|Range map of hippopotamus. Historic range is in red while current range is in green.|
The common hippopotamus (Hippopotamus amphibius), or hippo, is a large, mostly herbivorous mammal in sub-Saharan Africa, and one of only two extant species in the family Hippopotamidae, the other being the pygmy hippopotamus (Choeropsis liberiensis or Hexaprotodon liberiensis). The name comes from the ancient Greek for "river horse" (ἱπποπόταμος). After the elephant and rhinoceros, the common hippopotamus is the third-largest type of land mammal and the heaviest extant artiodactyl. Despite their physical resemblance to pigs and other terrestrial even-toed ungulates, the closest living relatives of the Hippopotamidae are cetaceans (whales, porpoises, etc.) from which they diverged about . The common ancestor of whales and hippos split from other even-toed ungulates around . The earliest known hippopotamus fossils, belonging to the genus Kenyapotamus in Africa, date to around .
Common hippos are recognisable by their barrel-shaped torsos, wide-opening mouths revealing large canine tusks, nearly hairless bodies, columnar-like legs and large size; adults average 1,500 kg (3,300 lb) and 1,300 kg (2,900 lb) for males and females respectively, making them the largest species of land mammal after the three species of elephants and the white and Indian rhinoceros. Despite its stocky shape and short legs, it is capable of running 30 km/h (19 mph) over short distances. The hippopotamus is a highly aggressive and unpredictable animal and is ranked among the most dangerous animals in Africa. Nevertheless, they are still threatened by habitat loss and poaching for their meat and ivory canine teeth.
The common hippopotamus is semiaquatic, inhabiting rivers, lakes and mangrove swamps, where territorial bulls preside over a stretch of river and groups of five to thirty females and young. During the day, they remain cool by staying in the water or mud; reproduction and childbirth both occur in water. They emerge at dusk to graze on grasses. While hippopotamuses rest near each other in the water, grazing is a solitary activity and hippos are not territorial on land.
- 1 Etymology
- 2 Taxonomy and origins
- 3 Description
- 4 Distribution
- 5 Behaviour and life history
- 6 Hippos and humans
- 7 References
- 8 External links
The Latin word "hippopotamus" is derived from the ancient Greek ἱπποπόταμος, hippopotamos, from ἵππος, hippos, "horse", and ποταμός, potamos, "river", meaning "horse of the river". In English, the plural is hippopotamuses, but "hippopotami" is also used; "hippos" can be used as a short plural. Hippopotamuses are gregarious, living in groups of up to thirty animals. A group is called a pod, herd, dale, or bloat.
In Africa, the hippo is known by various names, including seekoei (Afrikaans), mvuvu (Venda), kubu (Lozi) and mvubu (Xhosa, Siswati and Zulu) in the south; kiboko (Swahili), ensherre (Nkore), tomondo (Turu), nvubu (Luganda), ifuru (Luhya), emiria (Ateso), magawit (Sebei), kibei (Kalenjin) and olmakau (Maasai) in the African Great Lakes region;:256 and ጉማርረ/gumarre (Amharic) and jeer (Somali) in the Horn of Africa.
Taxonomy and origins
The hippopotamus is the type genus of the family Hippopotamidae. The pygmy hippopotamus belongs to a different genus in Hippopotamidae, either Choeropsis or Hexaprotodon. Hippopotamidae are sometimes known as hippopotamids. Sometimes, the subfamily Hippopotaminae is used. Further, some taxonomists group hippopotamuses and anthracotheres in the superfamily Anthracotheroidea.:39 Hippopotamidae are classified along with other even-toed ungulates in the order Artiodactyla. Other artiodactyls include camels, cattle, deer and pigs, although hippopotamuses are not closely related to these groups.
- Great northern hippopotamus or Nile hippopotamus H. a. amphibius – (the nominate subspecies) which stretched from Egypt, where they are now extinct, south up the Nile River to Tanzania and Mozambique
- East African hippopotamus H. a. kiboko – in Kenya in the African Great Lakes region, and in Somalia in the Horn of Africa. Broader nasals and more hollowed interorbital region
- Cape hippopotamus or South African hippopotamus H. a. capensis – from Zambia to South Africa, most flattened skull of the subspecies
- West African hippopotamus or Tchad hippopotamus H. a. tschadensis – throughout Western Africa to, as the name suggests, Chad, slightly shorter and wider face, with prominent orbits
- Angola hippopotamus H. a. constrictus – in Angola, the southern Democratic Republic of Congo and Namibia, named for its deeper preorbital constriction
The suggested subspecies were never widely used or validated by field biologists; the described morphological differences were small enough that they could have resulted from simple variation in nonrepresentative samples.:2 Genetic analyses have tested the existence of three of these putative subspecies. A study examining mitochondrial DNA from skin biopsies taken from 13 sampling locations, considered genetic diversity and structure among hippo populations across the continent. The authors found low, but significant, genetic differentiation among H. a. amphibius, H. a. capensis, and H. a. kiboko. Neither H. a. tschadensis nor H. a. constrictus has been tested.
Until 1909, naturalists grouped hippos with pigs, based on molar patterns. Several lines of evidence, first from blood proteins, then from molecular systematics and DNA  and the fossil record, show that their closest living relatives are cetaceans – whales, dolphins and porpoises. The common ancestor of hippos and whales branched off from Ruminantia and the rest of the even-toed ungulates; the cetacean and hippo lineages split soon afterwards.
The most recent theory of the origins of Hippopotamidae suggests that hippos and whales shared a common semiaquatic ancestor that branched off from other artiodactyls around  This hypothesized ancestral group likely split into two branches around . One branch would evolve into cetaceans, possibly beginning about , with the protowhale Pakicetus and other early whale ancestors collectively known as Archaeoceti, which eventually underwent aquatic adaptation into the completely aquatic cetaceans. The other branch became the anthracotheres, a large family of four-legged beasts, the earliest of which in the late Eocene would have resembled skinny hippopotamuses with comparatively small and narrow heads. All branches of the anthracotheres, except that which evolved into Hippopotamidae, became extinct during the Pliocene without leaving any descendants..
A rough evolutionary lineage can be traced from Eocene and Oligocene species: Anthracotherium and Elomeryx to the Miocene species Merycopotamus and Libycosaurus and the very latest anthracotheres in the Pliocene. Merycopotamus, Libycosaurus and all hippopotamids can be considered to form a clade, with Libycosaurus being more closely related to hippos. Their common ancestor would have lived in the Miocene, about . Hippopotamids are therefore deeply nested within the family Anthracotheriidae. The Hippopotamidae are believed to have evolved in Africa; the oldest known hippopotamid is the genus Kenyapotamus, which lived in Africa from 16 to . While hippopotamid species spread across Asia and Europe, no hippopotamuses have ever been discovered in the Americas, although various anthracothere genera emigrated into North America during the early Oligocene. From 7.5 to , an ancestor to the modern hippopotamus, Archaeopotamus, lived in Africa and the Middle East.
While the fossil record of hippos is still poorly understood, the two modern genera, Hippopotamus and Choeropsis (sometimes Hexaprotodon), may have diverged as far back as . Taxonomists disagree whether or not the modern pygmy hippopotamus is a member of Hexaprotodon – an apparently paraphyletic genus, also embracing many extinct Asian hippopotamuses, that is more closely related to Hippopotamus – or of Choeropsis, an older and basal genus.
Three species of Malagasy hippopotamus became extinct during the Holocene on Madagascar, one of them within the past 1,000 years. The Malagasy hippos were smaller than the modern hippopotamus, likely through the process of insular dwarfism. Fossil evidence indicates many Malagasy hippos were hunted by humans, a likely factor in their eventual extinction. Isolated members of Malagasy hippopotamus may have survived in remote pockets; in 1976, villagers described a living animal called the kilopilopitsofy, which may have been a Malagasy hippopotamus.
Three species of hippopotamus, the European hippopotamus (H. antiquus), H. major and H. gorgops, ranged throughout continental Europe and the British Isles. All three species became extinct before the last glaciation. Ancestors of European hippos found their way to many islands of the Mediterranean during the Pleistocene. The Pleistocene also saw a number of dwarf species evolve on several Mediterranean islands, including Crete (H. creutzburgi), Cyprus (H. minor), Malta (H. melitensis), and Sicily (H. pentlandi). Of these, the Cyprus dwarf hippopotamus survived until the end of the Pleistocene or early Holocene. Evidence from an archaeological site, Aetokremnos, continues to cause debate on whether or not the species was encountered, and was driven to extinction, by man.
Hippopotamuses are among the largest living land mammals, being only smaller than elephants and some rhinoceroses. Mean adult weight is around 1,500 kg (3,300 lb) and 1,300 kg (2,900 lb) for males and females respectively, very large males can reach 2,000 kg (4,400 lb) and an exceptional male weighing almost 2,700 kg (6,000 lb) has been reported. Male hippos appear to continue growing throughout their lives while females reach maximum weight at around age 25.
Different from all other large land mammals, hippos are of semiaquatic habits, spending the day in lakes and rivers.:3 The eyes, ears, and nostrils of hippos are placed high on the roof of their skulls. This allows these organs to remain above the surface while the rest of the body submerges.:259 Their barrel-shaped bodies have graviportal skeletal structures,:8 adapted to carrying their enormous weight, and their specific gravity allows them to sink and move along the bottom of a river. Hippopotamuses have small legs (relative to other megafauna) because the water in which they live reduces the weight burden. Though they are bulky animals, hippopotamuses can gallop at 30 km/h (19 mph) on land but normally trot. They are incapable of jumping but do climb up steep banks. Despite being semiaquatic and having webbed feet, an adult hippo is not a particularly good swimmer nor can it float. It is rarely found in deep water; when it is, the animal moves by porpoise-like leaps from the bottom.:3 The testes of the males descend only partially and a scrotum is not present. In addition, the penis retracts into the body when not erect. The genitals of the female are unusual in that the vagina is ridged and two large diverticula protrude from the vulval vestibule. The function of these is unknown.:28–29
The hippo's jaw is powered by a large masseter and a well-developed digastric; the latter loops up behind the former to the hyoid.:259 The jaw hinge is located far back enough to allow the animal to open its mouth at almost 180°.:17 A moderate folding of the orbicularis oris muscle allows the hippo to achieve such a gape without tearing any tissue. On the National Geographic Channel television program, "Dangerous Encounters with Brady Barr", Dr. Brady Barr measured the bite force of an adult female hippo at 8,100 newtons (1,800 lbf); Barr also attempted to measure the bite pressure of an adult male hippo, but had to abandon the attempt due to the male's aggressiveness. Hippopotamus teeth sharpen themselves as they grind together. The lower canines and lower incisors are enlarged, especially in males, and grow continuously. The incisors can reach 40 cm (1 ft 4 in), while the canines reach up to 50 cm (1 ft 8 in). The canines and incisors are used for combat and play no role in feeding. Hippos rely on their broad horny lips to grasp and pull grasses which are then ground by the molars.:259, 263 The hippo is considered to be a pseudoruminant; it has a complex three-chambered stomach but does not "chew cud".:22
Unlike most other semiaquatic animals, the hippopotamus has very little hair.:260 The skin is 6 cm (2 in) thick, providing it great protection against conspecifics and predators. By contrast, its subcutaneous fat layer is thin.:3 The animals' upper parts are purplish-gray to blue-black, while the under parts and areas around the eyes and ears can be brownish-pink.:260 Their skin secretes a natural sunscreen substance which is red-coloured. The secretion is sometimes referred to as "blood sweat", but is neither blood nor sweat. This secretion is initially colourless and turns red-orange within minutes, eventually becoming brown. Two distinct pigments have been identified in the secretions, one red (hipposudoric acid) and one orange (norhipposudoric acid). The two pigments are highly acidic compounds. Both pigments inhibit the growth of disease-causing bacteria; as well, the light absorption of both pigments peaks in the ultraviolet range, creating a sunscreen effect. All hippos, even those with different diets, secrete the pigments, so it does not appear that food is the source of the pigments. Instead, the animals may synthesize the pigments from precursors such as the amino acid tyrosine. Nevertheless, this natural sunscreen cannot prevent the animal's skin from cracking if it stays out of water too long.
A hippo's lifespan is typically 40–50 years.:277 Donna the Hippo was the oldest living hippo in captivity. She lived at the Mesker Park Zoo in Evansville, Indiana in the US until her death in 2012 at the age of 61. The oldest hippo ever recorded was called Tanga; she lived in Munich, Germany, and died in 1995 at the age of 61.
Hippopotamus amphibius was widespread in North Africa and Europe during the Eemian and late Pleistocene until about 30,000 years ago. Archaeological evidence exists of its presence in the Levant, dating to less than 3,000 years ago. The species was common in Egypt's Nile region during antiquity, but has since been extirpated. Pliny the Elder writes that, in his time, the best location in Egypt for capturing this animal was in the Saite nome; the animal could still be found along the Damietta branch after the Arab Conquest in 639. Hippos are still found in the rivers and lakes of the northern Democratic Republic of the Congo, Uganda, Tanzania and Kenya, north through to Ethiopia, Somalia and Sudan, west to The Gambia, and south to South Africa.
Genetic evidence suggests that common hippos in Africa experienced a marked population expansion during or after the Pleistocene epoch, attributed to an increase in water bodies at the end of the era. These findings have important conservation implications as hippo populations across the continent are currently threatened by loss of access to fresh water. Hippos are also subject to unregulated hunting and poaching. In May 2006, the hippopotamus was identified as a vulnerable species on the IUCN Red List drawn up by the International Union for Conservation of Nature (IUCN), with an estimated population of between 125,000 and 150,000 hippos, a decline of between 7% and 20% since the IUCN's 1996 study. Zambia (40,000) and Tanzania (20,000–30,000) possess the largest populations.
The hippo population declined most dramatically in the Democratic Republic of the Congo. By 2005, the population in Virunga National Park had dropped to 800 or 900 from around 29,000 in the mid-1970s. The decline is attributed to the disruptions caused by the Second Congo War. The poachers are believed to be Mai-Mai rebels, poorly paid Congolese soldiers, and local militia groups. Reasons for poaching include the belief that hippos are harmful to society, as well as financial gain. However, as of 2016, the Virunga hippo population appears to have increased, possibly due to greater enforcement and cooperation between fishermen and park authorities. The sale of hippo meat is illegal, but black-market sales are difficult for Virunga National Park officers to track. Hippo meat is considered a delicacy in some areas of central Africa and the teeth have become a valued substitute for elephant ivory.
In the late 1980s, Pablo Escobar kept four hippos in a private menagerie at his residence in Hacienda Nápoles, 100 kilometres (62 mi) east of Medellín, Colombia, after buying them in New Orleans. They were deemed too difficult to seize and move after Escobar's death, and hence left on the untended estate. By 2007, the animals had multiplied to 16 and had taken to roaming the area for food in the nearby Magdalena River. In 2009, two adults and one calf escaped the herd and, after attacking humans and killing cattle, one of the adults (called "Pepe") was killed by hunters under authorization of the local authorities. As of early 2014, 40 hippos have been reported to exist in Puerto Triunfo, Antioquia from the original four belonging to Escobar. The National Geographic Channel produced a documentary about them titled Cocaine Hippos.
Behaviour and life history
Hippos can be found in both savannah and forest areas. Proper habitat requires enough water to submerge in and grass nearby. Larger densities of the animals inhabit quiet waters with mostly firm, smooth sloping beaches. Males may be found in very small numbers in rapid waters in rocky gorges.:264 With the exception of eating, most of hippopotamuses' lives – from fighting with other hippos, mating, to parturition – occurs in the water. Hippos leave the water at dusk and travel inland, sometimes up to 10 km (6 mi), to graze on short grasses, their main source of food. They spend four to five hours grazing and can consume 68 kg (150 lb) of grass each night. Like almost any herbivore, they consume other plants if presented with them, but their diet in nature consists almost entirely of grass, with only minimal consumption of aquatic plants. Hippos are born with sterile intestines, and require bacteria obtained from their mothers' feces to digest vegetation. Hippos have (rarely) been filmed eating carrion, usually close to the water. There are other reports of meat-eating, and even cannibalism and predation. The stomach anatomy of a hippo is not suited to carnivory, and meat-eating is likely caused by aberrant behaviour or nutritional stress.:84
Hippo defecation creates allochthonous deposits of organic matter along the river beds. These deposits have an unclear ecological function. A 2015 study suggests that hippo dung provides nutrients from terrestrial material for fish and aquatic invertebrates. Because of their size and their habit of taking the same paths to feed, hippos can have a significant impact on the land across which they walk, both by keeping the land clear of vegetation and depressing the ground. Over prolonged periods, hippos can divert the paths of swamps and channels. Adult hippos move at speeds up to 8 km/h (5 mph) in water; typically resurfacing to breathe every three to five minutes. The young have to breathe every two to three minutes.:4 The process of surfacing and breathing is automatic. A hippo sleeping underwater rises and breathes without waking. A hippo closes its nostrils when it submerges into the water. As with fish and turtles on a coral reef, hippos occasionally visit cleaning stations and signal, by opening their mouths wide, their readiness for being cleaned of parasites by certain species of fishes. This is an example of mutualism in which the hippo benefits from the cleaning, while the fish receive food.
Studying the interaction of male and female hippopotamuses has long been complicated because hippos are not sexually dimorphic; thus females and young males are almost indistinguishable in the field. Although hippos lie close to each other, they do not seem to form social bonds except between mothers and daughters, and they are not social animals. The reason they huddle close together is unknown.:49 Hippopotamuses are territorial only in water, where a bull presides over a small stretch of river, on average 250 m (270 yd) in length, and containing 10 females. The largest pods can contain over 100 hippos.:50 Younger bachelors are allowed in a bull's stretch, as long as they behave submissively toward the bull. The territories of hippos exist to establish mating rights. Within the pods, the hippos tend to segregate by gender. Bachelors lounge near other bachelors, females with other females, and the bull on his own. When hippos emerge from the water to graze, they do so individually.:4
Hippopotamuses appear to communicate vocally, through grunts and bellows, and they may practice echolocation, but the purpose of these vocalizations is currently unknown. Hippos have the unique ability to hold their heads partially above the water and send out a cry that travels through both water and air; individuals respond above and under water.
Female hippos reach sexual maturity at five to six years and have a gestation period of eight months. A study of endocrine systems revealed that female hippopotamuses may begin puberty as early as three or four years. Males reach maturity at around 7.5 yr. A study of hippopotamus reproductive behaviour in Uganda showed that peak conceptions occurred during the end of the wet season in the summer, and peak births occurred toward the beginning of the wet season in late winter. This is because of the female's oestrous cycle; as with most large mammals, male hippopotamus spermatozoa is active year-round. Studies of hippos in Zambia and South Africa also showed evidence of births occurring at the start of the wet season.:60–61 After becoming pregnant, a female hippopotamus will typically not begin ovulation again for 17 months.
Mating occurs in the water, with the female submerged for most of the encounter,:63 her head emerging periodically to draw breath. Baby hippos are born underwater at a weight between 25 and 50 kg (55 and 110 lb) and an average length of around 127 cm (4.17 ft), and must swim to the surface to take their first breaths. A mother typically gives birth to only one calf, although twins also occur. The young often rest on their mothers' backs when the water is too deep for them, and they swim under water to suckle. They suckle on land when the mother leaves the water. Weaning starts between six and eight months after birth, and most calves are fully weaned after a year.:64 Like many other large mammals, hippos are described as K-strategists, in this case typically producing just one large, well-developed infant every couple of years (rather than many small, poorly developed young several times per year as is common among small mammals such as rodents).
Hippopotamuses are aggressive animals. Hippos that attack other animals are often either territorial bulls or females protecting their calves. Hippopotamus coexist with a variety of formidable predators. Nile crocodiles, lions and spotted hyenas are known to prey on young hippos.:273:118 However, due to their aggression and size, adult hippopotamus are not usually preyed upon by other animals. Cases where large lion prides or cooperating groups of Nile crocodiles have successfully preyed on adult hippopotamus have been reported; however, this predation is exceptionally rare. Crocodiles are frequent targets of hippo aggression, probably because they often inhabit the same riparian habitats; crocodiles may be either aggressively displaced or killed by hippopotamuses. Hippos are also very aggressive towards humans, whom they sometimes attack whether in boats or on land, commonly with no apparent provocation, and are widely considered to be one of the most dangerous large animals in Africa.
Hippos mark their territory by defecation. While depositing the faeces, hippos spin their tails to distribute their excrement over a greater area. "Yawning" serves as a threat display. When fighting, male hippos use their incisors to block each other's attacks and their large canines to inflict injuries.:260 When hippos become over-populated or a habitat is reduced, bulls sometimes attempt infanticide, but this behaviour is not common under normal conditions. Incidents of hippo cannibalism have been documented, but this is believed to be the behaviour of distressed or sick hippos.:82–83
Hippos and humans
The earliest evidence of human interaction with hippos comes from butchery cut marks on hippo bones at Bouri Formation dated around 160,000 years ago. Later rock paintings and engravings showing hippos being hunted have been found in the mountains of the central Sahara dated 4,000–5,000 years ago near Djanet in the Tassili n'Ajjer Mountains.:1 The ancient Egyptians recognised the hippo as a ferocious denizen of the Nile.
The hippopotamus was also known to the Greeks and Romans. The Greek historian Herodotus described the hippopotamus in The Histories (written circa 440 BC) and the Roman naturalist Pliny the Elder wrote about the hippopotamus in his encyclopedia Naturalis Historia (written circa 77 AD).
Zulu warriors preferred to be as brave as a hippopotamus, since even lions were not considered to match its courage. "In 1888, Captain Baden-Powell was part of a column searching for the Zulu chief Dinuzulu, who was leading the Usutu people in revolt against the British colonists. The column was joined by John Dunn, a white Zulu chief, who led an impi (army) of 2,000 Zulu warriors to join the British." 
The words of the Zulu anthem sounded like this:
"Een-gonyama Gonyama! Invooboo! Yah-bo! Yah-bo! Invooboo!
"John Dunn was at the head of his impi. [Baden Powell] asked him to translate the Zulu anthem his men had been singing. Dunn laughed and replied: 'He is a lion. Yes, he is better than a lion – he is a hippopotamus.'"
In the U.S., Representative Robert F. Broussard of Louisiana introduced the "American Hippo bill" in 1910 to authorize the importation and release of hippopotamus into the bayous of Louisiana. Broussard argued that the hippopotamus would eat the invasive water hyacinth that was clogging the rivers and also produce meat to help solve the American meat crisis. The chief collaborators and proponents of Broussard's bill were Major Frederick Russell Burnham and Captain Fritz Duquesne Former President Theodore Roosevelt backed the plan, as did the U.S. Department of Agriculture, The Washington Post, and The New York Times which praised the taste of hippopotamus as "lake cow bacon". The "American Hippo Bill" fell just short of being passed.
Attacks on humans
The hippopotamus is considered to be very aggressive and has frequently been reported as charging and attacking boats. Small boats can be capsized by hippos and passengers can be injured or killed by the animals or drown. In one case in Niger, a boat was capsized by a hippo and 13 people were killed. As hippopotamuses will often engage in raiding nearby crops if the opportunity arises, humans may also come in conflict with them on these occasions, with potential for fatalities on both sides.
Hippos in zoos
Hippopotamuses have long been popular zoo animals. The first zoo hippo in modern history was Obaysch, which arrived at the London Zoo on May 25, 1850, where he attracted up to 10,000 visitors a day and inspired a popular song, the "Hippopotamus Polka". Hippos have remained popular zoo animals since Obaysch, and generally breed well in captivity. Their birth rates are lower than in the wild, but this is attributed to zoos not wanting to breed as many hippos as possible, since hippos are large and relatively expensive animals to maintain.:129
Like many zoo animals, hippos were traditionally displayed in concrete exhibits. In the case of hippos, they usually had a pool of water and patch of grass. In the 1980s, zoo designers increasingly designed exhibits that reflected the animals' native habitats. One of these, the Toledo Zoo Hippoquarium, features a 360,000 gallon pool for hippos. In 1987, researchers were able to record for the first time an underwater birth as in the wild at the Toledo Zoo. The exhibit was so popular, the hippos became the logo of the Toledo Zoo.
A red hippopotamus represented the Ancient Egyptian god Set; the thigh is the "phallic leg of Set" symbolic of virility. Set's consort Tawaret was also seen as part hippo and was a goddess of protection in pregnancy and childbirth, because ancient Egyptians recognised the protective nature of a female hippopotamus toward her young. The Ijaw people of the Niger Delta wore masks of aquatic animals like the hippo when practicing their water spirit cults. The Behemoth from the Book of Job, 40:15–24 is thought to be based on a hippo.
Hippopotamuses have been the subjects of various African folktales. According to a San story; when the Creator assigned each animal its place in nature, the hippos wanted to live in the water, but were refused out of fear that they might eat all the fish. After begging and pleading, the hippos were finally allowed to live in the water on the conditions that they would eat grass instead of fish and would fling their dung so that it can be inspected for fish bones. In a Ndebele tale, the hippo originally had long, beautiful hair, but was set on fire by a jealous hare and had to jump into a nearby pool. The hippo lost most of his hair and was too embarrassed to leave the water.
Ever since Obaysch inspired the "Hippopotamus Polka", hippos have been popular animals in Western culture for their rotund appearance that many consider comical. Stories of hippos such as Huberta, which became a celebrity in South Africa in the 1930s for trekking across the country; or the tale of Owen and Mzee, a hippo and tortoise which developed an intimate bond; have amused people who have bought hippo books, merchandise, and many stuffed hippo toys. Hippos were mentioned in the novelty Christmas song "I Want a Hippopotamus for Christmas" that became a hit for child star Gayla Peevey in 1953. They also feature in the songs "The Hippopotamus" and "Hippo Encore" by Flanders and Swann, with the famous refrain "Mud, Mud, Glorious Mud". They even inspired a popular board game, Hungry Hungry Hippos.
Hippos have also been popular cartoon characters, where their rotund frames are used for humorous effect. For example, the Disney film Fantasia featured a ballerina hippopotamus dancing to the opera La Gioconda, and Hanna-Barbera created Peter Potamus.
- Lewison, R. & Oliver, W. (IUCN SSC Hippo Specialist Subgroup) (2008). "Hippopotamus amphibius". IUCN Red List of Threatened Species. Version 2008. International Union for Conservation of Nature. Retrieved 2009-04-05. Database entry includes a range map and justification for why this species is vulnerable.
- "ITIS on Hippopotamus amphibius". Integrated Taxonomic Information System. Archived from the original on 2014-08-26. Retrieved 2007-07-29.
- "Deadly 60: 15 Deadly Animal Facts". National Geographic. Archived from the original on 2014-10-16.
- ἱπποπόταμος, ἵππος, ποταμός. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project.
- "Hippopotamus". Merriam-Webster's Online Dictionary. Retrieved 2007-07-18.
- Harper, Douglas. "hippopotamus". Online Etymology Dictionary.
- "Plural of hippopotamus". Oxford English Dictionary. Retrieved 2007-07-18.
- Walker, C. (1997). Signs of the Wild. Struik. p. 140. ISBN 1-86825-896-3.
- Kingdon, J. (1988). East African Mammals: An Atlas of Evolution in Africa, Volume 3, Part B: Large Mammals. University Of Chicago Press. pp. 256–77. ISBN 0-226-43722-1.
- Kane, Thomas Leiper (1990). Amharic-English dictionary: H – N., Volume 1. Otto Harrassowitz Verlag. p. 1909. ISBN 3-447-02871-8.
- Saeed, John I. (1999). Somali. John Benjamins Publishing Company. p. 29. ISBN 90-272-3810-3.
- Eltringham, S.K. (1999). The Hippos. Poyser Natural History Series. Academic Press. ISBN 0-85661-131-X.
- Okello, J.B.A; Nyakaana, S.; Masembe, C.; Siegismund, H.R. an Arctander, P. (2005). "Mitochondrial DNA variation of the common hippopotamus: evidence for a recent population expansion.". Heredity. 95 (3): 206–215. doi:10.1038/sj.hdy.6800711. PMID 16030528.
- Meijaard, Erik, ed. (September 2005). "Suiform Soundings: The IUCN/SSC Pigs, Peccaries, and Hippos Specialist Group (PPHSG) Newsletter" (PDF). IUCN. 5 (1).
- Tsagkogeorga, Georgia; McGowen, Michael R.; Davies, Kalina T. J.; Jarman, Simon; Polanowski, Andrea; Bertelsen, Mads F.; Rossiter, Stephen J. (2015). "A phylogenomic analysis of the role and timing of molecular adaptation in the aquatic transition of cetartiodactyl mammals". Royal Society Open Science. 2 (9): 150156. doi:10.1098/rsos.150156. ISSN 2054-5703.
- Ursing, B.M.; Arnason U. (1998). "Analyses of mitochondrial genomes strongly support a hippopotamus-whale clade". Proceedings of the Royal Society. 265 (1412): 2251–5. doi:10.1098/rspb.1998.0567. PMC . PMID 9881471.
- Gatesy, J. (1 May 1997). "More DNA support for a Cetacea/Hippopotamidae clade: the blood-clotting protein gene gamma-fibrinogen" (PDF). Molecular Biology and Evolution. 14 (5): 537–543. doi:10.1093/oxfordjournals.molbev.a025790. PMID 9159931.
- Geisler, J. H.; Theodor, J. M. (2009). "Hippopotamus and whale phylogeny". Nature. 458 (7236): E1–4; discussion E5. doi:10.1038/nature07776. PMID 19295550.
- Sanders, Robert (2005-01-25). "Scientists find missing link between the dolphin, whale and its closest relative, the hippo". Science News Daily. Archived from the original on 2015-02-26. Retrieved 2011-01-08.
- Boisserie, Jean-Renaud; Lihoreau, Fabrice; Brunet, Michel (2005). "The position of Hippopotamidae within Cetartiodactyla". Proceedings of the National Academy of Sciences. 102 (5): 1537–1541. doi:10.1073/pnas.0409518102. PMC . PMID 15677331.
- Boisserie, Jean-Renaud; Lihoreau, Fabrice; Brunet, Michel (2005). "Origins of Hippopotamidae (Mammalia, Cetartiodactyla): towards resolution". Zoologica Scripta. 34 (2): 119–143. doi:10.1111/j.1463-6409.2005.00183.x.
- Boisserie, Jean-Renaud (2005). "The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis". Zoological Journal of the Linnean Society. 143: 1–26. doi:10.1111/j.1096-3642.2004.00138.x.
- Stuenes, Solweig (1989). "Taxonomy, habits and relationships of the sub-fossil Madagascan hippopotamuses Hippopotamus lemerlei and H. madagascariensis". Journal of Vertebrate Paleontology. 9 (3): 241–268. doi:10.1080/02724634.1989.10011761.
- Burney, David A.; Ramilisonina (1998). "The Kilopilopitsofy, Kidoky, and Bokyboky: Accounts of Strange Animals from Belo-sur-mer, Madagascar, and the Megafaunal "Extinction Window"". American Anthropologist. 100 (4): 957–966. doi:10.1525/aa.19184.108.40.2067. JSTOR 681820.
- Petronio, C. (1995). "Note on the taxonomy of Pleistocene hippopotamuses" (PDF). Ibex. 3: 53–55.
- Simmons, A. (2000). "Faunal extinction in an island society: pygmy hippopotamus hunters of Cyprus". Geoarchaeology. 15 (4): 379–381. doi:10.1002/(SICI)1520-6548(200004)15:4<379::AID-GEA7>3.0.CO;2-E.
- Owen-Smith, R. Norman (1995). Megaherbivores: The Influence of Very Large Body Size on Ecology. Cambridge University Press.
- Pienaar, U. de V.; Van Wyk, P.; Fairall, N. (1966). "An experimental cropping scheme of Hippopotami in the Letaba river of the Kruger National Park". Koedoe. 9 (1). doi:10.4102/koedoe.v9i1.778.
- Marshall, P.J.; Sayer, J.A. (1976). "Population ecology and response to cropping of a hippopotamus population in eastern Zambia". The Journal of Applied Ecology. 13 (2): 391–403. doi:10.2307/2401788. JSTOR 2401788.
- "Hippopotamus Hippopotamus amphibius". National Geographic. Archived from the original on 2014-11-25. Retrieved 2013-07-10.
- Exploring Mammals. Marshall Cavendish Corporation. 2008. p. 616. ISBN 9780761477280.
- Estes, R. (1992). The Behavior Guide to African Mammals: including hoofed mammals, carnivores, primates. University of California Press. pp. 222–26. ISBN 0-520-08085-8.
- Herring, S. W. (1975). "Adaptations for gape in the hippopotamus and its relatives". Forma et Functio. 8: 85–100.
- Barr, Brady. "Undercover Hippo," Dangerous Encounters, National Geographic Channel, January 20, 2008.
- Saikawa Y, Hashimoto K, Nakata M, Yoshihara M, Nagai K, Ida M, Komiya T (2004). "Pigment chemistry: the red sweat of the hippopotamus". Nature. 429 (6990): 363. doi:10.1038/429363a. PMID 15164051.
- Jablonski, Nina G. (2013). Skin: A Natural History. University of California Press. p. 34. ISBN 0-520-24281-5.
- "Oldest Hippo Turns 55!". Mesker Park Zoo. 2006-06-12. Archived from the original on 2007-09-27. Retrieved 2007-06-21.
- "Celebrate with Donna". Evansville Courier & Press. 2007-07-12. Retrieved 2007-07-15.
- Fears, Danika (2012-08-03). "Goodbye, Donna: World's oldest hippo in captivity dies at 61". Today.com. Retrieved 2013-09-12.
- "Old mother hippo dies". Agence France Press. July 12, 1995.
- van Kolfschoten, Th. (2000). "The Eemian mammal fauna of central Europe" (PDF). Netherlands Journal of Geosciences. 79 (2/3): 269–281.
- Horwitz, Liora Kolska; Eitan Tchernov (1990). "Cultural and Environmental Implications of Hippopotamus Bone Remains in Archaeological Contexts in the Levant". Bulletin of the American Schools of Oriental Research. 280: 67–76. doi:10.2307/1357310.
- Haas, Georg (1953). "On the Occurrence of Hippopotamus in the Iron Age of the Coastal Area of Israel". Bulletin of the American Schools of Oriental Research. 132: 30–34. doi:10.2307/1355798.
- Pliny the Elder. "Chapter 15, Book VIII". Naturalis Historia (in Latin original or English translation). ISBN 3-519-01652-4.
- "Hippo Haven". Smithsonian Magazine. 2006-01-01. Retrieved 2007-01-23.
- "DR Congo's hippos face extinction.". BBC. 2005-09-13. Retrieved 2005-11-14.
- Owen, James (2006-10-24). "Hippos Butchered by the Hundreds in Congo Wildlife Park". National Geographic News. Retrieved 2013-09-11.
- Sundaram, Anjan (2005-09-12). "Congo's Hippos Fast Disappearing". Toronto Star.
- Delaney, J.; Sautner, S. (2016-11-03). "After a Long Demise Due to Poaching, Virunga's Hippos Climbing Back". Wildlife Conservation Society. Retrieved 2016-11-25.
- Pearce, Fred (2003). "Poaching causes hippo population crash". New Scientist. Retrieved April 26, 2014.
- Kraul, Chris (2006-12-20). "A hippo critical situation". Los Angeles Times. Archived from the original on 2015-03-08. Retrieved 2008-03-27.
- "Colombia kills drug baron hippo". BBC News. 2009-07-11. Archived from the original on 2015-01-05. Retrieved 2009-07-11.
- "Crece controversia en el país por decisión de cazar a hipopótamos de Pablo Escobar". El Tiempo. Archived from the original on 2015-03-08. Retrieved 2009-07-11. English translation at Google Translate
- "Hipopótamos bravos". El Espectador. 2014-06-24. Archived from the original on 2014-05-09. Retrieved 2014-06-28. English translation at Google Translate
- "The Invaders: Cocaine Hippos". National Geographic Channel. Archived from the original on 2013-06-26.
- "Hippopotamus". Kruger National Park. Retrieved 2007-06-18.
- Grey, J.; Harper, D.M. (2002). "Using Stable Isotope Analyses To Identify Allochthonous Inputs to Lake Naivasha Mediated Via the Hippopotamus Gut". Isotopes in Environmental Health Studies. 38 (4): 245–250. doi:10.1080/10256010208033269. PMID 12725427.
- "BBC Nature — Dung eater videos, news and facts". Bbc.co.uk. Retrieved 2011-11-27.
- Dudley, J.P. "Reports of carnivory by the common hippo Hippopotamus Amphibius" (PDF). South African Journal of Wildlife Research. 28 (2): 58–59.
- McCauley, D. J.; et al. (2015). "Carbon stable isotopes suggest that hippopotamus-vectored nutrients subsidize aquatic consumers in an East African river". Ecosphere. 6 (4). doi:10.1890/ES14-00514.1.
- McCarthy, T.S.; Ellery, W. N.; Bloem, A (1998). "Some observations on the geomorphological impact of hippopotamus (Hippopotamus amphibius L.) in the Okavango Delta, Botswana". African Journal of Ecology. 36 (1): 44–56. doi:10.1046/j.1365-2028.1998.89-89089.x.
- "Hippopotamuses". PBS Nature. Retrieved 10 July 2013.
- Balcombe, Jonathan (2006). Pleasurable Kingdom: Animals and the Nature of Feeling Good. Palgrave Macmillan. pp. 132–33. ISBN 1-4039-8602-9.
- Beckwitt, R.; Shea, J.; Osborne, D.; Krueger, S. & Barklow, W. (2002). "A PCR-based method for sex identification in Hippopotamus amphibius" (PDF). African Zoology: 127–130.
- William E. Barklow (2004). "Low-frequency sounds and amphibious communication in Hippopotamus amphibious". The Journal of the Acoustical Society of America. 115 (5): 2555. doi:10.1121/1.4783854.
- Graham L.H.; Reid K.; Webster T.; Richards M.; Joseph S. (2002). "Endocrine patterns associated with reproduction in the Nile hippopotamus (Hippopotamus amphibius) as assessed by fecal progestagen analysis". General and Comparative Endocrinology. 128 (1): 74–81. doi:10.1016/S0016-6480(02)00066-7. PMID 12270790.
- Lewison, R (1998). "Infanticide in the hippopotamus: evidence for polygynous ungulates". Ethology, Ecology & Evolution. 10 (3): 277–286. doi:10.1080/08927014.1998.9522857.
- Novak, R. M. 1999. Walker's Mammals of the World. 6th edition. Johns Hopkins University Press, Baltimore. ISBN 0-8018-5789-9
- Hunter, Luke (2011). Carnivores of the World. Princeton University Press. ISBN 978-0-691-15228-8.
- Ross, S Charles A.; Garnett, Stephen (1989). Crocodiles and Alligators. Checkmark Books. ISBN 978-0-8160-2174-1.
- "Dangerous Encounters: Undercover Hippo". National Geographic Channel. Retrieved 2008-11-04.
- Hippo Specialist Group, World Conservation Union. (June 2008). In the News. Duke University. Retrieved 2009-9-04.
- National Geographic exhibit on different animals and their poop. News.nationalgeographic.com (October 28, 2010). Retrieved on 2012-05-12.
- Clark, JD; Beyene, Y; WoldeGabriel, G; Hart, WK; Renne, PR; Gilbert, H; Defleur, A; Suwa, G; et al. (2003). "Stratigraphic, chronological and behavioural contexts of Pleistocene Homo sapiens from Middle Awash, Ethiopia". Nature. 423 (6941): 747–52. doi:10.1038/nature01670. PMID 12802333.
- Herodotus. "Chapter 71, Book II". The Histories (in English translation). ISBN 0-19-521974-0.
- Ingonyama – he is a lion!. Scouting.org.za. Retrieved on 2011-03-29.
- Orans, Lewis P. (1997-06-17). "Scouting in South Africa 1884-1890". Pinetreeweb.com. Archived from the original on 2014-07-28. Retrieved 2011-03-29.
- Miller, Greg (2013-12-20). "The Crazy, Ingenious Plan to Bring Hippopotamus Ranching to America". Wired. ISSN 1059-1028. Archived from the original on 2015-01-17.
- Mooallem, John (2013). American Hippopotamus. New York: The Atavist. Retrieved March 7, 2014.
- Mooallem, Jon (August 10, 2014). "Lake Bacon: The Story of The Man Who Wanted Us to Eat Mississippi Hippos". The Daily Beast. The Newsweek Daily Beast Company. ISSN 0028-9604. Retrieved August 13, 2014.
- Eplett, Layla (March 27, 2014). "The hunger game meat: How hippos early invaded American cuisine". Scientific American. ISSN 0036-8733.
- Burnham, Frederick Russell (1944). Taking Chances. Los Angeles: Haynes Corp. pp. 11–23. ISBN 1-879356-32-5.
- Thomas, Pete (14 January 2015). "Massive hippo charges safari boat in Malawi, Africa". Pete Thomas Outdoor.
- "Hippopotamus attack kills 13 people, including 12 children, in boat near Niger's capital Niamey". Australian Broadcasting Corporation. 20 November 2014.
- Kendall, C. J. (2011). "The spatial and agricultural basis of crop raiding by the Vulnerable common hippopotamus Hippopotamus amphibius around Ruaha National Park, Tanzania". Oryx. 45 (1): 28–34. doi:10.1017/S0030605310000359.
- Root, N. J. (1993). "Victorian England's Hippomania". Natural History. 103: 34–39.
- Melissa Greene (December 1987). "No rms, jungle vu: a new group of "landscape-immersion" zoo designers are trying to break down visitors' sense of security by reminding them that wild animals really are wild.". The Atlantic Monthly.
- "Bubbles' legacy". www.toledozoo.org. Retrieved 2015-12-16.
- "Hippoquarium". Toledo Zoo. Archived from the original on February 11, 2007. Retrieved 2007-03-26.
- Cooper, J.C. (1992). Symbolic and Mythological Animals. London: Aquarian Press. p. 129. ISBN 1-85538-118-4.
- Hart, George (1986). A Dictionary of Egyptian Gods and Goddesses. Routledge. ISBN 0-415-05909-7.
- Hamilton, Janice (2003). Nigeria in Pictures. Twenty-First Century Books. p. 49. ISBN 0-8225-0373-5.
- Metzeger, Bruce M.; Coogan, Michael D. f, eds. (1993). The Oxford Companion to the Bible. Oxford, UK: Oxford University Press. p. 76. ISBN 0-19-504645-5.
- Greaves, N.; Clement, R. (2000). When Hippo Was Hairy: And Other Tales from Africa. Struik. pp. 67–71. ISBN 1-86872-456-5.
- Chilvers, H.A. (1931). Huberta Goes South, a Record of the Lone Trek of the Celebrated Zululand Hippopotamus. London: Gordon & Gotch.
- "A hippo and tortoise tale". NPR. July 17, 2005. Retrieved 2007-06-18.
- Hatkoff, Isabella; Hatkoff, Craig; Kahumbu, Paula (2006). Owen & Mzee; The True Story of a Remarkable Friendship. New York: Scholastic Press. ISBN 0-439-82973-9.
- "I Want A Hippopotamus For Christmas Lyrics". Christmas-lyrics.org. Retrieved 2007-12-20.
- "Childhood Trauma: Hungry Hungry Hippos". Newcastle Herald (Australia). May 2, 2006.
- "Fred Kroll, of Trouble and Hungry Hungry Hippos games, dead at 82". Associated Press. August 5, 2003.
- Markstein, Don (2008). "Peter Potamus". Don Markstein's Toonopedia. Archived from the original on 2015-03-08.
- "Weekly Candy: Kinder Happy Hippos". Philadelphia City Paper. 2011-11-22. Archived from the original on 2015-03-08.
- "Hippos: Wildlife summary". African Wildlife Foundation. Archived from the original on 2010-11-19.
- "Pablo Escobar's Fugitive Hippo Shot Dead". The Daily Telegraph. 2009-06-15. Archived from the original on 2009-07-17.
- "Pablo Escobar's fugitive hippos: zoologists called in to round up animals". The Daily Telegraph. 2009-08-27. Archived from the original on 2009-09-10.
- Kremer, William (2014-06-25). "Pablo Escobar's hippos: A growing problem". BBC News. Archived from the original on 2015-02-25.
- "Hippo Specialist Group of the World Conservation Union". International Union for Conservation of Nature. Archived from the original on 2015-03-08.
- "11 Things You May Not Know About Ancient Egypt: King Tut may have been killed by a hippopotamus". History. 2012-11-12. Archived from the original on 2014-12-17.