Pygmy hippopotamus

The Pygmy Hippopotamus (Choeropsis liberiensis or Hexaprotodon liberiensis) is a large mammal native to the forests and swamps of western Africa (the scientific species classification means "of Liberia", as this is where the vast majority lives). The pygmy hippo is reclusive and nocturnal. It is one of only two extant species in the hippopotamidae family, the other being its much larger cousin the common hippopotamus.

The pygmy hippopotamus displays many terrestrial adaptations, but like its larger cousin, it is semi-aquatic and relies on proximity to water to keep its skin moisturized and its body temperature cool. Behaviors such as mating and birth may occur in water or on land. The pygmy hippo is herbivorous, feeding on whatever ferns, broad-leaved plants, grasses and fruits it finds in the forests.

A rare nocturnal forest creature, the pygmy hippopotamus is a difficult animal to study in the wild; it also lives primarily in countries with a great degree of civil strife. Pygmy hippos lead mostly solitary lives; they are sometimes seen in pairs or threesomes, but never large pods like the common hippopotamus. Unlike the common hippopotamus, they are not known to be territorial.

Pygmy hippos were unknown outside of West Africa until the 19th century. Introduced to zoos in the early 20th century, they breed well in captivity and the vast majority of research is derived from zoo specimens. The survival of the species in captivity is more assured than in the wild: the World Conservation Union estimates that there are less than 3,000 pygmy hippos remaining in the wild. Pygmy hippos are primarily threatened by loss of habitat, as forests are logged and converted to farm land, and are also vulnerable to poaching, hunting, natural predators and war.

Taxonomy and origins

Nomenclature of the pygmy hippopotamus reflects that of the hippopotamus. The plural form is pygmy hippopotami (hippopotamuses is also accepted as a plural form by the OED, or pygmy hippos for short). A male pygmy hippopotamus is known as a bull, a female as a cow, and a baby as a calf.

The pygmy hippopotamus is a member of the family Hippopotamidae where it is classified as a member of either the genus Choeropsis ("resembling a hog") or, the genus Hexaprotodon ("six front teeth"). Hippopotamidae are sometimes known as Hippopotamids. Sometimes the sub-family Hippopotaminae is used. Further, some taxonomists group hippopotami and anthracotheres in the superfamily Anthracotheroidea or Hippopotamoidea.

A sister species of the pygmy hippopotamus may have been the little-studied Madagascan Pygmy Hippopotamus (Choeropsis madagascariensis, sometimes classified as Hexaprotodon madagascariensis or Hippopotamus madagascariensis), one of three recently-extinct species from Madagascar. C. madagascariensis was the same size as C. liberiensis and shared its terrestrial behaviour, inhabiting the forested highlands of Madagascar, rather than open rivers. It is believed to have gone extinct within the last 500 years.

The taxonomy of the genus of the pygmy hippopotamus has changed as understanding of the animal has developed. Samuel G. Morton initially classified the animal as Hippopotamus minor, but later determined it was distinct enough to warrant its own genus, and labeled it Choeropsis. In 1977, Coryndon proposed that the pygmy hippopotamus was closely related to Hexaprotodon, a genus that consisted of prehistoric hippos mostly native to Asia. This assertion was widely accepted, until Boisserie asserted in 2005 that the pygmy hippopotamus was not a member of Hexaprodoton, after a thorough examination of the phylogeny of hippopotamidae; he suggested instead that the pygmy hippopotamus was a distinct genus, and returned the animal to Choeropsis. All agree that the modern pygmy hippopotamus, be it H. liberiensis or C. liberiensis, is the only extant member of its genus.

Nigerian subspecies

A distinct subspecies of pygmy hippopotamus lived in Nigeria until at least the 20th century. The existence of the subspecies, makes Choeropsis liberiensis liberiensis (or Hexaprotodon liberiensis liberiensis under the old classification) the full trinomial nomenclature for the Liberian Pygmy Hippopotamus. The Nigerian Pygmy Hippopotamus subspecies was never studied in the wild and never captured. All research and all zoo specimens are the Liberian subspecies. The Nigerian subspecies is classified as C. liberiensis heslopi.

The Nigerian pygmy hippopotamus ranged in the Niger River Delta, especially near Port Harcourt, but no reliable reports exist after the collection of the museum specimens secured by I.R. P. Heslop, a British colonial officer, in the early 1940s. It is believed to be extinct. The subspecies was separated by over 1800 km and the Dahomey Gap, a region of desert that divides the forest regions of West Africa. The subspecies is named after I.R.P. Heslop, who claimed in 1945 to have shot a pygmy hippo in the Niger Delta region and collected several skulls. He estimated that perhaps no more than 30 pygmy hippos remained in the region.

Heslop reportedly sent four pygmy hippopotamus skulls he collected to the British Museum of Natural History in London. These specimens were not subjected to taxonomic evaluation, however, until 1969 when G.B. Corbet classified the skulls as belonging to a separate subspecies based on consistent variations in the proportions of the skulls. The Nigerian pygmy hippos were seen or shot in Rivers State, Imo State and Bayelsa State, Nigeria. While some local populations are aware that the species once existed, its history in the region is poorly documented.

Evolution

The evolution of the pygmy hippopotamus is most often studied in the context of its larger cousin. Both species were long believed to be most closely related to the family Suidae (pigs and hogs) or Tayassuidae ( peccaries), but research within the last 10 years has determined that pygmy hippos and hippos are most closely related to Cetaceans (whales and dolphins). Hippos and whales shared a common semi-aquatic ancestor that branched off from other Artiodactyls around 60 mya ago. This hypothesized ancestor likely split into two branches about six million years later. One branch would evolve into cetaceans, the other branch became the anthracotheres, a large family of four-legged beasts, whose earliest member, from the Late Eocene, would have resembled narrow hippopotami with comparatively small and thin heads.

Hippopotamids are deeply nested within the family Anthracotheriidae. The oldest known hippopotamid is the genus Kenyapotamus, which lived in Africa from 16 to 8 mya. Kenyapotamus is known only through fragmentary fossils, but was similar in size to C. liberiensis. The Hippopotamidae are believed to have evolved in Africa, and while at one point the species spread across Asia and Europe, no hippopotami have ever been discovered in the Americas. Starting 7.5 to 1.8 mya ago the Archaeopotamus, likely ancestors to the genus Hippopotamus and Hexaprotodon, lived in Africa and the Middle East.

While the fossil record of hippos is still poorly understood, the lineages of the two modern genera, Hippopotamus and Choeropsis, may have diverged as far back as 8 mya. The ancestral form of the pygmy hippopotamus may be the genus Saotherium. Saotherium and Choeropsis are significantly more basal than Hippopotamus and Hexaprotodon, and thus more closely resemble the ancestral species of hippos.

Extinct pygmy and dwarf hippos

Several species of small hippopotamidae have also become extinct in the Mediterranean in the late Pleistocene or early Holocene. Though these species are sometimes known as "Pygmy Hippopotami" they are not believed to be closely related to C. liberiensis. These include the Cretan Dwarf Hippopotamus (Hippopotamus creutzburgi) of Crete, the Sicilian Hippopotamus (Hippopotamus pentlandi) of Sicily, or the Maltese Hippopotamus (Hippopotamus melitensis) of Malta.

These species, though comparable in size to the pygmy hippopotamus, are considered dwarf hippopotamuses, rather than pygmies. They are likely descended from a full-sized species of European Hippopotamus, and reached their small size through the evolutionary process of insular dwarfism which is common on islands; the ancestors of pygmy hippopotami were also small and thus there was never a dwarfing process. There were also several species of pygmy hippo on the island of Madagascar (see Malagasy Hippopotamus).

Description

Pygmy hippos share the same general form as a hippopotamus. They have a graviportal skeleton, with four short legs and four toes on each foot, supporting a portly frame. The pygmy hippo, however, is only half as tall as the hippopotamus and weighs less than 1/4 as much as its larger cousin. Adult pygmy hippos stand about 75–83 cm (30–32 inches) high at the shoulder, are 150–177 cm (59–70 inches) in length and weigh 180–275 kilograms (400-600 pounds). Their lifespan in captivity ranges from 30 to 55 years, though it is unlikely that they live this long in the wild.

The skin is greenish-black or brown, shading to a creamy gray on the lower body. Their skin is very similar to the common hippo's, with a thin Epidermis over a dermis that is several centimeters thick. Pygmy hippos have the same unusual sweat as common hippos, that gives a pinkish tinge to their bodies, and is sometimes described as "blood sweat" though the secretion is neither sweat nor blood. The highly alkaline substance is believed to have antiseptic and sunscreening properties. The skin of hippos dries out quickly and cracks, which is why both species of hippos spend so much time in water.

The skeleton of C. liberiensis is more gracile than that of the common hippopotamus meaning their bones are proportionally thinner. The common hippo's spine is parallel with the ground; the pygmy hippo's back slopes forward, a likely adaptation to pass more easily through dense forest vegetation. Proportionally, the pygmy hippos legs and neck are longer and its head smaller. The orbits and nostrils of a pygmy hippo are much less pronounced, an adaptation from spending less time in deep water (where pronounced orbits and nostrils help the common hippo breathe and see). The feet of pygmy hippos are narrower, but the toes are more spread out and have less webbing, to assist in walking on the forest floor. Despite adaptations to a more terrestrial life than the common hippopotamus, pygmy hippos are still more aquatic than all other even-toed ungulates. The ears and nostrils of pygmy hippos have strong muscular valves to aid submerging underwater, and the skin physiology is dependent on the availability of water.

Behaviour

The behavior of the pygmy hippo differs from the common hippo in many ways. Much of its behaviour is more similar to that of a tapir, though this is an effect of convergent evolution. While the common hippopotamus is gregarious, pygmy hippos live either alone or in small groups, typically a mated pair or a mother and calf. Pygmy hippos tend to ignore each other rather than fight when they meet. Field studies have estimated that male pygmy hippos range over 1.85 km², while the range of a female is between 0.4 to 0.6 km².

Pygmy hippos spend most of the day hidden in rivers. They will rest in the same spot for several days in a row, before moving to a new spot. At least some pygmy hippos make use of dens or burrows that form in river banks. It is unknown if the pygmy hippos help create these dens, or how common it is to use them. Though a pygmy hippo has never been observed burrowing, other Artiodactyls, such as warthogs are burrowers.

Diet

Like the common hippopotamus, the pygmy hippo emerges from the water at dusk to feed. It relies on game trails to travel through dense forest vegetation. It marks the trails by spreading feces by vigorously waving its tail while defecating. The pygmy hippo spends about six hours a day foraging for food.

Pygmy hippos are herbivorous. They do not eat aquatic vegetation to a significant extent and rarely eat grass because it is uncommon in the thick forests they inhabit. The bulk of a pygmy hippo's diet consists of ferns, broad-leaved plants, and fruits that have fallen to the forest floor. The wide variety of plants pygmy hippos have been observed eating suggests that they will eat any plants available. This diet is of higher quality than that of the common hippopotamus.

Reproduction

A study of breeding behavior in the wild has never been conducted; the artificial conditions of captivity may cause the observed behaviour of pygmy hippos in zoos to differ from natural conditions. Sexual maturity for the pygmy hippopotamus occurs at between three to five years. The youngest reported age for giving birth is a pygmy hippo at the zoo in Basel, Switzerland which bore a calf at 3 years and 3 months. The oestrus cycle of a female pygmy hippo lasts an average of 35.5 days, with the oestrus itself lasting between 24-48 hours.

Pygmy hippos consort for mating, but the duration of the relationship is unknown. In zoos they breed as monogamous pairs. Copulation can take place on land or in the water, and a pair will mate one to four times during an oestrus period. In captivity, pygmy hippos have been conceived and born in all months of the year. The gestation period ranges from 190-210 days, and usually a single young is born, though twins are known to occur.

The common hippopotamus gives birth and mates only in the water, but pygmy hippos mate and give birth on both land and water. Young pygmy hippos can swim almost immediately. At birth, pygmy hippos weigh 4.5–6.2 kg (9.9–13.7 lbs) with males weighing about .25 kg (.55 lbs) more than females. Pygmy hippos are fully weaned between 6-8 months of age; before weaning they do not accompany their mother when she leaves the water to forage, but instead hide in the water by themselves. The mother returns to the hiding spot about three times a day and calls out for the calf to suckle. Suckling occurs with the mother lying on her side.

Distribution

The vast majority of pygmy hippos live in Liberia with smaller populations, mostly clustered around the Liberian border, in Côte d'Ivoire, Guinea and Sierra Leone. Though the range of the pygmy hippo as such has not been significantly reduced, populations are now fragmented. C. liberiensis lives exclusively in rivers running through forested regions.

Pygmy hippo populations occur in many forests within Liberia, the largest found in the country's largest protected region Sapo National Park. Other populations have been reported in Grand Kru County, Grand Cape Mount County, Grand Bassa County, Grand Gedeh County, Lofa County, Maryland County, Nimba County and Sinoe County. Studies of these populations, however, have been complicated by civil strife, such as the First Liberian Civil War and the Second Liberian Civil War. In Côte d'Ivoire the pygmy hippo ranges in several forests, including those in the Taï National Park. In Guinea the pygmy hippopotamuses live in the Reserve de Ziama on the border with Liberia. In Sierra Leone, several small populations exist—in the Gola Forest, on Tiwai Island in the Moa River and in the Loma Mountains.

The World Conservation Union's 1993 Action Plan estimated a population of between 2,000 and 3,000 pygmy hippos in the wild, with most in Liberia. The smallest population is that of Sierra Leone, estimated in 1993 to be only around 100. Due to deteriorating conditions in Liberia, the IUCN Red List estimated in 2006 that this number had likely declined, particularly due to loss of habitat.

Conservation

The greatest threat to the remaining pygmy hippopotamus population in the wild is loss of habitat. The forests in which pygmy hippos live have been subject to logging, settling and conversion to agriculture, with little efforts made to make logging sustainable. As forests shrink, the populations become more fragmented, leading to less genetic diversity in the potential mating pool. There are less than 3,000 in the wild.

Because of their reclusive lifestyle they are not a target of subsistence hunting, though they are hunted opportunistically by bush hunters. Their meat is said to be of excellent quality, like that of a wild boar; unlike the common hippo, the pygmy hippos teeth have no value. The effects of West Africa's civil strife on the pygmy hippopotamus are unknown, but unlikely to be positive. Adult common hippos have no natural predators, but the pygmy hippopotamus is capable of being killed by leopards, pythons and crocodiles. How often this occurs, however, is unknown.

C. liberiensis was identified as one of the top-10 "focal species" in 2007 by the Evolutionarily Distinct and Globally Endangered (EDGE) project. EDGE identifies species that are evolutionarily distinct and need better protection to prevent extinction. The pygmy hippo was placed in the top-10 along with Attenborough's long-beaked echidna, Hispaniolan solenodon, Bactrian camel, Yangtze River dolphin, Slender loris, Hirola antelope, Golden-rumped elephant shrew, Bumblebee bat, and the Long-eared jerboa.

Although threatened in the wild, pygmy hippos breed freely in zoos. Between 1970 and 1991 the population of pygmy hippos born in captivity more than doubled. The survival of the species in zoos is more certain than the survival of the species in the wild. In captivity, the pygmy hippo lives from 42-55 years, longer than in the wild. Since 1919, only 41 percent of pygmy hippos born in zoos have been male.

History and folklore

While the common hippopotamus was known to Europeans since classical antiquity, the pygmy hippopotamus was unknown outside of its range in West Africa until the 19th century. Due to their nocturnal, forested existence, they were poorly known within their range as well. In Liberia the animal was traditionally known as a water cow.

Early field reports of the animal misidentified it as a wild hog. Several skulls of the species were sent to the American natural scientist Samuel G. Morton, during his residency in Monrovia, Liberia. Morton first described the species in 1843. The first complete specimens were collected as part of a comprehensive investigation of Liberian fauna in the 1870s and 1880s by Dr. Johann Büttikofer. The specimens were taken to the Natural History Museum in Leiden, Netherlands.

The first pygmy hippo was brought to Europe in 1873 after being captured in Sierra Leone by a member of the British Colonial Service but died shortly after arrival. Pygmy hippos were successfully introduced to Europe in 1911. They were first shipped to Germany and then to the Bronx Zoo in New York City where they also thrived.

In 1927, Harvey Firestone of Firestone Tires presented Billy the pygmy hippo to U.S. President Calvin Coolidge. Coolidge donated Billy to the Smithsonian National Zoological Park. According to the zoo, Billy is a common ancestor to most pygmy hippos in U.S. zoos today.

Several folktales have been collected about the pygmy hippopotamus. One tale says that pygmy hippos carry a shining diamond in their mouths to help travel through thick forests at night; by day the pygmy hippo has a secret hiding place for the diamond, but if a hunter catches a pygmy hippo at night the diamond can be taken. Villagers sometimes believed that baby pygmy hippos do not nurse but rather lick secretions off the skin of the mother.