<p><strong>EAZA Best Practice Guidelines </strong><br><strong>Bonobo (Pan paniscus) </strong></p><p><strong>Editors: </strong>Dr Jeroen Stevens </p><p><strong>Contact information: </strong>Royal Zoological Society of Antwerp – K. Astridplein 26 – B 2018 Antwerp, Belgium <strong>Email: </strong><a href="mailto:[email protected]" target="_blank">[email protected] </a></p><p><strong>Name of TAG: </strong>Great Ape TAG </p><p><strong>TAG Chair: </strong>Dr. María Teresa Abelló Poveda – Barcelona Zoo <a href="mailto:[email protected]" target="_blank">[email protected] </a></p><p><strong>Edition: </strong>First edition - 2020 </p><p></p><ul style="display: flex;"><li style="flex:1">1</li><li style="flex:1">2</li></ul><p></p><p>EAZA Best Practice Guidelines disclaimer </p><p>Copyright (February 2020) by EAZA Executive Office, Amsterdam. All rights reserved. No part of this publication may be reproduced in hard copy, machine-readable or other forms without advance written permission from the European Association of Zoos and Aquaria (EAZA). Members of the European Association of Zoos and Aquaria (EAZA) may copy this information for their own use as needed. The information contained in these EAZA Best Practice Guidelines has been obtained from numerous sources believed to be reliable. EAZA and the EAZA APE TAG make a diligent effort to provide a complete and accurate representation of the data in its reports, publications, and services. However, EAZA does not guarantee the accuracy, adequacy, or completeness of any information. EAZA disclaims all liability for errors or omissions that may exist and shall not be liable for any incidental, consequential, or other damages (whether resulting from negligence or otherwise) including, without limitation, exemplary damages or lost profits arising out of or in connection with the use of this publication. Because the technical information provided in the EAZA Best Practice Guidelines can easily be misread or misinterpreted unless properly analysed, EAZA strongly recommends that users of this information consult with the editors in all matters related to data analysis and interpretation. </p><p>EAZA Preamble </p><p>Right from the very beginning it has been the concern of EAZA and the EEPs to encourage and promote the highest possible standards for husbandry of zoo and aquarium animals. For this </p><p>reason, quite early on, EAZA developed the “Minimum Standards for the Accommodation and Care of Animals in Zoos and Aquaria”. These standards lay down general principles of animal </p><p>keeping, to which the members of EAZA feel themselves committed. Above and beyond this, some countries have defined regulatory minimum standards for the keeping of individual species regarding the size and furnishings of enclosures etc., which, according to the opinion of authors, should definitely be fulfilled before allowing such animals to be kept within the area of the jurisdiction of those countries. These minimum standards are intended to determine the borderline of acceptable animal welfare. It is not permitted to fall short of these standards. How difficult it is to determine the standards, however, can be seen in the fact that minimum standards vary from country to country. Above and beyond this, specialists of the EEPs and TAGs have undertaken the considerable task of laying down guidelines for keeping individual animal species. Whilst some aspects of husbandry reported in the guidelines will define minimum standards, in general, these guidelines are not to be understood as minimum requirements; they represent best practice. As such the EAZA Best Practice Guidelines for keeping animals intend rather to describe the desirable design of enclosures and prerequisites for animal keeping that are, according to the present state of knowledge, considered as being optimal for each species. They intend above all to indicate how enclosures should be designed and what conditions should be fulfilled for the optimal care of individual species. </p><p>3</p><p><strong>EAZA Great Ape Taxon Advisory Group (2020): </strong></p><p><strong>Chair: </strong></p><p>María Teresa Abelló, Barcelona Zoo – <a href="mailto:[email protected]" target="_blank">[email protected] </a></p><p><strong>Vice-Chair: </strong></p><p>Sandra Reichler, Zoo Heidelberg –<a href="mailto:[email protected]" target="_blank">[email protected] </a></p><p><strong>Bonobo EEP Coordinator: </strong></p><p>Jeroen Stevens, Antwerp – <a href="mailto:[email protected]" target="_blank">[email protected] </a></p><p><strong>Bonobo EEP Vice-Coordinator: </strong></p><p>Zjef Pereboom, Antwerp – <a href="mailto:[email protected]" target="_blank">[email protected] </a></p><p><strong>International Studbook Keeper </strong></p><p>Zjef Pereboom, Antwerp – <a href="mailto:[email protected]" target="_blank">[email protected] </a><br>4</p><p>Contents </p><p><a href="#3_0">EAZA Best Practice Guidelines disclaimer ............................................................................................... 3 </a><a href="#3_1">EAZA Preambl</a><a href="#3_1">e</a><a href="#3_1">.</a><a href="#3_1">....................................................................................................................................... 3 </a><a href="#5_0">Contents .................................................................................................................................................. 5 </a><a href="#8_0">Introduction............................................................................................................................................. 8 </a><a href="#10_0">Section 1: Biology and Field data .......................................................................................................... 10 </a><br><a href="#13_0">1.1. Taxonom</a><a href="#13_0">y</a><a href="#13_0">.</a><a href="#13_0">.............................................................................................................................</a><a href="#13_0"> </a><a href="#13_0">13 </a><a href="#15_0">1.2. Morpholog</a><a href="#15_0">y</a><a href="#15_0">.</a><a href="#15_0">..........................................................................................................................</a><a href="#15_0"> </a><a href="#15_0">15 </a><br><a href="#15_1">1.2.1. </a><a href="#16_0">1.2.2. </a><br><a href="#15_1">Body Size........................................................................................................................ 15 </a><a href="#16_0">General appearance ...................................................................................................... 16 </a><br><a href="#18_0">1.3. Physiology..............................................................................................................................</a><a href="#18_0"> </a><a href="#18_0">18 </a><a href="#19_0">1.4. Longevity</a><a href="#19_0"> </a><a href="#19_0">............................................................................................................................... 19 </a><a href="#20_0">1.5. Zoogeography</a><a href="#20_0"> </a><a href="#20_0">/ Ecology/ Conservation status </a><a href="#20_0">/</a><a href="#20_0">.</a><a href="#20_0">.................................................................. 20 </a><br><a href="#20_1">1.5.1. </a><a href="#20_2">1.5.2. </a><a href="#22_0">1.5.3. </a><a href="#25_0">1.5.4. </a><a href="#25_1">1.5.5. </a><a href="#27_0">1.5.6. </a><br><a href="#20_1">Zoogeography................................................................................................................ 20 </a><a href="#20_2">Ecology/habitat ............................................................................................................. 20 </a><a href="#22_0">Field research ................................................................................................................ 22 </a><a href="#25_0">Conservation status....................................................................................................... 25 </a><a href="#25_1">Threats........................................................................................................................... 25 </a><a href="#27_0">Conservation action</a><a href="#27_0">s</a><a href="#27_0">.</a><a href="#27_0">.................................................................................................... 27 </a><br><a href="#0_0">1.6. Diet and feeding behaviou</a><a href="#0_0">r</a><a href="#0_0">.</a><a href="#0_0">....................................................................................................... 33 </a><br><a href="#0_1">1.6.1. </a><a href="#0_2">1.6.2. </a><a href="#0_0">1.6.3. </a><br><a href="#0_1">Gastro-intestinal system................................................................................................ 33 </a><a href="#0_2">Diet in the natural habita</a><a href="#0_2">t</a><a href="#0_2">.</a><a href="#0_2">............................................................................................ 33 </a><a href="#0_0">Feeding behaviou</a><a href="#0_0">r</a><a href="#0_0">.</a><a href="#0_0">........................................................................................................ 39 </a><br><a href="#0_3">1.7. Reproduction.........................................................................................................................</a><a href="#0_3"> </a><a href="#0_3">40 </a><br><a href="#0_0">1.7.1. Female reproductive characteristics ................................................................................... 41 </a><a href="#0_0">1.7.2. Male reproductive characteristics....................................................................................... 47 </a><br><a href="#0_4">1.8. Behaviour</a><a href="#0_4"> </a><a href="#0_4">.............................................................................................................................. 48 </a><br><a href="#0_5">1.8.1. </a><a href="#0_6">1.8.2. </a><a href="#0_7">1.8.3. </a><a href="#0_8">1.8.4. </a><a href="#0_9">1.8.5. </a><a href="#0_0">1.8.6. </a><br><a href="#0_5">Activity budget</a><a href="#0_5">s</a><a href="#0_5">.</a><a href="#0_5">............................................................................................................ 49 </a><a href="#0_6">Locomotion.................................................................................................................... 50 </a><a href="#0_7">Predatio</a><a href="#0_7">n</a><a href="#0_7">.</a><a href="#0_7">...................................................................................................................... 50 </a><a href="#0_8">Social behaviour ............................................................................................................ 51 </a><a href="#0_9">Sexual behaviour ........................................................................................................... 61 </a><a href="#0_0">Communication ............................................................................................................. 67 </a></p><p>5<br><a href="#0_0">Section 2: Management in zoos ............................................................................................................ 71 </a><br><a href="#0_10">2.0. Welfare of bonobos in zoos........................................................................................................ 71 </a><a href="#0_0">2.1. Enclosure .................................................................................................................................... 74 </a><br><a href="#0_11">2.1.1. Indoor enclosures................................................................................................................ 76 </a><a href="#0_12">2.1.2. Holding enclosures / quarantine enclosures....................................................................... 80 </a><a href="#0_13">2.1.3. Outdoor enclosure</a><a href="#0_13">s</a><a href="#0_13">.</a><a href="#0_13">............................................................................................................ 81 </a><a href="#0_14">2.1.4. Enclosure boundaries .......................................................................................................... 83 </a><a href="#0_0">2.1.5. Substrate ............................................................................................................................. 94 </a><a href="#0_15">2.1.6. Furnishing and maintenanc</a><a href="#0_15">e</a><a href="#0_15">.</a><a href="#0_15">.............................................................................................. 96 </a><a href="#0_0">2.1.7. Environmen</a><a href="#0_0">t</a><a href="#0_0">.</a><a href="#0_0">..................................................................................................................... 102 </a><a href="#0_16">2.1.8. Dimension</a><a href="#0_16">s</a><a href="#0_16">.</a><a href="#0_16">....................................................................................................................... 107 </a><br><a href="#0_0">2.2. Feeding ..................................................................................................................................... 108 </a><br><a href="#0_0">2.2.1. Basic diet ........................................................................................................................... 109 </a><a href="#0_17">2.2.2. Nutritional requirements .................................................................................................. 109 </a><a href="#0_0">2.2.3. Diets................................................................................................................................... 113 </a><a href="#0_0">2.2.4. Methods of feedin</a><a href="#0_0">g</a><a href="#0_0">.</a><a href="#0_0">.......................................................................................................... 117 </a><a href="#0_18">2.2.5. Water................................................................................................................................. 118 </a><br><a href="#0_0">2.3. Social Management.................................................................................................................. 119 </a><br><a href="#0_19">2.3.1. Managed fission-fusion ..................................................................................................... 120 </a><a href="#0_20">2.3.2. Transfers and introductions .............................................................................................. 123 </a><a href="#0_21">2.3.3. Sharing enclosure with other species................................................................................ 126 </a><br><a href="#0_0">2.4. Breeding ................................................................................................................................... 128 </a><br><a href="#0_12">2.4.1. Paternity diagnosis</a><a href="#0_12">............................................................................................................. 129 </a><a href="#0_22">2.4.2. Mating ............................................................................................................................... 129 </a><a href="#0_0">2.4.3. Female swelling cycle ........................................................................................................ 130 </a><a href="#0_23">2.4.4. Pregnanc</a><a href="#0_23">y</a><a href="#0_23">.</a><a href="#0_23">......................................................................................................................... 131 </a><a href="#0_24">2.4.5. Contraception possibilitie</a><a href="#0_24">s</a><a href="#0_24">.</a><a href="#0_24">............................................................................................... 133 </a><a href="#0_15">2.4.6. Birt</a><a href="#0_15">h</a><a href="#0_15">.</a><a href="#0_15">.................................................................................................................................. 136 </a><a href="#0_25">2.4.7. Development and care of young ....................................................................................... 138 </a><a href="#0_26">2.4.8. Hand-rearing...................................................................................................................... 143 </a><a href="#0_0">2.4.9. Population management................................................................................................... 152 </a><a href="#0_27">2.4.10. Sperm preservation and artificial inseminatio</a><a href="#0_27">n</a><a href="#0_27">.</a><a href="#0_27">............................................................. 152 </a><br><a href="#0_0">2.5. Behavioural enrichmen</a><a href="#0_0">t</a><a href="#0_0">.</a><a href="#0_0">.......................................................................................................... 153 </a></p><p>6<br><a href="#0_28">2.5.1. Food based enrichment..................................................................................................... 154 </a><a href="#0_29">2.5.2. Occupational enrichment.................................................................................................. 156 </a><a href="#0_30">2.5.3. Structural or Physical enrichment ..................................................................................... 158 </a><a href="#0_31">2.5.4. Sensory enrichment........................................................................................................... 159 </a><br><a href="#0_0">2.6. Handling.................................................................................................................................... 161 </a><br><a href="#0_0">2.6.1. Identification and sexing ................................................................................................... 162 </a><a href="#0_32">2.6.2. General handling ............................................................................................................... 162 </a><a href="#0_33">2.6.3. Catching/restraining.......................................................................................................... 165 </a><a href="#0_21">2.6.4. Transportation................................................................................................................... 165 </a><a href="#0_0">2.6.5. Safety................................................................................................................................. 170 </a><br><a href="#0_0">2.7. Veterinary considerations for health and welfare ................................................................... 173 </a><a href="#0_0">2.8. Specific challenges.................................................................................................................... 179 </a><a href="#0_0">2.9. Recommended research .......................................................................................................... 180 </a><br><a href="#0_0">Section 3: References / Bibliography .................................................................................................. 181 </a><a href="#0_0">Section 4: Appendices ......................................................................................................................... 253 </a><br><a href="#0_34">Appendix 1: Normal Haematological and biochemical blood values.............................................. 253 </a><a href="#0_0">Appendix 2: Bonobo Conservation Organisations........................................................................... 254 </a><a href="#0_0">Appendix 3: List of browse reported for bonobos in zoo</a><a href="#0_0">s</a><a href="#0_0">.</a><a href="#0_0">............................................................. 257 </a><a href="#0_0">Appendix 4: GREAT APE BEHAVIOURAL INFORMATION SHEET ...................................................... 259 </a><a href="#0_0">Appendix 5: Instructions for hair collection for paternity diagnosis............................................... 262 </a><a href="#0_0">Appendix 6: Swelling score sheet.................................................................................................... 263 </a><a href="#0_1">Appendix 7 : recorded pregnancy durations................................................................................... 264 </a><a href="#0_0">Appendix 8: A schedule for early infant introductio</a><a href="#0_0">n</a><a href="#0_0">.</a><a href="#0_0">.................................................................... 266 </a><a href="#0_35">Appendix 9: Sample ethogram for behavioural observations......................................................... 272 </a><a href="#0_10">Appendix 10: Design of blood sleeve as used by Columbus Zoo .................................................... 275 </a><a href="#0_0">Appendix 11: Protocol for basic post-mortem examination and sampling of the cardiovascular </a><a href="#0_0">system of great apes (EAZA)............................................................................................................ 281 </a></p><p>7</p><p>Introduction </p><p>The bonobo (<em>Pan paniscus</em>) was the last of the African ape species to be “discovered” by Western scientists. Unfortunately, nowadays this species is endangered, and as in the other species of great apes, direct threats include commercial hunting for bushmeat, that is exacerbated by habitat fragmentation through slash and burn agriculture and timber harvest. Worldwide there are 225 bonobos in managed populations, including 136 individuals in 10 EAZA zoos (December 2019). The Bonobo Eaza Ex Situ Program (EEP) population is managed in a global breeding program together with the Species Survival Program (SSP) population, which houses 95 individuals in nine institutions (December 2019). The individuals housed in </p><p>the sanctuary ‘Lola Ya Bonobo’; as well as a number of other individuals outside of the </p><p>managed population are included in the International Studbook, but are not managed by EEP or SSP. EAZA members have established Taxon Advisory Groups (TAGs) for different groups of animal species that are kept in zoos and aquariums. One of the main tasks of a TAG is to develop Regional Collection Plans that describe which species are recommended to be kept. The TAGs also identify which species need to be managed in a European breeding programme called EEP (EAZA, 2019). The mission statement of the EAZA Great Ape TAG [2018] is: </p><p><em>“The mission of the Great Ape TAG is to maintain self -sustaining and healthy populations of all </em></p><p><em>the taxa of great ape to sub specific level where possible, and to encourage and promote their conservation in the wild.The roles of zoo populations of great apes are primarily to be ambassadors for their wild counterparts for conservation-education, to support their global conservation and to be used for non-invasive research in line with the EAZA research standards. These zoo populations may also be necessary for future reinforcement of wild populations and should therefore be managed to preserve maximum genetic diversity and natural behaviour .” </em></p><p>In the regional collection plan of the EAZA Great Ape TAG [2011] the target population for bonobos, as defined by EAZA members needs/wishes (i.e. space allocated to the species) was set at approximately 200 individuals. This is more than the number that is strictly needed to maintain 90% of the current genetic diversity (which is approximately 111 individuals) for a minimum of a 100 years, but also takes into account the popularity of the species, as well as space needs for the other great ape species managed by the TAG. In addition to the Mission Statement of the EAZA Great Ape TAG, the Bonobo EEP, together with the Bonobo SSP, formulated the following mission statement for the Global Breeding Program: </p>
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