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156-447: The park's grasslands support a variety of large mammals, including tens of thousands of blue wildebeest , whose annual migration is Africa's second-largest. Frequently sighted large predators include the cheetah , spotted hyena , and lion , the most famous of which was a female resident called Lady Liuwa, who was the subject of a National Geographic documentary ( The Last Lioness ) before she died of natural causes in 2017. Lady Liuwa

312-508: A pompom in appearance. Unusually among hyaenids, and mammals in general, the female spotted hyena is considerably larger than the male. Both sexes have a pair of anal glands which open into the rectum just inside the anal opening. These glands produce a white, creamy secretion which is pasted onto grass stalks by everting the rectum. The odour of this secretion is strong, smelling of boiling cheap soap or burning, and can be detected by humans several metres downwind. The spotted hyena has

468-500: A "tigerwolf", because the settlers in southern Africa did not know of hyenas, and thus labelled them as "wolves". Bosman and Kolben's descriptions went largely unnoticed until 1771, when the Welsh naturalist Thomas Pennant used the descriptions and his personal experience with a captive specimen as a basis for consistently differentiating the spotted hyena from the striped in his Synopsis of Quadrupeds . The description given by Pennant

624-779: A behaviour which does not occur when they target other prey species. One of the signs of social intelligence is the ability to have a keen olfactory sense or nasal recognition. Spotted hyenas have unique scent signatures that help them distinguish themselves from other clans and/or individuals (i.e., males or females conspecific), which enables them to mark their territories with secretions from their scent glands. Also, spotted hyenas have been recorded to utilise deceptive behaviour, including giving alarm calls during feeding when no enemies are present, thus frightening off other hyenas and allowing them to temporarily eat in peace. Similarly, mothers will emit alarm calls when attempting to interrupt attacks on their cubs by other hyenas. The spotted hyena

780-415: A body mass range of 56–182 kg (123–401 lb), with a mode of 102 kg (225 lb). When hunting medium to large sized prey, spotted hyenas tend to select certain categories of animal; young animals are frequently targeted, as are old ones, though the latter category is not so significant when hunting zebras, due to their aggressive anti-predator behaviours. Small prey is killed by being shaken in

936-624: A broad spatial scale. Because plant productivity is limited by grassland precipitation, carbon stocks are highest in regions where precipitation is heaviest, such as the high grass prairie in the humid temperate region of the United States. Similarly, as annual temperatures rise, grassland carbon stocks decrease due to increased evapotranspiration . Grasslands have suffered large losses of organic carbon due to soil disturbances, vegetation degradation, fires, erosion, nutrient deficiencies, and water shortages. The type, frequency and intensity of

1092-758: A continental climate favourable to the evolution of grasslands. Around 5 million years ago during the Late Miocene in the New World and the Pliocene in the Old World, the first true grasslands occurred. Existing forest biomes declined, and grasslands became much more widespread. It is known that grasslands have existed in Europe throughout the Pleistocene (the last 1.8 million years). Following

1248-428: A discount in biodiversity as faster-growing plants outcompete others. A study of a California grassland found that global change may speed reductions in diversity and forb species are most prone to this process. Misguided afforestation efforts, for example as part of the global effort to increase carbon sequestration, can harm grasslands and their core ecosystem services. Forest centric restoration efforts can create

1404-430: A dominant animal, subordinate spotted hyenas will walk on the knees of their forelegs in submission. Greeting ceremonies among clan-members consist of two individuals standing parallel to each other and facing opposite directions. Both individuals raise their hind legs and lick each other's anogenital area . During these greeting ceremonies, the penis or pseudo-penis often becomes erect, in both males and females. Erection

1560-455: A firm grip on it. The head is wide and flat with a blunt muzzle and broad rhinarium . In contrast to the striped hyena, the ears of the spotted hyena are rounded rather than pointed. Each foot has four digits, which are webbed and have short, stout and blunt claws. The paw-pads are broad and flat, with the whole undersurface of the foot around them being naked. The tail is relatively short, being 300–350 mm (12–14 in) long, and resembles

1716-713: A food reward, successfully cooperating and learning the maneuvers quickly without prior training. Experienced hyenas even helped inexperienced clan-mates to solve the problem. In contrast, chimps and other primates often require extensive training, and cooperation between individuals is not always as easy for them. The intelligence of the spotted hyena was attested to by Dutch colonists in 19th-century South Africa, who noted that hyenas were exceedingly cunning and suspicious, particularly after successfully escaping from traps. Spotted hyenas seem to plan on hunting specific species in advance; hyenas have been observed to engage in activities such as scent marking before setting off to hunt zebras,

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1872-472: A hunt, spotted hyenas often run through ungulate herds to select an individual to attack. Once selected, their prey is chased over a long distance, often several kilometres, at speeds of up to 60 kilometres per hour (37 mph). The spotted hyena has a long history of interaction with humanity ; depictions of the species exist from the Upper Paleolithic period, with carvings and paintings from

2028-612: A kill by lions, spotted hyenas will either leave or wait patiently at a distance of 30–100 metres until the lions have finished eating. In some cases, spotted hyenas are bold enough to feed alongside lions, and may occasionally force lions off a kill. This mostly occurs during the nighttime, when hyenas are bolder. Spotted hyenas usually prevail against groups of lionesses unaccompanied by males if they outnumber them 4:1. In some instances they were seen to have taken on and routed two pride males while outnumbering them 5:1. The two species may act aggressively toward one another even when there

2184-418: A large degree of morphological geographic variation, which led to an equally extensive set of specific and subspecific epithets. It was gradually realised that all of this variation could be applied to individual differences in a single subspecies. In 1939, biologist L Harrison Matthews demonstrated through comparisons between a large selection of spotted hyena skulls from Tanzania that all the variation seen in

2340-429: A luxury lodge and helicopter service, to make the park more accessible for tourists. The safari company worked with African Parks to fund the lodge, worth US$ 1.6 million. King Lewanika Lodge, named after the former Litunga, can accommodate fifteen guests and includes six villas, one of which has two bedrooms. In 2016, Proflight announced plans for regular flights between Lusaka and Kalabo , further improving access to

2496-471: A male. There have also been cases in which a clan has been led by a male rather than a female. Cubs take the rank directly below their mothers at birth. So when the matriarch passes away (or, in rare instances, disperses into another clan ), their youngest female cub will take over as matriarch. It is typical for females to remain with their natal clan, thus large clans usually contain several matrilines , whereas males typically disperse from their natal clan at

2652-485: A massive loss of lowland habitats favoured by spotted hyenas, and a corresponding increase in mixed woodlands . Spotted hyenas, under these circumstances, would have been outcompeted by wolves and humans which were as much at home in forests as in open lands, and in highlands as in lowlands. Spotted hyena populations began to shrink roughly 20,000 years ago, completely disappearing from Western Europe between 14 and 11,000 years ago, and earlier in some areas. Historically,

2808-505: A mate. Hyenas travelling in another clan's home range typically exhibit bodily postures associated with fear, particularly when meeting other hyenas. An intruder can be accepted into another clan after a long period of time if it persists in wandering into the clan's territory, dens or kills. The spotted hyena is a non-seasonal breeder, though a birth peak does occur during the wet season . Females are polyestrous, with an estrus period lasting two weeks. Like many feliform species,

2964-460: A member of the Lozi tribe who lived and died in the park, and a grandmother to several staff who were still working in the park, as of 2016. Lady Liuwa was first reported to be present in the park in 2002, and was said to have visited the woodlands often where Mambeti was buried. In 2008, after no lions returned to the park naturally, African Parks attempted to introduce a male lion, but he died during

3120-505: A necessity, as is the case for brown and striped hyenas, and thus they became pack hunters as well as scavengers. They began forming increasingly larger territories, necessitated by the fact that their prey was often migratory and long chases in a small territory would have caused them to encroach into another clan's land. It has been theorised that female dominance in spotted hyena clans could be an adaptation in order to successfully compete with males on kills, and thus ensure that enough milk

3276-478: A pack. Kafue National Park ) translocated to Liuwa in late 2021. Smaller omnivores in Liuwa include the banded mongoose and side-striped jackal . Since 2005, the protected area is considered a Lion Conservation Unit. Lady Liuwa was the park's most prominent resident, and was the subject of a National Geographic documentary released in 2010. According to folklore, the lioness was a reincarnation of Mambeti,

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3432-412: A piece of meat unnoticed, though they may be mobbed by the dogs in the attempt. When operating in groups, spotted hyenas are more successful in pirating dog kills, though the dog's greater tendency to assist each other puts them at an advantage against spotted hyenas, who rarely work in unison. Cases of dogs scavenging from spotted hyenas are rare. Although wild dog packs can easily repel solitary hyenas, on

3588-422: A proportionately large heart , constituting close to 1% of its body weight, thus giving it great endurance in long chases. In contrast, a lion's heart makes up only 0.45–0.57 percent of its body weight. The now extinct Eurasian populations were distinguished from the modern African populations by their shorter distal extremities and longer humerus and femur . The skull of the spotted hyena differs from that of

3744-425: A rabies vector, thus indicating that the species catches the disease from other animals rather than from intraspecifics. The microfilaria of Dipetalonema dracuneuloides have been recorded in spotted hyenas in northern Kenya. The species is known to carry at least three cestode species of the genus Taenia , none of which are harmful to humans. It also carries protozoan parasites of the genus Hepatozoon in

3900-522: A range of marketed and non-marketed ecosystem services that are fundamental to the livelihoods of an estimated one billion people globally. Grasslands hold about twenty percent of global soil carbon stocks. Herbaceous (non-wooded) vegetation dominates grasslands and carbon is stored in the roots and soil underground. Above-ground biomass carbon is relatively short-lived due to grazing, fire, and senescence . Grassland species have an extensive fibrous root system, with grasses often accounting for 60-80% of

4056-489: A result of human activity. Hunting cultures around the world often set regular fires to maintain and extend grasslands and prevent fire-intolerant trees and shrubs from taking hold. The tallgrass prairies in the U.S. Midwest may have been extended eastward into Illinois , Indiana , and Ohio by human agency. Much grassland in northwest Europe developed after the Neolithic Period when people gradually cleared

4212-558: A subspecies of Crocuta crocuta , known as cave hyenas , roamed across Eurasia for at least one million years until the end of the Late Pleistocene . The spotted hyena is the largest extant member of the Hyaenidae, and is further physically distinguished from other species by its vaguely bear -like build, rounded ears, less prominent mane, spotted pelt, more dual-purposed dentition, fewer nipples , and pseudo-penis . It

4368-559: A symbiotic relationship between spotted hyenas and their scent gland bacteria, where the differences in fatty acids can be attributed to fermentation by different microbes. Clan boundaries are usually respected; hyenas chasing prey have been observed to stop dead in their tracks once their prey crosses into another clan's range. Hyenas will however ignore clan boundaries in times of food shortage. Males are more likely to enter another clan's territory than females are, as they are less attached to their natal group and will leave it when in search of

4524-576: Is nepotistic ; the offspring of dominant females automatically outrank adult females subordinate to their mother. However, rank in spotted hyena cubs is greatly dependent on the presence of the mother; low-ranking adults may act aggressively toward higher-ranking cubs when the mother is absent. Although individual spotted hyenas only care for their own young, and males take no part in raising their young, cubs are able to identify relatives as distantly related as great-aunts. Also, males associate more closely with their own daughters rather than unrelated cubs, and

4680-528: Is 12 years, with a maximum of 25 years. The clan's social life revolves around a communal den. While some clans may use particular den sites for years, others may use several different dens within a year or several den sites simultaneously. Spotted hyena dens can have more than a dozen entrances, and are mostly located on flat ground. The tunnels are usually oval in section, being wider than they are high, and narrow down from an entrance width of ½–1 metre (1.6–7.7 ft) to as small as 25 cm (9.8 in). In

4836-478: Is a hyena species, currently classed as the sole extant member of the genus Crocuta , native to sub-Saharan Africa . It is listed as being of least concern by the IUCN due to its widespread range and large numbers estimated between 27,000 and 47,000 individuals. The species is, however, experiencing declines outside of protected areas due to habitat loss and poaching. Populations of Crocuta , usually considered

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4992-631: Is a highly successful animal, being the most common large carnivore in Africa. Its success is due in part to its adaptability and opportunism ; it is primarily a hunter but may also scavenge , with the capacity to eat and digest skin, bone and other animal waste. In functional terms, the spotted hyena makes the most efficient use of animal matter of all African carnivores. The spotted hyena displays greater plasticity in its hunting and foraging behaviour than other African carnivores; it hunts alone, in small parties of 2–5 individuals or in large groups. During

5148-508: Is a lack of agreement on the amount of carbon that can be stored in grassland ecosystem. This is partly caused by different methodologies applied to measure soil organic carbon and limited respective datasets. Further, carbon accumulation in soils changes significantly over time and point in time measurements produce an insufficient evidence base. Grasslands are among the most threatened ecosystems. Global losses from grassland degradation are estimated to be over $ 7 billion per year. According to

5304-468: Is an area where the vegetation is dominated by grasses ( Poaceae ). However, sedge ( Cyperaceae ) and rush ( Juncaceae ) can also be found along with variable proportions of legumes , like clover , and other herbs . Grasslands occur naturally on all continents except Antarctica and are found in most ecoregions of the Earth . Furthermore, grasslands are one of the largest biomes on Earth and dominate

5460-424: Is at least one record of a young adult male leopard dying from a sepsis infection caused by wounds inflicted by a spotted hyena. Spotted hyenas will follow packs of African wild dogs to appropriate their kills. They will typically inspect areas where wild dogs have rested and eat any food remains they find. When approaching wild dogs at a kill, solitary hyenas will approach cautiously and attempt to take off with

5616-496: Is contracted by consuming already infected herbivores, rather than through direct infection from tsetse flies . It is known that adult spotted hyenas in the Serengeti have antibodies against rabies , canine herpes , canine brucellosis , canine parvovirus , feline calicivirus , leptospirosis , bovine brucellosis , rinderpest and anaplasmosis. During the canine distemper outbreak of 1993–94, molecular studies indicated that

5772-545: Is derived from the Sanskrit koṭṭhâraka , which in turn originates from kroshṭuka (both of which were originally meant to signify the golden jackal ). The earliest recorded mention of Κροκόττας is from Strabo 's Geographica , where the animal is described as a mix of wolf and dog native to Ethiopia. From Classical antiquity until the Renaissance , the spotted and striped hyena were either assumed to be

5928-751: Is home to a variety of mammals, including buffalo, common eland , common tsessebe , oribi , red lechwe , reedbuck , roan antelope , and migrating blue wildebeest , which gather in the tens of thousands. Liuwa's wildebeest migration is the second-largest in Africa. A survey conducted in 1991 recorded population estimates of 30,000 blue wildebeest, 800 tsessebe, 1,000 zebra, and 10,000 other large mammals, including buffalo, eland, oribi, red lechwe, reedbuck, and sitatunga . Subsequent surveys suggested major population declines, with possible eradication of buffalo, eland, Lichtenstein's hartebeest, and roan antelope. However, improved protections since 2003 has stabilized populations. Eland and buffalo have been reintroduced, and

6084-526: Is no food at stake. Lions may charge at hyenas and maul them for no apparent reason; one male lion was filmed killing two hyenas on separate occasions without eating them, and lion predation can account for up to 71% of hyena deaths in Etosha . Spotted hyenas have adapted to this pressure by frequently mobbing lions which enter their territories. Experiments on captive spotted hyenas revealed that specimens with no prior experience with lions act indifferently to

6240-513: Is not uncommon to see up to 20 cubs at a single site. The general form of a spotted hyena den is tunnel-shaped, with a spacious end chamber used for sleeping or breeding. This chamber measures up to 2 metres (6.6 feet) in width, the height being rather less. Females generally give birth at the communal den or a private birth den. The latter is primarily used by low status females to maintain continual access to their cubs, as well as ensure that they become acquainted with their cubs before transferral to

6396-627: Is often a plagioclimax ; it remains dominant in a particular area usually due to grazing , cutting, or natural or man-made fires, all discouraging colonization by and survival of tree and shrub seedlings . Some of the world's largest expanses of grassland are found in the African savanna, and these are maintained by wild herbivores as well as by nomadic pastoralists and their cattle , sheep or goats. Grasslands have an impact on climate change by slower decomposition rates of litter compared to forest environments. Grasslands may occur naturally or as

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6552-453: Is particularly apparent in same sexed litters, and can result in the death of the weaker cub. This neonatal siblicide kills an estimated 25% of all hyenas in their first month. Male cubs which survive grow faster and are likelier to achieve reproductive dominance, while female survivors eliminate rivals for dominance in their natal clan. Lactating females can carry 3–4 kg (6.6–8.8 lb) of milk in their udders . Spotted hyena milk has

6708-477: Is produced for their cubs. Another theory is that it is an adaptation to the length of time it takes for cubs to develop their massive skulls and jaws, thus necessitating greater attention and dominating behaviours from females. Both Björn Kurtén and Camille Arambourg promoted an Asiatic origin for the species; Kurtén focussed his arguments on the Plio-Pleistocene taxon Crocuta sivalensis from

6864-556: Is scarce or absent in tropical rainforests and coastal areas. Its preferred habitats in west Africa include the Guinea and Sudan savannahs, and is absent in the belt of dense coastal forest. In the Namib Desert , it occurs in riverine growth along seasonal rivers, the sub-desertic pro-Namib and the adjoining inland plateau. In ideal habitats, the spotted hyena outnumbers other large carnivores, including other hyena species. However,

7020-553: Is still controversial. A study in Brazilian Subtropical Highland Grasslands found that grasslands without traditional land management—which uses fire every two years and extensive cattle grazing—can disappear within 30 years. This study showed that grasslands inside protected areas , in which fire is not allowed and cattle grazing is banned, grasslands were quickly replaced by shrubs ( shrub encroachment ). Land cover has always changed during

7176-439: Is superimposed on the back and hind quarters. The spots, which are of variable distinction, may be reddish, deep brown or almost blackish. The spots vary in size, even on single individuals, but are commonly 20 mm (0.79 in) in diameter. A less distinct spot pattern is present on the legs and belly but not on the throat and chest. Some research groups (such as Ngorongoro Crater Hyena Project and MSU Hyena Project ) often use

7332-729: Is the largest extant member of the Hyaenidae. Adults measure 95–165.8 cm (37.4–65.3 in) in body length, and have a shoulder height of 70–91.5 cm (27.6–36.0 in). Adult male spotted hyenas in the Serengeti weigh 40.5–55.0 kg (89.3–121.3 lb), while females weigh 44.5–63.9 kg (98–141 lb). Spotted hyenas in Zambia tend to be heavier, with males weighing on average 67.6 kg (149 lb), and females 69.2 kg (153 lb). Exceptionally large weights of 81.7 kg (180 lb) and 90 kg (200 lb) are known. It has been estimated that adult members of

7488-606: Is the most carnivorous member of the Hyaenidae. Unlike its brown and striped cousins, the spotted hyena is primarily a predator; this has been shown since the 1960s. One of the earliest studies to demonstrate their hunting abilities was done by wildlife ecologist Hans Kruuk, and he showed through a 7-year study of hyena populations in the Serengeti and Ngorongoro Crater that spotted hyenas hunt as much as lions, with later studies showing this tendency in many other areas of Africa. There are however large differences between populations, depending on their habitat, available prey species and

7644-602: Is the only placental mammalian species where females have a pseudo-penis and lack an external vaginal opening. The spotted hyena is the most social of the Carnivora in that it has the largest group sizes and most complex social behaviours. Its social organisation is unlike that of any other carnivore, bearing closer resemblance to that of cercopithecine primates ( baboons and macaques ) with respect to group size, hierarchical structure, and frequency of social interaction among both kin and unrelated group-mates. However,

7800-473: Is the other way around, and lions will readily steal the kills of spotted hyenas. In the Ngorongoro Crater , it is common for lions to subsist largely on kills stolen from hyenas. Lions are quick to follow the calls of hyenas feeding, a fact demonstrated by field experiments, during which lions repeatedly approached whenever the tape-recorded calls of hyenas feeding were played. When confronted on

7956-427: Is traversed to its tip by a central urogenital canal, through which the female urinates, copulates and gives birth. The pseudo-penis can be distinguished from the males' genitalia by its somewhat shorter length, greater thickness, and more rounded glans . In both males and females, the base of the glans is covered with penile spines . The formation of the pseudo-penis appears largely androgen independent, as

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8112-580: Is usually a sign of submission, rather than dominance, and is more common in males than in females. It is said that feasting Hyaenas engage in violent fights, and there is such a croaking, shrieking and laughing at such times that a superstitious person might really think all the inhabitants of the infernal regions had been let loose. The spotted hyena has an extensive vocal range, with sounds ranging from whoops, fast whoops, grunts, groans, lows, giggles, yells, growls, soft grunt-laughs, loud grunt-laughs, whines and soft squeals. The loud who-oop call, along with

8268-671: The Dutch West India Company at the Gold Coast (modern-day Ghana ) from 1688 to 1701, wrote of Jakhals, of Boshond (jackals or woodland dogs) whose physical descriptions match the spotted hyena. Kolben, a German mathematician and astronomer who worked for the Dutch East India Company in the Cape of Good Hope from 1705 to 1713, described the spotted hyena in great detail, but referred to it as

8424-918: The Everglades of Florida , the Pantanal of Brazil , Bolivia and Paraguay or the Esteros del Ibera in Argentina , are classified with flooded savannas as the flooded grasslands and savannas biome and occur mostly in the tropics and subtropics. The species that live in these grasslands are well adapted to the hydrologic regimes and soil conditions. The Everglades—the world's largest rain-fed flooded grassland—is rich in 11,000 species of seed-bearing plants, 25 species of orchids , 300 bird species, and 150 fish species. Water-meadows are grasslands that are deliberately flooded for short periods. High-altitude grasslands located on high mountain ranges around

8580-477: The Lascaux and Chauvet Caves . The species has a largely negative reputation in both Western culture and African folklore . In the former, the species is mostly regarded as ugly and cowardly, while in the latter, it is viewed as greedy, gluttonous, stupid, and foolish, yet powerful and potentially dangerous. The majority of Western perceptions on the species can be found in the writings of Aristotle and Pliny

8736-463: The Pleistocene ice ages (with their glacials and interglacials ), grasslands expanded in the hotter, drier climates, and began to become the dominant land feature worldwide. Since the grasslands have existed for over 1.8 million years, there is high variability. For example steppe-tundra dominated in Northern and Central Europe whereas a higher amount of xerothermic grasslands occurred in

8892-630: The Russian Far East , and it has been theorised that the presence of hyenas there may have delayed the colonisation of North America . During the Last Glacial Maximum , the spotted hyena also roamed Southeast Asia . The causes of the species' extinction in Eurasia are still largely unknown. In Western Europe at least, the spotted hyena's extinction coincided with a decline in grasslands 12,500 years ago. Europe experienced

9048-522: The Siwaliks , a view defended by Arambourg, who nonetheless allowed the possibility of an Indo-Ethiopian origin. This stance was contested by Ficarelli and Torre, who referred to evidence of the spotted hyena's presence from African deposits dating from the early Pleistocene, a similar age to the Asian C. sivalensis . Modern scholarship supports an African origin for the genus, with the earliest fossils of

9204-671: The Transvaal Lowveld areas of South Africa . During the 1770s and 1780s the species was still widespread in southern and western South Africa, being recorded on the Cape Peninsula and Cape Flats , and near present-day Somerset West , Riviersonderend , Mossel Bay , George , Joubertina , Gamtoos River , Jansenville , Cannon Rocks , Alice , Onseepkans and Augrabies Falls . The species dwells in semi- deserts , savannah , open woodland , dense dry woodland, and mountainous forests up to 4,000 m in altitude. It

9360-446: The brown bear in bone-crushing ability, and free ranging hyenas have been observed to crack open the long bones of giraffes measuring 7 cm in diameter. A 63.1 kg (139 lb) spotted hyena is estimated to have a bite force of 565.7 newtons at the canine tip and 985.5 newtons at the carnassial eocene. One individual in a study was found to exert a bite force of 4,500 newtons on the measuring instruments. The spotted hyena

9516-480: The encroachment of woody species . Species richness is particularly high in grasslands of low soil fertility such as serpentine barrens and calcareous grasslands, where woody encroachment is prevented as low nutrient levels in the soil may inhibit the growth of forest and shrub species. Another common predicament often experienced by the ill-fated grassland creatures is the constant burning of plants, fueled by oxygen and many expired photosynthesizing organisms, with

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9672-871: The prairie and Pacific grasslands of North America , the Pampas of Argentina , Brazil and Uruguay , calcareous downland , and the steppes of Europe . They are classified with temperate savannas and shrublands as the temperate grasslands, savannas, and shrublands biome . Temperate grasslands are the home to many large herbivores , such as bison , gazelles , zebras , rhinoceroses , and wild horses . Carnivores like lions , wolves , cheetahs and leopards are also found in temperate grasslands. Other animals of this region include deer , prairie dogs , mice , jack rabbits , skunks , coyotes , snakes , foxes , owls , badgers , blackbirds, grasshoppers , meadowlarks , sparrows , quails , hawks and hyenas . Grasslands that are flooded seasonally or year-round, like

9828-561: The slaty egret ), the grey heron , pygmy geese , the spur-winged goose , and the three-banded plover . The black-winged pratincole , Denham's bustard , long-tailed widowbird , pink-billed lark , rosy-throated longclaw , secretary bird , sharp-tailed starling , swamp boubou , white-bellied bustard , and white-cheeked bee-eater are also present, as are clapper larks . Liuwa Plain National Park also provides habitat for various snake species. Grassland A grassland

9984-509: The tropical and subtropical grasslands, savannas and shrublands biome . The rainfall level for that grassland type is between 90 and 150 centimeters per year. Grasses and scattered trees are common for that ecoregion, as well as large mammals , such as wildebeest ( Connochaetes taurinus ) and zebra ( Equus zebra ). Notable tropical and subtropical grasslands include the Llanos grasslands of South America . Mid-latitude grasslands, including

10140-778: The African savannas or the Iberian deheza. As flowering plants and trees, grasses grow in great concentrations in climates where annual rainfall ranges between 500 and 900 mm (20 and 35 in). The root systems of perennial grasses and forbs form complex mats that hold the soil in place. Grasslands support the greatest aggregations of large animals on Earth, including jaguars, African wild dogs, pronghorn , black-footed ferret , plains bison , mountain plover , African elephant, Sunda tiger, black rhino, white rhino, savanna elephant, greater one-horned rhino, Indian elephant and swift fox . Grazing animals, herd animals, and predators in grasslands, like lions and cheetahs live in

10296-510: The American West—and introduction of invasive species , like cane toads in northern Australia, have disrupted the balance in these ecosystems and damaged a number of other species. Grasslands are home to a number of the foremost magnificent animals on the planet—elephants, bison, lions—and hunters have found them to be enticing prey. But when hunting is not controlled or is conducted illegally, species can become extinct. Grasslands provide

10452-508: The Elder , though in relatively unjudgmental form. Explicit, negative judgments occur in the Physiologus , where the animal is depicted as a hermaphrodite and grave-robber . The IUCN's hyena specialist group identifies the spotted hyena's negative reputation as detrimental to the species' continued survival, both in captivity and the wild. The spotted hyena's scientific name Crocuta

10608-539: The International Union for the Conservation of Nature (IUCN), the most significant threat to grasslands is human land use, especially agriculture and mining. The vulnerability of grasslands stems from a range of factors, such as misclassification, poor protection and cultivation. Grasslands have an extensive history of human activity and disturbance . To feed a growing human population, most of

10764-559: The Kalahari, where the brown outnumber the spotted. The two species typically encounter each other on carcasses, which the larger spotted species usually appropriate. Sometimes, brown hyenas will stand their ground and raise their manes while emitting growls. This usually has the effect of seemingly confusing spotted hyenas, which will act bewildered, though they will occasionally attack and maul their smaller cousins. Similar interactions have been recorded between spotted and striped hyenas in

10920-556: The Mediterranean area. Within temperate Europe, the range of types is quite wide and also became unique due to the exchange of species and genetic material between different biomes. The semi-natural grasslands first appeared when humans started farming. So for the use of agriculture, forests got cleared in Europe. Ancient meadows and pastures were the parts that were suitable for cultivation. The semi-natural grasslands were formed from these areas. However, there's also evidence for

11076-606: The Mushingashi Conservancy, the Zambia Carnivore Programme, and Zambia's Department of National Parks and Wildlife (DPNW) introduced another male lion to Liuwa Plain from Kafue National Park. The lion bonded with another in a boma for two months, and were then released. Lady Liuwa died of natural causes on August 9, 2017. 334 bird species, including various species of birds of prey , bustards , cranes (including grey crowned cranes and

11232-643: The Serengeti, Kenya and South Africa. Spotted hyenas may act as hosts in the life-cycles of various parasites which start life in herbivores; Taenia hyaenae and T. olnogojinae occur in hyenas in their adult phase. Trichinella spiralis are found as cysts in hyena muscles. The spotted hyena's distribution once ranged in Europe from the Iberian Peninsula to the Urals , where it remained for at least one million years. Remains have also been found in

11388-490: The Serengeti, where prey is migratory, clans are smaller than those in the Ngorongoro Crater, where prey is sedentary. Spotted hyena clans are more compact and unified than wolf packs, but are not as closely knit as those of African wild dogs . Females usually dominate males, including in cases where low-ranking females generally dominate over high-ranking males, but they will also occasionally co-dominate with

11544-417: The Serengeti. Though they readily take to water to catch and store prey, spotted hyenas will avoid crocodile-inhabited waters, and usually keep a safe distance from Nile crocodiles . Recent observations shows that African rock pythons can hunt adult spotted hyenas. Spotted hyenas have a complex set of postures in communication. When afraid, the ears are folded flat, and are often combined with baring of

11700-403: The addition of preferring males with whom they have had long and friendly prior relationships. Passive males tend to have greater success in courting females than aggressive ones. Copulation in spotted hyenas is a relatively short affair, lasting 4–12 minutes, and typically only occurs at night with no other hyenas present. The mating process is complicated, as the male's penis enters and exits

11856-487: The age of 2½ years. Höner et al. say that when a male co-dominates with a female or is otherwise able to lead, this is because the male was born to the matriarch of the clan and has taken the rank directly below his mother. The clan is a fission–fusion society , in which clan-members do not often remain together, but may forage in small groups. High-ranking hyenas maintain their position through aggression directed against lower-ranking clan-members. Spotted hyena hierarchy

12012-441: The age of one month. Within ten days of birth, they are able to move at considerable speed. Cubs begin to lose the black coat and develop the spotted, lighter coloured pelage of the adults at 2–3 months. They begin to exhibit hunting behaviours at the age of eight months, and will begin fully participating in group hunts after their first year. Spotted hyenas reach sexual maturity at the age of three years. The average lifespan in zoos

12168-506: The age of the earliest Crocuta specimens in Eurasia, which are around 2  million years ago from China. The nuclear genome results also suggest that the European and Asian populations were distinct from each other, but were more closely related to each other overall than to African Crocuta populations. Analysis of the nuclear genome suggests that there had been interbreeding between African and Eurasian populations for some time after

12324-461: The area. Sepo was captured, helicoptered back, and placed in a boma with Lady Liuwa. The pair bonded and were released after two months. After mating with Nakawa, Sepo gave birth to one male and two female cubs in December 2013. Nakawa was later killed, possibly from being poisoned. An unidentified lion was seen in the park in 2015. In September 2016, a collaborative project between African Parks,

12480-572: The atmosphere). It can have severe negative consequences on key ecosystem services, like land productivity and groundwater recharge. Despite growing recognition of the importance of grasslands, understanding of restoration options remains limited. Cost of grassland restoration is highly variable and respective data is scarce. Successful grassland restoration has several dimensions, including recognition in policy, standardisation of indicators of degradation, scientific innovation, knowledge transfer and data sharing. Restoration methods and measures include

12636-478: The biomass carbon in this ecosystem. This underground biomass can extend several meters below the surface and store abundant carbon into the soil, resulting in deep, fertile soils with high organic matter content. For this reason, soil carbon accounts for about 81% of the total ecosystem carbon in grasslands. The close link between soil carbon and underground biomass leads to similar responses of these carbon pools to fluctuations in annual precipitation and temperature on

12792-489: The communal den. Compared to other hyenas, the spotted hyena shows a greater relative amount of frontal cortex which is involved in the mediation of social behavior. Studies strongly suggest convergent evolution in spotted hyena and primate intelligence . A study done by evolutionary anthropologists demonstrated that spotted hyenas outperform chimpanzees on cooperative problem-solving tests; captive pairs of spotted hyenas were challenged to tug two ropes in unison to earn

12948-483: The den, consisting of small underground channels, is likely an effective anti-predator device which protects cubs from predation during the absence of the mother. Spotted hyenas rarely dig their own dens, having been observed for the most part to use the abandoned burrows of warthogs, springhares and jackals. Faeces are usually deposited 20 metres (66 feet) away from the den, though they urinate wherever they happen to be. Dens are used mostly by several females at once, and it

13104-417: The disturbance can play a key role in the soil organic carbon ( SOC ) balance of grasslands. Bedrock , irrigation practices, soil acidification , liming , and pasture management can all have potential impacts on grassland organic carbon stocks. Good grassland management can reverse historical soil carbon losses. The relationship of improved biodiversity with carbon storage is subject of research. There

13260-501: The economics that the world's grasslands have to offer, from producing grazing animals, tourism, ecosystems services such as clean water and air, and energy extraction. Vast areas of grassland are affected by woody encroachment , which is the expansion of woody plants at the expense of the herbaceous layer. Woody encroachment is caused by a combination of human impact (e.g. fire exclusion, overstocking and resulting overgrazing ) and environmental factors (i.e. increased CO 2 levels in

13416-409: The endangered wattled crane ), pelicans , pratincoles , and storks , have been recorded in Liuwa. Raptors include the bateleur , greater kestrel , martial eagle , palm-nut vulture , and Pel's fishing owl , as well as African fish eagles . Recorded water birds include the marabou , open-billed, saddle-billed , and yellow-billed stork , as well as the blacksmith lapwing , egrets (including

13572-595: The exact taxonomic nature of the hyena family in general, with most European travelers in Ethiopia referring to hyenas as "wolves". This partly stems from the Amharic word for hyena, ጅብ ( jɨbb ), which is linked to the Arabic word ذئب ( dhiʾb ) 'wolf'. The first detailed first-hand descriptions of the spotted hyena by Europeans come from Willem Bosman and Peter Kolbe . Bosman, a Dutch tradesman who worked for

13728-423: The eyes, the area around the rhinarium, the lips and the back portion of the chin are all blackish. The limbs are spotted, though the feet vary in colour, from light brown to blackish. The fur is relatively sparse and consists of two hair types; moderately fine underfur (measuring 15–20 mm (0.59–0.79 in)) and long, stout bristle hairs (30–40 mm (1.2–1.6 in)). European Paleolithic rock art depicting

13884-410: The female's reproductive tract through her pseudo-penis rather than directly through the vagina, which is blocked by the false scrotum and testes. These unusual traits make mating more laborious for the male than in other mammals, and also make forced copulation physically impossible. The female retracts her clitoris before the male's penis enters it by sliding beneath it, an operation facilitated by

14040-804: The following: For the period 2021–2030 the United Nations General Assembly has proclaimed the UN Decade on Restoration, involving a joint resolution by over 70 countries. It is led by the United Nations Environment Programme and the Food and Agriculture Organization . Grassland types by Schimper (1898, 1903): Grassland types by Ellenberg and Mueller-Dombois (1967): Formation-class V. Terrestrial herbaceous communities Grassland types by Laycock (1979): These grasslands can be classified as

14196-463: The forest to create areas for raising their livestock. Grasslands often occur in areas with annual precipitation is between 600 mm (24 in) and 1,500 mm (59 in) and average mean annual temperatures ranges from −5 and 20 °C. However, some grasslands occur in colder (−20 °C) and hotter (30 °C) climatic conditions. Grassland can exist in habitats that are frequently disturbed by grazing or fire, as such disturbance prevents

14352-439: The gazelle fawning season, as jackals and wolves are effective at tracking and catching young animals. Hyenas do not take to eating wolf flesh readily; four hyenas were reported to take half an hour in eating a golden wolf. Overall, the two animals typically ignore each other when there is no food or young at stake. Spotted hyenas dominate other hyena species wherever their ranges overlap. Brown hyenas encounter spotted hyenas in

14508-542: The genus Crocuta dating to the early Pliocene of Africa, around 3.63 to 3.85 million years ago. The earliest known fossil record of Crocuta crocuta in Africa is from the Cradle of Humankind , specifically from the Sterkfontein Member 4 which is dated approximately 2.1  million years ago in minimum, and it is also common in deposits dated between 2 to 1.5 million years ago . Analysis of

14664-513: The genus Crocuta still lacking. The ancestors of the genus Crocuta diverged from Hyaena (the genus of striped and brown hyenas ) 10 million years ago. The ancestors of the spotted hyena probably developed social behaviours in response to increased pressure from other predators on carcasses, which forced them to operate in teams. At one point in their evolution , spotted hyenas developed sharp carnassials behind their crushing premolars; this rendered waiting for their prey to die no longer

14820-419: The giant short-faced hyena. As there is no evidence of environmental change being responsible, it is likely that the giant short-faced hyena became extinct due to competition with the spotted hyena. The spotted hyena has a strong and well-developed neck and forequarters, but relatively underdeveloped hindquarters. The rump is rounded rather than angular, which prevents attackers coming from behind from getting

14976-759: The grassland areas have been turned to arable fields and disappeared again. The grasslands permanently became arable cropping fields due to the steady decrease in organic matter. Nowadays, semi-natural grasslands are rather located in areas that are unsuitable for agricultural farming. Grasslands dominated by unsown wild-plant communities ("unimproved grasslands") can be called either natural or "semi-natural" habitat. Although their plant communities are natural, their maintenance depends upon anthropogenic activities such as grazing and cutting regimes. The semi-natural grasslands contain many species of wild plants, including grasses, sedges, rushes, and herbs; 25 plant-species per 100 square centimeters can be found. A European record that

15132-612: The grassland type and on how strong it is affected by human impact. Dominant trees for the semi-natural grassland are Quercus robur , Betula pendula , Corylus avellana , Crataegus and many kinds of herbs. In chalk grassland , the plants can vary from very tall to very short. Quite tall grasses can be found in North American tallgrass prairie , South American grasslands, and African savanna . Woody plants, shrubs or trees may occur on some grasslands—forming savannas, scrubby grassland or semi-wooded grassland, such as

15288-443: The grasslands of the African savanna. Mites , insect larvae , nematodes , and earthworms inhabit deep soil, which can reach 6 metres (20 feet) underground in undisturbed grasslands on the richest soils of the world. These invertebrates, along with symbiotic fungi , extend the root systems, break apart hard soil, enrich it with urea and other natural fertilizers, trap minerals and water and promote growth. Some types of fungi make

15444-488: The highest protein and fat content of any terrestrial carnivore. Cubs will nurse from their mother for 12–16 months, though they can process solid food as early as three months. Mothers do not regurgitate food for their young. Females are protective of their cubs, and will not tolerate other adults, particularly males, approaching them. Spotted hyenas exhibit adult behaviours early in life; cubs have been observed to ritually sniff each other and mark their living space before

15600-840: The lack of rain pushing this problem to further heights. When not limited by other factors, increasing CO 2 concentration in the air increases plant growth, similarly as water use efficiency, which is very important in drier regions. However, the advantages of elevated CO 2 are limited by factors including water availability and available nutrients , particularly nitrogen. Thus effects of elevated CO 2 on plant growth will vary with local climate patterns, species adaptations to water limitations, and nitrogen availability. Studies indicate that nutrient depletion may happen faster in drier regions, and with factors like plant community composition and grazing. Nitrogen deposition from air pollutants and increased mineralization from higher temperatures can increase plant productivity, but increases are often among

15756-448: The land is also important, as it is then easier to fertilize, for example. For instance, if it is located near a road. With the development of technology, it is becoming increasingly easy to cultivate land with a steeper gradient, to the detriment of grasslands. The management of grasslands is also changing permanently. There is increased use of mineral fertilizers, furthermore borders and field edges are removed to enlarge fields and leveling

15912-437: The land, 49.7%, was covered with forest and there was also more semi-natural grassland (18.8%) than arable land (15.8%). In 2015 this has changed drastically. The forest cover has increased (50.8%) and arable land has also increased (20.4%), but the semi-natural grassland cover has decreased. Although it still covers a large area of the earth (10.6%). A quarter of semi-natural grassland was lost through intensification, i.e. it

16068-545: The landscape change due to agriculture of the last century. The original wild-plant communities having been replaced by sown monocultures of cultivated varieties of grasses and clovers, such as perennial ryegrass and white clover . In many parts of the world, "unimproved" grasslands are one of the most threatened types of habitat, and a target for acquisition by wildlife conservation groups or for special grants to landowners who are encouraged to manage them appropriately. Grassland vegetation can vary considerably depending on

16224-569: The landscape worldwide. There are different types of grasslands: natural grasslands, semi-natural grasslands, and agricultural grasslands. They cover 31–69% of the Earth's land area. Included among the variety of definitions for grasslands are: Semi-natural grasslands are a very common subcategory of the grasslands biome. These can be defined as: They can also be described as the following: There are many different types of semi-natural grasslands, e.g. hay meadows . The graminoids are among

16380-766: The latter favor their fathers by acting less aggressively toward them. Spotted hyena societies are more complex than those of other carnivorous mammals, and are remarkably similar to those of cercopithecine primates in respect to group size, structure, competition and cooperation. Like cercopithecine primates, spotted hyenas use multiple sensory modalities, recognise individual conspecifics, are conscious that some clan-mates may be more reliable than others, recognise third-party kin and rank relationships among clan-mates, and adaptively use this knowledge during social decision making. Also, like cercopithecine primates, dominance ranks in hyena societies are not correlated with size or aggression, but with ally networks. In this latter trait,

16536-408: The local persistence of natural grasslands in Europe, originally maintained by wild herbivores, throughout the pre-Neolithic Holocene. The removal of the plants by the grazing animals and later the mowing farmers led to co-existence of other plant species around. In the following, the biodiversity of the plants evolve. Also, the species that already lived there adapted to the new conditions. Most of

16692-573: The maniacal laughter, are among the most recognisable sounds of Africa. Typically, high-pitched calls indicate fear or submission, while loud, lower-pitched calls express aggression. The pitch of the laugh indicates the hyena's age, while variations in the frequency of notes used when hyenas make noises convey information about the animal's social rank. Dr. Hans Kruuk compiled the following table on spotted hyena calls in 1972: Spotted hyenas may contract brucellosis , rinderpest and anaplasmosis . They are vulnerable to Trypanosoma congolense , which

16848-551: The mitochondrial genomes of Eurasian Crocuta ( cave hyena ) specimens shows no clear separation from African lineages. It also suggests that modern spotted hyena populations descend from a widely ranging Afro-Eurasian population that was only recently reduced in range to Africa exclusively. However, analysis of the full nuclear genome suggests that Late Pleistocene African and Eurasian Crocuta populations were largely separate, having estimated to have diverged from each other around 2.5  million years ago , closely corresponding to

17004-544: The most versatile life forms . They became widespread toward the end of the Cretaceous period, and coprolites of fossilized dinosaur feces have been found containing phytoliths of a variety of grasses that include grasses that are related to modern rice and bamboo . The appearance of mountains in the western United States during the Miocene and Pliocene epochs, a period of some 25 million years, created

17160-470: The mouth, while large prey is eaten alive. Where spotted hyenas and lions occupy the same geographic area, the two species occupy the same ecological niche, and are thus in direct competition with one another. In some cases, the extent of dietary overlap can be as high as 68.8%. Lions typically ignore spotted hyenas, unless they are on a kill or are being harassed by them. There exists a common misconception that hyenas steal kills from lions, but most often it

17316-642: The national park was established, the area served as the hunting ground for Lubosi Lewanika (1842–1916), who was the Litunga (king or paramount chief ) of the Lozi people in Barotseland between 1878 and 1916. Lubosi Lewanika designated Liuwa Plain as a protected area in the early 1880s. National park designation was granted in 1972, and Zambia's government took over management. Despite gaining such status, increased human pressure led to an increase in poaching in

17472-436: The now extinct Eurasian populations weighed 102 kg (225 lb). Fur colour varies greatly and changes with age. Unlike the fur of the striped and brown hyena, that of the spotted hyena consists of spots rather than stripes and is much shorter, lacking the well defined spinal mane of the former two species. The base colour generally is a pale greyish-brown or yellowish-grey on which an irregular pattern of roundish spots

17628-614: The original diversity of plants having been destroyed by cultivation and by the use of fertilizers. Almost 90% of the European semi-natural grasslands do not exist anymore due to political and economic reasons. This loss took place during the 20th century. The ones in Western and Central Europe have almost disappeared completely. There are a few left in Northern Europe. Unfortunately, a large amount of red-listed species are specialists of semi-natural grasslands and are affected by

17784-564: The park in 2002. The organization has since led the introductions of several additional lions to reestablish a breeding pride in Liuwa, where there are 16 lions, as of 2022. In an effort to reestablish the species to the park, 11 African wild dogs (eight males from South Africa and three females from Kafue National Park ) were translocated to the park in late 2021. The animals were kept in a split compartment boma to bond, after which they were released in April 2022 and started to immediately move as

17940-723: The park. Liuwa Plain lies within the Barotse Floodplain , and is bounded by the Luambimba River to the north and Luanginga River to the south. The park is prominently made up of a grassland that measures approximately 72 by 32 km (45 by 20 mi), scattered with raffia palms and woodlands . Recorded grass species include Echinochloa stagnina and Vossia cuspidata , which are important for grazing herbivores, as well as Baikiaea plurijuga Guibourtia coleosperma , Peltophorum africanum , Terminalia sericea , and various types of Hyphaene . Liuwa

18096-739: The park. African Parks has managed the park in partnership with the Department of National Parks and Wildlife (DNPW) and the Barotse Royal Establishment (BRE) since 2003. Local interest in preserving the park and its wildlife has reportedly subsequently increased since management has improved and the area's connection to the Litunga was restored. Tourism to the park was limited until recently, with only 50 tourists visiting Liuwa in 2000, and less than 800 people reportedly visiting in 2014. African Parks and Norman Carr Safaris opened

18252-517: The penis's upward angle. The hyenas then adopt a typical mammalian mating posture and usually lick their genitals for several minutes after mating. Copulation may be repeated multiple times during a period of several hours. The length of the gestation period tends to vary greatly, but the average length is 110 days. In the final stages of pregnancy, dominant females provide their developing offspring with higher androgen levels than lower-ranking mothers do. The higher androgen levels –

18408-445: The plants more resistant to insect and microbial attacks. Grassland in all its form supports a vast variety of mammals, reptiles, birds, and insects. Typical large mammals include the blue wildebeest , American bison , giant anteater , and Przewalski's horse . The plants and animals that live in grasslands are connected through an unlimited web of interactions. But the removal of key species—such as buffalo and prairie dogs within

18564-427: The position of which allows it to crush bone with its premolars without blunting the carnassials. Combined with large jaw muscles and a special vaulting to protect the skull against large forces, these characteristics give the spotted hyena a powerful bite which can exert a pressure of 80  kgf/cm2 (1140 lbf/in² ), which is 40% more force than a leopard can generate. The jaws of the spotted hyena outmatch those of

18720-588: The presence of other predators. Thus the estimated contribution from hunting to their diet ranges from at least half up to 98%, the remainder being from scavenging. Wildebeest are the most commonly taken medium-sized ungulate prey item in both Ngorongoro and the Serengeti, with zebra and Thomson's gazelles , coming close behind. Cape buffalo are rarely attacked due to differences in habitat preference, though adult bulls have been recorded to be taken on occasion. In Kruger National Park, blue wildebeest , cape buffalo, plains zebra , greater kudu and impala are

18876-562: The pseudo-penis appears in the female fetus before differentiation of the fetal ovary and adrenal gland. When flaccid, the pseudo-penis is retracted into the abdomen, and only the prepuce is visible. After giving birth , the pseudo-penis is stretched, and loses many of its original aspects; it becomes a slack-walled and reduced prepuce with an enlarged orifice with split lips. Spotted hyenas are social animals that live in large communities (referred to as " clans ") which can consist of at most 80 individuals. Group-size varies geographically; in

19032-618: The raising of young. Giving birth is difficult for female hyenas, as the females give birth through their narrow clitoris, and spotted hyena cubs are the largest carnivoran young relative to their mothers' weight. During parturition, the clitoris ruptures to facilitate the passage of the young, and may take weeks to heal. Cubs are born with soft, dark brown hair, and weigh 1.5 kg on average. Unique among carnivorous mammals, spotted hyenas are also born with their eyes open and with 6–7 mm long canine teeth and 4 mm long incisors. Also, cubs will attack each other shortly after birth. This

19188-461: The relocation. Two other males were reintroduced in 2009. Both of them reportedly mated with Lady Liuwa, but she was infertile. The two male lions left Liuwa and made their way to Angola; one was killed by locals and the other, Nakawa, returned to the park on his own. Two lionesses were relocated from Kafue National Park to Liuwa in 2011; one of them was killed by trapping in 2012, and the other, Sepo, had to be returned to Liuwa after migrating out of

19344-506: The result of high concentrations of ovarian androstenedione  – are thought to be responsible for the extreme masculinization of female behavior and morphology. This has the effect of rendering the cubs of dominant females more aggressive and sexually active than those of lower ranking hyenas; high ranking male cubs will attempt to mount females earlier than lower ranking males. The average litter consists of two cubs, with three occasionally being reported. Males take no part in

19500-774: The risk of misreading and misclassifying of landscapes. A map created by the World Resources Institute in collaboration with the IUCN identifies 2 billion hectares for potential forest restoration . It is criticised for including 900 million hectares of grasslands. It is expected that non-native grasses will continue to outperform native species under warmer and drier conditions that occur in many grasslands due to climate change. The type of land management used in grasslands can also lead to grassland loss/degradation. Many grasslands and other open ecosystems depend on disturbances such as wildfires , controlled burns and/or grazing to persist, although this subject

19656-402: The rocky areas of East Africa and Congo, spotted hyenas use caves as dens, while those in the Serengeti use kopjes as resting areas in daylight hours. Dens have large bare patches around their entrances, where hyenas move or lie down on. Because of their size, adult hyenas are incapable of using the full extent of their burrows, as most tunnels are dug by cubs or smaller animals. The structure of

19812-447: The same clan, differences in scent gland compositions can help individuals differentiate the reproductive states and sex of their members. One study analyzed the scent gland secretions of several hyenas belonging to three different clans and discovered a high degree of similarity in fatty acid composition among members of the same clans, while hyenas of different clans had more dissimilar scent gland secretions. Further studies have proposed

19968-460: The same species, or distinguished purely on geographical, rather than physical, grounds. Hiob Ludolf , in his Historia aethiopica , was the first to clearly distinguish the Crocuta from Hyaena on account of physical and geographical grounds, though he never had any first hand experience of the species, having gotten his accounts from an Ethiopian intermediary. Confusion still persisted over

20124-504: The sight of them, but will react fearfully to the scent. Although cheetahs and leopards preferentially prey on smaller animals than those hunted by spotted hyenas, hyenas will steal their kills when the opportunity presents itself. Cheetahs are usually easily intimidated by hyenas, and put up little resistance, while leopards, particularly males, may stand up to hyenas. There are records of some male leopards preying on hyenas. Hyenas are nonetheless dangerous opponents for leopards; there

20280-443: The social system of the spotted hyena is openly competitive rather than cooperative, with access to kills, mating opportunities and the time of dispersal for males depending on the ability to dominate other clan -members. Females provide only for their own cubs rather than assist each other, and males display no paternal care. Spotted hyena society is matriarchal ; females are larger than males, and dominate them. The spotted hyena

20436-473: The species indicates that the Eurasian populations retained the spots of their modern-day African counterparts. The genitalia of the female closely resembles that of the male; the clitoris is shaped and positioned like a penis, a pseudo-penis , and is capable of erection . The female also possesses no external vagina (vaginal opening), as the labia are fused to form a pseudo- scrotum . The pseudo-penis

20592-454: The split, which likely explains the discordance between the nuclear and mitochondrial genome results, with the mitochondrial genomes of African and European Crocuta more closely related to each other than to Asian Crocuta, suggesting gene flow between the two groups after the split between the Asian and European populations. Its appearance in Europe and China coincided with the decline and eventual extinction of Pachycrocuta brevirostris ,

20748-410: The spot patterns to help identify individual hyenas. A set of five, pale and barely distinct bands replace the spots on the back and sides of the neck. A broad, medial band is present on the back of the neck, and is lengthened into a forward facing crest. The crest is mostly reddish-brown in colour. The crown and upper part of the face is brownish, save for a white band above both eyes, though the front of

20904-415: The spotted hyena further show parallels with primates by acquiring rank through coalition . However, rank reversals and overthrows in spotted hyena clans are rare. The social network dynamics of spotted hyenas are determined by multiple factors. Environmental factors include rainfall and prey abundance; individual factors include preference to bond with females and with kin; and topological effects include

21060-411: The spotted hyena is promiscuous , and no enduring pair bonds are formed. Members of both sexes may copulate with several mates over the course of several years. Males will show submissive behaviour when approaching females in heat, even if the male outweighs his partner. Females usually favour younger males born or joined into the clan after they were born. Older females show a similar preference, with

21216-516: The spotted hyena was widespread throughout Sub-Saharan Africa . It is present in all habitats save for the most extreme desert conditions, tropical rainforests and the top of alpine mountains. Its current distribution is patchy in many places, especially in West Africa. Populations are concentrated in protected areas and surrounding land. There is a continuous distribution over large areas of Ethiopia , Kenya , Tanzania , Botswana , Namibia and

21372-615: The spotted hyena's most important prey, while giraffe , impala, wildebeest and zebra are its major food sources in the nearby Timbavati area. Springbok and kudu are the main prey in Namibia 's Etosha National Park , and springbok in the Namib . In the southern Kalahari , gemsbok , common eland , lechwe , waterbuck , wildebeest and springbok are the principal prey. In Chobe , the spotted hyena's primary prey consists of migratory zebra and resident impala. In Kenya 's Masai Mara , 80% of

21528-618: The spotted hyena's prey consists of topi and Thomson's gazelle, save for during the four-month period when zebra and wildebeest herds migrate to the area. Bushbuck , suni and buffalo are the dominant prey items in the Aberdare Mountains , while Grant's gazelle , gerenuk , sheep , goats and cattle are likely preyed upon in northern Kenya. Unlike other large African carnivores, spotted hyenas do not preferentially prey on any species, and only African buffalo and giraffe are significantly avoided. Spotted hyenas prefer prey with

21684-560: The striped hyena by its much greater size and narrower sagittal crest . For its size, the spotted hyena has one of the most powerfully built skulls among the Carnivora. The dentition is more dual purposed than that of other modern hyena species, which are mostly scavengers; the upper and lower third premolars are conical bone-crushers, with a third bone-holding cone jutting from the lower fourth premolar. The spotted hyena also has its carnassials situated behind its bone-crushing premolars,

21840-424: The tail is curled below the belly. During an attack, or when excited, the tail is carried forward on the back. An erect tail does not always accompany a hostile encounter, as it has also been observed to occur when a harmless social interaction occurs. Although they do not wag their tails, spotted hyenas will flick their tails when approaching dominant animals or when there is a slight tendency to flee. When approaching

21996-411: The teeth and a flattening of the mane. When attacked by other hyenas or by wild dogs, the hyena lowers its hindquarters. Before and during an assertive attack, the head is held high with the ears cocked, mouth closed, mane erect and the hindquarters high. The tail usually hangs down when neutral, though it will change position according to the situation. When a high tendency to flee an attacker is apparent,

22152-743: The tendency to close triads in the network. Female hyenas are more flexible than males in their social bonding preferences. Higher ranking adult spotted hyenas tend to have higher telomere length and higher levels of some immune defense proteins in their blood serum. Territory size is highly variable, ranging from less than 40 km in the Ngorongoro Crater to over 1,000 km in the Kalahari. Home ranges are defended through vocal displays, scent marking and boundary patrols. Clans mark their territories by either pasting or pawing in special latrines located on clan range boundaries. Spotted hyenas use scent gland secretions to distinguish between members of their own clan and members of neighboring clans. Within

22308-413: The terrain to facilitate the use of agricultural machinery. The professional study of dry grasslands falls under the category of rangeland management , which focuses on ecosystem services associated with the grass-dominated arid and semi-arid rangelands of the world. Rangelands account for an estimated 70% of the earth's landmass; thus, many cultures including those of the United States are indebted to

22464-530: The then recognised subspecies could also be found in a single population, with the only set of characters standing out being pelage (which is subject to a high degree of individual variation) and size (which is subject to Bergmann's Rule ). When fossils are taken into consideration, the species displayed even greater variation than it does in modern times, and a number of these named fossil species have since been classed as synonymous with Crocuta crocuta , with firm evidence of there being more than one species within

22620-461: The viruses isolated from hyenas and lions were more closely related to each other than to the closest canine distemper virus in dogs. Evidence of canine distemper in spotted hyenas has also been recorded in the Masai Mara. Exposure to rabies does not cause clinical symptoms or affect individual survival or longevity. Analyses of several hyena saliva samples showed that the species is unlikely to be

22776-412: The whole, the relationship between the two species is a one sided benefit for the hyenas, with wild dog densities being negatively correlated with high hyena populations. Medium-sized canids like Black-backed and side-striped jackals , and African wolves will feed alongside hyenas, though they will be chased if they approach too closely. Spotted hyenas will sometimes follow jackals and wolves during

22932-593: The wild-plant diversity of the "unimproved" grasslands is usually a rich invertebrate fauna; there are also many species of birds that are grassland "specialists", such as the snipe and the little bustard . Owing to semi-natural grasslands being referred to as one of the most-species rich ecosystems in the world and essential habitat for many specialists, also including pollinators, there are many approaches to conservation activities lately. Agriculturally improved grasslands, which dominate modern intensive agricultural landscapes, are usually poor in wild plant species due to

23088-467: The world's grasslands are converted from natural landscapes to fields of corn, wheat or other crops. Grasslands that have remained largely intact thus far, such as the East African savannas , are in danger of being lost to agriculture. Grasslands are very sensitive to disturbances, such as people hunting and killing key species, or plowing the land to make more space for farms. Grassland vegetation

23244-528: The world, like the Páramo of the Andes Mountains . They are part of the montane grasslands and shrublands biome and can be tropical, subtropical, and temperate. The plants and animals, that can be found in the tropical montane, are able to adapt to cool, wet conditions as well as intense sunlight. Spotted hyena The spotted hyena ( Crocuta crocuta ), also known as the laughing hyena ,

23400-483: The years. The following relates to the changes between 1960 and 2015. There has been a decrease in semi-natural grasslands and an increase in areas with arable land , forest and land used for infrastructure and buildings. The line style and relative thickness of the lines indicates the percentage of the total area that changed. Changes less than 1% and land-cover classes with all changes less than 1% (i.e. semi-natural wetlands and water) are not included. In 1960 most of

23556-409: The zebra population has increased to over 4,000 individuals. Predators include the cheetah , leopard , lion , African wild dog and spotted hyena . According to the nonprofit conservation organization African Parks , all but one of the park's lions were eradicated during the 1990s due to poaching and trophy hunting . Liuwa's lone lioness, known as Lady Liuwa, was first reported to be present in

23712-412: Was converted into arable or pasture land and forests. It is more likely that intensification will occur in flat semi-natural grasslands, especially if the soil is fertile. On the other hand, grasslands, where the land is drought-prone or less productive, are more likely to persist as semi-natural grasslands than grasslands with fertile soil and low gradient of the terrain. Furthermore, the accessibility of

23868-627: Was found on a meadow in Estonia described 76 species of plants in one square meter. Chalk downlands in England can support over 40 species per square meter. In many parts of the world, few examples have escaped agricultural improvement (fertilizing, weed killing, plowing, or re-seeding). For example, original North American prairie grasslands or lowland wildflower meadows in the UK are now rare and their associated wild flora equally threatened. Associated with

24024-537: Was once widely thought to be derived from the Latin loanword crocutus , which translates as 'saffron-coloured one', in reference to the animal's fur colour. This was proven to be incorrect, as the correct spelling of the loanword would have been Crocāta , and the word was never used in that sense by Graeco-Roman sources. Crocuta actually comes from the Ancient Greek word Κροκόττας ( krokottas ), which

24180-489: Was precise enough to be included by Johann Erxleben in his Systema regni animalis by simply translating Pennant's text into Latin. Crocuta was finally recognised as a separate genus from Hyaena in 1828. Unlike the striped hyena, for which a number of subspecies were proposed in light of its extensive modern range, the spotted hyena is a genuinely variable species, both temporally and spatially. Its range once encompassed almost all of Africa and Eurasia , and displayed

24336-423: Was the only remaining lion in the area, following years of excess hunting, prior to African Parks' assuming management and introducing additional lions to encourage the re-establishment of a pride. More than 300 bird species have been recorded in Liuwa, which has experienced limited tourism until recently. Animal populations have since stabilized, despite declines and local extinctions during the 1990s–2000s. Before

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