The Study of Animal Behaviour

The aim of this book is to identify the main areas of active discussion about, and research into, the biology of animal behaviour, to describe and assess ways in which these can be studied and using selected examples, to illus­ trate the kinds of results which are emerging. It is not intended to provide an exhaustive review of all we know about animal behaviour, although the examples have been chosen to cover as many as possible of the things that animals do. XlV Preface Acknowledgements I would like to thank C. Swann, M.L.N. Murthy and the Superbrain for typing the manuscript; Linda Partridge, Pat Monaghan, Douglas Fraser and Richard Wilson for constructive criticism of earlier drafts; Alan Crowden for help in planning and producing the book and, particularly, Tim Huntingford for help and encouragement at all stages of its pro­ duction. Acknowledgements are gratefully made to Jim Tulley fot producing plates 1, 2, 3, 7 and 12 and to Michael Hansell for the remainder.

1 Introduction.- 1.1 The ethological approach to the study of behaviour.- 1.2 A brief outline of classical ethological theory.- 1.3 The modern study of animal behaviour.- 1.4 An outline of the book.- 2 The description and measurement of behaviour.- 2.1 Describing behaviour by its function.- 2.2 Describing behaviour by its form.- 2.2.1 Behaviour described as a continuous process.- 2.2.2 Behaviour described as a series of discrete events.- 2.3 Describing and measuring the relationship between an animal and its environment.- 2.3.1 Describing the way a single animal uses space.- 2.3.2 Describing the relationship between an animal and neighbouring conspecifics.- 2.4 What is the point of all this sophisticated analysis?.- 3 The study of the causes of behavioural change.- 3.1 What constitutes a causal explanation of behaviour?.- 3.2 The different kinds of causal explanation.- 3.3 Motivational models.- 3.3.1 Descriptive and explanatory models.- 3.3.2 General, theory-based and specific, data-based models.- 3.3.3 Deterministic and probabilistic models.- 3.3.4 Assessing models.- 3.4 Studying external influences on behaviour.- 3.4.1 Prey catching in toads; behavioural studies.- 3.4.2 Attack in Haplochromis burtoni.- 3.4.3 Incubation in herring gulls.- 3.4.4 Prey catching in toads; physiological studies.- 3.4.5 The role of external stimuli in the control of behaviour.- 3.5 Studying internal influences on behaviour.- 3.5.1 Identifying groups of causally related acts.- 3.5.2 The role of internal factors in the control of behaviour.- 3.6 What is the nature of the mechanisms which cause behavioural change?.- 3.6.1 A probabilistic model.- 3.6.2 A deterministic model.- 3.6.3 Modelling motivation as a hierarchy of decisions.- 3.6.4 Control theory models of motivation.- 3.6.5 Space-state models.- 3.7 Studying the physiological bases of behavioural change.- 3.7.1 Physiological explanations of complex behaviour.- 3.7.2 Alteration in behavioural theory as a result of physiological research.- 3.7.3 Identifying fruitful topics for physiological study by behavioural analysis.- 3.7.4 Ethological concepts and techniques can improve the quality of physiological research.- 3.7.5 Ethological ideas can help in the interpretation of physiological results.- 4 The development of behaviour.- 4.1 Problems with the instinct-learning dichotomy.- 4.1.1 Instincts are defined by exclusion.- 4.1.2 Interpreting deprivation experiments is not simple.- 4.1.3 Learning is not the only way the environment influences development of behaviour.- 4.1.4 The instinct-learning dichotomy confounds three distinct consequences of behavioural development.- 4.2 Why is the term innate still used?.- 4.3 Describing the ontogeny of behaviour.- 4.3.1 The emergence of recognizable behaviour patterns in young animals.- 4.3.2 The time of appearance of behaviour patterns during development.- 4.3.3 The development of motivational systems.- 4.3.4 Physiological changes accompanying behavioural development.- 4.4 Characterizing the factors which influence the development of behaviour.- 4.4.1 Deductions from descriptive studies.- 4.4.2 Experimental studies of behavioural development.- 4.4.3 Experimental studies of the development of song in passerines.- 4.4.4 A quantitative model of the development of social preferences in birds.- 4.5 Classifying the factors which influence the development of behaviour.- 4.6 Some general features of behavioural development.- 4.6.1 Sensitive periods.- 4.6.2 Constraints on learning.- 4.6.3 Functional aspects of behavioural development.- 5 The adaptive significance of behaviour.- 5.1 Sources of evidence about the adaptive significance of behaviour.- 5.1.1 Deduction from the theory of natural selection.- 5.1.2 The context in which behaviour is shown.- 5.1.3 The short-term consequences of behaviour.- 5.1.4 Comparisons between species and populations.- 5.1.5 Measuring the fitness of individuals which differ in their behaviour.- 5.1.6 Experimental manipulation of the consequences of behaviour.- 5.1.7 Deductions from the genetic architecture of behaviour.- 5.1.8 Models of the adaptive significance of behaviour.- 5.2 Difficulties in studying the adaptive significance of behaviour.- 5.2.1 Multiple benefits.- 5.2.2 Costs, benefits and optimality models.- 5.2.3 Frequency dependent costs and benefits; games theory.- 5.3 The state of the art.- 5.4 The adaptive significance of the way animals pattern their behavior in time.- 5.4.1 Rhythms of behaviour.- 5.4.2 The adaptive significance of behavioural rhythmicity.- 5.5 The adaptive significance of the way animals use space.- 5.5.1 Dispersal patterns.- 5.5.2 Deciding where to live.- 5.5.3 What to do where; optimal foraging theory.- 5.5.4 Problems with optimal foraging theory.- 5.5.5 Constraints on use of space resulting from the presence of conspecifics.- 5.5.6 Territorial behaviour.- 5.6 The adaptive significance of an animal’s aggressive responses.- 5.6.1 Games theory and the hawk-dove model.- 5.6.2 Weaknesses of the hawk-dove model.- 5.6.3 Games theory models in sexually reproducing species.- 5.6.4 Conflicts between doves; the War of Attrition model.- 5.6.5 Games with progressive escalation.- 5.6.6 Games with asymmetries between opponents.- 5.6.7 Games in which information is acquired during a fight.- 5.6.8 Games with non-random encounters.- 5.6.9 The contribution of games theory to our understanding of the adaptive significance of the way animals fight.- 5.7 The adaptive significance of an animal’s breeding habits; mating systems.- 5.7.1 Life history parameters.- 5.7.2 Questions about sex.- 5.7.3 Mating systems.- 5.8 Adaptive significance of behaviour accompanying mating.- 5.8.1 Identifying the correct species.- 5.8.2 Preventing mating with conspecifics of the wrong sex.- 5.8.3 Synchronizing the behaviour of potential mates.- 5.8.4 The impact ofintra-sexual competition for mates.- 5.8.5 The impact of selectiveness in potential mates.- 5.8.6 Alternative mating strategies.- 5.8.7 Darwin, natural selection and sexual selection.- 5.8.8 Conflicts of interest between the sexes.- 5.9 The adaptive significance of parental care.- 5.10 The adaptive significance of living in groups.- 5.10.1 Studies of the adaptive significance of group life.- 5.10.2 The benefits resulting from group life.- 5.10.3 The complex costs and benefits of group life; Caraco’s model (1979).- 5.11 Behaviour which cannot be explained by classic natural selection theory.- 5.11.1 An outline of the problem.- 5.11.2 Animals may ultimately benefit from their ‘altruistic’ behaviour.- 5.11.3 Group selection.- 5.11.4 Kin selection.- 5.11.5 Parental manipulation.- 5.11.6 Has altruism been explained?.- 5.12 Overview; sociobiology and behavioural ecology.- 5.12.1 Weaknesses in the sociobiological approach to behaviour.- 5.12.2 Reasons why behavioural differences may not be adaptive.- 6 The phylogeny of behaviour.- 6.1 Sources of evidence about the phylogeny of behaviour.- 6.1.1 The fossil record.- 6.1.2 Ontogeny.- 6.1.3 The behaviour of interspecific hybrids.- 6.1.4 Comparative studies.- 6.2 Some representative behavioural phylogenies.- 6.2.1 Tongue movements in reptiles.- 6.2.2 Predatory behaviour in gastropods.- 6.2.3 Aggressive interactions in fish.- 6.2.4 Hymenopteran social systems.- 6.3 Deriving general principles of behavioural evolution.- 6.3.1 Sound production in crickets.- 6.3.2 Courtship in peacocks.- 7 The role of behaviour in the evolutionary process.- 7.1 The behaviour of other animals as a major selective force.- 7.1.1 The behaviour of conspecifics.- 7.1.2 The behaviour of heterospecifics.- 7.2 Behaviour dictates the selection pressures to which an animal is exposed.- 7.3 The impact of behaviour on population structure.- 7.3.1 Behavioural influences on gene flow between populations.- 7.3.2 Behavioural influences on population size.- 7.3.3 Behavioural influences on panmixis.- 8 Behavioural genetics.- 8.1 The objectives of research into the inheritance of behaviour.- 8.2 Potential contributions of genetics to the study of animal behaviour.- 8.2.1 As an additional research tool in the analysis of behaviour.- 8.2.2 As a source of essential information about behaviour.- 8.3 Quantitative genetics.- 8.4 Techniques for identifying the genetic origin of a known behavioural difference.- 8.4.1 Crosses between behavioural variants.- 8.4.2 Selective breeding from extremes of a behavioural continuum.- 8.5 Screening known genetic variants for behavioural differences.- 8.5.1 Known single gene mutations which influence behaviour.- 8.5.2 Visible differences in chromosome structure with behavioural effects.- 8.5.3 Comparing naturally occurring, isolated populations.- 8.5.4 Correlating degree of relatedness and behavioural similarity.- 8.5.5 Inbred strains.- 8.5.6 Recombinant strains.- 8.6 Characterizing the precise behavioural effects of genetic differences.- 8.7 The mechanisms whereby genes influence behaviour.- 8.7.1 Effects on the structures necessary for performing the behaviour.- 8.7.2 Effects on muscles or neuromuscular junctions.- 8.7.3 Effects on the sense organs.- 8.7.4 Effects on the CNS.- 8.7.5 Effects on the endocrine system.- 8.7.6 Effects on some totally unknown mechanism.- 8.8 Genetic mosaics.- 8.9 Animal behaviour and behavioural genetics.- 9 Applied ethology.- 9.1 Clarification of terms; what is applied ethology?.- 9.2 Ways in which etiological research can be applied to practical problems.- 9.2.1 Direct application of the results of etiological research.- 9.2.2 Using the ideas and concepts of ethology.- 9.2.3 Using etiological techniques.- 9.3 Pest control.- 9.4 Increasing the productivity of commercially important species.- 9.4.1 Identifying suitable species for exploitation.- 9.4.2 Selection for desired behavioural attributes.- 9.4.3 Controlling dispersal and movement.- 9.4.4 Promoting survival.- 9.4.5 Promoting growth.- 9.4.6 Promoting successful reproduction.- 9.5 Animal welfare.- 9.5.1 Identifying suffering in animals.- 9.5.2 Alleviating animal suffering.- 9.6 Conservation.- 9.7 Human behaviour.- 9.7.1 Applying ethological techniques to human behaviour.- 9.7.2 Ethological concepts as a source of hypotheses about human behaviour.- 9.7.3 Making direct use of the facts collected by ethologists.- 9.7.4 Sociobiology and human behaviour.- References.- Author index.- Species index.