is
presented at the end of all chapters following the more traditional
textual summary. Like the chapter opening feature, the Visual
Summary presents the chapter topics in relation to each other and
expands on the Visual Outline with supporting details of the key
topics. The Visual Summary doesn’t contain all material in the
chapter, nor does it indicate all connections between topics, but
presents a conceptual scaffold. Students should be encouraged to
build upon the scaffold with additional information from the chapter.

xvi Preparing Students for the Future

 Fluid Cellular
mediate Transport
mosaic model membranes

Glycerol, 2 fatty acids, 3 types

Consists of phosphate
made of
Forms
Passive move Concentration move Active Bulk
spontaneously
Phospholipid transport down gradient up transport transport
bilayer
3 types types
Fluid defined as
Simple example
Transmembrane diffusion Concentration Movement
include Transporters into the cell
proteins difference
Sodium-
Osmosis
Enzymes requires potassium
Cytoskeleton Movement pump
Endocytosis
Receptors Facilitated of water
diffusion is specific
Cell surface
markers Identity Carrier
markers is proteins Energy Receptor-
mediated
types Adhesion endocytosis
Function Selective uses Carrier
proteins
proteins
Cytoskeleton Channel
Cell proteins
attachment Exocytosis
recognition

Glycoproteins
Movement
Glycolipids out of the
cell

Preparing Students for the Future xvii

Strengthen Problem-Solving Skills with Connect®
Detailed Feedback in Connect®
Learning is a process of iterative development, of making
mistakes, reflecting, and adjusting over time. The question and
test banks in Connect® for Biology, 13th edition, are more than
direct assessments; they are self-contained learning experiences
that systematically build student learning over time.
For many students, choosing the right answer is not
necessarily based on applying content correctly; it is more a
matter of increasing their statistical odds of guessing. A major
fault with this approach is that students don’t learn how to
process the questions correctly, mostly because they are
repeating and reinforcing their mistakes rather than reflecting
and learning from them. To help students develop problem-
solving skills, all higher-level Blooms questions in Connect are
supported with hints, to help students focus on important
information for answering the questions, and detailed feedback
that walks students through the problem-solving process, using
Socratic questions in a decision-tree-style framework to
xviii Preparing Students for the Future
scaffold learning, where each step models and reinforces the
learning process.
The feedback for each higher-level Blooms question
(Apply, Analyze, Evaluate) follows a similar process: Clarify
Question, Gather Content, Choose Answer, Reflect on Process.
Unpacking the Concepts
We’ve taken problem solving a step further. In each chapter, three
to five higher-level Blooms questions in the question and test
banks are broken out by the steps of the detailed feedback. Rather
than leaving it up to the student to work through the detailed
feedback, a second version of the question is presented in a
stepwise format. Following the problem-solving steps, students
need to answer questions about earlier steps, such as “What is the
key concept addressed by the question?” before proceeding to
answer the question. Found under the Coursewide Content section,
a professor can choose which version of the question to include in
the assignment based on the problem-solving skills of the students.
Data and Graphing Interactives
To help students develop analytical skills, Connect® for Biology,
13th edition, is enhanced with Data and Graphing Interactives
also found under the Coursewide Content. Students are
McGraw Hill
Virtual Labs and
Lab Simulations
While the biological sciences are hands-on disciplines, instructors
are now often being asked to deliver some of their lab components
online, as full online replacements, supplements to prepare for
in-person labs, or make-up labs.
These simulations help each student learn the practical and
conceptual skills needed, then check for understanding and
presented with a scientific problem and the opportunity to
manipulate variables in the interactive, producing different
results. A series of questions follows the activity to assess
whether the student understands and is able to interpret the
data and results.
provide feedback. With adaptive pre-lab assignment, found under
Adaptive Learning Assignment, and post-lab assessment avail-
able under Coursewide Content, instructors can customize each
assignment.
From the instructor’s perspective, these simulations may be
used in the lecture environment to help students visualize complex
scientific processes, such as DNA technology or Gram staining,
while at the same time providing a valuable connection between
the lecture and lab environments.
Preparing Students for the Future xix
Another random document with
no related content on Scribd:
much confidence in the people. I am convinced, however, that were
a representative of the Dutch Government to reside here, it would
soon become a highly prosperous place, and the harbour would
never be without vessels in it.
I cannot avoid giving a short description of the Pilandok or Arru
rabbits, an animal rather larger than the common rabbit, of a grey
colour, which, as they grow old, becomes quite grizzly. The forelegs
are short, and the hind ones, which resemble those of the hare, have
each three toes, provided with strong nails. The head is like that of a
weazel. These animals do not run very fast, and when resting they
usually sit upright on their hind legs. Their food consists of the leaves
of the yam plant and other greens, and they are easily tamed, when
they may be suffered to run around the house without their
attempting to escape. The flavour of their flesh is very agreeable.
On the 1st of September, when at Duryella, I appointed the Orang
Kayas to the islands of Wama and Maykor, to each of whom I
presented a silver-knobbed baton, besides other government gifts,
as pieces of cloth, flags, &c. Several of the chiefs of the more distant
islands were also confirmed in their authority on the same occasion,
many other affairs being also settled; among which was a
reconciliation between the people of two villages on Vorkay, who for
some time previously had lived in state of war with each other. The
quarrel originated in one of the parties fishing too long on a reef
which was their mutual property, and thus obtaining too large a share
of its produce. It had already been provisionally adjusted through the
mediation of the Christian chiefs, and was now submitted to me for
my final decision. The offenders solemnly promised that they would
not hereafter exceed their right, and the affair was settled to the
satisfaction of both parties, who declared that they would
henceforward live in peace and friendship with each other.
The greater portion of the Christian inhabitants of Maykor had come
to Duryella, partly to attend to their own interests, and partly for the
purpose of being present at the preachings of M. Kam, on which
occasions a large number of people were admitted as members of
the Christian Church. I occasionally invited the chiefs to dinner with
me, which was always considered by them as a great honour, and
they never failed to come attended by a numerous retinue, Arafuras
as well as Christians, who crept on board from all sides in an instant.
The chiefs of the more distant islands, not having been informed of
my arrival, had come in their usual dress, the tdejako, or waist-cloth,
as they never adorn themselves with their best clothing except on
festive occasions. These chiefs were much ashamed of their
deshabille, and made repeated excuses; but being informed of the
cause, I rejoiced them by having kabayas and breeches prepared for
them, when they made their appearance with much greater
confidence.
On one occasion a number of chiefs, who had dined on board,
remained to spend the evening with me, many of whom, particularly
the Arafuras, declared to me that they had never before been on
board a ship of war. To give them a good idea of such a vessel I
ordered several of the great guns to be fired, and although we had
informed them of what was coming, they were terribly frightened,
and it was truly ridiculous to see them jump into the air at every shot,
like harlequins. Many of them, indeed, jumped overboard, while
others hid themselves under the table which stood on the deck, and
in any corner they could find. These islanders generally can carry off
an inconceivable quantity of spirits. Although at our festive meetings
on board more than thirty jugs of arrack were often drunk, they would
still take a quantity on shore with them to finish the evening at home.
The prahus which are built upon these islands are very long, and sit
low on the water, their length being seldom less than seventy feet,
while their greatest breadth is not greater than ten feet. We saw
about thirty of these lying under the shore, and had good
opportunities for examining them. In the centre of them is a stage, on
which they make a fire to cook their provisions. Whenever these
vessels sail on a voyage they are adorned with a number of flags,
some being displayed from the stern and others planted along the
sides, the whole presenting a very gay appearance. These flags are
triangular (perhaps in imitation of those of the Chinese), and of
various colours, the prahus belonging to the chiefs being
distinguished by the superior size and number of those they display.
The inhabitants of the western Arrus obtain their prahus chiefly from
the Ki Islands, these being stronger and broader than those
constructed by the people of the Eastern Arrus. The Arafuras, when
at sea, protect themselves from the rain, which they dislike greatly,
by means of long bags open on one side, made from the bark of a
tree. When the weather is fine, they roll these up and use them as a
cushion to sit upon.
The abolition of the slave trade has not yet spread its beneficent
influence to these parts; indeed, the export and import of slaves
forms a considerable branch of commerce, &c.: so much so, that
were it to be discontinued many of the inhabitants would become a
prey to want. It must here be taken into consideration, that the words
"slave" and "slave trade" have a very different signification here from
that which they have in the West Indies, the people of the east
regarding the slave trade as being, in a certain point of view,
beneficial to the slaves themselves. Many, in fact, when in distress,
offer themselves as bondsmen that they may obtain the necessaries
of life. The natives, however, are well aware that this shameful trade
is much disapproved of by our Government, and that the greatest
object of their rule is to secure general freedom and prosperity to
mankind, for which reason the slaveholders treat their people with
humanity. These creatures are, therefore, less to be pitied than one
would suppose; and it must be taken into consideration, that so long
as agriculture and industry are so much neglected, the slave trade
must be a source of subsistence to a great part of these islanders,
by means of which they are held back from engaging in piracy, theft,
and other vices, to which want and necessity would otherwise force
them to resort.[32]
When M. Bik was here he was presented with two slave boys, to
whom, as I was informed by the natives, he gave their liberty, and
caused them to be sent back to their village. Shortly after the
departure of this gentleman these liberated slaves fell into a state of
great poverty, so that, to pay the debts they had incurred for
provisions and necessaries, they had again pledged or sold
themselves as slaves, and they now regard themselves as fortunate
in having found a good master. On my asking them if they wished to
become free again? they answered, that they would like it well if I
would take them with me, and provide for their future maintenance,
but that otherwise they would prefer living in their present state of
servitude. I allowed my interpreter to redeem a couple of orphans
who had been sold here by the Arafuras, and did not seem to have
derived much good from their master, for which he paid eighty
guilders. At their own request I took them with me to Amboyna,
where, with the cognizance of the Government, I set them at liberty,
so that they became in a condition to gain their livelihood by daily
labour.[33]
On the 4th of September the chiefs of Maykor and Wama came to
take their leave, and to present us with some stock and fruit, after
which we weighed anchor and steered for the island Wadia. A
number of prahus accompanied us a considerable distance, the
natives in them displaying their friendship by the music of their gongs
and songs of joy.

FOOTNOTES:
[31] So early as 1640, F. Corsten entered into a treaty with the
people of the Arrus, especially those of Wokan, Wadia, Wama,
Duryella and Maykor, in which the latter agreed to acknowledge
the supremacy of the Dutch East India Company. Afterwards, in
1645, the Fiscal Dorstman obtained from them an agreement to
trade exclusively with Banda, while the monopoly of the pearl
fishery was given to the East India Company.
[32] M. Kolff, whose opinion upon this point is totally at variance
with that of Sir Stamford Raffles, and of every other writer on the
Indian Archipelago, who has really taken an interest in the welfare
of the natives, appears to have mistaken effect for cause; as in
this part of the world it has invariably been found that the slave
trade alone has been sufficient to render a people, previously mild
and industrious, poor, idle and vicious. As to the slave trade
preventing the natives from engaging in piracy, M. Kolff must have
been aware that by far the greater number of piratical expeditions
fitted out by the natives are intended solely for the capture of
slaves, other plunder being in their eyes of minor importance only;
and this is especially the case in the parts visited by M. Kolff,
where the inhabitants possess nothing of sufficient value to tempt
the cupidity of pirates.—Ed.
[33] The inhabitants of the Arrus and the adjacent island, are glad
to obtain Dutch gold and silver coin, the greater part of which they
work up into ear-rings and other ornaments.
 CHAPTER XIII.
THE ARRU ISLANDS.—THE TENIMBER ISLANDS.
Arrival at the Island Wadia.—Particulars concerning the
Island and its Inhabitants.—Dispute between them and the
Orang Tua of Fannabel.—Sad Result of their Contentions.
—Departure from the Arru Islands.—Arrival at the
Tenimber Group.—Vordate.—Ignorance and Perplexity of
the Pilot.—Singular Customs.—Violent Conduct of the
People of Timor-Laut.—The Inhabitants of Watidal and
their Chiefs.
We soon entered the narrow strait which separates the islands of
Wassia and Wadia, the last of which was the object of our visit. The
people of Wakan, Samau, and the other villages on the sea shore
saluting us with shots from their lelahs, which were answered by the
brig with several guns. Wassia, the north-westernmost of the Arrus,
is overgrown with underwood, and the coral reefs extend some
distance from it on all sides, preventing vessels from approaching
the shore. The village of Wadia is situated on the west side of the
island of that name, on the banks of a salt-water creek, several other
villages being scattered over the island.
As soon as we had anchored in the strait, in ten fathoms sand, about
four miles from the village of Wadia, the Orang Tuas of the island
came on board in two prahus, to bid us welcome. My interpreter
accompanied them on shore, to inform the chiefs that I should land
the following day to transact the business which had brought me
here. With a view to add as much solemnity as possible to my visit, I
sent twenty-six armed men on shore early in the morning, and soon
after followed them with the gentlemen attached to the expedition.
After entering the creek we passed a temple and a number of tombs,
and soon arrived at the village, which lies about a cable's length and
a half from the mouth. The houses, which stand separated from each
other, are erected on both sides of the river, but by far the greater
number are to be found on the right-hand side, the dwelling of the
Orang Kaya forming a conspicuous object among them.
On our approach to the river we were saluted with seven shots from
a one-pounder gun, which had been taken on shore from the brig,
while our men were drawn up on the beach to receive us with
military honours, the natives themselves saluting us with numerous
shots from their lelahs, and displaying several Dutch flags, which
appeared to be of their own manufacture. The Orang Kaya
welcomed us to his house with many expressions of friendship,
when we found that one-half of his dwelling, (a new and rather large
building, similar to that of the Orang Kaya of Duryella), had been
arranged for our reception, another house having been appropriated
for the accommodation of our men. As the house of the Orang Kaya
is usually the place where the chiefs assemble, we thought it most
advisable to take up our quarters in a small building which had been
occupied by the foreign traders, but was now empty.
The population of Wadia is composed of Mohammedans, so that I
did not perceive among the chiefs the same affection towards the
Netherland's Government which those of the Christian islands
displayed. On conversing with them concerning the non-fulfillment of
the promise they made to M. Bik in 1824, to resort to Banda, I clearly
discovered that the Macassar traders had done all in their power to
make them fearful of us. These had made them believe that the sole
object of our Government consisted in increasing their revenue from
imposts, and many similar absurdities, which it cost me considerable
difficulty to drive from their minds. They also asserted, that owing to
the small amount of the population they dared not leave their villages
for a lengthened period, unless the Netherlands' Government should
have a post upon the Arru Islands.
This request for a Dutch Resident or Postholder, with a small
garrison, was made to me on all the islands, and the people
appeared to be well aware that it would contribute greatly to
maintaining order among them, and increasing their prosperity.
Christians, Mahommedans and Arafuras, all perfectly agreed in this
particular.
The creek on which this village is situated, runs right through the
island. Both banks are low and marshy, and the depth of water in it is
very small; indeed, many parts are quite dry at low water. It is
abundantly stocked with fish, which, in fact, is the case everywhere
among the group. Live stock, on the other hand, is very scarce: pigs
are only kept by the Arafuras, the Mohammedans contenting
themselves with goats and fowls, which last are to be purchased at
the rate of a rupee each. With the exception of these there are no
tame stock on the islands. Wild hogs are abundant, but they are
rarely hunted by the natives.
The pilandok is also to be met with, together with birds-of-paradise,
and immense numbers of cockatoos, red, green and black loories,
and other sorts of handsomely-plumaged birds. The Arafuras shoot
the birds-of-paradise with arrows, when they are opened, and
deprived of their entrails, and then wrapped in a thin leaf and hung to
dry in the smoke of their fires; this being the only preparation they
require to render them ready for exportation.
Sago and yams form the only vegetable productions which the island
affords. Rice is imported by the Macassars, and by the people of the
Ki and Goram Islands, who, on their part, obtain them from Banda.
The people have but few wants. Their dwellings are mean, and their
sole property consists in slaves, muskets, lelahs, gongs and
elephants' tusks. The first are obtained chiefly from New Guinea,
and from the back parts of the group.
The people of Wadia, and of the western islands of the Arru group in
general, are very desirous of obtaining the imitation blue, red and
black calicoes,[34] especially the last sort manufactured in the
Netherlands. Square checked, or closely flowered chintzes, are also
much in demand. These goods, however, would be too fine for the
Arafuras of the back of the group, who only care for those of coarser
materials, such as coarse chintzes, blue and white salempoors, red
coast-calicoes, red baftas, Samanap sarongs, axes, parangs,
chopping knives, brass wire, coarse Chinese porcelain, arrack,
aniseed, gongs, elephants' tusks, small red beads and other trifles of
a similar description.
The inhabitants of the western parts of the group carry on a constant
bartering trade among the group; the long and light vessels
previously described, which are from twelve to sixteen tons burthen,
being employed in transporting the goods from island to island. The
period which elapses between the cessation of one monsoon and
the commencement of the other, is the fittest period for these
voyages. They are fearful, however, of going to Banda or Amboyna,
not only from the reasons already given, but from a dread that the
Government would chastise them for some deeds they had
committed. I endeavoured, as much as lay in my power, to banish
this idea from their minds, promising them forgiveness on the part of
the Government; and my exertions were attended with so much
success, that during the following year, while I was remaining at
Amboyna, a greater number of their vessels arrived at that place,
and also at Banda, than had been the case previously.
The inhabitants of the villages on Wadia profess the Mohammedan
religion. In point of numbers they are far inferior to the Christians of
the neighbouring islands, with whom they live on very good terms,
and like them consider themselves as subjects to the Dutch
Government, although they appear to be more restless in their
characters than the others. A difference had long existed between
the people of Wadia and the Orang Tua of Fannabel; and the Orang
Kaya of Wokan, with the minor chiefs of that island, had followed me
hither to assist in inquiring into and deciding the dispute.
After I had made known to the inhabitants the philanthropic and
humane object of our Government, they gave me many proofs of
their wishes to pursue a proper course. I was subsequently
petitioned to examine into and decide the affair of the Orang Tua of
Fannabel, which, from the evidence given by both parties, appeared
to be as follows:—
The Orang Tua of Fannabel possessed a stone building, defended
by cannon, on the north-east side of Wokan, and being a skilful
trader he soon increased his property until he became one of the
wealthiest men in the islands. This aroused the envy of the people of
Wadia, of which island the Orang Tua had formerly been an
inhabitant. On the occasion of the marriage of one of his daughters,
there arose a difference between a native of Fannabel and another
of Wadia, of so violent a nature, that each threatened to take the life
of the other on the first opportunity. Some time after this, a number of
the people of Wadia visited Wokan to assist the Orang Kaya in the
construction of a church, and on their return home, one of the parties
engaged in the above-mentioned quarrel, with four others, were
murdered in their prahu. The offenders, who were soon after taken,
declared that they had been paid for committing the deed by the
people of Fannabel, on which those of Wadia, without enquiring
farther into the matter, went to the house of the Orang Tua of
Fannabel during his absence, and by way of taking revenge for the
murder, they shot several of his family, took away the remainder as
slaves, and then plundered and burned the house. The Orang Tua,
on his return to his home, seeing the destruction that had taken
place, fled to Wokan, where he was received with hospitality. The
Orang Kaya of that place, however, considering that he was not
sufficiently powerful to obtain redress for his friend by force, and
being also unwilling to go to war with the people of Wadia,
commenced negotiations, by which means a portion of the stolen
property was returned.
This affair had taken place four years previous to my visit. The ill-
treated Orang Tua assured me that he often thought of visiting
Banda to obtain redress through the interposition of the Government,
but had always been withheld by the hope that the latter would again
take possession of the Arru Islands, in which case he would have
brought the case before them, although he was himself aware that it
would be difficult, if not impossible, to obtain back all the goods of
which he had been plundered.
At the assembly which had been called together strong debates took
place, which lasted until four o'clock. Some of the arguments put
forth will tend to show their erroneous ideas of right and equity. A
man who had carried off three of the Orang Tua's slaves, was
unwilling to return more than one, or at most two, wishing to keep the
remainder to pay for the maintenance of the whole. I explained to
another, that as there was not the least proof of any offence on the
part of the Orang Tua of Fannabel (they themselves holding him
guiltless of the murder which had been committed,) while he had
already suffered severely from the loss of his son, and of all his
goods, it was nothing but just that he should have every thing
possible restored to him. "Here," answered he, "are four of our family
slain in their prahu; we must therefore have redress from somebody,
no matter whom." At length it was determined, that a portion of the
goods should be restored, and several gongs, elephants' tusks and
slaves were immediately given up.
After I had thus changed the inveterate hatred into friendship, and
had settled some other differences, I appointed a certain Abdul
Wahab as their Orang Kaya, to the universal satisfaction of the
people, and ceremoniously installed him in his office.
During our stay here I examined the neighbourhood of the village,
and met with some former strongholds, the remains of which proved
that they must have been extensive. We also found the traces of a
long street, enclosed by walls, running east and west through the
village, together with the ruins of many stone houses. The natives
could give us no clear information concerning them.
On the 8th of September we made ready for our departure. When
our people were on the point of embarking in the boats, the above-
mentioned Orang Tua of Fannabel came to me to complain that the
four slaves had not yet been given up to him, and that if we departed
before this took place he would never get them back. I therefore
caused it to be announced to the defaulter, that if he did not perform
his contract I would take the slaves from his house by force, on
which they were immediately given up. After having impressed him
with a sense of the injustice and impropriety of conduct he had been
guilty of, I caused my interpreter to draw up, in Arab characters, a
solemn agreement of mutual reconciliation and friendship, which was
signed by both parties, who promised in my presence that they
would henceforward live together in amity.
The Orang Kaya, with all his chiefs, accompanied us on board, and
took a friendly farewell of us, after having received some presents of
coffee and arrack. On the same day we left our anchorage off Wadia,
and stood to the southward under press of sail, towards the
Tenimber Islands.
The day after our departure we sighted the island Vordate to the
south-west. From a distance of about twenty miles on this side, this
island appears alternately hilly and flat, like irregular mountains,
while the neighbouring islands, Larrat, Timor-laut, &c., have a more
level appearance. Several reefs extend a considerable distance to
seaward from the south-east end of the island.
We continued during the night under easy sail, and the following
morning, under the guidance of the pilot, we passed along the west
side of Vordate, at a distance of about six miles from the shore. Off
the north point, a mile to seaward, lies a rock, for which reason the
point must not be made free with. The villages, in which were
displayed a number of small flags, alternating with the fertile valleys,
presented an alluring picture to us. As we passed a number of
prahus came rowing towards us, the people in which called out to us
to anchor off their village. The pilot, however, affirming that there was
good anchorage near Larrat, we continued our course without
accepting the invitation of the islanders. We crossed the extensive
reef of Larrat in irregular depths of three, four and five fathoms, and
ran along the north-west coast towards the mouth of a bay, which is
formed between the island of Larrat and the point of Timor-laut.
On approaching the entrance of the bay, I clearly perceived that
there was no possibility of the brig being able to enter. On perceiving
this, the pilot, as is usual with natives under similar circumstances,
became completely cast down, and all that I could learn from him
was that he had never visited the island before except during the
westerly monsoon, and then only in small vessels which they could
row over the reefs. This was all very true; but had he only given me
the slightest idea of the situation of the bay, I should certainly not
have come here; but it was now necessary to make a virtue of
necessity, and come to a speedy determination. The wind blew
directly out of the mouth of the bay, which was very narrow, so that
there was no possibility of working in. We could obtain no bottom
with the deep-sea lead and one hundred fathoms of line, while we
were everywhere surrounded by shoals. I therefore sent the boats to
lie on the edge of the coral reef which fronts the village of Watidal,
and then stood on, hoping that our anchor might hook a rock and
hold fast; but, although the anchor was let go in four fathoms water, it
had dragged into twelve fathoms before the ship was brought up,
while under the stern there was thirty-eight fathoms. A kedge was
then carried out astern, to prevent the tide from swinging the ship on
to the reef.
Before the sails could be furled, or the boarding nettings spread, the
brig was surrounded on all sides by a number of canoes, and I had
no sooner given permission to a couple of natives to come on board,
than the others all followed, so that, in a less space of time than five
minutes, the brig was full of men, who had crept on board through
the ports and every place where they could find admission. With a
view to inspire them with confidence I permitted them to enter, and
received them in as friendly a manner as possible, the distribution of
some glasses of arrack among them contributing not a little to
produce the desired effect.
The village off which we had anchored, like all on these islands, was
erected on a hill of limestone rock, near the sea shore. These
villages present a very picturesque appearance: they consist,
according to their population, of from twenty to fifty houses, erected
near each other, upon piles from six to eight feet high. These
dwellings are from twenty to twenty-five feet long by twelve to fifteen
broad. They are enclosed on all sides, and have a couple of long
holes cut in the walls to serve for windows. The roofs are covered
with a thatch of palm leaves, these being first arranged on small
sticks, and then placed neatly on the roof, overlapping each other.
The interior is usually kept in good order; but every part is blackened
with smoke from the fires they employ in cooking their provisions.
The house is entered by a door in the centre of the floor, to which the
inhabitants ascend by means of a ladder. Against the wainscoat,
immediately fronting the door, is placed a small scaffold of carved
wood, having upon it a large dish, containing the skull of one of the
forefathers of the owner of the house, whose weapons are also hung
around it. These consist of bows made of bamboo, bound round with
thread, and provided with strings of buffalo hide, and of arrows three
feet and a half long, made of thin reed or rattan, with sharp iron
points. With these they are able to bring down a hen or other bird at
the distance of sixty or seventy paces, rarely or never missing their
mark. In addition to these weapons they have muskets, lances with
iron points, old swords and long parangs. These last, which they
obtain from the Ceramese and Macassars, have their handles
covered with sharks' skin, and ornamented with handsome shells, a
small metal bell being usually appended to it. The combs and
necklaces of the inhabitants, which I shall describe more fully when
speaking of their mode of dress, are also displayed on this stage.
Round the interior of the house against the wall is erected a
framework, a foot and a half from the floor, which serves for beds by
night and for seats by day. These are occupied chiefly by the
women, the men squatting down on the floor in the centre of the
room.
On the right side of the entrance is placed a clay fire-place,
supported by a wooden stand, on which the provisions of the
inmates are cooked, these last being kept upon a large shelf above
the fire-place. Their more valuable property is hung around on the
wall, or preserved under the benches which line the wall. Upon the
whole, their dwellings present an appearance of greater neatness
than those of the inhabitants of the islands to the westward.
Before reaching the village it is necessary to pass the spot where
their dead is deposited, the interment of whom is attended with
several curious ceremonies. The body is wrapped in cloth, usually
coarse Mirzapore chintz, and put into a coffin made of rough planks,
fastened with wooden pins, a dish being placed under the head. The
coffin is then taken to the burial ground, and wrapped round with the
above-mentioned coarse chintz, after which it is deposited on a
stage raised four or five feet from the ground. A piece of Mirzapore
chintz, fastened to a long bamboo stick, is hoisted over the burial
place, and a paling is erected round it, on which are hung baskets
containing fruits, cooked yams, fowls, rice and other provisions,
which are occasionally renewed. The heat of the sun, and the rains,
soon cause the chintz in which the coffin has been enveloped, to
decay and fall off, but it is immediately renewed, and another dish
placed under the skull. Should the body be that of a rich man, or of
the relative of one of the village chiefs, the skull is removed to the
house, and placed in a conspicuous spot, as I have before
mentioned. If, on the other hand, the body is that of a poor man, the
remains are thrown together into a cleft in a rock, or some such
place, as soon as the coffin has decayed. The bodies of malefactors
are buried at once, without anything being erected to mark the place
of interment. The remains of human beings are never to be seen
scattered about, and it is extremely difficult to obtain a skull from the
natives. I have frequently seen the entire skeleton through the sides
of the coffin, when it has become full of holes through age and
decay. During still weather an intolerable smell arises from these
cemeteries; but the natives appear not to suffer the least
inconvenience from it.
The vessels of the natives are usually hauled up on the beach near
these burial places, and protected from the weather by sheds similar
to our boat-houses. These vessels are chiefly of the kind called
"jonkos," which are either built here or on the island Baba. They are
constructed of rough planks, fastened to each other by wooden pins,
and bound to the timbers with bands of rattan. Saws are never
employed in cutting the planks, neither are iron nails ever used in the
construction of their vessels. The sides of the jonkos are held apart
by thwarts, upon which a deck of bamboos is laid, so that, when
complete, they may be considered as tight and seaworthy. The
gunwale is usually two or three feet above the level of the water.
Their length is from forty to fifty feet; breadth from ten to twelve feet;
and they carry from twelve to sixteen tons. A roof of atap covers the
whole.
Each village usually possesses two or three of these vessels, the
property of the entire community; but they have also numerous
canoes with outriggers, some of which are large enough to contain a
thatched habitation. These craft are employed in fishing, or in the
transport of goods from island to island.
On making enquiries concerning the trading vessels which came
here, I learned, among other things, that during the previous year a
vessel under the English flag had arrived at the east extremity of
Timor-laut, the master of which sent his boat on shore to purchase
provisions, giving out that he had been necessitated to visit the place
from a great mortality and sickness having occurred among his crew.
In how far this was the case, I will leave to the judgment of those
who are aware of the artifices of the English, in order to carry on a
smuggling trade in these parts,[35] and, above all, to import
prohibited goods, such as gunpowder and ammunition. Thus was the
brig, in name (dakleijen), loaded only with roofing slates and iron,
but, in fact, she had a large number of muskets, swords and other
weapons on board. The captain agreed with the natives to give
weapons in exchange for stock, and went on shore with half his
crew, which consisted of ten Europeans. In the mean time the
natives considered this to be a fine opportunity to overpower those
who remained on board, and gain possession of the vessel. They,
therefore, boarded her unexpectedly in great numbers, and
murdered the people in her, while at the same time those on shore
were made away with, with the exception of two boys, who owed
their preservation to the interposition of the women. After this crime
had been committed, they hauled the brig on shore, stripped her of
all they could carry away, and burned her. The plundered goods
were shared out among the inhabitants, and part of them sold to the
traders who visited them, the remainder being kept, and now,
perhaps, serving as finery for the inhabitants of the east coast of
Timor-laut. One of the village chiefs stated to me on this occasion,
that he had himself seen the chain cable of the brig hanging around
the village, and that two iron carronades which had belonged to her
lay there upon the ground, the natives not having, yet mustered
sufficient courage to fire them off.
I fixed the day subsequent to our arrival for a general meeting of the
people, at which I might make known the benevolent intentions of
the Government, and the object of my visit. One of the Orang Kayas
of Serra was accidentally here, making preparations for his return
home, so that I had a good opportunity of sending an invitation to all
the chiefs of that island to come and see me, as I could not promise
with confidence to visit them in their own villages.
We have already seen proofs of the desire shown by the natives to
adopt, in some particular or other, the mode of dress of our
countrymen. As may be supposed, they have no idea whatever of
making clothes, and this is the chief reason of their having such a
ludicrous appearance when clad in their European vestments. They
were, therefore, not a little delighted at finding that I had had several
coats made on board, of red, blue and green cloth, which I gave to
them, in addition to the Government presents. Garments of this
description are not only viewed as a dress of state, but are
considered as distinguished tokens of honour; and it will, therefore,
readily be conceived that they are preserved with the greatest care
as heirlooms, and used only on festive occasions.
Early on the morning of the 12th, I went on shore with a numerous
escort, and under the customary salute from the guns. In the middle
of the village of Watidal a booth had been erected, and covered with
the sails of vessels, under which were numerous seats of plank, the
part assigned to me, as Vakeel or Embassador of the Dutch
Government, being arranged with especial care, and covered with
fine coloured calico. Voluntary homage to our Government was
apparent in all their actions. The people not only filled the booth, but
surrounded it on all sides, leading me to believe that the entire
population, young and old, were present. The presents were
received with the loudest expressions of joy and thankfulness. The
entire day was spent in installing the village chiefs, and making
arrangements for preserving peace and unity among them. The
villages are all independent of one another, each having its own
territory, consisting of a portion of land, and contiguous trepang
banks, and being governed by its own chief. So long, therefore, as
they respect the property of their neighbours, and live according to
the old usages of their forefathers, they remain in perfect peace with
each other. The upper chief, whom I appointed for Watidal, was an
aged man, not at all deficient in good sense and ability. He spoke
very good Malayan, although the language of the natives consists of
a peculiar dialect, in which point there is a remarkable difference
between them and the people of the neighbouring islands.
I could no where discover any remains which would point out that an
officer of the East India Company had resided here, whence I
concluded that the Dutch had not visited these islands for a long
series of years.
The authority of our Government here was more necessary than one
would be led to suppose, from the state of society which exists
among these people. The chiefs are respected by the common men
as the oldest, the most experienced, and the richest among the
inhabitants; they are consulted when differences occur, and their
decision is usually respected; but they still want, to give force to their
authority, an act of appointment from our Government. The
considerate reader, who will already have perceived in the
particulars previously related, how fond these simple people are of
outward show, will not be surprised to find that such an installation is
considered as an important epoch in the history of their country, and
that they wished to celebrate the day on which they were visited by
an officer of the Netherlands Government, after so long a period of
neglect, with all joy and solemnity, as a national festivity.
The want of a formal act of appointment from the Dutch Government
has been attended with results very detrimental to the welfare of the
natives, especially those of Watidal. In the time of the Ceramese
Hongie-togten,[36] the chiefs of Watidal had been presented with
sticks provided with knobs, on which the arms of the Company were
engraven. This token of alliance with us was not sufficient to prevent
the covetous Ceramese from practising extortion upon them, and
when the chiefs complained openly to the Company, and stedfastly
refused all irregular contributions, the Ceramese did not hesitate to
plunder and rob them in open enmity. This would not have taken
place had the chiefs been appointed with more solemnity, and
provided with regular acts of appointment, in which case their
subjects would have been more subordinate to their control, and the
Ceramese rovers would have been withheld from plundering by
dread of punishment.

FOOTNOTES:
[34] These are called imitation calicoes, from their being
European imitations of the pattern and material of the cotton
manufactures of India.—Ed.
[35] In Marsden's Malayan Grammar, among the letters given in
the Appendix, will be found a remarkable example of these
English tricks. An English captain came to the King of Tringanu,
with a pretended letter of recommendation from the Governor-
General of India; asked permission to enter the river to repair, and
smuggled several chests of opium on shore!
[36] Expedition for the extirpation of the spice trees, undertaken in
the time of the old (Dutch) East India Company.
 CHAPTER XIV.
THE TENIMBER ISLANDS.
Visit to Watidal.—Respect shown to the Dutch Flag.—The
English supposed by the Natives to be Orang-gunung, or
Mountaineers.—The Prosperity of the People inseparable
from the Rule of the Dutch over these Countries.—Traces
of the Christian Religion having formerly obtained here.—
Departure from Larrat to Vordate.—Allurements of the
latter Island.—The Inhabitants of the Tenimber Islands.—
Their Manners and Customs.—Mode of Warfare.—Striking
Proofs of their Attachment to the Dutch Government.
The English, during their short possession of the Moluccas, sent a
British flag to the people of Watidal, who had preserved it well, but
without making any use of it. When I demanded the reason of this, I
was answered that they plainly perceived that this was not a
Company's flag, and that they would not recognize any other nation
than the Dutch as their allies. Their ideas of the different European
nations are very singular. From their questions, it appeared that they
considered the Orang Compania (the Dutch) only as true white
people, and that they were unacquainted with other Europeans.
They asked me if the English were not the inland mountaineers, or
Orang Gunung, like the inhabitants of Timor-laut, and other
barbarous people.
The natives call themselves "Anak Compania," literally, Children of
the Company; and they often expressed to me their surprise at the
difficulties the traders who arrived at the island, threw in their way to
prevent them from visiting Banda. They also expressed themselves
much discontented at the arbitrary mode of trading adopted by the
native merchants from Banda, which had often necessitated them to
be their own judges, as they dared not nourish a hope that the Dutch
would again visit and defend them. This state of affairs caused them