DOMESTIC CW ana ~ = BEHAVIOUR and WELFARE ~ sth Edition @)..Domestic Animal Behaviour and Welfare 5th Edition Donald M. Broom Department of Veterinary Medicine and St Catharine’s College, University of Cambridge, UK and Andrew F. Fraser formerly Memorial University of Newfoundland, Canada @) werw.cabl.orgCABI is atrading name of CAB International ABI ABI Nowwonthy Way ry Stet Wallingford Suite 100 Oxgirdibire OXI 8DE Maou UK USA Tels +44 (0)1491 832111 Tak +1 800 $52 3083 (ol ie) Fee: +6 (091491 833508 Enmailsesbionsoeabiong E musi infoweabions Website wunncabions ar 2015, All ghes reserved, No part ofthis plication may be reprodced in any’ hem or by any means DM. Broom and AK clectronieally mechanially; by photocepying cording orctherwiss, without the pre permission of the copyright omnes, ‘A catalogue ecoed foe this baok isavalable from the British Library, Landoa, UK, cation Data Litwary of Congress Cataloging-ia-Pul Broom, Donald M. author ‘Domnzstic animal bekaviour and wellre/ Dondid M. Brooms and Andeew E Fuser. ~ sh edo. pacm, Tacs bibiogeaph ISBN 978.1-78064.539-1 (hardback + alk. piper) — ISBN 978.1-78064.563-6 (pbk alk. paper) 1. Domestic animals-Behavir. 2.Pete-ehavior. 3. Animal welfare. I. Fras, Andrew E, aushor. Il, CA.B, Intemational, isuing body. Il. Tile IDNLM: |. Animals Domestic. 2. lehavios, Animal. 3. Animal Welfare 4. Vaerinary Medicine~mthods. SF 756.7] al rfereoves and indexes S#756.1.876 2015 2014041431 ISN-131 978 1 78064 539 1 (hb) 978 1 78064 563 6 (pbk) Commisioning editor: Caroline Makepeace EdiotilemieantsAlemandra Laimbury Production ofitar Tracy Head Typeset y SPF, Povlicherny, Ilia Printedand bound by Gutenberg Press Taren, Malta Copyrighted materialContents Preface ‘Acknowledgements Section 1 Introduction to Ideas and Measurement | Introduction and Concepts 2. Descibing, Recoiding and Measuring Behaviour Section 2 Fundamental Topics Learning, Cognition and Behaviour Development Motivation Evolution and Opsimalicy Welfare Assessment Section 3. Organization of Behaviour 7 Behaviour'Towards Predators and Social Atackers: Ami-predator Strategies 8 Feeding 9° Body Care 19 Locomotion and Space Occupied 11 Exploration 12. Spacing Behasiour 13 Rest and Sleep Section 4 Reproductive and Social Behaviour 14 General Social Behaviour 15 Human—Domestic Animal Interactions 16. Seasonal and Reproductive Behaviour 17 Sexual Behaviour Section 5 Early and Parental Behaviour 18 Fecal and Parcuriene Behaviour 19 Maternal and Neonasal Behaviour 20. Juvenile and Play Behaviour Section 6 Welfare Topics 21 Handling, Transport and Humane Contol of Domestic Animals 22. Scunaingand Slaughter 23. Welfare and Behaviour in Relation to Disease 24 Abnormal Behaviour 1: Stereorypies 31 46 ol 58 85 90 106 116 122 125 133 143 153, 156 161 7 188 202 215. 233 237 246 Copyrighted material23 2 28 Contents Abnormal Behaviour 2: Seldirected and Eavizonment-ditected Abnormal Behaviour 3: Addressed to Another Individual Abnormal Behaviour 4: Failure of Function Abnormal Behaviour 5: Anomalous Reactivity Section 7 Welfare of Various Animals 29 30 31 3 33 34 35 36 Be 38 39 Glossary Referenc Ihe Welfare of Cattle The Welfare of Sheep and Goats “the Welfire of Pigs ‘The Welfire of Poultry “The Welfise of Farmed and Pet Fish ‘The Welfare of Animals Kept for Fur Production The Welfare of Horses, Other Equids and Ocher Draught Animals “The Welfare of Farmed and Pet Rabbits “The Welfire af Dogs “The Welfare of Cats “The Welfare of Other Pet Animals es Subject Index Author I index 234 258 267 274 279 203 297 318 328 335 340 344 37 353. 358 361 371 437 449Preface All of those who have an interes in livestock produc- tion oF companion animal management and breeding, including every farmes, pet owner and veterinary sur- goon, need «© know about domestic animal behaviour im order that they ean carry out their jobs and care for their animals properly. All of these people and all con- sumers of farm animal products have to consider their moral stance in relation to domestic animal welfare and require precise information about that welfare in order to do %, This book isa comprehensive guide to the be- haviour and welfare of domestic animals. Ie provides practical information for those involved with farming, pet animals and veterinary work and reviews scientific information about the assessment of animal welfare, ard the evaluation of the eflects on animals of genetic selection and of different management methods and housing conditions Assessing welfare necessarily in- vyelves measuisment of physiology, slisease sate and production as well as behaviour. Ethology developed rapidly after 1950 and became fully established as a par of academic courses on bio- logical subjects hy 19810-1999, Fshology is the obser tion and derailed description of behaviour with the ob- jective of finding out how biological mechanisms function. Some yeats fier its expansion in zoology and psychology teaching, animal behaviour has become part of veterinary and animal science or agriculture courses. Inxiced, it is an essential part of professional knowledge forall who use and eare for animals and isa subject of widespread interest to the general public “There have been exciting developments in our know- ledge of the behaviour of domestic animals and these are incorporated in chis new edition, Animal wellare science arose as a scientific discipline afier 1980 and the major increase in its academic study is sill occurring, The welfie of an individual i ite state ax regards its attempts 9 cope wich its environment. This in- cludes the sate of mechanisms in its brain and other parts ofthe body including feelings and system for dealing with dlisewse. Coping with the environment involves a wide range of interacting biclogical systems and so the subject is a fundamental biological science. les development kas been rapid. “There have long been rexearchers working on animal health, which isa key pare of animal welfare. Other than these, there were only 20-30 animal welfare scientists before 1990 but there are now several thousand, In recent years, che EU Welfare Quality and Animal Welfare Indica tors (AWIN) projects have provided much valuable infor- mation abour welfare outcome asessment in practical situations, The use of animul-based welfare outcome indi= caters by wellare inspectors, veterinarians and farmers has developed, ficilisted by the European Food Safety Au- thority pons and guidelines. This dynamic ares of in- creasing knowledge has changed what we know ahouit the many animal welliretopies discussed in this book The evaluation of behaviour and welfare is also im- portant in human biology and mediciae. ‘he detailed scientific study of behaviour was developed lor non-humans hefore ie was applied to humans. It prom vides information that cannot be accurately obsained by asking people questions. The methodology devel- oped in animal welfare science is also being applied vo human psychiatry and medicine. ‘The rapidly develop- ing field of anchrozoology, which concerns inceraetions beeween humans and ocher species, has depended greatly on studies involving che behaviour and physi- ology of petsand their owners, ‘A further area of rapid development in science has been the study of brain function. Since the brain has a controlling effect on all behaviour, and alll methods of coping with the environment of an individual ate cenired in the bra . neuroscience ~ including, studies of cognitive functioning ~ is of central importance to the subject of this book. Seucles of hov clever animals are, and of the representations of events that they have in their brains, ater the way in which people think of the animal species. These studies are direetly elevant to understanding animal needs and hence welfare. Some of Copyrighted materialFreface the burgeoning literature on animal eognision feelings, emotion and sentience is discussed in this new edition, This book extends the coverage of the fourth edition im that much new information is provided, for example con equids other than horses and on draught animals. Chapters have been added on welfare during scunaing and slaugheer, the welfae of sheep and goats, and the welfare of addicional pet species. Hence all major farmed animals and companion animals are considered. Both authors have phnned this updated edition and provided new illustrations, while the text cevision has heen ear ried out by Donald Broom. The concepts of behaviour Department of Veterinary Maticine and StC and welfare are introduced, and then behaviour de- scription, learning, cognition, motivation, evolution and welfare assessment are considered. Sections on the varinus aspects of individual, social and reproductive behaviour then follow. Welfare during transport and slaughter, welfare and disease, and the various kinds of abnormal behaviour are then described. ln the final 11 chapters, the wellare of different species is discussed, “The book js illustrated with many photographs and, since an understanding of the meaning of concepts is sive glossary. Donald M. Broom MA, PhD, ScD, Hon DSc, HonDe Profesor of Animal Wi fare (Emeritus) Centre for Animal Welfare and Anthrozoology svine's College University of Cambridge, UK Andrew F. Fraser MRCVS, MYSc, FIBiol Formerly Professor of Susgery (Veterinary) Memorial University of Newfoundland, CanadaWe thank first the many authors, mentioned in the reference list, whose published work forms the basis for this book. We thank especially che following, some now lcceased, who, for one or more of the editions of the book, have given encourgement 10 the authors, provided illastraive material or helped in book preparation: A.M. Atchison, Jack Albright, Graham Arnold, Sob Baldwin, DD, Bieger, M. Bicget, Harty Bradshaw, Sally Broom, Alex Brownlee, DD. Dooley Ingvar Ekesbo, Michael Fox, David Acknowledgements Frases, C. George, Temple Grandin, Scephen Hall, Ruth Hanrison, H. Hastie, Ken Johason, Ron Kilgous, Erina Kirby, A. Lidejoho, Frank Loew, Mike Mend, Maia-Alice Moreira, B. Payton, Anthony Podberscek, Irene Rachie, Hans-Hinrich Sambraus, James Serpel, Flizabeth Shilo Waker, Barbara Sommerville Alex Stolba, Ray Swicklin, ‘Sue Tennang,TarjeiTennessen, C. Thome, KlausVestergaani, Piet Wiepkema and David Wood-Gush, We are gracefil for the efforts of the staff at CABI to improve this book, Copyrighted materialAsignificane part ofour interest in the world around ws is focused on non-human animals, not jusc as sources bur also because of our empathy for them and our fas- ination with what chey can do and how they function, (Podberscek etal, 2000;Serpell. 2004: D. Fraser, 2008: Broom, 20145). ‘The farming of animals has played an imporrant pare in the development of human civiliza- tion, Food, clothing and transport are obtained by man from a wide variety of species. It is thought thae hu mans have had an even longer relationship with wolves, or dogs as we now call one form of the Middle-Eastern grey wolf (Clutton-Brock, 1999; von Holdt et al, 2010). This relationship, and we may speak of wolves domesticating humans just as correctly as humans do- mesticating wolver (Broom, 2006a), seems likely +0 have been mutually beneficial to both species, as has other domestication (see Chapter 5). Dogs, cats, other Jong been treated ascompanionsand viewed with affec- tion by chose whose job it was to care for them, Good stockmanship has always involved knowing how to re spond to the behaviaurof animals when handling thers or identifying their problems By the sartof the 20:h century, farm animal usehad increased with the expansion of the human population and consumption of animal products. Animals began to bbe kept in concentrated populations and, priorto 1970, inensive animal husbandry had arrived in the form of close confinement for cattle, pigs and poultey under new husbandry systems, The innovations in management ate characterized principally by larger livestock numbers kept cogether in markedly reduced space. Such condt- tions have effects on disease transmission and they re= quite considerable physiological and behavioural adap- tation by the animals (Broom, 2006a). It was assumed that the animals could adapt to the environmental re strictions, bur both adapration ang failure to adjust have come 0 be recognizable when welfare is assessed, As Mason (2010) explains, animals kepr in captivity vary ©DM.Brocm and AF. Fraser 2015. Domestic Animal Behaviour and Welfare, Sth Edition (D.M. Broom and AF. Feser) Introduction and Concepts greatly in how well they can adape and in the extent of poor welfare resulting from the captivity. Knowledge of the behaviour of livestock under intensive husbandry systems is needed to asvess these systems of manage- ment, just as information about behaviour is neeced to manage animals extensively. This knowledge can then be applied in the agriculture industry in order to improve production and welfare. Many animal husbandry prob- lems are nor soluble by investigation of nutrition, body physiology or disease contol, hur require investigations of the behaviour of the animals before progress can be made towards a solution The attirude of people wo dogs (Serpell, 1995) ranges fiom viewing the dog as a cause of vicious and unprovoked attacks on childien, of pollution of our screets and of serious disease risks, co considering the dog as « family member, an archetype of affeccionate fidelity and a source af unconditional love. People who use animals as companions. of for some form of work orenteriainment, are aware of the behaviour of the ani- mab, In some cases, the behaviour is not what people wane and is viewed as « problem. In other cases the behaviour is the reason why the animal is usefal, shether or not this use results in good welfare in the animal, Behaviour can bean indicator af good or poor welfare in aay animal. The term ethology means the observation and detailed desexiption of behaviour with the abjective of finding ovr how biological mechanisms The scientific study of animal behaviour has pro- ceeded very rapidly during the lax 50 years. Some of the changes in ideas that have occurred during this de- velopment are described hy Jensen (2009). There ha been substantial recent advances in the precision of be~ haviour description and the understanding of behav ious ongeniattion in lation to physiological and av tionary processes. Modern techniques in ethology and in experimental psychology mean that we now have a much moze extensive knowledge of sensoxy analysis,Domestic Animal Behaviour and Welfare motor control, hormonal effects, motivation, body: maintenance behaviour in good and difficult condi- tions, reproductive behaviour and soeial structure. This knowledge and many other methods of animal welfare assessment are used in animal welfare science and ap- plied to domestic animals, Sentience and Animal Protection Animals vary in the extent to which they are aware of themselves (DeGratia, 1996) and of cheir incr with their environment, including dete abi perience pleasurable states such as happiness and aversive states such as pain, fear and grief. This cap= acity may be referred to as their degree of sentience The term sentience has generally been used to mean that the individual has the capacity to bave feelings (Kirkwood, 2006). ‘This raises the question of what abilities are needed in order to have this capacity Sentience implies a range of abilities, noe just having feelings. A definition is: a sentient being ft one thas har tome ability to evaluate the actions af others in relation 10 itself and third parties: to remember some of its own actions and their consequences; to asses risks and bene fit; to have some foelingss and t» have some degree of awareness. This. definition, slightly modified after Broom (2006¢), and various aspects of sentience are discussed further by Broom (2014). Human: opinion ‘as to. which, individuale-of our own and other species are sentient has generally changed over time in well-educated societies to encom- poss firs all humans instead of just a subsct oFumans, and then: certain mammals that were kept as compan ions; animals that seemed most similar to humans such as monkeys; the larger mammals; all_ mammals, all warm-blooded animals: then all vertebrates: and now some invertebrates, The general public has been ready to accept some who have collected information about the abilities and functioning of che animals. Animals that are shown to be comples in their organization, capable of sophisti- cated learning and aware are generally respected mare than those that ate aot, and such animals are ess likely to be treated badly. However, some people view animals solely on the basis of their effects on, or perecived (cx- trinsic) value to, humans and have little concern for them as individuals. Ethics Something is moral if i perasins to right rather than sera and ethics is the saudy of moral iauer Humans and other animals, especially social animals, have many biological mechanisms that enable them co behave in a. moral way. It is not possible to live successfully in a social group unless the individuals have the ability « avokd harming others (see Fig. 11) and pethops to collborate. Asa consequence. morality has evolve oured genes that promote abilities such as recognition of individ and natural selection bas f ualsandl memory of moral and immoral sctions (de Wal, 1995; Ridley, 1996; Broom, 2003, 2006). Two other bio logical mechanisms which may promote moral actions are empathy and compasion (Wirbel, 2009). People are more likely to feel empathy with fellow humans andl other animals peresived to have a capacity for feelings similar co those of humans. ‘They are likely 10 show compasion co those whom they perceive to need compassion ‘Two underlying approaches in thinking about how to behave towards people or other animals are known as deontological and consequentialist. The deontalogical in which the structure is 2 set of daties pertinent co all individuals. Hence, the individual should assess what action duty dietstes using rational thought and cary cout that action Consequentialism in ethics implies thatthe extent 0 which an act is morally ight is determined solely by the goodness of the act's consequences. This approach was extended into utilitarianism by J. Mill (1843), who ar- sgued thatthe right actor policy is that which will resulcin the maximum utliey or expected balance of satisfaction fected. minus dissatisfaction, in all che sentient beings a Although many aspects of utilitarianism are helpial when deciding what is morally right, sa general approach viewed as incomplete (Broom, 2003). Acting in such a way that general happiness or genctal good is pro mowed will be entirely desirable in some circumstances, but following such a philosophy implies that decisions ate taken only on the basis of the average ot overall good of collections of individuals. This view does not ake account of the fact that humans and other animals interact with Jhiake. The’ mechaniene underlying moral codes are based on effects on individuals as well aon collections of individual, An example of the flaw in the extrem utilitarianism appro is dat, allow ing chis approach, an individual could be caused extreme pain or ether poor weliare, or could be killed if che overallIntroduction and Concepts Fig. 1.1. These Longhorn cattle carry dangerous weaponsand spend much time close together but theyare very carefu not io harm fone another (phetagiesh DM. Broa) fice on a collection of individuals was good. This ndivid- ual might be a dangstous criminal or an entirely innovent person, but should they he tortured, caused prolonged misery or killed? Most people would noc wish an innocent person to be killed, however great the resulting good, and those who heststed on the inte might be swayed comands thac view ifthe person were their neighbour, their mother or themseles. ‘Criticisms of the utilitarian position have been made by many, including Williams (1972) and Midgley (1978). Those who would consider themselves deonto- logical ethicists would maintain that certain rights, rules, prineiplesor obligations take prexedenee over util ity. However, a wholly deontological approach also has flaws. Elements of both deontological and utilitarian ap- proaches are necessary in order to act in a moral way. Ieis my view (Broom, 2003, 2006b, 2010, 20146) thac all human behaviour and laws should be based on. the obligations of each person to act in an acceptable way towards each other person and to each animal with which we interact. IF we use a living animal in a way thae gives us some benefit, we have some obliga- tions to that animal, We have some obligations to any individual considered to have an intrinsic value and other obligations whenever we have concern for the individual’ welfare. We should avoid causing poor welfare in the animal excepe where the action leads to anet benefit to that animal. In some cases, we might cause poor welfare because of a net benefit when we also take account of otheranimals, including humans, or of the environment, Ie is beter for stiategjes for living vo be based on our obligations rather than to involve the concept of rights. This is because many so-called rights can result in harm to others. Argumentsabout the importance of freedom to contiol one's life led to the idea thac such, freedom is aright’ which all should have. Strong pro: ponents of rights structare for determining what are proper actions regard the stated rights as absolute, so they cannot be mitigated by other citcumstances. A key issue here isthe establishment of what is & right. There are few so-called rights that would be accepted as valid in all circumstances. ‘The oft-proclaimed rig free speech can cause great harm to certain individuals and hence can be morally wrong, in my view, as can the ‘right’ to drive a car at fast as you wish, ar to camry a gun, or to determine the sex of your children, ‘The concept of rights causes many problems. All behaviour and laws should be based on the obligations of each person to act in an acceptable way towards each other person or other sentient individual. Arguments based on obligations are better than any attempts to assert a ‘right’, Lawsand other such statements should provide guidelines for the behaviour of each person rather than stating what the individual who is the object of an ac~ tion can demand, ‘The ethics aspects of animal behaviour research are discussed by M. Dawkins and Gosling (1992).Domestic Animal Behaviour and Welfare Sustainability and Animal Welfare Members ofthe public now ask whether or not any system for exploiting resources is sustainable (Aland and Madec, 2009). "The fact that something. is profitable and these is demand for the product is not now sufficient rason for the continuation of production (Broom, 2010a). A gstem recur is sustainable if 1 18 acceptable wore and if its expected future efecs ave acceptable. in particular in relation so resource availbiley comequences of functioning aid mor- abty of actim (Broom, 20014, 2014b), An animal wage spccrn might not he nussinable because: (i) ix ievolver 40 much depletion of a resource that his will become une available to the system; (i) a product of the system accu mutates co a degree that prevents the functioning of the system: or fii) members of the public find an action in- volved in it unacceptable. Where there is depletion of a reouree of accumulation of a product, the level at which this is unacceproble, and hence the poin at which the sys tem is unsustainable, is usually considerably lower than that at whieh che production system itself fa A.gystem could be unsustainable because of harms to the perpetrator other people, the environment or other animals. Agriculture is the main aetiviy on mest of the land in the world and has resulted in much loss of bio~ diversity, so ie must be considered in most discussions of wildlife conservation (Balmfoxl ef al, 2012) and the sus tainabiity of many of its practices is questionable. Cur- rently in the world, animal welfwre is one of the major reasons why certain methods used in animal production and management are unsustainable. Aspects of produc tion, such asthe welfue of theanimals, are now imporant sehen evaluating the quality of animal products, Con: sumefs expect supermarket companies and restaurant chains to guarantee that the methods used 0 produce products are nor negative in relation to animal welfae or any other aspect of sustainability and product quality BF fortsare now being made o design animal production sys tems that ate sustainable in all ways. Welfare Concepts The wientiie swidy of animal welfare hardly existed 40 years ago, with the exception of studies of animal eisease, bbe has developed rapidly during the las 30 years(Broom, 2011). The concepts have been refined and a range of methods of asessment have been developed. Substantial challenges to animal functioning include chose resulting fiom: (9 pathogens (i) tissue damage (i) attack ot threat of attack by 2 conspecific or predator; (iv) other socal competision: (¥) complexity of information procesing in a situation where an individual receives excessive simula- tions (vi) lack of key stimuli such as a teat for a young, mammal or social eomact cues; (vi lack of overall stimu: lavion; and (i) inability to control interactions with the environment. Hence, potentially damaging challenges may come fiom the environment outside the body (eg, many pathogens or causes of tissue damage), or from within it e.g, aniety, boredom or frustation that come from the environment of a control system). Systems that respond to or prepare for challenges are coping systems and coping means having control of mental ara boily tabity (Broom and johnson, 1993), Coping, attempts may be unsuccessful in that such control is not achieved but, as soon as there ie control, the individual is coping. Systems for attempting 1 cope with challenge may respond to short-term ot long-term problems, or sometimes to both The responses «chal lenge may involve activity in parts of the brain and vari ous endocrine, immunological or other physialogical responses as well as behaviour. However, the more that thac these various types of response ate intenlependent For example, noc only do brain changes regulate bodily coping responses, bur adrenal changes have sevetal com sequeness for brain functions lymphocytes have opioid receptors and a potential for altering brain activity; and heart rate changes can be used to regulate mental state hence, farther responses, Some coping systems include feelings as a part of their functioning, for example pain, fetr and the vari ouskinds of pleasute, all of which are adaptive (Broom, 1998: D, Fraser, 2008: Chapter 4). Bad feelings that continue for more than a short period are referred 10 as suffering. Other high or low-level brain processes and other aspects of body functioning are aso a par of at- tempts © cope with challenge. In order to understand coping systems in humansand other species, itis neces- sary to study a wide range of mechanisms including complex brain functioning, as well as simpler systems. Investigations of how easy oF difficult isis for the indi- vidual to cope with the environment, and of how great is the impactof positive or negative aspects af the envir~ ‘onmenton the individual, are investigacions of welfare. If, ec some particular time, an individual has ne prob- lems to deal with, clue individual is likely 1 be in a good state, including good feelings and indicated by