Michelle Keel-Stocker (2014) Behavioural Enrichment in Captive Male Lions by ALERT

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of

conservation breeding male African lions (Panthera leo).

By Michelle Keel-Stocker {2013-2014}

A Project submitted in partial fulfilment of the requirements for the Bachelors Degree in Animal Conservation and Biodiversity for the University of Greenwich.

ACADEMIC SESSION: 2013-2014 PROJECT DECLARATION â&#x20AC;&#x153;This project report is the result of the independent work of Michelle Keel-Stocker. All other work reported in the text has been attributed to the original authors and is fully referenced in the text, and listed in the Reference sectionâ&#x20AC;?.

Student Name:- Michelle Keel-Stocker

Student Signature:-

Date:- 02/05/2014

Research Project dedicated to Antelope Park and the African Lion Environmental Research Trust (ALERT)

ACKNOWLEDGEMENTS This research was only possible due to the support and dedicated commitment from a number of people from all walks of life. First and foremost I would like to thank my project supervisor Rena Boothe for providing me with encouragement and inspiration throughout my project. Through all stages of the project Rena gave me sincere advise, guidance and support. This Project would not have been possible without the African lion Environmental Research trust (ALERT) and Antelope Park. I would like to give special thanks to the research manager Rae Kokes for her support while formulating my project and Dan Matthews for his attentiveness in my project and for arranging trips for me to collect the materials I needed for my project. I would also like to thank Alertâ&#x20AC;&#x2122;s researcher Megan Evershed for taking me and picking me up from the breeding programme grounds of which my research was carried out at. I would also like to thank Tinifo Jiri and Markson Ncube for the enthusiasm and help with the lion movements and allowing me access to lion enclosures to position my Behavioural Enrichments. I would like to pay my respects to the people at Antelope Park for welcoming me throughout the duration of my stay. I would also like to give my most sincere appreciation to my father William Keel-Stocker for his never ending support and interest in my project for the time during my data collection and for accompanying me on my trip to Zimbabwe to make my project a reality. Finally I would like to thank all my family and friends that have shown their support and belief in me throughout my project.

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ABSTRACT A behaviour study was carried out at Antelope Park in Zimbabwe on 16 captive male breeding Africa lions (Panthera leo) in order to evaluate the effectiveness of three low cost behavioural enrichments and their effects on territorial behaviour. The behaviours that were observed for the study included active locomotion, vocalisation, resting, behaviour enrichment (BE) feeding related, BE- touch related, social behaviour, territorial behaviour, grooming, aggression and stereotypical behaviour. The study was carried out over an 18 day period producing 108 hours of observations in total. The three different types of enrichments which were tested with the lions in the investigation on a rotational basis included a log covered in zebra (Equus quagga) dung, a Flat White Boer Ford pumpkin (Cucurbita maxima) and a log with Cow blood. The hypotheses that were formulated for the behavioural study included: “Lions will interact most with the food related enrichment (cow blood log) more than the other two behavioural enrichments”. “Zebra dung and cow blood behavioural enrichments will receive most interactions throughout the day”. “More territorial behavioural displays will be exhibited with the presence of the Zebra dung log and cow blood log enrichments” and “Zebra dung log and cow blood log will increase activity levels”. The results from the investigation concluded that the presence of the log with zebra dung enrichment significantly increased behaviours such as active locomotion, vocalisation and territorial behaviour. The statistical tests carried out showed an increase in these behaviours with a significant P-Value of <0.05, indicating a significant difference between data. Both behavioural enrichment interaction categories; BE- feeding related and BE- touch related was also increased with the presence of the log with zebra dung enrichment, leading to the conclusion that this enrichment type was the most successful treatment in this investigation.

LIST OF TABLES Table 1: African Lion classification.................................................................................................................3 Table 2: Enrichment availability and costs................................................................................................23 Table 3: Different measurements between Pumpkin and Watermelon.........................................32 Table 4: Lion groups used in the investigation composition and ages............................................33 Table 5: Details of Lion group 1....................................................................................................................33 Table 6: Details of Lion group 2....................................................................................................................34 Table 7: Details of Lion group 3....................................................................................................................34 Table 8: Measurements of the Lion Groupâ&#x20AC;&#x2122;s enclosures.......................................................................35 Table 9: Measurements of Acacia logs used for the BE:LD enrichment.........................................44 Table 10: Measurements of Acacia logs used for the BE:LB enrichment.......................................44 Table 11: Measurements of the Pumpkins used for BE:P enrichment.............................................44 Table 12: Ethogram and definitions used in the investigation...........................................................45 Table 13: Results from the chi square test for BE- feeding related interactions.........................63 Table 14: Results for the chi square test for BE- touch related interactions.................................63 Table 15: Results from the chi square test for resting behaviour displays....................................64 Table 16: Results for the chi square test for Active locomotion displays.......................................64 Table 17: Results from the chi square test for Vocalisation displays...............................................65 Table 18: Results from the chi square test for Social behavioural displays..................................65 Table 19: Results from the chi square test for Territorial behavioural displays.........................66 Table 20: Results from the chi square test for grooming behavioural displays..........................67 Table 21: Results from the chi square test for Aggressive behavioural displays.........................67 Table 22: Results from the chi square test for Stereotypical behavioural displays....................68

LIST OF FIGURES Figure 1: Current and Historical Range of the African Lion............................................................................5 Figure 2: Population estimates of the African Lion across the continent of Africa.......................5 Figure 3: Location of Zimbabwe on the African continent.................................................................28 Figure 4: Location in Gweru, Zimbabwe....................................................................................................29 Figure 5: Location of Antelope Park in Gweru, Zimbabwe.................................................................29 Figure 6: Stopwatch used for the investigation........................................................................................30 Figure 7: Size difference between Pumpkin and Watermelon...........................................................32 Figure 8: Pumpkin cut open to determine concentration value........................................................32 Figure 9: Breeding program grounds site plan at Antelope Park ....................................................34 Figure 10: Lion enclosures and Lion groups used in investigation..................................................35 Figure 11: Zebra dung used in the investigation....................................................................................38 Figure 12: Process of applying the Zebra dung to the log...................................................................39 Figure 13: BE:LD fully prepared and ready for Lion enclosure.........................................................39 Figure 14: One of the Pumpkins used in the investigation.................................................................40 Figure 15: Cow blood samples that were used for the BE:LB in the investigation......................42 Figure 16: A BE:LB that was used in the investigation..........................................................................42 Figure 17: A BE:LB that was used in the investigation..........................................................................43 Figure 18: BE:LB showing how the end surface area was covered with the Cow blood..........43 Figure 19: Graph 1: Results from Pilot study Watermelon VS Pumpkin........................................47 Figure 20: Graph 2: All behaviours observed in all behavioural enrichment types..................48 Figure 21: Graph 3: Results from the controlled observations..........................................................49 Figure 22: Graph 4: Behaviour frequencies observed with the presence of the BE:LD............50 Figure 23: Graph 5: Behaviour frequencies observed with the presence of the BE:P...............51 Figure 24: Graph 6: Behaviour frequencies observed with the presence of the BE:LB.............52 vi

Figure 25: Graph 7: Observations of all behaviours with the Zebra dung log enrichment throughout all time periods............................................................................................................................53 Figure 26: Graph 8: Observations of all behaviours with the Pumpkin enrichment throughout all time periods............................................................................................................................54 Figure 27: Graph 9: Observations of all behaviours with the Cow blood log enrichment throughout all time periods............................................................................................................................55 Figure 28: Graph 10: Frequencies recorded for the BE- feeding related interactions for all the Behavioural Enrichment types...............................................................................................................56 Figure 29: Graph 11: Frequencies recorded for the BE- touch related interactions for all the Behavioural Enrichment types.......................................................................................................................57 Figure 30: Graph 12: Frequencies recorded for Active Locomotion displays for all Behavioural Enrichment types.......................................................................................................................58 Figure 31: Graph 13: Frequencies recorded for vocalisation behaviour for all Behavioural Enrichment types.................................................................................................................................................59 Figure 32: Graph 14: Frequencies recorded for Territorial behaviour displays for all Behavioural Enrichment types.......................................................................................................................60 Figure 33: Graph 15: Frequencies recorded for Aggressive behaviour for all Behavioural Enrichment types.................................................................................................................................................61 Figure 34: Graph 16: Frequencies recorded for Stereotypical behaviour for all Behavioural Enrichment types.................................................................................................................................................62 Figure 35: Lion displaying touch related interaction with the BE:LD.............................................69 Figure 36: Lion displaying feeding related interaction with BE:LD.................................................69 Figure 37: Lion displaying touch related interaction with BE:P........................................................70 Figure 38: Lion displaying feeding related interaction with the BE:P.............................................70 Figure 39: Lion displaying touch related interaction with BE:P playful nature..........................71 Figure 40: BE:P after being given to a group of lions for the duration of one day.....................71 Figure 41: Lion displaying touch related interaction with the BE:LB..............................................72 Figure 42: lion displaying feeding related interaction with the BE:LB...........................................72 vii

LIST OF ABBEVIATIONS AP: Antelope Park ALERT: African Lion Environmental Research Trust BE: Behavioural Enrichment BE:LD: Behavioural Enrichment Log with Dung BE:P: Behavioural Enrichment Pumpkin BE:LB: Behavioural Enrichment Log with Blood

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CONTENTS ACKNOWLEDGEMENTS...................................................................................................................................III ABSTRACT...............................................................................................................................................................IV LIST OF TABLES......................................................................................................................................................V LIST OF FIGURES..........................................................................................................................................VI-VII LIST OF ABBREVIATIONS...............................................................................................................................VIII 1. INTRODUCTION..........................................................................................................................................1-2 2. LITERATURE REVIEW......................................................................................................................................3 2.1 Classification....................................................................................................................................................3 2.1.2 African lions.................................................................................................................................................3 2.1.3 Distribution.............................................................................................................................................4-5 2.1.4 Threats to lions.......................................................................................................................................6-8 2.1.5 Reproduction & relationships to young........................................................................................8-9 2.2 Adaptations............................................................................................................................................10-11 2.3 Wild Diet.................................................................................................................................................12-13 2.3.1 Captive Diet........................................................................................................................................13-14 2.4 Communication....................................................................................................................................14-16 2.5 Territorial behaviour..........................................................................................................................16-17 2.6 Natural behaviour...............................................................................................................................17-18 2.6.1 Resting..................................................................................................................................................17-18 2.6.2 Movement activity...................................................................................................................................18 2.6.3 Grooming....................................................................................................................................................18 2.6.4 Agonistic behaviour................................................................................................................................18 2.7 Social Structure....................................................................................................................................18-19 2.8 Lions in Captivity.................................................................................................................................19-21 2.9 Environmental Enrichment.....................................................................................................................21 ix

2.9.1 Olfactory Enrichment............................................................................................................................22 2.9.2 Food Enrichment..............................................................................................................................22-23 2.9.3 Low cost Enrichment..............................................................................................................................23 3. Antelope Park...........................................................................................................................................23-24 3.1 African Lion Environmental Research Trust (ALERT)....................................................................24 3.2 Previous Research................................................................................................................................24-25 4. Aims and objectives of the investigation...............................................................................................26 4.1 Hypotheses.....................................................................................................................................................26 4.2 Null Hypotheses...........................................................................................................................................26 4.3 Alternative Hypotheses..............................................................................................................................27 5. METHODOLOGY...........................................................................................................................................28 5.1 Site descriptions....................................................................................................................................28-29 5.2 Behavioural enrichment...........................................................................................................................30 5.3 Materials for the investigation...............................................................................................................30 5.3.1 Controlled Study......................................................................................................................................31 5.3.2 Pilot study............................................................................................................................................31-32 5.4 Lions and enclosures..........................................................................................................................33-34 5.4.1 Site area measurements.........................................................................................................................35 5.5 Design of study.............................................................................................................................................35 5.5.1 Procedure....................................................................................................................................................36 5.5.2 Data analysis..............................................................................................................................................37 5.6 Preparing enrichment Log with Zebra Dung...........................................................................38-39 5.6.1 Preparing enrichment Pumpkin........................................................................................................40 5.6.2 Preparing enrichment Log with Cow blood..........................................................................41-43 5.6.3 Measurements of logs used in the investigation..........................................................................44 5.6.4 Measurements of Pumpkins used in the investigation..............................................................44

6. Ethogram...........................................................................................................................................................45 7. RESULTS.............................................................................................................................................................46 7.1 Graphs & Graph Analysis.................................................................................................................46-62 7.2 Statistical Analysis of Results...........................................................................................................63-68 7.3 Photographs of African lions interacting with the BE treatments.....................................69-72 8. DISCUSSION....................................................................................................................................................73 8.1 Behavioural Enrichment effectiveness................................................................................................73 8.2 Log with Zebra dung..........................................................................................................................73-75 8.2.1 Pumpkin..............................................................................................................................................75-76 8.2.2 Log with Cow blood........................................................................................................................76-77 8.3 Findings from the investigation.............................................................................................................78 8.3.1 Territorial behaviour Outcomes.................................................................................................78-79 8.3.2 Activity outcomes.............................................................................................................................79-80 8.3.3 Stereotypical outcomes..........................................................................................................................80 8.3.4 Aggressive behaviour outcomes.........................................................................................................81 8.3.5 Natural Enrichment vs. Novel Enrichment............................................................................81-82 8.3.6 Behavioural enrichment interactions..............................................................................................82 9. CONCLUSION.................................................................................................................................................83 10. FUTURE WORK............................................................................................................................................84 10.1 Limitations...................................................................................................................................................84 10.2 Further investigations.............................................................................................................................84 11. REFERENCES...........................................................................................................................................85-90 12. APPENDICES..................................................................................................................................................91 12.1 Appendix Raw Data.........................................................................................................................92-97 12.2 Data collection sheet...............................................................................................................................98 12.3 Raw Statistical Data.......................................................................................................................99-102

12.4 Risk Assessment............................................................................................................................103-104

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Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

1. INTRODUCTION African Lions in the wild typically spend 20-21 hours resting (Schaller, 1972) and tend to become more active during late afternoon (Estes, 1991). Their activity peaks are after 17.00 and before 08.00 (Schaller, 1972) often displaying a flurry of activity around dusk 18.00. Activity often exhibited during this time includes stretching, defecating, grooming, scent marking and social interactions (Schaller, 1972). More often than not they then recline again for an hour or so before a similar activity pattern will be repeated again (Schaller, 1972). Commonly outbreaks of walking can last for an hour or more, with frequent intervals of resting or remaining stationary (Schaller, 1972). African Lions in captivity are often faced with ecological problems; as their enclosure and environment my limit or even restrict their natural behaviours; often leading to stereotypical behaviours (Szokalski et al., 2012). Often natural behavioural instincts in Panthera leo can be reduced within a captive environment such as territorial behaviour displays (Boler, 2009), often as a result of habituation (Morgan et al., 2007). The loss of natural behaviours and like the renowned stereotypic pacing in lions like many other big cat species; proposes more undesired behaviours (Morgan et al., 2007). Yet relatively little knowledge is known about optimal enrichment for the species (Szokalski et al., 2012), therefore there is a need for behavioural enrichment for indivuals in captivity; to encourage and promote natural behaviours and reduce stereotypical behaviour (Ncube et al., 2010). Behavioural enrichment has been used in a number of studies with big cats in captivity however little is documented for Panthera Leo and often mostly for Panthera leo cubs; not adults. A Focused study whereby behavioural enrichment has been used to increase activity and reduce stereotypical behaviour was a study carried out by (Quirke et al., 2011) whereby the effects of different enrichment types were tested to see if they effected the behaviour of cheetahs (Acinonyx jubatus). A typical Panthera Leo diet in the wild is comprised of a diverse intake, with their common choice of prey including hares, baboons, various antelopes, zebra and buffalo (Schaller, 1972) and of course other species of fauna such as giraffe and crocodile (Schaller, 1972) however usually less regularly and often dependent upon opportunity and scavenging chance (Talbot, 1

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

1963). In captivity their diet is less varied and usually consists of horse and cow meat (Black et al., 2010). In contrast to that of the wild diet there is an enormous difference, not to mention that in the wild they would consume the majority of the whole carcass (Rodrigo, 1999) but often are not even given this opportunity in captivity (Hine, 2014), this in itself is considered a lack of natural enrichment (Morgan et al., 2007). The investigation was carried out on 16 adult male Lions, observations were collected for six hours per day for 18 days over the period of one month (July). For the study three different enrichments were tested. The behavioural enrichments for the study comprised of Acacia log smeared with zebra dung, a whole pumpkin and an Acacia log smeared with cow blood. A significant feature of the investigation was to provide enrichment items that were naturally available in the wild and to compare it to that of a novel object (pumpkin) that would not be. The investigation measured the following behaviours; active locomotion, vocalisation, resting, enrichment feeding related, enrichment touch related, social behaviour, territorial behaviour, grooming, aggression and stereotypical behaviour.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

2. LITERATURE REVIEW 2.1 Classification Lions are classified in the following structure: Table1: shows the African Lion’s Classification (Alert, 2013)

Kingdom

Animalia

Phylum

Chordata

Subphylum

Vertebrata

Class

Mammalia

Order

Carnivora

Family

Felidae

Subfamily

Patherinae

Genus

Panthera

Species

Panthera leo

2.1.2 African lions African Lions (Panthera Leo) are one of the most social of all the Panthera species; being the only big cat that lives in a social organisation known as a pride (Schaller, 1975). A pride is a fairly cohesive social unit that bares a hierarchical structure (Schaller, 1972) which usually comprises of; a fully grown adult male, six to 10 lionesses, a coalition of juvenile males, and multiple cubs (Rudnai, 1973). African lions are native to central and Southern Africa (Server, 1993), this is where the annual average temperature in central Africa (Central African Republic, Birao) is 26.5◦ (Climate, 2013) and the annual average precipitation is 842mm of rainfall per year (Climate, 2013). In Southern Africa (Johannesburg) the annual average temperature is 22.4◦ (Climate, 2013) and the annual average precipitation is 543mm of rainfall per year (Climate, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

2.1.3 Distribution African Lions were formally once widespread across the African continent, with the exception of the rainforests and Sahara Deserts (Riggio, 2012). Although their numbers in earlier years are not clearly defined it is apparent from research that both the African lionâ&#x20AC;&#x2122;s distribution and abundance have severely declined (Mason, 2013). Today the African lion now lives in less than a quarter of its historic range (Alert, 2013) and research estimates a dramatic loss of almost half of the lions in only the past two decades (Alert, 2013). Recent research from 2012 confirms numbers could be as low as 20,000 individuals (Panthera, 2013) and currently in Southern Africa only 12,036 individuals (Riggio, 2012), the overall assumed continent-wide lion population is 25,000 indivuals (Panthera, 2013). Due to these substantial declines, conservation organisations proposed to list the African Lion (Panthera Leo) as endangered under the U.S Endangered Species Act (Mason, 2013) and also request to upgrade the species on CITES protection from Appendix II to Appendix I (Mason, 2013). Today, the African lion have disappeared from 80% of its historical range (Riggio, 2012). Now only existing in 28 countries in Africa (Riggio, 2012), and are confirmed extinct in 26 countries (Alert, 2013). At present day lions now only cover 10% of the continent, primarily in southern and eastern Africa (Alert, 2013). The population of African lions estimated in the country Zimbabwe is now approximately 850 individuals (Alert, 2013). It is thought that a major contribution to reduction in numbers within Zimbabwe is due to sport hunting (Davidson et al., 2011). A study carried out by Loveridge between the years of 1999 - 2004 examined the ecological study of African lions (Panthera leo) in Hwange National Park, western Zimbabwe. The study measured the impact of sport-hunting beyond the park on the lion population within the park, using radiotelemetry and direct observation (Davidson et al., 2011). Results showed 34 of 62 tagged lions died during the study; of which 24 were shot by sport hunters- 13 adult males, 6 sub-adult males and 5 adult females (Davidson et al., 2011).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Figure 1: shows the Current and Historical Range of the African Lion (Panthera, 2013).

Figure 2: shows the current population estimates of the African Lion (Panthera leo) across the continent of Africa (Alert, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

2.1.4 Threats to lions The issues that threaten the African Lion are of similar threats that many other species encounter (Mcdonald et al., 2013), however being a top predator species; lions are a main source of human conflict (Bauer et al., 2003). The issues arose as human populations continue to grow throughout Africa (Mcdonald et al., 2013); contributing to the transformation of natural landscape into urban expansions and agricultural land use developments (Bauer et al., 2003). This in effect has indirect and direct consequences impacting upon lion populations (Hemson et al., 2009). As human inhabitancies continue to take over areas that were once wilderness, areas are becoming more urbanised and segregated, which often makes lionâ&#x20AC;&#x2122;s natural habitat and often a whole prides territory range severely fragmented (Bauer et al., 2003). This subsequently leads to lion human conflict (Hemson et al., 2009), as lions still continue to return and revisit the areas that were once a part of their former range (Patterson et al., 2004), only to discover changes from wilderness to development areas (Patterson et al., 2004). This in turn raises a negative perception amongst the local people, as lions are regarded as trespassing into human inhabitancies and are considered a fearful dangerous animal (Bauer et al., 2003). In context lions are carnivores so therefore raise an instant awareness of fear (Mcdonald et al., 2013), but are regarded as harmless in most areas if not provoked or challenged (Alert, 2013). However this form of unwanted presence often instantly withstands low tolerance levels from the locals (Kolowski et al., 2006); resulting in attacks and deaths of what is to be considered troublesome lions (Bauer et al., 2003). Human Development also leads to fragmented areas, thus reducing the areas for lions to naturally disperse and range in (Hemson et al., 2009), this causes extreme concealed damage to not only lion populations but the entire species (Alert, 2013); as inbreeding trends can rapidly increase when prides are confined to small, disconnected areas of land (Bauer et al., 2003). With less opportunities of genetic diversity, lion populations can become far more susceptible to future birth defects and diseases (Bauer et al., 2003). Agricultural developments usually inspire livestock keeping, which has a profound effect to lion human conflict (Dar et al., 2009). Even though lions naturally favour hunts with indivuals 6

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

from within their pride usually in the structure of several females or in small groups (Schaller, 1972) and usually hunt sick or injured prey; acting advantageous to them reducing time expenditure and energy spent physically hunting (Rudnai, 1973). Lions will still cease a prime opportunity such as an abundance of livestock in an area (Kolowski et al., 2006). They will undoubtedly acknowledge this prospect, and take an easy break through by venturing closer for and opportunistic kill (Mason, 2013). It is estimated that lions are responsible for the killing and damage to approximately 700 cattle and over 1000 small livestock annually (Bauer et al., 2003); with finance figures valued at approximately US $140 000 per year (Bauer et al., 2003). Commercial ranches in south-eastern Kenya found that lions were responsible for 86% of attacks on livestock annually (Kolowski et al., 2006). It is also determined that not necessarily all these attacks are killing for consumption, but as a result of an easy target (Patterson et al., 2004). The fact that lions do not always consume prey does not justify natural killing instincts (Hemson et al., 2009), however when livestock farming areas are erected in former lion home ranges it should surely be determined and expected that indents such as these are inevitable (Maclennan et al., 2009). Incidences like these influence the locals to take immediate physical action (Mason, 2013) in the plan of attacking the lions. A study in Maasai steppe, Tanzania revealed that 85 lions were killed in one month as a result of retaliatory killing of lions (Kissui, 2008). This is a common devastating incident carried out by the locals, however is done as a result of protecting their livestock (Mason, 2013). Another main threat to lions is Poaching and Trophy hunting (Alert, 2013), contributing to an averaged 665 Lion trophy exports per year (Alert, 2013,) which is non-sustainable due to the given fact that the whole African lion population estimate is only 25,000 (Loverideg et al., 2007). In areas that trophy hunting is permitted by government authorities, it is often due to the prospect to deliver capital; though research has exposed severe decline (Croes et al., 2011). Regarding the fact that males are the desirable trophy as an indication of size, power and strength, from the overview of whole population figures statistics indicate that there are about 3,000 adult trophy males in Africa (Alert, 2013), an estimate of 40% live in strictly protected areas (Alert, 2013), leaving an estimate of 1,800 male lions essentially â&#x20AC;&#x153;huntableâ&#x20AC;? (Alert, 2013). Consequently with the given figure of 665 individualâ&#x20AC;&#x2122;s trophy hunted per year 7

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

and that some areas are permitted this will continue to an unruly decline (Davidson et al., 2011). Prey depletion also contributes to be another threat to lion populations (Hemson et al., 2009) as in areas whereby development is established, the lions prey species also create an attraction by the local people for human consumption (Patterson et al., 2004) and this depletes the abundance of natural prey available for lion consumption within these areas (Patterson et al., 2004); contributing to competition for food resources (Schuette et al., 2013). The utilization of game species throughout Africa has always been an ongoing impact (Loveridge et al., 2007), however it is thought to be an ever growing trend due to promoted safari game experiences (Davidson et al., 2011).

2.1.5 Reproduction and relationships to young Young female lions usually become sexually mature at around three years old and males usually just a few months longer (Schaller,1972). Females are polyestrous which contributes to a high reproductive potential (Packer et al., 1983). An average length for gestation periods was estimated at 109.7 days (Cooper, 1942) with variation of 100 to 114 days (Cooper, 1942). The osetrus cycle can last for a week (Schaller, 1972) and recurs at intervals of three weeks (Schaller, 1972), however in captive lions past records show that both the oestrus period and the time between these periods have more variation (Schaller, 1972). Mating takes place between a mature male and lioness usually after they remain close by to each other for several days (Schaller, 1972). Once the lioness is relaxed with the maleâ&#x20AC;&#x2122;s presence she will allow him to mount her for copulation (Cooper, 1942), which usually only lasts between 6-10 seconds (Asdell, 1964). This cycle can repeat as often as every 20 minutes (Schaller, 1972) and a pair of lions can copulate as often as 40 times in a day (Asdell, 1964). The number of cubs in a litter size can vary from 1- 6 individuals (Schaller, 1972), litter size is thought to be influenced by food abundance within the area (Schaller, 1972), as past studies carried out in the Serengeti on Masai lion prides showed that prides living within an area with large populations of resident prey species had larger litter sizes (Stevenson-Hamilton, 1954). 8

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

There is no birth season for lions, however reports from Kruger National Park developed a trend in that most births were observed during the months of March to July (StevensonHamilton, 1954). The relationship between a lioness and her cubs is a finely balanced relationship between care and abandonment (Schaller, 1972), as a lioness has her own desires and needs as well as ensuring the survival of her offspring whenever conditions threaten them (Schaller, 1972). Cubs are born helpless and fully dependent upon their mother, their eyes are closed and they weigh about 2kg (Carvalho, 1968), their eyes usually open around 10 days old and first incisors appear at around 20 days old (Carvalho, 1968). Although cubs are able to crawl they cannot walk steadily until around three weeks of age (Schaller, 1972), therefore lionesses regularly carry their cubs, picking them up with their teeth by the neck and moving them to a safe hide (Adamson, 1963); this can happen as often as a lioness feels necessary to do so for her cubs protection (Schaller, 1975). Lionesses with new born often become separated from the rest of the pride not for any reason other than the rest of the pride continue their usual routine while she has to attend to her cubs (Adamson, 1963). As a result of this occurrence Lionesses can often leave the cubs to attend back to the pride (Schaller, 1972), during this time they usually keep their cubs hidden in dense pockets of vegetation to imitate that of a hide (Schaller, 1972). It is important for lionesses to remain social with the rest of the pride members as sometimes several lions within one pride may have young offspring to care for at the same time, which can at times lead to companionship and multiple mothering between females to each otherâ&#x20AC;&#x2122;s cubs (Adamson, 1963); for instance communal suckling duties, which long term is an advantage to group living and ultimately cub survival (Cooper, 1942). When cubs are around 6-7 weeks old they are no longer carried by the lioness (Schaller, 1972), but they still remain close by to the females within the pride (Schaller, 1972). Cubs often seek out a male and attempt to interact with him with greeting and play behaviour (Cooper, 1942), however males tend to show an unresponsiveness in return (Schaller, 1972). Though males do play a role in the nurture and protection of the cubs as males watch over cubs during the time females in the pride feed (Cooper, 1942). 9

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

2.2 Adaptations Lions are physiologically adapted to their environment and their way of living with mechanisms and strategies in order to survive (Wroe et al., 2007). Lions have adapted retractable claws with four claws on their back feet and five on the front (Alert, 2013), the extra digit on the front is referred to as the dew claw (Hopwood, 2008); this acts like a thumb which is used to hold down prey (Hopwood, 2008). The retractable claws main function is to grab and hold prey (Alert, 2013), the structure of the claws are sharp, strong and curled (Alert, 2013); serving the purpose needed of impaction into their preyâ&#x20AC;&#x2122;s skin immediately upon the impact of capture (Wroe et al., 2007), enabling a firm grip of the muscle tissue (Wroe et al., 2007). This function gives lions enough time to position their mouths around the preyâ&#x20AC;&#x2122;s neck for a successful and efficient kill (Wroe et al., 2007). The purpose of the claws being retractable is when claws are not in use they are able to pull the claws into the paw in order to prevent them from damage (Alert, 2013), the retractable nature also aids the claw strength and protection for the time the claws need to be strongest during hunting (Wroe et al., 2007). Lions have a barbed covered tongue (Wroe et al., 2007), a feeding strategy adaptation, as the tongueâ&#x20AC;&#x2122;s surface is covered in sharp backward pointing spikes called Papillae (Wroe et al., 2007), the structure of their tongue makes the surface abrasive (Alert, 2013) which plays a vital role in stripping fur from prey and meat from bones (Alert, 2013). The barbed tongue also plays an essential role as a grooming tool (Wroe et al., 2007), as they use their tongue and saliva to moisten their fur. This not only keeps them clean, but also helps to remove parasites such as ticks and mosquitoes, to remove lose fur; acting as a detangling mechanism (Wroe et al., 2007) as well as helping to regulate body temperature and homeostasis (Wroe et al., 2007). Panthera leo are distinguished from the other Panthera lineage by the males mane (Hopwood, 2008), although all Panthera genus are thought to all share the common ancestor (Hopwood, 2008), it is thought that Panthera leo evolved to have manes after they diverged from the other members of the Panthera lineage (Hopwood, 2008) and that selection pressure and genetic response led to mane development within the species (Hopwood, 2008). Group living within a pride proposes several pressures; sexual selection competition and competition between rival males for territorial range (Schaller, 1972). The presence of the 10

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

mane induces several functions to the male lion, firstly it acts as a shield when male to male competition takes place (Wroe et al., 2007) as the neck is a vulnerable area, thus having a dense mane covering this area provides some protection against rival males that compete for pride and territory takeover (Packer et al., 1983). The mane also serves as a signal of fighting ability and physiological condition (Alert, 2013) as it is thought that mane size and dark colouration indicates the level of dominance, age and strength in a male lion (Hopwood, 2008). A study on Serengeti male lions proved this theory, as the study showed that over time both the length and pigmentation of mane changed (Schaller, 1972); manes grew longer and darkened due to an increase of testosterone levels during adolescence development (Schaller, 1972) and (Rudnai, 1973). Furthermore a signal of dark mane and long length indicates to females a male with good fitness (Alert, 2013), which is a benefit to her as when she reproduces with him, not only will his genetics be passed on to a future generation but he will also be a strong candidate to protect their offspring and pride (Schaller, 1972). Males are also an important asset to providing the pride with food (Funston et al., 1998) as although females are the main cooperative hunters, male strength is needed rather than speed to take down large prey like Cape buffalo (Syncerus caffer) (Funston et al., 1998) which the pride also require (Rudnai, 1973). A lionâ&#x20AC;&#x2122;s sense of smell is well developed with a specialized olfactory organ located in the roof of their mouth called the Jacobsonâ&#x20AC;&#x2122;s organ (Wroe et al., 2007); this is used to detect smells within their environment (Alert, 2013). Lions demonstrate the use of this organ with what is known as a Flehman response (Jackman et al., 1982). Lions smell a scent by passing it over the organ (Wroe et al., 2007), this is exhibited in a motion that shows a lions mouth firmly open (Schaller, 1972). The function of this organ gives lions an exceptional sense of smell (Schaller, 1972) and provides males with the ability to receive a femaleâ&#x20AC;&#x2122;s reproductive status within a few metres of her presence (Rudnai, 1973).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

2.3 Wild Diet A wild lion’s diet consists of a diverse range of prey species (Rudnai, 1973) and the prey they kill ranges from large to small mammals (Schaller, 1972). Preferred prey includes; Wildebeest (Connochaetes), Zebra (Equus quagga), Thompsons gazelle (Eudorcas thomsonii), Eland (Taurotragus oryx), Topi (Damaliscus korrigum), Hartebeest (Alcelaphus buselaphus), Grants gazelle (Nanger granti), Impala (Aepyceros melampus), Cape buffalo (Syncerus caffer), Warthog (Phacochoerus africanus) and Giraffe (Giraffa camelopardalis) (Schaller, 1972). A lions diet is influenced by four main factors in the wild; size of prey, availability, density and scavenging. Species such as elephant (Loxodonta africana) and Hippopotamus (Hippopotamus amphibious) are too large a species to prey on and even young are usually vigorously defended by the adults (Grzimek et al., 1960); therefore lions usually do not make an attempt to prey on them (Grzimek et al., 1960). It was reported however that on rare occasions in Virunga National Park, Ituri, Congo; lions were known to kill young (Hippopotamus amphibious) (Bourlière, 1963). Apart from their size, this species have a habit of staying in water for the majority of their day (Jackman et al., 1982), which also makes them unavailable to lions (Schaller, 1972). Lions usually prey upon herbivores similar to the same weight and size as themselves (Rudnai, 1973), however their ability to use cooperate hunting enables them to succeed in taking down prey larger than themselves such as (Syncerus caffer) (Bourlière, 1963). Prey availability has a profound influence upon the lions diet, for instance prey species such as (Connochaetes), which are a migratory species; following seasonal movements within a region (Talbot, 1963), was found that (Connochaetes) over the course of three years were available to lions in the Serengeti region for an average of only 35% of the time (Schaller, 1972), (Equus quagga) 63% (Schaller, 1972) and (Eudorcas thomsonii) 68% (Schaller, 1972). Lion populations do not follow migratory herds (Rudnai, 1973) and so therefore their diet can vary dramatically as a result of seasons (Rudnai, 1973), although Lions readily have (Syncerus caffer), (Damaliscus korrigum) and (Aepyceros melampus) available most of the year (Schaller, 1972). Though (Equus quagga) in Seronera, Tanzania were recorded to have been killed every month of the year with peaks in the months of September and November in 1966 (Schaller, 1972). 12

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Density also effects their diet as densities of prey constantly change as a result of movement patterns (Rudnai, 1973), consequently giving a continual adjustment to lions diet (Schaller, 1972). Prides shift to greater roaming areas across the ecosystem when a prey species density in an area decreases through emigration (Schaller, 1972); vulnerability decreases also (Schaller, 1972), which encourages lions to work harder to meet their dietary requirements (Rudnai, 1973). Lions will partake in scavenging behaviour (Rudnai, 1973), usually when prey have died from disease or injury within the local area (Schaller, 1972) or as a result of prey killed by spotted hyenas (Crocuta crocuta) within their range (Rudnai 1973). Field studies have estimated that mature wild lions consume on average between 10-25 pounds of prey a day (Hine, 2014), this is between 4-7% of their body weight (Hine, 2014). 2.3.1 Captive Diet A lions captive diet is profoundly dissimilar to that of their wild diet (Hine, 2014); a captive diet is structured by taking into consideration the species specific needs such as nutritional and ecological requirements (Black et al., 2010), this is to ensure that the species natural feeding patterns and behaviours are stimulated (AZA, 2008). In captivity lions are usually fed a selection of either cow or horse meat and occasionally chicken (Altman et al., 2005). Captive lions eat considerably less due to their sedentary life style (Altman et al., 2005) as they burn fewer calories (Hine, 2014), they eat between 4-6% of their body weight (Hine, 2014), which is between 9-18 pounds of meat; usually fed five days a week (Hine, 2014). Lions require a high protein and fat diet which must be replicated in captivity (Black et al., 2010), It has been determined that vitamin A, Arachadonic acid, Taurine, Vitamin D and Calcium are essential components to their diet (Hine, 2014). Vitamin A is required for the lining of the digestive and respiratory system (Kohn et al., 2011), the provision of Vitamin A prevents respiratory and digestive tract infections (Kohn et al., 2011), as well as other common diseases such as blindness and conjunctivitis (Hine, 2014), it also benefits sperm quality and fertility (Barros et al., 2007), which in some cases is essential to captive breeding programmes (Hine, 2014). Arachadonic acid is needed in the diet as it is considered a fatty acid (Hind, 2014), found in membranes and cellular tissue (Hind, 2014) being an essential part of their dietary 13

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

requirements (Kohn et al., 2011). It is crucial that lions receive a sufficient amount of Taurine in their diet, as a lack may result in vision problems, heart failure, immune system dysfunction, blood clotting disorders and initially a reduced life span (Black et al., 2010). Vitamin D and Calcium are vital for all stages of life for growth development, these needs can be met with a carcass feed or meat containing bones (Bashaw et al., 2007) as they ingest calcium and phosphorus in a ratio of approximate two parts calcium to one part phosphorus (Hind, 2014), this measurement is ideal for growing indivuals (hind, 2014). Vitamin D and calcium deficiencies can lead to numerous problems such as; metabolic bone disease, reproductive failure, bowed leg development and arthritis (Kohn et al., 2011). These problems can be prevented by providing a well-balanced diet in captivity (Hind, 2014) ensuring bones and muscle tissue are fed regularly and a combination of vitamin A and D is given (Black et al., 2010), this can be administered via a supplemental powder which can be embedded into the meat thats fed (Black et al., 2010). It is not possible to fed captive lions live prey as defined by the Secretary of stateâ&#x20AC;&#x2122;s standards for modern zoo practice (Altman et al., 2005), backed by the Animal welfare Act 2006 also (Defra, 2012). This itself restricts lions activity levels in a captive environment (Bashaw et al., 2007), as they do not follow the same hunting and feeding regime to that of a wild lion (Bashaw et al., 2007), however feeding tools such as pole apparatusâ&#x20AC;&#x2122;s can encourage an energy level increase to obtain their meat (Bashaw et al., 2007), thus burning off more calories (Kohn et al., 2011).

2.4 Communication The successful functioning of pride living is dependent on communication (Rudnai, 1973), lions rely on vocalisation, facial expressions, scent marking, olfactory communication, tactile communication and posture to determine signalling between members (Schaller, 1972). Lions use long range signals to communicate consisting of olfactory and scent markings within their environment and vocalisation (Server, 1993). An adult males roars can be heard up to eight kilometres (five miles) away (Rudnai, 1973). Close range signals between members of the pride are complex and are built of intricate patterns (Scenkel, 1966), these include the use of 14

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

teeth, lips, ears, tail and posture (Schaller, 1972) and lions constant communication behaviour is intensified by the use of vocal, olfactory and tactile messages (Schaller, 1972). Vocal communication is considered a territorial display (Schaller, 1972), whilst also promoting group cohesion (Labuschagne et al., 1963), vocalisation communication is expressed in a repertoire of snarls, moans, growls, grunts and roars (Grinnell et al., 2001). However there are less distinctive expressions which include puffing, bleating and humming (Schenkel, 1966), by the age of one month old a lion cub is capable to form all vocal sounds that an adults make, other than a full roar which is not exposed until sub-adulthood (Schaller, 1972). Facial expressions in lions are distinguished by aggressive threat, defence facial expressions and relaxed expressions (Schaller, 1972). Aggressive communication exhibits the display of the mouth almost closed, ears erect but backward facing and pupils small (Schaller, 1972), communication such as this can occur between members as a display of dominance (Rudnai, 1973). With defensive facial expressions canines are exposed and lips are drawn back, ears are flattened and pupils are large (Fox, 1969) an expression of this nature would be exhibited between male to male competition (Fox, 1969). Relaxed expressions distinguishes features in a neutral position (Van Hooff, 1967), ears are erect; pointing laterally, eyes are closed and mouth is either shut or slightly open with lips hanging (Adamson, 1963) usually this expression is apparent during social resting activity (Fox, 1969). The function of scent marking and olfactory communication is used to familiarize living ranges (Schenkel, 1966) and as a deterrent to ward off intruders (Kleiman, 1966). Besides lions spray scent marking they also scent mark by rubbing their heads against objects within their environment such as bushes or trees (Kleiman, 1966). Head rubbing is also exhibited within pride memberâ&#x20AC;&#x2122;s interactions (Schaller, 1972), a lion usually approaches from the side or back of another lion touching cheeks as it gently passes (Schaller, 1972) this act is also considered as a social greeting to re-establish social bonds in the pride by scent marking (Rudnai, 1973) as well as cub protection (Rudnai, 1973). Studies carried out in the Serengeti confirmed 11% of females scent rub cubs (Schaller, 1972). Any tracks left by a lion such as urine or faeces in the environment serves as a signal to other lions within the area (Schenkel, 1966), these markings are usually left in a manner of moving 15

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

through their own territory as during scent marking male lions demonstrate a â&#x20AC;&#x2DC;proud` posture with active looking around vigilance (Schaller, 1972); this is a visual territorial pattern of marking among male lions (Schenkel, 1966). Nomad lions use olfactory senses as part of their routine communication (Grinnell et al., 2001) to gain understanding of resident lions within the area before approaching (Bauer et al., 2003). Tactile communication in lions involves brief interactions between members (Schaller, 1972), this occurs in a variety of situations such as play fights, when crowded around a kill and during resting times (Schenkel, 1966) as often lions sleep closely together with other pride members (Rudnai, 1973), positioning paws over each other or leaning on each other (Schaller, 1972). Other means of initiating contact as mentioned previously is the interaction of head rubbing as well as social licking (Schaller, 1972) these two interactions are the two most common and peaceful tactile gestures (Schaller, 1972), these communication signals help assist with group cohesion (Sparks, 1967). Body posture signals most commonly used by lions in their daily routine are those of walking and resting (Schaller, 1972), these display an element of relaxation (Rudnai, 1973), however a range of postures such as stalking walking, crouch walking and crouch depict more concentrated communication signals (Schaller, 1972). These can be focused towards predation behaviour (Schaller, 1972) as well as play behaviour within lion groups (Rudnai, 1973). These postures usually encourage other lions to respond to these postures thus by looking in the same direction (Schaller, 1972).

2.5 Territorial behaviour Panthera leo are renowned for their sociable life-style and perhaps one of their most important pride shared activity is joint defence of the prideâ&#x20AC;&#x2122;s territory against other females and infanticidal males (Heinsohn, 1997). Male lions advertise their territory by vocalising and by using the number of roars they hear to gauge the strength of the opposition male (Heinsohn, 1997). The majority of territorial vocalisation occurs at dawn and dusk as during this time lions become most active and protective of their range (Mosser et al., 2009). Overall males are significantly more territorial than females (Mosser et al., 2009), this is expected due 16

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

to female mortality and wounding injuries that females get when associated with neighbouring rival males (Mosser et al., 2009). Males are stronger and better protected with their mane covering the sensitive neck area (West et al., 2006). Males often use lethal aggression if necessary to obtain power over outside males (West et al., 2006), in the favour of the pride; as ultimately males display territorial behaviour to protect their pride and offspring (Rudnai, 1973), which primarily benefits group living in the Panthera leo species (Grinnell et al., 1995). Although vocalisation is perhaps the most profound territorial behaviour exhibited by male lionâ&#x20AC;&#x2122;s, olfaction flehmen response and scent marking are also territorial displays (Schaller, 1972); the flehmen response lets lions detect other scent (Schaller, 1972), as mentioned before for the use of a females reproductive state, however males patrolling their territory and pride range use this response to determine the tracks or scent marks left by other males within the area (Rudnai, 1973), consequently encouraging them to vocalise their kinship and dominance within that range (Grinnell et al., 1995). Scent marking in males is as a result of them outlining their occupied territory (Mosseret et al., 2009), males usually sent mark upon objects within their territory for instance trees (Mosseret et al., 2009), this in relation is projected at the height of any rival males (Mosseret et al., 2009); this act in itself is to ward off nomad males and to ensure protection of the pride (Heinsohn, 1997). Protection of the pride is reliant upon the dominant male of the pride (Heinsohn, 1997), as his territorial behaviour is the strongest mechanism a male can provide to reassure the females and offspringâ&#x20AC;&#x2122;s protection and survival (Mosser et al., 2009). This is an essential concept of the Panthera leo group living (West et al., 2006), as infanticide of cubs is a certain occurrence should a nomad male enter an established prideâ&#x20AC;&#x2122;s territory and defeat the fathering male (Mosser et al., 2009). 2.6 Natural behaviour 2.6.1 Resting On average lions spend about 20-21 hours of their day inactive and resting (Schaller, 1972). Resting behaviour has been described by Schaller (1972) as; a lion resting on its stomach, sides or sometimes on its back with hind legs spread (Schaller, 1972). Lions usually choose a prominent resting place, such as a rock or shady patch (Rudnai, 1973), often when seeking 17

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

shade lions will rest together, however males often rest alone, as far as 100m from the rest of the group (Schaller, 1972). 2.6.2 Movement Activity Active locomotion activities include walking, ranging and climbing (Schaller, 1972), lions only spend around two hours during the day actively walking (Schaller, 1972); however 6 hours of this ranging activity was known to be recorded by Schaller (1972). On the other hand lions can spend up to 4-5 hours during the night walking (Schaller, 1972), males are more likely to display this level of activity during the night due to territory patrols (Rudnai, 1973). 2.6.3 Grooming As mentioned earlier male lions tend to remain a distance from the rest of the pride, therefore less allo-grooming takes place between males and females (Schaller, 1972), However females will make attempts to groom males even when they are uncertain of the response they may recieve (Schaller, 1972). Most allo-grooming takes place between female members and females with cubs (Jackman et al., 1982), this social contact is usually present with the behaviour of resting (Rudnai, 1973). 2.6.4 Agonistic behaviour The majority of the time lions live harmoniously together (Rudnai, 1973), this is supported in observations of a group of 7 lions by Schaller (1972) as only one account of aggression was recorded over the course of four days (Schaller, 1972). However agonistic behaviour is natural and often short; residing in a slap, baring of teeth and vocalisation towards another group member (Schaller, 1972). This behaviour is most influenced during a kill (Rudnai, 1973) as lions seem at their most anti-social state; as a result of bolting for meat (Schaller 1972). This can be a vigorous performance leading to snarls of aggression to any group member that feels threatened to share (Schaller, 1972); especially when the kill is small (Schaller, 1972), such as a Thompsonâ&#x20AC;&#x2122;s gazelle (Eudorcas thomsonii). 2.7 Social Structure Lions social structure is extremely complex and prides can include up to 35 individuals (Server, 1993), within this group there are usually up to four non-permanent males (Rudnai, 1973), lionessâ&#x20AC;&#x2122;s make up the greater part of the pride along with their offspring (Rudnai, 1973), which almost always remain a part of the pride their entire life (Server, 1993). The non-permanent 18

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

males are usually driven out of the pride by the age of three (Packer, 1987) and they usually live with brothers or are solitary until the age of five (Server, 1993) where they will then find their own pride in which they will have to drive off the resident male in order to gain leadership themselves (Server, 1993). Pride living provides cooperative hunting ability, which ultimately benefits the group as a whole (Bauer et al., 2003), strength in numbers when hunting provides a 30-40% increased success rate of bringing down large prey such as Cape buffalo (Syncerus caffer) (Packer, 1987), thus appertaining maximum food resources to the pride (Packer, 1987). Group living provides concentrated protection as both the resident male and females protect pride members and offspring working as a joint defence system (Packer, 1987). Perhaps the most important aspect of lion sociality is that pride living has a direct effectiveness over habitat and land resources (Bauer et al., 2003), for instance a well-established pride in terms of group numbers, male dominance, reproductive fitness and fitness in general permits that pride to dominate and include in their territory essential resources such as watering holes (Bauer et al., 2003). Not only for the supply of water but prey abundance, as prey species are attracted by such resources (Mosser et al., 2009); thus by protecting these resources ultimately safeguards the prides survival and the survival of future generations (Mosser et al., 2009).

2.8 Lions in Captivity Panthera leo are commonly kept in zoological collections as they are suggested a popular attraction for visitors (Boler, 2009). However large mammals such as Panthera leo are often provided with enclosures and environments that propose un-natural lifestyles (Breton, 2014). It has been shown that animals that occupy large territorial ranges in the wild, such as lions; are more vulnerable to problems associated with stress in captivity (Morgan et al., 2007) including the abnormal behaviour pacing (Breton, 2014). Pacing is defined as the action of walking back and forth over the same paths in an enclosure (Bashaw et al., 2007), this action is most commonly observed in terrestrial Carnivores including Panthera leo (Breton, 2014). Although lions can spend the majority of their day resting and living in harmony with each other (Schaller, 1972), when housed in captivity lions notability develop increased agonistic

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

behaviours and in worst cases infanticide between members they are housed with (Breton, 2014). Lions in the wild are top of the food chain and they can travel up to 50km per day to find prey (Rudnai, 1973), therefore captivity has a direct impact upon ranging behaviour (Clubb et al., 2003). Lions in captivity require a multitude of different necessities in order to allow them to express natural behaviours (Boler, 2009); these include an outdoor enclosure that contains a diversity of flora species (Boler, 2009) to allow them to scent mark and seek cover (Clubb et al., 2006), an indoor house to have privacy from the public (Clubb et al., 2006), correct nutritional values and methods of feeding (Black et al., 2010) including a starve period to replicate natural biological functions (Clubb et al., 2006), breeding management and enrichment provisions (Clubb et al., 2006). It has been suggested that lions can live longer in captivity (Alert, 2013); this has been suggested as a result of veterinary care (Kleiman et al., 1966). However lions are susceptible to problems when kept in captivity (Breton, 2014); including teeth problems, due to a lack of carcass or meat on bone feeds (Hinds, 2014), as the texture of their diet when not containing bone material is to soft (Clubb et al., 2006) and can also severely reduce their calcium, mineral and vitamin intake (Clubb et al., 2006) in this case supplements should be provided to maintain welfare standards (Clubb et al., 2006). The social structure of lions when kept in captivity is very different from that of the wild (Breton, 2014), as mentioned earlier natural dispersal takes place in a wild pride with males moving out of the pride at maturity (Schaller, 1972), members are able to leave and move on to create a new pride structure elsewhere (Server, 1993), captive lions do not have this ability, often agonistic aggression can be observed in lions housed within small living environments and can sometimes result in fatal attacks between group members or individuals having to be removed (Clubb et al., 2006). Abnormal repetitive behaviours such as pacing are rather prevalent, and often reflect the inadequacy of the lionâ&#x20AC;&#x2122;s enclosure and living conditions (Clubb et al., 2003). A study by Breton (2014) examined if enclosure size had any impact on the display of stereotypical behaviour in 38 captive Tigers (Panthera Tigris), the results showed that enclosure size and ability to range 20

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

greater distance decreased the stereotypical behaviour pacing (Breton, 2014). Therefore it is considered compromised living space can trigger pacing behaviour (Breton, 2014).

2.9 Environmental Enrichment Environmental enrichment has changed over the last 20 years; behaviour enrichment used to be referred to as something keepers could provide if they had time, to now it being an essential part of captive husbandry (Young, 2003). Enrichment provides animals with physical stimulation which is beneficial to their welfare, and provides appropriate ‘naturalistic` or ‘wild` behavioural opportunities (Hill et al., 2009). Zoo enrichment most commonly includes physical enrichment- a structure or object to provide change for interactions within an animal’s enclosure (Young, 2003); such as substrate (Hosey et al., 2009), Cognitive enrichment- to enhance an animal’s learning ability or problem solving (Hosey et al., 2009), social enrichment- pairings or grouping of individuals or species (Hoesy et al., 2009), sensory enrichment- manipulating light, sound, smell and touch (Hosey et al., 2009) and nutritional enrichment- providing different types of food and in different forms of presentation (Hosey et al., 2009). Environmental enrichment success is measured by three main characteristics; Behavioural, physiological and neurological signs (Young, 2003). Enrichment is suggested to improve the biological functioning of animals within a captive environment (Mason et al., 2007) and also stimulate desirable behaviours, increase activity and reduce stereotypical or undesirable behaviours (Young, 2003) and (Hosey et al., 2009). Enrichment for captive animals is broken down into the categories mentioned and is provided using the following methods; visual, olfactory, auditory, tactile, gustatory stimuli and feeding enrichment (Hill et al., 2009).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

2.9.1 Olfactory Enrichment In the wild scent and scent marking is used to distinguish territories, locate prey and group members as well as identifying reproductive status (Young, 2003). Schaller (1972) stated that lions use olfactory stimulation with their Jacobsonâ&#x20AC;&#x2122;s organ in many routine daily life including reproductive state, identifying their home range and identifying other males located close by (Schaller, 1972). Olfactory enrichment has been suggested to encourage high levels of play (Wells, 2009) and interactions that can be rarely observed as a result of a captive environment and habituation (Hill et al., 2009). Olfactory enrichment is said to benefit large cats as they display a direct response to odours within their environment (Wells, 2003), thus increasing the time spent on active behaviours such as locomotion, exploratory and grooming activity (Wells, 2003) Olfactory enrichment is thought to increase natural scent marking behaviour and territorial behaviour in Panthera species (Szokalski et al., 2012).

2.9.2 Food Enrichment Food related enrichment is used to encourage natural behaviours and exploration of the environment through the use of an object (Hill et al., 2009), an animalâ&#x20AC;&#x2122;s food can be presented in a variety of ways to stimulate feeding related behaviours (Young, 2003). Given the fact that lions devote a large proportion of their active time to hunting prey, it should be addressed that Panthera leo when kept in captivity require environmental food related enrichment to preserve and increase their natural hunting and feeding behavioural patterns (Szokalski et al., 2012). As mentioned before wild lions consume a range of prey (Schaller, 1972), after they make a kill lions spend a substantial proportion of time seizing, pulling, gripping, biting and licking their prey (Szokalski et al., 2012), these tactics are needed in order to tear through the preyâ&#x20AC;&#x2122;s tough fur and skin and to strip meat from bones (Schaller, 1972). Food related enrichment given to the Panthera genus can contribute to an increase in natural hunting behaviours such as capturing prey, mock killing and fur plucking (Szokalski et al., 2012), typically food related enrichment is suggested to decrease stereotypical behaviour as demonstrated in a study carried out by Szokalski (2012); as results showed that food related enrichment of fish and bones when given to captive tigers (Panthera tigris) significantly 22

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

reduced stereotypical behaviour by 30% from the original controlled baseline observations (Szokalski et al., 2012). As Panthera leo practice the same natural hunting behaviours and strategies as that of the Panthera tigris food related enrichment is thought to be a useful stimuli for encouraging the display of natural behavioural traits (Szokalski et al., 2012).

2.9.3 Low cost Enrichment The concept behind the choice of the Behavioural enrichment treatments used for the investigation is that they are kept to the lowest cost possible and only contain natural products. As Zimbabwe is a less economically developed country, Antelope Park would benefit from using enrichment treatments which are otherwise less expensive to produce compared to that of enrichments that can be purchased for big cat species, for instance boomer balls. Table 2: shows the availability and costs of the behavioural enrichment types used in the investigation in Zimbabwe currency.

Enrichment Type Log with Zebra Dung Pumpkin Log with Cow blood

Available at All products at Antelope Park Local Village

Cost $0.00

All products at Antelope Park

$0.00

3. Antelope Park Antelope Park (AP) is home to one of the African lion environmental research trust (ALERTâ&#x20AC;&#x2122;s) lion rehabilitation programs; the larger of the two that exist (Alert, 2013). The park is located in over 3000 acres of open savannah grassland habitat (Alert, 2013) and the reintroduction program is structured of four stages (Alert, 2013). Stage 1 begins at the breeding program grounds; which has 82 lions resident (Alert, 2013), these lions are the individuals that provide offspring for the rehabilitation program (Alert, 2013). Records are kept of all the breeding lionsâ&#x20AC;&#x2122; at the park including origin and parentage (Alert, 2013) within a stud book to ensure 23

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

genetic diversity management with the controlled breeding. Stage 2 consists of a structured pride of lions being released into a 403 acre wild managed environment with wild prey available for them to hunt independently (Alert, 2013); thus encouraging the development of social structure and self-sustainability within the pride. Stage 3 is where the rehabilitated pride will be taken to a larger area with greater diversity of game species to hunt and competitive species present such as hyenas (Crocuta crocuta) (Alert, 2013), during this stage it is expected for the pride to reproduce (Alert, 2013). Stage 4 is the process of releasing lion prides back into wild areas with government protection such as reserves (Alert, 2013); population management would be necessary as it is with most lion populations throughout Africa (Alert, 2013). Currently AP is at stage 2 of the program, however a stage 3 site has been secured with only funding holding the lionâ&#x20AC;&#x2122;s translocation process back (Alert, 2013) and reproduction has already occurred within the stage 2 pride (Alert, 2013).

3.1 African Lion Environmental Research Trust (ALERT) ALERT was founded in 2005 at AP (Alert, 2013), it is a non-profit organisation that is dedicated to the facilitation of African lion conservation and management (Alert, 2013). The organisation works with local governments, wildlife authorities and communities in order to build sustainable relationships for the reintroduction program and to promote ecotourism (Alert, 2013). The trustâ&#x20AC;&#x2122;s main objectives is to support the four stage African lion rehabilitation program, to provide a long term solution to the decreasing lion populations; through the release of wild born offspring from rehabilitated captive bred lions (Alert, 2013) and to raise awareness and funds to secure suitable release sites for lions (Alert, 2013).

3.2 Previous Research Past research for behavioural enrichment trailed in Panthera leo is mostly documented in juvenileâ&#x20AC;&#x2122;s or infant cubs, however It has been suggested that in all life stages captive Panthera leo benefit with behavioural enrichment in order to influence social upbringing (Ncube et al., 2010). A previous investigation carried out by Ncube (2010) at Antelope Park studied how 24

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

three groups of lion cubs were stimulated with the presence of sticks, grass, fresh dung from elephant (Loxodonta africana), Wildebeest (Connochaetes), Zebra (Equus quagga), Impala (Aepyceros melampus) Giraffe (Giraffa camelopardalis) and cardboard boxes (Ncube et al., 2010). These enrichments were utilised to encourage active behaviours (Ncube et al., 2010). The results from the study showed that the presence of behavioural enrichment increased activity levels (Ncube et al., 2010). A previous investigation carried out by Quirke (2011) examined the effect of differnet types of behavioural enrichment on the behaviour of captive Cheetahs (Acinonyx jubatus), the study tested temporal variation in feeding, spatial variation in feeding and olfactory enrichment using Scimitar-horned Oryx (Oryx dammah) faeces (Quirke et al., 2011). The results from the study showed that the presence of the (Oryx dammah) faeces determined the most significant increase in locomotion and exploratory behaviour, as well as decreasing inactivity (Quirke et al., 2011). This study in comparison to this investigation proposes similarity in that the use of prey dung is expected to increase activity levels.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

4. Aims and Objectives of the Investigation The aim of the investigation is to determine if the presence of the behavioural enrichments promote natural territorial behaviour displays and if they increase activity levels; as well as reducing stereotypical behaviours in 16 conservation breeding male African lions at Antelope Park, Zimbabwe, Gweru. Observations will evaluate if there is a significant increase or decrease in territorial behaviour displays and general activity levels with the presence of the enrichment types tested. The investigation will also examine which behavioural enrichment (BE) the lions interact with most frequently and which BE type they interact with throughout the whole six hour period; (7.00am-9.00am), (10.00am-12.00pm) and (4.00pm-6.00pm) in order to evaluate the success of each BE type. 4.1 Hypothesis -

Lions will interact most with the food related enrichment (cow blood log) more than the other two behavioural enrichments.

Zebra dung log and cow blood log behavioural enrichments will receive most interactions throughout the day.

More territorial behavioural displays will be exhibited with the presence of the Zebra dung log and cow blood log enrichments.

Zebra dung log and cow blood log will increase activity levels.

4.2 Null Hypothesis -

There will be no significant difference in the lionâ&#x20AC;&#x2122;s interactions between all three behavioural enrichments.

There will be no significant difference between all enrichment types receiving interactions throughout the day.

There will be no significant difference in the amount of territorial behaviour displays between all three behavioural enrichment types.

There will be no significant difference in activity levels with all three behavioural enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

4.3 Alternative Hypothesis -

The lions will prefer the two other enrichments suppose to the food related enrichment cow blood log.

Pumpkin enrichment will receive the most interactions throughout the day.

More territorial behavioural displays will be exhibited with the presence of the behavioural enrichment Pumpkin.

Pumpkin will increase activity levels.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5. METHODOLOGY

5.1 Site descriptions Antelope Park is located 10 km from Gweru in the Highveld of Zimbabwe (19⁰28’S, 29⁰09’E) (Alert, 2013). The mean annual temperature is 24-30⁰C (Climate, 2013) and the annual rainfall is 948-1420 mm (Climate, 2013). Antelope Park is a private game reserve involved in the conservation of African Lions (Jenkins et al., 2013); having its own Lion breeding facility. The breeding and reintroduction program is structured of four key stages from breeding to the final release (Alert, 2013). The programs aim is to captive breed lions, and through careful management and group dynamics composition successfully structure prides that are equipped with the vital behaviours that will enable them to survive in the wild after release (Alert, 2013). ALERT founded in 2005 at Antelope Park, it is a non-profitable organisation dedicated to the promotion of conservation and management plans for the African Lion (Alert, 2013). Its aim is to carry out vital research and investigations at Antelope Park; to gain an understanding of lion ecology, behaviour and dynamics for future references (Alert, 2013).

Figure 3: shows the location of Zimbabwe on the African continent; highlighted in green (ccissues, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Figure 4: shows the location of Gweru, Zimbabwe (Worldtravels, 2014).

Figure 5: shows the location of Antelope Park in Gweru, Zimbabwe (Jenkins et al., 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.2 Behavioural Enrichment The Breeding Male lions at Antelope Park were behaviourally enriched with three natural low cost treatments being the following; a Acacia log smeared with Zebra (Equus burchelli) dung, a Pumpkin - Flat White Boer Ford (Cucurbita maxima) whole as a novel enrichment and a Acacia log smeared with cow (Bos primigenius) blood. The behavioural enrichments aimed to increase a repertoire of natural stimuliâ&#x20AC;&#x2122;s and reduce any abnormal behaviours classified as stereotypical; such as pacing. A main objective of the investigation was to increase territorial behaviour displays in the lions, as being housed in enclosures with only other males can often reduce the triggers and natural displays of territorial behaviours, this is often as a result of habituation to their environment, surrounding settings and each otherâ&#x20AC;&#x2122;s presence (Szokalski et al., 2012). The main aim of increasing territorial behaviours in the breeding male lions at the park is thought to be an important prospect; as territorial behavioural traits are believed to be vital behaviours to be passed on to future generations, especially if those generations are consequently to be the individuals who will be introduced into wild areas with the prospect to survive and protect their pride equally to that of a fully wild pride. 5.3 Materials for the investigation Acacia Logs, Zebra dung, Cow blood, Pumpkins, Ethogram, recording data sheets, pens, waterproof lithium battery stop watch (additional spare one), water, spare paper for additional notes and watermelons.

Figure 6: shows the stopwatch used for the investigation (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.3.1 Controlled study A controlled study was carried out on each lion group that was used in the investigation to record behaviour frequencies displayed without the influence of any behavioural enrichment treatments refer to figure 19. These were unfortunately carried out on days amongst the main investigation; only due to time constraints, as in an ideal situation a controlled study would have been carried out before the main investigation was started. However when the controlled studies were carried out it was insured that they were performed amongst a post treatment stage with each lion group. Controlled studies were as follows; Lion group one on the 21/07/13, lion group two on the 07/07/13 and lion group three on the 18/07/13; thus during the investigation period. 5.3.2 Pilot study A Pilot study was constructed and carried out of the 4th of July 2013 to ensure that the format and structure of data collection should run as smoothly as possible. The original behavioural enrichment program that was first constructed included the use of a Watermelon as the novel enrichment as proposed to the Pumpkins that were used in the final investigation. The pilot study was performed using the Watermelon as a trail to test and evaluate its overall effectiveness. The pilot study was trailed with one of the lion groups that was then distinguished as a final group to be used throughout the investigation, known as Lion Group one. After the pilot study was constructed it was apparent from analysis that the Watermelon was only effective up until the second recording time period categorised as the mid-morning session (10.00am-12.00pm). After this outcome was confirmed, further examinations of both the watermelon and pumpkins was required. Measurements and weights were taken of both and both were cut open to test the difference in density. After this procedure It was determined that the pumpkin was slightly larger, a lot heavier and also had a tougher Skin and fleshy centre; which was thought to be an advantage over the watermelon; thus suggesting it would possibly be more robust and stay whole for longer, therefore encouraging more interactions. After this was decided it was concluded that the pumpkin would be the

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

third enrichment type used in the investigation; acting as the novel enrichment replacing the original concept of using a watermelon. Table 3: Shows the difference in measurements and weights between the Watermelon and Pumpkin.

Enrichment Type Watermelon Pumpkin

Measurements 160x160mm 275x110mm

Weight 2.0Kg 3.6Kg

Figure 7: shows the size difference between the Flat White Boer Ford pumpkin (Cucurbita maxima) and the Watermelon (Citullus lanatus) which concluded why it thought best to use the pumpkins over the watermelon due to size differences (Keel-Stocker, 2013).

Figure 8: shows a pumpkin that was not used in the investigation but cut open to determine concentration value (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.4 Lions and enclosures The investigation was carried out during July 2013, for the period of one month using three groups of adult male breeding lions; with groups added all together a total of 16 lions were exposed to the three behavioural enrichment treatments. All three groups were housed in three separate enclosures, but within close proximity of each other. For easily identification the groups were labelled as Lion group 1, 2 and 3. Each of the enclosures contained grass, tall trees and numerous small shrubs and bushes; unknown identification. Group 1 and 3 also had platforms present in there enclosure, whereas group 2 had no platform level. All of the lion groups used in the investigation were housed with a fence line directly next to another group of lions that was also used in the study and group 3 was also housed next to another group of adult male lions that was not used in the investigation on the far side of their enclosure refer to figure 9. On one side of all three enclosures the fence line faced open grassland of the parks protected fenced perimeter area which holds a range of fauna species most of which are natural prey species to lions; these included Zebra (Equus burchelli), Giraffe (Giraffa camelopardalis), Impala (Aepyceros melampus), Wildebeest (Connochaetes) Kudu (Tragelaphus strepsiceros) and jackals (Canis aureus).

Table 4: shows the lions groups used in the investigation; group composition and ages.

Lion Group 1

Composition Five males

2 3

Four males Seven males

Age 6 years x2, 5 years x2, unknown x1 9 years x3, 10 years x1 9 years x5, 8 years x 2

Table 5: shows the details of the Lions in Group 1

Lion Group 1: Name Damisi Echo Etosha Kosey Batoka

D.O.B 15/09/2008 12/04/2007 12/04/2007 Unknown 15/03/2008

Mother Unknown Elsa Elsa Unknown Unknown

Father Unknown Casper Casper Unknown Unknown

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Table 6: shows the details of the lions in Group 2

Lion Group 2: Name Arthur Casper Mufasa Maxwell

D.O.B 26/01/2004 17/06/2004 09/01/2004 14/03/2003

Mother Anna Unknown Mafuza Unknown

Father Mara Unknown Duma Mara

Table 7: shows the details of the lions in Group 3

Lion Group 3: Name Achillies Apollo Kwezi Mambo Mamba Phoenix Penduka

D.O.B 08/11/04 08/11/04 31/10/04 27/09/04 27/09/04 20/02/05 20/02/05

Mother Anna Anna Kitty Mafuta Mafuta Penga Penga

Father Teddy Teddy Teddy Teddy Teddy Teddy Teddy

Figure 9: shows the Breeding Program grounds at Antelope Park with the three groups of lions used in the investigation- Group 1, 2 and 3 highlighted in blue (keel-Stocker, 2014).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.4.1 Site area measurements All of the lion groups used in the investigation were housed in approximately the same size enclosures; when selecting the lion groups for the investigation this was an aspect taken into consideration, as lions housed within the same living space was thought to limit any variables as regards to space and enrichment presence. Table 8: shows the measurements in metres of the lion groups enclosures used in the investigation.

Lion Group

Enclosure length

Enclosure Width

55m

15m

50m

15m

50m

Figure 10: shows the lion enclosures and lion groups used in the investigation (Keel-Stocker, 2014)

5.5 Design of the study The investigation involved the introduction of three natural low cost behavioural enrichment treatments to three groups of adult African male breeding lions at the breeding program grounds at Antelope Park. Behavioural enrichment treatments have been abbreviated in the following way. Behavioural enrichment Log with Zebra dung (BE:LD), Behavioural enrichment Pumpkin (BE:P) and Behavioural enrichment Log with cow blood (BE:LB).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.5.1 Procedure Ten behaviours were recorded during the investigation; active locomotion, vocalisation, resting, behaviour enrichment feeding related, behaviour enrichment touch related, social behaviour, territorial behaviour, grooming, aggression and stereotypical behaviour; refer to table 12. Each behavioural enrichment (BE) types were introduced to a group of lions for a period of three days in the order of BE:LD, BE:P, BE:LB once they had be tested with the three day enrichment treatment programme they were given a post-treatment period of six days prior to the next enrichment schedule. The enrichment program followed in the same order with each lion group. Each BE was positioned in the enclosures at approximately 6:50am every morning to ensure that each lion group could be let into the enclosure promptly at 7:00am; so that data collection could begin. With each lion group, each of the BE treatments were all positioned in the same location within each of the enclosures to limit any variables such as preferences to any areas within the enclosure. Scan sampling with an interval of 60 seconds was the method used throughout the investigation for data collection and a total of six hours was recorded per day. The data recording start time was from 7:00am until 6:00pm the observational recording time was divided up into three time periods throughout the day in order to determine the effectiveness of the behavioural enrichments over time, also to allow comparisons against the different time periods. These recording sessions were categorized as; Early morning (7:00am – 9:00am) Midday (10:00am – 12:00pm) and Afternoon (4:00pm – 6:00pm). A total of 36 hours was obtained from each of the lion groups, giving a total of 180 hours overall. Each lion group tested each BE treatment twice, therefore a total of 12 hours data was collected for each behavioural enrichment treatment for each lion group. Giving a concluded total of 6 samples and 36 hours per each enrichment type and 18 samples overall.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.5.2 Data analysis The frequencies recorded for each behaviour with each enrichment type gave a total of six samples per each BE type. The samples were then expressed into an overall total for each enrichment type; making all samples combined for each BE type. For each behaviour observed in the investigation the three BE type results for every behaviour were tested against each other by conducting a Chi Square goodness of fit test. This was to determine a P-value in order to determine if there was a significant difference between the data sets; to gain a conclusion of which BE type observed the most interactions and encouraged the most active locomotion and territorial displays.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.6 Preparing Behavioural Enrichment: Log with Zebra Dung For the enrichment log with Zebra dung a few procedures took place. An Acacia log was chosen within the measurement of no greater than 1metre long; refer to table 9. The logs that were chosen had an outward dry layer of bark covering them to allow the dung used in the investigation better grip. Zebra dung was collected from the surrounding grassland within the fenced area of Antelope Park, Zebra dung was collected approximately three days before it was used for each study and stored under cool conditions of approximately 5â&#x2014;ŚC -6â&#x2014;ŚC in order for it to retain its flexibility and aroma. Once needed for the study two ample handfuls were used for each log. In order to apply the dung to the log and to encourage it to bond to the log 250ml of water was poured over the dung beforehand; this stimulated several effects; the dung to be more apply-able to the surface of the log and the scent of the dung to be refreshed. The dung was then applied to the log using fingers to smear and rub the substance into the bark along the length surface and surface ends of the log.

Figure 11: shows some of the Zebra dung that was used in the investigation (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Figure 12: shows the process of applying the Zebra dung to the log (Keel-Stocker, 2013).

Figure 13: shows the BE:LD fully prepared with a layer of zebra dung, ready to be transferred into a Lion enclosure (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.6.1 Preparing Behavioural Enrichment: Pumpkin The Novel Behavioural Enrichment Pumpkin was measured and weighed beforehand in order to maintain similarity between all pumpkins used in the investigation; refer to table 11. This was the only procedure that took place in order to prepare the pumpkins for use in the investigation.

Figure 14: shows one of the pumpkins that was used in the investigation (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.6.2 Preparing Behavioural Enrichment: Log with Cow Blood To prepare the behavioural enrichment log with cow blood, the first step was to obtain enough cow blood for the investigation. The Lion handler staff collected the blood for the investigation from the cattle carcasses that were used for Lion feeds. The Blood was drained out of several carcasses into some plastic bottles at roughly equal measures. Once enough blood had been collected the bottles of blood were stored in the meat freezer, this was to ensure that the blood remained fresh throughout the investigation. The night before the BE:LB was to be tested the following day; a bottle of blood was collected from the freezer and left over night to defrost at room temperature of approximately 6â&#x2014;ŚC. The Acacia logs that were selected for the BE:LB also measured approximately 1metre long, refer to table 10; this was to ensure likeness to the other Behavioural enrichment treatment BE:LD, also to limit size variables between the two enrichments. Ensuring that the size of logs were similar would suggest that any preference determined after the investigation in the results of the investigation would not be as a result of size preference to logs between the two enrichment treatments especially as they are using in the same mechanism. On the morning the BE:LB was to be prepared for testing 100ml of cow blood was drained from the bottle of blood it was then mixed with 50ml of water, the blood mixture was then poured slowly over the log, being patted and smeared into the bark along the length surface and ends surface of the log using fingers to ensure some abortion into the bark; in order to safeguard that the blood was retained within the log and to maximise the duration of blood presence once in the Lion enclosure.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Figure 15: shows some of the Cow blood samples that were used for the BE:LB in the investigation (Keel-Stocker, 2013).

Figure 16: shows a BE:LB that was used in the investigation (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Figure 17: shows a BE:LB that was used in the investigation (Keel-Stocker, 2013).

Figure 18: shows the end of a BE:LB to demonstrate how the end surface area was covered with the Cow blood (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

5.6.3 Measurements of Acacia logs used in the investigation Table 9: shows the measurements of the logs used in the investigation for the BE:LD; length, diameter and weight. (Keel-Stocker, 2013)

Logs used for the Behavioural Enrichment: Log with Zebra Dung Date Log was used 05/07/2013 10/07/2013 13/07/2013 17/07/2013 20/07/2013 24/07/2013

Lion Group 1 2 3 1 2 3

Length (mm) 830 810 825 790 795 800

Diameter (mm) 130 140 135 140 135 140

Weight (kg) 7.25 8.16 7.25 7.25 6.80 8.16

Table 10: shows the measurements of the logs used in the investigation for the BE:LB; length, diameter and weight. (Keel-Stocker, 2013)

Logs used for the Behavioural Enrichment: Log with Cow Blood Date Log was

used 09/07/2013 12/07/2013 16/07/2013 19/07/2013 23/07/2013 26/07/2013

Lion Group 1 2 3 1 2 3

Length (mm) 810 790 805 815 800 785

Diameter (mm) 140 135 135 130 140 130

Weight (kg) 7.25 7.70 7.70 7.70 8.16 7.25

5.6.4 Measurements of the Flat White Boer Ford Pumpkins used in the investigation Table 11: shows the measurements of the Pumpkins used in the investigation for BE:P; diameter, height and weight. (Keel-Stocker, 2013)

Pumpkins used for the Behavioural Enrichment: Pumpkin Date Pumpkin was used 08/07/2013 11/07/2013 15/07/2013 18/07/2013 22/07/2013 25/07/2013

Lion Group 1 2 3 1 2 3

Diameter (mm) 275 270 280 270 290 265

Height (mm) 110 110 105 115 100 120

Weight (kg) 3.60 3.50 3.60 3.60 3.40 3.50

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

6. Ethogram and definitions used in the investigation

Table 12: shows the list of behaviours used in the ethogram with their descriptions defined.

Behaviour Category Active Locomotion

Behaviour Descriptions Walking, running or climbing

Vocalisation

Roaring or grunting

Resting

Sitting, standing sleeping (Non vigilant)

Behavioural Enrichment- Feeding Related

Licking, biting chewing or swallowing

Behavioural Enrichment- Touch Related

Interaction with BE object, sniffing, moving over, carrying (movements erratic, over exaggerated and spontaneous)

Social Behaviour

Social play, greeting, rubbing interactions and social grooming

Territorial Behaviour

Olfaction flehmen response and Scent marking

Grooming

Self-grooming

Aggression

Aggressive growling, pawing, biting another lion and chasing another lion

Stereotypical Behaviour

Pacing

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

7. RESULTS 7.1 Graphs and Graph Analysis: Raw data can be seen in the Appendix 12.1. The results from the investigation show a clear outcome that the behaviour enrichment Zebra dung log was the most successful enrichment treatment. BE:LD encouraged the most activity with active locomotion levels at 24%, as a result of this the least amount of resting activity was observed at 49%. BE:LD encouraged more territorial behaviour than the BE:P and BE:LB enrichment types; with 5%. Overall the BE:LD was the enrichment treatment that was interacted with most frequently in both of the behaviour enrichment interaction categories; Behavioural enrichment feeding related 5% and Behavioural enrichment touch related 7%.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Pilot study results comparison between watermelon & Pumpkin enrichment: Figure 19 Graph 1 shows the results from the Pilot study with the Watermelon trialling as a behavioural enrichment and the first trail in the investigation using the pumpkin enrichment. Both of these observations were recorded from the same group of Lions; Group 1. Although this data cannot be examined to determine if there is a significant difference between the successes of these enrichments; it is apparent from the results displayed in Graph 1 that the pumpkin enrichment exhibited an increase in territorial behaviour and both of the enrichment interaction categories; BE feeding related and BE touch related. The graph suggests that the watermelon enrichment did substantially encourage Active locomotion, however as this is the results from only one six hour observation these results could simply be a outcome of random chance. The watermelon also seemed to increase stereotypical behavioural displays with the frequency of 7 recorded for the six hour pilot study, again this could be down to random chance. However the pumpkin enrichment seemed to discourage stereotypical behaviour with the frequency of only 1 recorded for the first six hour observation; this was later confirmed in the investigation as the BE:P significantly observed the least stereotypical behavioural displays with its presence.

Frequency

Pilot study: Watermelon enrichment vs Pumpkin enrichment 400 350 300 250 200 150 100 50 0

Behaviours Watermelon

Pumpkin

Figure 19: Graph 1, showing the results from the Pilot study using Watermelon enrichment and the first observation results of using the Pumpkin enrichment in the investigation with the same group of lions.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Summary Graph showing behaviour frequencies for all behaviour enrichment types: Figure 20 Graph 2 shows all behavioural frequencies displayed for all the behavioural enrichment types throughout the investigation. The graph shows that the behaviours active locomotion, vocalisation, territorial behaviour, BE feeding related, BE touch related and stereotypical behaviour was displayed the most with the presence of the BE:LD. The graph shows that the BE:P also significantly encouraged territorial behaviours and vocalisation as well as ultimately being a successful enrichment that was interacted with second most in BE feeding related and BE touch related; compared to that of the BE:LB, which was least successful in encouraging territorial behaviours, vocalisation and both the enrichment related interaction categories.

Bar Graph showing all Behaviour Enrichment types and behaviour Frequencies observed 2500

Frequency

2000 1500 1000 500 0

Behaviours Zebra Dung

Pumpkin

Blood

Figure 20: Graph 2; shows all behaviour frequencies observed in all behavioural enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Controlled study behaviour frequencies compared to behaviour frequencies recorded with the presence of BE types: Figure 21 Graph 3 shows the results from the three day controlled observational studies. Behaviours such as active locomotion, vocalisation and territorial behaviour are significantly displayed less compared to the frequency of these behaviours displayed with the behavioural enrichment treatments. When controlled behaviour frequencies were compared to the behaviour frequencies with the BE treatments there was a considerable difference. Although there is only three samples obtained for controlled observations when comparing graph 3 to graphs 4, 5 and 6 containing 6 samples it is apparent that across the observations from the controlled conditions active locomotion, vocalisation and territorial behaviours were expressed considerably less under these conditions despite having only three samples worth of data.

Behaviours observed from the controlled observations 1200

1060

Frequency

1000 800 600 400

244

200 33

Behaviours Figure 21: Graph 3; shows the results from the controlled observations.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Behaviour frequencies observed with the presence of log with zebra dung enrichment: Figure 22 Graph 4 shows the behaviour frequencies observed with the presence of the Zebra dung log enrichment. The graph shows that the highest frequency was seen in resting behaviour. High frequencies were recorded for the behaviours active locomotion, vocalisation, territorial behaviour, stereotypical behaviour and both BE categories; BE- feeding related and BE-touch related compared to the other two behavioural enrichments refer to Graph 5 and 6.

Behaviour Frequencies observed with the presence of the Zebra dung log Enrichment 2500 1958

Frequency

2000 1500 1000

952

500 164

206

277 67

183

Behaviours

Figure 22: Graph 4; shows the behaviour frequencies observed with the presence of the BE:LD.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Behaviour frequencies observed with the presence of Pumpkin enrichment: Figure 23 Graph 5 shows the behaviour frequencies observed with the presence of the pumpkin enrichment. The graph shows that Resting was the most frequently observed activity and was the highest observed frequency for this behaviour compared to BE:LD and BE:LB. The graph shows that there was still a relatively high frequency observed for active locomotion compared to that observed in the BE:LB refer to graph 6. The highest frequency of Social behaviour was observed with the presence of the BE:P compared to that of the other two BE types. Territorial behaviour and both BE categories; BE- feeding related and BE-touch related observed the second highest frequency of interactions second to the BE:LD. The presence of the BE:P recorded the lowest frequency of Aggressive behaviour out of all three BE types.

Behaviour Frequencies observed with the presence of the Pumpkin Enrichment 2500 2045

Frequency

2000 1500 1000

789

500 153

174

213

124

151

Behaviours

Figure 23: Graph 5; shows the behaviour frequencies observed with the presence of the BE:P.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Behaviour frequencies observed with the presence of log with cow blood enrichment: Figure 24 Graph 6 shows the behaviour frequencies observed with the presence of the Cow blood log enrichment. The graph shows that the highest frequency was observed in resting behaviour. Compared to BE:LD and BE:P there is a considerable decrease in observations of vocalisation, social behaviour, territorial behaviour and both the BE interaction categories BE- feeding related and BE- touch related. The graph shows that the BE:LB received the highest frequency in aggressive behaviour observations when compared to graphs 4 and 5.

Behaviour Frequencies observed with the presence of the Cow blood log Enrichment 2500 2026

Frequency

2000 1500 1000

864

500 113

110

135

146

Behaviours Figure 24: Graph 6; shows the behaviour frequencies observed with the presence of the BE:LB

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Behaviour frequencies throughout the early morning, midday and afternoon time periods with Log with Zebra dung enrichment: Figure 25 Graph 7 shows the observations of all behaviours recorded for the BE:LD. The graph shows that BE:LD was interacted with throughout the whole day; with the highest frequencies of both interaction categories; BE- feeding related and BE- touch related recorded with the presence of this enrichment type. The highest frequency of interactions with the behavioural enrichment took place during the morning period, yet the enrichment was also interacted with throughout the midday and afternoon period, however considerably less in the afternoon period. In correlation with this the highest frequency of territorial behaviour was observed during the morning period with a slow decrease throughout the day. Territorial observations were exhibited across the whole day with the presence of the BE:LD.

Observations of behaviours with the Zebra dung log enrichment throughout the morning, midday and afternoon time periods 800 700

Frequency

600 500 400 300 200 100 0

Behaviours Dung (Morning)

Dung (Midday)

Dung (Afternoon)

Figure 25: Graph 7; shows the observations of all behaviours with the Zebra dung log enrichment throughout all time periods.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Behaviour frequencies throughout the early morning, midday and afternoon time periods with Pumpkin enrichment: Figure 26 Graph 8 shows the observations of all behaviours with the BE:P. The graph shows that BE:P was interacted with throughout the whole day. The most interactions took place during the morning period and almost identical measures were recorded for both behavioural enrichment categories; BE- feeding related and BE- touch related during the midday and afternoon periods. This makes the BE:P the most successful enrichment in terms of stable interactions throughout the whole day. In correlation with the most behavioural enrichment interactions taking place during the morning period the highest frequency of territorial behaviour displays were recorded. Territorial behaviour displays remained in constant balance with the enrichment interactions.

Frequency

Observations of behaviours with the Pumpkin enrichment throughout the morning, midday and afternoon time periods 800 700 600 500 400 300 200 100 0

Behaviours Pumpkin (Morning)

Pumpkin (Midday)

Pumpkin (Afternoon)

Figure 26: Graph 8; shows the observations of all behaviours with the Pumpkin enrichment throughout all time periods.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Behaviour frequencies throughout the early morning, midday and afternoon time periods with log with cow blood enrichment: Figure 27 Graph 9 shows the observations of all behaviours with the BE:LB. The graph shows that the BE:LB observed interactions throughout the whole day but with a dramatic decrease in the midday and afternoon periods. The most interactions with the behavioural enrichment took place during the morning period in both behaviour enrichment categories; BE- feeding related and BE- touch related. Behaviour enrichment interactions were closely in equilibrium for both behaviour enrichment categories during the midday period, however received the lowest interactions compared to that of the BE:LD and BE:P. The afternoon period only received BEtouch related interactions with an extremely low frequency compared to the behavioural enrichment interaction frequencies observed in the other two behavioural enrichment types during this time period. Territorial behaviour frequencies were recorded the most during the morning period, least during the midday period and a slight increase during the afternoon period. Overall territorial behaviour was the least observed with the presence of the BE:LB compared to the other two behaviour enrichment types. Observations of behaviours with the Cow blood log enrichment throughout the morning, midday and afternoon time periods 800 700

Frequency

600 500 400 300 200 100 0

Behaviours Blood (Morning)

Blood (Midday)

Blood (Afternoon)

Figure 27: Graph 9; shows the observations of all behaviours with the Cow blood log enrichment throughout all time periods.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Behaviour enrichment feeding related interaction for all BE types: Figure 28 Graph 10 shows the frequencies that were recorded for all behavioural enrichment types for the BE- feeding related category. BE:LD was the enrichment type that was interacted with the most for BE- feeding related throughout the whole investigation. The BE:P was second to be interacted with in this Behaviour enrichment category and BE:LB observed the least interactions for this category.

Behaviour Enrichment - Feeding Related interation frequencies recorded for all Behavioural Enrichment types 250 206 200

Frequency

174 150 110 100

0 Dung

Pumpkin

Blood

Enrichment Type

Figure 28: Graph 10; shows the frequencies recorded for the BE- feeding related interactions for all the Behavioural Enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Behaviour enrichment touch related interaction for all BE types: Figure 29 Graph 11 shows the frequencies that were recorded for all behavioural enrichment types for the BE- touch related category. BE:LD was the behaviour enrichment that was interacted with the most for this category, BE:P received the second most interactions under this category and blood received the least amount of interactions under this category.

Enrichment - Touch Related interaction frequencies recorded for all Behavioural Enrichment types 300

277

250 213

Frequency

200 135

150 100 50 0 Dung

Pumpkin

Blood

Enrichment Type

Figure 29: Graph 11; shows the frequencies recorded for the BE- touch related interactions for all the Behavioural Enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Active locomotion frequencies recorded for all BE types: Figure 30 Graph 12 shows the active locomotion frequencies recorded for all behavioural enrichment types. The graph shows that BE:LD recorded the highest frequency of active locomotion displays throughout the investigation; which was considerably more than the other two behavioural enrichment types. The BE:LB recorded the second most active locomotion displays throughout the investigation and the BE:P recorded the least amount of active locomotion displays.

Active Locomotion frequencies recorded for all Behavioural Enrichment types 1000

952 864

900 789

800

Frequency

700 600 500 400 300 200 100 0 Dung

Pumpkin

Blood

Enrichment Type Figure 30: Graph 12; shows the frequencies recorded for Active Locomotion displays for all Behavioural Enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Vocalisation frequencies recorded for all BE types: Figure 31 Graph 13 shows the frequencies recorded for vocalisation behaviour in all behavioural enrichment types. The graph shows that vocalisation behaviour was observed the most frequently with the presence of BE:LD, second most in the BE:P and the least observed in the BE:LB.

Vocalisation frequencies recorded for all Behavioural Enrichment types 180

164 153

160 140

113

Frequency

120 100 80 60 40 20 0 Dung

Pumpkin

Blood

Enrichment Type

Figure 31: Graph 13; shows the frequencies recorded for vocalisation behaviour for all Behavioural Enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Territorial behaviour frequencies recorded for all BE types: Figure 32 Graph 14 shows the frequencies recorded for territorial behaviour in all behavioural enrichment types. The graph shows that BE:LD observed the most territorial behaviour displays compared to the other two behavioural enrichment types. The BE:P received the second most observed territorial behaviour displays with its presence and the BE:LB observed the least amount of territorial behaviour displays.

Territorial Behaviour frequencies recorded for all Behavioural Enrichment types 200

183

180 160

Frequency

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124

120 100

80 60 40 20 0 Dung

Pumpkin

Blood

Enrichment Type Figure 32: Graph 14 shows the frequencies recorded for Territorial behaviour displays for all Behavioural Enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Aggressive behaviour frequencies recorded for all BE types: Figure 33 Graph 15 shows the observations that were recorded for Aggressive behaviour for all behavioural enrichment types. The graph shows that BE:LB observed the most aggressive behavioural displays throughout the investigation. BE:LD received the second most aggressive behavioural frequencies and the BE:P received a considerable reduced frequency of aggressive behavioural displays.

Aggressive Behaviour frequencies recorded for all Behavioural Enrichment types 16

Frequency

10 8 6 4 4 2 0 Dung

Pumpkin

Blood

Enrichment Type Figure 33: Graph 15; shows the frequencies recorded for Aggressive behaviour for all Behavioural Enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Stereotypical behaviour frequencies recorded for all BE types: Figure 34 Graph 16 shows the observations for Stereotypical behaviour observed in all behavioural enrichment types. The graph shows that BE:LD observed the most stereotypical behaviour displays during the investigation, BE:LB observed the second most frequent display of stereotypical behaviour and the BE:P received the least amount of stereotypical behaviour displays during the investigation.

Stereotypical Behaviour frequencies recorded for all Behavioural Enrichment types 80

70 59

Frequency

60 50 40 29

30 20 10 0 Dung

Pumpkin

Blood

Enrichment Type Figure 34: Graph 16; shows the frequencies recorded for Stereotypical behaviour for all Behavioural Enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

7.2 Statistical Analysis of results Table 13: shows the results from the chi square test for BE- feeding related interactions.

Enrichment

Behaviour

Zebra Dung log

BE – feeding related

Pumpkin

BE – feeding related

Blood log

BE – feeding related

P-value

0.0004

Significance

Significant Difference

Table 14: shows the results for the chi square test for BE- touch related interactions.

Enrichment

Behaviour

Zebra Dung log

BE – touch related

Pumpkin

BE –touch related

Blood log

BE – touch related

P-value

0.0003

Significance

Significant Difference

Null Hypothesis: There will be no significant difference in the lion’s interactions between all three behavioural enrichments. Alternative Hypothesis: The lions will prefer the two other enrichments suppose to the food related enrichment cow blood log. Table’s 13 and 14 shows the Chi square results of both BE interaction categories between all BE types, as shown by the P-value for both of the BE interaction categories; BE- feeding related and BE-touch related there is a significant difference in the data showing that with the presence of BE:LD the most interactions occurred. Although BE:P also received a high frequency of interactions throughout both the BE categories there was still a significant difference between the two BE types; proving BE:LD was the BE most interacted with. The P63

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

value therefore supports the alternative hypothesis and rejects the null hypothesis as the BE:LD received the most BE interactions. Table 15: shows the results from the chi square test for resting behaviour displays.

Enrichment

Behaviour

Zebra Dung log

Resting

Pumpkin

Resting

Blood log

Resting

P-value

0.353053909

Significance

No Significant Difference

Table 15 shows the results from the Chi square test to determine if there was a significance difference between all BE types effecting the display of resting behaviour. According to the Pvalue >0.05 there was no significant difference between all three BE types affecting the amount of resting behaviour displayed.

Table 16: shows results for the chi square test for Active locomotion displays.

Enrichment

Behaviour

Zebra Dung log

Active Locomotion

Pumpkin

Active Locomotion

Blood log

Active Locomotion

P-value

0.000468655

Significance

Significant Difference

Hypothesis: Zebra dung log and cow blood log will increase activity levels. Null Hypothesis: There will be no significant difference in activity levels with all three behavioural enrichment types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Alternative Hypothesis: Increased activity levels will be displayed most frequently with the presence of the behavioural enrichment Pumpkin. Table 16 shows that the results from the Chi square test for active locomotion displays between all BE types. According to the P-value result <0.05 there was a significant difference between the presence of the different BE types contributing to active locomotion displays subsequently BE:LD observed the greatest frequency of active locomotion displays.

Table 17: shows the results from the chi square test for Vocalisation displays.

Enrichment

Behaviour

Zebra Dung log

Vocalisation

Pumpkin

Vocalisation

Blood log

Vocalisation

P-value

0.006567767

Significance

Significant Difference

Table 17 shows the results from the Chi square test for Vocalisation displays with all BE types. According to the P- value <0.05 there was a significant difference between the amount of vocalisation displays with the presence of BE:LD as this BE type positively influenced an increase in vocalisation displays.

Table 18: shows the results from the chi square test for Social behavioural displays.

Enrichment

Behaviour

Zebra Dung log

Social

Pumpkin

Social

Blood log

Social

P-value

0.060611012

Significance

No Significant Difference

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Table 18 shows the results from the Chi square test for social displays between all BE types. As shown by the P- value there was no significant difference between all three BE types affecting the frequency of social displays.

Table 19: shows the results from the chi square test for Territorial behavioural displays.

Enrichment

Behaviour

Zebra Dung log

Territorial

Pumpkin

Territorial

Blood log

Territorial

P-value

0.0003

Significance

Significant Difference

Hypothesis: More territorial behavioural displays will be exhibited with the presence of the Zebra dung log and cow blood log enrichments. Null Hypothesis: There will be no significant difference in the amount of territorial behaviour displays between all three behavioural enrichment types. Alternative Hypothesis: More territorial behavioural displays will be exhibited with the presence of the Pumpkin enrichment. Table 19 shows the results from the Chi square test for territorial displays between all BE types. As shown by the P-value <0.05 there is a significant difference between the presence of the BE types and the frequency of territorial behavioural displays observed. The presence of the BE:LD increased territorial displays; this supports the hypothesis and rejects both the null and alternative hypothesis.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Table 20: shows the results from the chi square test for grooming behavioural displays.

Enrichment

Behaviour

Zebra Dung log

Grooming

Pumpkin

Grooming

Blood log

Grooming

P-value

0.0002

Significance

Significant Difference

Table 20 shows the results from the Chi square test for grooming displays between all BE types. According to the P- value <0.05 there is a significant difference between the data; confirming that the BE:P increased the frequency of grooming displays more than the other two BE types.

Table 21: shows the results from the chi square test for Aggressive behavioural displays.

Enrichment

Behaviour

Zebra Dung log

Aggressive

Pumpkin

Aggressive

Blood log

Aggressive

P-value

0.05

Significance

No Significant Difference

Table 21 shows the results from the Chi square test for aggressive behavioural displays between all BE types. According to the P- value >0.05 there is no significant difference between all BE types upon influencing aggressive behavioural displays.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Table 22: shows the results from the chi square test for Stereotypical behavioural displays.

Enrichment

Behaviour

Zebra Dung log

Stereotypical

Pumpkin

Stereotypical

Blood log

Stereotypical

P-value

0.0004

Significance

Significant Difference

Table 22 shows the results from the Chi square test for stereotypical behavioural displays between all BE types. The P- value <0.05 determined that there is a significant difference between the BE types upon influencing stereotypical behavioural displays. According to the results BE:LD increased the display of stereotypical behavioural significantly more than the other two BE types.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

7.3 Photographs of African lions interacting with the BE treatments

Figure 35: shows a lion displaying touch related interaction with the BE:LD (Keel-Stocker, 2013).

Figure 36: shows a lion displaying feeding related interaction with BE:LD (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Figure 37: shows a lion displaying touch related interaction with BE:P (Keel-Stocker, 2013).

Figure 38: shows a lion displaying feeding related interaction with the BE:P (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Figure 39: shows a lion displaying touch related interaction with BE:P in a playful nature (Keel-Stocker, 2013).

Figure 40: shows the BE:P after being given to a group of lions for the duration of one six hour observational day (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Figure 41: shows a lion displaying touch related interaction with the BE:LB (Keel-Stocker, 2013).

Figure 42: shows a lion displaying feeding related interaction with the BE:LB (Keel-Stocker, 2013).

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

8. DISCUSSION The aim of the investigation was to determine if the presence of the BE types tested promoted natural territorial behaviour displays and if they increased activity levels; as well as reducing stereotypical behaviours. The investigation was also to examine which low cost behavioural enrichment the lions would interact with most throughout the six hour period. Observations were recorded to evaluate the success of each BE type. The three different BE types were put into each of the three lion group enclosures twice on a rotational basis with a post treatment stage of six days between each enrichment schedule. Each BE type was observed for a total of 36 hours, to see if the BE type had an effect on the lions behaviour. The results from the investigation carried out at Antelope Park, Zimbabwe showed that there was a significant difference in many of the observed behaviours with the presence of the three BE types during the 18 day investigation. 8.1 Behaviour Enrichment effectiveness 8.2 Log with Zebra dung Enrichment The results from four of the Chi square tests showed that the presence of the BE:LD produced a significant P-value of <0.05 upon the display of several behaviours; including active locomotion, vocalisation, territorial behaviour and stereotypical behaviour. The P-values for all of these behaviours were significantly <0.05; concluding that this BE type significantly influenced the display of these behaviours more frequently than the other two BE types refer to tables 16, 17, 19 and 22. A significant P-value was also expressed in both of the BE interaction categories (BE- feeding related and BE-touch related), as the BE:LD produced a Pvalue significantly <0.05 for both of these categories; refer to tables 13 and 14. BE:LD also received interactions throughout the whole six hour period (morning, midday and afternoon). These results determined that BE:LD was the BE that was not only the BE interacted with most frequently but also interacted with for the duration of the whole day when tested; therefore accomplishing one of the main aims of the investigation; to examine which BE the lions interact with most frequently and which BE type they interact with throughout the whole six hour period in order to evaluate the success of each BE type.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Active locomotion behaviour is whereby a lion is either displaying walking, running or climbing activity (Schaller, 1972), this was expressed most frequently in the morning and midday time periods with the BE:LD, perhaps as a result of the lions interacting with the BE:LD most frequently during these time periods, refer to figure 25. This is a positive outcome as effective BE is expected to increase activity levels (Young, 2003). Vocalisation (Roaring or grunting) was expressed the most with the presence of this enrichment refer to figure 31 and throughout the whole day more so than the other two BE types, this suggests that the presence of this BE is stimulating vocalisation- a territorial behaviour. This could be as a result of this BE type containing living prey related olfactory smell; thus encouraging a stimulus to vocalise as a territorial measure, to protect territory and prey within their range (Rudnai, 1973). Lions vocalise to advertise territorial ownership, intimidate rivals, to locate other pride members and to strengthen social bonds between other group members (Schaller, 1972). Vocalisation is most commonly expressed when lions are most active at dusk (Rudnai, 1973), which explains why a correlation is seen between most frequent active locomotion levels and the highest frequency of vocalisation displayed in both the midday time period for BE:LD refer to figure 25. Territorial behaviour (Olfaction flehmen response and Scent marking) was observed most during the morning time period with the BE:LD; refer to figure 25, as were the most interactions recorded with this enrichment type for both BE categories, proposing that this BE type had a profound effect upon stimulating territorial behaviours. As the P-value was considerably <0.05 for territorial behaviour and both of the BE interactive categories it is rational to suggest that the presence of the BE:LD increased territorial behaviour displays. Therefore making it the most effective BE type in the investigation, as these results accomplish one of the main aims of the investigation; to determine if the presence of the behavioural enrichments promotes natural territorial behaviour displays. Stereotypical behaviour was observed most with the presence of the BE:LD producing a Pvalue significantly <0.05, however although this behaviour was recorded most with the presence of this BE treatment during all observations additional notes were made which indicate that the majority of all stereotypical displays were as a result of the presence of

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

vehicles entering the breeding programme site area, this is further discussed under the stereotypical outcomes section. The results from resting behaviour observations found that there was no significant difference between all BE types influencing this display thus producing a P-value of >0.05 refer to table 15. As resting behaviour was mostly observed throughout the afternoon time period this could just be an outcome of natural circadian rhythms, as lions spend up to 22 hours a day resting (Schaller, 1972) and more predominantly during late afternoon when temperatures are highest (Rudnai 1973) therefore the high occurrence of this behaviour is natural. 8.2.1 Pumpkin Enrichment The highest behaviour frequency that was observed in the investigation was Resting behaviour with the presence of the BE:P. Resting behaviour is whereby the lion is sitting, standing or sleeping (Non vigilant) (Schaller, 1972). Resting was observed with BE:P most during the afternoon time period; refer to figure26, as was resting behaviour in the other two BE types during the afternoon refer to figures 25, 26 and 27. As a result of the high frequency of resting, active locomotion was the least observed with the BE:P throughout the investigation. Grooming behaviour was observed most with BE:P, this could be as a result of the high amount of resting behaviour observed as grooming behaviour usually occurs with the behaviour of relaxation in Panthera species (Server, 1993). BE:P received the second highest frequencies to BE:LD for both the BE interaction categories, however a greater frequency in the touch related category; refer to figure 26, this was as a result of the pumpkin being more frequently played with refer to figure 39. The BE:P encouraged natural playful behaviour, similar to play behaviour that can be exhibited when lions play and toss their prey (Schaller, 1972), this behaviour encouraged the use of the lions retractable claws also, which is a use that can become castoff in captivity due to a lack of hunting ability (Hinds, 2014). Vocalisation was observed second most with the presence of the BE:P; refer to figure 31, however vocalisation was recorded at the highest frequency during the afternoon period when both BE categories recorded the lowest frequency of interactions; possibly suggesting that there is not a significant relationship between vocalisation and the BE:P. Territorial behaviour was also the second most observed with the presence of the BE:P throughout the 75

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

investigation; refer to figure 32, with the highest frequency being recorded during the morning time period. Both BE interaction categories recorded the highest frequencies during the morning time period also; refer to figure 26, which could suggest a possible association between the territorial displays and interaction with the BE:P therefore it is thought that BE:P increased territorial behaviour displays. Social behaviour between lions was observed the most with the presence of the BE:P refer to figure 20, as was the highest frequencies recorded in the morning and midday time periods refer to figure 26, which were similarly the two time periods that recorded the highest frequency in both the BE interaction categories. From the additional notes that were made throughout the investigation it was noted that lions expressed social interactions whilst interacting with the BE:P such as; social greetings, rubbing and social play with the pumpkin, these types of interactions are important from group cohesion and bonds (Rudnai, 1973). The least amount of both aggressive and stereotypical behaviour was observed with the presence of the BE:P; refer to figures 33 and 34 compared to both the other two BE types for both these categories. However the statistical analysis for aggressive behaviour determined a P-value of >0.05 stating that there was no significant difference between all three BE types effecting the display of aggressive behaviour. However the statistical results for stereotypical behaviour produced a P-value that was significantly <0.05 determining that stereotypical behaviour was decreased with the presence of BE:P.

8.2.2 Log with Cow blood Enrichment The lowest frequency was recorded with the presence of BE:LB for a range of behaviours in the investigation such as; vocalisation, territorial behaviour and both the BE interactive categories; refer to figures 28,29,31 and 32. Vocalisation was observed the most during the afternoon time period and considerably less during the other two time periods (morning and midday) refer to figure 27; this suggests that the increase for vocalisation in the afternoon could just be as a result of natural circadian rhythms as lions often vocalise territorial calls at dusk (Schaller, 1972 and Rudnai 1973), which coincides with the afternoon time period (4:00pm â&#x20AC;&#x201C; 6:00pm). The highest frequency of both BE interactive categories took place during 76

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

the morning period, therefore the vocalisation results for BE:LB does not determine a relationship. However Territorial behaviour was observed the most during the morning time period so could propose that there is a slight relationship between BE:LB interactions and an increase in territorial behavioural displays as often territorial behaviour is displayed with the presence of food close by (Rudnai, 1973). As BE:LB was the only BE that contained a related food source that the lions are used to, it was suggested in the hypothesis of the investigation that BE:LB would receive the most interactions yet interestingly it received the least. The statistical results revealed a P-value of <0.05, thus concluding a significant difference between the three BE types therefore concluding that BE:LB received an extremely low frequency for both of the BE interaction categories; making it the least effective BE in the investigation. Active locomotion was observed the second most frequently with the presence of the BE:LB and the frequency levels throughout all time periods during the day were more equal to one another than that compared to the other two BE types which observed peak times of high and low frequencies, this suggests that this could of just been the lions natural routine, or this could suggest that the BE:LB made lions active throughout the whole day, maybe as a result of anticipation for food due to the nature of the BE containing cow blood, a common meat given in captivity (Hinds, 2014). Aggressive behaviour was observed most with the presence of the BE:LB refer to figure 33, however the statistical analysis produced a P-value of >0.05 showing that there was no significant difference. It could be thought that the increase of aggressive behaviour could be as a result that this was the only food related enrichment used in the investigation; containing the scent of blood which could naturally induce aggression towards members within the group; possibly due to dominance and the fact BE:LB resembled food. This conclusion could also bare reason to the lower frequency of social behaviour observed with the presence of this BE as even though the statistical analysis stated that there was no significant difference refer to table 18, it is apparent from figure 20 that BE:LB received less social interactions when compared to the similar frequencies observed between that of BE:LD and BE:P.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

8.3 Findings from the Investigation The findings from the investigation suggest that the most effective BE type was the log with Zebra dung as according to the P-value results this BE type significantly increased the lions activity levels, vocalisation and territorial behaviour. Previous research testing the effect of prey dung upon the activity levels in cheetahs (Acinonyx jubatus) discovered that the prey dung of Scimitar-horned Oryx (Oryx dammah) faeces when used as an enrichment received the highest percentages of locomotion activity and exploratory behaviour; locomotion defined as movement from one location to another at any speed and exploratory behaviour defined as olfaction and scent marking (Quirke et al., 2011) in comparison these outcomes are similar to this investigation. 8.3.1 Territorial behaviour Outcomes Similarly territorial behaviour defined in this investigation for Panthera leo species was defined as olfaction, flehman response and scent marking and consequently received the highest frequency with the presence of the BE containing prey dung BE:LD. Vocalisation was likewise observed most frequently with the BE:LD refer figure 31 which remained constant throughout all time periods throughout the day refer to figure 25, this could suggestively be territorial vocalisation or a result of communication between group members linked with the changes in their environment (Schaller, 1972), perhaps the presence of the enrichment. This results from the investigation supports two of the proposed hypotheses being; “More territorial behavioural displays will be exhibited with the presence of the Zebra dung log and cow blood log enrichments” and “Zebra dung log and cow blood log will increase activity levels”. However territorial displays were least observed with the presence of BE:LB as was vocalisation behaviour, a behaviour associated with territorial behaviour (Rudnai, 1973), therefore the BE:LB was least effective throughout this investigation for encouraging territorial behaviour displays. Observations showed that the presence of the BE:P recorded the second highest frequencies in both territorial behaviour and vocalisation; therefore making it a successful low cost enrichment influencing territorial behaviours second to the BE:LD.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

The importance of encouraging territory behaviour in the male lions used in the investigation at Antelope park is that they are part of a breeding programme, as mentioned earlier territorial behaviour is an essential behavioural repertoire of male lions as these behaviours in the wild protect the rest of the pride members and offspring (Schaller, 1972). Territorial behaviours like many other instinctive behaviours can be lost in captivity due to habituation (Hoesy et al., 2009), thatâ&#x20AC;&#x2122;s why it is vital to provide BE to stimulate natural behaviours (Young, 2003). The offspring fathered by the male lions used in this investigation are introduced once mature into the stage 2 wild managed area; in the prospect that their offspring will be the candidates for the final release programme at stage 4 (Alert, 2013). It is thought to be essential that offspring have instinctive territorial behavioural traits in order to live life independently, protect their offspring, other pride members, pride range and ultimately survive as wild lions (Alert, 2013). If the breeding lions are not stimulated with BE to encourage and retain valuable instinctive behaviours such as territorial behaviours and to reduce stereotypical behaviours, habitation and natural behaviours could be reduced and lost, which ultimately are a behavioural traits that should be passed down to future generations. It is essential that these behavioural traits are not lost if future captive bred lion generations are to once become wild lions and have to survive within a wild ecosystem. 8.3.2 Activity outcomes Encouraging activity levels is important for lions in captivity as they often range over large areas in the wild; up to 12km a day (Schaller, 1972). Captivity can often result in animals living in the same enclosure and conditions for long periods of time (Young, 2003), which can lead to boredom, in-activity and stress behaviours (Morgan et al., 2007). The use of enrichment treatments can stimulate natural behaviours that are essential to animals in captivity and animals that are part of a breeding programme (Mason et al., 2007). Pacing is a very common stereotype observed in carnivores (Mason et al., 2007), as stated by Mason (2007) stereotypies are often displayed in carnivores due to food anticipation. The BE:LD significantly observed the most active locomotion displays with a significant P-value of <0.05, making it the most successful BE for increasing activity levels. The Log with cow blood enrichment encouraged a high frequency of active locomotion, this could have been a 79

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

result of this being the food related enrichment; thus encouraging exploration of the environment through the use of this object (Young, 2003). This BE type could have encouraged activity due to the lions anticipating food as this enrichment was the only enrichment containing food related material (Cow blood). The BE:P observed the least active locomotion displays; refer to figure 30, therefore BE:P recorded the highest frequency of resting behaviour. Furthermore these results support the hypothesis that â&#x20AC;&#x153;Zebra dung log and cow blood log will increase activity levelsâ&#x20AC;?. 8.3.3 Stereotypical outcomes The lions at Antelope park displayed stereotypical behaviour most frequently with the presence of the BE:LD, however when the additional notes were examined from the observations a repetitive occurrence associated with the trigger of stereotypical pacing was in fact continually influenced by the presence of vehicles entering the breeding programme grounds. The lionâ&#x20AC;&#x2122;s meat is transported to the enclosures by some of these same vehicles, suggestively giving reason for the lions associating feeding time with the noise or visualisation of vehicles. To make this claim more robust vehicles entering the site that did not have as association with meat delivery also triggered the stereotypical behaviour of pacing instantly when present, thus concluding the noise associated with vehicles triggered stereotypical behaviour. When compared to the observations of stereotypical behaviour recorded with the presence of the other two BE types stereotypical behaviour was significantly less with the BE:P as the P-value produced was <0.05; refer to figure 34 and table 22. Although this could simply be to random chance, for instance less vehicles entering the site during observations with the BE:P. Although BE:LD observed the most stereotypical displays both the other two BE types also recorded frequencies of stereotypical behavioural displays; giving reason to the vehicle noise trigger concept and that possibly the statistical results could just be a coincidence.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

8.3.4 Aggressive behaviour outcomes Aggressive behaviour was observed the most with the presence of the BE:LB, this could be as a result of the association of food related to this BE type, the aggressive behaviour was agonistic towards members within the group, however this is thought to be a natural behaviour (Schaller, 1972), occurrences of aggression lasted for a minimal amount of time which has been stated by Schaller (1972) that agonistic behaviour is often shown for short intervals. Agonistic behaviour is most influenced during a kill (Schaller, 1972), which could explain why aggressive behaviour was recorded most with the presence of the BE:LB. Agonistic behavioural displays are usually in the form of snarls (Rudnai, 1973) and are displayed to another member of the group if an individual feels threatened (Schaller, 1972) this is most apparent during the time a small kill is made such as a Thompsons gazelle (Eudorcas thomsonii) (Schaller, 1972). This could suggest why aggression was observed

significantly more with the BE:LB as the lions may have envisaged the food related enrichment type as a small prey species leaking blood. Nevertheless agonistic behaviour is a natural occurrence (Schaller, 1972) and so perhaps this should not be thought as, as a negative outcome but a positive one, as the lions displayed natural behaviour similar to the occurrence that would be displayed between a wild group of lions.

8.3.5 Natural Enrichment vs. Novel Enrichment The results from the investigation showed that the alternative hypothesis was supported; â&#x20AC;&#x153;The lions will prefer the two other enrichments suppose to the food related enrichment (cow blood log)â&#x20AC;?. The purpose of the investigation was to provide the lions with low cost behavioural enrichments to stimulated natural behaviours such as locomotion activity and territorial behaviour. The concept of using the behavioural enrichments BE:LD and BE:LB was to provide the lions with natural materials that would be present within their wild environment at a low cost price. The purpose of the BE:P is that it would resemble a novel enrichment; an object that lions would not naturally come across within their wild environment, but it still be made of natural material and be low cost. Both the novel BE:P and BE:LD were the two most successful enrichments, however the natural BE:LD more so. 81

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

The results from the investigation show that the BE:P was the second most successful BE type receiving the second highest frequencies to the BE:LD in behaviours; vocalisation and territorial behaviour; refer to figures 31 and 32. Therefore BE:P is a successful enrichment for stimulating territorial behaviour in male lions, however not as successful as the other two BE types at encouraging activity, as BE:P recorded the lowest frequency for active locomotion.

8.3.6 Behavioural enrichment Interactions Overall the behavioural enrichment that received the most interaction in both of the interaction categories (BE-feeding related) and (BE- touch related) was the BE:LD, refer to figures 28 and 29, these interactions were displayed throughout the course of the whole day in all three time periods (Morning, Midday and Afternoon); refer to figure 25, and most frequently compared to the other two enrichments; especially against the BE:LB which the lions lost interest in and stopped interacting with after the first time period (early morning) refer to figure 27. As already mentioned BE:P received the second most interactions in both behaviour categories, like the BE:LD it also received both forms of interaction throughout all time periods ( Morning, Midday and Afternoon); refer to figure 26. As both the BE:LD and BE:P observed interactions throughout all time periods (the duration of a day) this makes these two BE types successful long lasting enrichment treatments.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

9. CONCLUSION Typically captive animals display repetitive behaviours that are otherwise not seen in the wild (Mason et al., 2007) stereotypical behaviour is often present in large mammals that are constrained in enclosures (Mason et al., 2007). Behavioural enrichment is used to stimulate natural and desirable behaviours that are reduced when in captivity (Morgan et al., 2007), environmental enrichment is essential for breeding animals as the reduction of stress in captivity proposes benefits to successful mating and production of healthy offspring (Morgan et al., 2007), as well as vital behavioural characteristics and traits being passed on to future generations (Mason et al., 2007). In conclusion to the investigation carried out at Antelope Park, Zimbabwe the BE types BE:LD and BE:P had the most positive effect on increasing natural behaviours in Panthera leo, furthermore with success in promoting territorial behaviours and increased activity levels as well as providing mental stimulation for the lions while interacting with these BE types. Therefore this indicates that the Null hypotheses “There will be no significant difference in the amount of territorial behaviour displays between all three behavioural enrichment types” was proved incorrect and the alternative hypothesis “The lions will prefer the two other enrichments suppose to the food related enrichment cow blood log” can be accepted. Overall the BE:LD increased active locomotion, vocalisation and territorial behaviour most frequently, it was also the most effective BE type thus observing the most interactions in both interaction categories as well as the BE type that received interactions throughout all time periods; proving its effectiveness as a long lasting enrichment. In summary BE:LD was the BE type that observed most interactions in both BE interaction categories and it also increased the most activity with the highest frequency observed in active locomotion. BE:LD was the most successful BE tested in the investigation as it increased the desirable territorial behaviours; vocalisation, scent marking and flehmen response. BE:LB observed the least interactions and also had no effect upon territorial behaviour displays; making it the BE that was least effective. BE:P recorded the second highest frequencies to the BE:LD in both BE interaction categories, vocalisation and territorial behaviour and the lowest frequency in Stereotypical behaviour, therefore overall a successful BE treatment.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

10. FUTURE WORK 10.1 Limitations It could be suggested that there were various variables that were out of the recorders control such as the occurrence of vehicles entering the breeding program grounds on a daily basis, this created increased noise levels at the site and as mentioned earlier seemed to trigger the occurrence of pacing activity; an abnormal behaviour that is classified as a stereotypical behaviour. This could have had an effect on the amount of observations recorded for stereotypical behaviour and in set terms may not be a true reflection of lions behaviour with the presence of any of the BE tested in the investigation. Time may have also affected the investigation as only 36 hours of observations was recorded with each BE type. If the investigation was carried out over a longer period of time, it could possibly determine the behavioural enrichment effectiveness further and examine if the lions interaction with each of the behavioural enrichments declines over time. 10.2 Further Investigations Future investigations could be carried out to investigate if the presence of vehicle noise is the main trigger for the stereotypical displays in the breeding lions at Antelope Park, as an investigation of this nature could benefit the park in reducing the lionâ&#x20AC;&#x2122;s amount of stereotypical behaviour, which ultimately is a benefit for the lionâ&#x20AC;&#x2122;s welfare long term. As the findings from this investigation conclude that the BE:P reduced stereotypical behaviour as the least frequency was recorded with this BE type and a significant p-value was produced from the data, an investigation could be carried out or combined with the vehicle concept investigation to determine if the BE:P truly reduces the display of stereotypical behaviour evaluated against a controlled investigation.

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

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Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

12. APPENDICES

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

12.1 Appendix Raw data

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

12.2 Data collection Sheet

Copy of the recording sheet that was used for data collection throughout the investigation. (Keel-Stocker, 2013)

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

12.3 Raw statistical data Enrichment

Behaviour

Zebra Dung log

BE – feeding related

Pumpkin

Blood log

Zebra Dung log

BE – touch related

Blood log

Total

174

Formatted: 163.3333 0.0004

110 490

Behaviour

Chi Square: 4.43498E-07

163.3333

BE – feeding related

Enrichment

Chi-Square

206

BE – feeding related

Total

Pumpkin

Observed Expected

Significance

Significant Difference

163.3333

Observed Expected

Chi-Square Chi Square: 2.86691E-11

277

208.3333

213

Formatted: 208.3333 0.0003

135

208.3333

BE –touch related BE – touch related

625

Significance

Significant Difference

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Enrichment

Behaviour

Zebra Dung log

Active Locomotion

Pumpkin

Active Locomotion

Blood log

Active Locomotion

Total Enrichment

Behaviour

Zebra Dung log

Vocalisation

Pumpkin

Blood log

952

868.3333

789

868.3333

864 2605

868.3333

Observed Expected

164

143.33333

153

143.33333

113 430

143.33333

Vocalisation

Chi-Square

Chi Square: 0.000468655

Chi-Square

Chi Square: 0.006567767

Significance

Significant Difference

Significance

Significant Difference

Vocalisation

Total

Enrichment

Behaviour

Zebra Dung log

Resting

Pumpkin

Resting

Blood log

Resting

Total

Observed Expected

1958

2009.667

2045

2009.667

2026 6029

2009.667

100

Chi-Square

Significance

Chi Square: 0.353053909

No Significant Difference

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Enrichment

Behaviour

Zebra Dung log

Social

Pumpkin

Blood log

46 183

Behaviour

Zebra Dung log

Territorial

Observed Expected

Chi Square: 0.060611012

No Significant Difference

Chi-Square

Significance

Chi Square: 131.6667 2.58518E-08

124

Formatted: 131.6667 0.0003

Territorial 88 395

Enrichment

Behaviour

Zebra Dung log

Grooming

Pumpkin

Grooming

Significant Difference

131.6667

Observed Expected

Total

Significance

183 Territorial

Total

Blood log

Chi-Square

Social

Enrichment

Blood log

67 Social

Total

Pumpkin

Observed Expected

Chi-Square

127

Chi Square: 1.72361E-05

151

127 Formatted: 0.0002

146 381

127

Grooming

101

Significance

Significant Difference

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

Enrichment

Behaviour

Zebra Dung log

Aggressive

Pumpkin

Blood log

Behaviour

Zebra Dung log

Stereotypical

Total

15 30

Aggressive

Enrichment

Blood log

11 Aggressive

Total

Pumpkin

Observed Expected

54.33333

59 163

54.33333

Stereotypical

102

Chi-Square

Significance

Chi Square: 0.045049202

No Significant Difference

Formatted: 0.05

Chi-Square

Chi Square: 4.37592E-05 Formatted: 0.0004

Significance

Significant Difference

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

12.4 Risk Assessment

RISK ASSESSMENT Assessment Area: Antelope Park, Ngamo Road, Gweru. Zimbabwe Date of Assessment: 29/06/13 Assessment Completed by: Michelle Keel-Stocker Activity: Dissertation Research Location: Antelope Park, Ngamo Road, Gweru. Zimbabwe Equipment: Ethogram Recording chart, pens, stopwatch, spare paper each day for additional notes, logs, Zebra Dung, Cow blood and Pumpkins. IDENTIFY HAZARDS: 

Working in a close proximity with Lions,



Entering enclosures to position enrichment each morning



Carrying heavy logs



Using knives to extract prey blood from carcass

IDENTIFY WHO MAY BE HARMED (tick): Employees :

Students: X 

Members of Public: Contractors: Others:

EVALUATE RISK: List Existing Controls then assess risk to give residual risk rating: 

Working in close proximity with lions always had a mobile phone on me for safety.



When entering lion enclosures I was always accompanied by a Lion handler.



When carrying heavy logs my father was there for assistance and help.



When using knives they were always used facing away from myself and stored in a safe place.

103

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

RESIDUAL RISK RATING (tick) Likelihood

Severity

Risk Rating

Trivial Injury

= 1

X X

0–5

Low

Possible Occurrence =

Minor Injury

= 2

6 – 11

Medium

XXX

= 3

12 – 16

High

Unlikely Occurrence 1

2 =

Major Injury

Probable Occurrence =

Fatal

Likely Occurrence 3

= 4

Additional Controls Measures (needed if risk rating above ‘LOW’) then reassess for final risk rating: 

Needed to be on guard at all times while working in close proximity with Lions and also have my mobile phone on me for security purposes.

FINAL RISK RATING (tick) High

Medium

Supervisor responsible for controlling work activity: Staff at Antelope park and Lion Handers

104

Low

Echo - “My favourite Lion at Antelope Park”