Striped Hyaena
Hyaena (Hyaena) hyaena (Linnaeus, 1758)
Based on Heribert Hofer, in Status Survey and Conservation Action Plan - Hyaenas

  DISTRIBUTION
PHYSICAL DESCRIPTION AND SUBSPECIES
HABITAT
DIET AND FORAGING BEHAVIOR
REPRODUCTION AND DENNING BEHAVIOR
SOCIAL BEHAVIOR
COMPETITION
THREATS
CONSERVATION STATUS OF GLOBAL POPULATION AND SUBPOPULATIONS - CONSERVATION MEASURES

  

Physical Description and Subspecies
This medium-sized, dog-like animal has a back sloping downwards towards the tail, and black vertical stripes on the sides. Its general colour is pale grey or beige. It has a black patch on the throat, five to nine more or less distinct vertical stripes on the flanks and clearer black transverse and horizontal stripes on the fore and hind legs. The head is roundish with a pointed muzzle and pointed ears. It has a mane along the mid-dorsal line which can be held erect. Its black and white tail is long and bushy, with hair that is generally coarse and long. The feet have four toes and short, blunt, non-retractable claws.

Five subspecies are distinguished, mainly by their differences in size and pelage, although this classification is provisional:
H. h. barbara from northwest Africa
H. h. dubbah from northeast Africa
H. h. sultana from Arabia
H. h. syriaca from Syria, Asia Minor and the Caucasus
H. h. hyaena from India
H. h. sultana from the Arabian peninsula 

Body mass varies between 26 and 41kg for males and 26 and 34kg for females. Total body length excluding tail varies between 1.0 and 1.15m and shoulder height between 0.66-0.75m. Amongst the provisional subspecies, body mass and body size are only well studied in H. h. syriaca in Israel (Mendelssohn 1985, Mendelssohn and Yom-Tov 1988). 

Habitat
In most of its range the striped hyaena occurs in open habitat or light thorn bush country. In North Africa it prefers open woodlands and bushy and mountainous regions. Both the centre of the Arabian desert and the Sahara are avoided (Rieger 1979a). In central Asia it also avoids high altitudes and dense thickets and forests (Heptner and Sludskij 1980). The maximum altitudes recorded are 2,250m in Iran, 2,500m in India (sources in Rieger 1979a) and 3,300m in Pakistan (Roberts 1977). In the Caucasus region, Turkmenistan, Tadzhikistan, and Uzbekistan, prime habitats include savannah and semi-desert regions up to an altitude of 2,100m, mountain areas with a strong relief and valleys and slopes (even with little or no vegetation) with plenty of caves or other resting sites and riverine areas. Other preferred habitats are thickets of tamarisks, the periphery of sand deserts, and the special pistachio (Pistacia vera) savannahs characteristic of the Badhyz area of southeast Turkmenistan (Heptner and Sludskij 1980). Because of its limited ability to thermoregulate, the striped hyaena stays south of the January isotherms of 1 �C, and avoids areas with minimum temperatures of less than -15 to -20�C and more than 80-120 days of frost per year (Heptner and Slodskij 1980). 

In Israel it is present even close to dense human settlements. Individuals have recently been recorded 19km south of Tel Aviv, 5km east of the international airport and on the Tel Aviv-Haifa highway near Mount Carmel (Mendelssohn 1985, Mendelssohn and Yom-Tov 1988). In India it used to be common in open country especially where low hills and ravines were available. (Prater 1948). 

In west Africa the striped hyaena occurs in the Sahel and Sudan savannas. 

Diet and Foraging Behavior
The striped hyaena scavenges carrion and the remains of kills of other predators (wolf, spotted hyaena, cheetah, leopard, lion, tiger). It also consumes a wide variety of vertebrates, invertebrates, vegetables and fruits, including the fruits of Balanites trees, and human-associated organic matter (Kruuk 1976, Rieger 1979a, Heptner and Sludskij 1980, Osborn and Helmy 1980, Kerbis-Peterhans and Horwitz 1992). The massive cheek teeth and supporting musculature easily permit the gnawing and breaking of bones and carapaces. The striped hyaena may also kill smaller vertebrates including livestock (see section on Damage to agriculture and livestock, below). 

The proportion of scavenged and killed prey items is still a matter of debate as there are no detailed studies on the diet of the striped hyaena. Rieger (1979a) suggests that only individuals from the three larger subspecies H. h. barbara, H. h. syriaca and H. h. hyaena (Middle East, Asia minor, central Asia and the Indian subcontinent, and North Africa) kill larger prey animals including livestock, as there is no evidence that the smaller subspecies H. h. dubbah and H. h. sultana (east Africa and Arabian peninsula) attack larger herbivores. In Turkmenistan it has been recorded to feed on wild boar, kulan, porcupine, and particularly tortoises. In Uzbekhistan and Tadzhikistan, seasonal abundance of oil willow fruits (Eleagnus angustifolia) is an important contribution to their diet, while in the Caucasus region it is grasshoppers (Heptner and Sludskij 1980). In Israel it feeds on garbage, carrion, and fruits, particularly dates and melons (Macdonald 1978, Mendelssohn 1985, Mendelssohn and Yora-Tov 1988). In eastern Jordan near the Azraq oasis, the main sources of food are carcasses of feral horses and water buffalo, and refuse from local villages (Al Younis 1993). The striped hyaena is able to drink water of very variable quality, from freshwater to soda and salt water, but it may also fulfil its water requirements with melons (Heptner and Sludskij 1980). 

Foraging behaviour
Less is known about the sensory capacities, prey location and hunting behaviour of the striped hyaena than of either the spotted or brown hyaena. Seasonal influxes of striped hyaenas accompanying migrations of large herds of domestic and wild ungulates in Turkmenistan suggest that it may cover long distances on foraging trips, or at least part of the time live a nomadic existence in this region (Heptner and Sludskij 1980). In Egypt it is known to move along ancient caravan roads where the chance of locating a dead camel is high (Osborn and Helmy 1980). In the Serengeti, the greater part of its nocturnal activity is spent searching for food or moving between established foraging sites. It covers a total of 7-27km (mean 19km) per night, either following established animal tracks or zig-zagging cross-country (Kruuk 1976). While walking at a speed of two to four km/h (Kruuk 1976), it stops to investigate the bases of tree trunks, dense shrubs, clumps of grass, old holes, etc. It is apparently able to memorise the location of fruiting trees, garbage dumps and other established feeding sites. It is able to locate tortoises in their hiding places during periods of aestivation and hibernation (Kullman 1965, Gaisler et al. 1968); one hyaena was observed locating and digging out three tortoises in two and a half hours in one night (Heptner and Sludskij 1980). Observed hunts were a simple chase and grab procedure (Kruuk 1976). Food storage is practised commonly; the relevant food item may be stored in tall bushy or marshy clumps or at the base of dense shrubby vegetation (Kruuk 1976). 

Damage to Agriculture and Livestock
Summary of the information from the Status Survey and Conservation Action Plan questionnaires on damage to agricultural produce and livestock killed by the striped hyaena suggests that goats, sheep, dogs, and poultry are the most commonly recorded items. Larger animals are also occasionally reputed to be killed, although the possibility cannot be excluded that cases of scavenging were mistakenly identified as kills. In most cases of damage to larger livestock it is unclear whether the targeted individual was adult or young, healthy or sick. The records suggest that attacks typically occur at low frequencies. Exceptions of more frequent livestock damage are reputed to occur in Egypt, Ethiopia, India, Iraq, and possibly Morocco. 

In Turkmenistan the striped hyaena is known to kill dogs, whereas in the Caucasus region it is reported to kill dogs, sheep and other small domestic animals (Heptner and Sludskij 1980). In Iraq reports from the 1950s indicate that the striped hyaena may attack horses and donkeys (Hatt 1959). In Africa, dogs, sheep and goats are occasionally at risk (Ronnefeld 1969). Records of attacks on sheep and goats originate from North Africa, Israel, Iran, Pakistan, and India, on donkeys from North Africa, Israel, Iran, Pakistan, and India, on horses in Iran, and on dogs in India (Roberts 1977, Rieger 1979a, Johnson 1987). Older records of attacks on sheep and goats also come from the Sinai and Somalia (Osborn and Helmy 1980). 

The striped hyaena also occasionally causes damage to melon fields and to date palms in date plantations in Israel (H. Mendelssohn unpublished data) and Egypt (Osborn and Helmy 1980), and to water and honey melon plantations in Turkmenistan (Heptner and Sludskij 1980). 

Reproduction and Denning Behaviour
In the wild, litter size varies from one to four (median of three) throughout the year, after a gestation period of 90-91 days (Pocock 1941, Ronnefeld 1969, Heptner and Sludskij 1980).. Average litter size in captivity is 2.4, with a range of one to five (Rieger 1979a). Parturition is preceded by intensive digging behaviour by the female and often followed by a one-day post-partum oestrus three weeks later (Rieger 1981). 

Cubs are born blind, with closed ear tubes and white to grey fur with clear black stripes. Eyes first open after seven to eight days, and teeth erupt from day 21 onwards. Cubs begin to eat meat at the age of 30 days (Rieger 1979a). Weaning in captivity takes place after eight weeks (Heptner and Sludskij 1980). In the wild cubs have been observed suckling until four to five months of age (Rieger 1981), or up to 10-12 months (Kruuk 1976). Both the male and female bring food to the cubs (Kruuk 1976, Davidar 1985, 1990). 

Various ages of sexual maturity have been reported. A striped hyaena was four years old when she gave birth to her first litter in the zoo of Tashkent (Heptner and Sludskij 1980), but most females mature by the age of two to three years in other zoos (Rieger 1979a). Mendelssohn (1985) reported three free-living individuals in Israel of approximately 15 months of age with three large embryos.

The striped hyaena prefers to den in caves. Den entrances are fairly narrow and may be hidden by large boulders. Measurements of two dens in the Karakum desert yielded a width of 0.67m and 0.72m for the entrance. The dens lead 3m and 2.5m down and extended over a distance of 4.15m and 5m. There were no lateral extensions or special chambers (Heptner and Sludskij 1980). These simple constructions contrast with much more elaborate designs exceeding 27m in length discovered in Israel (Kerbis-Peterhans and Horwitz 1992). 

Social Behavior
Rieger (1979a, 1981) has argued that across subspecies, differences in body size, proportion of killed prey items in the diet, and group sizes (sociality) co-vary. The two smaller subspecies, H. h. dubbah and H. h. sultana, formerly sympatric with the spotted hyaena, are supposed to be more solitary and are not known to kill larger wild or domestic herbivores. The larger subspecies H. h. syriaca, H. h. hyaena and H. h. barbara, however, kill larger herbivores and have been repeatedly observed in small groups. Current information is inadequate to test this data. 

Typical group sizes are one or two in all subspecies (Rieger 1979a), although groups of up to seven animals have been reported in H. h. dubbah in Libya (Hufnagi 1972). In Israel, H. h. syriaca is generally solitary, but occasionally several are seen together at a carcass, apparently males and females, or females and large cubs (Macdonald 1978). H. h. syriaca has been recorded as monogamous in central Asia (Heptner and Sludskij 1980). 

Home range sizes of one female and one male in the Serengeti were 44km² and 72km² respectively, with little evidence of territorial behaviour (Kruuk 1976). Van Aarde et al. (1988) calculated a home-range size for a single female in the Negev desert in Israel to be approximately 61km² over a period of seven months, which partly overlapped with two other individuals. 

When striped hyaenas fight they bite at the throat and legs, not the mane. The mane serves as a signalling device during social interactions. During meetings, striped hyaenas investigate and lick the mid-back region where the mid-dorsal crest is situated. Greetings also involve sniffing of the nose and extruded anal pouch, and repeated pawing of the throat of the greeting partner (Fox 1971, Rieger 1978, Macdonald 1978). In aggressive encounters, the black patch near the thoracic and lumbar vertebrae is erected (Rieger 1979a). The striped hyaena scent marks (pastes) on grass stalks, stones, tree trunks and other objects, with secretions from the anal pouch (Kruuk 1976). The striped hyaena uses a smaller variety of calls than the spotted hyaena (Kruuk 1976, Peters 1984). 

Competition
In Israel the striped hyaena may encounter wolves, red foxes and caracals at carcasses. On a one-to-one basis it is dominant over the wolf, but a group of four wolves has been observed driving a single hyaena from a carcass (H. Mendelssohn unpublished data). A caracal may drive a subadult striped hyaena away from a carcass (Skinner and Ilani 1979). Competitors in central Asia include leopards, wolves, golden jackals, red and corsac foxes and vultures (Heptner and Sludskij 1980). The striped hyaena frequently scavenges from kills of tiger, leopard, cheetah, and wolf- a major component of the striped hyaena's diet in central Asia are scavenged carcasses killed by wolves (Heptner and Sludskij 1980, Lukarevsky 1988). In India, the striped hyaena usually wins one-to-one encounters over carcasses with leopards, tiger cubs and domestic dogs but may be dominated by adult tigers (observations in Action Plan questionnaires, Pocock 1941, Rieger 1979 and references therein). In east Africa, the striped hyaena is dominated by the spotted hyaena and sometimes the leopard, yet in turn it may dominate the leopard and the domestic dog (Kruuk 1976). When attacked by domestic dogs or dug out by humans, the striped hyaena may use "shamming", i.e. the animal pretends to be dead, even if repeatedly bitten (Pocock 1941, Heptner and Sludskij 1980). 

Threats
A bounty system operating in Algeria during the 1880s contributed to a decline in population size; in 1881 and 1882 alone, 196 individuals were killed (Kowalski and Rzebik-Kowalska 1991). In the Arabian peninsula, the majority of museum specimens were collected dead, hanging in trees and on sign posts (Gasperetti et al. 1985).

The striped hyaena appears to be very susceptible to poisoning as it readily accepts strychnine-poisoned bait. In many cases it is not the target, as the bait is laid out for other carnivores such as wolves or leopards suspected of killing livestock, or because wolves are wanted by fur trappers in central Asia (Heptner and Sludskij 1980). Along the Mediterranean coast in Israel, the striped hyaena was exterminated by strychnine poisoning during the rabies eradication campaign administered by the British government between 1918 and 1948. The striped hyaenas ate poisoned donkey carcasses that were provided to control golden jackals, then the main carrier of rabies. Further large-scale poisoning occurred here between 1950 and 1970. Today, large-scale reduction by strychnine poisoning also threatens the striped hyaena throughout Niger (Millington and Tiega 1990, 1991). 

In the Caucasus and in central Asia, a major source of mortality over the past 100 years has been persecution, as the striped hyaena was held responsible for the disappearance of unattended small children. In the 1880s alone the striped hyaena was held responsible for the kidnapping or injuring (biting) of 25 children and three adults who slept outside in the district of Jerewan in the Caucasus. The government paid a substantial bounty (100 rubles) for every hyaena killed. Further cases of striped hyaenas killing or kidnapping children in this area were reported in the 1890s and 1900s, as well as in Azerbaidjan in the 1930s and 1940s (Heptner and Sludskij 1980). Today in India, the government still organises killings of wolves and striped hyaenas (even in conservation areas) in places where carnivores are suspected of child lifting. In recent times this has happened in Karnataka, Bihar state. Attacks on children have been reported as recently as 1974 when 19 children up to the age of four years were reported killed at night (Rieger 1979a). 

Striped hyaenas have rarely been hunted for their fur (although this has occured in the Caucasus countries), but have incidentally been caught in traps set by fur trappers for other species. In Russia, the striped hyaena was not even considered a fur species but was bought and sold as "minor quality wolf and "fox". Nevertheless, in the areas covered by the Commonwealth of Independent States (i.e. the Caucasus region, Tadzhikistan, Turkmenistan and Uzbekistan) a total of 200 skins were bought by the government in the 1930s. In the 1950s less than 100 were bought and none have been bought since 1970. In Turkmenistan alone between 1931 and 1937 up to 130 skins were offered by trappers every year. However, since 1948 this number has been reduced to a few dozen and since 1970 none have been offered (Heptner and Sludskij 1980). 

Conservation Status of Global Population and Subpopulations - Conservation Measures
Questionnaire surveys in the Status Survey and Conservation Action Plan and an evaluation of published information suggest that the striped hyaena is already extinct in many localities and that populations are generally declining throughout its range. The major reasons for this decline are decreasing natural and domestic sources of carrion due to declines in the populations of other large carnivores (wolf, cheetah, leopard, lion, tiger) and their prey, and changes in livestock practices. Moreover, the low densities and associated large home ranges are likely to increase the chances of fragmentation of populations into small, non-viable units. This must be considered a key problem for the future. The striped hyaena evokes many superstitious fears because of reputed and documented cases of injuries to humans sleeping outside, snatching and killing of children, and grave robbery. In addition, it is widely exploited as an aphrodisiac, utilised for traditional healing, and killed because of suspected or real damage inflicted on agricultural produce and livestock. The striped hyaena has been widely hunted through poisoning, baiting traps, pits, or with the help of dogs. A tentative estimate of the total worldwide population size is 5,000 to 14,000 individuals. 

The upper estimate of the global population size of the striped hyaena exceeds 10,000 individuals. However, fragmentation of the world population into many subpopulations is suspected even though the actual degree of fragmentation is unknown. In addition, a degree of habitat loss and population decline is taking place at an unknown rate, and the minimum population estimate is less than 10,000 individuals. We therefore agree with the status of striped hyaena as Lower Risk: near threatened (IUCN, 2000).