Husbandry Manual for the Spotted Hyena,

Crocuta crocuta

Jaimee Foote Monarto Zoo

November 2014

IUCN Status: Least Concern

Husbandry Manual for the Spotted Hyena,

Crocuta crocuta

Jaimee Foote

Monarto Zoo

Cover Image: Forest and cub, Spotted Hyena, Monarto Zoo, South Australia

Spotted Hyena at Monarto Zoo. © Jerome Kalvas.

Table of Contents1. Introduction ………………………………………………………………………………………………………………………………7

2. Taxonomy………………………………………………………………………………………………………………………………….7

2.1 Nomenclature…………………………………………………………………………………………………………………………..7

2.2 Other Hyena Species…………………………………………………………………………………………………………………7

2.3 Other common names………………………………………………………………………………………………………………7

3. NaturalHistory……………………………………………………………………………………………………………………………..7

3.1 Morphometrics…………………………………………………………………………………………………………………………..7

3.2 Distribution…………………………………………………………………………………………………………………………………8

3.3 Habitat……………………………………………………………………………………………………………………………………….8

3.4 Conservational Status…………………………………………………………………………………………………………………9

3.5 Wild diet…………………………………………………………………………………………………………………………………….9

3.6 Longevity........................................................................................................................................9

3.6.1 Wild……………………………………………………………………………………………………………………………………….9

3.6.2 Captivity………………………………………………………………………………………………………………………………..9

3.6.3 Techniques to determine the age of adults………………………………………………………………………….10

4. Housing.............................................................................................................................................11

4.1 Exhibit design…………………………………………………………………………………………………………………………11

4.2 Holding area design………………………………………………………………………………………………………………..11

4.3 Spatial requirements………………………………………………………………………………………………………………12

4.4 Position of enclosure………………………………………………………………………………………………………………13

4.5 Weather protection………………………………………………………………………………………………………………..13

4.6 Cubbing den and bedding materials……………………………………………………………………………………….14

4.7 Heating requirements…………………………………………………………………………………………………………….14

4.8 Substrates………………………………………………………………………………………………………………………………14

4.9 Enclosure furnishings……………………………………………………………………………………………………………..15

5. General Husbandry.......................................................................................................................18

5.1 Hygiene and cleaning…………………………………………………………………………………………………………….18

5.2 Record keeping………………………………………………………………………………………………………………………18

5.3 Methods of identification………………………………………………………………………………………………………19

5.4 Routine data collection………………………………………………………………………………………………………….19

6. Feeding Requirements..................................................................................................................20

6.1 Captive diet……………………………………………………………………………………………………………………………20

6.2 Supplements………………………………………………………………………………………………………………………….21

6.3 Presentation of food………………………………………………………………………………………………………………22

7. Handling and Transport…………………………………………………………………………………………………………..23

7.1 Timing of capture and handling……………………………………………………………………………………………..23

7.2 Capture and restraint techniques…………………………………………………………………………………………..23

7.3 Weighing and examination…………………………………………………………………………………………………….23

7.4 Release…………………………………………………………………………………………………………………………………..23

7.5 Transport requirements………………………………………………………………………………………………………….23

7.5.1 Box design……………………………………………………………………………………………………………………………23

7.5.2 Furnishings………………………………………………………………………………………………………………………….25

7.5.3 Water and food…………………………………………………………………………………………………………………..25

7.5.4 Animal per box……………………………………………………………………………………………………………………25

7.5.5 Timing of transport……………………………………………………………………………………………………………..25

7.5.6 Release from box………………………………………………………………………………………………………………..25

8. Health Requirements……………………………………………………………………………………………………………….26

8.1 Daily health checks…………………………………………………………………………………………………………………26

8.2 Detailed physical examination………………………………………………………………………………………………..26

8.3 Routine treatments and vaccinations……………………………………………………………………………………..27

8.4 Known health problems………………………………………………………………………………………………………….27

8.5 Quarantine requirements……………………………………………………………………………………………………….27

9. Behaviour ……………………………………………………………………………………………………………………………….28

9.1 Activity…………………………………………………………………………………………………………………………………..28

9.2 Social behaviour…………………………………………………………………………………………………………………….28

9.3 Reproductive behaviour…………………………………………………………………………………………………………28

9.4 Bathing…………………………………………………………………………………………………………………………………..29

9.5 Behavioural problems…………………………………………………………………………………………………………….29

9.6 Signs of stress…………………………………………………………………………………………………………………………29

9.7 Behavioural enrichment………………………………………………………………………………………………………….29

9.8 Operant conditioning……………………………………………………………………………………………………………..38

9.9 Introductions and removals……………………………………………………………………………………………………38

9.10 Interspecific compatibility……………………………………………………………………………………………………39

9.11 Intraspecific compatibility……………………………………………………………………………………………………39

10. Breeding………………………………………………………………………………………………………………………………..40

10.1 Mating system……………………………………………………………………………………………………………………..40

10.2 Ease of breeding…………………………………………………………………………………………………………………..40

10.3 Reproductive condition………………………………………………………………………………………………………..41

10.3.1 Females…………………………………………………………………………………………………………………………….41

10.3.2 Males………………………………………………………………………………………………………………………………..41

10.4 Techniques used to control breeding……………………………………………………………………………………41

10.5 Occurrence of hybrid…………………………………………………………………………………………………………...41

10.6 Timing of breeding……………………………………………………………………………………………………………….41

10.7 Age of first breeding and last breeding…………………………………………………………………………………42

10.8 Ability to breed every year…………………………………………………………………………………………………..42

10.9 Ability to breed more than once per year…………………………………………………………………………….42

10.10 Denning requirements……………………………………………………………………………………………………….42

10.11 Breeding diet……………………………………………………………………………………………………………………..42

10.12 Oestrus cycle and gestation period…………………………………………………………………………………….42

10.13 Litter size……………………………………………………………………………………………………………………………42

10.14 Age at weaning………………………………………………………………………………………………………………….43

10.15 Removal from parents……………………………………………………………………………………………………….43

10.16 Growth and development………………………………………………………………………………………………….44

11. Artificial Rearing……………………………………………………………………………………………………………………44

11.1 Housing………………………………………………………………………………………………………………………………..44

11.2 Temperature requirements………………………………………………………………………………………………….45

11.3 Diet and feeding routine………………………………………………………………………………………………………45

11.4 Specific requirements………………………………………………………………………………………………………….47

11.5 Identification methods…………………………………………………………………………………………………………48

11.6 Hygiene………………………………………………………………………………………………………………………………..48

11.7 Behavioural considerations………………………………………………………………………………………………….48

11.8 Use of a foster species………………………………………………………………………………………………………….49

11.9 Weaning………………………………………………………………………………………………………………………………49

12. Acknowledgements……………………………………………………………………………………………………………….50

13. References……………………………………………………………………………………………………………………………..51

15. Appendix……………………………………………………………………………………………………………………………….53

1. Introduction

The Spotted Hyena, Crocuta crocuta, are the largest of all hyena species. According to the IUCN Threatened Species List they are: Least Concern as the species remain widespread in Africa (IUCN Red List 2008).They are a heavy set dog like carnivorous mammal whose active prey are large ungulates. They are distinguished by their canny cackle and features of spotted brown short and rough fur, which has a woolly undercoat, and dark brown fur which covers their face and legs (Brottman 2012). The forequarters are large with a massive neck, the back is sloping, and the hindquarters are rather weakly developed. The forelegs and hind legs have four digits and their claws are not retractable (WAZA) .Their massive jaws and teeth are usually capable of cracking large bones and ribs (Husbandry Standards Hyena). They are a social species who live in large clans mainly confined to sub-Saharan Africa (Planet wildlife 2010). The Spotted Hyena are one of the most misunderstood animals, and are often mislabelled as cowardly scavengers. They are in fact a highly intelligent species and are both successful hunters and opportunists.

‘Zoologist Hans Kruuk, author of The Spotted Hyena (1972), believes ‘there is a magic about hyenas’, and those who spend time with them cannot help but ‘fall for the spell’ of these animals that are ‘so totally different’. ‘Get to know a hyena up close, and you will encounter a fabulous, enchanted beast, neither cat nor dog, with a touch of ferret thrown in for bad luck. There is a sublime and hideous grandeur to this big-eared, red-eyed, corpse-eating creature that has rarely been recognised or acknowledged’ (Brottman 2012)

2. Taxonomy2.1 NomenclatureClass: MammaliaOrder: CarnivoraFamily: HyaenidaeGenus Species: Crocuta crocuta

2.2 Other Hyena SpeciesThe Striped Hyena – Hyaena hyaenaThe Brown Hyena – Hyaena brunneaAardwolf – Proteles cristata

2.3 Other common namesSometimes referred to as the Laughing Hyena (Alongi 2011-2014)

3. Natural History3.1 MorphometricsHead and body size: 120-150cm (WAZA)

Shoulder height: 75-85cm (WAZA)

Tail: 25-35 cm (National Geographic 1998-2014)

Weight: ranges in males from 45-60kg; and females weigh between 55-70kg. The matriarchal female is generally 10% heavier than males (WAZA).

Weight at birth: average 1.4kg (San Diego Zoo 2014)

3.2 DistributionThere have been confirmed recordings of Spotted Hyena from Tanzania, Kenya, Zambia, Zimbabwe, Mozambique, South Africa, Botswana, Namibia, Congo, Sudan, Central African Republic, Cameroon, Nigeria, Burkina Faso, Ivory Coast, Mali, Senegal, and Sierra Leone. Although Spotted Hyena occur throughout sub-Saharan Africa, high densities occur in the Serengeti ecosystem, including Ngorongoro Crater in Tanzania, and in the Aberdare Mountains of Kenya (IUCN Hyena Specialist Group 1997-2014).

http://

designeranimals2011.wikispaces.com/Hyena-Vivian+LinFigure 1: Distribution of the Spotted Hyena

3.3 HabitatThe Spotted Hyena occurs everywhere in Africa south of the Sahara except the Congo basin; they are also seen eastwards through Arabia to India (National Geographic (1996-2014). They are found in all habitats from Savannahs, grasslands, woodlands, forest edges, sub deserts and mountains up to 4,000 metres (African Wildlife Foundation), from extremely hot and arid in the north and south to the cold heights of dense mist Forest Mountains in east Africa and Ethiopia. Their greatest densities occur in fairly flat open country with few or no trees and short grass. Their optimal habitat occurs at Ngorongoro Crater and Serengeti National Park (Kruuk 1972).

3.4 Conservational StatusAccording to the IUCN Redlist (Honer, Holekamp, K.E. & Mills 2008) the Spotted Hyena is currently listed as Least Concern as the species remains widespread in Africa and the total world population well exceeds 10,000 mature individuals. There is a continued decline to their population outside and within protected areas mainly due to human destruction and habitat loss. These will be explored briefly below.

Persecution: Outside protected areas the Spotted Hyena are subject to human destruction through shooting, trapping and poisoning (IUCN Redlist). The adult mortality rate is very high due to wire snares which kill around 400 adult Spotted Hyena each year. Only since the mid 1970’s has game meat hunting rapidly expanded (Hofer 1998). Numbers killed during sport hunting are low as they are not considered an attractive species but are also killed for food and medicine (Hofer and Mill 1998b).

Habitat Loss: A threat is posed by the decline in densities of Spotted Hyenas due to habitat loss caused by increased human settlement and overgrazing by livestock (IUCN Redlist).

3.5 Wild DietSpotted Hyena have long been misinterpreted as cowardly scavengers, where in fact they are among the most abundant and successful carnivores in sub-Saharan Africa with a better kill rate than Lions (Brottman 2012).

They usually hunt in clans of 10-15 animals lead by the alpha female. They will usually prey on gazelle, wildebeest, water buffalo or zebra and sometimes young giraffe, rhino and hippo. Spotted Hyena have also been known to prey on farm animals including horse, donkey, camels, poultry (even cats and dogs) and the prey is known to be eaten alive. However they will sometimes scavenge from human camps, dumps and landfill in South Africa.

When hunting alone they tend to prey on smaller animals like birds, hares, foxes, jackals, fish, snakes and carrion. To supplement their diet they will often eat fruit, eggs and invertebrates. They are known to consume around 3-6kg of meat per day. With their powerful jaws they can crush through bone and feed on the marrow inside. Their stomach contains concentrated hydrochloric acid which allows them to digest the bones they eat (The Animal Files). There are some items that hyenas can’t digest, such as hair and horns, although they still eat them. They regurgitate what they can’t digest in the form of a pellet (San Diego Zoo 2014)

3.6 Longevity

3.6.1 WildLifespan is approximately 20 years in the wild (Law J 2004).

3.6.2 CaptivityOn average 25 years in captivity (National Geographic 1996-2104). One has been recorded living to 41 years of age (Jones, 1982).

3.6.3 Techniques to Determine the Age of Adults Reliable age estimation of Spotted Hyena has proven difficult (Kruuk 1972). Techniques that have been used include distinguishing between five tooth wear classes using the third mandibular premolar and generalised these to absolute age in years. (Figure 2 the amount of wear indicates age class). (Figure 3 indicates the relation between spot pattern and age of tooth wear).

The Spotted Hyena – A study of predation and social behaviour (Hans Kruuk 1972) page 33Figure 2: Third premolar in the lower jaw of a spotted hyena. The amount of wear indicates age class.

The Spotted Hyena - A study of predation and social behaviour (Hans Kruuk 1972) page 210Figure 3: The relation between spot pattern and age of tooth wear.

4. Housing Requirements

4.1 Exhibit Design

Spotted Hyena exhibits should consist of the following design elements (Foote unpublished data 2014, Shoemaker 2006 and Bennett, pers. comm. August 2014).

A secure boundary with electric fencing and an underground mesh skirt. Weld mesh and or cyclone mesh could be used.

Enclosure should be designed in a way that mimics their natural habitat The size of outdoor enclosures should be large enough to prevent destruction of grasses and

plantings Must provide a good view of their surroundings, the ability to see advantage points, enough

space to flee from other clan members if needed and no corners, instead more open spaces Due to their social dominance the enclosure should have multiple entrances, escape routes and

exits to accommodate aggression To avoid stereotypical behaviour enclosures should be irregular in shape and without corners to

prevent aggressive individuals from cornering subordinate individuals Enclosures should be provisioned with objects to enhance their marking behaviours and change

the route of their movements Exhibit must have multiple dens including dens for cubbing Enclosure must display a bathing pond so the hyena can display natural bathingbehaviours Animals must have access to fresh drinking water Animals must always have access to shade and cool areas in hot weather Animals must have access to shelter and warm areas in cold weather Trees, stumps, deadfalls and rocks placed irregularly throughout the exhibit will greatly enhance

their daily routine and provide visual barriers for lower ranking individuals Enclosures must contain objects 5-30 centimetres above the substrates for the animals to paste

markings from their anal glands Instillation and display points for environmental enrichment Enclosure must be safely accessible to keeper, also provide means of securing the animals in a

holding area Enclosure should have vehicle access for maintenance work or heavy machinery if space permits Enclosure must allow for the animals to escape out of public view and remove themselves from

the clan if needed

The design of exhibits must provide animals with the “Five Freedoms” of animal welfare as per the Animal Welfare Act (1999). These are:

1. Freedom from hunger and thirst2. Freedom from discomfort3. Freedom from pain, injury or disease.4. Freedom to express normal behaviour5. Freedom from fear or distress.

4.2 Holding area design

Holding areas in which animals are kept for long periods of time should allow for enough space to accommodate the social and behavioural needs of the Spotted Hyena. Monarto Zoo has an off display holding area which consists of four separate lock-away areas, each approximated 220 m². This is a minimum guideline that could be followed (Foote, unpublished 2014, Washford, pers. comm. November

2014 and Brooks, pers. comm. November 2014). Spotted Hyena holding facility should consist of the following design elements:

Holding area must be safe and secure, with the use of weld mesh and buried mesh around exhibit perimeter with varied substrates

Ability to safely secure hyena off exhibit Ability to safely separate individuals away from each other with the ability of mesh contact so

aggression is avoided when re introduced Ability for a transport box to be incorporated into the design or in the area for transport and

medical procedures Designated area where animals can be safely trained and conditioned by keepers A crush system designed to habituate hyena by permitting daily movement from one space to

another through the crush area Holding should have multiple entrances, escape routes and exists to accommodate aggression Holding areas to mimic their natural habitat with appropriate substrates and furnishings Provide fresh drinking water and places for feeding Provide bathing ponds Access to multiple dens including dens for cubbing Controlled environment such as shelter, shade and any necessary heating Ability for enrichment to be installed and displayedHolding areas must also provide animals with the “Five Freedoms” of animal welfare as previously mentioned in Section 4.1.

4.3 Spatial Requirements

In general, outdoor exhibits will display Spotted Hyena best, and enclosures suitable for small to medium sized bears or large felids will meet most of their behavioural and spatial needs. Public observation seems to cause stress for single animals and more room such as (20.9m²) may be necessary if solitary animals are on display (Shoemaker, Dulaney,Noble 2006). The spatial requirements shown in table 1 and 2 are only a minimum standard that are set for bears and lions and must be achieved as per the legislation in New South Wales, Australia. These requirements are not Spotted Hyena specific but can be a minimum guideline.

Name Surface area (SQ-M)

Max no animals for enclosure size

Additionarea for each extra animal in exhibit (m2)

Addition area for each extra animal in medium term holding yard (m2)

Height above water of wall adjoin moat (m)

Moat width (m)

Moat depth (m)

Smooth faced unclimbable wall (m)

Mesh fence height (m)

Wire mesh dia (mm)

Mesh spac (mm)

Depth of Inhang (m)

Inhang (Degrees above horiz)

FAMILY UrsidaeAmerican Black BearUrsus americanus

300 (30) 2 20 15 2.0 3.0 4.0 - 4.0 5 75x50 1.0 45

FAMILYFelidaeLionPanthera leo

300 (30) 2 20 15 1.8 8 1.8 - 4.5 5 75x50 1.0 45

Table 1: Highlights the carnivore biology and exhibit guide as per the legislation in New South Wales. This information can also be a guideline for the spotted hyena.

Species Social Peak Level Mesh Roof Wire Solid Piano wire Glass front Moat Water

group activity o/hang needed

needed under substrate

walls advised

permissible permissible feature advised

FAMILYUrisidaeBlack BearUrsus arctos sytacus

S D/C T/A Yes - S - - Yes Wet/dry Yes

FAMILYFelidaeLIONPanthera Leo

G C/N T Yes - S - - Yes - -

Table 2: Biology and exhibit guide for Black Bear and lion (Department of Primary Industries).

Monarto Zoo, South Australia has had experience in housing Spotted Hyena in many different scenarios. Allowing the most space possible is recommended due to the social dynamics of a clan. If only a small exhibit is available, housing a minimum number of animals is recommended to reduce any behavioural problems. The following dimension used at Monarto Zoo can be followed as a guide. Fence vertically 2.7 metres high, top fence 700 millimetres of overhang at approximately 45 degrees and a horizontal skirt around the entire perimeter of the exhibit, buried 30 centimetres under and going 1 metre inside the exhibit (Bennett, pers. comm. September 2014 and Brooks, pers. comm. August 2014).

Table 3: Minimum guide for housing Spotted Hyena (Bennett, pers. comm. September 2014).

Perth Zoo, Western Australia, house 2.0 Spotted Hyena. Their night quarter area is 149.57 m² and the exhibit area is 600.07 m². They have two public viewing areas which has glass panels and below that inside the exhibit a 1 metre wide x 5 metres long electric fence. The exhibit walls are partially steel mesh and the rest is concrete approximately 3.5-4 metres high (Thomasson, pers. comm. October 2014).

4.4 Position of EnclosuresSpotted Hyenas being large active carnivores are best exhibited outdoors in enclosures that mimic their natural habitat (Shoemaker, Dulaney, Noble 2006). The exhibit should be positioned in a way where they have a good visual of their surroundings, the ability to see advantage points, enough space to flee from other clan members if needed, no corners, instead more open spaces, many den options and the ability to hide and remove themselves from the public and other clan members. Spotted Hyenas seem to live fine alongside other carnivores. Monarto Zoo, South Australia houses their Spotted Hyena and African Painted Dogs alongside each other which is great social and mental stimulation and no issues have been seen. There is also the option for the two species to swap exhibits and cross paths (Bennett, pers. comm. August 2014).

4.5 Weather Protection

Number of Spotted Hyena Exhibit size

2 -3 4000 m²3 or more 6000-8000 m² ( biggest space possible)

Sufficient shelter from weather conditions, such as wind, rain, and extremes in temperature must be provided. If shelters on exhibit are not available, animals must be provided with access to their holding areas to seek protection (Foote, unpublished data 2014).

4.6 Cubbing Den and Bedding Materials

A den or cave should be provided for sleeping and cubbing needs. Hyenas do fine on hard substrates (Shoemaker, Dulaney, Noble 2006). Monarto Zoo, South Australia have found that a smaller den attached to the large den works well as the cub can feel protected in a smaller confined area and venture out to the bigger den when ready; this mimics a wild situation. The small den also allows no adults to enter, as it only has a 15cm x 15cm opening, making it a safe and protected space. Peep holes are located on the small and large den for staff to look in and monitor if needed. Dirt has only been used as den bedding material, to avoid unnecessary tampering/cleaning of maternal den areas. (Foote, unpublished data 2014, Bennett, pers. comm. September 2014).

Figure 4: Large and small Spotted Hyena den at Monarto Zoo, South Australia (Foote 2014).

4.7 Heating Requirements

Spotted Hyenas encounter climates of hot, over 40°C and cool to 0°C during some parts of the year. As a result, most individuals are tolerant of a wide range of daytime temperatures. Regardless, animals kept outside should always have access to shade, especially during warmer parts of the year. When acclimatised, most species without young require only minimal shelter at night if temperatures are only expected to reach 0°C, although heated shelters or indoor enclosures are needed for animals kept in cooler climates. For exhibit purposes, basking areas will provide a great deal of warmth in cooler weather, and the use of 'hot rocks' will significantly increase their exhibit use in winter (Shoemaker, Dulaney, Noble 2006).

4.8 Substrate

Large den

Small den attached to large den, arrow highlights peep hole

Peep hole in large den. Large den has 3 peep holes in total

Outdoor enclosures employing natural soils are best. Turf and natural grasses, as well as bushes, should be planted and left to grow. In areas of high humidity or rainfall, mixes of sand and pea gravel also do well to keep skin problems such as dry skin to a minimum. Hyenas being active diggers will dig shallow depressions and holes. Wire mesh buried below substrates will still encourage digging but prevent animals from escaping. (Shoemaker, Dulaney, Noble 2006)

4.9 Enclosure Furnishings

Furnishings must accommodate an array of locomotory and foraging behaviours as well as resting and sleeping points. Their enclosure should be irregular in shape and enhanced with rocks, tree stumps, bushes, fallen logs and branches to change the pattern of their movements to increase their level of daily investigation and scent marking. Structures could be moved daily, weekly or periodically for best stimulation. Hyenas are strong-jawed animals with a tendency for chewing. All objects in their enclosure must be able to withstand their destructive tendencies or be easily replaced. Rubber tyres are inappropriate as furnishings as they may be chewed into pieces and swallowed (Shoemaker, Dulaney, Noble 2006).

http://

www.panoramio.com/photo/98745420Figure 5: Enclosure structures in the Spotted Hyena exhibit at Colchester Zoo

http://www.zoochat.com/57/striped-hyena-spotted-hyena-enclosure-colchester-64972/Figure 6: Enclosure structures in the Spotted Hyena/Stripped Hyena exhibit at Colchester Zoo

http://www.zoochat.com/581/spotted-hyena-exhibit-56811/Figure 7: Enclosure structures in the Spotted Hyena/Stripped Hyena exhibit at Miami Zoo

5. General Husbandry

5.1 Hygiene and cleaning

Hyenas are very scent oriented and obsessive cleaning/disinfecting can cause stress and reluctance to shift or enter holding or exhibit areas. Logs and play objects as well as food containers should be kept clean but enough scent left on them for the animals to recognise them. Dirt and grass substrates in outdoor exhibits should be spot-cleaned daily. (Shoemaker, Dulaney, Noble 2006). Water bowls should be cleaned daily so a constant supply of fresh water is available, including water troughs as they use these for bathing (Foote, unpublished data 2014).

5.2 Record Keeping

Records must be kept current and up to date of all Spotted Hyena and contain the following information (Foote, unpublished data 2014 and ZIMS)

Common name Scientific name Individuals name Distinct markings or behaviour Veterinary records and examination with advice and treatment provided Sex/contraception status Rearing type Dam/Sire Birth type Birth location Birth date Transponder or identification method Changes in behaviour Physical changes Breeding information Health and behaviour problems Training information Dietary changes Anything significant Date of death

5.3 Methods of Identification

Individual sex can be hard to determine by sight as the female has a cliteriform appendage pseudo-penis (Brottman 2012) which looks very similar to the male’s genitalia. The matriarchal female is generally 10% heavier than males (WAZA). As the Spotted Hyena ages they also become lighter in colour and their spots fade. Ear notch identification has been used in the wild to identify individual animals (Bennett, pers. comm. August 2014).

http://www.hyaenidae.org/hyena-myths.html

Figure 8: Highlights identification differences between male and female Spotted Hyena

5.4 Routine Data Collection

Those items mentioned in section 5.2 should all be checked regularly to ensure information is accurate and up to date. Daily records should be taken and stored safely especially for future reference and guidance (Foote, unpublished data 2014).

6. Feeding requirements

6.1 Captive Diet

In the wild, hyenas are typically gorge feeders and feed only once or twice a day. Due to problems with obesity, hyenas that are not pregnant or lactating are recommended to have one or two starve days a week. Hyenas are highly adapted to consuming bones, and these should be included in their diet several times a week (Berger & Frank 1992). The Spotted Hyena will often hide food in water including dams and shallow pools for short periods so providing appropriate places for cache food is essential (Wildlifesouthafrica). Spotted hyenas are naturally aggressive feeders so individually separating for feeding is essential in captivity to ensure both dominate and subordinate animals have adequate feeding time (Shoemaker, Dulaney, Noble 2006). A variety of different meat should be offered as part of their diet including bone and whole animal carcasses and only one main feed is required per feed day (Foote, unpublished date 2014).

At Denver Zoo, they feed their spotted hyena 5 times a week, the other two days only a knuckle bone or carcass feed is given. If the carcass is larger than 1.5-2kg an extra fast day is given.

Meat Bone Carcass feed 1.5-2kg

1.1 kg Milliken brand canine diet, once a day 5 times a week

One day a week they only receive a knuckle bone

One day a week they only receive either a whole:

Rabbit Guinea pig Large rat Lamb/beef shanks Goat

Table 4: Captive diet for the 1.1 Spotted Hyena at Denver Zoo.

Fruit/Vegetables Meats Items not allowed

Food Enrichment Items Melons Brussel sprouts Green beans

Mice Quail Fish Liver Heart Dog biscuits Dog and cat

chow Pig ears

Onion Garlic Citrus

Table 5: Food enrichment items offered to the Spotted Hyena at Denver Zoo.

At Monarto Zoo South Australia, they feed their Spotted Hyena 5 times a week with two fast days on Monday and Thursday. The meat portion is given with bone and hide. Each animal receives a certain amount depending on their life stage. Beef and horse makes up majority of their diet but kangaroo and turkey is also offered. All meat is frozen for 7 days prior to feeding to eliminate any potential infection from pathogens. A supplement is also added to their daily meat portion.

Animal Amount Beef/Horse Big Cat Supplement(10g per kg of meat)

Kigali 1.7kg 17gForest 2.5kg (Currently lactating) 25g

Piltengi 1.7 kg 17gGamba 2kg 20gPinduli 2kg 20gZuberi 2kg 20g

Table 6: Captive diet for the 4.2 Spotted Hyena at Monarto Zoo.

Fruit/Vegetables Meat Eggs

Food enrichment Items

Melons Pumpkins Apples

Whitebait/pilchards Tin sardines Whiskas cat food Anchovy paste Dog biscuits Ice blocks with blood

and meat pieces Blood trails Mince Bones Blood jelly

Whole ostrich egg

Boiled or raw chicken egg

Table 7: Food enrichment items offered to the Spotted Hyena at Monarto Zoo.

6.2 Supplements

Monarto Zoo, South Australia adds a big cat supplement to their hyenas meat diet. This supplement is locally made in Adelaide, South Australia by Wombaroo (Passwell). To supplement an all meat diet 10g (1 level tablespoon) is needed per kilogram of meat. Best added with water to make a paste and applied by slicing open a portion of meat and inserting the supplement into the cut (Foote, unpublished data 2014).

NUTRIENT COMPOSITION PER kg COMPOSITION PER MJ OF

PREPARED MEAT

Retinol 34mg 144gCholecalciferol 750g 1.05gd-a-tocopherol 5300mg 7.8mgPhytomenadione 4.3mg 8mgPantothenic Acid 170mg 1.2mgPyridoxine 140mg 0.8mgRiboflavin 110mg 0.4mgThiamine 170mg 0.4mgFolic Acid 36mg 60gBiotin 860g 4.2gCyanocobalamin 690g 6.0gCalcium 300g 440mgPhosphorus 32g 360mgMagnesium 3.6g 35mgIron (Fe++) 2100mg 6.0mgZinc 360mg 4.5mgManganese 400mg 600gCopper 120mg 300gIodine 36mg 90gLinoleic acid 1800mg 230mglinolenic acid 5000mg 85mgArachidonic acid 28mg 110mgEicosapentaenoic acid 690mg 46mgDocosahexaenoic acid 370mg 16mgTaurine 38g 53mgTotal Carotenoids 3.6mg 5.0g

Table 8: Ingredients list and nutritional quantities of the Big Cat Supplement (Wombaroo Passwell).

6.3 Presentation of food

When food items are hidden throughout their exhibits, enough items placed at widely placed locations are necessary to reduce aggression while providing adequate time for both dominate and subordinate animals to find food. In situations where more than one animal must be housed within the same “feeding” enclosure, at least two feeding areas should be used to reduce aggression.

Hyenas naturally eat as quickly as possible, bolting their food in order to reduce the likelihood that it will be stolen. If circumstances permit, whole or partial carcasses can be offered to simulate natural feeding and to enhance educational value of the exhibit. Escape routes must be present for subordinate individuals to escape aggression during feeding. (Shoemaker, Dulaney, Noble 2006).

Monarto Zoo, South Australia individually separates each clan member for a brief time every feed day for feeding; this allows each member to eat their correct meat portion. Their meat portion is given through a protected feed chute and consists of meat, bone and hide. Food type enrichment is regularly provided to the clan which is offered periodically and either hidden, spread throughout the exhibit to stimulate activity or provided at keeper presentations for public interactions. The food may be fresh or frozen and presented by spiking, scattering, burying and smearing. Monarto also houses a single male hyena which allows keepers to present his food in many different ways as he has no competition for food and requires the stimulation. Food presentation includes hiding small meat chunks throughout the exhibit, hanging a kangaroo tail or leg by chain, giving a whole ostrich egg to crack open and placing apples stuffed with cat food in ponds for fetching. (Foote, unpublished data 2014).

7. Handling and Transport

7.1 Timing of Capture and Handling

The time of capture will depend on the situation, destination and means of transport. Weather conditions must be considered so the animal is not being transported through the heat of the day and a transfer is best avoided in the extreme hot months (Bennett, pers. comm. September 2014).

7.2 Capture and Restraint Techniques

Spotted Hyena can be successfully conditioned to walk into a crate for transportation. This is a successful technique which can easily be achieved to avoid stress on the animal. Two or more weeks prior to transport the animal should be conditioned to the crate to make sure it is relaxed and settled prior to departure (Bennett, pers. comm. September 2014).

7.3 Weighing and Examination

Spotted Hyena can be conditioned to step onto scales and be regularly weighed. If conducting a detailed physical examination prior to transportation please refer to section 8.2 (Foote, unpublished data 2014).

7.4 Release

They should be released into a smaller area that must be quiet and safe for the animal with no dangerous obstacles as they may be frightened and stressed on release. Only once the animal is calm with its new surroundings should they be given full access to their exhibit, this process must be monitored (Foote, unpublished data 2014).

7.5 Transport RequirementsHyenas can be safely shipped by air without staff accompanying them. Hyenas normally transport easily, and few problems are likely to be encountered. Overland transportation requiring several days is acceptable although arrangements for feeding will be necessary (Shoemaker, Dulaney, Noble 2006).

7.5.1 Box Design

The International Air Transport Association’s Live Animals Regulations (2001) recommendations for the transportation of hyena species are as follows:

Materials:The box should be constructed with wood external, metal lined and with weld mesh doors or solid metal doors.

Dimensions:The height of the container must allow the animal to stand in a natural position with its head extended and the width must permit it to turn around and lie down comfortably.

The frame:The frame must be made from solid wood or metal parts bolted or screwed together. It must be constructed so that it cannot be damaged from continual biting or scratching it must be metal lined. If the total weight of the container plus animal exceeds 60kg metal bracing must be added to the frame.

Sides:The sides and doors must be made of metal or solid wood with metal lining. The front of the container must be constructed of weld mesh. The mesh must have a diameter that will prevent the animal protruding its nose or paws outside. The whole front must be covered by a sliding shutter which can be raised and lowered to permit feeding and watering. It must have two observation holes of at least 10cm in the upper part and ventilation holes, with a minimum diameter of 2.5cm spread over the remainder of the surface in order to give good ventilation but at the same time leave the animal in semi-darkness.

Floor and roof:The floor must be slatted, over a leak-proof droppings tray and the roof must be solid wood or metal with ventilation openings over its surface.

Doors:A sliding door must be provided which can be made from the weld meshed ventilation front if required. It must have a secure means of fastening so that it cannot be opened accidently.

Ventilation:The main ventilation front must be supplemented by meshed openings along the upper part of the container walls and/or holes with a minimum diameter of 2.5cm over the top third of the sides and the whole of the back and top. These holes must be spaced both horizontally and vertically at intervals of approximately 10cm centre to centre. It is essential some ventilation is provided in the lower third of the sides for the removal of harmful gases. Total ventilation must be at least 20% of the total area of the surface of all four sides.

Space bars/handles:Must be made to a depth of 2.5 cm to allow for fingers to fit in handles and must be present on the sides of the container.

Forklift spacers:Forklift spacers must be provided if the total weight of the container plus the animal exceeds 60kg.

Special Requirements:The container must be completely lined with sheet iron or other hard metal sheeting and through ventilation holes cut into it. Palletised shipments must have containers made entirely of weld mesh of a suitable dimension that no part of the animal can protrude in order to ensure good ventilation.

Figure 9: IATA box design suitable for transporting hyena species (IATA 2001).

7.5.2 Furnishings

Straw or wood shavings should be included in the container for increased comfort by the animal during transport as well as absorption of urine. (Shoemaker, Dulaney, Noble 2006). This will be dependent on quarantine regulations for destinations (Bennett, pers. comm. October 2014).

7.5.3 Water and Food

Spotted Hyena do not normally require additional feeding or watering during transport for periods of up to 24 hours. Assess weather conditions for transportation and if required a metal water bowl can be attached to the corner of the crate but it must be secure as they may well destroy it (Shoemaker, Dulaney, Noble 2006), (Kalvas, pers. comm. August 2014).

7.5.4 Animals per Box

Spotted Hyena of all ages must be shipped in individual containers (Shoemaker,Dulaney,Noble 2006).

7.5.5 Timing of Transport

Please refer to section 7.1.

7.5.6 Release from Box

Must be released into a restricted area first and monitored for health and response to new environment. If multiple animals have arrived, each animal must be released into individual areas. Once individual health has been monitored, mesh introductions must occur and once positive behaviours are seen from the separated animals, they can then be reintroduced (Bennett, pers. comm. September 2014).

8. Health Requirements

8.1 Daily Health Checks

With any captive animal, a thorough health check should be carried out first thing in the morning, with a number of checks throughout the day, including a final check before leaving at the end of the day. Abnormal signs that should be looked for which could indicate poor health or injury include (Foote, unpublished data 2014):

Lameness Unable to move around freely such as running and climbing Any gain or loss of appetite or lack of motivation towards food Eyes clear and fully open General appearance of body, coat and skin condition, any fur loss Any obvious wounds or scratches Discharges from the eyes, nose or mouth or heavily panting Changes in individual behaviour Level of activity – knowing the animals normal behaviour will give good indication Any change in faecal or urine colour or consistency or sign of vomit. Any abnormalities or difficulties with defecating and urinating Changes in genital appearance Any signs of blood They are being harassed by other clan members

The progression of a training plan can help assist keepers in daily health checks of the animal. The Spotted Hyena are proven to be very intelligent species who respond well to training. Hyenas have successfully participated in training and conditioning which has helped with veterinary procedures including blood draw and injections (Foote, unpublished data 2014).

8.2 Detailed Physical Examination

Due to the nature of the Spotted Hyena, a detailed physical examination can only be carried out if the animal is under anaesthetic. The following list highlights the various aspects of the Spotted Hyena that should be checked or collected during a routine physical examination (Kalvas, pers. comm. August 2014).

Eyes Ears Dental examination Body condition score Weight Blood – Serum/EDTA Faeces Urine Abdominal palpation X-rays if required Microchip if required

Any routine vaccinations or antibiotics if required Further examination of concern areas if any issues arise during the procedure

Spotted Hyena normal parameters that must be checked and maintained during the physical examination:

Heart rate: 40-60 bpm Respiration rate: 20-40 Temperature: 38 c ̊ Packed Cell Volume : 26-57 Total Protein : 54-86

8.3 Routine Treatments and Vaccinations

The Spotted Hyena housed at Monarto Zoo South Australia receive the following routine treatments and vaccinations (Kalvas, pers. comm. August 2014).

F3 vaccinations at 8 weeks and 12 weeks and then biannually Faecal float every 6 months, worm as required

8.4 Known Health Problems

Historically, Distemper is known as the main concern with no reports seen with Spotted Hyena in Australia. Gastrointestinal worms including hook, whip and round worm have been reported including external parasites such as fleas and mites. Due to some hierarchical aggression within a clan wounds are readily observed but due to their well-developed immune system wounds seem to heal well with no real issues. Spotted Hyenas are known as quite robust animal with no real known health issues. (Kalvas, pers. comm. August 2014).

8.5 Quarantine Requirements

Quarantine requirements differ depending on whether the animal arrives interstate or international and will depend on individual institution requirements or DAFF requirements for international acquisitions. (Kalvas and Bennett, pers. comm. August 2014).

International/ Domestic quarantine requirements include:

International arrivals require a long term minimum holding of up to 90 days Domestic arrivals require a minimum holding of up to 30 days Obtaining previous institution medical records on the animal is desired Animals must be housed completely separate from any other animals especially other hyena Keepers must follow quarantine procedures such as footbath, protective clothing and attending

to the quarantine animals last in the day after all other animals have been seen 3 faecal exams are taken after minimum holding time Followed by a physical examination under anaesthetic conducting a general health check

including x rays and bloods Animals can only be removed from quarantine after the 90/30 days if all results are clear

9. Behaviour

9.1 Activity

The Spotted Hyena is predominately nocturnal and most active during the hours of darkness due to their foraging strategy as they frequently feed on large food items and rest thereafter. Their day is divided into active and resting periods which include variation of feeding, social, territorial and sexual behaviours. Observation of a single female Spotted Hyena showed she was only active 15.8% of her day with 84.2% laying down, but this estimate is also quite low. Main activity is seen hunting, feeding and foraging and they have been known to travel on average 10.1km per day. In the wild kills have been witnesses to happen at the same time of day in sequence then change fairly suddenly to a different time of day. During their resting period they choose many places which depend on individuals, clans, areas and seasons. Female with cubs will spend all day resting around the den generally in a cool, shady place usually alone. Others generally go down an unused hole, lie at the edge of a lake or stream, in dense shrub, bushy vegetation or in shade high on sloping land and during the colder months will sleep out in the open plains. (Hans 1972)

9.2 Social Behaviour

The Spotted Hyena is one the most highly social of all carnivores; it lives in groups containing up to 90 individuals, and exhibits very complex social behaviours. These animals live in social groups called clans that defend group territories. The society is characterised by a strict matriarchal dominance hierarchy. Females are dominant over males, and even the lowest ranking female is dominant to the highest ranking male (IUCN 2007-2014). Males typically disperse from their natal clans when they are between two and six years of age. Females usually remain in their natal clan, so large clans may contain several different matrilines. Although cubs of both sexes 'inherit' their mothers' social ranks, males voluntarily forsake those to assume much lower ranks in the neighbouring clans to which they disperse. (Smale et al 1997; Holekamp & Smale 1998).

Spotted Hyena clan members frequently wander alone or in small groups and only sometimes meet in large numbers. This occurs at kills, at the communal den, or when clan members rally together to defend individual carcasses or group territories (Kruuk 1972)

Territory size in the Spotted Hyena is highly variable, ranging from less than 40 km2 in the Ngorongoro Crater to over 1,000 km2 in the Kalahari (Mills 1990). Clans defend communal territories through vocal displays, scent marking and boundary patrols. Clan members also cooperate in defending territories during boundary disputes with neighboring clans. Long distance calls, particularly whoops, are used to quickly rally clan members to such sites of conflict. Spotted Hyenas scent mark their territories by pasting a secretion from the interdigital gland onto grass stalks, and by depositing a secretion from anal glands when they scratch the ground. Spotted Hyenas also scent mark their territories by defecating in communal latrines (Kruuk 1972a. Mills 1990).

9.3 Reproductive Behaviour

Males perform a bowing display to females before mating. The male lowers his muzzle to the ground, advances quickly toward the female, bows again, and then paws the ground close behind the female. The dominance of females assures that males are timid and will retreat immediately if the female shows any aggression. The female's reproductive tract makes mating somewhat difficult and requires full cooperation from the female. Male hyenas approach and slide their haunches under the female to achieve intromission. Once intromission is achieved they move to a more typical mating posture, with

the male's underside resting on the female's back. The female psudeo penis is completely slack during mating (Law, J. 2004). Males have also been witnessed tapping their leg before mating and showing aggression towards other male clan members (Bennett, pers. comm. August 2014).

9.4 Bathing

Water plays an important part of the Spotted Hyena’s life not only for bathing but also practical social playing and cooling off. Whether housed inside or outside a water source must be provided in the form of pools, streams or a large water feature. Many hyenas enjoy bathing in shallow pools and will also utilise tanks as deep as 3 metres. If housed outside most water features are placed in highly visible locations as they will spend a lot of time around the source especially in the warmer months. To avoid aggression and allow the clan to bath and play together the pool should allow multiple entry points (Foote unpublished data 2014 and Shoemaker, Dulaney, Noble 2006). A party of ten hyenas were witnessed in Uganda bathing on a lakeshore, not swimming but wading and chasing each other around (Hans 1972).

9.5 Behaviour Problems

Pacing has been witnessed in captive situations where different groups of hyena are housed separate sharing direct fence contact and where a single male is housed alone and has visual of other hyenas. Destructive behaviours have been seen in attempts at getting to one another such as destruction to fences and digging holes under the fence. To avoid any unnecessary behavioural problems, having the ability to separate the animals for feeding will reduce any chasing and aggression from dominant animals.

No direct aggression has been seen towards keepers or public, but it has been witnessed even though they enjoy tactile contact, a single male has changed his behaviours towards keepers due to neighbouring animal’s behaviours and females can become protective of their new cubs. The Spotted Hyena tends to be more wary and nervous around keepers especially in confined raceways, unsure areas and with sudden movements. Desensitisation programs have been implemented to familiarise hyena to walk through raceways and new areas. Females in season can also have some effects on the group dynamics (Bennett, pers. comm. September 2014).

9.6 Signs of Stress

The Spotted Hyena will vocalise in times of stress. “Laughing” is a high pitched cackle, which is generally produced when the hyena is stressed or nervous. When being chased by another hyena in pursuit of food, or when attempting to attack a much larger animal, such as a lion, the Spotted Hyena will often vocalise this "laughter". The intensity and volume of the sounds is thought to correlate to the distress or excitement a hyena is feeling (vocalization in Spotted Hyena). Signs of stress may include their body posture to change to be very submissive towards other animals, excessive pacing even to the level where feet start to bleed, seeking distance away for the stressful situation, and a low pitched nervous stutter vocalisation which they will make when nervous or scared in a situation. (Bennett, pers. comm. September 2014).

9.7 Behavioural Enrichment

Environmental enrichment is the process of providing stimulating environments for zoo animals in order for them to demonstrate their species-typical behaviours, to allow them to exercise control or choice over their environment, and to enhance their well-being (National Zoological Park).

Enrichment includes the design of stimulating and naturalistic enclosures, the housing of appropriate social groups in zoos, and the introduction of objects, sounds, smells and other stimuli in the animal’s environment. Enrichment can be provided in a variety of ways, such as exhibit design, training and conditioning, olfactory, auditory, food related and novel objects (National Zoological Park). Bathing can also be incorporated as part of behavioural enrichment, please refer to section 9.4 (Foote, unpublished data 2014).

The provision of enrichment should become a part of every captive animal’s daily husbandry needs. A daily enrichment schedule should be in place for all captive animals and be monitored and changed in accordance to their responses, as some provisions may not appeal to every animal (Foote, unpublished data 2014). A new study shows the African carnivores are adept at problem-solving, and can even count. This makes scientists believe that Spotted Hyena could have comparable intelligence levels to some primates (SciTech daily 1998-2014) which highlights the need of providing a great deal of mental stimulation in captivity for the Spotted Hyena.

When providing enrichment for the hyena, it is very important to keep in mind their jaw capacity as many novel items may not be suitable as they can be easily chewed into small pieces and swallowed. Veterinary approval must always be sought first and providing sufficient enrichment for all animals housed together is also necessary to avoid any aggression within the clan (Foote, unpublished data 2014).

For enrichment on or off exhibit, successful items include bones, rawhide, whole prey, carcasses or partial carcasses, eggs, fruits and vegetables and dog biscuits. Hiding food in the exhibit or in papier-mâché animal forms, paper bags, Boomer Balls, boxes or tubes will stimulate natural behaviours and provide exercise, as will Christmas trees, flowers and other horticultural cuttings and scents from natural prey (urine or faeces) or commercially available animal scents ( Shoemaker, Dulaney, Noble 2006). Provided below are a number of different enrichment ideas currently in use by other institutions. Section 9.8 highlights the use of operant conditioning as a means of enrichment.

Fruit/Vegetables Meats Items not allowed

Food Enrichment Items

Melons Brussel sprouts Green beans

Mice Quail Fish Liver Heart Dog biscuits Dog and cat

chow Pig ears

Onion Garlic Citrus

Table 9: Food enrichment items offered to the Spotted Hyena at Denver Zoological foundation.

Table 10: Monarto Zoo, Spotted Hyena enrichment schedule.

http://www.zoochat.com/1659/spotted-hyena-enrichment-295307/Figure 10: Spotted Hyena with ostrich egg – able to break open

http://gardenwalkgardentalk.com/2012/02/22/w4w-time/Figure 11: Spotted Hyena moving a tough ball structure

http://perezhilton.com/teddyhilton/2013-10-31-cute-baby-hyena-plays-with-pumpkin-at-berlin-zooFigure 12: Spotted Hyena cub eating a pumpkin

http://www.oaklandzoo.org/blog/author/zena-the-zookeeper/Figure 13: Spotted Hyenas enjoying a cool off in water bath

http://spzkeeperblog.files.wordpress.com/2013/03/lou-with-ball.jpgFigure 14: Spotted Hyena with ball

http://www.givengain.com/cause/2626/posts/107454/Figure 15: Spotted Hyena tug-o-war with horse tail

9.8 Operant Conditioning

There are many great benefits for training animals in captivity, particularly as a means of enrichment and to assist in the husbandry needs of the animal (Foote, unpublished data 2014). A new study shows

the African carnivores are adept at problem-solving, and can even count. This makes scientists believe that Spotted Hyena could have comparable intelligence levels to some primates (SciTech daily 1998-2014).

Monarto Zoo, South Australia has implemented a training program which has proven success with the Spotted Hyena achieving a range of behaviours. The following list comprises of the behaviours that have been achieved (Bennett, pers. comm. September 2014).

Up Leg L/R Shoulder L/R Target Mouth open + hold Down Sit Belly (side on) Station + hold Foot Stand front on Vocalise Neck Onto box Ear L/R Recall Scales Gate desensitisation Race desensitisation Novel items: Painting, fetch stick, touch ball and block into basket

The training program must be well thought out and planned before initiated. A training program outline must be written with aims and justification of the behaviours wanting to be achieved and clear and precise steps of how the behaviours will be achieved. Primary and secondary trainers must be allocated along with any cue, bridge or rewards that will be used. The program must be consistent, progress documented and reviewed accordingly (Foote, unpublished data 2014).

Appendix 1 shows an outline of a training program and a list of behaviours that can be achieved.

9.9 Introductions and Removal

The Spotted Hyena has a very structured clan with females being the highest dominant member. When introducing hyena there are many options that can be undertaken and should be undertaken with time

and care to allow for the best chance of success. Any introduction must start with fence contact first and monitoring of the behaviours.

Female/Male introduction: Fence contact first for as long as required to monitor the behaviour. You want to see the male submissive and no aggression from the female.

Female/female and male/male introduction: Fence contact first for as long as needed to monitor the behaviour. You must be prepared for fighting and aggression when allowing them together and must only separate if absolutely necessary such as one of the animals is requiring veterinary assistance.

Perth Zoo, Western Australia house two males only, they have had numerous aggression issues between this pair. Contraception was trialled on both males which seemed to have minimised the aggression between the two as the hormone levels were altered.

When housing a clan of Spotted Hyena you must have a plan in place for when there is a need to separate and re-introduce. The female clan members especially can be separated for feeding and various procedures but must only be for a maximum of 1 hour. Re-introducing will all depend on the age, nature and hierarchy of the animals involved.

At Monarto Zoo, South Australia, the female Spotted Hyena at various times have been separated for days and in one instance months, and on reintroducing, high levels of aggression and greetings were seen, as the hierarchy was altered and needed re-establishment.

At Monarto Zoo, the clan are required to be housed in a more confined area at times so splitting the clan is necessary to avoid aggression. When separation is required, females should remain together, and likewise with males, to avoid hierarchy changes between the sexes. All clan mates can then be reintroduced back together into their larger area.

Many situations have been trialled at Monarto Zoo, South Australia, and are happily contactable if needing any further clarification or advice (Bennett, pers. comm. September 2014).

9.10 Interspecific Compatibility

Due to the carnivorous nature of the Spotted Hyena, they should not be housed with any other species. At Monarto Zoo, Interspecies rotations of animals in exhibits and fence contact with the African Painted Dogs have proven successful with stimulating benefits (Bennett, pers. comm. August 2014).

9.11 Intraspecific Compatibility

Spotted Hyenas naturally live together in a clan of both male and females. Depending on the hierarchy and age of the animals will determine how sexes are put together. This process should be gradual and monitored intensely. Introductions should be one on one to begin with and worked up from there based on individual behaviours. Natural aggression is to be expected when hierarchies are determined and should be left to sort unless intervention is absolutely necessary (Bennett, pers. comm. August 2014).

10. Breeding

10.1 Mating System

Spotted Hyenas are highly polygynous meaning that males will mate with multiple females and mating is aseasonal (Law 2004).

10.2 Ease of Breeding

Experienced breeders have proven to breed well in captivity. Evidence suggests that first time breeders only have a 20% chance of success which results in either the female dying or the cub being stillborn due to the un-ruptured penileclitoris. The long and circuitous path of the urogenital canal and the extraordinary external genitalia of the female appears to present a difficult obstacle to the descending neonate. In a captive environment the intervention of caesarean section has been successfully performed when labour has not progressed (L.G Frank and S. E Glickman 1994). A clan of hyenas is made of up of many breeding females and multiple cubs of different ages at the den. This however can present the chance of cubs being killed by other females in the clan if there is a hierarchical issue occurring. Adequate space and den options are a necessity for a breeding clan (Bennett, pers. comm. September 2014).

Figure 16: Calculation of the reduction in potential reproductive success caused by maternal and neonatal mortality in first time pregnant Spotted Hyena (Nature 1995).

10.3 Reproductive Condition

10.3.1 Females (Bennett, pers. comm. August 2014).

1) Juveniles – No signs of a torn pseudo penis and nipples are not clearly visible and animal weight is below that of average adults.

2) Oestrus - No real visual signs the animal is in oestrus, but you may see some female to female aggression and fighting. Male will show interest in females in oestrus and different behaviours between males in the clan.

4) Pregnancy – Can be determined in later stages of pregnancy by enlarged nipples and belly, and female to female aggression and fighting. Hyena mating tends to occur at night so a visual may not necessarily be witnessed.

5) Lactation - Nipples large and elongated.

6) Proven female breeder – Have reached adult weight and will have a torn cliteriform appendage which is not always obvious and may only be seen during a greeting or when the appendage is extended.

10.3.2 Males (Bennett, pers. comm. August 2014).

Males show no real physical change, apart from reaching adult size. They are known to show some breeding behaviours including leg tapping, following the female around very closely and aggression between male clan members.

Appendix 2 shows physical changes to a female Spotted Hyena while pregnant with twins

Appendix 3 shows a torn cliteriform appendage of a proven breeder

10.4 Technique Used to Control Breeding

The following methods can be used to prevent the captive breeding of Spotted Hyena (Kalvas, pers. comm. 2014).

Separating males and females until breeding is desired Contraceptive implant of Deslorelin 9.4mg (Suprelorin) suitable for both male and female, given

subcut under anaesthetic and lasts around 12-18 months Irreversible ovarian hysterectomy or castration

The contraceptive implant has been used on a female Spotted Hyena as young as 18 months at Monarto Zoo which lasted approx. 2 years. (Bennett, pers. comm. August 2014).

10.5 Occurrence of Hybrid

Not applicable (Kalvas, pers. comm. 2014).

10.6 Timing of Breeding

Spotted Hyena breeding can occur throughout the year (Law, J. 2004).

10.7 Age of First Breeding and Last Breeding

Sexual maturity in the female Spotted Hyena occurs around the age of 3 years; on average 40 months (Law, J. 2004). Age at first parturition varies substantially between two and five years (Frank et al. 1995a, Holekamp et al 1996; Hofer & East 1996). No literature was found on last breeding age but considering the oldest living hyena reported was 41 years with an average lifespan of 25 years, it’s predicted Hyenas could breed up until their early to mid-twenties if in overall good health and condition (Kalvas, pers. comm. September 2014).

10.8 Ability to Breed Every Year

Spotted Hyenas breed about every 16 months, with a range from 11 to 21 months (Law 2004).

10.9 Ability to Breed More Than Once Per Year

The only instance a Spotted Hyena may breed more than once per year is if the cub has died, or has been removed for hand rearing, in which the female will enter oestrus and could conceive again (Kalvas, pers. comm. September 2014).

10.10 Denning Requirements

In the wild two to six weeks after birth, the mother transports young from the burrow in which they were born, to a communal den. Communal denning seems to be an important part of Spotted Hyena social behaviour, but no communal care of young takes place (Law 2004). In order to successfully parent-rear cubs in a captive situation, it is recommended that females have access to many denning options, and a communal den situation is probably not feasible under captive conditions for risk of injury to young (Foote, unpublished data 2014 and Shoemaker, Dulaney, Noble 2006).

10.11 Breeding Diet

For pregnant females, the diet should increase during different stages of pregnancy including early and late gestation and lactation period. Dietary requirements are calculated as a multiple of Basal Metabolic Rate (kcal/day). During breeding the diet should increase from maintenance (BMR x 1.8) to a multiple of 2.1 for breeding and increasing to a multiple of 2.3 during late gestation. Periods of lactation should see an increase of up to BMRx4.0. In summary, an overall diet increase of 10-20% is required for a breeding animal with requirements being adjusted in accordance to the individual’s weight. (Kalvas, pers. comm. September 2014).

10.12 Oestrus Cycle and Gestation Period

Spotted Hyena gestation period is on average 110 days, (4 months) (Law 2004) and they have a 14 day long oestrus cycle at intervals of 1- 1.5 years (MaasaiMara). Males will generally mate with the female a couple of times while she is in season (Kay E Holekamp 2001).

10.13 Litter Size

Spotted Hyenas have 1-3 cubs, usually 2. Siblicide can occur, where a dominant cub can kill subordinate cubs or supress other cubs from emerging from the den. There is competition from birth to reach mothers milk when she returns to the den (Bennett, pers. comm. October 2014 and the Animal files).

10.14 Age at Weaning

Cubs are nursed for a prolonged period and not weaned until they are between 14 and 18 months of age (Hofer & East 1993c; Holekamp et al. 1996). Sexual maturity in the female Spotted Hyena occurs around the age of 3 years; on average 40 months (Law, J. 2004). Age at first parturition varies substantially between two and five years (Hofer & East 1996). Evidence suggests that first time breeders only have a 20% chance of success which results in either the female dying or the cub being stillborn due to the long and circuitous path the cub must pass through (L.G Frank and S. E Glickman 1994).

A recent study by Holekamp and her colleagues suggests that status of cubs begins in the womb. They discovered that in the final weeks of pregnancy, high-ranking females produce a flood of testosterone and related hormones. These chemicals saturate the developing cubs both males and females and make them more aggressive. They’re born with a drive to dominate. By contrast, a pregnant subordinate female produces a smaller spike of hormones, and her descendants become submissive. Holekamp says this is the first evidence in mammals that traits related to social status can be “inherited” through a mother’s hormones rather than genetics (Londolozi 2014).

Young Spotted Hyenas are dependent on maternal milk that contains a high fat, protein and energy content for approximately the first 12 months of life, (Hofer and East 1995). This long dependence on maternal milk is likely to favour selection for strong sibling rivalry, leading to siblicide when ecological conditions such as low food resources prevent the delivery of sufficient maternal milk to sustain a twin litter. When twin littermates competed for access to maternal milk supplies, the rate of sibling aggression increases as the interval between nursing visits by the mother increased. This escalated sibling rivalry resulted in increased exclusion of the subordinate sibling from access to maternal nipples and milk supply (Hofer and East 1993). It has also been witnesses from captive hyena births how infrequent the cub seems to nurse (Foote, unpublished data 2014).

Escalated sibling conflict of this kind that persisted for weeks caused the development of large irregularities in body mass between littermates and, in some instances, the death from starvation of the subordinate member of a litter (Hofer and East 1997). Siblicide in twin Spotted Hyena litters has onlybeen observed in populations that forage on low or fluctuating levels of prey (Hofer and East 1997. In Spotted Hyenas, siblicide normally occurs before a twin litter reaches 3 months of age (Hofer and East 1997). Siblicide aggression between Spotted Hyena neonates results in mortality of up to 25% of offspring (Nature 1995).

10.15 Age at Removal from Parents

Males disperse from their natal clan shortly after becoming reproductively mature, which is usually when they are two years old. Females however, remain in their natal clan (Kay E. Holekamp 2007-2014).

10.16 Growth and Development

The newborn Spotted Hyena weighs from 1 to 1.6 kilograms and is quite advanced, being born with their eyes open and a full set of teeth. Newborns are almost entirely black. If siblings are the same sex, they begin fighting violently soon after birth, which can result in the death of one cub. Since single young receive more food and mature faster, this behaviour is probably adaptive. A diversity of calls becomes apparent during the first week, the cub can utter a high-pitched whimpering begging call and a series of very high pitched but clearly recognisable whoops. The black colour of the cub gradually changes into the spotted coat of the adults, beginning at one to two months, with the head turning light grey . Two to six weeks after birth, the mother transports young from the burrow in which they were born, often to an abandoned aardvark, warthog or bat-eared fox burrow, to a communal den. The major source of food for the young during this time is milk from the mother (Law 2004).

The milk of the Spotted Hyena has the highest protein content (mean 14.9%) recorded for any terrestrial carnivore, a fat content (mean 14.1%) exceeded only by that of paleoarctic bears and the sea otter, and a higher gross energy density than the milk of most terrestrial carnivores (Hofer & East 1995a). Due to their milk's high energy content and the long nursing period, Spotted Hyenas have the highest energetic investment per litter of any carnivore (Oftedal & Gittleman 1989). Weaning occurs from about the age of 12 to 16 months, which is extremely late. By the weaning age, juvenile Hyenas already have completely erupted adult teeth, which is also very rare (Law 2004).

Cubs begin to eat meat from kills near the den at about 5 months, but they are suckled for as long as 12 to 18 months, an unusually long time for carnivores. This is probably a necessity, as most kills are made far from the den, and hyenas, unlike jackals and hunting dogs, do not bring back food and regurgitate it for their young. At about 1 year, cubs begin to follow their mothers on their hunting and scavenging forays. Until then, they are left behind at the den with a babysitting adult (Janssen 2010).

11. Artificial Rearing

11.1 Housing

Housing will be dependent on the age and health of the cub. Preferably, you want to keep the cub in the same area as other hyena. It's best that they grow up with the sounds, smells and sights of others as this will help promote natural behaviours. Setting up an area with no high steps or benches in the hyena holding area is best, or if this is not possible, taking the cub to the hyena area every day is essential. The area needs to be kept warm enough for the cub to maintain body temperature if they are very young (less than a few weeks). The holding area must provide a suitable den for the cub such as half a dog kennel, flipped upside down. Cubs must be provided with a den to replicate how they would be raised in the wild. Bedding can be soft blankets and towels with no loose strings or holes; newspaper and shavings are not recommended. Unsoiled towels can be left in so their scent remains (Genter SSP, pers. comm. October 2014).

The replicate den should be kept in place for as long as the cub uses it. It will become a place of comfort and security. It is highly recommended to pair lone cubs with other hand reared cubs if possible. A lone hyena cub may not develop the behaviours it needs to display to other hyena as an adult, which could result in severe attacks and a solitary life for that animal. The sooner they are housed with other hyena, the better (Genter SSP, pers. comm. October 2014).

11.2 Temperature requirements

A heat lamp can be provided for added heat, this allows the cub to seek warmth and move away as desired. The temperature should be checked and recorded on a daily basis. As a baseline the temperature could be 20 degrees under the heat and 15-18 degrees away. The height of the lamp can be moved and re positioned to seek a suitable temperature (Bennett, pers. comm. August 2014, Foote, unpublished data and Monarto Zoo, Spotted Hyena hand rearing notes 2007). Heated floors and heaters can also provide extra warmth if needed. If the cub is weak or sick they may need an incubator to help them maintain body temperature. Rectal temperature can be taken throughout the day for the first couple of weeks until you are sure that they can maintain their temperature without extra care (Genter SSP, pers. comm. October 2014).

11.3 Diet and Feeding Routine

Young hyena can be hand raised on Esbilac or milk formulas that are developed for kittens. At the University of California at Berkeley, neonates are fed 70 ml of formula four times a day, gradually changing to 200 ml of formula twice a day (Shoemaker,Dulaney,Noble 2006).

Monarto Zoo, South Australia hand raised a Spotted Hyena cub in 2007. Considering the high protein and fat content of a female hyena’s milk, a special formula was designed by a local supplier (Wombaroo Passwell) to meet the cub’s needs. Table 12 highlights how much milk is needed in accordance to cub’s weight.

The behaviours witnessed at Monarto Zoo, South Australia from a breeding female in the first few weeks after giving birth shows the female to be very attentive to her cub spending most of her time in and around the den not moving far at any time. After 2-3 weeks the female seems to spend more time with the clan and only attends to the cub a few times throughout the day. From observations and previous hand rearing experience seen at Monarto Zoo, the following feeding routine could be followed as a guide (Bennett, pers. comm. August 2014 and Foote, unpublished data).

The following information will vary in accordance with the cub’s weight and response to rearing. Follow table 12 as a guide for milk quantity. The following information was taken from previous hyena hand rearing records. Under feeds per day, comments in bold highlight what would be undertaken differently if another hyena was hand raised (Bennett, pers. comm. October 2014).

Age of cub Feeds per day Food type and amount

0-21 days 4 milk feeds per day Milk, amount will depend on cub’s weight

(7-8 feeds per day)21-51 days 3 milk feeds per day

(4-6 feeds per day)Milk, amount will depend on cub’s weight

52-74 days 2 milk feeds per day + 1 meat/bone feed(2-3 feeds per day)

Milk, amount will depend on cub’s weight.During this age bracket 60g mince meat and small bones can be given once per day.

75- 90 days 1 milk feed per day + 1 meat/bone feed

Milk, amount will depend on cub’s weight.During this age bracket 60g mince meat and small bones can be given, including turkey drumsticks

90- 100 days 1 milk feed per day + 1 meat/bone feed

Milk, amount will depend on cub’s weight.During this age bracket 210g mince meat or meat pieces and small bones can be given, including turkey drumsticks

100-110 days 1 milk feed per day + 1 meat/bone feed

Milk, amount will depend on cub’s weight.During this age bracket 300g meat pieces and small bones can be given, including turkey drumsticks.

Table 11: Number of feeds and food type for hand rearing Spotted Hyena (Monarto Zoo, Spotted Hyena hand rearing notes 2007).

Table 12: Highlights (Wombaroo Passwell) milk replacement composition and feeding guide for the Spotted Hyena (Foote, unpublished data 2014). Denver Zoo, Colorado has experienced hand raising of Spotted Hyena cubs. From their experience the following can be used as a guide.

Feeding times will depend on the cub. Usually start with feeds every 2-3 hours for the first few days or weeks. Let the cub tell the care taker when they are due for a feed and don’t wake them for feeding. Initially start with at least 7-8 feeds a day spaced by 3 hours. As the cub grows and sleeps for longer periods, start to remove late night feeds until you are just feeding between 7am and 10pm. Then you can start removing select feeds during the day, adding the removed milk to other feeds being careful not to feed more than 7% of their body weight at a time (Genter SSP, pers. comm. October 2014).

Around 2 months old, start exposing them to meat, particularly bones, this is a great way to encourage them to like meat. Try adding meat to the top of bones to encourage them to eat meat. Pay attention to faecal consistency and if they become too loose or odd in colour you may need to slow down the transition, this transition is best undertaken slowly. Esbilac milk formula was used and when around 2.5 months old half a jar of Gerber baby food (meat flavours such as turkey, beef and chicken) were added to the formula (Genter SSP, pers. comm. October 2014).

The cubs should be fed about 10 to 15% of their total body weight daily. To calculate their diet, this formula can be used:

Body weight in grams x 0.2 = total amount of grams to feed. 1 gram = 1ml.

The total amount of formula should be divided into the number of feeds for the day. For example, if the cub weighs 8 pounds, convert to kg by dividing by 2.204 = 3.6kg, convert to grams 3.6 x 1000 = 3636g then multiple by 15%. 3636 x 0.2 = 727.2g which equals 727.2ml.

This would be divided into the number of feeds per day. If you are doing 6 feeds per day they would receive 121.2ml of formula at each feed (Genter SSP, pers. comm. October 2014).

Milk is fed with the use of regular plastic baby bottles, and appropriate size nipples for feeding. Hyenids feed sternally (laying on abdomen) with neck extended and head tilted up slightly. Milk should be warm (about body temperature) but never hot. Milk made up should be used within 24 hours, both constructed formula and opened powder must be refrigerated (Genter SSP, pers. comm. October 2014).

11.4 Specific Requirements

Toileting – A small towel must be moistened with warm water and stimulate the cubs genitals by patting/rubbing immediately after each feed to allow them to defecate and urinate. This process mimics the mother’s natural behaviour of licking the genitals for the same purpose (Kalvas, pers. comm. September 2014 and Aecvets 1975). The process and choice of stimulation will depend on the cubs temperament and caretaker, consistency is very important. Cubs will likely urinate during each stimulation, but not always defecate. Normal defecation should occur at least 2-3 times a day ( K.Grant 2004).

Weighing - The cub should be weighed on a daily basis to monitor progress, particularly during the early stages of hand-rearing. Once stable, this can be decreased to once every 2 days (Foote, unpublished data).

Supplements - Lectade was used at Monarto Zoo as an oral rehydration to help with diarrhoea and dehydration. Lectade comes in powder form which is dissolved in warm water (Monarto Zoo, Spotted Hyena hand rearing notes 2007).

Physical/mental stimulation – Cubs should be introduced to enrichment to encourage natural behaviours and active play as soon as possible. Smaller items like Kong’s, smaller boomer balls and cardboard can be provided initially. As the cub grows in size and strength the size and weight of enrichment can increase (Genter SSP, pers. comm. October 2014). Please refer to Section 9.7 which highlights behavioural enrichment ideas.

11.5 Identification Methods

Identification methods are the same as those mentioned in section 5.3. As a young cub the hyena is fully chocolate brown/black and over time starts to develop white/grey markings and spots. In the early stages, cubs look very different in comparison to adults (Foote, unpublished data 2014). Determining the sex of a cub can be difficult and the easiest approach is taking a blood test or DNA sample. Keepers may notice behaviours that will help to determine the sex. Females tend to be independent, confident and assertive. Males are more nervous, shy, and submissive even at a few days old. The male penis is slightly more pointed than the females, which tends to be rounded at the tip (Genter SSP, pers. comm. October 2014).

11.6 Hygiene

Sanitisers such as Milton anti-bacterial solutions should be used to clean and disinfect all hand rearing equipment such as bottles and teats and to be rinsed with pre boiled water before use. Gloves, hand wash, footbaths and utilising non-carnivore staff must be used in the early stages until the cub is stabilised. The milk replacement must be made as per packet instructions to ensure consistency, and timing of feeds must also be consistent (Bennett and Kalvas, pers. comm. September 2014). It is extremely important that a high level of hygiene be maintained throughout the hand-rearing process (Foote, unpublished data 2014). Blankets and towels should be washed frequently to prevent bacterial build up. (Genter SSP, pers. comm. October 2014).

11.7 Behavioural Considerations

Best approach to encourage and imprint natural behaviours on the young cub would be to house the hand raised animal in the close vicinity of the other hyena, so the cub can watch interactions and hear vocalisations. If this is not achievable, the cub (beginning from a young age), should be taken to the hyena area each day to socialise with the other hyenas safely by mesh contact. Over time the duration of visits should increase in hours to overnight stays. Behaviour should be closely monitored and once positive behaviours have been witnessed through mesh contact, an introduction back into the clan can occur. Please refer to section 9.9 for introductions (Bennett, pers. comm. September 2014).

As little interaction as possible with the cub is recommended. The more humans interfere, the less likely the animal will behave normally and therefore may not be readily accepted by others. It will also make the transition of going "hands off" easier in the long run. Even at a very young age, their bite is extremely powerful and high care must always be taken before entering their area. Encourage play with toys and conspecifics but do not let them practice on you or you may get serious bite wounds. Their

nails are also quite sharp at this age. You can trim them with nail clippers to prevent scratches and prevent their nails from getting caught in bedding (Genter SSP, pers. comm. October 2014).

11.8 Use of Foster Species

The best approach, considering the hyenas social structure, would be close contact with other hyenas, (please refer to section 11.7) (Bennett, pers. comm. September 2014). Potentially a domestic dog could be used for companionship; however there are no other foster species appropriate for the cub to suckle from (Kalvas, pers. comm. September 2014).

11.9 Weaning

Observations from Monarto Zoo suggest the cub can be introduced to solids as young as 2 months old due to their advance development and that they are born with teeth. Mincemeat can be introduced around 75 days (2.5 months old) and slowly progressing to meat chunks and bone at around 90 days (3 months old). The introduction of solids must be monitored to make sure they are capable of chewing well. From 110 days old (4 month), cubs can be introduced to more solids in conjunction with milk feeds (Monarto Zoo, Spotted Hyena hand rearing notes 2007 and Foote, unpublished data 2014). As the cub begins consuming solid foods in a measurable amount, the volume of formula can be reduced proportionately. Many times cubs will take the bottle at some feedings but refuse it at others. Eliminating specific feedings rather than reducing the volume at each feeding allow the cubs to get hungry enough to explore other food options available, such as a bowl of meat (Genter SSP, pers. comm. October 2014).

12. Acknowledgements

Thank you to Anna Bennett (Team leader of Carnivours and Ungulates Monarto Zoo) for your assistance, knowledge and firsthand experience with many aspects of the manual. Also thanks for the use of your pictures. Your passion for Spotted Hyena is truly inspiring.

Thank you to Jerome Kalvas (veterinarian at ZOOSSA) for advice on medical, dietary, breeding and quarantine components of captive Spotted Hyena. Also thanks for the use of many of your pictures also.

Thank you to Sarah Washford (Senior Keeper Carnivores Monarto Zoo) for your assistance and advice.

Thank you to Heather Genter (Spotted Hyena SSP Coordinator and Studbook Keeper Denver Zoo) for advice and experience with hand raising and dietary information.

Thank you to Gordon Rich (Wombaroo Food Products / Passwell Pty Ltd) for Hyena milk replacement information.

Thank you to Jodi Buchecker (Species Management Officers at Monarto Zoo) for advice with spatial and transport requirements.

Thank you to Gert Skipper (Curator Collection Development Adelaide Zoo) for advice with spatial requirements.

Thank you to Nicky Tomkinson for information on hand raising Spotted Hyena.

And, thank you to Becky Thomasson (Senior Technical Officer Zoology Carnivore/Ungulates Perth Zoo) for information on housing 2.0 Spotted Hyena.

13. References

Adam Bannister, May 15, 2012, Londolozi Est 1926, http://blog.londolozi.com/2012/05/why-are-female-hyenas-bigger-than-males/, Accessed 19 September 2014.

Alan Shoemaker Riverbanks Zoological Park, Mike Dulaney Cincinnati Zoo and Botanical Garden, 1992, Minimum AZA Guidelines For Keeping Hyenas And Aardwolves In Captivity, http://www.species.net/Carnivora/Canidae/GuideHy.htm, Accessed 15 September 2014.

Alan Shoemaker, Mike Dulaney and Gary Noble, 2006, Standardized Animal Care Guidelines for Hyaenas and Aardwolves, viewed and cited pages 1-29.

Animal Emergency Centre, 1975, http://www.aecvets.com.au/docs/Hand-Rearing-of-Puppies-NATIONAL.pdf, Accessed 25 September 2014.

Brottman, M, 2012, Hyena, Reaktion Books LTD, London, UK.

Conservation and Education Oakland Zoo, 2014, http://www.oaklandzoo.org/blog/author/zena-the-zookeeper/, Accessed 9 September 2014.

CC 2006 - 2014 theanimalfiles, http://www.theanimalfiles.com/mammals/carnivores/hyena_spotted.html , Accesses 24 April 2014.

Designer animals 2001 Spotted Hyena, 2014, http://designeranimals2011.wikispaces.com/Hyena-Vivian+Lin Accessed 24 April 2014.

Garden Walk Garden Talk, 2010-2014, http://gardenwalkgardentalk.com/2012/02/22/w4w-time/,Accessed 9 September 2014.

Gyuyeong Han, 2012, Zoo Chat, http://www.zoochat.com/1659/spotted-hyena-enrichment-295307/, Accessed 9 September 2014.

Honer, O., Holekamp, K.E. & Mills, G. 2008. Crocuta crocuta. The IUCN Red List of Threatened Species. Version 2014.2 <http://www.iucnredlist.org/details/5674/0 > Accessed 24 April 2014.

Jenise Alongi, Animal Facts Encyclopedia, 2011-2014, http://www.animalfactsencyclopedia.com/Hyena-facts.html, Accessed 13 October 2014.

Kay. E. Holekamp, 2007-2014, ICUN Hyaena Specialist Group, http://www.hyaenidae.org/the-hyaenidae/spotted-hyena-crocuta-crocuta/crocuta-distribution-and-habitat.html Accessed 24 April 2014.

Kay E. Holekamp , 2011, Scientists at work, http://scientistatwork.blogs.nytimes.com/2011/07/19/how-spotted-hyenasmate/?_php=true&_type=blogs&_r=0, Accessed 3 September 2014.

Kruuk. H, 1972, The Spotted Hyena, The University of Chicago Press LTD, Chicago, USA.

Law, J. 2004. "Crocuta crocuta" (On-line), Animal Diversity Web, http://animaldiversity.ummz.umich.edu/accounts/Crocuta_crocuta/, Accessed 27 July 2014.

International Air Transport Association (IATA), 2001, Live Animal Regulations 28th Edition, Montreal Geneva.

Maasai Mara, Africa’ s Greatest Wildlife Reserve, http://www.maasaimara.com/entries/spotted-hyena, Accessed 3 and 12 September 2014.

Mills, M.G.L, 1990, Kalahari Hyenas, Comparative Behavioral Ecology of two Species, The Blackburn Press, New Jersey, USA.

National Geographic, Spotted Hyena, Crocuta crocuta, 1996-2014 http://animals.nationalgeographic.com.au/animals/mammals/hyena/ Accessed 24 April 2014

National Zoological Gardens of South Africa, 2001-2013, http://www.givengain.com/cause/2626/posts/107454/, Accessed 12 September 2014.

Paul Janssen, Out of Africa with Ellen and Paul, http://www.outtoafrica.nl/animals/enghyena.html, Accessed 12 September 2014.

PerezHilton.com, Berlin Zoo,2005-2014, http://perezhilton.com/teddyhilton/2013-10-31-cute-baby-hyena-plays-with-pumpkin-at-berlin-zoo, Accessed 9 September 2014.

Planet Wildlife 2010-2013, http://www.planetwildlife.com/information/species/spotted-hyena , Accessed 24 April 2014.

San Diego Zoo Global, 2014, http://animals.sandiegozoo.org/animals/spotted-hyena, Accessed 10 May 2014

Smithsonian Conservation Biology Institutions, 2014, http://nationalzoo.si.edu/SCBI/AnimalEnrichment/, Accessed 12 September 2014.

Wildlife South Africa, http://wildlifesouthafrica.com/factfiles/spotted-hyena.htm, Accessed 5 September 2014.

Wombaroo Food Products, 2014, http://www.wombaroo.com.au/about_us.htm, Accessed 27 July and 15 August 2014.

World Association of Zoos and Aquariums WAZA, United for Conservation, http://www.waza.org/en/zoo/visit-the-zoo/dogs-and-hyenas/crocuta-crocuta , Accesses 10 May 2014.

15. Appendix

Appendix 1 Example of operant conditioning programs used by Spotted Hyena keepers at Monarto Zoo (Bennett 2011).

MZ Hyena Training/Conditioning Program record sheet – 2011

Date: ______________________ Time: ______________________

Keeper: ______________________ Keeper: ______________________

Place a tick/comment in the box to indicate the result of the individual’s session. Table of cues on reverse.

Behaviour Piltengi Gamba Forest Kigali PinduliUpLeg L/RShoulder L/RTargetOpen + HoldDownSitBelly (side on)Station + HoldFootStand front onVocaliseNeckBoxEar L/RRecallScalesGate desensitisationRace desensitisationNovel items

- Painting- Stick- ‘touch’ ball- ‘Block’ in basket

Comments:

Monarto Zoo Hyena (Crocuta crocuta) Training/Conditioning Program

Monthly update – January 2011

Aims:

Behaviours:

1 Up (stand up on fence to expose underside & pads of front feet)2 Leg (stand parallel to fence so hip & shoulder pushes into fence, poke thigh) left &

right sides3 Shoulder (stand parallel to fence so hip & shoulder pushes into fence, poke shoulder)

left & right sides4 Target (touch end of target stick with nose)5 Open (open mouth wide so teeth, gums & teeth can be inspected)6 Down (sternal recumbency)7 Sit8 Foot (presents foot up on/through mesh and allows contact)9 Stand front on (at 90 degrees to fence – front on to keeper)

10 Vocalisation (capture opportunistically)11 Box (jumps up onto box)12 Ear (allows contact with ears i.e. fly repellent application)13 Recalls (work with two trainers in same session (does a few behaviours with one

trainer, then called over to work with other trainer, etc...)14 Swabs/topical treatments on body15 Crate16 Raceway work

Background:

The hyena training/conditioning program began formally on 1st December 2010, therefore, this is the first monthly update for the program. At the time of writing, Monarto Zoo holds 6.2 Spotted hyena. 4.2 are adults and 2.0 are juveniles. The facility consists of one exhibit, and six yards, interconnected by a central raceway.

Training history: none except Piltengi have had any formal training. In the past, Piltengi has been trained to present his right leg/thigh (pressed close and parallel to fence line) for intramuscular injection.

Roles and responsibilities:

Primary trainers: Anna Bennett (Senior Keeper of Carnivore section) and Linda Gardiner (Team Leader of Carnivore/Ungulate section).

Secondary trainers: none appointed as yet due to program just starting. As the hyenas’ training progresses, secondary trainers will be incorporated into the program to maintain established behaviours.

Summary:

The total number of training sessions for December was 18 out of a possible 31. The sessions were conducted once a day. Due to the housing/holding configurations (yard and exhibit) of the hyenas on a day to day basis, not all hyenas could be trained each session. The adult females and the juveniles (Amani and Makhali) were sometimes exhibited and yarded as a group. The adult males (Kigana and Mwenka) and the juveniles were sometimes exhibited and yarded as a group. The adult males Piltengi and Gamba were held in yards as a single animal. Due to the layout of the facility, the hyenas in the yards for the day had a clear view of the training sessions as they occurred.

Results:

The table below shows the number of training sessions for each individual hyena with the results also expressed as a percentage (number of actual sessions divided by the total number of training sessions conducted for the month shown, multiplied by 100)

Hyenas Piltengi♂

Gamba♂

Kigana♂

Mwenka♂

Forest(Msitu) ♀

Kigali♀

Amani♂

Makhali♂

HY3a

Exhibit

HY1a

HY3b

HY2a HY2bHY1b

HY4bHY4a

Hyena night-yards

Dec 10Sessions

(16)16 16 7 7 6 6 2 2

Dec 10As % 89 89 39 39 33 33 11 11

Jan 11Sessions

(12)10 11 6 5 4 4 8 8

Jan 11As %

92 92 50 42 33 33 67 67

As the results indicate in the table above, Piltengi and Gamba participated in the highest number of training sessions. This is because they are both housed singularly and not put onto exhibit as often as the other two groups, as a single animal is not as visible as a group (school holidays) and have a history of not reliably coming back into the night yards as required.

Bridge association: initially, a morsel of meat was offered to the individual (with tongs) simultaneously with a sharp toot of the training whistle. Every hyena grasped the ‘bridge’ concept after 3-5 repetitions in their first session.

Cues: hand signals and verbal cues given, also shaping using the reward to orient/manipulate body position (see ‘Table of training cues’)

Individual’s training progress for the month:Note: all behaviours commented on in the table below are at the preliminary stage and will be rated according to initial established response (i.e. 100% established/responsive for mouth open, duration of behaviour to be increased over time). The behavioural repertoire is growing all the time as the hyenas show/offer the trainers more and more in training sessions.

PiltengiBehaviour Progress Success

to date %Mouth open Grasped this behaviour almost straight away in the first session.

Working on duration of time mouth held open.100

Leg/thigh Knew this behaviour already & offered it almost every time without being cued at the start of the program. He will only offer the right side at this point.

100

Up Shaped the behaviour by encouraging him to follow the food reward (luring). Working on him responding to the given cues while slowly removing the ‘lure’

100

Stand front on

Due to offering the ‘leg’ behaviour without being cued almost constantly at the beginning, this behaviour was introduced to get him standing at a 90 degree angle to the trainer to break the association between what he was offering and what he was being trained (i.e. he would present his leg while being asked to open his mouth resulting in both occurring at the same time).

100

Box In order to add more to his behavioural repertoire, this behaviour (jump up onto the box) was added to give him more to do during sessions

100

Vocalise Often vocalises (giggles) when excited during sessions, especially when a food/large reward item is presented – capturing vocalisations and progressively rewarding for louder, longer vocalisations when cued

80

Target Introduced in last week of the month, he picked up the concept quickly although will try to bite the end of the target stick. Working on rewarding him for touches only.

100

Recalls in yard

Began training him with two trainers per session to keep it fresh and interesting for him – one trainer situated at one side of his yard and another on the opposite side – one trainer did a few behaviours at a time then the other trainer would call him and do a few behaviours with him – back and forth about 3-5 times during a session – he responds very enthusiastically to the added stimulation

100

Recalls from exhibit to yards

Due to his inconsistent/unreliable response to coming back into the yards from exhibit, he has not been let onto exhibit much as a result. To improve his exhibit to yard recall reliability so he can be let out onto exhibit more often, some gate training onto the exhibit was carried out. This one session strengthened the recall response rate dramatically.

100

Ear Learning to present his ears for topical applications (i.e. fly repellent) – new 25

behaviour introduced at end of monthComment He is very food motivated and eager to participate in sessions, is very animated

and fast to respond during training sessions & frequently vocalises excitedly – he learns new behaviours almost straight away in most cases. Piltengi responds well to behaviour sequences (i.e. ask for & bridge 3 behaviours in quick succession, then reward at end of sequence)

GambaBehaviour Progress Success

to date %Mouth open Took a few sessions to get the mouth open behaviour, but once he knew what

was expected, he offered the behaviour without hesitation (he seemed to graspthe concept more easily if was in the ‘up’ position when asked to open hismouth)

100

Leg/thigh Watched Piltengi do this and offered it in the first session! He will present this behaviour on both left and right sides

100

Up Shaped the behaviour by encouraging him to follow the food reward (luring). 100Sit Captured initially, but will sit when cued on most occasions 90-100Target Grasped the concept immediately – almost as if he knew the behaviour! 100Recalls He respond well to working between two trainers in one session as long as he is

given a bit of time to respond to the change – he tends to get confused by fast changes

100

Recalls from exhibit to yards

Due to his inconsistent/unreliable response to coming back into the yards from exhibit, he has not been let onto exhibit mush as a result. To improve his exhibit to yard recall reliability so he can be let out onto exhibit more often, some gate training onto the exhibit was carried out. His level of co-operation and relaxed behaviour was reinforced in the raceway also with great results. This one session strengthened the recall response rate dramatically.

100

Comment Gamba is very quick to learn new behaviours, he is relatively calm during sessions and he even learned a more advanced behaviour just by watching Piltengi in his first ever training session! He does get a bit confused if behaviours are asked for too quickly

KiganaBehaviour Progress Success

to date %Mouth open Working on getting him to offer wider mouth presentation 75Up Shaped the behaviour by encouraging him to follow the food reward (luring). 100Foot Offers foot up on mesh and will allow pad contact (both front feet) 90Stand front on

Leans into fence while offering foot, so trained to stand front on to separate this behaviour from other behaviours being trained

100

Leg Will present leg/thigh on right side 100Target Will offer this behaviour readily, working on getting him to touch the stick rather

than trying to bite it100

Separate Will readily separate from conspecifics to participate in training sessions 100Comment Very focussed during sessions, offers various positions to try to earn reward.

MwenkaBehaviour Progress Success

to date %Mouth open Working on getting him to offer wider mouth presentation 50Up Shaped the behaviour by encouraging him to follow the food reward (luring). 100Stand front on

Easily shaped this behaviour 100

Target Will offer this behaviour readily 100Separate Will readily separate from conspecifics to participate in training sessions 100Comment Not overly motivated during sessions, but is responsive

ForestBehaviour Progress Success

to date %Mouth open Working on getting her to offer wider mouth presentation 50Up Shaped the behaviour by encouraging her to follow the food reward (luring). 100

Target Introduced in last week of the month, she picked up the concept quickly although will try to bite the end of the target stick. Working on rewarding her for touches only.

Separate Will readily separate from conspecifics to participate in training sessions 100Comment A bit flighty during sessions, but over time it is anticipated she will become more

relaxed and focused.

KigaliBehaviour Progress Success

to date %Mouth open Working on getting her to offer wider mouth presentation 25Up Shaped the behaviour by encouraging her to follow the food reward (luring). 100Target Introduced in last week of the month, she picked up the concept quicklySeparate Will readily separate from conspecifics to participate in training sessions 100Comment Wild caught individual, very wary/nervous, will participate in sessions but, will

lose focus if too much activity is going on around her or she isn’t grasping the behaviour straight away. With more exposure to training sessions, it is anticipated that she will become a lot more relaxed and focused during sessions

AmaniBehaviour Progress Success

to date %Mouth open Introduced at end of month 25Up Shaped the behaviour by encouraging him to follow the food reward (luring). 100Separate Will readily separate from conspecifics to participate in training sessions 100Comment A sedate individual, learns slowly but steadily

MakhaliBehaviour Progress Success

to date %Mouth open Introduced at end of month 25Up Shaped the behaviour by encouraging him to follow the food reward (luring). 100Down Introduced at end of month 100Separate Will readily separate from conspecifics to participate in training sessions 100Comment Very bright, eager individual, learns quickly

Section items of note

For most of December the two cubs were being housed either with the two females or with males Mwenka and Kigana. This meant that the cubs were out on exhibit everyday and were not involved in the training as much as the other hyenas. Towards the end of December, it was decided that the cubs would then be housed with the two males permanently and could then be part of the training program more often.

Physical health of animals

There were no major health issues during the month of December.

Fly bites on ears were becoming a problem which was the reason we started to condition for ear presentation for topical treatment. Piltengi grasped this concept well from the start.

Issues affecting program

As often as possible, groups of people (tour members/visitors/other staff) were invited to watch a training session. This created a different environment and different distractions for the hyenas during their training sessions. All hyenas were very focussed despite the varying audience.

Table of training cues

Verbal cue Visual cue Response to cueUp Raise hand palm up parallel with ground

(with fingers together)Stands up against fence on back legs exposing underside/belly and paws of front feet

Leg Orient head position/direction with tongs/food and point to thigh, touch/poke thigh with finger/target stick

Presses body/thigh against fence line

Shoulder Orient head position/direction with tongs/food and point to shoulder, touch/poke shoulder with finger/target stick

Presses body/shoulder against fence line

Target Present target stick Touches end of target stick with noseOpen Make ‘L’ shape with thumb and forefinger Opens mouth wideDown Lower hand palm down parallel with ground

(with fingers together)Lowers body into sternal recumbency position

Sit Make a fist and raise fist up slightly Sits on tail end, front legs uprightFoot Point at (left or right front) foot Lifts foot and rests foot up on meshStraight Hand held vertically, fingers together,

fingers pointed toward hyenaStands with body at 90degrees to trainer

Say Show large enticing food morsel Excited chatter/giggling/vocalisationBox Point (and position body toward box) Jumps up onto boxEar Point to ear Leans ear toward trainer (against mesh)Come here (recall)

No cue other than trainer in position to train Approaches trainer

Appendix 2 shows physical changes to a female Spotted Hyena while pregnant with twins (Bennett 2014).

Appendix 3 shows a females torn cliteriform appendage of a proven breeder (Kalvas 2014).