Many animals, including humans, acquire valuable skills and knowledge by copying others. Scientists refer to this as social learning. It is one of the most exciting and rapidly developing areas of behavioral research and sits at the interface of many academic disciplines, including biology, experimental psychology, economics, and cognitive neuroscience. Social Learning provides a comprehensive, practical guide to the research methods of this important emerging field. William Hoppitt and Kevin Laland define the mechanisms thought to underlie social learning and demonstrate how to distinguish them experimentally in the laboratory. They present techniques for detecting and quantifying social learning in nature, including statistical modeling of the spatial distribution of behavior traits. They also describe the latest theory and empirical findings on social learning strategies, and introduce readers to mathematical methods and models used in the study of cultural evolution. This book is an indispensable tool for researchers and an essential primer for students.

• Provides a comprehensive, practical guide to social learning research
• Combines theoretical and empirical approaches
• Describes techniques for the laboratory and the field
• Covers social learning mechanisms and strategies, statistical modeling techniques for field data, mathematical modeling of cultural evolution, and more

Acknowledgments ix
1 Introduction 1
What Is Social Learning? 3
Social transmission 5
Imitation 9
Innovation 9
Why Study Social Learning? 11
Summary of the Book 14
2 A Brief History of Social Learning Research 16
The Evolution of Mind 17
Social Learning Mechanisms 20
Animal "Culture" 24
The Diffusion of Innovations 27
Child Development 28
Cultural Evolution 30
Conclusions 31
3 Methods for Studying Social Learning in the Laboratory 33
Traditional Social Learning Studies 35
Alternative Experimental Approaches 38
Transmission chain studies 40
Diffusion studies 47
Innovation 50
The Biological Bases of Social Learning 52
Neuroendocrinological studies 53
Social learning of fear 54
Neural mechanisms of observational learning 56
Conclusions 61
4 Social Learning Mechanisms 62
A Classification of Social Learning Mechanisms 63
Stimulus enhancement 65
Local enhancement 65
Observational conditioning 68
Response facilitation 69
Social facilitation 70
Imitation 70
Observational R-S learning 77
Emulation 77
Opportunity providing 78
(Inadvertent) coaching 79
Distinguishing Social Learning Mechanisms 80
Stimulus enhancement 80
Local enhancement 81
Observational conditioning 83
Response facilitation 85
Contextual imitation 87
Production imitation 89
Observational R-S learning 92
Emulation 93
Opportunity providing 94
(Inadvertent) coaching 96
A Pragmatic Approach to Characterizing Mechanisms of Social Transmission 96
Teaching 98
Summary 104
5 Statistical Methods for Diffusion Data 105
Diffusion Curve Analysis 106
Network-Based Diffusion Analysis (NBDA) 108
Inclusion of individual-level variables 112
Model selection and inference 114
Modeling multiple diffusions 115
Choosing a social network 120
"Untransmitted" social effects 121
Related methods 122
Is NBDA realistic? 123
Examples 125
Spatial Spread of a Behavioral Trait 125
Wave of advance models 125
Other approaches for spatial data 127
Summary 128
6 Repertoire-Based Methods for Detecting and Quantifying Social Transmission 129
The Group-Contrasts Approach 130
The Method of Exclusion 133
Basic and advanced methods of exclusion 133
Methods for assessing the genetic hypothesis 134
Methods for assessing the ecological hypothesis 143
Further problems with the method of exclusion 149
A Model-Fitting Approach 150
A matrix regression approach 150
Examples 151
A return to group comparisons 154
A Causal Modeling Framework 155
Relationship to the matrix regression approach 164
Ruling out homophily 164
Ruling out unknown ecological variables 167
Relationship to the method of exclusion 168
Conclusions 168
7 Developmental Methods for Studying Social Learning 172
Observational Data 173
Describing the developmental process 173
Modeling probability of acquisition 177
Modeling time of acquisition 179
Modeling proficiency of trait performance 184
Modeling option choice 185
Limitations of observational data 188
Experimental Manipulations 188
Diffusion experiments 189
Manipulation of social experience 191
Translocation studies 193
Conclusions 195
8 Social Learning Strategies 196
Why Social Learning Is Strategic 196
"When" Strategies 203
Copy when established behavior is unproductive 203
Copy when asocial learning is costly 205
Copy when uncertain 210
Copy when prior information is outdated 212
Copy when dissatisfied 213
Is copying a first or last resort? 214
"Who" Strategies 215
Frequency-dependent biases 216
Success biases 221
Kin and age biases 225
"What" Strategies 226
Random Copying 227
Statistical Methods for Detecting Social Learning Strategies 229
Meta-strategies, Best Strategies, and Hierarchical Control 232
9 Modeling Social Learning and Culture 235
Introduction 235
Why model? 235
Operationalizing the culture concept 237
Parallels between biological and cultural evolution 238
Theoretical Approaches to Social Learning and Cultural Evolution 239
Population-genetic style models of cultural evolution 239
Population-genetic style models of gene-culture coevolution 241
Neutral models and random copying 246
Social foraging theory 250
Spatially explicit models 251
Reaction-diffusion models 253
Agent-based models 254
Phylogenetic models 255
Conclusions 258
10 Conclusions 260
References 265
Index 301