A multifactorial approach to improving captive primate welfare and enclosure usage Thesis submitted in accordance with the requirements of the University of Liverpool for the degree of Doctor in Philosophy by Colleen Goh June 2016 1 Table of Contents Chapter 1: Introduction .............................................................................................. 1 1.1 Background information ................................................................................... 1 Chapter 2: Tackling Stereotypical Behaviour in Captive Western Lowland Gorillas . 12 2.1 Introduction .................................................................................................... 12 2.2 Materials and methods ................................................................................... 14 2.2.1 Study sites and subjects ........................................................................... 14 2.2.3 Sampling method and variables recorded ............................................... 18 2.2.4 Changes made at Zoo A ........................................................................... 20 2.2.5 Statistical analysis .................................................................................... 21 2.3 Results ............................................................................................................. 23 2.3.1 Description and comparison of behaviour profiles at Zoo A and B in first part of study ...................................................................................................... 23 2.3.2 Identification of forms of enrichment that encourage species-typical behaviour .......................................................................................................... 27 2.3.3 Behaviour profile at Zoo A in second part of study .................................. 28 2.3.4. Differences in behaviour profiles at Zoo A between first and second part of study ............................................................................................................. 28 2.3.5 Behavioural comparison between hand reared and parent reared individuals for both zoos during both parts of study ........................................ 29 2.4 Discussion ....................................................................................................... 29 2.4.1 Differential frequencies in activity between captivity and wild may eXplain R&R eXpression in captivity .................................................................. 29 2.4.2 Differential frequencies of ingestion and social behaviour between zoos associated with R&R ......................................................................................... 30 2.4.3 Social structure in captive gorillas influences amount of mental stimulation and subsequently affects R&R frequency ...................................... 31 2.4.4 Differential frequencies of ingestion and social behaviour between adults and juveniles likely eXplain R&R at Zoo B ......................................................... 32 2.4.5 Dietary fibre levels highly likely to be associated with R&R in Zoo A gorillas ............................................................................................................... 32 2.4.6 Tool-use based enrichment reduces AAB ................................................ 34 2 2.4.7 Different types of AAB stem from different reasons ............................... 35 2.5 Conclusions ..................................................................................................... 36 Chapter 3: The role of habitat structure in the locomotor repertoire of captive and wild gorillas ............................................................................................................... 39 3.1 Introduction .................................................................................................... 39 3.2 Methods .......................................................................................................... 44 3.2.1 Data collection and study sites ................................................................ 44 3.2.2 Variables recorded ................................................................................... 47 3.2.3 Statistical analysis .................................................................................... 57 3.3 Results ............................................................................................................. 58 3.3.1 Comparison of profiles of locomotion and posture between both zoos . 58 3.3.2 Vertical space usage in relation to behaviour/posture/locomotion ........ 62 3.3.3 Support usage in bipedal locomotion and posture .................................. 67 3.3.4 Support usage in relation to vertical space usage and behaviour ........... 68 3.3.5 Wild gorilla positional behaviour profile .................................................. 69 3.3.6 Wild and captive gorilla support usage .................................................... 71 3.3.7 Support diameter and positional behaviour in wild and captive western lowland gorillas ................................................................................................. 73 3.4 Discussion ....................................................................................................... 74 3.4.1 Comparison of arboreal locomotor behaviour profile between captive and wild datasets .............................................................................................. 74 3.4.2 Gorillas in both zoos adopt terrestrial feeding postures similar to that of the wild ............................................................................................................. 76 3.4.3.Vertical space usage was somewhat similar to that of wild .................... 76 3.4.4 Differences in vertical space usage between zoos ................................... 77 3.4.5 Zoos provide similar support size distribution to that of the wild ........... 78 3.4.6 Differences in support availability influence support preference between wild and captive environments, but not between wild sites ............................ 78 3.4.7 Support use in bipedalism ........................................................................ 79 3.4.8 Between species comparison of arboreal positional behaviour in wild dataset .............................................................................................................. 81 3.4.9 SeXual differences in support size usage ................................................. 81 3 3.5 Conclusion ....................................................................................................... 82 Chapter 4: 3D Musculoskeletal Model of a Western Lowland Gorilla Hindlimb ....... 85 4.1 Introduction .................................................................................................... 85 4.2 Methods .......................................................................................................... 93 4.2.1 Dissection and subject data ..................................................................... 94 4.2.2 Building the 3D musculoskeletal model ................................................... 95 4.2.3 Collecting moment arm data ................................................................. 100 4.2.4. Collecting torque data ........................................................................... 103 4.2.5 Incorporating kinematic and joint angle data into our moment arm and torque graphs .................................................................................................. 104 4.2.6 Collecting moment arm data with varying support sizes ....................... 104 4.2.7 Comparison with Payne’s data ............................................................... 106 4.3 Results ........................................................................................................... 106 4.3.1 Hypothesis 1 ........................................................................................... 106 4.3.2 Hypothesis 2 ........................................................................................... 113 4.3.3 Hypothesis 3 ........................................................................................... 121 4.3.4 Hypothesis 4 ........................................................................................... 122 4.4 Discussion ..................................................................................................... 130 4.4.1 Hypothesis 1 ........................................................................................... 130 4.4.1 Hypothesis 2 ........................................................................................... 134 4.4.3 Hypothesis 3 ........................................................................................... 137 4.4.4. Hypothesis 4 .......................................................................................... 140 4.6 AppendiX ....................................................................................................... 142 Chapter 5: Using a 3D Computer Model of the Enclosure to Enclosure and Support Usage in Siamangs .................................................................................................. 145 5.1 Introduction .................................................................................................. 145 5.2 Materials and methods ................................................................................. 147 5.2.1 Building Lidar model .............................................................................