Chapter 1

Studies of Behaviour, Welfare and Ecology in Captive

To begin at the beginning Dylan Thomas (1954)

COPYRIGHTED MATERIAL

Fig. 1.1 North American porcupine (Erethizon dorsatum).

Studying Captive Animals: A Workbook of Methods in Behaviour, Welfare and Ecology, First Edition. Paul A. Rees. © 2015 John Wiley & Sons, Ltd. Published 2015 by John Wiley & Sons, Ltd. Companion Website: www.wiley.com/go/rees/captiveanimals

0002254143.indd 3 2/25/2015 10:06:15 PM 2/25/2015 10:06:15 PM ; trans. trans. bc; History 350 (Aristotle, Animals of Introduction Studies HistoryA Short and Incomplete Captive of re Are Animals? What Captive Of all wild animals the most easily tamed and the gentlest is the elephant. It can be and the gentlest is the elephant. Of all wild animals the most easily tamed for instance, the drift and meaning of which it understands; as, taught a number of tricks, It is very sensitive and possessed of an the king. it can be taught to kneel in presence of intelligence superior to that of other animals. in private collections for study and for The Roman emperors kept many animals Park Zoo in London, The first zoo created for scientific purposes was the Regent’s concerned times was largely Victorian The scientific interest in zoo animals in The male camel declines intercourse with its mother; if his keeper tries compulsion, he its mother; if his keeper tries compulsion, The male camel declines intercourse with the by declined being was intercourse when occasion, one On disinclination. evinces covered over the mother and put the young male to her; but, the keeper young male, though the operation was had been removed, when after the intercourse the wrapping still by and by he bit his keeper to death. completed and could not be revoked, of elephants: Aristotle even refers to the training 1910). Alexander the Great captured many exotic animals on his military expeditions many exotic animals on his military Alexander the Great captured 1910). Most of the city they were kept in menageries. and sent them back to Greece where Aristotle the and these would have given states of Greece had such menageries The would not have been able to see in the wild. opportunity to study animals that he History of animal behaviour that of Animals contains descriptions of various aspects in the camel: he discusses mating example, can only relate to captive animals For use in the Roman Games, where they perished in very large numbers. These animals where they perished in very large numbers. use in the Roman Games, the countries were supplied by the Roman military and acquired from the rulers of Thousands of wild animals were 2001). where the animals were endemic (Epplett, slaughtered when the Colosseum was inaugurated in ad 80. years earlier in 1826 Two It was founded in 1828. now known as ZSL London Zoo. The first Stamford Raffles had founded the Zoological Society of London (ZSL). year the first scientific meetings of the Society were held in 1830 and in the same Initially the zoo was open Proceedings of the Zoological Society of London were published. ‘order’ from a only to Fellows of the Zoological Society and others with a written It was fully opened to the public some 20 years later in 1847. Fellow. or any with anatomy and taxonomy and did not extend to concern for their welfare Early enclosures were barren and many detailed consideration of their behaviour. species were held behind iron bars (Fig. 1.2). 1.1 1.1.1 or have those that are, captive animals as I have defined of this book For the purposes animals, zoo and farm This includes humans. the control of under been, recently - but not animals kept in labora and free‐ranging feral animals, companion animals in relatively small areas (or volumes) In most cases they are confined tory conditions. some cases they are In are unable to escape. and walls and other barriers, by fences, remain closely associated with humans. free‐ranging but still 1.1.2 Over 2300 years ago the kept in captivity have a long history. Studies of animals his wrote Aristotle philosopher Greek

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5 Studies of Behaviour, Welfare and Ecology in Captive Animals - . The Act made it an offence to: The . ) enclosure at the former Belle Vue Zoo, Zoo, Vue ) enclosure at the former Belle Ursus maritimus Ursus The polar bear (

An Act to prevent the improper Treatment of Cattle Treatment the improper Act to prevent An The twentieth century saw the founding of many new organisations concerned The twentieth century saw the founding of many new organisations Aquariums (AZA) was founded in 1924 – originally Association of Zoos and The …wantonly and cruelly beat, abuse or ill treat any Horse, Mare, Gelding, Mule, Ass, Ox, Ox, Ass, Mule, Gelding, Mare, abuse or ill treat any Horse, …wantonly and cruelly beat, Cattle… Sheep or other Steer, Heifer, Cow, was founded Animals (RSPCA) of Cruelty to The Royal Society for the Prevention Animal welfare legislation is not yet 200 years old. In England in 1882 Richard In England 200 years old. Animal welfare legislation is not yet ­publish research. – and in Aquariums (AAZPA) Association of Zoological Parks and American as the Welfare Animal 1926 Major Charles Hume founded the University of London Welfare Animal for Federation became the Universities which (ULAWS), Society Both organisations publish their own journals. in 1938. (UFAW) Queen Victoria. The Society has been funding animal welfare research for many Victoria. Queen It now has a Science Group whose funded my own PhD in 1977. indeed, years and, animals, staff study and provide expert advice on welfare issues relating to companion research animals and wildlife. farm animals, and many of these now undertake and with the keeping and welfare of animals, This Act was the first national legislation in the world that punished cruelty to Act was the first national legislation in the world that This animals. as the Society for the Prevention in 1824 in London, more than half a century earlier, was added in 1840 with the permission of ‘R’ The Animals (SPCA). of Cruelty to Martin MP (who earned the nickname ‘Humanity Dick’) was responsible for the pass ‘Humanity Dick’) was responsible Martin MP (who earned the nickname ing of Fig. 1.2 Fig. Manchester Chetham’s permission, with Reproduced Source: c.1850. constructed (UK), Manchester. Library, 0002254143.indd 5 2/25/2015 10:06:16 PM Report of Behaviour Animal , and its and , Behaviour Animal of Journal British ) was first published in 1974. The first issue Animal Behaviour Science) was first published in 1974. Applied Serious studies of animals living in zoos and circuses date from the 1950s and were animals living in zoos and circuses date Serious studies of the of the middle in behaviour appeared concerned with animal Academic journals established a research institute – the Institute The Zoological Society of London there was great concern for the welfare of farm in the UK, Around the same time, Aquariums (BIAZA) was founded in Association of Zoos and The British and Irish of Farm Welfare and published the first edition of Management In 1971 UFAW The new Zoo in Whipsnade has just proved the means of fulfilling the very first and means of fulfilling just proved the Whipsnade has Zoo in The new designed of acclimatization, It was to be a garden Society. of the Zoological original idea animal selected The very first England. to wild life in part to introduce animals at least in the guinea‐fowl. for experiment was The Spectator The in 1931, published Experiments’ ‘Zoo entitled short article In a pioneered by the Swiss zoologist Professor Heini Hediger. He is considered to be the He is considered zoologist Professor Heini Hediger. pioneered by the Swiss Hediger published Zoo. and was once the Director of Zurich ‘father of zoo ’ and Studies of the Psychology books on captive animals including a number of early Animals Wild 1955), in Circuses (Hediger, Animals in Zoos and Behaviour of Captive 1964) and Psychology (Hediger, Zoological Gardens An Outline of the Biology of Captivity: 1969). Circuses (Hediger, Animals in Zoos and and Behaviour of W.H. and Tinbergen The journal Behaviour was founded by Niko last century. and van Iersel (1947) Tinbergen first issue carried a paper by The Thorpe in 1948. the stickleback’ along with three‐spined the in ‘“Displacement reactions” entitled and communication conducted by Schenkel classic study of wolf social behaviour Altmann Basle Zoological Gardens (see Fig. in (1948) years later, Over 25 4.12). often quoted paper in Behaviour in which she (1974) published an important and for observational studies of social behav- described the seven major types of sampling journal The literature. the in used being time that at were that iour the as 1953 in published originally was problems in laboratory rats (Lane‐Petter, first issue included articles on behaviour 1953). cattle (Castle and Halley, 1953) and behaviour in dairy who was of the efforts of Lord Solly Zuckerman, of Zoology – in 1960–61 as a result and served as both Secretary and President a pioneer in the study of primate behaviour Crandall published American zoologist Lee S. the A few years later, of the Society. Captivity while working at the Mammals in Wild of the classic text The Management 1964). Bronx Zoo (Crandall, entitled welfare farm animal into investigation an of report The animals. Intensive under Animals kept of Welfare the into the Committee to Enquire Technical Husbandry Conditions (the Brambell Report) was published in 1965 and led Livestock ‘five freedoms’ that are now widely used as the basis for to the establishment of the 1965). zoos and other captive environments (HMSO, good animal welfare in farms, National Zoo created its zoological In the same year in the USA the Smithsonian’s behaviour and ecology of zoo species. research division to study the reproduction, It is the 1966 as the Federation of Zoological Gardens of Great Britain and Ireland. in Britainprofessional organisation that represents the zoo and aquarium community it promotes research within its member institutions. and Ireland; among other things, The academic journal Applied Animal Handbook. Farm The UFAW Animals: (now deer (Espmark, roe of behaviour social the from ranging subjects on papers contained 1974). 1974) to agonistic and mating behaviour in chickens (Craig and Bhagwat, (Anon., 1931) reported the release of a number of guinea fowl from Whipsnade, Whipsnade, from fowl guinea of number a of release the reported 1931) (Anon., that: proclaiming

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7 Studies of Behaviour, Welfare and Ecology in Captive Animals was first published. It was the first the first It was first published. was Zoo Biology the journal in 1982, years later, Eight in the world, Welfare Animal became the first Professor of In 1986 Mammals in Captivity: Principles and Wild earlier work, Building on Crandall’s European Union April 2002 zoos located within the Member States of the Since increased gained has zoos in living animals of welfare the in interest Scientific concerns about the keeping of some species in zoos have recently stimu- Welfare journal to concern itself exclusively with scientific studies of zoos and zoo animals. zoos and zoo animals. studies of with scientific concern itself exclusively journal to by was co‐authored very first paper and the Maple, Its Terry editor was Dr first behaviour in on social effects of spatial crowding It concerned the Waal. de Dr Frans and Netherlands (Nieuwenhuijsen The in Arnhem Zoo colony at a chimpanzee of otters enrichment study an account of an second paper was The 1982). de Waal, - the engineering of active envi who pioneered Hal Markowitz, co‐authored by Dr 1982). and Markowitz, living in zoos (Foster‐Turley ronments for animals He has of Cambridge. Medicine at the University Veterinary of in the Department Farm 1981), (Broom, of books including the Biology of Behaviour published a number 1990) and Domestic Animal (Fraser and Broom, Behaviour Welfare Animal Behaviour and was Animal Welfare The academic journal 2007). (Broom and Fraser, and Welfare The first volume contained papers first time in 1992. for the ­published by UFAW device for gorillas at London of a puzzle feeder as an enrichment describing the use hens to carriage on the behavioural responses of laying 1992) and ., Zoo (Gilloux et al was Science The Journal of Applied Animal Welfare 1992). conveyors (Scott and Moran, issue included papers on abnormal behaviour in first published in 1998 and its first for use of toys as enrichment the and 1998) Cate, Ten and (van Hoek birds caged 1998). chimpanzees (Brent and Stone, 1996) and then revised as Wild et al., was first published in 1996 (Kleiman Techniques (2nd edition), for Zoo Management Techniques Mammals in Captivity: Principles and 2010). published in 2010 (Kleiman et al., 1999/22/EC of 29 March1999 on the keep- have been required by Council Directive Directive – to have a conservation function. ing of wild animals in zoos – the Zoos discharge this obligation is to participate in One of the ways in which they may Although research is not a of species. research whose results benefit the preservation incentive for zoos this legislation has acted as an important mandatory requirement, research and to co‐operate with universities and in Europe to undertake their own other research organisations. The first conference on the welfare of zoo animals (From attention in recent years. Science and Policy) Welfare Animal Advancing Zoo – Welfare Good Care to Great 2011 under the August Society at Detroit Zoo in was held by the Detroit Zoological Papers from the symposium Welfare. Animal Center for Zoo auspices of the zoo’s were published in 2013 in a special issue of the Journal of Applied Animal Welfare Maple published a major text entitled Zoo Terry In 2013 issue 4). 16, Science (vo1. In June 2013 the Chicago Zoological 2013). (Maple and Perdue, Animal Welfare organised a second (CSAW) Welfare Animal Center for the Science of Society’s Innovations and Future Directions. Welfare: Animal international symposium: Zoo The welfare challenges associated with the keep- lated considerable research effort. of years, ing of elephants in zoos have been of international concern for a number The first was conducted by Clubb and resulting in two major studies in the UK. of Cruelty to Mason (2002) and funded by the Royal Society for the Prevention 2008) and was Animals; the second was conducted by Bristol University (Harris et al., Affairs (Defra) Food and Rural commissioned by the Department for Environment, for Fund International the and RSPCA BIAZA, from funding received also and 0002254143.indd 7 2/25/2015 10:06:17 PM - ypes of Studies Behaviour Studies T The repertoire of responses that an animal may make to changes in its environment. The repertoire of responses that an animal A specific response made by an organism to a specific stimulus. inter the and world external an animal senses the which by processes those ‘…all 1972). (Manning, perceives’ it which changes to responds and body, its of state nal Behaviour may be defined in a number of different ways, for example: of different ways, Behaviour may be defined in a number of theOne of the main functions of the nervous system is to control the activity that they Keeping animals in captivity inevitably restricts the range of behaviours There has been concern for some time about the keeping of a number of other a number of other the keeping of some time about been concern for There has 1.2 this However, wide range of scientific studies. Captive animals may be used for a very undertaken by observation alone and falls into book focuses on research that may be welfare and ecology. one of three types: behaviour, 1.2.1 are often concerned with some aspect of Studies undertaken with captive animals These animals represent an important resource for students and their behaviour. animals in the wild and also because there are researchers who are unable to study occur in captive situations – especially on farms certain aspects of behaviour that only welfare implications. and in zoos – and many of these have 1 2 3 may attempt to explain this We body – along with hormones – and produce behaviour. study thebehaviour in terms of neurophysiological mechanisms or we may simply ofthe behaviour study and manipulate to both possible It is behaviour itself. observed mechanisms. animals without fully understanding the underlying physiological many behaviour studies make little reference to the underlying physiology. Indeed, we do not know what is going on ‘black box’ (i.e. This approach treats the animal as a the causes ofinside) and is useful if we are primarily concerned with understanding behaviour rather than the underlying neural mechanisms (Fig. 1.3). knowledge own sake, for its apart from being of interest However, able to exhibit. are Animal Welfare (IFAW). A review of the latter by the Zoos Forum (now the Zoos the (now Forum Zoos the by latter the of review A (IFAW). Welfare Animal by Group Welfare of an Elephant in the formation resulted Expert Committee) It was charged 2010). (Zoos Forum, UK government the request of the BIAZA at UK of elephants in the welfare and care improvements in a series of with proposing collections. zoological bears and marine mammals – and about – especially chimpanzees, species in captivity which pigs is also unease about the conditions in There in circuses. the use of animals has been addressed by legisla- on intensive farms – much of which and hens are kept of Laying Hens Welfare Pigs Directive and the the in the European Union, tion (e.g. companion in some animals kept in zoos and some in about obesity Directive) – and the amount of space There is even concern about cats and dogs. especially animals, Assembly has Welsh the of the public for their pet rabbits; provided by members of the under section 14 2009) of Rabbits (Anon., Welfare for the issued a Code of Practice many provides welfare animal in interest increased This 2006. Act Welfare Animal may contribute to the debate to conduct new research that opportunities for scientists under human care. about how we should treat animals

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9 Studies of Behaviour, Welfare and Ecology in Captive Animals - - - - stimuli Internal box Black Behaviour ). stimuli External Eudyptes moseleyi Eudyptes The animal as a black box. It is not necessary to understand all of the processes that The animal as a black box.

Causation – What causes an animal to exhibit a particular behaviour? What mech What causes an animal to exhibit a particular behaviour? Causation – What is its motivation? anisms underlie the behaviour? behaviour develop during the lifetime of the Development – How did a particular what is the ontogeny of the behaviour? i.e. animal, Why does an animal exhibit a particular behaviour? How Function (survival value) – fitness? What are the consequences for the animal’s does it help it to survive? Evolution – How did evolution produce a particular behaviour? Over 50 years ago, Tinbergen (1963) published an important paper in which he Tinbergen Over 50 years ago, to considerWhen we undertake a study of a particular behaviour it is important

abnormal behaviour might help a zoo to prevent it, and understanding how a normal abnormal behaviour might help a zoo to prevent it, for normalbehaviour develops might help a zoo to provide appropriate conditions Understanding the function of behaviours is important in appreciating development. and the possible conse the relationship between animals and their environment, backquences of releasing poorly adjusted animals from captive breeding programmes into the wild. 4 In a zoo context we would prob which of the four questions above is being addressed. development and function of behaviour and ably be more interested in the causation, the cause of an understanding For example, less interested in its evolution (Fig. 1.4). 1 2 3 of behaviour is important if we are to understand many of the factors that affect of behaviour is important if we are poor husbandry may have an adverse effect on the For example, ­animal welfare. or the pro breeding programme in a zoo, reproduction of animals used in a captive ductivity of animals kept on a farm. ‘four whys’: the study of animal behaviour: the outlined the four major questions in Fig. 1.3 Fig. be treated like a black The nervous system can occur in the nervous system to study behaviour. input of stimuli and causes the generation of a behavioural response. which receives an box, Rockhopper penguin ( 0002254143.indd 9 2/25/2015 10:06:17 PM - - ., 2007), assessing 2007), ., et al ) shows her teeth to visitors who approach who visitors to teeth her shows ) Pongo pygmaeus Pongo Keeper knowledge may be essential in understanding the origin of some unusual Keeper knowledge may be essential in

Ecological Studies Animal Welfare Studies Animal Welfare

., 2014); the first time such behaviour has been recorded in an invertebrate. 2014); the first time such behaviour has been recorded in an ., The relationship between behaviour and welfare is complex. Mason and Mendl The relationship between behaviour and welfare is complex. is a branch of applied animal behaviour. Welfare studies may Welfare animal behaviour. is a branch of applied Animal welfare science Fig. 1.4 Fig. ( orangutan Bornean This behaviours. She was trained to do this by zoo staff when she had a dental the window to her enclosure. problem and needed regular checkups. (1993) have noted that some responses used to assess animal welfare can be elicited (1993) have noted that some responses used to assess animal welfare can specificity. lack they ones; aversive by as well as events pleasant even or neutral by This often makes behavioural studies very difficult to interpret. 1.2.3 other and Ecology is the study of the relationship between organisms and each captive other and zoos, in living animals Although environment. their and organisms clearly do not have the same ecological relationships that they would have animals, companion animals and animals living in zoos. Most of this concern has been focused Most of this concern in zoos. companion animals and animals living have been recently crayfish However, birds. extent, to a lesser and, on mammals, behaviour following exposure to stress (Fossat demonstrated to exhibit anxiety‐like et al provided that they do not compromise their well be undertaken with captive animals assessing an measuring behaviour under various conditions, They may involve being. or assessing its between different conditions, preference when given a choice animal’s involve comparing the behav for example, Such studies might, physical condition. iours observed when different floor types are available (Meller bodyscoring or 2002) Schmid, and (Jenny device enrichment an of effectiveness the 1998). condition (Gregory and Robins, 1.2.2 of academic study and as a legitimate discipline is now established welfare Animal recent years about the welfare of farm animals, there has been increasing concern in

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Studies of Behaviour, Welfare and Ecology in Captive Animals 11 subjects Animals ossible Study Locations ossible Study dvantages and Disadvantages of Studying Captive Captive of Studying and Disadvantages Advantages P Where might we find animals suitable for study (Figs. 1.5 and 1.6)? Traditional zoos Traditional 1.6)? and (Figs. 1.5 study for suitable animals find we might Where espe- cats make interesting subjects of study, Companion animals such as dogs and of some species that are kept in First‐rate opportunities are available for the study Some studies are concerned with the interface between ecology and behaviour. behaviour. and ecology between interface the with concerned are studies Some There are advantages and disadvantages associated with using captive animals in There are advantages and disadvantages associated with using captive Finding animals in the wild is time‐consuming, behavioural and ecological studies. Often wild ­ and clearly this is not a problem when captive animals are studied. zoo or companion animals is have an unknown history whereas the history of farm, keepers or owners) are likely to likely to be well known and their caretakers (farmers, Natural environments be able to assist with the identification of individual animals. 1.4 Many students undertake studies of captive animals as part of their studies in zoology, in zoology, studies as part of their animals captive of studies undertake students Many Indeed, related discipline. animal welfare or some psychology, animal behaviour, animals under human control for their work. many research scientists routinely use Zoo are obvious places where a wide range ofsuch as ZSL London Zoo or San Diego to these large facilities, In addition researcher. species is potentially available to the specialist collections contain interesting species, there are many other smaller zoos that UK) and aquari- in the Trust Wetlands and Wildfowl of birds (such as those kept by the Some zoos and which keep aquatic species. seal sanctuaries and marine parks, ums, suffi- a have Some colleges purposes. research and teaching for animals keep colleges that they are required to have a zoo licence. cient number and range of exotic species other goats and pigs, horses, cattle, dairy or beef of herds flocks of sheep, have Others research opportunities. farms also offer Working livestock including poultry. goats and horses, as do feral feral populations, cially where they have established along with the deer populations on deer farms and those estab- other feral species, lished in the grounds of many stately homes in the UK. National Primate Research Center, Yerkes specialist research facilities such as the USA (see Section 2.2.3). which is operated by Emory University in Georgia, 1.3 - of inves is worthy and this have an ecology they nevertheless habitat, natural in their the animals. observation of more than mere studies involve Many ecological tigation. of faeces of samples require the collection of parasites may studies For example, and blood; confined and food animals to be ecology may require of feeding studies Such 1983). 1982a, Rees, measured (e.g. to be carefully faeces production intake and in a perfectly normal environment but in a zoo may be problematic ­confinement observational ecological studies that involve purely However, farm environment. example, for might, studies Field environments. captive in possible are methods are and studies of population biology within an enclosure, examine habitat selection from zoo records and studbooks. possible using data evolutionary basis of animal is the study of the ecological and Behavioural ecology Hypotheses in organism to its environment. and its role in adapting an behaviour, are Captive environments assume that behaviour is optimised. behavioural ecology environment can be artificially behavioural ecology because the useful for studying controlled. 0002254143.indd 11 2/25/2015 10:06:17 PM ) ) Pan ) at Reaseheath ) at Reaseheath ); aquariums – Sealife ); aquariums – Sealife Giraffa camelopardalis Giraffa Ailuropoda melanoleuca Bos taurus Ovis aries ), Galloway Forest, Scotland; domestic pets – rabbits – rabbits Scotland; domestic pets Galloway Forest, ), ); free‐ranging farm animals – hill sheep ( ) at Dunham Massey Park, UK; traditional zoos – chimpanzees ( UK; traditional zoos – chimpanzees ) at Dunham Massey Park, Capra aegagrus hircus aegagrus Capra Sources of animals for study. Centre: safari parks – a giraffe ( giraffe a – parks safari Centre: study. for animals of Sources ) at Chester Zoo, UK; college farms – a herd of dairy cattle ( Chester Zoo, ) at

Dama dama Oryctolagus cuniculus Oryctolagus Centre, Blackpool, UK; specialist research centres – giant pandas ( UK; specialist research Blackpool, Centre, at China (photograph courtesy of Chengdu Panda Breeding Research Centre, the Prof. Geoff Hide, Reproduced with permission); deer parks – fallow of Salford. University deer ( ­troglodytes UK. College, Fig. 1.5 Fig. top left: feral animals – a herd of British Clockwise from UK. Midlands Safari Park, West at primitive goats ( (

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Studies of Behaviour, Welfare and Ecology in Captive Animals 13 ave Been Conducted Have ypes of Research T What What There appears to be no published analysis of the nature of the research conducted There appears to be no published analysis (2010) noted that the type of research con- A recent study by Maple and Bashaw papers This study analysed 353 Hardy (1996). undertaken by A similar study was However, there are disadvantages inherent in using captive animals as research using captive animals inherent in are disadvantages there However, so that it is not may be restricted due to safety considerations, Access to animals on farm or companion animals. However, a number of accounts have been published a number of accounts have been published However, on farm or companion animals. on the nature of zoo research. and consists of stud- is primarily non‐experimental, ducted in zoos is relatively stable, the Within mammals. of reproduction and genetics nutrition, behaviour, the on ies (2008) Anderson et al. primates and carnivores are the most popular taxa. Mammalia, found that 35.5% of the 991 articles published in Zoo Biology between 1982 and 2006 13.9% on 23.4% on Carnivora, were conducted on members of the order Primates, Most articles focused 8.8% on Proboscidea and 6.0% on Perissodactyla. Artiodactyla, and longitudinal trends in the content on the behaviour and reproduction of animals, and studies of diet and of the journal included an increase in experimental studies, while studies of behaviour and population biology declined in number. nutrition, published in Zoo Biology between 1982 and 1992 and found that of the 287 papers on a further 5.9% 29.6% were studies of behaviour or behavioural ecology, mammals, of repro- involved behavioural/environmental enrichment and 20.2% were studies Only 3.8% of papers were concerned with genetics or population ductive biology. Captive animals offer opportunities to study a wide range of aspects of their biology, to study a wide range of aspects of their biology, Captive animals offer opportunities paren- hierarchies, social behaviour and dominance such as behavioural enrichment, cog- food selection and feeding strategies, activity budgets, aggression, tal behaviour, and animals between interactions and biology population use, enclosure nition, people (Figs. 1.7 and 1.8). 1.5 nimals Living in Zoos? on Animals Living are fairly unpredictable and outside of the researcher’s control. In contrast, captive captive In contrast, control. researcher’s of the and outside unpredictable fairly are to manipu- be relatively easy which may conditions, live in controlled animals often bedding the types of or altering enrichment device, adding an for example, late by, in a stall on a farm. available this may be natural behaviour is to investigate of the study If the purpose subjects. and normal social are not kept in naturalistic environments impossible if the animals of presence The behaviours. stereotypic developed have individuals if or groups, during keepers (e.g. disturbed by animals are In zoos, may affect behaviour. humans and on farms the animals are disturbed cleaning) and visitors, feeding and enclosure for studies, may be of interest for some Although these interactions by farm workers. interfere with data collection. many others they Researchers night. during the times of the day or at certain recordings possible to make when the zoo is open to the may only be allowed on the premises working in a zoo 10:00 to 18:00), hours a day in the summer (e.g. A zoo may only be open for 8 public. The cost of to 16:00). 10:00 in winter (e.g. and this may be reduced to just 6 hours some For example, a barrier to data collection. studying captive animals may also be to purchase a research pass for access to the zoos require students and researchers (and of travelling the cost near the zoo not live the researcher does if In addition, zoo. possibly accommodation) may be prohibitive. 0002254143.indd 13 2/25/2015 10:06:17 PM s mp s S concerned stables RM Circuses Racehorse farms FA Traditional Ranches & Elephant ca boar, etc. cattle stations FACILITIES Specialist farms, e.g. deer, ostrich, Breeding centre Pre-release pens Pre-release CONSERVATION WORKING ENVIRONMENTS Deer parks s s s rk s ms S ZOO fari pa between 1996 and mid‐2004 and mid‐2004 and 1996 between Biology Zoo Aviarie FERAL STUDIES Aquariu Sa Oceanarium SITUATIONS CAPTIVE ANIMAL Zoological garden Zoological Cats, dogs, pigeons, etc. goats, horses, & FACILITIES RESEARCH & EDUCATIONAL s al anim Specialist shelters Cat & dog Cat & centres SANCTUARIES sanctuaries, e.g. for Sources of captive animals for study. RESCUE CENTRES wolves, elephants, etc. panion Wildlife rescue DOMESTIC farms Primate Center Research ENVIRONMENTS College farms & Com Specialist facilities, animal collections Laboratories e.g. Yerkes National I examined 349 papers published in published papers 349 examined I Fig. 1.6 Fig. biology, and just 2.3% involved wildlife management. The remainder were ­ and just 2.3% involved wildlife management. biology, veterinary med- exhibit design and evaluation (1.2%), with nutrition and diet (3.5%), captive management (24%) and morphology and development (5.6%). icine (5.6%), zoos during identified a significant change in emphasis in the research carried out by Reproductive studies replaced behaviour as the largest cat- 2005). this period (Rees, growth and development (19%) followed by studies of nutrition, egory (34%), field biology, Studies concerned with ecology, and enrichment (17%). and behaviour but there ­conservation and reintroduction only accounted for some 2% of the total, genetics and population biology (10%). was an increase in papers on taxonomy,

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Studies of Behaviour, Welfare and Ecology in Captive Animals 15 Escape reactions hierarchies Parenting behaviour Dominance Abnormal aggression coprophagia Coprophilia & behaviour, e.g. Abnormal sexual mounting objects mating Social behaviour Courtship & Communication apathy Hypoactivity/ behaviour behaviour Reproductive Development of Mate selection Hyperactivity Personality Vomiting & NATURAL regurgitation ABNORMAL behaviour, e.g. BEHAVIOURS BEHAVIOURS Prolonged infantile excessive vocalisation Activity patterns Feeding disorders Intelligence Cognition Activity budgets Self-mutilation . (1992) analysed 302 research projects carried out on mammals by zoo zoo by mammals on out carried projects research 302 analysed (1992) al. et Types of studies of (a) normal and (b) abnormal behaviour that may be undertaken of studies Types e.g. & territoriality Abnormal mother- Memory infant relationships, Stereotypic Ranging behaviour over-preening rocking, pacing Over-grooming/ behaviours, e.g. Biological rhythms Semple (2002) took a different approach and analysed 904 research projects con- projects research 904 and analysed approach different took a (2002) Semple Wiese (b) (a) Fig. 1.7 Fig. on captive animals. ducted in British and Irish zoos. Studies of behaviour were most popular (40%), fol- (40%), were most popular of behaviour Studies zoos. British and Irish ducted in than 5% Fewer (8%). and reproduction (18%) environmental enrichment lowed by genetics or conservation. with ecology, were concerned of projects with of studies conducted compared the number zoos and American North staff in 40 and They found that behavioural within each subject area. the number published undertaken but only 5.3% of studies made up 22.8% of studies behavioural ecology of reproductive physiology accounted Studies in the same period. studies published while studies, published all of 31% almost but undertaken studies of 19.5% only for 0002254143.indd 15 2/25/2015 10:06:18 PM relating Habitat selection Feeding efficiency FEEDING ECOLOGY RESOURCE UTILISATION Food use selection Enclosure ECOLOGY structure Population competition Intraspecific dynamics Population ECOLOGY ECOLOGY COMMUNITY POPULATION competition Interspecific Types of ecological studies that may be undertaken on captive animals. of ecological studies that may be undertaken on Types Parasites density and disease Population What Sort of Study Should I Undertake? Sort of Study What Lankard (2001) listed and categorised 957 publications produced in 1999–2000 by Lankard (2001) listed and categorised taxa being poorly other studies of mammals, Zoo research is heavily biased towards 1.8 Fig. What should you study? As we have seen, a great deal of the research conducted on As we have seen, What should you study? Behavioural research may captive animals is concerned with behaviour and welfare. a phenom- A scientist may wish to study take place for a number of reasons (Fig. 1.7). of wild‐ enon that occurs in the wild but be unable to gain access to a suitable group Although captive animals by for example tool‐use in primates. living animals, it is nevertheless possible to conduct definition do not live in a natural environment, a practical problem ­ A zoo may wish to solve studies of their ecology (Fig. 1.8). 1.6 studies of natural history or fieldwork represented 23.1% of all published studies but studies of natural history or fieldwork only 16.6% of studies conducted. Association (now the Aquarium American Zoo and the member institutions of the conservation/reintroduction ecology/field was category research largest The AZA). behaviour/ethology and (15%) medicine/physiology veterinary by followed (27%), to just 7% of the Studies of reproductive physiology/technology amounted (9%). and nutrition accounted for only 3%. total, and amphib- American reptile A study of the research activity of North represented. to conduct ian departments concluded that they were not realising their potential 1998). formal research and conservation projects (Card et al.,

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Studies of Behaviour, Welfare and Ecology in Captive Animals 17 welfare, for example determining the effects of visitor visitor effects of the determining for example welfare, Some simple studies may not involve the testing of a hypothesis as such. They may They such. as hypothesis a of testing the involve not may studies simple Some to animal husbandry or ­ husbandry to animal wish or a student may reclusive species; of a particularly on the behaviour ­presence of behavioural data. and analysis practise the collection simply to ‘How do such as research question fairly straightforward out to answer a simply set particular the group have individuals within ‘Do or spend their day?’ the animals are that those is, that studies, best the However, individuals?’ other with friendships The next chapter more hypotheses. test one or most scientific, considered to be design. ­considers experimental 0002254143.indd 17 Chapter No.: 1 Title Name: Rees 0002254143 0002254143.indd 18 2/25/2015 10:06:18 PM Comp. by: Vijayakumar E Date: 25 Feb 2015 Time: 10:06:14 PM Stage: Printer WorkFlow:CSW Page Number: 18