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Table of Contents
PREFACE XI DARWIN, WALLACE, AND THE “MEANS

ACKNOWLEDGMENTS xxii OF NATURAL SELECTION” 40
ABOUT THE COVER xxiii VOYAGES TO NATURAL SELECTION 41
ABOUT THE AUTHORS xxiii What Is Natural Selection? 45
Missing Links 46
Key Ideas 47
Part one Deep Currents Key Terms 48
Key Questions to Ask Myself 48
Chapter 1
Key Critical Thinking Questions 48
Introduction to Biological Anthropology 2 Key Things to Do Next 48
Prologue: In the Beginning … 4
Chapter 3
Biological Anthropology: A Diversity of The Biological Basis of Human Variation 50
Interests and an Interest in Diversity 4
Prologue: The Gene Scene 52
Anthropology’s Scope: Who
We Are, What We Are, Why We Are 5 The Building Blocks of Life 52
Fields within Fields: The Scope of Biological The Functions of DNA 55
Anthropology 8 CELL DIVISION 56
Skeletal Biology 10 Mitosis 56
Paleoanthropology 12 Meiosis 57
Human Biology 13
MECHANISMS OF INHERITANCE 59
Molecular Anthropology 14
Pleiotropy 63
Primatology 14
Polygenic Traits 63
Applied Biological Anthropology 16
BEHAVIOURAL GENETICS 64
A Science Old and New 18
TAMPERING WITH MOTHER NATURE: THE
Key Ideas 22
ETHICS OF GENETIC MANIPULATION 64
Key Terms 22
Key Questions to Ask Myself 22 MAPPING THE HUMAN GENOME 65
Key Critical Thinking Questions 22 ANCIENT DNA 66
Key Things to Do Next 22
Key Ideas 67
Chapter 2 Key Terms 68
Science and the Development of Evolutionary Key Questions to Ask Myself 68
Theory 24 Key Critical Thinking Questions 68
Prologue: World views and the
Elephant’s Child 26
Chapter 4
From Variant to Species 70
What is Science? 27
What Is Theory? 28 Prologue: Sweet Accidents of
From Hypothesis to Law 28 History 72
Normal Science and Paradigm Shifts 30 Variant, Variation, Variability 73
EVOLUTIONARY THOUGHT FROM PLATO TO A New Synthesis in the Making? 74
DARWIN 32 Evolutionary Developmental Biology 74
Essence, Order, and Purpose: Plato, Aristotle, From Germ Plasm Theory to Epigenetic
and the “Ladder of Creation” 32 Inheritance Systems 75
From Renaissance to Revolution 33 SPECIES NOW AND THEN 76
Time 34 Species and Species Concepts 77
Diversity 36 Are Other Species Concepts More Useful? 79
Mutability 38 What about Species in the Fossil Record? 80
vi NEL
Copyright 2015 Nelson Education Ltd. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part.
Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s).
How Do New Species Evolve? 81 Hominidae 111
MECHANISMS OF EVOLUTION 82
Mapping Nonhuman Primate Genomes 112
How Does Selection Operate to Produce PRIMATE CONSERVATION 113
Adaptation? 82 Key Ideas 116
Beyond Natural Selection 84 Key Terms 116
Gene Flow 84 Key Questions to Ask Myself 117
Founder Effect and Genetic Drift 86 Key Critical Thinking Questions 117
Is Macroevolution Simply the Sum of Key Things to Do Next 117
M icroevolution? 88
Chapter 6
WRESTLING WITH DIVERSITY 89 Primate Behavioural Ecology 118
The Name Game 89
Prologue: Primates “R” Us 120
Key Ideas 92
Key Terms 92 The Roots of Primatology 120
Key Questions to Ask Myself 92 Social Living 123
HUNTING AND MEAT EATING 125
DO NONHUMAN PRIMATES HAVE
CULTURE? 127
Part TWO Tropical Currents Primate Medicine: Is There
a Doctor in the Forest? 129
Chapter 5
What It Means to Be a Primate 94 AGONISTIC BEHAVIOURS 130
AFFILIATIVE BEHAVIOURS 131
Prologue: Through the Looking Glass 96
Grooming 131
Defining a Primate 96 Alliances 131
Locomotory Features 97
Sensory Adaptations 98 SEXUAL BEHAVIOUR AND REPRODUCTIVE
STRATEGIES 132
Dietary Adaptations 99
Life History and Reproductive Features 100 Sexual Strategies 133
Female Strategies 133
THE ORIGIN OF PRIMATE Male Strategies 134
CHARACTERISTICS 100
LANGUAGE AND COMMUNICATION 136
CLASSIFYING PRIMATES 101
NONHUMAN PRIMATES AS MODELS
PRIMATE BIOGEOGRAPHY 102 OF EARLY HOMININ BEHAVIOUR 138
Primate Locomotion 102
Feeding and Foraging 103 Key Ideas 139
Key Terms 139
THE LIVING PRIMATES 105 Key Questions to Ask Myself 139
Strepsirhini 105 Key Critical Thinking Questions 139
Lemuriformes 106 Key Things to Do Next 139
Lorisiformes 106
Chapter 7
Haplorhini 107
Tarsiiformes 107 Primate Evolution 140
Platyrrhini 107 Prologue: The Family Tree 142
Catarrhini 108 How Primates Become Fossils 142
Cercopithecinae 109 Dating Primate Fossils 142
Colobinae 109 Relative Dating Methods 143
Hominoidea 110 Absolute Dating Methods 143
Hylobatidae 110 Geologic Time Scale 145
NEL Table of Contents vii
THE EARTH AT A GLANCE 146 Key Ideas 192
Key Terms 192
Reconstructing Ancient
Environments 146
Classifying Fossil Primates 148 Key Things to Do Next 192
RECONSTRUCTING EARLY PRIMATE
Chapter 9
DIETS AND SOCIAL BEHAVIOUR 150
Hominin Origins: From Ape to
Dietary Reconstruction 150
Social Behaviour 151 Australopithecine 194
PROLOGUE: WE ARE ALL AFRICANS 196
PALEOCENE: AGE OF THE
PLESIADAPIFORMS 151 A PLACE IN TIME 199
EOCENE: AGE OF THE STREPSIRHINES 153 Hominoids, Habitats, and Hypotheses 200
Climate, Habitat, and Selection 201
OLIGOCENE: AGE OF THE ANTHROPOIDS 155
Origin and Evolution of New World Monkeys 157 A PLETHORA OF PROTOHOMININS 202
Origin and Evolution of Old World Sahelanthropus tchadensis—First Twig on the
Monkeys 157 Bush? 204
Why a New Hominin and Why a
MIOCENE: PLANET OF THE APES 158
New Taxon? 205
Early Miocene (23–16 Ma) 159 Orrorin tugenensis—Sound on the Ground and at
Middle Miocene (16–12 Ma) 159 Ease in the Trees 206
Late Miocene (12–5.3 Ma) 160 Why a New Hominin and Why
MOLECULAR SYSTEMATICS 163 a New Taxon? 207
Key Ideas 163
The Ardipithecines: Hominins, Yes, But Whose
Key Terms 164 Ancestor? 208
Key Questions to Ask Myself 164 Why a New Hominin and Why a
Key Critical Thinking Questions 164 New Taxon? 208
Key Things to Do Next 164 THE AUSTRALOPITHECINES 209
Australopithecine Origins 209
More Skeletons in the Australopithecine Closet 214
Part THREE Ancient Currents Australopithecus sediba—A Study in Mosaic
Chapter 8 Evolution 216
What It Means to Be a Hominin 166 Robust Australopithecines and the Megadont
Adaptation 218
PROLOGUE: A LOOK IN THE MIRROR 168
FROM BUSH TO BRANCH: RECONSTRUCTING
WHAT MAKES A HOMININ A HOMININ? 170 PLIOCENE HOMININ PHYLOGENY 220
Four Legs Good, Two Legs Better? 170
What Are the Benefits of Bipedal Key Ideas 221
Key Terms 221
Locomotion? 174
Bipedalism Also Has Significant Drawbacks 176
Do We Know Why Hominins Adopted Bipedal Key Things to Do Next 222
Behaviour? 178
Chapter 10
CARRYING: WHAT AND WHY? 180
The Emergence of the Genus Homo 224
FEEDING: HOW AND WHEN? 181
From Walking to Running 182 PROLOGUE: 35% MAKES ALL THE
DIFFERENCE 226
TALES TOLD BY TEETH 183
OUR ANCESTORS COME TO LIGHT 226
Canine Honing 184
The Anatomy of Early Homo 227
Enamel Thickness 185
One Species or Two? 228
Why Is My Forehead So Large? 186
BEHAVIOURAL ADAPTATIONS 229
The Brain and Its Organization 186
The First Stone Tools 229
ENERGETICS 189 Dietary Adaptations 230
viii Table of Contents NEL
HOMO ERECTUS: ENIGMA OR ARCHETYPE? 232 Chapter 12
An Anatomy for All Occasions 232 The Emergence of Anatomically Modern
Homo erectus in Africa 233 Humans 274
Homo erectus in Indonesia 236
PROLOGUE: DANCING WITH
Homo erectus in China 237
NEANDERTALS? 276
Homo erectus in Georgia 237
One Species or Two? 239 MODERN HUMANS TAKE THE STAGE 276
BEHAVIOURAL ADAPTATIONS 239 THE ORIGINS OF ANATOMICALLY

Technological Leaps and Bounds 239 MODERN HUMANS 276
The First Big-Game Hunters? 241 Recent Out-of-Africa Model 277
The First Voyager: Hominins Out of Africa 242 Assimilation Model 277
The Multiregional Evolution Model 277
The Hobbits of Flores Island 243 What Do the Fossils Tell Us? 278
Another Piece of the Puzzle, What Does the Genetic Evidence Tell Us? 281
but Which Piece? 245
THE EMERGENCE OF MODERN HUMAN
Key Ideas 246 BEHAVIOUR 283
Key Terms 247 Technology and Subsistence 283
LATER STONE AGE/UPPER PALEOLITHIC 284
Key Things to Do Next 247 IN SEARCH OF NEW LANDS 287
Colonization of Australia 287
Chapter 11 Coming to the Americas 288
The Advent of Humanity 248
Key Ideas 293
PROLOGUE: SKELETONS IN THE Key Terms 293
CLOSET 250 Key Questions to Ask Myself 293
THE EMERGENCE OF ARCHAIC
HOMININS 250
African Archaic Hominins 250
European Archaic Hominins 251
Asian Archaic Hominins 253
Part FOUR Modern Currents
One Species or Several? 253 Chapter 13
Contemplating Modern Human Diversity 296
BEHAVIOURAL ADAPTATIONS OF
ARCHAIC HOMININS 255 PROLOGUE: WE ARE ALL HUMAN 298
THE APPEARANCE OF NEANDERTALS 256 RACE, PLACE, AND FACE 298
Neandertal Morphology 256 If People Are Not Packages, Then Why Are
Neandertal Growth and Development 260 Asians from Asia, Africans from Africa ...? 300
Reflections of a Hard Life 261 What about Spaces between “Races”? 301
NEANDERTAL BEHAVIOURAL SIX FALLACIES CONCERNING RACE 304
ADAPTATIONS 262 Fallacy 1: Human Populations Are
Technology 262 Homogenous 304
Diet and Subsistence Practices 264 Fallacy 2: Polygenic Traits Can Be Measured
Ritual and Symbolic Behaviour 265 Accurately 304
Neandertal Language Capabilities 266 Fallacy 3: Continuously Varying Traits
Deciphering the Neandertal Can Be Marked by Discrete Boundaries 305
Genome 268 Fallacy 4: Traits Used in Racial Classification Are
The Fate of Neandertals 271 Linked 306
Fallacy 5: A Specific Number of Traits
Key Ideas 272 Can Define a Race 306
Key Terms 272
Fallacy 6: Between-Group Genetic Diversity
Distinguishes Geographic Races 307
Key Things to Do Next 272 DIFFERENCE ABUSED AND USED 307
NEL Table of Contents ix
“The Great Problem of Civilization ...” 308 Key Ideas 345
Race as Advantage 310 Key Terms 345
Defining and Measuring Intelligence 311 Key Questions to Ask Myself 346
Do Populations Differ in Intelligence? 311
One Intelligence or Several
Intelligences? 312 Chapter 15
Biological Anthropology as Applied Science 348
RACE AND BIOMEDICINE 313
PROLOGUE: REAL WORLDS, REAL PROBLEMS 350
Key Ideas 314
Key Terms 315 MEDICAL ANTHROPOLOGY 350
Key Questions to Ask Myself 315 Community-Based Research: Fostering
Key Critical Thinking Questions 315 Respectful Collaboration 353
Key Things to Do Next 315 CBR and Indigenous Peoples 353
Chapter 14 EVOLUTIONARY MEDICINE 355
Biology of Contemporary and Past ANTHROPOMETRY AND ERGONOMICS 358
Populations 316
FORENSIC ANTHROPOLOGY 360
PROLOGUE: “THREE OF THESE THINGS From Field to Lab 360
BELONG TOGETHER ...” 318 A New Forensic Anthropology? 365
BIOLOGY OF CONTEMPORARY Key Ideas 367
POPULATIONS 318 Key Terms 367
INFECTIOUS DISEASES 319 Key Questions to Ask Myself 367
Emerging and Re-emerging Infectious Key Critical Thinking Questions 367
Diseases 319 Key Things to Do Next 367
THE HUMAN LIFE COURSE 322 Chapter 16

The Early Years: Breastfeeding, Weaning, Human Legacies, Human Prospects 368
and Child Growth 323 PROLOGUE: PAST, PRESENT, FUTURE 370
The Anthropology of Aging 324
POPULATION GROWTH 371
NUTRITIONAL ANTHROPOLOGY 325 Measuring Population Growth 371
Undernutrition 325 Future Tense? 375
Nutritional Excess: Overnutrition and Obesity 327 A Neo-Malthusian Dilemma? 375
What Exactly Are We Putting into
Our Bodies? 328 BIOLOGIES OF EXCESS AND NEGLECT 378
Nutritional Adaptation 329 Children and Canaries 381
Diabetes and the Thrifty Genotype 329 THE END OF HUMAN EVOLUTION? 383
Lactose Intolerance 331 Blondes Had More Fun? 384
HUMAN ADAPTABILITY 332 Technology, Genetic Load, and Adaptation 384
The Adaptive Significance of Skin Colour 332 Evidence for Recent Human Evolution 385
Living at High Altitude 333 Key Ideas 388
Adaptation to Temperature Extremes 335 Key Terms 388
Heat Stress 335 Key Questions to Ask Myself 388
Cold Stress 337 Key Critical Thinking Questions 389
INVESTIGATING THE BIOLOGY
OF PAST POPULATIONS 337
Appendix A Human and Nonhuman Primate
Health and Disease 338 Comparative Anatomy 390
Paleonutrition 341
Appendix b The Hardy-Weinberg Equilibrium
Activity Patterns 342
Principle 395
Residential Histories 343
SELECTED BIBLIOGRAPHY 397
The Ethics of Studying Human
GLOSSARY 421
Remains 344
INDEX 435
x Table of Contents NEL
Preface
The important thing is not to stop questioning. Curiosity has its own reason for existing.
—Albert Einstein (1879–1955)
Why This Book Now?

There seems to be no shortage of textbooks in the field of biological (also called physical)
anthropology; many have been released in new editions. As university teachers, we have field-
tested several of these over the past 20 years or more. So you might reasonably ask: Why yet
another? The simple answer is that the time has arrived for this particular book; indeed, it is
possibly well overdue. When Nelson proposed the somewhat brash idea of writing the first
biological anthropology book written by Canadian biological anthropologists for Canadian
postsecondary students, we welcomed the opportunity (perhaps with more than a little
naïveté). And thus A Human Voyage was launched!
The growth of the field in Canada is such that it is clearly time for such a text. The
professional association for the discipline, known as CAPA/ACAP (a rather long acronym for the
much lengthier title Canadian Association for Physical Anthropology/l’Association Canadienne
d’Anthropologie Physique), has grown considerably since its inception in the early 1970s and
boasted well over 160 members in 2012, representing several generations of teachers and their
students. As well, university and college courses in the field (and in related subjects such as
human ecology, human adaptability, medical anthropology, and forensic anthropology) have seen
enrolment increase steadily over the past several decades; also growing is the number of graduate
programs leading to advanced degrees (M.A. or Ph.D.). This growth is reflected in a broad range of
nationally and internationally recognized academic scholarship, carried out not just in Canada but
around the world, which covers virtually all facets of the discipline from the latest developments in
evolutionary theory to interpretations of the fossil record, insights into nonhuman primate ecology
and behaviour, modern population biology, and applied biocultural anthropology.
As teachers, one lesson we have taken from our own students over the years is that their
conceptual and theoretical learning is greatly enhanced when practical examples and references
resonate geographically, socially, and culturally. As we suspect many of our colleagues across
Canada have done in the past, we often bring material into class highlighting Canadian
scholarship to supplement that provided by the text, even in the recently evolved Canadian
editions of existing (non-Canadian) texts. The intent of A Human Voyage is to put that
knowledge and experience “between the covers.” This book, then, is unabashedly Canadian
in its focus and content, but not to the point of being parochial. An important point we
make throughout this second edition of A Human Voyage is that human biocultural diversity
crosses national boundaries: research done in Canada has significance around the world. At
the same time we acknowledge that these boundaries and the political and economic entities
that create and maintain them can have profound impacts on human biology—the history
of colonization and the health of Aboriginal populations in Canada and elsewhere stands as
one of many such examples, as does the unique geographic distribution of rare traits such as
Tay-Sachs disease (see Chapter 13), which has followed population migration over the ages.
Connecting the Dots ...

Textbooks in biological anthropology typically adhere to a particular structure, beginning
with an overview of the discipline’s history, a bit of evolutionary theory, and some dabbling (at
times outright wading) into the genetic mechanisms underlying population variation. This is
NEL xi
then followed by a number of chapters discussing the evolutionary history and biobehavioural
diversity of our closest living relatives, the nonhuman primates. This overview sets the
stage for introducing the story of our own evolving lineage of the past several million years,
beginning in Africa and tracing the global expansion of human migration. Eventually we end
up in the here and now, and conclude with several chapters discussing modern population
biology, its historical antecedents and geographic patterning. This is a logical, tried-and-true
model, and in writing this text we have chosen to follow a similar structure. The benefit in
doing so is that it provides a text that will have a familiar feel for the instructor, which in turn
will make for an easy transition when adapting its content for the student.
In keeping with this model, A Human Voyage is presented in 16 chapters divided into four
parts: Deep Currents introduces the history of biological anthropology as a field of study
and its development in Canada, as well as the theoretical foundation and structure of human
variation; Tropical Currents traces the evolution of our nearest primate relatives and the
fascinating adaptations and behaviours expressed by those species still among us; Ancient
Currents delves into the 7-million-year-old story of how our particular ancestors came to
be human, some of it speculative and controversial; and Modern Currents focuses on the
complex variations that exist among living human populations, how they came about, and
their importance for humanity as we look forward into the 21st century and beyond.
A Few Comments for the Student

This is your book—it was written for students with little or no background in the field and
in such a way as to make the story of human evolution not only accessible but also enjoyable.
However, in reading this second edition of A Human Voyage, you will see that the path to our
past is not always clearly defined, and you may well wonder how we know anything at all!
Do not be deterred—this is simply science at work. The possibility that we could have several
plausible explanations for how we came to be as we are today is a cornerstone of modern
science (a classic example being the adoption of upright walking, known as bipedalism,
discussed in Chapter 8). However, it is also true that while a number of credible scenarios
may be proposed based on available evidence, there was in fact only one pathway taken by
those many generations of your ancestors and their descendants—step by step by step. The
journey along that pathway culminated in the diverse, complex global species—modern
Homo sapiens—of which you are a member. It is quite possible, if not actually probable, that
none of our current interpretations is an entirely accurate account of that voyage, although
the weight of evidence may favour one over other reasonable explanations. This is why
we have entitled this book A Human Voyage rather than The Human Voyage, which would
presume that somehow we possessed a complete and precise understanding of the past. This
is a claim no one can justifiably make.
Our hope is that this second edition and the course you are taking will encourage a
critical perspective and sense of wonder in each of you. Each chapter begins with specified
learning outcomes—these are not meant to be exhaustive but to illustrate some of the
objectives we hope you will achieve in working through the text. Each chapter ends with a
concise summary (Learning Keys) that recounts the key ideas we have covered in the chapter,
the significant key terms associated with these essential points, and a few questions you might
want to ask yourself about what you have learned. We also pose critical thinking questions
to have you challenge what you have just read in the previous pages. The Learning Keys
and questions are your best bet to understanding the substance of each chapter’s content, and
perhaps doing very well in the course you are taking. In that regard, A Human Voyage is as
much about the art of questioning as it is about our current understanding of our species—its
diversity, its history, and its possible future. As the authors of A Human Voyage, we will gauge
its success by the degree to which it leaves you, the student, feeling somewhat dissatisfied
xii Preface NEL
with the descriptions, interpretations, and arguments provided, considerably more aware that
there is a very robust—although still imperfect—body of evidence detailing humankind’s
evolutionary story, and much less complacent and accepting of the current state of our species
and its impact on this planet.
What’s New in the Second Edition?

This second edition of A Human Voyage has been revised cover to cover in light of several
reviewers’ excellent suggestions to improve clarity and student accessibility. Each chapter
has been updated to reflect the current state of the literature, with over 200 new references
added to an already extensive bibliography. We have also made significant improvements to
both chapter-opening material (specifying Learning Objectives according to Bloom’s Revised
Taxonomy) as well as ending material in the form of informative and visually accessible
Learning Keys (ideas, key terms, and questions). The Learning Keys presented in each chapter
are a metaphor—as keys, they “unlock” the essential learning points covered; they link to the
Key Concepts and the Key Terms, thus tying together all of the major elements covered in
the chapter. The following provides a brief chapter-by-chapter synopsis:
Chapter 1 (“Introduction to Biological Anthropology”) sees the addition of two new

features: an expanded description of the significant role of molecular anthropology in the
discipline, and a new Profile box highlighting the research of Dr. Tracy Prowse (McMaster
University) in the use of archaeological isotopes to explore patterns of diet and migration
in prehistory. We have also rewritten the historical accounts of the discipline of biological
anthropology, making it more accessible to students.
Chapter 2 (“Science and the Development of Evolutionary Theory”) provides a review of the
practice of science as a way of knowing and the foundations for understanding evolutionary
theory, from Plato to Darwin. In this second edition, we have revised and shortened a number
of sections, improving clarity and understanding of important concepts including uniformi-
tarianism, Linnaean classification, Lamarckian inheritance, and Darwinian evolution.
Chapter 3 (“The Biological Basis of Human Variation”) features expanded sections on the
functions of DNA and on ancient DNA.
Chapter 4 (“From Variant to Species”) presents a new section covering the principles of the
Hardy-Weinberg Equilibrium (explored further in Appendix B, a new addition to this edition).
In a new table, we clearly outline the assumptions under which evolution (changes in gene
frequency) would not occur as specified by the tenets of the Hardy-Weinberg Equilibrium.
Also new to Chapter 4 is a revised treatment of epigenetic inheritance, making this important
although complex topic much more accessible to students. New material describing the ideas
of homology and homoplasy has been added, preparing students for subsequent discussion of
primate and human evolution.
Chapter 5 (“What It Means to Be a Primate”) includes new content featuring research by

a number of well-known Canadian primatologists. The sections on mapping nonhuman
primate genomes and primate cloning have been amalgamated and updated. The section on
primate feeding and foraging has been expanded to include a discussion of the relationship
between body size, molar cusp morphology, foraging group size and diet, sex differences in
diet, and methods used to study primate diets. The discussion of primate conservation has
been shifted from Chapter 15 in the first edition to the end of this chapter.
NEL Preface xiii
Chapter 6 (“Primate Behavioural Ecology”) is characterized by the addition of two new
sections, one focusing on hunting and meat eating, and the other on the use of nonhuman
primates as models for early hominin behaviour. The discussion of social living has been
shifted from Chapter 5 in the first edition to this chapter and has been expanded. The sections
on culture among nonhuman primates and language and communication studies have both
been expanded to include more examples, and in the case of the latter, to highlight some of
the controversies surrounding ape language studies. A new box contributed by Dr. Linda
Fedigan (University of Calgary) profiles the history of primatology in Canada.
Chapter 7 (“Primate Evolution”) features a revised section on primate classification that

includes a discussion of the difference between crown and stem primates. The section on
dietary reconstruction has been expanded to include a discussion of the techniques used to
explore primate diets. Updated sections on Paleocene, Eocene, Oligocene, and Miocene
primates include new fossil primate discoveries, and a new Profile box highlights research by
Dr. Mary Silcox (University of Toronto) on plesiadapiforms.
Chapter 8 (“What It Means to Be a Hominin”) outlines the major developments in

hominin evolution, focusing on bipedalism, dental reduction, and encephalization. New
findings regarding variation in the bony labyrinth, the midtarsal break in the foot, and the
debate regarding the endurance running hypothesis have been added. More informative
illustrations of the major anatomical changes accompanying bipedal locomotion have been
developed to enhance student understanding of these often complex changes. The discussion
of the obstetric dilemma is now contrasted with Dunsworth’s recently advanced concept of
the Energetics of Gestation and Growth. Our revised discussion of bipedalism emphasizes the
emerging consensus that there may have been many different styles of bipedal locomotion
explored by our hominin ancestors over the past 5 million years.
Chapter 9 (“Hominin Origins: From Ape to Australopithecine”) begins the story of

hominin evolution, from Sahelanthropus to Australopithecus. Each section has been updated to
include the most recent analyses (such as the recently described hominin molar from Ishango),
and we have completely revised our discussion of Ardipithecus, emphasizing its unique bipedal
morphology. An important new addition centres on the incredible finds from Malapa, South
Africa, attributed to the new species Australopithecus sediba. A new Profile box is included,
contributed by Dr. Tracy Kivell (University of Kent), one of the principal researchers of the
Au. sediba fossil sample, focusing on the hand bones and noting that Au. sediba would have
been quite capable of the fine precision grips needed for stone tool manufacture.
Chapter 10 (“The Emergence of the Genus Homo”) includes an expanded discussion of the
Dmanisi fossils.
Chapter 11 (“The Advent of Humanity”) features a new Profile box highlighting research by
Dr. Mirjana Roksandic (University of Winnipeg) on Middle Pleistocene hominins in Eastern
Europe. The section on the Neandertal genome has been significantly expanded to include a
box highlighting the most recent genetic studies of these hominins. Similarly, the discussion of
the Denisovan fossils has been lengthened to include recent advances in the study of their DNA.
Chapter 12 (“The Emergence of Anatomically Modern Humans”) presents new genetic

findings from Siberia that cast light on the ancestry of the first inhabitants of North and South
America.
Chapter 13 (“Contemplating Modern Human Diversity”) explores the basis of modern

human diversity by first examining the historic and contentious concept of race. New
xiv Preface NEL
weblinks are provided, along with updates to our model of the “Six Fallacies Concerning
Race.” We have also revised the Race and Biomedicine section to consider the practice
of “direct-to-consumer” genetic engineering, an idea that is becoming increasingly of
interest in the realm of so-called personalized medicine.
Chapter 14 (“Biology of Contemporary and Past Populations”) features two new Profile
boxes, one contributed by Dr. Stacie Burke (University of Manitoba) highlighting her
research on tuberculosis in Canada, and the other written by Dr. Helen Kurki (University
of Victoria) profiling her studies of body shape variation among small-bodied populations.
Expanded discussions of nutrition in contemporary populations and activity patterns of past
populations are also included.
Chapter 15 (“Biological Anthropology as Applied Science”) examines several biological

anthropology fields that fall under the rubric of applied science. Collaborators Tina Moffat
(McMaster University), Dan Sellen (University of Toronto), and Warren Wilson (University
of Calgary) describe their work, looking at breastfeeding practices of immigrant mothers
in a new Focus On box. A new section, written by Sylvia Abonyi from the University of
Saskatoon, explores the practice of community-based research with Canada’s Indigenous
peoples. Finally a new Profile box contributed by Tanya Peckmann (St. Mary’s University)
examines one of the most frequently asked of student questions: “Can I have a career doing
forensic anthropology?”
Chapter 16 (“Human Legacies, Human Prospects”) includes new information in the sections
on demography, population growth, and political economy. As in the first edition, we return
to another very common student question “Are humans still evolving,?” with updated
research on the proposed role of the CCR5D32 gene and immune response.
Appendix A is new to this edition and provides a comparative depiction of the human
and nonhuman primate skeleton, labelling the major bones and illustrating the significant
differences related to the evolution of unique human features such as bipedal locomotion,
craniodental reduction, and encephalization.
Appendix B is also a new addition to this text. It outlines the major tenets of the Hardy-
Weinberg Equilibrium theorem introduced in Chapter 4. This appendix models the effect
of selection on changing allele frequencies in a population using the standard binomial
formula p2 + 2pq + q2 = 1. The Hardy-Weinberg Equilibrium theorem is illustrated with
a real-world example, modelling allelic variation in the CYP2C8 gene that moderates drug
effectiveness in treating malarial infection in Africa.
NEL Preface xv
SPECIAL FEATURES OF THE BOOK
Chapter Openers
Each chapter begins with an Overview section that summarizes the content that will
be covered in that chapter. A Key Concepts section introduces students to central ideas
that will be discussed in the chapter. These key concepts are followed by a series of Key
Learning Objectives based on Revised Bloom’s taxonomy to guide the student learner
through the chapter content.
Maps, Photographs, and Illustrations

Colourful visuals, many of which are unique
to this text, are used to illustrate key concepts
and information and to help students master the
material.
xvi NEL
Pedagogical Boxes
Each chapter includes one or more boxed features that delve more
deeply into the subject matter and history of biological anthropology.
These boxes take three forms:
Retrospection boxes emphasize key ideas

or seminal developments in the field’s litera-
ture and provide insight into how the disci-
pline has taken shape over the years.
Focus On boxes provide in-depth analysis

of particular topics covered in the chapter.
Profile boxes—written by some

of our colleagues—illustrate the
exceptional scholarship typifying
biological anthropology in Canada.
We could have included many more of
these, but for some reason, our editors
felt that a 1,000-page textbook might
not be appropriate!
NEL Special Features of the Book xvii
Key Terms and Glossary
Each chapter features key terms and their definitions
in the margin, beside where they appear in the text. A
cumulative glossary is provided at the back of the book.
End-of-Chapter Learning Keys

Connecting to the key concepts and learning objectives
listed in the chapter-opening pages, each chapter ends
with the learning keys shown below and on the next
page:
Key Ideas A bulleted summary of the

chapter content highlights the major
themes, issues, and concepts covered.
Key Terms A listing of the most

important key terms in the chapter
helps student remember important
concepts.
xviii Special Features of the Book NEL
Key Questions to Ask
Myself Student-centred questions are written
from the learner’s perspective, inviting the stu-
dent to engage in a novel way with the chapter
content.
Key Critical Thinking

Questions Provocative questions are
designed to critically engage students with
the chapter content and to promote indi-
vidual learning as well as seminar-style
discussion.
Key Things to Do Next This is a student

reminder of, and an invitation to, the Biolo-
gical Anthropology CourseMate website,
where a wealth of support content is available.
This website brings course concepts to life
with interactive learning and exam preparation
tools that integrate with the printed textbook.
Students activate their knowledge through
quizzes, games, and flashcards, among many
other tools. Visit NELSONbrain.com to start
using CourseMate. Enter the Online Access
Code from the card included with your text. If
a code card is not provided, you can purchase
instant access at NELSONbrain.com.
Bibliography
In-text citations throughout the book provide the sources from which the materials are
drawn. Sources from 2004 to date, along with a few of the most important “classic” works,
are listed in the Selected Bibliography at the back of the book, while earlier sources are
available on the CourseMate site for students (accessible through NELSONbrain.com;
an additional charge may apply), with the Instructor Resources at www.nelson.com/
humanvoyage2e, and on the Instructor’s Resource CD (ISBN 978-0-17-656068-3).
NEL Special Features of the Book xix
ANCILLARIES
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All NETA and other key instructor ancillaries are provided on the Instructor’s Resource CD (ISBN
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NETA Test Bank: This resource was written by Alexis Dolphin, Western University. It
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xx Ancillaries NEL
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NETA PowerPoint: Microsoft® PowerPoint ® lecture slides for every chapter have been
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Student Ancillaries
Biological Anthropology CourseMate: The more you study, the better the results. Make
the most of your study time by accessing everything you need to succeed in one place.
Biological Anthropology CourseMate includes
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NEL Ancillaries xxi
ACKNOWLEDGMENTS
This labour of love would not have been possible without the help and support of numerous
friends and colleagues. At Nelson, we wish to thank former sales representative Erin Carlson
for planting in our heads the idea to do this project, and Executive Editor Maya Castle for her
tireless effort in promoting the text and guiding us from the first through the second edition.
Special thanks go to our developmental editor, Katherine Goodes, for her patience and
persistence in keeping us on schedule and on budget! Thanks also go to Nelson’s permissions
editor, Carrie McGregor; production project manager, Natalia Denesiuk Harris; copy editor,
June Trusty; and project manager, Naman Mahisauria.
We also acknowledge the support of our colleagues across Canada and beyond who shared
many of their stories, insights, and images to make this a better book. We are especially grateful
to the following colleagues who contributed written material and photographs for the book
or the associated CourseMate website: David Begun (University of Toronto), Stacie Burke
(University of Manitoba), Julie Cormack (Mount Royal University), Alan Cross (Simon
Fraser University), Jerry Cybulski (formerly with the Canadian Museum of History),
Michelle Drapeau (Université de Montréal), Linda Fedigan (University of Calgary), Tracey
Galloway (University of Manitoba), Todd Garlie (U.S. Army Research, Development
and Engineering Command, Natick, Massachusetts), Tracy Kivell (University of Kent),
Helen Kurki (University of Victoria), Carol MacLeod (Langara College), Tina Moffat
(McMaster University), Mary Pavelka (University of Calgary), Tanya Peckmann (St. Mary’s
University), Tracy Prowse (McMaster University), Dan Sellen (University of Toronto),
Mirjana Roksandic (University of Winnipeg), Mary Silcox (University of Toronto), Mark
Skinner (Simon Fraser University), Matt Skinner (University College London), Matt
Tocheri (National Museum of Natural History, Smithsonian Institution), and Warren
Wilson (University of Calgary).
We also thank Henry Schwarcz (McMaster University) and Jane Evans (NERC
Isotope Geosciences Laboratory, United Kingdom) for their helpful comments, and the
following individuals for contributing many of the wonderful photographs in the book: Ian
Colquhoun (University of Western Ontario), Tosha Dupras (University of Central Florida),
Lisa Gould (University of Victoria), Kayla Hartwell (Wildlife Care Center of Belize,
Belmopan, Belize), Bonnie Kahlon (McMaster University), Shannon McPherron (Max
Plank Institute for Evolutionary Anthropology), Mary Pavelka (University of Calgary),
Tracy Prowse (McMaster University), Amy Scott (University of Manitoba), Pascale
Sicotte (University of Calgary), Travis Steffens (researcher/photographer, Ankarafantsika,
Madagascar), and Andrzej Weber (University of Alberta). Thank you to Leah Andrews
(Adam Scott Collegiate, Peterborough) for doing some of the background research for
the book, and to our students for their encouragement. Special thanks go to those who
participated in the reviewing process for this second edition and whose comments helped
bring this text to fruition:
Julie Cormack, Mount Royal University
Paul Erickson, Saint Mary’s University
Samanti Kulatilake, Mount Royal University
Carol MacLeod, Langara College
Bob Muckle, Capilano University
Katherine Patton, St. Francis Xavier University
Dennis Sandgathe, Simon Fraser University
Mary Silcox, University of Toronto Scarborough
xxii Acknowledgments NEL
Finally, we are eternally grateful for the support given and the sacrifices made by family and
friends, who never let us forget that biological anthropologists are human, too!
ABOUT THE COVER

Humans share two biological voyages, recounted in this text and reflected on the cover. The
first is a developmental voyage, from conception to death, during which our unique biology
is expressed, challenged, and modified by the diverse circumstances and environments in
which we grow and age. The second is the continuing evolutionary voyage, written in our
primate heritage from ancestor to descendent, upon which we can look back with fascination
and only wonder at the future.
ABOUT THE AUTHORS

Anne Keenleyside
Anne Keenleyside is an Associate Professor and Chair of the Department of Anthropology at
Trent University in Peterborough, Ontario, where she has been based since 2002. She received
her Ph.D. in anthropology from McMaster University and also holds a Bachelor of Education
degree from the Ontario Institute for Studies in Education. Trained as a bioarchaeologist, she
has conducted fieldwork in Nunavut, Siberia, Russia, Romania, and Tunisia, and for the past
decade has been investigating the health and diet of an ancient Greek colonial population on
the Black Sea coast of Bulgaria. Most recently, she has embarked on the analysis of newly
recovered skeletal remains from the last expedition of Sir John Franklin (1845–1848) to the
Canadian Arctic.
Richard Lazenby
Richard Lazenby is Professor and founding member of the Anthropology Department at the
University of Northern British Columbia, Prince George, British Columbia. Richard arrived
at UNBC in 1994 after receiving his Ph.D. at McMaster University (1992) and a brief sojourn
as a Post-Doctoral Fellow at the University of Guelph. His overarching research interest is to
understand how life history (growth, aging, diet, activity) shapes both external and internal
skeletal morphology. Most recently he has been working with colleagues at the University of
Calgary and at the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany,
exploring the application of 3D microcomputed tomographic imaging to investigate patterns
of asymmetry related to hand use in humans, nonhuman primates, and fossil hominins.
NEL About the Authors xxiii
Another random document with
no related content on Scribd:
for black bees, and seven dollars for pure Italians. If you purchase in
the fall, require 33⅓ per cent, discount on these rates.
WHERE TO LOCATE.
If apiculture is an avocation, then your location will be fixed by
your principal business or profession. And here I may state, that if
we may judge from reports which come from nearly every section of
the United States, from Maine to Texas, and from Florida to Oregon,
you can hardly go amiss anywhere in our goodly land.
If you are to engage as a specialist, then you can select first with
reference to society and climate, after which it will be well to secure
a succession of natural honey-plants (Chap. XVI.), by virtue of your
locality. It will also be well to look for reasonable prospects of a good
home market, as good home markets are, and must ever be, the
most desirable. It will be desirable, too, that your neighborhood is not
overstocked with bees. It is a well-established fact, that apiarists with
few colonies receive relatively larger profits than those with large
apiaries. While this may be owing in part to better care, much
doubtless depends on the fact that there is not an undue proportion
of bees to the number of honey-plants, and consequent secretion of
nectar. To have the undisputed monopoly of an area reaching at
least four miles in every direction from your apiary, is unquestionably
a great advantage.
If you desire to begin two kinds of business, so that your dangers
from possible misfortune may be lessened, then a small farm—
especially a fruit farm—in some locality where fruit-raising is
successfully practiced, will be very desirable. You thus add others of
the luxuries of life to the products of your business, and at the same
time may create additional pasturage for your bees by simply
attending to your other business. In this case, your location becomes
a more complex matter, and will demand still greater thought and
attention. Some of Michigan's most successful apiarists are also
noted as successful pomologists.
For position and arrangement of apiary see Chapter VI.
CHAPTER V.
HIVES AND BOXES
An early choice among the innumerable hives is of course

demanded; and here let me state with emphasis, that none of the
standard hives are now covered by patents, so let no one buy rights.
Success by the skillful apiarist with almost any hive is possible. Yet,
without question, some hives are far superior to others, and for
certain uses, and with certain persons, some hives are far preferable
to others, though all may be meritorious. As a change in hives, after
one is once engaged in apiculture, involves much time, labor and
expense, this becomes an important question, and one worthy
earnest consideration by the prospective apiarist. I shall give it a first
place, and a thorough consideration, in this discussion of practical
apiculture.
BOX-HIVES.
I feel free to say that no person who reads, thinks, and studies—
and success in apiculture can be promised to no other—will ever be
content to use the old box-hives. In fact, thought and intelligence,
which imply an eagerness to investigate, are essential elements in
the apiarist's character. And to such an one a box-hive would be
valued just in proportion to the amount of kindling-wood it contained.
A very serious fault with one of our principal bee-books, which
otherwise is mainly excellent in subject matter and treatment, is the
fact that it presumes its readers to be box-hive men. As well make
emperors, kings, and chivalry the basis of good government, in an
essay written for American readers. I shall entirely ignore box-hives
in the following discussions, for I believe no sensible, intelligent
apiarists, such as read books, will tolerate them, and that, supposing
they would, it would be an expensive mistake, which I have no right
to encourage, in fact, am bound to discourage, not only for the
benefit of individuals, but also for the art itself.
To be sure of success, the apiarist must be able to inspect the
whole interior of the hive at his pleasure, must be able to exchange
combs from one hive to another, to regulate the movements of the
'bees: by destroying queen-cells, by giving or withholding drone-
comb, by extracting the honey, by introducing queens, and by many
other manipulations to be explained, which are only practicable with
a movable-frame hive.
MOVABLE-COMB HIVES.
There are, at present, two types of the movable-comb hive in use
among us, each of which is unquestionably valuable, as each has
advocates among our most intelligent, successful and extensive
apiarists. Each, too, has been superseded by the other, to the
satisfaction of the person making the change. The kind most used
consists of a box, in which hang the frames which hold the combs.
The adjacent frames are so far separated that the combs, which just
fill them, shall be the proper distance apart. In the other kind, the
frames are wider than the comb, and when in position are close
together, and of themselves form two sides of a box. When in use,
these frames are surrounded by a second box, without a bottom,
which, with them, rests on a bottom board. Each of these kinds is
represented by various forms, sizes, etc., where the details are
varied to suit the apiarist's notion. Yet, I believe that all hives in
present use, worthy of recommendation, fall within one or the other
of the above named types.
THE LANGSTROTH HIVE.
This (Fig, 29) is the hive most in use among Americans and
Britons, if not among all who practice improved apiculture. It is stated
that the late Major Munn was first to invent this style of hive. He
states (see Bevan, p. 37) that he first used it in 1834. But, as
suggested by Neighbour in his valuable hand-book, the invention
was of no avail to apiarists, as it was either unknown, or else ignored
by practical men. This invention also originated independently with
Rev. L. L. Langstroth, who brought it forth in 1851, so perfect, that it
needed scarce any improvement; and for this gift, as well as his able
researches in apiculture, as given in his invaluable book, "The
Honey-Bee," he has conferred a benefit upon our art which cannot
be over-estimated, and for which we, as apiarists, cannot be too
grateful. It was his book—one of my old teachers, for which I have
no word of chiding—that led me to some of the most delightful
investigations of my life. It was his invention—the Langstroth hive—
that enabled me to make those investigations. For one, I shall
always revere the name of Langstroth, as a great leader in scientific
apiculture, both in America and throughout the world. His name must
ever stand beside that of Dzierzon and the elder Huber. Surely this
hive, which left the hands of the great master in so perfect a form,
that even the details remain unchanged by many of our first bee-
keepers, should ever bear his name. Thus, though I prefer and use
the size of frame first used, I believe, by Mr. Gallup, still I use the
Langstroth hive. (See Appendix, page 287).
Fig. 29.
CHARACTER OF THE HIVE.

The main feature of the hive should be simplicity, which, would
exclude doors, drawers, and traps of all kinds. The body should be
made of good pine or white-wood lumber, one inch thick, thoroughly
seasoned, and planed on both sides. It should be simply a plain box
(Fig, 30), without top or bottom, and of a size and form to suit the
apiarist. The size will depend upon our purpose. If we desire no
comb-honey, or desire comb-honey in frames, the hive may contain
4,000 cubic inches. If we desire honey in boxes, it should not contain
over 2,000, and may be even smaller. If the hive is to be a two-story
one—that is, one hive above a similar hive below (Fig, 29)—I prefer
that it should be eighteen inches long, twelve inches wide, and
twelve inches deep, inside measure. If simply small frames or boxes
are to be used above, I would have the hive at least two feet long. A
three-fourths inch rabbet should be cut from the top of the sides or
ends as the apiarist prefers, on the inside (Fig, 30, c).
Fig. 30
The rabbet may equal a little more than one-half the thickness of
the board. Heavy tin strips (Fig, 33), three-fourths of an inch wide,
should be tacked to the side below the rabbet, so as to reach one-
fourth of an inch above the shoulder. These are to bear the frames,
and are convenient, as they prevent the frames from becoming glued
to the hive. We are thus able to loosen the frames without jarring the
bees. I would not have hives without such tin rabbets, though some
apiarists, among whom is Mr. James Heddon, of this State, whose
rank as a successful apiarist is very high, do not like them. The
objection to them is cost, and liability of the frames to move when the
hive is moved. But with their use we are not compelled to pry the
frames loose, and are not so likely to irritate the bees, while making
an examination of the contents of the hive, which arguments are
conclusive with me.
Any one who is not a skilled mechanic, especially if he has not a
buzz-saw, had better join the sides of his hives after the style of
making common dry-goods boxes (Fig, 30). In this case, the sides
not rabbeted should project by, else the corners will have to be
stopped up where they were rabbeted.
Fig. 31.
Bevel-Gauge.
The mechanic may prefer to bevel the ends of the boards, and
unite them by a miter-joint (Fig, 33). This looks a little better,
otherwise is not superior to the other method. It is difficult to form
accurate joints—and as everything about the hive should be
accurate and uniform—this style is not to be recommended to the
general apiarist. To miter with a hand-saw unless one is very skillful,
requires a perfect miter-box, and, even then, much care is required
to secure perfect joints. With a buzz-saw this is easier. We have only
to make a carrier as follows: Take two boards (Fig, 31. a, b), each
one foot in length, and dove-tail them together, as though with two
others you meant to make a square box. Be sure that they form a
perfect right-angle. Then bevel the ends opposite the angle, and
unite these with a third board (Fig, 31, c), firmly nailed to the others.
We thus have a triangular pyramid. Through one of the shorter faces
make longitudinal slits (Fig, 31, d), so that this can be bolted firmly to
the saw-table. In use, the longer face will reach the saw, and from
thence will slant up and back. Along the back edge of this a narrow
board (Fig, 31, e) should be nailed, which will project an inch above
it. This will keep the board to be beveled in line with the carrier, and
will retain the right angles. Of course the boards for the hive must be
perfect rectangles, and of just the right length and width, before the
bevels are cut.
Such a carrier (Fig, 31) I ordered for my Barnes' saw, from a
cabinet-maker. It was made of hard wood, all three joints dove-tailed,
and nicely finished, at a cost of $1.50.
In sawing the ends and sides of the hive, whether by hand or with
a buzz-saw, use should be made of a guide, so that perfect
uniformity will be secured.
THE BOTTOM BOARD.
For a bottom board or stand (Fig, 32), we should have a single

one-inch board (Fig, 32, b) just as wide as the hive, and four inches
longer, if the bees are to enter at the end of the hive, and as long,
and four inches wider, if the bees are to enter at the side. This is
nailed to two pieces of two by four, or two by two scantling (Fig, 32,
a, a). Thus the hive rests two or four inches from the ground. These
scantlings should extend at one end eight inches beyond the board,
and these projections be beveled from the edge of the board, to the
lower outer corner of the scantling. Upon these beveled edges nail a
board (Fig, 32>, d), which shall reach from the edge of the bottom
board to the ground. We thus have the alighting-board, whose upper
edge should be beveled, so as to fit closely to the bottom board. If
the hives are to be carried into a cellar to winter, this alighting-board
(Fig, 31, d) had better be separate, otherwise it is more convenient
to have it attached. It may be made separate at first, or may be
easily separated by sawing off the beveled portion of the scantlings.
Should the apiarist desire his bees to enter at the side of the hive,
the scantling (Fig, 32, a, a) should run the other way, and the
alighting-board (Fig, 32, d) should be longer, and changed to the
side. I have tried both, and see no difference, so the matter may be
controlled by the taste of the apiarist.
Fig. 32.
For an opening to the hive (Fig, 32, c), I would bevel the middle
of the edge of the bottom board, next to the inclined board. At the
edge, this bevel should be three-quarters of an inch deep and four
inches wide. It may decrease in both width and depth as it runs back,
till at a distance of four inches, it is one-half an inch wide and five
thirty-seconds of an inch deep. This may terminate the opening,
though the shoulder at the end may be beveled off, if desired.
With this bottom board the bees are near the ground, and with
the slanting board in front, even the most tired and heavily-laden will
not fail to gain the hive, as they come in with their load of stores. In
spring, too, many bees are saved, as they come in on windy days,
by low hives and an alighting-board. No hive should be more than
four inches from the ground, and no hive should be without the
slanting alighting-board. With this opening, too, the entrance can be
contracted in case of robbing, or entirely closed when desired, by
simply moving the hive back.
Some apiarists cut an opening in the side of the hive, and
regulate the size by tin slides or triangular blocks (Fig, 29); others
form an opening by sliding the hive forward beyond the bottom board
—which I would do with the above in hot weather when storing was
very rapid—but for simplicity, cheapness and convenience, I have
yet to see an opening superior to the above. I think, too, I am a
competent judge, as I have at least a half-dozen styles in present
use.
I strongly urge, too, that only this one opening be used. Auger
holes about the hive, and entrances on two sides, are worse than
useless. By enlarging this opening, we secure ample ventilation,
even in sultry August, and when we contract the entrance, no bees
are lost by finding the usual door closed.
Some of our best bee-keepers, as Messrs. Heddon, Baldridge,
etc., prefer that the bottom board be nailed to the hive (Fig, 39). I
have such hives; have had for years, but strongly object to them.
They will not permit a quick clearing of the bottom board, when we
give a cleansing flight in winter, or when we commence operations in
spring, which, especially if there is a quart or more of dead bees, is
very desirable. Nor with their use can we contract the opening in cold
weather, or to stop robbing, without the blocks (Fig, 29), tins or other
traps. Simplicity should be the motto in hive-making. The arguments
in favor of such fastening are: Convenience in moving colonies, and
in feeding, as we have not to fasten the bottoms when we desire to
ship our bees, and to feed we have only to pour our liquids into the
hives.
Of course, such points are not essential—only matters of
convenience. Let each one decide for himself, which experience will
enable him to do.
THE COVER OF THE HIVE.
The cover (Fig, 33, a) should be about six inches high, and like
the lid of a trunk. The length and breadth may be the same as the
body of the hive, and fit on with beveled edges (Fig, 33), the body
having the outer edge beveled, and the cover the inner. If we thus
join the cover and hive with a mitered-joint, we must not be satisfied
with anything less than perfection, else in case of storms, the rain
will beat into our hives, which should never be permitted. Such
covers can be fastened to the hives with hinges, or by hooks and
staples. But unless the apiarist is skilled in the use of tools, or hires a
mechanic to make his hives, it will be more satisfactory to make the
cover just large enough (Fig, 29) to shut over and rest on shoulders
formed either by nailing inch strips around the body of the hive, one
inch from the top, or else inside the cover (Fig, 29). If it is preferred
to have a two-story hive, with the upper story (Fig, 33, b) just like the
lower (Fig, 33, c), this (Fig, 53) may join the lower by a miter-joint,
while a cover (Fig, 33, a) two inches high, may join this with a similar
joint.
Fig. 33.
If the upper story shuts over the lower and rests on a shoulder
(Fig, 29) it may still be made to take the same sized frame, by nailing
pieces one-half an inch square to the corners, whose length shall
equal the distance from the rabbet in the lower story to the bottom
board. Now nail to these upright pieces, parallel to the rabbeted
faces below, a three-eighths inch board as wide as the pieces are
long. The top of these thin boards will take the place of the rabbet in
the lower story. This style, which is adopted in the two-story hives as
made by Mr. Langstroth (Fig, 29), will permit in the upper story the
same frames as used in the lower story, while two more can be
inserted. Upon this upper story a shallow cover will rest. Such
covers, if desired, may be made roof-like (Fig, 34), by cutting end
pieces, (Fig 34, b) in form of the gable of a house. In this case there
will be two slanting boards (Fig, 34, a, a), instead of one that is
horizontal, to carry off the rain. The slanting boards should project at
the ends (Fig, 34, d), for convenience in handling. In such covers we
need thin, narrow ridge-boards (Fig, 34, c), to keep all perfectly dry.
These covers look neat, are not so apt to check, and will dry much
quicker after a rain.
Fig. 34.
If we secure comb-honey in crates, and winter out-doors—in

which case we shall need to protect in the Northern States—it will be
convenient to have a box of the same general form as the main body
of the hive, from six to eight inches deep, just large enough to set
over the body of the hive and rest on shoulder-strips, and without top
or bottom; this to have such a cover as just described. Such is the
arrangement of Southard and Ranney, of Kalamazoo, which, on the
score of simplicity and convenience, has much to recommend it.
In the above I have said nothing about porticos (Fig, 29). If hives
are shaded as they should be, these are useless, and I believe that
in no case will they pay. To be sure, they are nice for spider-webs,
and a shady place in which bees may cluster; but such are
inconvenient places to study the wondrous fabrics of the spider, even
were he a friend of the bees, and the most successful apiarist will not
force his bees to hang in idle clusters about the hive.
THE FRAMES.
The form and size of frames, though not quite as various as the
persons who use them, are still very different. Some prefer large
frames. I first used one ten by eighteen inches, and afterward a
shallow frame about seven by eighteen (Fig, 29). The advantage
claimed for large frames is that there are less to handle, and time is
saved; yet may not smaller frames be handled so much more
dexterously, especially if they are to be handled through all the long
day, as to compensate, in part at least, for the number? The
advantage of the shallow frame is, as claimed, that the bees will go
into boxes more readily; yet they are not considered so safe for out-
door wintering. This is the style recommended and used by Mr.
Langstroth, which fact may account for its popularity in the United
States. Another frame in common use, is one about one foot square.
I use one eleven inches square. The reasons that I prefer this form
are, that the comb seldom breaks from the frame, the frames are
convenient for nuclei, and save the expense of constructing extra
nucleus hives, and that these frames permit the most compact
arrangement for winter and spring, and thus enable us to economize
heat. By use of a division board, we can, by using eight of these
frames, occupy just a cubic foot of space in spring, and by repeated
experiments I have found that a hive so constructed that the bees
always cover the combs during the early cold weather, always gives
the best results. As the honey season comes on more can be added,
till we have reached twelve, as many, I think, as will ever be needed
for brood. This was the size of frame preferred by Mr. Gallup, and is
the one used by Messrs. Davis and Doolittle, and many others of our
most successful apiarists. That this size is imperative is, of course,
not true; that it combines as many desirable points as any other, I
think, is true. For apiarists who are not very strong, especially for
ladies, it is beyond question superior to all others.
HOW TO CONSTRUCT THE FRAMES.
In this description, I shall suppose that the frames desired are of

the form and size (Fig, 35) which I use. It will be easy, for any who
may desire, to change the form at pleasure. For the top-bar (Fig, 35,
a) of the frame, use a triangular strip twelve and three-quarter inches
long, with each face of the triangle one inch across. Seven-eighths of
an inch from each end of this, form a shoulder, by sawing from one
angle to within one-fourth of an inch of the opposite face, so that
when the piece is split out from the end, these projections shall be
just one-fourth of an inch thick throughout. For the side pieces (Fig,
35, b, b), take strips eleven inches long, seven-eighths of an inch
wide, and one-fourth of an inch thick. Tack with small brads the end
of two of these strips firmly to the shoulder of the top-bar, taking
pains that the end touches squarely against the projection. Now tack
to the opposite ends or bottoms, the ends of a similar strip (Fig. 35,
d) eleven and a half inches long. We shall thus have a square frame.
Fig. 35.
If comb-foundation is to be used, and certainly it will be by the

enterprising apiarist, then the top-bar (Fig, 36, a) should be twelve
and three-quarters inches by one-quarter by one inch, with a
rectangular, instead of a triangular, projection below (Fig, 36, b),
which should be one-fourth by one-eighth inch, the longest direction
up and down. This should be entirely to one side of the centre, so
that when the foundation (Fig, 36, c) is pressed against this piece it
will hang exactly from the centre of the top-bar. If preferred, the
bottom of the frame (Fig, 36, e) need not be more than half as wide
or thick as described above.
The timber should be thoroughly seasoned, and of the best pine
or white-wood. Care should be taken that the frame be made so as
to hang vertically, when suspended on the rabbets of the hive. To
secure this very important point—true frames that will always hang
true—they should always be made around a guide.
Fig. 36.
Frame, also Cross-Section

of Top-Bar.
A BLOCK FOR MAKING FRAMES.
This may be made as follows: Take a rectangular board (Fig, 37)

eleven and a quarter by thirteen and a half inches. On both ends of
one face of this, nail hard-wood pieces (Fig, 37, e, e) one inch
square and eleven inches long, so that one end (Fig, 37, g, g) shall
lack one-fourth inch of reaching the edge of the board. On the other
face of the board, nail a strip (Fig, 37, c) four inches wide and eleven
and a quarter inches long, at right-angles to it, and in such position
that the ends shall just reach to the edges of the board. Midway
between the one inch square pieces, screw on another hard-wood
strip (Fig, 37, d) one inch square and four inches long, parallel with
and three-fourths of an inch from the edge. To the bottom of this,
screw a semi-oval piece of hoop-steel (Fig, 37, b, b), which shall
bend around and press against the square strips. The ends of this
should not reach quite to the bottom of the board. Near the ends of
this spring, fasten, by rivets, an inch strap (Fig, 37, a), which shall be
straight when thus riveted. These dimensions are for frames eleven
inches square, inside measure, and must be varied for other sizes.
Fig. 37.
To use this block, we crowd the end-bars of our frames between

the steel springs (Fig, 37, b, b) and the square strips (Fig, 37, e, e);
then lay on our top-bar and nail, after which we invert the block and
nail the bottom-bar, as we did the top-bar. Now press down on the
strap (Fig, 37, a), which will loosen the frame, when it may be
removed, all complete and true. Such a gauge not only insures
perfect frames, but demands that every piece shall be cut with great
accuracy. And some such arrangement should always be used in
making the frames.
The projecting ends of the top-bar will rest on the tins (Fig, 33),
and thus the frame can be easily loosened at any time without jarring
the bees, as the frames will not be glued fast, as they would in case
they rested on the wooden rabbets. The danger of killing bees is also
abolished by use of the tins.
When the frames are in the hive there should be at least a three-
sixteenths inch space between the sides and bottom of the frames,
and the sides and bottom of the hive. Even doubling this would do no
harm; though a much wider space would very likely receive comb,
and be troublesome. Frames that fit close in the hive, or that reach to
the bottom, are very inconvenient and undesirable. To secure
against this, our lumber must be thoroughly seasoned, else when
shrinkage takes place our frames may touch the bottom-board.
The distance between the frames may be one-fourth of an inch,
though a slight variation either way does no harm. Some men, of
very precise habits, prefer nails or wire staples in the side of the
frames, at top and bottom, which project just a quarter of an inch, so
as to maintain this unvarying distance; or staples in the bottom of the
hive to secure the same end. Mr. Langstroth so arranged his frames,
and Mr. Palmer, of Hart, Michigan, whose neatness is only
surpassed by his success, does the same thing. I have had hives
with these extra attachments, but found them no special advantage. I
think we can regulate the distance with the eye, so as to meet every
practical demand, and thus save the expense and trouble which the
above attachments cost.
COVER FOR FRAMES.
Nothing that I have ever tried is equal to a quilt for this purpose. It
is a good absorbent of moisture, preserves the heat in spring and
winter, and can be used in summer without jarring or crushing the
bees. This should be a real quilt, made of firm unbleached factory,
duck, or cambric—I have used the first with entire satisfaction for
four years—enclosing a thick layer of batting, and hemmed about the
edges. My wife quilts and hems them on a machine. The quilting is in
squares, and all is made in less than fifteen minutes. The quilt
should be a little larger than the top of the hive, so that after all
possible shrinkage, it will still cover closely. Thus, when this is put
on, no bees can ever get above it. When we use the feeder, it may
be covered by the quilt, and a flap cut in the latter, just above the
hole in the feeder, enables us to feed without disturbing the bees,
though I place the feeder at the end of the chamber, wherein are the
bees, and have only to double the quilt back when I feed. The only
objection that I know to the quilt is, that the bees will fasten propolis,
and even comb, between it and the frames, and this looks bad. A
little care 'will make this a small objection. Mr. Langstroth used a
board above the frames, which Mr. Heddon uses even now. Perhaps
Mr. Heddon never used the quilts. Perhaps his love of order and
neatness caused him to discard them. Still, I feel to thank Mr. A. I.
Root for calling my attention to quilts.
Fig. 38.
DIVISION BOARD.
A close-fitting division board (Fig, 38) for contracting the

chamber, is very important, and though unappreciated by many
excellent apiarists, still no hive is complete without it. I find it
especially valuable in winter and spring, and useful at all seasons.
This is made the same form as the frames, though all below the top-
bar—which consists of a strip thirteen inches long by one inch by
three-eighths, and is nailed firmly to the board below—is a solid inch
board (Fig, 38, b), which is exactly one foot square, so that it fits
closely to the inside of the hive. If desired, the edges (Fig, 38, e, e)
can be beveled, as seen in the figure. When this is inserted in the
hive it entirely separates the chamber into two chambers, so that an
insect much smaller than a bee could not pass from the one to the
other. Mr. A. I. Root makes one of cloth, chaff, etc. Yet, I think few
apiarists would bother with so much machinery. Mr. W. L. Porter,
Secretary of the Michigan Association, makes the board a little
loose, and then inserts a rubber strip in a groove sawed in the
edges. This keeps the board snug, and makes its insertion easy,
even though heat may shrink or damp may swell either the board or
hive. I have not tried this, but like the suggestion.
The use of the division board is to contract the chamber in winter,
to vary it so as to keep combs covered in spring, to convert the hive
into a nucleus hive, and to contract the chamber in the upper-story of
a two-story hive, when first adding frames to secure surplus comb-
honey.
THE HUBER HIVE.
The other type of hives originated when Huber hinged several of

his leaf or unicomb hives together, so that the frames would open
like the leaves of a book; though it has been stated that the Grecians
had, in early times, something similar.
In 1866, Mr. T. F. Bingham, then of New York, improved upon the
Huber hive, securing a patent on his triangular frame hive. This, so
far as I can judge, was the Huber hive made practical.
In 1868, Mr. M. S. Snow, then of New York, now of Minnesota,
procured a patent on his hive, which was essentially the same as the
hives now known as the Quinby and Bingham hives.
Soon after, the late Mr. Quinby brought forth his hive, which is
essentially the same as the above, only differing in details. No patent
was obtained by Mr. Quinby, whose great heart and boundless
generosity endeared him to all acquaintances. Those who knew him
best, never tire of praising the unselfish acts and life of this noble
man. If we except Mr. Langstroth, no man has probably done so
much to promote the interest and growth of improved apiculture in
the United States. His hive, his book, his views of wintering, his
introduction of the bellows-smoker—a gift to apiarists—all speak his
praise as a man and an apiarist.
The fact that the Bingham hive, as now made, is a great favorite
with those who have used it, and is pronounced by so capable a
judge as Mr. Heddon, to be the best movable-comb hive in
existence, that Mr. Quinby preferred this style or type of hive, that the
Quinby form is used by the Hetherington brothers. Captain J. E., the
prince of American apiarists, and O. J., whose neatness, precision,
and mechanical skill are enough to awaken envy; that the Russell
hive is but a modification of the same type, are surely enough to
awaken curiosity and bespeak a description.
Fig. 39.
Frame, Bottom-Board and Frame-Support of

Quinby Hive.
The Quinby hive (Fig, 39), as used by the Hetherington brothers,
consists of a series of rectangular frames (Fig, 39) twelve by
seventeen inches, outside measure. The ends of these frames are
one and a half inches wide and half an inch thick. The top and
bottom one inch wide and half an inch thick. The outer half of the
ends projects one-fourth of an inch beyond the top and bottom. This
projection is lined with sheet iron, which is inserted in a groove which
runs one inch into each end of the end-pieces and are tacked by the
same nails that fasten the end-bars to the top and bottom-bars. This
iron at the end of the bar bends in at right-angles (Fig, 39, a, a), and
extends one-fourth of an inch parallel with the top and bottom-bars.
Thus, when these frames stand side by side, the ends are close,
while half-inch openings extend between the top and bottom-bars of
adjacent frames. The bottom-bars, too, are one-fourth of an inch
from the bottom-board. Tacked to the bottom-board, in line with the
position of the back end-bars of the frames is an inch strip of sheet-
iron (Fig, 39, b, b) sixteen inches in length. One-third of this strip,
from the front edge back, is bent over so it lies not quite in contact
with the second third, while the posterior third receives the tacks
which hold it to the bottom-board. Now, when in use this iron flange

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PREFACE XI DARWIN, WALLACE, AND THE “MEANS

NEL Table of Contents vii

viii Table of Contents NEL

BEHAVIOURAL ADAPTATIONS 239 THE ORIGINS OF ANATOMICALLY

NEL Table of Contents ix

THE HUMAN LIFE COURSE 322 Chapter 16

x Table of Contents NEL

Why This Book Now?

Connecting the Dots ...

A Few Comments for the Student

xii Preface NEL

What’s New in the Second Edition?

Chapter 1 (“Introduction to Biological Anthropology”) sees the addition of two new

Chapter 5 (“What It Means to Be a Primate”) includes new content featuring research by

NEL Preface xiii

Chapter 7 (“Primate Evolution”) features a revised section on primate classification that

Chapter 8 (“What It Means to Be a Hominin”) outlines the major developments in

Chapter 9 (“Hominin Origins: From Ape to Australopithecine”) begins the story of

Chapter 12 (“The Emergence of Anatomically Modern Humans”) presents new genetic

Chapter 13 (“Contemplating Modern Human Diversity”) explores the basis of modern

xiv Preface NEL

Chapter 15 (“Biological Anthropology as Applied Science”) examines several biological

Maps, Photographs, and Illustrations

Retrospection boxes emphasize key ideas

Focus On boxes provide in-depth analysis

Profile boxes—written by some

NEL Special Features of the Book xvii

End-of-Chapter Learning Keys

Key Ideas A bulleted summary of the

Key Terms A listing of the most

xviii Special Features of the Book NEL

Key Critical Thinking

Key Things to Do Next This is a student

NEL Special Features of the Book xix

NEL Ancillaries xxi

xxii Acknowledgments NEL

ABOUT THE COVER

ABOUT THE AUTHORS

NEL About the Authors xxiii

An early choice among the innumerable hives is of course

THE LANGSTROTH HIVE.

CHARACTER OF THE HIVE.

THE BOTTOM BOARD.

For a bottom board or stand (Fig, 32), we should have a single

THE COVER OF THE HIVE.

If we secure comb-honey in crates, and winter out-doors—in

HOW TO CONSTRUCT THE FRAMES.

In this description, I shall suppose that the frames desired are of

If comb-foundation is to be used, and certainly it will be by the

Frame, also Cross-Section

A BLOCK FOR MAKING FRAMES.

This may be made as follows: Take a rectangular board (Fig, 37)

To use this block, we crowd the end-bars of our frames between

COVER FOR FRAMES.