Stereotypic pacing in Siberian tiger (Panthera tigris altaica) and the effect of interspecies presence: A case study

Similar documents
ORIGINAL RESEARCH PAPER

Tigers Stereotypic Pacing and Enrichment

February 1, 2018 Robert Gibbens, D.V.M. Director, Animal Welfare Operations USDA/APHIS/Animal Care

Meredith J. Bashaw a, Angela S. Kelling b, Mollie A. Bloomsmith b & Terry L. Maple b a TECHlab, Zoo Atlanta, and Center for

Interactions between large felids and humans in zoos: animal, keeper and visitor perspectives. Monika Szokalski. Doctor of Philosophy (Psychology)

Comparison of Several Types of Enrichment for Captive Felids

Role of Temperature and Shade Coverage on Behavior and Habitat Use of Captive African Lions, Snow Leopards, and Cougars

Abnormal Repetitive Behaviours

Assessment and Opinion of Health and Welfare of Animals at Spring River Park and Zoo Christine Capaldo, DVM March 9, 2017

Frustrated Appetitive Foraging Behavior, Stereotypic Pacing, and Fecal Glucocorticoid Levels in Snow Leopards (Uncia uncia) in the Zurich Zoo

Zurich Open Repository and Archive. Feeding live prey to zoo animals: response of zoo visitors in Switzerland

Shelter Guidelines Project. Shelter Guidelines - Content

Director Stearns Zoological Rescue & Rehab Ctr. Inc Meridian Ave. Dade City, FL 33525

If they don t have a box they do their best to hide

Outline CANINE ENRICHMENT: STRESS REDUCTION FOR HAPPY AND HEALTHY SHELTER DOGS. What is Environmental Enrichment?

Waiting for a Forever Home

NORMAL AND ABNORMAL BEHAVIOUR. Course in Animal Welfare 2017

Effects of interactive visitor encounters on the behaviour and welfare of animals commonly housed in Australian zoos

Unit 3 Sustainability and interdependence Sub Topic 3.4: Animal welfare

Keywords: 1ZA/Acinonyx jubatus/cheetah/hair sample/method/monitoring/scent

Behavior of Captive Humboldt Penguin (Spheniscus humboldti) Chicks in Response to Environmental Enrichment

3/14/2017. Shelter Cat or Sheltered Cat? The Road Map. April 9, Miranda K. Workman, MS CABC, CBCC-KA, CPDT-KSA. How Training Play Case Studies

EXOTIC PETS The landscape has changed

TOMPKINS COUNTY SOCIETY FOR THE PREVENTION OF CRUELTY TO ANIMALS

Enrichment for Indoor Cats Ilona Rodan, DVM, ABVP Dipl., Feline Milwaukee Veterinary Medical Association November 2014

God s Grace Canine Rescue, Inc. Adoption Questionnaire. Your Name: Address: Home Phone: Cell Phone: Employer: Work Phone:

Effects of construction noise on behaviour of and exhibit use by Snow leopards Uncia uncia at Basel zoo

Mental stim ulation it s not just for dogs!! By Danielle Middleton- Beck BSc hons, PGDip CABC

Animal Welfare Assessment Transfers Checklist

Social Housing and Environmental Enrichment Policy

Council for Innovative Research

Keywords: Acinonyx jubatus/breeding/captivity/cheetah/management/off-exhibit

Scientifically evaluating welfare in commercial breeding kennels: does high volume preclude good welfare?

AN INVESTIGATION INTO THE WELFARE STANDARDS OF ZOOS IN MALAYSIA

March 16, Guide's space recommendations as a minimum while always recognizing that performance standards also must be met.

What is a tiger? Tigers are felids (members of the cat family). They are in the genus Panthera.

Animal Study: Adelaide Zoo

Behaviour of cats and dogs

Talking about zoos, animal well-being and education with Jon Coe

RURAL INDUSTRIES RESEARCH AND DEVELOPMENT CORPORATION FINAL REPORT. Improvement in egg shell quality at high temperatures

Use of the Animal Welfare Assessment Grid to assess the life time experience of animals and cumulative severity of procedures

Intact Carcasses as Enrichment for Large Felids: Effects on On- and Off-Exhibit Behaviors

FEAR-FREE HOSPITAL DESIGN GUIDELINE Heather E. Lewis, AIA, NCARB. Draft / January 2015 PREPARED BY. architecture animals people

Kathleen Krafte, Lincoln Larson, Robert Powell Clemson University ISSRM: June 14, 2015

AnimalShelterStatistics

The Humane Society of the Southeast, Inc.

Community Cats and the Ecosystem

Daily Animal Health Monitoring Program

2017 ANIMAL SHELTER STATISTICS

Social and Reproductive Behaviours in the Cheetah (Acinonyx jubatus) in A Captive Population

Effects of Olfactory Enrichments on African Cheetahs (Acinonyx Jubtus)

AnimalShelterStatistics

AFRICAN CATS AT DISNEY S ANIMAL KINGDOM THEME PARK. Educator s Background Information

Care For Us Arc$c Wolf (Canis lupus arctos)

Karen Pryor. Getting Started: Clicker Book. with Your Rabbit. Joan Orr and Teresa Lewin

Animal Welfare. Tag Words: animal welfare; animals; welfare; animal health. Authors: Gralat Xue & Soo Jin Park with Julie M. Fagan, Ph.

Management of bold wolves

Animal Enrichment Best Practice Series

Ali s Journey. German Shepherd Rescue of The Rockies. May Newsletter Issue 2.

ZOO GUIDELINES FOR KEEPING LARGE FELIDS IN CAPTIVITY

Adoption Application Form

NEW VOLUNTEER GUIDELINES

To link to this article: PLEASE SCROLL DOWN FOR ARTICLE

It s a dog s life: vet nursing at Dogs Trust centre, Leeds

Distressed Animal Behaviors and Some Recommendations for Improvements at the Kuala Lumpur Zoo, Malaysia Amber Haque Published online: 04 Jun 2010.

Copyright is owned by the Author of the thesis. Permission is given for a copy to be downloaded by an individual for the purpose of research and

the Environment and Feeding Regime on the Behavior of Captive Felids

DRAFT. Code of Practice for the Care of Dogs in New Brunswick. The New Brunswick Society for the Prevention of Cruelty to Animals.

United States Department of Agriculture Animal and Plant Health Inspection Service. Inspection Report

Why should environmental enrichment be used to improve welfare on mink farms?

INTRODUCTION & MEASURING ANIMAL BEHAVIOR

Enrichments for captive Andean Condor (Vultur gryphus) in Zuleta, North Ecuador. Yann Potaufeu (2014)

Strengthening the Human Animal Connection

Shelter Intake Cats 16,000 14,000 12,000 10,000 8,000 6,000 4,000 2, All Other Zips. Total

Angela S. Kelling a b c, Meredith J. Bashaw a b d, Mollie A. Bloomsmith a e & Terry L. Maple a b f a Center for Conservation and Behavior, School of

Home Phone Business or Cell Phone Fax Number

Species Fact Sheets. Order: Gruiformes Family: Cariamidae Scientific Name: Cariama cristata Common Name: Red-legged seriema

Establishing a routine

An Evaluation of Respondent Conditioning Procedures to Decrease Barking in an Animal Shelter

How do we assess for welfare of individuals? Can we have a generic welfare assessment? Will it work for all shelters?

Behaviour of Hens in Cages

Dog Adoption Application Form

Play-Aggression in Kittens or Cats

Cat Math A math lesson on pet overpopulation

Adopting a rescue dog

Failing the Grade. Alberta Zoos Five Years On. September 2005

Zoo Animal Welfare Research. David Shepherdson Ph.D. Oregon Zoo

What do the experts think? Highlights from Expert Statements

TRAINING & BEHAVIOR QUESTIONNAIRE

Dog BehaviouristFor Barking Dog Complaints Trial. Brenton Thomass Community Safety Leader

PIGEON DISCRIMINATION OF PAINTINGS 1

Back to basics - Accommodating birds in the laboratory setting

2016 Community Report

The Effect of Zoo Visitors on the Behaviour and Welfare of Zoo Mammals. Alexandra Farrand

Applied Animal Behaviour Science

DOG FOR LIFE ADOPTION APPLICATION

Anthony Richard/Kendra Richard 6885 Mesa Ridge Pkwy. #169 Fountain, CO Phone: March 19, 2018

10/13/2016 CAT STATS STRESS & ITS EFFECTS ON SHELTER CATS YES, YOU CAN TRAIN A CAT!

SP6B NUISANCE BARKING IN DOGS AND THE WELFARE IMPLICATIONS OF DIFFERENT TREATMENT MODALITIES

HOW TO INTRODUCE A NEW DOG TO YOUR CURRENT RESIDENT DOG

Transcription:

Stereotypic pacing in Siberian tiger (Panthera tigris altaica) and the effect of interspecies presence: A case study Table of Contents List of Figures... 2 List of Tables... 3 Abstract... 4 1.0 Introduction... 5 2.0 Materials and Methods... 9 Cage Set up... 9 Data Collection... 9 3.0 Results... 11 4.0 Discussion... 14 5.0 Conclusion... 17 6.0 References... 18 Appendix... 20 1

List of Figures Figure 1 Study area coding for Siberian tiger (Panthera tigris altaica) case study...9 Figure 2 Observations with trainer present (n=30) and absent (n=666) in Siberian tiger (Panthera tigris altaica) enclosure with behaviour displayed by tiger.11 Figure 3 Observations with lion present (n=342) and absent (n=264) in Siberian tiger (Panthera tigris altaica) enclosure with behaviour displayed by tiger.12 Figure 4 Observations with trainer and lion present (n=27) and absent (n=346) in Siberian tiger (Panthera tigris altaica) enclosure with behaviour displayed by tiger 13 2

List of Tables Table 1 Behaviours recorded during observation of Siberian tiger (Panthera tigris altaica) during case study..10 3

Abstract Stereotypic behaviours are repetitive, apparently functionless behaviours displayed by captive species. These behaviours can be in response to suboptimal environments or as a method of coping with the inability to display highly motivated behaviours. In captive tigers this frustration most often is manifested by the display of stereotypic pacing. Methods of environmental enrichment have been used in an attempt to decrease these unwanted behaviours; however, social enrichment has not been studied in tigers and could be beneficial. This project is a case study of Hana, a 4 year old Siberian tiger (Panthera tigris altaica) at the Greater Vancouver Zoo in Aldergrove, British Columbia. The aim of this study was to determine if stereotypic pacing exhibited by Hana was effected by interspecies presence. One-zero sampling at one minute increments was collected over two days, with each day consisting of 6 hours of data collection in two hour increments (9:00-11:00; 11:30-13:30; 14:00-16:00). Hana s behaviour and location in her enclosure were recorded. The interspecies subjects in this study were the male lion in an adjacent enclosure and the trainer; their presence was also recorded for each observation. Of total observations 35.92% (n=250) were spent pacing. Comparatively, when the trainer and lion were both present (n=27) 7.4% were spent pacing, with only lion present (n=342) 34.8% were spent pacing and with trainer presence (n=30) 6.67% observations were spent pacing. The results of this study show that trainer presence had an effect on Hana s behaviour during the observed period. These results suggest that trainer-animal interaction could act as a form of social enrichment in tigers; however, more research is needed before this claim can be made. Keywords: Siberian tiger (Panthera tigris altaica), enrichment, social enrichment, zoo, animal welfare 4

1.0 Introduction The concept of stereotypic behaviour has become very important when exploring zoo animal welfare; with stereotypic behaviours defined as repetitive behaviours that serve no obvious purpose or function (Mason et al., 2007). Many captive species have developed stereotypic behaviours when in less than satisfactory environments and the development of these stereotypies can take many forms (Swaisgood and Shepherdson, 2005). They are often triggered by a behavioural response (Swaisgood and Shepherdson, 2005; Mason et al., 2007) or as a method of coping with stress or frustration due to the inability to perform a highly motivated behaviour (Iossa et al., 2009). Additionally, these behaviours can be the result of an animals past environment that has subsequently affected brain development, resulting in consistent abnormal behavioural responses (Mason et al., 2007). For many years, zoo enclosures have provided less than satisfactory environments; however, in recent years there has been growing public concern surrounding improving the welfare of captive species (Whitham and Wielebnowski, 2013). This includes decreasing the prevalence of stereotypic behaviours, which in large cat species displays primarily as stereotypic pacing. Though the prevalence of this behaviour varies between facilities and individuals, in extreme cases it has been documented that captive tiger species can pace for up to 60% of their daily time budget (Iossa et al., 2009). This is in large contrast to wild activity budgets in which tigers spend 18 hours a day sleeping with very few bouts of activity (Nevill and Friend, 2006). Factors that influence the development of this stereotypy are not fully understood (Krawczel et al., 2005), although it is known that these behaviours do not arise from a single state and in some cases may be a scar from a previous environment (Swaisgood and Shepherdson, 2005; Mason et al., 2007). 5

Not only is it important for zoological institutions to focus on decreasing the display of stereotypic behaviours for the welfare of the animals, it also has a benefit from a business and conservation standpoint (Miller, 2012). Miller (2012) reported that when visitors were shown videos of both pacing and resting tigers, they were more likely to view pacing tigers as receiving lower care when compared to those that were at rest. Even more interestingly, people were less likely to spend time standing at exhibits that housed pacing tigers and less likely to support the zoological institution in the future (Miller, 2012). There have been many attempts to change or decrease stereotypic behaviours including pharmaceutical drugs, positive reinforcement of alternative behaviour, physical prevention or punishment and environmental enrichment (Mason et al., 2007). Environmental enrichment is one of the most common methods (Szokalski et al., 2012), defined as one or more factors added to an environment to improve the physical or psychological welfare of an animal (Ellis, 2009). In tigers, the most common environmental enrichment devices used are those that stimulate natural hunting behaviours, varied feeding behaviours and olfactory stimulation (Szokalski et al., 2012). Studies have been done that use zebra dung and frozen blood balls (Van Metter, et al., 2008) as well as feeding enrichment using live fish (Bashaw et al., 2003). Success has been found with these devices although new objects must be introduced to avoid habituation (Szokalski et al., 2012) One area of enrichment that has been notably absent from tiger based scientific research is that of social enrichment (Swaisgood and Sherpherdson, 2005). Tigers have primarily been believed to be solitary animals, typically only housed with genetically related conspecifics or if there is mutual tolerance (Tilson et al., 2010). While in the wild this may be the case, when in captive environments tigers may benefit from interaction with conspecifics. One study conducted 6

by Miller et al. (2008) reported that tigers paced significantly more when they see conspecifics in adjacent enclosures, suggesting a frustration with the inability to interact with the conspecifics. While in many facilities group housing may not be a possibility, largely due to the limited numbers of these species, the benefit of interaction may extend to other interspecific interactions as well. Therefore, another option for social enrichment may be that of human- animal interactions (ie. trainers, general public). The level of training and interaction with trainer in different facilities varies greatly, but in large cat species training is often limited (personal communication, March 3, 2015). However, in a solitary housed tiger, any training or social stimulation may have an impact on boredom experienced by these tigers and the subsequent stereotypic pacing. The subject of this study is Hana, the solitarily housed Siberian tiger at the Greater Vancouver Zoo. Hana is a 4 year old female Siberian tiger who was housed in a pair with her sister Hani, until Hani died in April 2014 of a genetic heart defect. Hana and Hani were originally from the Grand Park Zoo in South Korea and are the result of their onsite tiger breeding program, part of the species survival plan. The Greater Vancouver Zoo is a 120 acre CAZA (Canada`s Accredited Zoos and Aquariums) accredited facility located in Aldergrove, British Columbia (BC) (Greater Vancouver Zoo, 2014) and is home to over 600 animals. Hana s enclosure is one of four in a block of enclosures that all house different species (African lions, Jaguar, Yellow baboons). The Greater Vancouver Zoo does not employ any hands on training practices with their large carnivores (personal communication, February 22, 2015) although other enrichment programs are unknown. The aim of this study was to determine if the stereotypic pacing exhibited by Hana changes with interspecies presence (trainer and lion). It is 7

hypothesized that stereotypic pacing during interspecies presence will decrease during the observation period. 8

2.0 Materials and Methods Cage Set up The study area is made up of the two enclosures (lion and tiger) of unknown size (Figure 1) and for observations has been divided into 6 quadrants (3 tiger enclosure; 3 lion enclosure). Within the tiger s enclosure there is a pond, 2 trees, a cement enclosure (Figure 1; 0 in scoring) and one red ball (hypothetically for enrichment). The enclosure is surrounded by chain link fence and there is approximately a ten foot path between the lion and the tiger enclosures. Data Collection Interspecies presence for this project is defined as presence or absence of a trainer and cage location of the male lion. The location of the male lion was Tiger Lion 0 0 1 2 1 2 recorded using the coding seen in Figure 1. While there are three lions in the enclosure, only the location of the male lion was recorded as the male is more active than the females in the enclosure Public Viewing Area Figure 2 Study area coding for Siberian tiger (Panthera tigris altaica) case study and appears to have influence on where the females are located, as seen through personal observations. These two interspecies subjects were chosen as they are the most commonly observed, as well as they both provide ease of visibility for the observer. Observer sat on the bleachers in the public viewing area (Figure 1), which allowed for visibility of lion and tiger enclosures as well as trainer entrance. One-zero sampling data at one minute increments was 9

collected over two days, with each day consisting of 6 hours of data collection in two hour increments (9:00-11:00; 11:30-13:30; 14:00-16:00). The choice of hours was based on the Greater Vancouver Zoo winter hours of operation (http://gvzoo.com/visitor-info/rates-andhours/index.php; February 5, 2015). This observation resulted in 360 data points each day, 720 data points total for the project. At each one minute interval the quadrant location of the lion and tiger was recorded as well as the presence (1) or absence (0) of the trainer. In addition, the behaviour displayed by the tiger at the time of observation was recorded. The three possible recorded behaviours for the tiger are pacing, resting, and other (Table 1). Other behaviours are not separated into more detail as they are not required to answer the study question. Behaviour Resting Pacing Other* Smell Walk Description Individual is lying down with no movement; eyes can be open or closed Walking back and forth along the same path repeatedly; appears without intention Investigating object, ground, or air by sniffing at it; requires uptake of air through nostrils Directional moving from one place to another * The few examples categorized under other do not cover all of the possible behaviours. This provides some possible examples of behaviours displayed during observation. Table 1 Behaviours recorded during observation of Siberian tiger (Panthera tigris altaica) during case study. 10

3.0 Results There were 696 total observation points, with the remaining data points (n=24) excluded if the tiger was out of sight of the observer. Of the total number of behavioural observations (n=696), Hana spent 35.92% (n=250) pacing, 42.24% (n=294) resting and 21.84% displaying other behaviours. Some of the other behaviours displayed included standing vigilant, eating, drinking, etc. Trainers were present for 30 (4.31%) observations. Of the observations with the trainer present, Hana spent 6.67% (n=2) of the observations pacing, 10% (n=3) resting and 83.3% (n=25) displaying other behaviours (Figure 2). 100% 90% 80% Percentage of Observa0ons 70% 60% 50% 40% 30% 20% 10% Pacing Res>ng Other 0% Not Present (n=666) Present (n=30) Figure 2 Observations with trainer present (n=30) and absent (n=666) in Siberian tiger (Panthera tigris altaica) enclosure with behaviour displayed by tiger. The lion was considered present when located in quadrant 1 (Figure 1) while Hana was in quadrant 2. This definition of present is only used for the lion, not the trainer. This was chosen as 11

from the opposite end of the lion and tiger enclosures, quadrant 2 and 1 respectively, presence could not be accurately assessed. The lion was present for 342 (49.14%) observations. Of the observations with the lion present, Hana spent 34.80% (n=119) pacing, 48.83% (n=167) resting and 16.37% (n=56) displaying other behaviours (Figure 3). 100% 90% Percentage of Observa0ons 80% 70% 60% 50% 40% 30% 20% 10% Pacing Res>ng Other 0% Lion Absent (n=264) Lion Present (n=342) Figure 3 Observations with lion present (n=342) and absent (n=264) in Siberian tiger (Panthera tigris altaica) enclosure with behaviour displayed by tiger. Lion and trainer were both present for 27 (3.88%) observations. When both trainer and lion were present 7.4% (n=2) observations were spent pacing, 7.4% (n=2) were spent resting and 85.2% (n=23) were spent displaying other behaviours (Figure 4). 12

Percentage of Observa0ons 100% 90% 80% 70% 60% 50% 40% 30% 20% 10% Pacing Res>ng Other 0% Trainer and Lion Absent (n=346) Trainer and Lion Present (n=27) Figure 4 Observations with trainer and lion present (n=27) and absent (n=346) in Siberian tiger (Panthera tigris altaica) enclosure with behaviour displayed by tiger. 13

4.0 Discussion Of the total observations, Hana spent 35.92% pacing. While this value seems high it is significantly lower than other research in which tigers in zoo and circus settings have been known to spend up to 60% of their day pacing (Iossa et al., 2009). Comparatively, Hana spent 42.24% resting; this value is much lower than resting behaviour found in wild tiger population, with 75% of their time spent resting or sleeping (Nevill and Friend, 2006). Resting was chosen as opposed to sleeping in this study as sleeping can be more difficult to assess in cat species due to their ability to display feigned sleeping or fake sleeping (Dybdall et al., 2007). Trainer Presence Trainer presence did appear to have an impact on pacing behaviour as it decreased to 6.67% when the trainer was present, compared to37.24% of observations spent pacing with the trainer absent. This shift in behaviour could be a result of anticipatory behaviours due to feeding times. Although this is speculation as Hana s feeding schedule is unknown. However, whether this was anticipatory behaviour or not there was a change in behaviour with trainer presence, direct contact was not required. This result suggests that trainer-animal interaction could have a large impact on stereotypic pacing and potentially act as a social enrichment for solitary animals. As mentioned, social enrichment is not a common area of study in captive species and one that should be explored further. The training program currently in place for Hana is unknown although it is a no contact program (personal communication, February 22, 2015). However, by increasing trainer presence or introducing new training programs, trainers may decrease the prevalence of pacing in Hana. The concern is that there is a limited number of data points with the trainer present. Therefore, while there does appear to be an effect, a larger sample size would strengthen the impact of these results. 14

Lion Presence Presence of the male lion does not appear to have an effect on Hana`s behaviour. There are many reasons that this could be the case although the most predominant reason would be that of habituation (Anderson et al., 2010). Hana did not appear interested in the presence of the lion in the same way she did with the presence of the trainer, and that was reflected in the results. This could suggest that while at some point the presence of the lion could have been a form of social enrichment, there is the concern of habituation which is a problem that all enrichment programs face (Anderson et al., 2010). Lion and Trainer Presence The results when both the lion and trainer were present did appear to have an effect; however this was driven primarily by the trainer presence observation points. While this may mean that trainer presence has a larger impact than that of lion presence, again this could be due to the small sample size. Limitations While some conclusions can be made from this project there were many limitations to the experimental design. The first of which was the limited amount of time for observation. Observation days took place during open hours of the Greater Vancouver Zoo; however, in ideal circumstances a longer observation period would have strengthened the project. Additionally, only one season was observed and seasonality could have had an effect on both Hana`s behaviour as well as the number of visitors to the facility. Similarly, the presence of the public was not taken into account but could have had significant impact on Hana`s behaviour. The cage behind Hana s, which houses Yellow Baboons, was not included in this study although Hana 15

appeared to be very interested in any activity from that enclosure. This could be providing additional social stimulation that was not otherwise accounted for. Another limitation of this study is the unknown enrichment programs currently employed by the Greater Vancouver Zoo staff, they were contacted for information but none was given. As mentioned, there is a red ball in Hana s enclosure theoretically for enrichment purposes. However, this item was in the same location on visits three months apart which suggests that the item is not providing any enriching qualities for Hana. 16

5.0 Conclusion Stereotypic pacing is a very prevalent welfare concern in captive cat species and determining new ways to decrease its prevalence is of great importance. While environmental enrichment has been used most notably to date, social enrichment is a method that has not been widely explored. While this study is specific to the one individual at the Greater Vancouver Zoo, it could suggest that trainer-animal interaction could play a part in trying to alleviate stereotypic pacing. Future research is needed that explores these interactions as well as how various training practices, both hands on and hands off can effect stereotypic pacing in tigers. Similarly, research is needed to determine the impact of the general public on these tigers as well as species in surrounding enclosures. Decreasing stereotypic pacing in zoo settings is of great importance to ensure the greatest quality of life for tigers and other species in captivity. Social enrichment and more specifically trainer-animal interactions may be the next step to achieve this. 17

6.0 References Anderson, C., Shanmugam Arun, A., Jensen, P., 2010. Habituation to environmental enrichment in captive sloth bears- effect on stereotypies. Zoo Biology. 29, 705-714. Bashaw, M.J., Bloomsmith, M.A., Marr, M.J., Maple, T.L., 2003. To hunt or not to hunt? A feeding enrichment experiment with captive large fields. Zoo Biology. 22, 189-198. Dybdall, K., Strasser, R., Katz, T., 2007. Behavioural differences between owner surrender and stray domestic cats entering an animal shelter. Applied Animal Behaviour Science. 104, 85-94. Ellis, S.L.H., 2009. Environmental enrichment: practical strategies for improving feline welfare. Journal of Feline Medicine and Surgery. 11, 901-912. Greater Vancouver Zoo. 2014. About the zoo. Available at: http://gvzoo.com/about/about-thezoo/index.php. Accessed November 27, 2014. Iossa, G., Soulsbury, C.D., Harris, S., 2009. Are wild animals suited to travelling circus life? Animal Welfare. 18, 129-140. Krawczel, P.D., Friend, T.H, Windom, A., 2005. Stereotypic behavior in circus tigers: Effects of performance. Applied Animal Behaviour Science. 95,189-198. Latham, N., Mason, G., 2010. Frustration and preservsation in stereotypic captive animals: is a taste of enrichment worse than none at all? Behavioural Brain Research. 211, Lyons, L., Young, R.J., Deaq, J.M., 1997. The physical characteristics of the environment and feeding regime on the behavior of captive felids. Zoo Biology. 16, 71-83. Mason, G., Clubb, R., Latham, N., Vickery, S., 2007. Why and how should we use environmental enrichment to tackle stereotypic behaviour? Applied Animal Behaviour Science. 102, 163-188. Miller, L.J., 2012. Visitor reaction to pacing behaviour: influence on the perception of animal care and interest in supporting zoological institutions. Zoo Biology. 31, 242-248. Miller, L.J., Bettinger, T., Mellen, J., 2008. The reduction of stereotypic pacing in tigers (Panthera tigris) by obstructing the view of neighbouring individuals. Animal Welfare. 17,255-258. Mohapatra, R.K., Panda, S., Acharya, U.R., 2014. Study on the activity pattern and incidence of stereotypic behavior in captive tigers. Journal of Veterinary Behavior. 9, 172-176. 18

Nevill, C. H., Friend, T.H., 2006. A preliminary study on the effects of limited access to an exercise pen on stereotypic pacing in circus tigers. Applied Animal Behaviour Science. 101, 355-361. Swaisgood, R.R., Shepherdson, D.J., 2005. Scientific approaches to enrichment and stereotypies in zoo animals: What s been done and where should we go next? Zoo Biology. 24,499-518. Szokalski, M.S., Litchfield, C.A., Foster, W.K., 2012. Enrichment for captive tigers (Panthera tigris): current knowledge and future directions. Applied Animal Behaviour Science. 139, 1-9. Tilson, R. Harris, T., Goodrowe Beck, K., Morris, C., Napier, J., Skurski, M., Armstrong, D., 2010. Tiger Care Manual. Silver Spring, MD: Association of Zoos and Aquariums. Van Metter, J.E., Harriger M.D., Bolen, R.H. Environmental enrichment utilizing stimulus objects for African Lions (Panthera leo leo) and Sumatran tigers (Panthera tigris sumatrae). Bios. 79, 7-16. Whitham, J.C., Wielebnowski, N. 2013. New directions for zoo animal welfare science. Applied Animal Behaviour Science. 147, 247-260. 19

Appendix Data Collection Sheet Time: Date: Observer: Behaviours Presence Tiger Location R P O Lion Trainer Notes 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 Tiger Location: 0, 1,2 (location on Figure 1) Behaviours: (No: 0, Yes:1) R: Resting P: Pacing O: Other behaviours Presence: Lion: position of lion in enclosure (0,1,2) (location on Figure 1) Trainer: absence (0) or presence (1) of trainer 20