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Hippopotamidae
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| Hippopotamidae Temporal range: Late Miocene-Recent
| |
|---|---|
| Common hippopotamus | |
| Pygmy hippopotamus | |
| Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Artiodactyla |
| Infraorder: | Ancodonta |
| Superfamily: | Hippopotamoidea |
| Family: | Hippopotamidae Gray, 1821 |
| Subtaxa | |
|
†Trilobophorus Geze, 1985
| |
Hippopotamidae is a family of stout, naked-skinned, and semiaquatic artiodactyl mammals, possessing three-chambered stomachs and walking on four toes on each foot. While they resemble pigs physiologically, their closest living relatives are the cetaceans. They are formally referred to as hippopotamids.
There are two living species of hippopotamid in two genera; the pygmy hippo, Choeropsis liberiensis of the forests of west Africa, and the common hippo, Hippopotamus amphibius. The term hippopotamus can also be applied to hippopotamids in general, although it is most frequently used for the common hippo and its respective genus.
Characteristics
[edit]Hippopotamids are large mammals, with short, stumpy legs, and barrel-shaped bodies. They have large heads, with broad mouths, and nostrils placed at the top of their snouts. Like pigs, they have four toes, but unlike pigs, all of the toes are used in walking. Hippopotamids are unguligrade, although, unlike most other such animals, they have no hooves, instead using a pad of tough connective tissue on each foot. Their stomachs have three chambers, but they are not true ruminants.
The living species are smooth-skinned and lack both sebaceous glands and sweat glands. The outer epidermis is relatively thin, so hippos dehydrate rapidly in dry environments.[1]
Both the incisors and canines are large and tusk-like, although the canine tusks are by far the larger. The tusks grow throughout life. The postcanine teeth are large and complex, suited for chewing the plant matter that comprises their diets. The number of incisors varies even within the same species, but the general dental formula is given in the table below:
| Dentition |
|---|
| 2–3.1.4.3 |
| 1–3.1.4.3 |
Evolution
[edit]
The hippopotamids are descended from the anthracotheres, a family of semiaquatic and terrestrial artiodactyls that appeared in the late Eocene, and are thought to have resembled small- or narrow-headed hippos. The hippos split off from the anthracotheres some time during the Miocene. The oldest records of Hippopotamidae are from Afro-Arabia and date to the late Miocene, approximately 7.4 million years ago, expanding into Eurasia around 6 million years ago.[2] It has been theorised that this Late Miocene radiation of hippopotamids represents the coevolution of hippopotamids with the expansion of C4 grasslands,[3] a phenomenon known as the hippopotamine event (HE).[4] After the appearance of the hippopotamids, the remaining anthracotheres went into a decline brought about by a combination of climatic change and competition with their descendants, until the last genus, Merycopotamus, died out in the early Pliocene of India.
There were once many species of hippopotamid, but only two survive today: Hippopotamus amphibius, and Choeropsis liberiensis. They are the last survivors of two major evolutionary lineages, the hippos proper and the pygmy hippos, respectively; these lineages could arguably be considered subfamilies, but their relationship to each other – apart from being fairly distant relatives – is not well resolved.
The enigmatic Miocene Kenyapotamus is insufficiently known to be assigned a place in the hippo phylogeny with any degree of certainty. In addition, the genus Hexaprotodon, which is now largely restricted to South Asia and Southeast Asia, formerly included many fossil hippopotamuses that are now thought to be unrelated.[5]
Taxonomy
[edit]Hippopotamidae's placement within Artiodactyla can be represented in the following cladogram:[6][7][8][9][10]
| Artiodactyla |
| ||||||||||||||||||||||||
Analogous structures
[edit]The lower canine teeth of hippopotamids are similar in function and structure to the tusks of elephants. While hippopotamids and elephants are only very distantly related within the Mammalia, the lower canine teeth of both groups are long and have a slight curve, and species of both families use this structure when fighting.
Species
[edit]The systematics and taxonomy used here mostly follows a review by J.-R. Boisserie[11][a] and the American Society of Mammalogists.[12]
Recent species
[edit]- Genus Hippopotamus – true hippopotamuses
- Hippopotamus amphibius – hippopotamus (Africa)
- †Hippopotamus lemerlei – Lemerle's dwarf hippopotamus (Madagascar; Holocene)
- †Hippopotamus madagascariensis – Madagascan dwarf hippopotamus (Madagascar; Holocene)
- Genus Choeropsis – pygmy hippopotamuses
- Choeropsis liberiensis – pygmy hippopotamus (west Africa)
Fossil species
[edit]- Genus Hippopotamus – true hippopotamuses
- †Hippopotamus antiquus – mainland Europe & British Isles; Pleistocene
- †Hippopotamus creutzburgi – Crete; Pleistocene
- †Hippopotamus minor[a] – Cyprus; Pleistocene to Holocene
- †Hippopotamus melitensis – Malta; Pleistocene
- †Hippopotamus pentlandi – Sicily; Pleistocene
- †Hippopotamus laloumena – Madagascar; Holocene
- †Hippopotamus gorgops – Africa Late Miocene–Middle Pleistocene
- Tentatively placed into Hippopotamus:
- †Hippopotamus aethiopicus – Kenya & Ethiopia; Pliocene -Pleistocene
- †Hippopotamus afarensis – Ethiopia, formerly genus Trilobophorus; Pliocene
- †Hippopotamus behemoth – Israel, Early Pleistocene
- †Hippopotamus kaisensis – Central Africa; Pliocene
- †Hippopotamus sirensis – Morocco & Algeria; Pleistocene
- Genus Hexaprotodon – hexaprotodons or Asian hippopotamuses
- †Hexaprotodon bruneti – Ethiopia; Pliocene
- †Hexaprotodon coryndoni – Ethiopia; Pliocene
- †Hexaprotodon crusafonti – Spain; Late Miocene (syn. Hexaprotodon primaevus)
- †Hexaprotodon hipponensis – Algeria
- †Hexaprotodon imagunculus – Uganda and Congo; Pliocene
- †Hexaprotodon iravaticus – Myanmar; Pliocene – Pleistocene
- †Hexaprotodon karumensis – Kenya and Eritrea; Pleistocene
- †Hexaprotodon namadicus – India; (possibly same as Hex. palaeindicus)
- †Hexaprotodon palaeindicus – India;
- †Hexaprotodon pantanellii – Italy; Pliocene
- †Hexaprotodon protamphibius – Kenya and Chad; Pliocene
- †Hexaprotodon siculus –
- †Hexaprotodon sivajavanicus – Indonesia
- †Hexaprotodon sivalensis – India
- †Hexaprotodon sp. (undescribed) – Myanmar
- Genus Archaeopotamus – formerly included in Hexaprotodon
- †Archaeopotamus harvardi – Arabian Peninsula and Central Africa; Miocene – Pliocene
- †Archaeopotamus lothagamensis – Kenya; Miocene
- One or two undescribed species
- Genus Saotherium – formerly included in Hexaprotodon
- †Saotherium mingoz Chad; Pliocene
See also
[edit]Footnotes
[edit]References
[edit]- ^ Laws, Richard (1984). Macdonald, D. (ed.). The Encyclopedia of Mammals. New York: Facts on File. pp. 506–511. ISBN 0-87196-871-1.
- ^ Martino, R.; Pandolfi, L. (3 July 2022). "The Quaternary Hippopotamus records from Italy". Historical Biology. 34 (7): 1146–1156. Bibcode:2022HBio...34.1146M. doi:10.1080/08912963.2021.1965138. ISSN 0891-2963. S2CID 239713930.
- ^ Boisserie, Jean-Renaud; Zazzo, Antoine; Merceron, Gildas; Blondel, Cécile; Vignaud, Patrick; Likius, Andossa; Mackaye, Hassane Taïsso; Brunet, Michel (27 May 2005). "Diets of modern and late Miocene hippopotamids: Evidence from carbon isotope composition and micro-wear of tooth enamel". Palaeogeography, Palaeoclimatology, Palaeoecology. 221 (1–2): 153–174. doi:10.1016/j.palaeo.2005.02.010. Retrieved 18 August 2025 – via Elsevier Science Direct.
- ^ Boisserie, Jean-Renaud; Merceron, Gildas (1 August 2011). "Correlating the success of Hippopotaminae with the C4 grass expansion in Africa: Relationship and diet of early Pliocene hippopotamids from Langebaanweg, South Africa". Palaeogeography, Palaeoclimatology, Palaeoecology. 308 (3–4): 350–361. doi:10.1016/j.palaeo.2011.05.040. Retrieved 20 August 2025 – via Elsevier Science Direct.
- ^ 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.
- ^ Beck, N.R. (2006). "A higher-level MRP supertree of placental mammals". BMC Evol Biol. 6: 93. doi:10.1186/1471-2148-6-93. PMC 1654192. PMID 17101039.
- ^ O'Leary, M.A.; Bloch, J.I.; Flynn, J.J.; Gaudin, T.J.; Giallombardo, A.; Giannini, N.P.; et al. (2013). "The placental mammal ancestor and the post-K-Pg radiation of placentals". Science. 339 (6120): 662–667. Bibcode:2013Sci...339..662O. doi:10.1126/science.1229237. hdl:11336/7302. PMID 23393258. S2CID 206544776.
- ^ Song, S.; Liu, L.; Edwards, S.V.; Wu, S. (2012). "Resolving conflict in eutherian mammal phylogeny using phylogenomics and the multispecies coalescent model". Proceedings of the National Academy of Sciences. 109 (37): 14942–14947. Bibcode:2012PNAS..10914942S. doi:10.1073/pnas.1211733109. PMC 3443116. PMID 22930817.
- ^ dos Reis, M.; Inoue, J.; Hasegawa, M.; Asher, R.J.; Donoghue, P.C.J.; Yang, Z. (2012). "Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny". Proceedings of the Royal Society B: Biological Sciences. 279 (1742): 3491–3500. doi:10.1098/rspb.2012.0683. PMC 3396900. PMID 22628470.
- ^ Upham, N.S.; Esselstyn, J.A.; Jetz, W. (2019). "Inferring the mammal tree: Species-level sets of phylogenies for questions in ecology, evolution, and conservation". PLOS Biology. 17 (12) e3000494. doi:10.1371/journal.pbio.3000494. PMC 6892540. PMID 31800571; (see fig S10).
- ^ a b 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 (HTML abstract). 143 (1): 1–26. doi:10.1111/j.1096-3642.2004.00138.x.
- ^ Upham, Nathan; Burgin, Connor; Widness, Jane; Liphardt, Schuyler; Parker, Camila; Becker, Madeleine; et al. (10 August 2021) [2004]. "Mammal Diversity Database" (vers. 1.6). American Society of Mammalogists. doi:10.5281/zenodo.5175993. Retrieved 28 August 2021.
Further reading
[edit]- "Hippopotamidae". Integrated Taxonomic Information System. Retrieved 11 August 2004.
- Petronio, C. (1995). "Note on the taxonomy of Pleistocene hippopotamuses" (PDF). Ibex. 3: 53–55. Archived from the original (PDF) on 12 September 2008.
Hippopotamidae
View on GrokipediaTaxonomy and Classification
Etymology and nomenclature
The name Hippopotamidae is derived from the Ancient Greek words hippos (ἵππος), meaning "horse," and potamos (ποταμός), meaning "river," thus translating to "river horse family," a nomenclature that aptly reflects the semi-aquatic habits of its members.[4] The family was formally established by British zoologist John Edward Gray in 1821, with Hippopotamus designated as the type genus.[5][6] Within Hippopotamidae, the living members are classified under the subfamily Hippopotaminae (also established by Gray in 1821), which encompasses the two extant genera: Hippopotamus and Choeropsis.[5] The genus Hippopotamus includes the species H. amphibius (common hippopotamus), named by Carl Linnaeus in 1758, where amphibius denotes its dual terrestrial and aquatic lifestyle.[7] The genus Choeropsis, meaning "pig-like face" from Greek choiros (hog) and opsis (appearance), contains the species C. liberiensis (pygmy hippopotamus), described by Samuel G. Morton in 1849 and named for its Liberian origins.[4][8] Extinct subfamilies include Kenyapotaminae, proposed by Martin Pickford in 1983 to accommodate early Miocene forms like Kenyapotamus, highlighting the family's African origins.[9]Phylogenetic position
Hippopotamidae belongs to the order Artiodactyla, the even-toed ungulates, and is positioned within the monophyletic clade Whippomorpha, which unites it with the order Cetacea (whales, dolphins, and porpoises).[10] This placement reflects a shared evolutionary history where hippos represent the closest living relatives to cetaceans among terrestrial mammals, supported by both molecular and morphological data.[11] Key evidence derives from DNA sequencing studies, particularly analyses of mitochondrial genes such as cytochrome b, which consistently show Hippopotamidae as the sister group to Cetacea.[12] These molecular phylogenies indicate that the divergence between the common ancestor of hippos and cetaceans occurred around 55 million years ago, during the early Eocene.[13] Morphological synapomorphies reinforcing this relationship include the even-toed foot structure characteristic of artiodactyls—manifested in hippos as four toes with two primary weight-bearing digits—and a multi-chambered stomach enabling foregut fermentation, akin to ruminant digestion but with three chambers in hippos.[14] Such traits underscore the deep ties within Whippomorpha, distinguishing it from other artiodactyl lineages like ruminants and suines.[15] Historically, Hippopotamidae was misclassified as closely related to suines (pigs and peccaries) based on superficial morphological similarities in dentition and digit reduction.[16] This view persisted until the 1990s, when molecular studies, including pioneering work on cytochrome b sequences, resolved the phylogeny by demonstrating the stronger affinity with cetaceans over suines.[12] Subsequent phylogenomic analyses have further solidified this consensus, integrating fossil and extant data to affirm Whippomorpha's position within Artiodactyla.[17]Living species
The family Hippopotamidae encompasses two extant genera, each containing a single living species: the genus Hippopotamus, represented by the common hippopotamus (H. amphibius), and the genus Choeropsis, represented by the pygmy hippopotamus (C. liberiensis). These two species form sister taxa within the family, sharing a close phylogenetic relationship that diverged during the Pliocene–Early Pleistocene.[18] The common hippopotamus (Hippopotamus amphibius) is classified as Vulnerable on the IUCN Red List.[19] In contrast, the pygmy hippopotamus (Choeropsis liberiensis) is classified as Endangered.[3]Physical Description
Morphology and anatomy
Hippopotamids exhibit a distinctive body plan suited to their semi-aquatic lifestyle, featuring a massive, barrel-shaped torso supported by short, sturdy pillar-like legs that end in broad, hoofed feet with four functional toes. The head is characterized by a broad, flattened muzzle housing a wide gape, while the overall form is nearly hairless in adults, with only sparse bristles on the tail, mane, and mouth edges; juveniles retain finer body hair that is largely shed as they mature. This robust morphology emphasizes weight-bearing efficiency over agility on land.[20] The skin of hippopotamids is exceptionally thick, with dermal layers reaching up to 6 cm in depth across much of the body, providing robust protection against abrasions, sunburn, and microbial invasion in their watery habitats; however, the outermost epidermal layer remains relatively thin and prone to cracking if not kept moist. Embedded within this skin are specialized subdermal mucous glands distributed across the body, particularly densely on the flanks and back, which secrete a viscous, reddish fluid often called "blood sweat." This secretion contains hipposudoric acid and norhipposudoric acid, pigments that rapidly polymerize upon exposure to air, offering natural UV absorption to prevent sunburn and exhibiting broad-spectrum antibiotic activity against bacteria such as Pseudomonas and Escherichia coli.[21][22] Skeletally, hippopotamids display graviportal adaptations, with dense, pachyosteosclerotic bones in the limbs and ribs that enhance structural support for their enormous mass and facilitate neutral buoyancy underwater by increasing overall body density. The limbs are short and columnar, with robust femora and humeri featuring pronounced muscle attachment sites to accommodate powerful propulsion both terrestrially and aquatically. Cranially, the eyes, ears, and nostrils are dorsally positioned high on the skull, enabling the animals to remain mostly submerged while maintaining vigilance and respiration; this arrangement is accentuated in the common hippopotamus, where forward-facing eyes contrast with the more laterally placed orbits in the pygmy species. The permanent dental formula for the common hippopotamus is I 2/2, C 1/1, P 3–4/3–4, M 3/3 (totaling 36–40 teeth), while the pygmy hippopotamus varies slightly as I 2/1, C 1/1, P 4/4, M 3/3 (totaling 38 teeth); both feature continuously growing tusklike canines and incisors adapted for cropping vegetation and occasional defense.[23][24][20][25] Sexual dimorphism is evident in body proportions, with males typically exhibiting larger overall size and more pronounced canine tusks compared to females.[20]Size, weight, and sexual dimorphism
Hippopotamids exhibit significant variation in size and weight between the two living species, with the common hippopotamus (Hippopotamus amphibius) being one of the largest terrestrial mammals. Adult males typically measure 3.5–4.5 m in total length and reach a shoulder height of about 1.5 m, while females are slightly smaller at around 1.4 m in shoulder height.[26] Males weigh 1,300–1,800 kg on average, whereas females range from 1,200–1,500 kg, reflecting moderate sexual dimorphism in body size.[27] This dimorphism is evident in linear dimensions as well, with males averaging 3.12 m in head-body length compared to 3.0 m for females.[26] In contrast, the pygmy hippopotamus (Choeropsis liberiensis) is considerably smaller and displays minimal sexual dimorphism, with sexes appearing nearly alike in size. Adults measure 1.5–1.75 m in length and 0.75–1.0 m in shoulder height, with weights ranging from 180–275 kg for both males and females.[28] This reduced dimorphism contrasts with the common species, where males are consistently larger and heavier, often by 10–20% in mass.[26] Growth in hippopotamids is rapid during the juvenile phase, with calves gaining substantial mass in the first few months; for instance, pygmy hippo young can reach ten times their birth weight (about 4–6 kg) within five months. Both species attain near-adult size by 5–7 years, coinciding with sexual maturity, though common hippo males continue growing incrementally throughout life while females stabilize around age 25.[29][28] Allometric scaling in hippopotamids results in a disproportionate increase in body girth relative to length, yielding a barrel-shaped form that enhances buoyancy in aquatic environments despite their substantial mass.[30] This adaptation supports their semi-aquatic lifestyle, with mass-length relationships emphasizing robustness over elongation.[31]Evolutionary History
Origins and timeline
The clade Whippomorpha, encompassing hippopotamids and cetaceans, diverged from other cetartiodactyls approximately 59 million years ago during the Paleocene-Eocene transition, with the hippopotamid lineage separating from cetaceans around 55 million years ago in the post-Eocene period.[32] This deep phylogenetic split positioned hippopotamids within the broader artiodactyl radiation, evolving from terrestrial ancestors amid the diversification of even-toed ungulates following the Cretaceous-Paleogene extinction.[11] Hippopotamidae emerged in Africa during the early Miocene, around 21 million years ago, originating from anthracothere ancestors within the paraphyletic family Anthracotheriidae, specifically advanced selenodont forms like bothriodontines.[33] The earliest known fossils, such as Morotochoerus ugandensis from Uganda, document this origin in East African ecosystems, marking the family's initial radiation into two subfamilies: the more terrestrial Kenyapotaminae (spanning ~21–15 million years ago) and the later Hippopotaminae around 7.5 million years ago.[33] Fossil evidence extends the family's record to the early Miocene, with diversification accelerating in the late Miocene as anthracothere-like traits transitioned toward the semi-aquatic adaptations seen in later forms.[34] Major evolutionary events included the first dispersal out of Africa to Eurasia approximately 6–7 million years ago during the late Miocene, likely via connections across the Arabian Peninsula, followed by a return migration to Africa and the eventual extinction of non-African lineages in the late Pleistocene, around 40,000–50,000 years ago. Recent paleogenomic studies from 2025 confirm the survival of the common hippopotamus (Hippopotamus amphibius) in central Europe during the last ice age, with remains dated to approximately 47,000 years ago, highlighting greater adaptability to cold climates than previously thought.[33][35] These movements coincided with Miocene climatic shifts toward cooler, drier conditions in Africa, where expanding savannas and seasonal water scarcity drove the evolution of a semi-aquatic niche, enabling hippopotamids to exploit riverine refugia amid habitat fragmentation.[33] This adaptation, briefly tied to shared aquatic traits with cetacean relatives, underscores the family's resilience in fluctuating paleoenvironments.[32]Fossil record
The fossil record of Hippopotamidae extends back to the early Miocene, with the family's emergence in Africa marking a key phase in artiodactyl evolution.[36] Early forms include Kenyapotamus, a genus known from the middle Miocene of East Africa, particularly sites in Kenya such as those yielding dental remains dated to approximately 16 million years ago.[9][37] This small-bodied taxon, estimated at 170–220 kg, exhibited more terrestrial adaptations compared to later relatives, with bunodont dentition suggesting a diet suited to forested environments.[36] Kenyapotamus represents a transitional stage in hippopotamid evolution, bridging primitive anthracothere-like ancestors and more derived semi-aquatic forms.[9] Advanced genera appeared during the late Miocene to Pleistocene, exemplified by Hexaprotodon, which ranged across Eurasia and Africa from roughly 7 million years ago into the late Pleistocene.[38] Fossils of Hexaprotodon, such as H. lothagamensis from late Miocene deposits in Kenya, indicate larger body sizes—up to several tons—and enhanced aquatic traits, including broader snouts and limb modifications for wading.[38] This genus also demonstrates island dwarfism in isolated populations, notably the Malagasy species that evolved reduced sizes, with adults weighing around 200–500 kg, likely as an adaptation to limited island resources during the Pleistocene.[39] Late survivors among extinct hippopotamids include island-endemic species like Hippopotamus lemerlei from Madagascar, which persisted until approximately 1,000 years ago based on radiocarbon-dated subfossil remains showing human modification.[40] These dwarf forms, confined to wetlands and rivers, highlight the vulnerability of insular populations to environmental changes and human arrival.[41] Significant discoveries have illuminated the family's paleobiology, particularly at Toros-Menalla in Chad, a 7-million-year-old late Miocene site that has produced postcranial fossils of early hippopotamids such as Archaeopotamus, revealing semi-aquatic locomotor adaptations and supporting links to cetacean ancestry through shared artiodactyl traits.[42] These finds underscore Africa's role as the cradle of hippopotamid diversification.[43]Relationships to other artiodactyls
Hippopotamidae forms the sister group to Cetacea within the clade Whippomorpha, a relationship supported by molecular and morphological evidence indicating that cetaceans are nested within Artiodactyla as the closest living relatives of hippos.[44] This phylogenetic linkage is reinforced by shared derived traits related to semi-aquatic lifestyles, such as streamlined body forms, multilayered skin with reduced hair follicles, and adaptations for subcutaneous fat storage, though some features like epidermal gland secretions appear to have evolved convergently rather than from a common aquatic ancestor.[45] Unlike fully aquatic cetaceans, hippopotamids retain terrestrial mobility while exhibiting these modifications, highlighting a transitional evolutionary stage.[46] The evolutionary origins of Hippopotamidae trace back to anthracothere lineages, a group of extinct semiaquatic artiodactyls that exhibited hippo-like dental specializations, including bunodont molars suited for processing a mix of vegetation and possibly animal matter.[36] Transitional forms such as Bothriogenys, a late Eocene to early Oligocene anthracothere from Africa, display dentition and cranial features foreshadowing those in modern hippos, suggesting anthracotheres as stem hippopotamoids that migrated into aquatic niches.[34] Fossil evidence from these ancestors supports the anthracothere-hippopotamid connection, with progressive adaptations toward larger body sizes and broader premolars observed across Miocene specimens.[47] In contrast to other artiodactyl families like Suidae (pigs) and Camelidae (camels), hippopotamids lack advanced grazing mechanisms such as multi-chambered stomachs or specialized hypsodont teeth for abrasive forage, instead relying on a simpler digestive system that accommodates occasional omnivory alongside herbivory.[48] Suids exhibit broader omnivorous diets with carnassial-like premolars for meat consumption, while camels, as pseudoruminants, possess foregut fermentation but elongated limbs for arid browsing, features absent in the stocky, riverine-adapted hippos.[49] This divergence underscores hippopotamids' unique emphasis on aquatic foraging over terrestrial grazing dominance seen in suiform or tylopod artiodactyls. Molecular clock analyses estimate the divergence of the hippopotamid-cetacean lineage (Whippomorpha) from other cetartiodactyls, including ruminants, around 59-67 million years ago, aligning with post-Cretaceous radiation of even-toed ungulates.[10][50] These timelines, calibrated using Bayesian relaxed clock models on genomic data, indicate that hippopotamids split from ruminant ancestors approximately 60 million years ago, preceding the Eocene diversification of anthracothere forebears.[33]Distribution and Habitat
Geographic range
The family Hippopotamidae, comprising the common hippopotamus (Hippopotamus amphibius) and the pygmy hippopotamus (Choeropsis liberiensis), is primarily distributed across sub-Saharan Africa today. The common hippopotamus occupies a broad range spanning 37 countries, from Senegal in the west to Somalia in the east, and south to South Africa, inhabiting rivers, lakes, and associated wetlands throughout this region.[19] Its distribution includes key populations in countries such as Angola, Botswana, the Democratic Republic of the Congo, Ethiopia, Kenya, Tanzania, Uganda, Zambia, and Zimbabwe, where it remains relatively abundant in protected areas.[19] In contrast, the pygmy hippopotamus is restricted to a much narrower range in West Africa, with fragmented populations surviving in only four countries: Côte d'Ivoire, Guinea, Liberia, and Sierra Leone, primarily in forested riverine habitats.[51] Historically, hippopotamids exhibited a more extensive distribution beyond their current African stronghold. Fossil evidence indicates that species within the family expanded into Eurasia during the late Miocene, with persistence into the Pleistocene epoch, including records of Hippopotamus antiquus and related forms in Europe until the Late Pleistocene, approximately 31,000 to 47,000 years ago, across regions from the British Isles to the Mediterranean and as far east as Anatolia.[35] In Africa, endemic dwarf hippopotamids, such as Hippopotamus lemerlei, were once widespread on Madagascar but became extinct around 1,000 years ago, following human colonization approximately 2,000 years ago, as evidenced by dated modified bones and associated archaeological layers.[52] The common hippopotamus also formerly occurred in North African regions like Egypt and Algeria, from which it has been extirpated.[19] Over the past century, hippopotamid ranges have undergone significant contractions due to habitat fragmentation, with the common hippopotamus experiencing an estimated 7–20% population decline over the last three generations (as of 2016), leading to isolated subpopulations increasingly confined to protected wetlands.[19] The pygmy hippopotamus has faced even more severe habitat loss, with over 90% of its original forest range in the Upper Guinean ecoregion disappearing since 1900, exacerbating its fragmented distribution.[53] These changes have isolated populations and heightened vulnerability, though core areas in eastern and southern Africa maintain the largest remaining extents for the common species.[19]Environmental preferences and adaptations
Hippopotamids exhibit a strong preference for semi-aquatic environments characterized by shallow, slow-moving freshwater bodies such as rivers, lakes, and swamps, which provide essential refuge during the day. These habitats are typically situated in tropical and subtropical regions of sub-Saharan Africa, where access to permanent or seasonal water sources is critical for thermoregulation and skin protection from solar radiation. Adjacent grasslands or floodplains are necessary to support their terrestrial foraging activities, ensuring a balance between aquatic safety and land-based resource availability.[54][55] The common hippopotamus (Hippopotamus amphibius) thrives in open savannas and wetlands with depths of 1–2 meters, allowing full submersion to avoid overheating in climates where temperatures often exceed 30°C. In contrast, the pygmy hippopotamus (Choeropsis liberiensis) favors densely forested wetlands and riverine areas in West African rainforests, where shaded, moist microhabitats near streams offer cover and reduce exposure to drier conditions. Both species are highly sensitive to droughts, which can fragment habitats and limit water access, leading to stress and reduced viability in altered environments.[56][57] Key aquatic adaptations enable hippopotamids to exploit these watery habitats effectively. Valvular nostrils, positioned dorsally, can seal tightly during submersion, permitting the common hippopotamus to remain underwater for up to 5 minutes while resting or evading threats. Similarly positioned eyes and ears allow sensory monitoring of the surface without full exposure. Their barrel-shaped bodies, supported by dense bones rather than substantial subdermal fat, confer negative buoyancy that facilitates bounding along the substrate at speeds up to 0.5 m/s, rather than true swimming, conserving energy in shallow waters.[23][58][59] On land, hippopotamids depend on proximity to water to mitigate dehydration risks, as their thick, hairless skin secretes a mucous-based sunscreen but lacks functional sweat glands for prolonged cooling. Foraging excursions are thus limited, with individuals typically venturing less than 2 km from aquatic refuges during nocturnal activities to prevent overheating and desiccation. This constraint underscores their vulnerability in fragmented landscapes where water bodies are isolated from suitable grazing areas.[60][30]Behavior and Ecology
Social structure and daily activities
Hippopotamuses, particularly the common hippopotamus (Hippopotamus amphibius), form gregarious social groups known as herds or pods, typically consisting of 10 to 30 individuals, though sizes can reach up to 100 in favorable habitats.[61][26] These herds are polygynous and primarily comprise adult females with their offspring, supplemented by a single dominant territorial male who maintains access to the group for mating purposes.[26] Within the herd, females often occupy the central positions in resting pools and exhibit leadership in group movements, while subadult males and non-territorial adults may associate loosely on the periphery.[27] Unattached adult males typically live solitarily or form temporary bachelor groups away from established territories, avoiding direct competition with dominant individuals.[26] In contrast, the pygmy hippopotamus (Choeropsis liberiensis) is generally solitary, though it may form temporary pairs or small family units consisting of a female and her offspring; males maintain larger territories and wander between areas occupied by females, with limited social interactions outside of mating.[62][63] Pygmy hippos exhibit minimal territorial aggression and do not form large groups like their common relatives.[64] Territorial behavior is prominent among adult males and is confined almost exclusively to aquatic environments, where they defend linear stretches of river or lakeshore ranging from 50 to 100 meters.[61] To demarcate boundaries, males engage in dung showering, rapidly spinning their tails while defecating to disperse feces along paths leading to and from the water, creating distinctive middens that serve as olfactory markers.[61][27] Displays of aggression include wide-mouthed yawns that expose their large canines, up to 50 cm long, as a visual threat to rivals, often accompanied by rearing on hind legs or charging.[27] These behaviors peak during the dry season when water levels concentrate groups, intensifying competition for prime resting sites.[26] Daily activities follow a distinct circadian rhythm adapted to their semi-aquatic lifestyle, with hippopotamuses spending the daylight hours submerged in rivers, lakes, or pools for thermoregulation and protection from the sun, emerging only briefly to bask or reposition.[61][27] At dusk, herds exit the water along well-trodden paths, dispersing to graze nocturnally for 4 to 5 hours, covering distances of 3 to 5 km individually in search of vegetation, though this activity is largely solitary despite the social nature of daytime resting.[61][27] In areas of resource scarcity, individuals may travel up to 8 km or more, returning to water before dawn to avoid dehydration and predation risks.[61] Pygmy hippopotamuses are primarily nocturnal and secretive, spending days resting or submerged in rivers, forested swamps, or wallows to avoid heat and predators, before emerging at night to forage for several hours (typically around 6 hours) on land within their territories.[63][65] Communication among hippopotamuses relies on a repertoire of vocalizations, visual signals, and olfactory cues to maintain group cohesion and deter intruders over distances exceeding 1 km.[61][27] Key sounds include deep grunts, wheeze-honks—loud calls reaching 115 decibels—and bellows produced both above and below water, with about 80% of vocalizations occurring submerged for efficient transmission in aquatic environments.[27] These low-frequency calls, with fundamentals as low as 20–60 Hz, incorporate infrasonic components that propagate far through air and water, enabling long-distance recognition of familiar group members versus strangers.[66][67] Visual displays, such as yawning or open-mouth threats, reinforce these signals during close encounters, while dung and urine marking provide persistent chemical communication.[61]Diet, feeding, and interactions
Hippopotamids are strictly herbivorous, with diets dominated by plant matter that reflects their respective habitats. The common hippopotamus (Hippopotamus amphibius) primarily consumes grasses, comprising over 80% of its intake, as evidenced by stable isotope analysis of tusks from historical specimens. Adults forage nocturnally, consuming 35–50 kg of vegetation per night, equivalent to about 1–2% of their body mass. In contrast, the pygmy hippopotamus (Choeropsis liberiensis) maintains a more varied diet as a generalist intermediate feeder, incorporating shrubs, grasses, ferns, broad-leaved plants, herbaceous shoots, and fallen fruits, with minimal grass consumption compared to its larger relative.[68][69][70][71] Feeding occurs primarily on land during nocturnal excursions from aquatic refuges, where individuals use their broad, muscular lips to grasp vegetation and crop it close to the ground by swinging their heads side to side, aided by lower incisors for shearing. Unlike ruminants, hippopotamids exhibit minimal rumination, relying instead on foregut fermentation in a multi-chambered stomach where microbial activity breaks down fibrous plant material. This digestive strategy supports efficient processing of low-quality forage but results in lower digestibility rates than in true ruminants, with ingesta retention times of 24–48 hours.[68][72][73] As large grazers, hippopotamids play a key role in shaping savanna and wetland vegetation through selective cropping, which prevents overgrowth and maintains short-grass landscapes favorable to other herbivores. Their dung, rich in nutrients like phosphorus and nitrogen, facilitates nutrient cycling by transporting terrestrial organic matter into aquatic systems, subsidizing primary production and supporting food webs in rivers and pools—though excessive loading in high-density areas can lead to hypoxic conditions.[69][74] Ecological interactions include conflicts with humans, particularly crop raiding by common hippos, which damage agricultural fields near water bodies and account for the majority of reported human-wildlife conflicts in affected regions. Predation on hippopotamids is rare due to their size and aggression, but Nile crocodiles (Crocodylus niloticus) occasionally target calves or weakened individuals in shared aquatic habitats.[75][76]Reproduction and development
Hippopotamids exhibit a polygynous mating system, in which dominant males secure access to multiple females within their territories, influenced by social hierarchies that determine reproductive success.[77] Mating occurs primarily in water and is not strictly seasonal, though conceptions peak during the dry season, aligning births with the wet season for optimal conditions.[54] In the common hippopotamus (Hippopotamus amphibius), gestation lasts approximately 8 months (227–240 days), after which females typically give birth to a single calf in shallow water to protect against predators.[78] The newborn calf weighs 25–55 kg and can swim immediately, though it relies on the mother for buoyancy and nursing.[78] Females isolate themselves from the group for several days before and after birth to ensure safety, providing exclusive maternal care while males play no role in rearing.[79] Nursing continues for about 8 months, after which calves begin grazing but remain dependent on the mother for up to 2 years.[78] Sexual maturity is reached by females at 3–6 years and males at 6–13 years, with wild individuals living 40–50 years.[80] The pygmy hippopotamus (Choeropsis liberiensis) shares a similar reproductive pattern but with adaptations to its more solitary, forest-dwelling lifestyle. Gestation spans 6–7 months (184–204 days), resulting in litters of one calf, rarely two, born on land in dense vegetation.[81] At birth, calves weigh 3.4–6.4 kg and receive intensive maternal care, including nursing for 6–8 months, as females remain highly protective and males are uninvolved.[62] Sexual maturity occurs around 4–5 years for both sexes, and wild lifespan extends up to 42 years.[82]Conservation
Current status and threats
The common hippopotamus (Hippopotamus amphibius) is classified as Vulnerable on the IUCN Red List, with a global population estimated at 125,000–148,000 individuals as of 2025 and an overall decreasing trend.[83] Assessments indicate a suspected decline of more than 20% over the past three generations (approximately 30 years), though recent data from the 2020s suggest range-wide annual declines of 6–8% in many regions, equating to roughly 7–20% over a decade in affected areas.[19][84] The pygmy hippopotamus (Choeropsis liberiensis) is listed as Endangered, with a population of 2,000–3,000 mature individuals also showing a decreasing trend, driven by ongoing habitat fragmentation and low reproductive rates.[3][83] Primary threats to both species include habitat loss from agricultural expansion, dam construction, and deforestation, which reduce access to rivers, wetlands, and grazing areas essential for their semi-aquatic lifestyle.[19][3] Poaching for bushmeat and ivory from their large canines persists, particularly in conflict zones, contributing to localized population crashes.[19] Human-wildlife conflict exacerbates declines, as hippos raiding crops lead to retaliatory killings; hundreds of common hippos are shot annually across Africa due to such incidents.[85] Diseases like anthrax outbreaks further impact populations, with recent events in 2025 killing at least 50 common hippos in Virunga National Park, Democratic Republic of Congo.[86] Pollution from mining and agricultural runoff degrades water quality, increasing vulnerability to infections and disrupting aquatic habitats.[3] Regional variations intensify risks, with pygmy hippo populations fragmented across West Africa and a total estimate of 2,000–3,000 individuals remaining, many in small, isolated groups due to intensive logging and forest clearance for cash crops.[3][87] Common hippo subpopulations in the same region have declined by over 30% in the last decade from similar pressures, compounded by civil unrest facilitating poaching.[19] These factors fragment habitats across their sub-Saharan ranges, limiting dispersal and genetic exchange.[84]Protection measures and future outlook
Conservation efforts for Hippopotamidae focus on establishing and maintaining protected areas to safeguard habitats critical for both the common hippopotamus (Hippopotamus amphibius) and the pygmy hippopotamus (Choeropsis liberiensis). Significant portions of their ranges overlap with national parks and reserves, including Virunga National Park in the Democratic Republic of the Congo, which supports recovering hippo populations through anti-poaching enforcement, and Niokolo-Koba National Park in Senegal, a UNESCO World Heritage site preserving riverine habitats for the species.[88][89] Both species are regulated under CITES Appendix II, which controls international trade in hippo parts like teeth and skins to prevent overexploitation.[90] Key initiatives include anti-poaching patrols and community-based education programs implemented across more than 20 African countries where hippos occur, aiming to reduce human-wildlife conflicts and illegal hunting.[85][91] In 2025, the IUCN SSC Hippo Specialist Group published a report emphasizing the need for enhanced monitoring of declining pygmy populations and local abundance in common hippos, and a workshop developed an action plan for common hippos in West Africa.[87][92] Reintroduction efforts have shown promise, particularly in South Africa, where hippos have been successfully relocated to reserves such as Amakhala Game Reserve and Botlierskop Game Reserve to bolster local populations and restore ecological roles in aquatic systems.[93][94] Ongoing research emphasizes genetic studies to evaluate inbreeding risks and population connectivity, informing breeding programs in both wild and captive settings to enhance genetic diversity.[95] Habitat restoration projects target wetland rehabilitation, such as those in community-managed sanctuaries like Wechiau Hippo Sanctuary in Ghana, which integrate local involvement to protect riverine environments essential for hippo survival.[96] Future projections indicate continued vulnerability, with common hippo populations declining at 6-8% annually in some regions due to ongoing threats like poaching, potentially leading to substantial losses by 2050 without intensified interventions.[84] However, optimistic scenarios suggest population recovery is achievable through expanded protected areas, climate-resilient habitat management, and transfrontier conservation efforts that promote connectivity across borders.[97][98]References
- https://en.wiktionary.org/wiki/Choeropsis