Hippopotamus: Difference between revisions
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{{Short description|Large semi-aquatic mammal native to sub-Saharan Africa}} |
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{{redirect|Hippo}} |
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{{Redirect|Hippo|its genus|Hippopotamus (genus)|other uses|Hippopotamus (disambiguation)|and|Hippo (disambiguation)}} |
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{{Featured article}} |
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{{Taxobox |
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{{Use British English|date=March 2022}} |
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| name = Common hippopotamus |
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{{Use dmy dates|date=March 2022}} |
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{{Speciesbox |
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| italic_title = no |
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| name = Hippopotamus |
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| fossil_range = {{fossilrange|Pleistocene|Recent}} |
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| image = Portrait Hippopotamus in the water.jpg |
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| image_caption = A hippopotamus in [[Saadani National Park]], Tanzania |
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| status = VU |
| status = VU |
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| status_system = |
| status_system = IUCN3.1 |
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| status_ref = <ref name=iucn>{{cite iucn |author=Lewison, R. |author2=Pluháček, J. |date=2017 |title=''Hippopotamus amphibius'' |page=e.T10103A18567364 |doi=10.2305/IUCN.UK.2017-2.RLTS.T10103A18567364.en |access-date=19 November 2021}}</ref> |
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| status_ref = <ref name="Redlist"/> |
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| status2 = CITES_A2 |
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| status2_system = CITES |
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| image = Hippopotamus - 04.jpg |
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| status2_ref = <ref name=iucn/> |
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| image_width = 250px |
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| genus = Hippopotamus |
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| species = amphibius |
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| phylum = [[Chordate|Chordata]] |
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| authority = [[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]<ref name=Linnæus>{{cite book |author=Linnæus, C. |year=1758 |title=Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis |publisher=Salvius |location=Holmiae |chapter=''Hippopotamus amphibius'' |page=74 |chapter-url=https://archive.org/details/mobot31753000798865/page/n87/mode/2up}}</ref> |
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| classis = [[Mammal]]ia |
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| range_map = Hippopotamus_distribution.png |
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| ordo = [[Even-toed ungulate|Artiodactyla]] |
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| range_map_caption = Current distribution of the hippopotamus |
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| familia = [[Hippopotamidae]] |
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}} |
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| genus = ''[[Hippopotamus (genus)|Hippopotamus]]'' |
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The '''hippopotamus''' ('''''Hippopotamus amphibius''''') ({{IPAc-en|ˌ|h|ɪ|p|ə|ˈ|p|ɒ|t|ə|m|ə|s}}; {{plural form}}: '''hippopotamuses'''; often shortened to '''hippo''' ({{plural form}}: '''hippos'''), further qualified as the '''common hippopotamus''', '''Nile hippopotamus''' and '''river hippopotamus''', is a large [[semiaquatic]] mammal native to [[sub-Saharan Africa]]. It is one of only two [[extant taxon|extant]] [[species]] in the [[scientific classification|family]] [[Hippopotamidae]], the other being the [[pygmy hippopotamus]] (''Choeropsis liberiensis'' or ''Hexaprotodon liberiensis''). Its name comes from the [[ancient Greek]] for "river horse" ({{lang|grc|ἱπποπόταμος}}). |
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| genus_authority = [[Carl Linnaeus|Linnaeus]], 1758 |
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| species = '''''H. amphibius''''' |
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| binomial = ''Hippopotamus amphibius'' |
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| binomial_authority = [[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]<ref>{{cite web|url = http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=625024 |title= ITIS on Hippopotamus amphibius|accessdate = 2007-07-29|work = ''[[Integrated Taxonomic Information System]]''|archiveurl= https://web.archive.org/web/20140826161420/http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=625024|archivedate= 2014-08-26|deadurl= no}}</ref> |
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| range_map=hippo_distribution.gif |
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| range_map_caption= Range map of hippopotamus. Historic range is in red while current range is in green.<ref name="Redlist"/> |
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| range_map_width = 230px}} |
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After [[elephant]]s and [[rhinoceros]]es, the hippopotamus is the next [[Largest mammals#Even-toed Ungulates (Artiodactyla)|largest land mammal]]. It is also the largest extant land [[artiodactyl]]. Despite their physical resemblance to [[pig]]s and other terrestrial even-toed ungulates, the closest living relatives of the hippopotamids are [[cetacea]]ns ([[whale]]s, [[dolphin]]s, [[porpoise]]s, etc.), from which they diverged about 55 million years ago. Hippos are recognisable for their barrel-shaped torsos, wide-opening mouths with large [[Canine tooth|canine]] tusks, nearly hairless bodies, pillar-like legs, and large size: adults average {{cvt|1500|kg}} for bulls (males) and {{cvt|1300|kg|}} for cows (females). Despite its stocky shape and short legs, it is capable of running {{cvt|30|km/h}} over short distances. |
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Hippos inhabit rivers, lakes, and [[mangrove]] swamps. [[Territorial animal|Territorial]] bulls each preside over a stretch of water and a group of five to thirty cows and calves. [[Mating]] and birth both occur in the water. During the day, hippos remain cool by staying in water or mud, emerging at dusk to graze on grasses. While hippos rest near each other in the water, grazing is a solitary activity and hippos typically do not display territorial behaviour on land. Hippos are among the most dangerous animals in the world due to their aggressive and unpredictable nature. They are threatened by [[habitat loss]] and [[poaching]] for their meat and [[ivory]] (canine teeth). |
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Common hippos are recognizable by their barrel-shaped torsos, wide-opening mouths revealing large canine tusks, nearly hairless bodies, columnar-like legs and large size; adults average {{convert|1500|kg|lbs|abbr=on}} and {{convert|1300|kg|lbs|abbr=on}} for males and females respectively. Despite its stocky shape and short legs, it is capable of running {{convert|30|km/h|abbr=on}} over short distances. The hippopotamus is a highly aggressive and unpredictable animal and is ranked among the most dangerous animals in Africa.<ref>{{cite web |url= http://animals.nationalgeographic.com/animals/wild/shows-deadly-60/fun-facts/|title= Deadly 60: 15 Deadly Animal Facts|publisher= ''[[National Geographic (magazine)|National Geographic]]''|archiveurl= https://web.archive.org/web/20141016145736/http://animals.nationalgeographic.com/animals/wild/shows-deadly-60/fun-facts/|archivedate= 2014-10-16|deadurl= no}}</ref> Nevertheless, they are still threatened by habitat loss and poaching for their meat and [[ivory]] canine teeth. |
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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 30 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 [[Territory (animal)|territorial]] on land. |
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==Etymology== |
==Etymology== |
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The word |
The [[Latin]] word {{lang|la|hippopotamus}} is derived from the ancient [[Greek language|Greek]] {{lang|grc|ἱπποπόταμος}} ({{Transliteration|grc|hippopótamos}}), from {{lang|grc|ἵππος}} ({{Transliteration|grc|híppos}}) {{gloss|horse}} and {{lang|grc|ποταμός}} ({{Transliteration|grc|potamós}}) {{gloss|river}}, together meaning {{gloss|horse of the river}}.<ref>{{LSJ|i(ppopo/tamos|ἱπποπόταμος}}, {{LSJ|i(/ppos|ἵππος}}, {{LSJ|potamo/s|ποταμός|ref}}.</ref><ref>{{cite encyclopedia|url=http://www.merriam-webster.com/dictionary/hippopotamus|title=Hippopotamus|dictionary= [[Webster's Dictionary|Merriam-Webster's Online Dictionary]]|access-date=18 July 2007}}</ref><ref>{{OEtymD|hippopotamus}}</ref> In English, the [[English plural|plural]] is "hippopotamuses".<ref>{{cite encyclopedia|url=http://www.askoxford.com/asktheexperts/faq/aboutgrammar/plurals?view=uk|title=Plural of hippopotamus|dictionary=[[Oxford English Dictionary]]|access-date=18 July 2007|archive-date=13 October 2008|archive-url=https://web.archive.org/web/20081013041855/http://www.askoxford.com/asktheexperts/faq/aboutgrammar/plurals?view=uk|url-status=dead}}</ref> |
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In Africa, the hippo is known by various names, including ''seekoei'' ([[Afrikaans]]), ''mvuvu'' ([[Venda language|Venda]]), ''kubu'' ([[Lozi language|Lozi]]) and ''mvubu'' ([[Xhosa language|Xhosa]], [[Swazi language|Siswati]] and [[Zulu language|Zulu]]) in the south;<ref>{{cite book|author=Walker, C.|year=1997|title=Signs of the Wild|publisher=Struik|page=140|isbn=1-86825-896-3}}</ref> ''kiboko'' ([[Swahili language|Swahili]]), e''nsherre'' ([[Nkore language|Nkore]]), ''tomondo'' ([[Turu language|Turu]]), ''nvubu'' ([[Luganda]]), ''ifuru'' ([[Luhya language|Luhya]]), ''emiria'' ([[Teso language|Ateso]]), ''magawit'' ([[Elgon languages|Sebei]]), ''kibei'' ([[Nandi–Markweta languages|Kalenjin]]) and ''olmakau'' ([[Maasai language|Maasai]]) in the [[African Great Lakes]] region;<ref name=kingdon/>{{rp|256}} and ጉማርረ/''gumarre'' ([[Amharic language|Amharic]])<ref name="Lkane">{{cite book|last=Kane|first=Thomas Leiper|title=Amharic-English dictionary: H – N., Volume 1|year=1990|publisher=Otto Harrassowitz Verlag|isbn=3-447-02871-8|page=1909|url=http://books.google.com/books?id=H6tnix8o0mwC}}</ref> and ''jeer'' ([[Somali language|Somali]]) in the [[Horn of Africa]].<ref name="Saeed">{{cite book|last=Saeed|first=John I.|title=Somali|year=1999|publisher=[[John Benjamins Publishing Company]]|isbn=90-272-3810-3|page=29|url=http://books.google.com/books?id=EqCNXXEcoBgC&pg=PA29}}</ref> |
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==Taxonomy and origins== |
==Taxonomy and origins== |
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===Classification=== |
===Classification=== |
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{{multiple image |
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The hippopotamus is the [[Types in zoology|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.<ref name="The Hippos"/>{{rp|39}} Hippopotamidae are classified along with other [[even-toed ungulates]] in the [[order (biology)|order]] [[Artiodactyla]]. Other artiodactyls include [[camels]], [[cows|cattle]], [[deer]] and [[pigs]], although hippopotamuses are not closely related to these groups. |
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| direction = vertical |
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| image1 = Hipopótamo (Hippopotamus amphibius), parque nacional de Chobe, Botsuana, 2018-07-28, DD 60.jpg |
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|alt1=Closeup photo, top of head, feeding while partially submerged |
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| image2 = Hippopotamus amphibius 3d scan Natural History Museum University of Pisa C 228.stl |
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|alt2=Interactive 3D partial skull scan |
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| image3 = Hippopotamus amphibius (mandible) 3d scan Natural History Museum University of Pisa C228.stl |
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|alt3=Interactive 3D scan of the remaining mandible |
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| footer = Head, skull and mandible of ''Hippopotamus amphibius'' |
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}} |
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The modern hippopotamus and the [[pygmy hippopotamus]] are the only living members of the family Hippopotamidae. Some taxonomists place hippos and [[anthracotheres]] in the superfamily Anthracotheroidea. Hippopotamidae are classified along with other [[even-toed ungulates]] in the [[order (biology)|order]] [[Artiodactyla]].<ref name="The Hippos"/>{{rp|39–40}} |
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Five subspecies of hippos have been described based on [[morphology (biology)|morphological]] differences in their skulls as well as differences in geographical range:<ref name="The Hippos"/>{{rp|3}}<ref>{{cite book|author=Lydekker, R|year=1915|title=Catalogue of the Ungulate Mammals in the British Museum of Natural History, vol. 4|volume=4 |publisher=British Museum|pages=389–392|url=https://www.biodiversitylibrary.org/item/114771#page/417/mode/1up}}</ref><ref name="Okello"/><ref>{{Cite journal|title = Suiform Soundings: The IUCN/SSC Pigs, Peccaries, and Hippos Specialist Group (PPHSG) Newsletter|volume = 5|issue = 1|date=September 2005|url = https://www.scribd.com/document/65917875/Newsletter-5-1 |format=PDF |journal = [[World Conservation Union|IUCN]]|editor = Meijaard, Erik}}</ref> |
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[[File:Pygmy Hippopotamus (Hexaprotodon liberiensis) (cropped).jpg|thumb|right|A [[pygmy hippopotamus]] (''Choeropsis liberiensis'').]] |
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*''H. a. amphibius'' – (the nominate subspecies) ranges from [[Gambia]] east to [[Ethiopia]] and then south to [[Mozambique]] and historically ranged as far north as Egypt; its skull is distinguished by a moderately reduced preorbital region, a bulging dorsal surface, elongated [[mandibular symphysis]] and larger chewing teeth. |
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*''H. a. kiboko'' – found in [[Kenya]] and [[Somalia]]; was noted to be smaller and more lightly coloured than other hippos with wider nostrils, somewhat longer snout and more rounded and relatively raised [[Orbit (anatomy)|orbit]]s with the space between them being incurved. |
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Five subspecies of hippos have been described based on [[morphology (biology)|morphological]] differences in their skulls and geographical differences:<ref name="The Hippos"/>{{rp|3}} |
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*''H. a. capensis'' – found in [[Zambia]] and [[South Africa]]; distinguished by wider orbits. |
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*''H. a. amphibius'' – (the nominate subspecies) which stretched from [[Egypt]], where they are now extinct, south up the [[Nile River]] to [[Tanzania]] and [[Mozambique]] |
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*''H. a. |
*''H. a. tschadensis'' – ranges between [[Chad]] and [[Niger]]; featured a slightly shorter but broader face, and pronounced, forward-facing orbits. |
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*''H. a. constrictus'' – ranged from the southern [[Democratic Republic of Congo]] to [[Angola]] and [[Namibia]]; skull characterised by a thicker preorbital region, shorter snout, flatter dorsal surface, reduced mandibular symphysis and smaller chewing teeth. |
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*''H. a. capensis'' – from Zambia to [[South Africa]], most flattened skull of the subspecies |
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The suggested subspecies above 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.<ref name="The Hippos">{{cite book |last=Eltringham |first=S. K. |title=The Hippos: Natural History and Conservation |year=1999 |publisher=Academic Press |isbn=978-0-85661-131-5 |series=Poyser Natural History Series}}</ref>{{rp|2}} A study examining [[mitochondrial DNA]] from skin biopsies taken from 13 sampling locations 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'' have been tested.<ref name=Okello>{{Cite journal |author=Okello, J. B. A. |author2=Nyakaana, S. |author3=Masembe, C. |author4=Siegismund, H. R. |author5=Arctander, P. |year=2005 |title=Mitochondrial DNA variation of the common hippopotamus: evidence for a recent population expansion |journal=[[Heredity (journal)|Heredity]] |pmid=16030528 |volume =95 |issue=3 |pages=206–215 |doi=10.1038/sj.hdy.6800711 |doi-access=free}}</ref> |
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*''H. a. tschadensis'' – throughout Western Africa to, as the name suggests, [[Chad]], slightly shorter and wider face, with prominent orbits |
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*''H. a. constrictus'' – in [[Angola]], the southern [[Democratic Republic of Congo]] and [[Namibia]], named for its deeper preorbital constriction |
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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.<ref name="The Hippos">{{cite book |last=Eltringham |first=S.K. |title= The Hippos|year=1999 |publisher=Academic Press |isbn=0-85661-131-X |series= Poyser Natural History Series}}</ref>{{rp|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.<ref name="Okello">{{Cite journal|author = Okello, J.B.A, Nyakaana, S., Masembe, C., Siegismund, H.R. an Arctander, P.|year = 2005|title = Mitochondrial DNA variation of the common hippopotamus: evidence for a recent population expansion.|journal = [[Heredity (journal)|Heredity]]|pmid = 16030528|volume = 95|issue = 3|pages = 206–215|doi = 10.1038/sj.hdy.6800711}}</ref><ref>{{Cite journal|title = Suiform Soundings: The IUCN/SSC Pigs, Peccaries, and Hippos Specialist Group (PPHSG) Newsletter|volume = 5|issue = 1|date=September 2005|url = http://www.iucn.org/themes/ssc/sgs/pphsg/Suiform%20soundings/Newsletter%205(1).pdf |format=PDF |archiveurl=http://web.archive.org/web/20080308192646/http://www.iucn.org/themes/ssc/sgs/pphsg/Suiform+soundings/Newsletter+5%281%29.pdf |archivedate=2008-03-08|work = [[World Conservation Union|IUCN]]|author = Meijaard, Erik (ed.)}}</ref> |
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===Evolution=== |
===Evolution=== |
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[[File:Evolutionary relationships among laurasiatherian mammals as used in molecular evolution analyses.jpg|thumb|Evolutionary relationships among hippo and Cetacea (whales, dolphins)<ref name="TsagkogeorgaMcGowen2015">{{cite journal |last1=Tsagkogeorga |first1=G. |last2=McGowen |first2=M. R. |last3=Davies |first3=K. T. J.|last4=Jarman |first4=S. |last5=Polanowski |first5=A. |last6=Bertelsen |first6=M. F. |last7=Rossiter |first7=S. J. |title=A phylogenomic analysis of the role and timing of molecular adaptation in the aquatic transition of cetartiodactyl mammals |journal=Royal Society Open Science |volume=2 |issue=9 |year=2015 |page=150156 |doi=10.1098/rsos.150156 |pmid=26473040 |pmc=4593674 |bibcode=2015RSOS....250156T}}</ref>]] |
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Until 1909, [[natural history|naturalists]] grouped hippos with [[pig]]s, based on [[molar (tooth)|molar]] patterns. Several lines of evidence, first from [[blood]] [[protein]]s, then from [[molecular systematics]]<ref name="Genomes"/> and [[DNA]] |
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<ref name= |
Until 1909, [[natural history|naturalists]] classified hippos together with pigs based on [[molar (tooth)|molar]] patterns. Several lines of evidence, first from blood proteins, then from [[molecular systematics]],<ref name=Genomes/> [[DNA]]<ref name=DNA>{{Cite journal |title=More DNA support for a Cetacea/Hippopotamidae clade: the blood-clotting protein gene gamma-fibrinogen |author=Gatesy, J. |journal=[[Molecular Biology and Evolution]] |date=1997 |volume=14 |pages=537–543 |pmid=9159931 |issue=5 |doi=10.1093/oxfordjournals.molbev.a025790 |doi-access=free}}</ref><ref name=Nature2009>{{Cite journal |title=Hippopotamus and whale phylogeny |author1=Geisler, J. H. |author2=Theodor, J. M. |journal=[[Nature (journal)|Nature]] |year=2009 |pmid=19295550 |volume=458 |doi=10.1038/nature07776 |bibcode=2009Natur.458....1G |issue=7236 |pages=E1–E4; discussion E5 |s2cid=4320261}}</ref> and the [[fossil record]], show their closest living relatives are [[cetacean]]s ([[whale]]s, [[dolphin]]s, and [[porpoise]]s).<ref name=ScienceNews>{{cite web |first=R. |last=Sanders |title=Scientists find missing link between the dolphin, whale and its closest relative, the hippo |date=2005 |archive-url= https://web.archive.org/web/20150226060713/http://www.innovations-report.com//html/reports/life-sciences/report-39309.html |archive-date= 26 February 2015 |url-status=live |access-date=8 January 2011 |url=http://www.innovations-report.com/html/reports/life_sciences/report-39309.html |work=Science News Daily}}</ref><ref name=Cetartiodactyla/> 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.<ref name="DNA"/><ref name=Cetartiodactyla>{{cite journal |author1=Boisserie, J.-R. |author2=Lihoreau, F. |author3=Brunet, M. |year=2005 |title=The position of Hippopotamidae within Cetartiodactyla |journal=[[Proceedings of the National Academy of Sciences]] |volume=102 |issue=5 |pages=1537–1541 |doi=10.1073/pnas.0409518102 |pmid=15677331 |pmc=547867 |bibcode=2005PNAS..102.1537B |doi-access=free}}</ref> |
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{{Cetartiodactyla Cladogram}} |
{{Cetartiodactyla Cladogram}} |
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[[File: |
[[File:Anthracotherium magnum.jpg|thumb|right|''[[Anthracotherium]] magnum'' from the Oligocene of Europe]] |
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The most recent theory of the origins of Hippopotamidae suggests |
The most recent theory of the origins of Hippopotamidae suggests hippos and whales shared a common semiaquatic ancestor that branched off from other artiodactyls around {{mya|60|million years ago}}.<ref name=DNA/><ref name=ScienceNews/> This hypothesised ancestral group likely split into two branches again around {{mya|54|million years ago}}.<ref name=Genomes>{{Cite journal |title=Analyses of mitochondrial genomes strongly support a hippopotamus-whale clade |volume=265 |issue=1412 |year=1998 |journal=[[Proceedings of the Royal Society]] |pmid=9881471 |author1=Ursing, B. M. |author2=Arnason U. |pmc=1689531 |doi=10.1098/rspb.1998.0567 |pages=2251–2255}}</ref> |
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One branch would [[Evolution of cetaceans|evolve into cetaceans]], possibly beginning about {{mya|52|million years ago}}, with the protowhale ''[[Pakicetus]]'' and other early whale ancestors collectively known as [[Archaeoceti]]. This group eventually underwent [[aquatic adaptation]] into the completely aquatic [[cetacea]]ns.<ref name=Cetartiodactyla/> The other branch became the [[anthracotheriidae|anthracotheres]], a large family of four-legged beasts, the earliest of which in the late [[Eocene]] would have resembled skinny hippos with comparatively smaller, narrower heads. All branches of the anthracotheres, except that which evolved into [[Hippopotamidae]], became extinct during the [[Pliocene]], leaving no descendants.<ref name=ScienceNews/><ref name=Cetartiodactyla/> |
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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]].<ref name="Origins">{{cite journal|last= Boisserie|first=Jean-Renaud|author2=Lihoreau, Fabrice|author3=Brunet, Michel |year=2005 |title=Origins of Hippopotamidae (Mammalia, Cetartiodactyla): towards resolution |journal=[[Zoologica Scripta]] |volume= 34|issue= 2|pages=119–143|doi = 10.1111/j.1463-6409.2005.00183.x}}</ref> ''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 {{Mya|20|million years ago}}. 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 {{mya|8|million years ago}}. 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 {{mya|1.8|million years ago}}, an ancestor to the modern hippopotamus, ''[[Archaeopotamus]]'', lived in Africa and the Middle East.<ref name="Linnean">{{cite journal |last=Boisserie |first=Jean-Renaud |year=2005 |title=The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis |journal=[[Zoological Journal of the Linnean Society]] |volume= 143 |pages= 1–26|doi = 10.1111/j.1096-3642.2004.00138.x}}</ref> |
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A rough evolutionary lineage of the hippo can thus be traced from Eocene and [[Oligocene]] species: from ''[[Anthracotherium]]'' and ''[[Elomeryx]]'' to the [[Miocene]] species ''[[Merycopotamus]]'' and ''[[Libycosaurus]]'' and finally the very latest anthracotheres in the [[Pliocene]].<ref name=Origins>{{cite journal |last=Boisserie |first=J.-R. |author2=Lihoreau, F. |author3=Brunet, M. |year=2005 |title=Origins of Hippopotamidae (Mammalia, Cetartiodactyla): towards resolution |journal=[[Zoologica Scripta]] |volume=34 |issue=2 |pages=119–143 |doi=10.1111/j.1463-6409.2005.00183.x |s2cid=83768668}}</ref> These groups lived across Eurasia and Africa. The discovery of ''Epirigenys'' in East Africa, which was likely a descent of Asian anthracotheres and a [[sister taxon]] to Hippopotamidae, suggests that hippo ancestors entered Africa from Asia around {{Mya|35|million years ago}}.<ref>{{cite journal|author1=Lihoreau, F. |author2=Boisserie, J.-R. |author3=Manthi, F. K. |author4=Ducrocq. S. |year=2015 |title=Hippos stem from the longest sequence of terrestrial cetartiodactyl evolution in Africa |journal=Nature Communications |volume=6 |page=6264 |doi=10.1038/ncomms7264 |pmid=25710445 |bibcode=2015NatCo...6.6264L |doi-access=free}}</ref><ref>{{cite web |author=Lihoreau, F. |date=2015 |title=New Fossils from Kenya Precise Hippo Origin |publisher=Serious Science |access-date=5 June 2021|url=http://serious-science.org/new-fossils-from-kenya-precise-hippo-origin-2675}}</ref> An early hippopotamid is the genus ''[[Kenyapotamus]]'', which lived in Africa from 15 to {{mya|9|million years ago}}.<ref name=Origins/> Hippopotamid species would spread across Africa and Eurasia, including the modern pygmy hippo. From 7.5 to {{mya|1.8|million years ago}}, a possible ancestor to the modern hippo, ''[[Archaeopotamus]]'', lived in Africa and the Middle East.<ref name=Linnean>{{cite journal |last=Boisserie |first=J.-R. |year=2005 |title=The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis |journal=[[Zoological Journal of the Linnean Society]] |volume=143 |pages=1–26 |doi=10.1111/j.1096-3642.2004.00138.x |doi-access=free}}</ref> The oldest records of the genus ''[[Hippopotamus (genus)|Hippopotamus]]'' date to the [[Pliocene]] (5.3–2.6 million years ago).<ref name=VanDerMadeEtAl2017/> The oldest unambiguous records of the modern ''H. amphibius'' date to the [[Middle Pleistocene]], though there are possible [[Early Pleistocene]] records.<ref>{{Cite journal |last1=Pandolfi |first1=L. |last2=Martino |first2=R. |last3=Belvedere |first3=M. |last4=Martínez-Navarro |first4=B. |last5=Medin |first5=T. |last6=Libsekal |first6=Y. |last7=Rook |first7=L. |date=2023 |title=The latest Early Pleistocene hippopotami from the human-bearing locality of Buia (Eritrea) |journal=Quaternary Science Reviews |volume=308 |pages=108039 |doi=10.1016/j.quascirev.2023.108039|bibcode=2023QSRv..30808039P |s2cid=258024770}}</ref> |
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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 {{mya|8|million years ago}}. Taxonomists disagree whether or not the modern [[Pygmy Hippopotamus|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 (phylogenetics)|basal]] genus.<ref name="Origins"/><ref name="Linnean"/> |
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[[File:Malagasy Hippopotamus.jpg|thumb|right|''Choeropsis madagascariensis'' skeleton with a modern hippopotamus skull |
[[File:Malagasy Hippopotamus.jpg|thumb|right|''Choeropsis madagascariensis'' skeleton with a modern hippopotamus skull]] |
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===Extinct species=== |
===Extinct species=== |
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Three species of [[ |
Three species of [[Malagasy hippopotamus]] became extinct during the [[Holocene]] on [[Madagascar]], the last of them within the past 1,000 years. The Malagasy hippos were smaller than the modern hippo, a likely result of the process of [[insular dwarfism]].<ref name=Stuenes>{{Cite journal |doi=10.1080/02724634.1989.10011761 |author=Stuenes, S. |year=1989 |title=Taxonomy, habits and relationships of the sub-fossil Madagascan hippopotamuses ''Hippopotamus lemerlei'' and ''H. madagascariensis'' |journal=[[Journal of Vertebrate Paleontology]] |volume=9 |pages=241–268 |issue=3}}</ref> Fossil evidence indicates many Malagasy hippos were hunted by humans, a factor in their eventual extinction.<ref name=Stuenes/> Isolated individual Malagasy hippos may have survived in remote pockets; in 1976, villagers described a living animal called the ''kilopilopitsofy'', which may have been a Malagasy hippo.<ref name=Kidoky>{{Cite journal |title=The Kilopilopitsofy, Kidoky, and Bokyboky: Accounts of Strange Animals from Belo-sur-mer, Madagascar, and the Megafaunal "Extinction Window" |journal=[[American Anthropologist]] |year=1998 |author=Burney, D. A. |jstor=681820 |volume=100 |issue=4 |pages=957–966 |author2=Ramilisonina |doi=10.1525/aa.1998.100.4.957}}</ref> |
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''[[Hippopotamus gorgops]]'' from the Early Pleistocene to the early Middle Pleistocene of Africa and West Asia grew considerably larger than the living hippopotamus, with an estimated body mass of over {{Cvt|4000|kg}}.<ref name=VanDerMadeEtAl2017>van der Made J, Sahnouni M & Boulaghraief K. 2017. [https://www.mncn.csic.es/sites/default/files/inline-files/2017%20Van%20der%20Made%20ea%20-%20Hippopotamus%20-%20El%20Kherba%20-%20ProcIIMeetingAfrPrehist.pdf ''Hippopotamus gorgops'' from El Kherba (Algeria) and the context of its biogeography]. In Proceedings of the II Meeting of African Prehistory: Burgos 15–16 April 2015, Sahnouni M, Semaw S, Rios Garaizar J (eds). CENIEH: Burgos; 135–169.</ref><ref>Chaix L, Faure M, Guérin C, Honegger M. [https://kerma.ch/documents/Publications_PDF/Chaix_Faure_Guerin_Honegger_Kaddanarti_2000.pdf Kaddanarti, a Lower Pleistocene Assemblage from Northern Sudan]. In: Krzyżaniak L, Kroeper K, Kobusiewicz M, editors. Recent Research into the Stone Age of Northeastern Africa. Poznań: Poznań Archaeological Museum; 2000. p. 33–46.</ref><ref>{{Cite journal |last=Hutchinson |first=J. R. |date=2021 |title=The evolutionary biomechanics of locomotor function in giant land animals |journal=Journal of Experimental Biology |volume=224 |issue=11 |doi=10.1242/jeb.217463 |issn=0022-0949 |pmc=8214834 |pmid=34100541|bibcode=2021JExpB.224B7463H }}</ref> ''[[Hippopotamus antiquus]]'' ranged throughout Europe, extending as far north as Britain during the [[Early Pleistocene|Early]] and [[Middle Pleistocene]] epochs, before being replaced by the modern ''H. amphibius'' in Europe during the latter part of the Middle Pleistocene.<ref>{{Cite journal |last1=Adams|first1=N. F. |last2=Candy |first2=I. |last3=Schreve |first3=D. C.|title=An Early Pleistocene hippopotamus from Westbury Cave, Somerset, England: support for a previously unrecognized temperate interval in the British Quaternary record |journal=Journal of Quaternary Science |year=2022 |volume=37 |issue=1 |pages=28–41 |doi=10.1002/jqs.3375 |bibcode=2022JQS....37...28A |s2cid=244179438 |doi-access=free}}</ref> The Pleistocene also saw a number of dwarf species evolve on several Mediterranean islands, including [[Crete]] (''[[Hippopotamus creutzburgi]]''), [[Cyprus]] (the [[Cyprus dwarf hippopotamus]], ''Hippopotamus minor''), [[Malta]] (''[[Hippopotamus melitensis]]''), and [[Sicily]] (''[[Hippopotamus pentlandi]]''). Of these, the Cyprus dwarf hippo survived until the end of the Pleistocene or early Holocene. Evidence from the archaeological site [[Aetokremnos]] continues to cause debate on whether or not the species was driven to extinction or even encountered by humans.<ref>{{Cite journal |author=Simmons, A. |year=2000 |title=Faunal extinction in an island society: pygmy hippopotamus hunters of Cyprus |journal=[[Geoarchaeology]] |volume=15 |issue=4 |pages=379–381 |doi=10.1002/(SICI)1520-6548(200004)15:4<379::AID-GEA7>3.0.CO;2-E |bibcode=2000Gearc..15..379G}}</ref><ref name="Taxonomy of Pleistocene">{{cite journal |author=Petronio, C. |year=1995 |title=Note on the taxonomy of Pleistocene hippopotamuses |journal=Ibex |volume=3 |pages=53–55 |url=http://www.mountainecology.org/IBEX3/pdf/Art_Capitolo1/note_taxonomy_pleistocene.pdf |url-status=dead |archive-url=https://web.archive.org/web/20080912012236/http://www.mountainecology.org/IBEX3/pdf/Art_Capitolo1/note_taxonomy_pleistocene.pdf|archive-date=12 September 2008}}</ref> |
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==Characteristics== |
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[[File:Hippo skull dark.jpg|thumb| |
[[File:Hippo skull dark.jpg|thumb|Hippopotamus [[skull]], showing the large canines and incisors used for fighting]] |
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The hippopotamus is a [[megaherbivore]] and is exceeded in size among land animals only by [[elephant]]s and some [[rhinoceros]] species. The mean adult weight is around {{cvt|1480|kg}} for bulls and {{cvt|1365|kg}} for cows. Exceptionally large males have been recorded reaching {{cvt|2660|kg}}.<ref name="Owen-Smith_1992">{{cite book|last1=Owen-Smith|first1=R. N. |title=Megaherbivores: The Influence of Very Large Body Size on Ecology |date=1992 |pages=6–14|publisher=Cambridge University Press|isbn=9780521426374}}</ref> Male hippos appear to continue growing throughout their lives, while females reach maximum weight at around age 25.<ref>{{Cite journal |author1=Marshall, P. J. |author2=Sayer, J. A. |title=Population ecology and response to cropping of a hippo population in eastern Zambia |jstor=2401788 |volume=13 |issue=2 |year=1976 |journal=The Journal of Applied Ecology |pages=391–403 |doi=10.2307/2401788}}</ref> It is {{cvt|2.90|to|5.05|m}} long,<ref>{{cite web |url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Hippopotamus_amphibius.html#physical_description|title = Physical Description |access-date=2020-10-22}}</ref> including a tail of about {{cvt|35|to|56|cm}} in length and {{cvt|1.30|to|1.65|m}} tall at the shoulder,<ref>{{cite web|url=http://www.ultimateungulate.com/artiodactyla/hippopotamus_amphibius.html|title = ''Hippopotamus amphibius''|access-date=2020-10-22}}</ref><ref name="Hippopotamus">{{cite web|url=http://sarkive.com/mammals/hippopotamus-amphibius/|title=Hippopotamus|access-date=2020-10-22|archive-date=8 August 2020|archive-url=https://web.archive.org/web/20200808175302/http://sarkive.com/mammals/hippopotamus-amphibius/|url-status=dead}}</ref> with males and females ranging {{cvt|1.40|to|1.65|m}} and {{cvt|1.30|to|1.45|m}} tall at the shoulder respectively.<ref name="Hippopotamus"/> The species has a typical head–body length of {{cvt|3.3|–|3.45|m}} and an average standing height of {{cvt|1.4|m}} at the shoulder.<ref>{{Cite book|title=The Encyclopedia of Mammals|publisher=[[Oxford University Press]]|year=2001|isbn=978-0-7607-1969-5 |edition=2nd |page=491 |author=Eltringham, S. K.|contribution=Hippopotamuses|editor=MacDonald, D.}}</ref> |
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Hippos have barrel-shaped bodies with short tails and legs, and an hourglass-shaped skull with a long snout.<ref name=estes/><ref name="The Hippos"/>{{rp|3, 19}} Their skeletal structures are [[wikt:graviportal|graviportal]], adapted to carrying their enormous weight,<ref name="The Hippos"/>{{rp|8}} and their dense bones and low [[centre of gravity]] allows them to sink and move along the bottom of the water.<ref>{{cite journal|author1=Coughlin, B. L.|author2=Fish, F. E.|year=2009 |title=Hippopotamus underwater locomotion: Reduced-gravity movements for a massive mammal |journal=Journal of Mammalogy|volume=90|issue=3|pages=675–679|doi=10.1644/08-MAMM-A-279R.1 |s2cid=51686926 |doi-access=free}}</ref> Hippopotamuses have small legs (relative to other [[megafauna]]) because the water in which they live reduces the weight burden.<ref>{{cite book|title=Exploring Mammals|publisher=Marshall Cavendish Corporation |page=616 |url=https://books.google.com/books?id=k8o0HTqvwH0C&q=hippo+legs+water&pg=PA616 |isbn=9780761477280 |year=2008}}</ref> The toes are webbed and the [[pelvis]] rests at an angle of 45 degrees.<ref name="The Hippos"/>{{rp|3, 9}} Though chubby-looking, hippos have little fat.<ref name="The Hippos"/>{{rp|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 is submerged.<ref name=kingdon/>{{rp|259}} The nostrils and ears can close when underwater while [[nictitating membrane]]s cover the eyes.<ref name="The Hippos"/>{{rp|4, 116}} The [[vocal fold]]s of the hippo are more horizontally positioned, much like baleen whales. Underneath are throat tissues, where vibrations are transmitted to produce underwater calls.<ref>{{cite journal|last1=Reidenberg|first1=J. S. |year=2017|title=Terrestrial, semiaquatic, and fully aquatic mammal sound production mechanisms |journal=Acoustics Today |volume=13|issue=2|pages=35–43 |url=https://acousticstoday.org/wp-content/uploads/2017/05/Reidenberg.pdf}}</ref> |
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Hippopotamuses are among the largest living land mammals being only smaller than [[elephant]]s and some [[rhinoceros]]es. Head-and-body length is from {{convert|2.8|to|4|m|ftin|abbr=on}}, a tail of about {{convert|35|to|50|cm|in|abbr=on}} and shoulder height averages about {{convert|1.4|to|1.5|m|ftin|abbr=on}}.<ref>Carawadine, M. (1995) ''Natural History Museum: Animal Records''. Guinness Publishing (Sterling). ISBN 1402756232.</ref><ref>{{cite book|last1 = Stuart|first1 = Chris|last2 = Stuart|first2 =Mathilde|title = Field Guide to the Mammals of Southern Africa|publisher = [[Penguin Random House|Struik Publishers]]|year = 2011|pages = |isbn = 978-1770074040}}</ref> Mean adult weight is around {{convert|1500|kg|lbs|abbr=on}} and {{convert|1300|kg|lbs|abbr=on}} for males and females respectively,<ref name=Owen-Smith_1992>{{cite book|last1=Owen-Smith|first1=R. Norman|title=Megaherbivores: The Influence of Very Large Body Size on Ecology|date=1995|publisher=Cambridge University Press}}</ref><ref name=Piennar_etal_1966>{{cite journal|last1=Pienaar|first1=U. de V.|last2=Van Wyk|first2=P.|last3=Fairall|first3=N.|title=An experimental cropping scheme of Hippopotami in the Letaba river of the Kruger National Park|journal=Koedoe|date=1966|volume=9|issue=1|doi=10.4102/koedoe.v9i1.778|url=http://www.koedoe.co.za/index.php/koedoe/article/view/778}}</ref> very large males can reach {{convert|2000|kg|lbs|abbr=on}} and an exceptional male weighting almost {{convert|2700|kg|lbs|abbr=on}} has been reported.<ref name=Owen-Smith_1992/> Male hippos appear to continue growing throughout their lives while females reach maximum weight at around age 25.<ref>{{Cite journal|author = Marshall, P.J., Sayer, J.A.|title = Population ecology and response to cropping of a hippopotamus population in eastern Zambia|jstor=2401788|volume = 13|issue = 2|year = 1976|journal = The Journal of Applied Ecology|pages = 391–403|doi = 10.2307/2401788}}</ref> |
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[[File:Hipopótamos (Hippopotamus amphibius), parque nacional de Chobe, Botsuana, 2018-07-28, DD 79.jpg|thumb|Characteristic "yawn" of a hippo]] |
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Different from all other large land mammals, hippos are of semiaquatic habits, spending the day in lakes and rivers.<ref name="The Hippos"/>{{rp|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.<ref name=kingdon/>{{rp|259}} Their barrel-shaped bodies have [[wikt:graviportal|graviportal]] skeletal structures,<ref name="The Hippos"/>{{rp|8}} adapted to carrying their enormous weight, and their [[specific gravity]] allows them to sink and move along the bottom of a river.<ref>{{cite web|title=Hippopotamus ''Hippopotamus amphibius''|publisher=''National Geographic''|accessdate= 2013-07-10|url= http://animals.nationalgeographic.com/animals/mammals/hippopotamus/|archiveurl= https://web.archive.org/web/20141125041546/http://animals.nationalgeographic.com/animals/mammals/hippopotamus/|archivedate= 2014-11-25|deadurl= no}}</ref> Hippopotamuses have small legs (relative to other [[megafauna]]) because the water in which they live reduces the weight burden.<ref>{{cite book|title=Exploring Mammals|publisher=Marshall Cavendish Corporation|page=616|url=http://books.google.com/?id=k8o0HTqvwH0C&pg=PA616&dq=hippo+legs+water#v=onepage&q=hippo%20legs%20water&f=false|isbn=9780761477280|year=2008}}</ref> Though they are bulky animals, hippopotamuses can gallop at {{convert|30|km/h|abbr=on}} on land but normally trot. They are incapable of jumping but do climb up steep banks.<ref name=estes/> 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.<ref name="The Hippos"/>{{rp|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 [[erection|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.<ref name="The Hippos"/>{{rp|28–29}} |
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The hippo's jaw is powered by huge [[masseter muscle|masseter]] and [[digastric muscle]]s which give them large, droopy cheeks.<ref name=kingdon>{{Cite book|author=Kingdon, J. |year=1988|title=East African Mammals: An Atlas of Evolution in Africa |volume=((3, Part B: Large Mammals)) |pages=256–277|publisher=University Of Chicago Press |isbn=978-0-226-43722-4}}</ref>{{rp|259}} The jaw hinge allows the animal to open its mouth at almost 180°.<ref name="The Hippos"/>{{rp|17}} A folded [[orbicularis oris muscle]] allows the hippo to attain an extreme gape without tearing any tissue.<ref>{{cite journal|author=Herring, S. W.|year=1975|title=Adaptations for gape in the hippopotamus and its relatives |journal=Forma et Functio |volume=8 |pages=85–100}}</ref> On the lower jaw, the [[incisor]]s and [[Canine tooth|canines]] grow continuously, the former reaching {{cvt|40|cm}}, while the latter can grow to up to {{cvt|50|cm}}. The lower canines are sharpened through contact with the smaller upper canines.<ref name=estes>{{Cite book |title=The Behavior Guide to African Mammals: including hoofed mammals, carnivores, primates |author=Estes, R. |publisher=University of California Press |pages=222–226 |year=1992 |isbn=978-0-520-08085-0 |url=https://archive.org/details/isbn_0520080858/page/222}}</ref> The canines and incisors are used mainly for combat instead of feeding. Hippos rely on their flattened, horny lips to grasp and pull grasses which are then ground by the [[Molar (tooth)|molars]].<ref name=kingdon/>{{rp|259, 263}} The hippo is considered to be a [[pseudoruminant]]; it has a complex three-chambered stomach, but does not "chew [[cud]]".<ref name="The Hippos"/>{{rp|22}} |
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[[File:Hippo mouth opening.jpg|thumb|right|Characteristic "yawn" of a hippo.]] |
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[[File:Hippopotamus in San Diego Zoo.jpg|thumb| |
[[File:Hippopotamus in San Diego Zoo.jpg|thumb|Completely submerged hippo ([[San Diego Zoo]])]] |
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The hippo's jaw is powered by a large [[Masseter muscle|masseter]] and a well-developed [[Digastric muscle|digastric]]; the latter loops up behind the former to the [[hyoid]].<ref name=kingdon>{{Cite book|author=Kingdon, J. |year=1988|title=East African Mammals: An Atlas of Evolution in Africa, Volume 3, Part B: Large Mammals|pages=256–77|publisher=University Of Chicago Press |isbn=0-226-43722-1}}</ref>{{rp|259}} The jaw hinge is located far back enough to allow the animal to open its mouth at almost 180°.<ref name="The Hippos"/>{{rp|17}} 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 {{convert|8100|newtons|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.<ref>[[Brady Barr|Barr, Brady]]. "Undercover Hippo," ''Dangerous Encounters'', [[National Geographic Channel]], January 20, 2008.</ref> Hippopotamus teeth sharpen themselves as they grind together. The lower [[Canine tooth|canines]] and lower [[incisor]]s are enlarged, especially in males, and grow continuously. The incisors can reach {{convert|40|cm|ft|abbr=on}}, while the canines reach up to {{convert|50|cm|ft|abbr=on}}.<ref name=estes>{{Cite book|title=The Behavior Guide to African Mammals: including hoofed mammals, carnivores, primates|author=Estes, R.|publisher=University of California Press|pages=222–26|year=1992|isbn=0-520-08085-8}}</ref> 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 [[Molar (tooth)|molars]].<ref name=kingdon/>{{rp|259, 263}} The hippo is considered to be a [[pseudoruminant]], it has a complex three- or four-chambered stomach but does not "chew [[cud]]".<ref name="The Hippos"/>{{rp|22}} |
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Hippo skin is {{cvt|6|cm}} thick across much of its body with little hair.<ref name=estes/><ref name=kingdon/>{{rp|260}} The animal is mostly purplish-grey or blue-black, but brownish-pink on the underside and around the eyes and ears.<ref name=kingdon/>{{rp|260}} Their skin secretes a natural, red-coloured [[sunscreen]] substance that 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 highly acidic pigments have been identified in the secretions; one red [[hipposudoric acid]] and one orange norhipposudoric acid, which inhibit the growth of disease-causing bacteria and their light-absorption profile peaks in the [[ultraviolet]] range, creating a sunscreen effect.<ref name="Distillations">{{cite journal |last1=Kean |first1=Sam |title=Sweating blood |url=https://www.sciencehistory.org/distillations/magazine/sweating-blood |journal=Distillations |date=2018|volume=4 |issue=2 |page=5 |access-date=20 August 2018}}</ref><ref name="Saikawa"/> Regardless of diet, all hippos secrete these pigments so food does not appear to be their source; rather, they may be synthesised from [[Protein precursor|precursors]] such as the [[amino acid]] [[tyrosine]].<ref name="Saikawa">{{cite journal|author=Saikawa, Y.|author2= Hashimoto, K.|author3= Nakata, M.|author4= Yoshihara, M.|author5= Nagai, K.|author6= Ida, M.|author7= Komiya, T. |title=Pigment chemistry: the red sweat of the hippopotamus|journal=[[Nature (journal)|Nature]] |volume=429 |issue=6990 |year=2004 |page=363 |pmid=15164051 |doi=10.1038/429363a |bibcode=2004Natur.429..363S |s2cid=4404922 |doi-access=free}}</ref> This natural sunscreen cannot prevent the animal's skin from cracking if it stays out of water too long.<ref>{{cite book|author=Jablonski, N. G.|year=2013 |title=Skin: A Natural History|publisher=University of California Press |page=34 |isbn=978-0-520-24281-4 |url=https://archive.org/details/skinnaturalhisto00jabl/page/34}}</ref> |
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The testes of the males do not fully descend and a scrotum is not present. In addition, the penis retracts into the body when not [[erection|erect]]. The genitals of the female hippos are unusual in that the vagina is ridged and the [[vulval vestibule]] has two large, protruding [[diverticula]]. Both of these have an unknown function.<ref name="The Hippos"/>{{rp|28–29}} |
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A hippo's lifespan is typically 40–50 years.<ref name=kingdon/>{{rp|277}} Donna the Hippo was the oldest living hippo in captivity. She lived at the [[Mesker Park Zoo]] in [[Evansville, Indiana]] in the US<ref>{{cite web|url = http://www.evansvillecvb.org/media/index.tpl?ID=39&Display=Detail|title = Oldest Hippo Turns 55!|work = [[Mesker Park Zoo]]|date = 2006-06-12|accessdate = 2007-06-21 |archiveurl = http://web.archive.org/web/20070927120505/http://www.evansvillecvb.org/media/index.tpl?ID=39&Display=Detail |archivedate = 2007-09-27}}</ref><ref>{{Cite news|url = http://www.courierpress.com/news/2007/jul/12/celebrate-with-donna|title = Celebrate with Donna|work = [[Evansville Courier & Press]]|accessdate = 2007-07-15|date = 2007-07-12}}</ref> until her death in 2012 at the age of 61.<ref>{{cite web|author=Fears, Danika|date=2012-08-03|title=Goodbye, Donna: World's oldest hippo in captivity dies at 61|publisher=Today.com|accessdate=2013-09-12|url=http://www.today.com/pets/goodbye-donna-worlds-oldest-hippo-captivity-dies-61-922081}}</ref> The oldest hippo ever recorded was called Tanga; she lived in [[Munich]], [[Germany]], and died in 1995 at the age of 61.<ref name=AFP>{{Cite news|title = Old mother hippo dies|work = [[Agence France Press]]|date = July 12, 1995}}</ref> |
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A hippo's lifespan is typically 40 to 50 years.<ref name=kingdon/>{{rp|277}} Donna the Hippo was one of the oldest living hippos in captivity. She lived at the [[Mesker Park Zoo]] in [[Evansville, Indiana]], in the US<ref>{{cite web |title=Oldest Hippo Turns 55! |url=http://www.evansvillecvb.org/media/index.tpl?ID=39&Display=Detail |publisher=[[Mesker Park Zoo]] |date=2006 |access-date=21 June 2007 |archive-url=https://web.archive.org/web/20070927120505/http://www.evansvillecvb.org/media/index.tpl?ID=39&Display=Detail |archive-date=27 September 2007}}</ref><ref>{{Cite news |url=http://www.courierpress.com/news/2007/jul/12/celebrate-with-donna |title=Celebrate with Donna |work=[[Evansville Courier & Press]] |access-date=15 July 2007 |date=2007 |archive-date=16 January 2014 |archive-url=https://web.archive.org/web/20140116201843/http://www.courierpress.com/news/2007/jul/12/celebrate-with-donna/ |url-status=dead}}</ref> until her death in 2012 at the age of 61.<ref>{{cite web |author=Fears, D. |date=2012 |title=Goodbye, Donna: World's oldest hippo in captivity dies at 61 |publisher=Today.com|access-date=12 September 2013|url=http://www.today.com/pets/goodbye-donna-worlds-oldest-hippo-captivity-dies-61-922081 |url-status=dead |archive-url=https://web.archive.org/web/20130604043252/http://www.today.com/pets/goodbye-donna-worlds-oldest-hippo-captivity-dies-61-922081 |archive-date=4 June 2013}}</ref> The oldest hippo ever recorded was called Bertha; she had lived in the [[Manila Zoo]] in the Philippines since it first opened in 1959. When she died in 2017, her age was estimated to be 65.<ref>{{Cite news |title=Bertha, the world's 'oldest' hippo, dies at 65 |url=https://www.bbc.com/news/world-asia-40558057 |work=BBC News |date=2017 |access-date=29 November 2020}}</ref> The oldest living hippopotamus in captivity is Lu the Hippo, from the Ellie Schiller [[Homosassa Springs Wildlife State Park]]. As of 2024, he is 64 years old.<ref>{{Cite web |last=Lentz-Janney |first=M. |date=2021 |title=Our Favorite Homosassa Springs Resident: Lu the Hippo |url=https://authenticflorida.com/lu-the-hippo/ |access-date=2024-01-31 |website=Authentic Florida}}</ref> |
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==Distribution== |
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''Hippopotamus amphibius'' was widespread in [[North Africa]] and [[Europe]] during the [[Eemian]]<ref>{{cite journal |last=van Kolfschoten |first=Th. |year=2000 |title=The Eemian mammal fauna of central Europe |journal=Netherlands Journal of Geosciences |volume=79 |issue=2/3 |pages=269–281 |url=http://www.njgonline.nl/publish/articles/000099/article.pdf}}</ref> 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.<ref>{{cite journal|last=Horwitz|first=Liora Kolska|author2=Eitan Tchernov|title=Cultural and Environmental Implications of Hippopotamus Bone Remains in Archaeological Contexts in the Levant|journal=Bulletin of the American Schools of Oriental Research|year=1990|volume=280|pages=67–76|doi=10.2307/1357310}}</ref><ref>{{cite journal|last=Haas|first=Georg|title=On the Occurrence of Hippopotamus in the Iron Age of the Coastal Area of Israel|journal=Bulletin of the American Schools of Oriental Research|year=1953|volume=132|pages=30–34|doi=10.2307/1355798}}</ref> The species was common in [[Egypt]]'s [[Nile]] region during [[Classical antiquity|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 [[Sais, Egypt|Saite nome]];<ref name="Pliny">{{Cite book|title = [[Naturalis Historia]]|author = [[Pliny the Elder]]|chapter = Chapter 15, Book VIII|language = [http://penelope.uchicago.edu/Thayer/L/Roman/Texts/Pliny_the_Elder/8*.html Latin original] or [http://penelope.uchicago.edu/holland/pliny8.html English translation]|isbn = 3-519-01652-4}}</ref> 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 [[Gambia]], and south to [[South Africa]]. They inhabit both savanna and forest areas.<ref name="Redlist"/> |
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== |
==Distribution and status== |
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[[File:Hippovictora.png|thumb |
[[File:Hippovictora.png|thumb|right|Ugandan tribespeople with hippo slain for food (early 20th century)]] |
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[[File:Incised hippopotamus ivory tusk, upper canine. Four holes around top. From Naqada Tomb 1419, Egypt. Naqada period. The Petrie Museum of Egyptian Archaeology, London.jpg|thumb|Incised hippopotamus ivory tusk (upper canine) with four holes around top (Naqada Tomb 1419, Egypt; Naqada period)]] |
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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.<ref name="Okello"/> 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 [[World Conservation Union]] (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.<ref name="Redlist">{{IUCN2008|assessors=Lewison, R. & Oliver, W. (IUCN SSC Hippo Specialist Subgroup)|year=2008|id=10103|title=Hippopotamus amphibius|archiveurl= https://web.archive.org/web/20141103123159/http://www.iucnredlist.org/details/10103/0|archivedate= 2014-11-03|deadurl= no|downloaded= 2009-04-05}} Database entry includes a range map and justification for why this species is vulnerable.</ref> |
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''Hippopotamus amphibius'' arrived in Europe around 560–460,000 years ago, during the Middle Pleistocene.<ref>{{Cite journal |last1=Mecozzi |first1=Beniamino |last2=Iannucci |first2=Alessio |last3=Mancini |first3=Marco |last4=Tentori |first4=Daniel |last5=Cavasinni |first5=Chiara |last6=Conti |first6=Jacopo |last7=Messina |first7=Mattia Yuri |last8=Sarra |first8=Alex |last9=Sardella |first9=Raffaele |date=2023-11-22 |editor-last=Carnevale |editor-first=Giorgio |title=Reinforcing the idea of an early dispersal of Hippopotamus amphibius in Europe: Restoration and multidisciplinary study of the skull from the Middle Pleistocene of Cava Montanari (Rome, central Italy) |journal=PLOS ONE |language=en |volume=18 |issue=11 |pages=e0293405 |doi=10.1371/journal.pone.0293405 |issn=1932-6203 |pmc=10664965 |pmid=37992018 |doi-access=free|bibcode=2023PLoSO..1893405M }}</ref> The distribution of ''Hippopotamus amphibius'' in Europe during the Pleistocene was largely confined to Southern Europe, including the Iberian Peninsula,<ref name=":0">{{Cite journal |last1=Fidalgo |first1=Darío |last2=Madurell-Malapeira |first2=Joan |last3=Martino |first3=Roberta |last4=Pandolfi |first4=Luca |last5=Rosas |first5=Antonio |date=2024-01-10 |title=An Updated Review of The Quaternary Hippopotamus Fossil Records from the Iberian Peninsula |journal=Quaternary |language=en |volume=7 |issue=1 |pages=4 |doi=10.3390/quat7010004 |doi-access=free |issn=2571-550X|hdl=10261/345195 |hdl-access=free }}</ref> Italy,<ref>{{Cite journal |last1=Briatico |first1=Giuseppe |last2=Gioia |first2=Patrizia |last3=Bocherens |first3=Hervé |date=10 December 2023 |title=Diet and habitat of the late Middle Pleistocene mammals from the Casal de' Pazzi site (Rome, Italy) using stable carbon and oxygen isotope ratios |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618223003336 |journal=[[Quaternary International]] |language=en |volume=676 |pages=53–62 |doi=10.1016/j.quaint.2023.11.002 |bibcode=2023QuInt.676...53B |access-date=27 April 2024 |via=Elsevier Science Direct}}</ref><ref name=":02">{{Cite journal |last1=Martino |first1=R. |last2=Pandolfi |first2=L. |date=2022-07-03 |title=The Quaternary Hippopotamus records from Italy |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2021.1965138 |journal=Historical Biology |language=en |volume=34 |issue=7 |pages=1146–1156 |doi=10.1080/08912963.2021.1965138 |bibcode=2022HBio...34.1146M |issn=0891-2963 |s2cid=239713930}}</ref> and Greece,<ref name=":1">{{Citation |last=Athanassiou |first=Athanassios |title=The Fossil Record of Continental Hippopotamids (Mammalia: Artiodactyla: Hippopotamidae) in Greece |date=2022 |work=Fossil Vertebrates of Greece Vol. 2 |pages=281–299 |editor-last=Vlachos |editor-first=Evangelos |url=https://link.springer.com/10.1007/978-3-030-68442-6_9 |access-date=2024-01-26 |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-030-68442-6_9 |isbn=978-3-030-68441-9|s2cid=239839913 }}</ref> but extended into northwestern Europe, including Great Britain (as far north as [[North Yorkshire]]), the Netherlands, and western Germany during [[interglacial]] periods, such as the [[Last Interglacial]] (130–115,000 years ago).<ref>{{Cite journal |last=Pushkina |first=Diana |date=July 2007 |title=The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian Palaeoloxodon antiquus assemblage |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1365-2907.2007.00109.x |journal=Mammal Review |language=en |volume=37 |issue=3 |pages=224–245 |doi=10.1111/j.1365-2907.2007.00109.x |issn=0305-1838 }}</ref><ref>{{Cite journal |last=van Kolfschoten |first=Th. |date=August 2000 |title=The Eemian mammal fauna of central Europe |url=https://www.cambridge.org/core/product/identifier/S0016774600021752/type/journal_article |journal=Netherlands Journal of Geosciences |language=en |volume=79 |issue=2–3 |pages=269–281 |doi=10.1017/S0016774600021752 |bibcode=2000NJGeo..79..269V |issn=0016-7746}}</ref><ref>{{Cite journal |last=Schreve |first=Danielle C. |date=January 2009 |title=A new record of Pleistocene hippopotamus from River Severn terrace deposits, Gloucester, UK—palaeoenvironmental setting and stratigraphical significance |url=https://linkinghub.elsevier.com/retrieve/pii/S0016787809000054 |journal=Proceedings of the Geologists' Association |language=en |volume=120 |issue=1 |pages=58–64 |bibcode=2009PrGA..120...58S |doi=10.1016/j.pgeola.2009.03.003}}</ref> The youngest records of the species in Europe are from the Late Pleistocene of Greece, dating to around 40–30,000 years ago.<ref name=":1" /><ref name=":0" /> |
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Archaeological evidence exists of its presence in the [[Levant]], dating to less than 3,000 years ago.<ref>{{cite journal|last=Horwitz|first=Liora Kolska|author2= Tchernov, Eitan|title=Cultural and Environmental Implications of Hippopotamus Bone Remains in Archaeological Contexts in the Levant|journal=Bulletin of the American Schools of Oriental Research|year=1990|volume=280|issue=280|pages=67–76|doi=10.2307/1357310|jstor=1357310|s2cid=163871070}}</ref><ref>{{cite journal|last=Haas|first=Georg|title=On the Occurrence of Hippopotamus in the Iron Age of the Coastal Area of Israel|journal=Bulletin of the American Schools of Oriental Research|year=1953|volume=132|issue=132|pages=30–34|doi=10.2307/1355798|jstor=1355798|s2cid=163758714}}</ref> The species was common in [[Egypt]]'s [[Nile]] region during [[Classical antiquity|antiquity]], but it has since been driven out. According to [[Pliny the Elder]], in his time, the best location in Egypt for capturing this animal was in the [[Sais, Egypt|Saite nome]];<ref name="Pliny">{{Cite book|title = Naturalis Historia|author = Pliny the Elder|author-link = Pliny the Elder|chapter = Chapter 15, Book VIII|language=en |isbn = 978-3-519-01652-6|title-link = Naturalis Historia|date = 1 January 1987| publisher=Vieweg+Teubner Verlag }} ([http://penelope.uchicago.edu/holland/pliny8.html English translation]; [https://penelope.uchicago.edu/Thayer/L/Roman/Texts/Pliny_the_Elder/8*.html Latin original])</ref> the animal could still be found along the [[Damietta]] branch of the Nile after the Arab Conquest in 639. Reports of the slaughter of the last hippo in [[Natal Province]] were made at the end of the 19th century.<ref>{{Cite news|url=http://nla.gov.au/nla.news-article14137838|title=The Hippopotamus Going - Extinction of the huge species at hand|date=7 October 1898|work=The Sydney Morning Herald|access-date=14 July 2017}}</ref> 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]].<ref name="iucn" /> |
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The hippo population declined most dramatically in the Democratic Republic of the Congo.<ref name="HippoHaven">{{cite news |url = http://www.smithsonianmagazine.com/issues/2006/january/hippos.php |title = Hippo Haven |work = [[Smithsonian Magazine]] |date = 2006-01-01 |accessdate = 2007-01-23}}</ref> The population in [[Virunga National Park]] had dropped to 800 or 900 from around 29,000 in the mid-1970s.<ref name="BBC"/> The decline is attributed to the disruptions caused by the [[Second Congo War]].<ref name="BBC">{{Cite news| title = DR Congo's hippos face extinction.|url = http://news.bbc.co.uk/2/hi/africa/4240420.stm|date = 2005-09-13|accessdate = 2005-11-14|work = [[BBC]]}}</ref> The poachers are believed to be [[Mai-Mai]] rebels, poorly paid Congolese soldiers, and local militia groups.<ref name="BBC"/><ref name=natgeo>{{cite web|author=Owen, James|date=2006-10-24|title=Hippos Butchered by the Hundreds in Congo Wildlife Park|publisher=National Geographic News|accessdate=2013-09-11|url=http://news.nationalgeographic.com/news/2006/10/061024-hippo-congo.html}}</ref> Reasons for poaching include the belief that hippos are harmful to society, as well as financial gain.<ref name="Star"/> The sale of hippo meat is illegal, but black-market sales are difficult for Virunga National Park officers to track.<ref name=natgeo/><ref name="Star">{{Cite news |last=Sundaram |first=Anjan |title = Congo's Hippos Fast Disappearing |newspaper=[[Toronto Star]] |date=2005-09-12}}</ref> Hippo meat is considered a delicacy in some areas of central Africa and the teeth have become a valued substitute for elephant ivory.<ref name="newscientist2003">{{cite journal | first= Fred | last = Pearce | title = Poaching causes hippo population crash | journal = New Scientist | year = 2003 | url = http://www.newscientist.com/article/dn4109-poaching-causes-hippo-population-crash.html#.U1vYUPldWz4 | accessdate = April 26, 2014}}</ref> |
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Genetic evidence suggests common hippos in Africa experienced a marked population expansion during or after the [[Pleistocene]], 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.<ref name="Okello"/> Hippos are also subject to unregulated hunting and [[poaching]]. The species is included in Appendix II of the [[Convention on International Trade in Endangered Species]] (CITES) meaning international export/import (including in parts and derivatives) requires CITES documentation to be obtained and presented to border authorities.<ref name=iucn/><ref name="Appendices {{!}} CITES">{{Cite web|title=Appendices {{!}} CITES|url=https://cites.org/eng/app/index.php|access-date=2022-11-13|website=cites.org}}</ref> |
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===Invasive potential=== |
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In the late 1980s, [[Pablo Escobar]] kept four hippos in a private menagerie at his residence in [[Hacienda Nápoles]], {{convert|100|km}} 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]].<ref>{{cite news |url=http://www.latimes.com/news/nationworld/world/la-fg-hippos20dec20,0,5373140.story |title=A hippo critical situation |accessdate=2008-03-27|first= Chris|last= Kraul|date= 2006-12-20 |work=[[Los Angeles Times]]|archiveurl= http://www.webcitation.org/6WsgZRAjS|archivedate= 2015-03-08|deadurl= no}}</ref><ref name=baron/> 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.<ref name=baron>{{cite news|url=http://news.bbc.co.uk/2/hi/americas/8145676.stm|title = Colombia kills drug baron hippo|publisher = BBC News|accessdate=2009-07-11|date=2009-07-11|archiveurl= https://web.archive.org/web/20150105105021/http://news.bbc.co.uk/2/hi/americas/8145676.stm|archivedate= 2015-01-05|deadurl= no}}</ref><ref>{{cite web|url= http://elarcadepitusa.blogspot.com/2009/07/crece-controversia-en-colombia-por-la.html|title = Crece controversia en el país por decisión de cazar a hipopótamos de Pablo Escobar |publisher = ''[[El Tiempo (Colombia)|El Tiempo]]''|archiveurl= https://web.archive.org/web/20150308182559/http://elarcadepitusa.blogspot.com/2009/07/crece-controversia-en-colombia-por-la.html|archivedate= 2015-03-08|deadurl= yes|accessdate=2009-07-11}} [https://translate.google.com/translate?sl=auto&tl=en&js=y&prev=_t&hl=en&ie=UTF-8&u=http%3A%2F%2Felarcadepitusa.blogspot.com%2F2009%2F07%2Fcrece-controversia-en-colombia-por-la.html&edit-text= English translation] at [[Google Translate]]</ref> As of early 2014, 40 hippos have been reported to exist in Puerto Triunfo, [[Antioquia]] from the original four belonging to Escobar.<ref>{{cite news|url=http://www.elespectador.com/opinion/editorial/hipopotamos-bravos-articulo-487824|title=Hipopótamos bravos|date=2014-06-24|publisher= ''[[El Espectador]]''|archiveurl= https://web.archive.org/web/20140509133359/http://www.elespectador.com/opinion/editorial/hipopotamos-bravos-articulo-487824|archivedate= 2014-05-09|deadurl= no|accessdate=2014-06-28}} [https://translate.google.com/translate?sl=auto&tl=en&js=y&prev=_t&hl=en&ie=UTF-8&u=http%3A%2F%2Fwww.elespectador.com%2Fopinion%2Feditorial%2Fhipopotamos-bravos-articulo-487824&edit-text= English translation] at Google Translate</ref> The [[National Geographic Channel]] produced a documentary about them titled ''Cocaine Hippos''.<ref>{{cite web |url= http://natgeotv.com/asia/cocaine-hippos/about|title= The Invaders: Cocaine Hippos|publisher= [[National Geographic Channel]]|archiveurl= https://web.archive.org/web/20130626213741/http://natgeotv.com/asia/cocaine-hippos/about|archivedate= 2013-06-26|deadurl= no}}</ref> |
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As of 2017, the [[IUCN Red List]] drawn up by the [[International Union for Conservation of Nature]] (IUCN) lists the species as [[vulnerable species|vulnerable]], with a stable population estimated between 115,000 and 130,000 animals.<ref name=iucn/> The hippo population has declined most dramatically in the Democratic Republic of the Congo.<ref name="HippoHaven">{{cite news |url = http://www.smithsonianmagazine.com/issues/2006/january/hippos.php |title = Hippo Haven |work = [[Smithsonian Magazine]] |date = 1 January 2006 |access-date = 23 January 2007}}</ref> By 2005, the population in [[Virunga National Park]] had dropped to 800 or 900 from around 29,000 in the mid-1970s.<ref name="BBC"/> This decline is attributed to the disruptions caused by the [[Second Congo War]].<ref name="BBC">{{Cite news| title = DR Congo's hippos face extinction.|url = http://news.bbc.co.uk/2/hi/africa/4240420.stm|date = 13 September 2005|access-date = 14 November 2005|work = [[BBC]]}}</ref> The poachers are believed to be [[Mai-Mai]] rebels, underpaid Congolese soldiers, and local militia groups.<ref name="BBC"/><ref name=natgeo>{{cite web|author=Owen, James|date=24 October 2006|title=Hippos Butchered by the Hundreds in Congo Wildlife Park|publisher=National Geographic News|access-date=11 September 2013|url=http://news.nationalgeographic.com/news/2006/10/061024-hippo-congo.html|url-status=dead| archive-url=https://web.archive.org/web/20140220005151/https://news.nationalgeographic.com/news/2006/10/061024-hippo-congo.html|archive-date=20 February 2014}}</ref> Reasons for poaching include the belief hippos are harmful to society, as well as financial gain.<ref name="Star"/> As of 2016, the Virunga hippo population appears to have increased again, possibly due to better protection from park rangers, who have worked with local fishermen.<ref>{{cite web|author1=Delaney, J. |author2=Sautner, S. |title=After a Long Demise Due to Poaching, Virunga's Hippos Climbing Back|publisher=Wildlife Conservation Society|access-date=25 November 2016|date=3 November 2016|url=https://newsroom.wcs.org/News-Releases/articleType/ArticleView/articleId/9373/After-a-Long-Demise-Due-to-Poaching,-Virunga%E2%80%99s-Hippos-Climbing-Back.aspx}}</ref> The sale of hippo meat is illegal, but black-market sales are difficult for Virunga National Park officers to track.<ref name=natgeo/><ref name="Star">{{Cite news |last=Sundaram |first=Anjan |title = Congo's Hippos Fast Disappearing |newspaper=[[Toronto Star]] |date=12 September 2005}}</ref> Hippo meat is highly valued in some areas of central Africa and the teeth may be used as a replacement for elephant ivory.<ref name="newscientist2003">{{cite journal | first= Fred | last = Pearce | title = Poaching causes hippo population crash | journal = New Scientist | year = 2003 | url = https://www.newscientist.com/article/dn4109-poaching-causes-hippo-population-crash.html#.U1vYUPldWz4 | access-date = 26 April 2014}}</ref> |
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==Behavior== |
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[[File:Hippo at dawn.jpg|thumb|left|A hippo out of water just after sunrise]] |
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A population of hippos [[Hippopotamuses in Colombia|exists in Colombia]], descended from captive individuals that escaped from [[Pablo Escobar]]'s estate after his death in 1993. Their numbers grew to 100 by the 2020s and ecologists believe the population should be eradicated, as they are breeding rapidly and are an increasing menace to humans and the environment. Attempts to control them include sterilisation and [[culling]].<ref>{{cite news |last1=Charles |first1=Matthew |title=Colombia's rapidly breeding 'cocaine hippos' must be stopped, scientists say |url=https://www.yahoo.com/news/colombias-rapidly-breeding-cocaine-hippos-153839192.html |access-date=January 18, 2021 |work=news.yahoo.com |publisher=The Telegraph |date=January 17, 2021}}</ref> |
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With the exception of eating, most of hippopotamuses' lives – from childbirth, fighting with other hippos, to reproduction – occurs in the water. Hippos leave the water at dusk and travel inland, sometimes up to {{convert|10|km|0|abbr=on}},<ref name=estes/> to graze on short grasses, their main source of food. They spend four to five hours grazing and can consume {{convert|68|kg|abbr=on}} of grass each night.<ref>{{cite web|url = http://www.krugerpark.co.za/africa-hippopotamus.html|title = Hippopotamus|accessdate = 2007-06-18|work = [[Kruger National Park]]}}</ref> 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.<ref name="Grey"/> [[Hippo]]s are born with sterile intestines, and require bacteria obtained from their mothers' feces to digest vegetation.<ref>{{cite web|url=http://www.bbc.co.uk/nature/adaptations/Coprophagia |title=BBC Nature — Dung eater videos, news and facts |publisher=Bbc.co.uk |accessdate=2011-11-27}}</ref><ref>https://www.princeton.edu/~achaney/tmve/wiki100k/docs/Coprophagia.html</ref> Hippos have (rarely) been filmed eating [[carrion]], usually close to the water. There are other reports of meat-eating, and even [[cannibalism]] and [[predation]].<ref>{{Cite journal|title = Reports of carnivory by the common hippo Hippopotamus Amphibius|author = Dudley, J.P. |journal = South African Journal of Wildlife Research|volume = 28|issue = 2|pages = 58–59 |url=http://search.sabinet.co.za/WebZ/images/ejour/wild/wild_v28_n2_a4.pdf?sessionid=0&format=F}}</ref> The stomach anatomy of a hippo is not suited to carnivory, and meat-eating is likely caused by aberrant behavior or nutritional stress.<ref name="The Hippos"/>{{rp|84}} |
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==Behaviour and ecology== |
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[[File:Hippo.ogv|thumb|right|Video of hippos in the wild]] |
[[File:Hippo.ogv|thumb|right|Video of hippos in the wild]] |
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Hippos are [[semiaquatic]] and require enough water to immerse in, while being close to grass.<ref name=estes/> They mostly live in freshwater habitat, but can be found in [[estuaries]].<ref name=iucn/> They prefer relatively still waters with gently sloping shores, though male hippos may also be found in very small numbers in more [[Rapids|rapid]] waters with rocky slopes.<ref name=kingdon/>{{rp|264}} Like most herbivores, hippos will consume a variety of plants if presented with them in captivity, but their diet in nature consists almost entirely of grass, with only minimal consumption of aquatic plants.<ref name="Grey"/> Hippos spend most of the day in water to stay cool and hydrated. Just before night begins, they leave the water to forage on land. A hippo will travel {{cvt|3|–|5|km}} per night, eating around {{cvt|40|kg}} of grass. By dawn, they are back in the water.<ref name=estes/> |
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Despite being semiaquatic, an adult hippo is not a particularly good swimmer, nor can it float. It rarely enters deep water; when it does, the animal moves by bouncing off the bottom. An adult hippo surfaces every four to six minutes, while young need to breathe every two to three minutes.<ref name="The Hippos"/>{{rp|3–4}} Hippos move on land by [[trot]]ting, and limb movements do not change between speeds. They can reach an airborne stage (a stage when all limb are off the ground) when they move fast enough. Hippos are reported to reach {{cvt|30|km/h}} but this has not been confirmed.<ref>{{cite journal|last1=Hutchinson|first1=J. R.|last2=Pringle|first2=E. V.|year=2024|title=Footfall patterns and stride parameters of Common hippopotamus (''Hippopotamus amphibius'') on land|journal=PeerJ|volume=12|page=e17675|doi=10.7717/peerj.17675|doi-access=free |pmid=38974416 |pmc=11227274}}</ref> They are incapable of jumping but can walk up steep banks.<ref name=estes/> The hippopotamus sleeps with both hemispheres of the brain resting, as in all land mammals, and usually sleeps on land or in water with the nostrils exposed. Despite this, it may be capable of sleeping while submerged, intermittently surfacing to breathe without waking. They appear to transition between different phases of sleep more quickly than other mammals.<ref>{{cite journal|author1=Dell, Leigh-Anne|author2=Patzke, Nina|author3=Spocter, Muhammad A.|author4=Bertelsen, Mads F.|author5=Siegel, Jerome M.|author6=Manger, Paul R.|year=2016|title=Organization of the sleep-related neural systems in the brain of the river hippopotamus (''Hippopotamus amphibius''): A most unusual cetartiodactyl species|journal=Journal of Comparative Neurology|volume=524|issue=10|pages=2036–2058|doi=10.1002/cne.23930|pmid=26588600 |pmc=8716328 }}</ref> |
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Hippo defecation creates [[allochthonous]] deposits of organic matter along the river beds. These deposits have an unclear ecological function.<ref name="Grey">{{Cite journal|title = Using Stable Isotope Analyses To Identify Allochthonous Inputs to Lake Naivasha Mediated Via the Hippopotamus Gut| author = Grey, J., Harper, D.M.|journal = Isotopes in Environmental Health Studies|volume = 38|issue = 4|year = 2002|pages = 245–250|doi = 10.1080/10256010208033269| pmid = 12725427| last2 = Harper}}</ref> 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.<ref name="Botswana">{{Cite journal|journal = African Journal of Ecology|volume = 36|issue = 1|pages = 44–56|year = 1998|author = McCarthy, T.S., Ellery, W. N., Bloem, A|title = Some observations on the geomorphological impact of hippopotamus (Hippopotamus amphibius L.) in the Okavango Delta, Botswana|doi = 10.1046/j.1365-2028.1998.89-89089.x}}</ref> |
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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, keeping the land clear of vegetation and depressing the ground. Over prolonged periods, hippos can divert the paths of swamps and channels.<ref name="Botswana">{{Cite journal|journal = African Journal of Ecology|volume = 36|issue = 1|pages = 44–56|year = 1998|author1=McCarthy, T. S. |author2=Ellery, W. N. |author3=Bloem, A |title = Some observations on the geomorphological impact of hippopotamus (''Hippopotamus amphibius'' L.) in the Okavango Delta, Botswana|doi = 10.1046/j.1365-2028.1998.89-89089.x| bibcode=1998AfJEc..36...44M }}</ref> By defecating in the water, the animals also appear to pass on microbes from their gut, affecting the [[biogeochemical cycle]].<ref>{{cite journal|author1=Dutton, C. L.|author2=Subalusky, A. L.|author3=Sanchez, A.|author4=Estrela, S.|author5=Lu, N.|author6=Hamilton, S. K.|author7=Njoroge, L.|author8=Rosi, E. J.|author9=Post, D. M.|year=2021|title=The meta-gut: community coalescence of animal gut and environmental microbiomes|journal=Scientific Reports|volume=11|issue=1|page=23117|doi=10.1038/s41598-021-02349-1|pmid=34848778 |pmc=8633035 |bibcode=2021NatSR..1123117D }}</ref> On occasion, hippos have been filmed eating [[carrion]], usually near the water. There are other reports of meat-eating and even [[cannibalism]] and [[predation]].<ref>{{Cite journal|title = Reports of carnivory by the common hippo ''Hippopotamus amphibius''|author = Dudley, J. P.|journal = South African Journal of Wildlife Research| date=January 1998 |volume = 28|issue = 2|pages = 58–59|url = https://journals.co.za/content/wild/28/2/EJC117046}}</ref> Hippos' stomach anatomy lacks adaptions to carnivory and meat-eating is likely caused by lack of nutrients or just an abnormal behaviour.<ref name="The Hippos"/>{{rp|82–84}} |
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Adult hippos move at speeds up to {{convert|8|km/h|0|abbr=on}} in water; typically resurfacing to breathe every three to five minutes. The young have to breathe every two to three minutes.<ref name="The Hippos"/>{{rp|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.<ref>{{cite web|title=Hippopotamuses|url=http://www.pbs.org/wnet/nature/critters/hippo.html|publisher=PBS Nature|accessdate=10 July 2013}}</ref> 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 (biology)|mutualism]] in which the hippo benefits from the cleaning, while the fish receive food.<ref>{{cite book|author=Balcombe, Jonathan|year=2006|title=Pleasurable Kingdom: Animals and the Nature of Feeling Good|publisher=Palgrave Macmillan|pages=132–33|isbn=1-4039-8602-9}}</ref> |
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===Social life=== |
===Social life=== |
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[[File:Hippo pod edit.jpg|thumb|right| |
[[File:Hippo pod edit.jpg|thumb|right|Hippopotamus pod]] |
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It is challenging to study the interaction of bulls and cows because hippos are not [[Sexual dimorphism|sexually dimorphic]], so cows and young bulls are almost indistinguishable in the field.<ref>{{Cite journal |journal=[[African Zoology]] |volume=37 |issue=2 |author1=Beckwitt, R. |author2=Shea, J. |author3=Osborne, D. |author4=Krueger, S. |author5=Barklow, W. |year=2002 |title=A PCR-based method for sex identification in ''Hippopotamus amphibius'' |pages=127–130 |url=http://www.framingham.edu/~rbeckwitt/hippo2002.pdf |url-status=dead |archive-url=https://web.archive.org/web/20100617172906/http://www.framingham.edu/~rbeckwitt/hippo2002.pdf |archive-date=17 June 2010 |doi=10.1080/15627020.2002.11657167 |s2cid=88102117 }}</ref> Hippo pods fluctuate but can contain over 100 hippos. Although they lie close together, adults develop almost no social bonds. Males establish [[Territory (animal)|territories]] in water but not land, and these may range {{cvt|250|–|500|m}} in lakes and {{cvt|50|–|100|m}} in rivers. Territories are abandoned when the water dries up. The bull has breeding access to all the cows in his territory. Younger bachelors are allowed to stay as long as they defer to him. A younger male may challenge the old bull for control of the territory. Within the pods, the hippos tend to segregate by sex and status. Bachelor males lounge near other bachelors, females with other females, and the territorial male is on his own. When hippos emerge from the water to graze, they do so individually.<ref name="The Hippos"/>{{rp|4–5, 49–50}} |
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[[File:Hippo fight.jpg|thumb|left|Male hippos fighting]] |
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Hippopotamuses are territorial only in water, where a bull presides over a small stretch of river, on average {{convert|250|m|yards|abbr=on}} in length, and containing 10 females. The largest pods can contain over 100 hippos.<ref name="The Hippos"/>{{rp|50}} Other 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.<ref name="The Hippos"/>{{rp|4}} |
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Hippos engage in "muck-spreading" which involves defecating while spinning their tails to distribute the faeces over a greater area. Muck-spreading occurs both on land and in water and its function is not well understood. It is unlikely to serve a territorial function, as the animals only establish territories in the water. They may be used as trails between the water and grazing areas.<ref name="The Hippos"/>{{rp|5, 51–52}} "Yawning" serves as a threat display.<ref name=estes/> When fighting, bulls use their incisors to block each other's attacks and their large canines as offensive weapons.<ref name=kingdon/>{{rp|259–260}} When hippos become over-populated or a habitat shrinks, bulls sometimes attempt [[Infanticide (zoology)|infanticide]], but this behaviour is not common under normal conditions.<ref name="Infanticide"/> |
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The most common hippo vocalisation is the "wheeze honk", which can travel over long distances in air.<ref name=wheeze-honk/> This call starts as a high-pitched squeal followed by a deeper, resonant call.<ref name="The Hippos"/>{{rp|5}} The animals can recognise the calls of other individuals. Hippos are more likely to react to the wheeze honks of strangers than to those they are more familiar with.<ref name=wheeze-honk>{{cite journal|last1=Thévenet|first1=J.|last2=Grimault|first2=N.|last3=Fonseca|first3=P.|last4=Mathevon|first4=N.|year=2022|title=Voice-mediated interactions in a megaherbivore|journal=Current Biology|volume=32|issue=2|pages=R70–R71|doi=10.1016/j.cub.2021.12.017|pmid=35077689 |bibcode=2022CBio...32..R70T |s2cid=246242737 |url=https://hal.archives-ouvertes.fr/hal-03543609/file/Pre-Print-CURRENT-BIOLOGY.pdf |archive-url=https://ghostarchive.org/archive/20221009/https://hal.archives-ouvertes.fr/hal-03543609/file/Pre-Print-CURRENT-BIOLOGY.pdf |archive-date=2022-10-09 |url-status=live }}</ref> When threatened or alarmed, they produce exhalations,<ref name=estes/> and fighting bulls will bellow loudly.<ref name="The Hippos"/>{{rp|5}} Hippos are recorded to produce clicks underwater which may have [[Animal echolocation|echolocative]] properties.<ref>{{cite journal|last1=Maust-Mohl|first1=M|last2=Soltis|first2=J|last3= Reiss|first3=D|year=2018|title=Underwater click train production by the hippopotamus (''Hippopotamus amphibius'') suggests an echo-ranging function|journal=Behaviour|volume=155|issue=2–3|pages=231–251|doi=10.1163/1568539X-00003484|jstor=26488527}}</ref> They 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 both above and below water.<ref>{{Cite journal|title = Low-frequency sounds and amphibious communication in ''Hippopotamus amphibious''|url = http://scitation.aip.org/getabs/servlet/GetabsServlet?prog=normal&id=JASMAN000115000005002555000001&idtype=cvips&gifs=yes|journal = The Journal of the Acoustical Society of America|author = Barklow, William E.|year = 2004|volume = 115|issue = 5|page = 2555|doi=10.1121/1.4783854|bibcode = 2004ASAJ..115.2555B}}</ref> |
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===Reproduction=== |
===Reproduction=== |
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[[File: |
[[File:Hippofetus.jpg|thumb|right|Preserved hippopotamus fetus]] |
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[[File:Mother and Young Hippo, Uganda (15397037561).jpg|thumb|right|Cow and calf]] |
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Female hippos reach sexual maturity at five to six years of age and have a [[pregnancy (mammals)|gestation period]] of eight months. A study of [[endocrine system]]s revealed that female hippopotamuses may begin puberty as early as three or four years of age.<ref name="Endocrine">{{Cite journal|title = Endocrine patterns associated with reproduction in the Nile hippopotamus (Hippopotamus amphibius) as assessed by fecal progestagen analysis|journal=General and Comparative Endocrinology|volume=128|issue=1|year=2002|pages=74–81|author=Graham L.H., Reid K.; Webster T.; Richards M.; Joseph S.|doi=10.1016/S0016-6480(02)00066-7|pmid = 12270790}}</ref> Males reach maturity at around 7.5 yr. A study of hippopotamus reproductive behavior 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 [[estrous 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.<ref name="The Hippos"/>{{rp|60–61}} After becoming pregnant, a female hippopotamus will typically not begin ovulation again for 17 months.<ref name="Endocrine"/> |
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Cows reach sexual maturity at five to six years of age and have a [[pregnancy (mammals)|gestation period]] of eight months.<ref>{{cite journal|last1=Brown|first1=C. Emerson|title=Rearing Hippopotamuses in Captivity|journal=Journal of Mammalogy|date=November 1924|volume=5|issue=4|pages=243–246|doi=10.2307/1373731|jstor=1373731}}</ref> A study of [[endocrine system]]s revealed cows may begin puberty at as early as three or four years.<ref name="Endocrine">{{Cite journal|title = Endocrine patterns associated with reproduction in the Nile hippopotamus (''Hippopotamus amphibius'') as assessed by fecal progestagen analysis|journal=General and Comparative Endocrinology|volume=128|issue=1|year=2002|pages=74–81|author1=Graham, L. H.|author2= Reid, K. |author3=Webster, T. |author4=Richards, M. |author5=Joseph, S. |doi=10.1016/S0016-6480(02)00066-7|pmid = 12270790}}</ref> Bulls reach maturity at around 7.5 years. Both conceptions and births are highest during the [[wet season]]. Male hippos always have mobile [[spermatozoa]] and can breed year-round.<ref name="The Hippos"/>{{rp|59–61, 66}} After becoming pregnant, a female hippo will typically not begin ovulation again for 17 months.<ref name="Endocrine"/> Hippos mate in the water, with the cow remaining under the surface,<ref name="The Hippos"/>{{rp|63}} her head emerging periodically to draw breath. Cows give birth in seclusion and return within 10 to 14 days. Calves are born on land or shallow water<ref name=estes/> weighing on average {{cvt|50|kg}} and at an average length of around {{cvt|127|cm}}. The female lies on her side when nursing, which can occur underwater or on land. The young are carried on their mothers' backs in deep water.<ref name="The Hippos"/>{{rp|4, 64}} |
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Mother hippos are very protective of their young, not allowing others to get too close.<ref name=estes/> One cow was recorded protecting a calf's carcass after it had died.<ref>{{cite journal|author1=Inman, V. L.|author2=Leggett, K. E. A.|year=2020|title=Observations on the response of a pod of hippos to a dead juvenile hippo (''Hippopotamus amphibius'', Linnaeus 1758)|journal=African Journal of Ecology|volume=58|issue=1|pages=123–125|doi=10.1111/aje.12644|bibcode=2020AfJEc..58..123I |s2cid=191169281 }}</ref> Calves may be temporarily kept in [[Crèche (zoology)|nurseries]], guarded by one or more adults, and will play amongst themselves.<ref name=estes/> Like many other large mammals, hippos are described as [[k-selection|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).<ref name="Endocrine"/><ref name="Infanticide">{{Cite journal|doi = 10.1080/08927014.1998.9522857|title = Infanticide in the hippopotamus: evidence for polygynous ungulates|author = Lewison, R.|journal = Ethology, Ecology & Evolution|volume = 10|issue = 3|pages = 277–286|year = 1998| bibcode=1998EtEcE..10..277L |url = http://www.fupress.net/index.php/eee/article/viewFile/805/751|access-date = 11 July 2010|archive-date = 6 March 2011|archive-url = https://web.archive.org/web/20110306033554/http://www.fupress.net/index.php/eee/article/viewFile/805/751|url-status = dead}}</ref> Calves no longer need to suckle when they are a year old.<ref name="The Hippos"/>{{rp|64}} |
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===Interspecies interactions=== |
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===Aggression=== |
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[[File:Hippo |
[[File:Kruger-Park-Hippo-And-Crocodile.jpg|thumb|right|A hippopotamus and Nile crocodile side by side in Kruger National Park]] |
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Hippos coexist alongside a variety of large predators in their habitats. [[Nile crocodile]]s, [[lion]]s, and [[spotted hyena]]s are known to prey on young hippos.<ref name=estes/> Beyond these, adult hippos are not usually preyed upon by other animals due to their aggression and size. Cases where large lion prides have successfully preyed on adult hippos have been reported, but it is generally rare.<ref>{{cite book|last = Hunter|first = Luke|title = Carnivores of the World|publisher = [[Princeton University Press]]|year = 2011|isbn = 978-0-691-15228-8}}</ref> Lions occasionally prey on adults at [[Gorongosa National Park]] and calves are sometimes taken at Virunga.<ref>{{Cite book | last= Schaller |first= George B. |year= 1972 |title= The Serengeti lion: A study of predator–prey relations|publisher= University of Chicago Press|pages=208–209|isbn = 978-0-226-73639-6 }}</ref> [[Crocodile]]s are frequent targets of hippo aggression, probably because they often inhabit the same [[riparian]] habitats; crocodiles may be either aggressively displaced or killed by hippos.<ref>{{cite book|last1 = Ross|first1 = Charles A.|last2 = Garnett|first2 = Stephen|title = Crocodiles and Alligators|publisher = Checkmark Books|year = 1989|isbn = 978-0-8160-2174-1}}</ref> In turn, very large Nile crocodiles have been observed preying occasionally on calves, "half-grown" hippos, and possibly also adult female hippos. Groups of crocodiles have also been observed finishing off still-living male hippos that were previously injured in mating battles with other males.<ref name= Cott>{{cite journal|author=Cott, H. B. |title=Scientific results of an inquiry into the ecology and economic status of the Nile crocodile (''Crocodilus niloticus'') in Uganda and Northern Rhodesia|journal=The Transactions of the Zoological Society of London|volume=29|issue=4|pages=211–356|doi=10.1111/j.1096-3642.1961.tb00220.x|year=2010}}</ref><ref name= Guggisberg>{{cite book | author = Guggisberg, C. A. W. | title = Crocodiles: Their Natural History, Folklore, and Conservation| year = 1972 | isbn = 978-0-7153-5272-4 | page = 195| publisher = David & Charles}}</ref> |
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Hippos occasionally visit [[cleaning stations]] in order to be cleaned of parasites by certain species of fishes. They signal their readiness for this service by opening their mouths wide. This is an example of [[Mutualism (biology)|mutualism]], in which the hippo benefits from the cleaning while the fish receive food.<ref>{{cite book|author1 =Balcombe, Jonathan|year=2006|title=Pleasurable Kingdom: Animals and the Nature of Feeling Good|url =https://archive.org/details/pleasurablekingd00balc_653|url-access =limited|publisher=Palgrave Macmillan|pages=[https://archive.org/details/pleasurablekingd00balc_653/page/n140 132]–133|isbn=978-1-4039-8602-3}}</ref> Hippo defecation creates [[allochthonous]] deposits of organic matter along the river beds. These deposits have an unclear ecological function.<ref name="Grey">{{Cite journal|title = Using Stable Isotope Analyses To Identify Allochthonous Inputs to Lake Naivasha Mediated Via the Hippopotamus Gut| author = Grey, J.| author2 = Harper, D. M.|journal = Isotopes in Environmental Health Studies|volume = 38|issue = 4|year = 2002|pages = 245–250|doi = 10.1080/10256010208033269| pmid = 12725427| bibcode = 2002IEHS...38..245G| s2cid = 216152807}}</ref> A 2015 study concluded hippo dung provides nutrients from terrestrial material for fish and aquatic invertebrates,<ref>{{cite journal|author=McCauley, D. J.|year=2015|title=Carbon stable isotopes suggest that hippopotamus-vectored nutrients subsidize aquatic consumers in an East African river|journal=Ecosphere|volume=6|issue=4|pages=1–11|doi=10.1890/ES14-00514.1|display-authors=etal|doi-access=free}}</ref> while a 2018 study found that their dung can be toxic to aquatic life in large quantities, due to absorption of dissolved oxygen in water bodies.<ref>{{cite journal|author1=Dutton, C. L. |author2=Subalusky, A. L. |author3=Hamilton, S. K. |author4=Rosi, E. J. |author5=Post, D. M. |year=2018|title=Organic matter loading by hippopotami causes subsidy overload resulting in downstream hypoxia and fish kills|journal=Nature Communications|volume=9|issue=1951|page=1951 |doi=10.1038/s41467-018-04391-6|pmid=29769538 |pmc=5956076|bibcode=2018NatCo...9.1951D }}</ref><ref>{{cite web |url=https://www.theatlantic.com/science/archive/2018/05/hippos-poop-so-much-that-sometimes-all-the-fish-die/560486/ |title=Hippos Poop So Much That Sometimes All the Fish Die |last=Yong |first=Ed |work=The Atlantic |date=16 May 2018 |access-date=23 June 2021 }}</ref> |
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Hippos [[territorial marking|mark their territory]] by defecation. While depositing the faeces, hippos spin their tails distribute their excrement over a greater area.<ref>[http://news.nationalgeographic.com/news/2006/11/061110-poop-scoop_2.html National Geographic exhibit on different animals and their poop]. News.nationalgeographic.com (October 28, 2010). Retrieved on 2012-05-12.</ref> "Yawning" serves as a threat display.<ref name=estes/> When fighting, male hippos use their incisors to block each other's attacks and their large canines to inflict injuries.<ref name=kingdon/>{{rp|260}} When hippos become over-populated or a habitat is reduced, bulls sometimes attempt [[infantacide]], but this behavior is not common under normal conditions.<ref name="Infanticide"/> Incidents of hippo cannibalism have been documented, but this is believed to be the behavior of distressed or sick hippos.<ref name="The Hippos"/>{{rp|82–83}} |
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The parasitic [[monogenea]]n [[flatworm]]'' Oculotrema hippopotami'' infests hippopotamus eyes, mainly the nictitating membrane. It is the only monogenean species (which normally live on fish) documented to live on a mammal.<ref>{{cite journal|author1=Rubtsova, N. Y.|author2=Heckmann, R. A.|author3=Smit, W. J.|author4=Luus-Powell, W. J.|author5=Halajian, A.|author6=Roux, F.|year=2018|title=Morphological studies of developmental stages of ''Oculotrema hippopotami'' (Monogenea: Polystomatidae) infecting the eye of ''Hippopotamus amphibius'' (Mammalia: Hippopotamidae) ssing SEM and EDXA with notes on histopathology|journal=The Korean Journal of Parasitology|volume=56|issue=5|pages=463–475|doi=10.3347/kjp.2018.56.5.463|pmid=30419732 |pmc=6243182 |s2cid=53289954 }}</ref> |
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==Hippos and humans== |
==Hippos and humans== |
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[[File:Hippopotamus |
[[File:Standing Hippopotamus MET DP248993.jpg|thumb|Hippopotamus ([[William the Faience Hippopotamus|"William"]]), [[Middle Kingdom of Egypt]], {{circa}} 1961–1878 BC]] |
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[[File: |
[[File:NMW - Römisches Mosaik 1b.jpg|thumb|[[Roman mosaic]] showing a hippopotamus in the river Nile, [[Roman Egypt]], 1st century AD]] |
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Cut marks on bones of ''H. amphibius'' found at [[Bolomor Cave]], a site in Spain preserving fossils dating from 230,000 to 120,000 years ago, provides evidence for [[Neanderthal]] butchery of hippopotamuses.<ref>{{Cite journal |last=Pawłowska |first=K. |date=2017 |title=Large mammals affected by hominins: Paleogeography of butchering for the European Early and Middle Pleistocene |journal=[[Quaternary International]] |volume=438 |pages=104–115 |doi=10.1016/j.quaint.2017.03.043|bibcode=2017QuInt.438..104P }}</ref><ref>{{Cite journal |last1=Blasco |first1=Ruth |last2=Rosell |first2=Jordi |last3=Fernández Peris |first3=Josep |last4=Arsuaga |first4=Juan Luis |last5=Bermúdez de Castro |first5=José María |last6=Carbonell |first6=Eudald |date=June 2013 |title=Environmental availability, behavioural diversity and diet: a zooarchaeological approach from the TD10-1 sublevel of Gran Dolina (Sierra de Atapuerca, Burgos, Spain) and Bolomor Cave (Valencia, Spain) |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379113001133 |journal=Quaternary Science Reviews |language=en |volume=70 |pages=124–144 |doi=10.1016/j.quascirev.2013.03.008|bibcode=2013QSRv...70..124B }}</ref> The earliest evidence of modern human interaction with hippos comes from butchery cut marks on hippo bones found at the [[Bouri Formation]] and dated to around 160,000 years ago.<ref name=Clark>{{cite journal |doi=10.1038/nature01670 |last1=Clark |first1=J. D. |last2=Beyene |first2=Y. |last3=WoldeGabriel |first3=G. |last4=Hart |first4=W. K. |last5=Renne |first5=P. R. |last6=Gilbert |first6=H. |last7=Defleur |first7=A. |last8=Suwa |first8=G. |last9=Katoh |first9=S. |last10=Ludwig |first10=K. R. |last11=Boisserie |first11=J.-R. |last12=Asfaw |first12=Berhane |last13=White |first13=T. D. |year=2003 |title=Stratigraphic, chronological and behavioural contexts of Pleistocene Homo sapiens from Middle Awash, Ethiopia |journal=Nature |volume=423 |issue=6941 |pages=747–752 |pmid=12802333 |bibcode=2003Natur.423..747C |s2cid=4312418}}</ref> 4,000–5,000 year art showing hippos being hunted have been found in the [[Tassili n'Ajjer|Tassili n'Ajjer Mountains]] of the central [[Sahara]] near [[Djanet]].<ref name="The Hippos"/>{{rp|1}} The [[ancient Egypt]]ians recognised the hippo as ferocious, and representations on the tombs of nobles show humans hunting them.<ref>{{cite book |author=Atiya, F. |year=2008 |title=Ancient Egypt |publisher=The American University in Cairo Press |page=164|isbn=978-9771744399}}</ref> |
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[[File:Taweret statue.jpg|thumb|right|Egyptian god [[Taweret]] statue]] |
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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.<ref name="Clark">{{cite journal|doi = 10.1038/nature01670|last1 = Clark|first1 = JD|last2 = Beyene|first2 = Y|last3 = WoldeGabriel|first3 = G|last4 = Hart|first4 = WK|last5 = Renne|first5 = PR|last6 = Gilbert|first6 = H|last7 = Defleur|first7 = A|last8 = Suwa|first8 = G|last9 = Katoh|first9 = S et al.|last10 = Ludwig|first10 = Kenneth R.|last11 = Boisserie|first11 = Jean-Renaud|last12 = Asfaw|first12 = Berhane|last13 = White|first13 = Tim D.|year = 2003|title = Stratigraphic, chronological and behavioural contexts of Pleistocene Homo sapiens from Middle Awash, Ethiopia|journal = Nature|volume = 423|issue = 6941| pages = 747–52|pmid = 12802333|display-authors = 8}}</ref> 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|Tassili n'Ajjer Mountains]].<ref name="The Hippos"/>{{rp|1}} The [[ancient Egyptians]] recognized the hippo as a ferocious denizen of the [[Nile]]. |
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The |
The hippo was also known to the [[Ancient Greece|Greeks]] and [[Ancient Rome|Romans]]. The Greek historian [[Herodotus]] described the hippo in ''[[Histories (Herodotus)|The Histories]]'' (written ''[[Wiktionary:circa|circa]]'' 440 BC) and the Roman naturalist [[Pliny the Elder]] wrote about the hippo in his encyclopedia ''[[Naturalis Historia]]'' (written ''circa'' 77 AD).<ref name="Pliny"/><ref>{{Cite book |title= The Histories |author=Herodotus|author-link=Herodotus |chapter=Chapter 71, Book II |isbn = 978-0-19-521974-6|title-link = Histories (Herodotus)|date =2003| publisher=Oxford University Press, Incorporated |chapter-url=https://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Aabo%3Atlg%2C0016%2C001&query=2%3A71%3A1}}</ref> The [[Yoruba people]] called the hippo ''erinmi'', which means "elephant of the water".<ref>{{cite book |author=Drewal, M. T. |year=1992|title=Yoruba Ritual: Performers, Play, Agency |publisher=Indiana University Press|page=74|isbn=978-0253112736}}</ref> Some individual hippos have achieved international fame. [[Huberta (hippopotamus)|Huberta]] became a celebrity during the Great Depression for trekking a great distance across [[South Africa]].<ref>{{Cite book |title=Huberta Goes South, a Record of the Lone Trek of the Celebrated Zululand Hippopotamus |author=Chilvers, H. A. |year=1931 |publisher=Gordon & Gotch|location=London}}</ref><ref name=reaktion/>{{rp|111–112}} |
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[[File:Kalabari masks BM.jpg|thumb|150px|left|Ijaw hippopotamus masks]] |
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[[Zulu people|Zulu]] warriors preferred to be as brave as a hippopotamus, since even lions were not considered to match its courage. "In 1888, [[Robert Baden-Powell, 1st Baron Baden-Powell|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 Robert Dunn|John Dunn]], a white Zulu chief, who led an ''[[impi]]'' (army) of 2,000 Zulu warriors to join the British." <ref>[http://www.scouting.org.za/seeds/ingonyama.html Ingonyama – he is a lion!]. Scouting.org.za. Retrieved on 2011-03-29.</ref> |
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===Attacks on humans=== |
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The words of the Zulu anthem sounded like this: |
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The hippo is considered to be extremely aggressive and has frequently been reported charging and attacking boats.<ref>{{cite web |url=http://www.petethomasoutdoors.com/2015/01/massive-hippo-charges-safari-boat-in-malawi-africa-.html|title=Massive hippo charges safari boat in Malawi, Africa|last=Thomas|first=Pete|publisher=Pete Thomas Outdoor|date=14 January 2015}}</ref> Small boats can easily be capsized by hippos and passengers can be injured or killed by the animals, or drown in the water. In one 2014 case in [[Niger]], a boat was capsized by a hippo and 13 people were killed.<ref>{{cite news|url=http://www.abc.net.au/news/2014-11-20/hippopotamus-attack-kills-13-in-boat-in-niger/5904646|title=Hippopotamus attack kills 13 people, including 12 children, in boat near Niger's capital Niamey|publisher=Australian Broadcasting Corporation|date=20 November 2014}}</ref> Hippos will often raid farm crops if the opportunity arises, and humans may come into conflict with them on these occasions. These encounters can be fatal to either humans or hippos.<ref>{{Cite journal |author=Kendall, C. J. |year=2011 |title=The spatial and agricultural basis of crop raiding by the Vulnerable common hippopotamus ''Hippopotamus amphibius'' around Ruaha National Park, Tanzania |journal=Oryx |volume=45 |issue=1 |pages=28–34 |doi=10.1017/S0030605310000359 |doi-access=free}}</ref> |
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According to the [[Ptolemaic kingdom|Ptolemaic]] historian [[Manetho]], the [[pharaoh]] [[Menes]] was carried off and then killed by a hippopotamus.<ref>{{Cite book |last=Elder |first=E. |year=1849 |title=Dictionary of Greek and Roman biography and mythology |editor-last=Smith |editor-first=W. |volume=2 |place=Boston |publisher=[[Little, Brown and Company|Charles C. Little & James Brown]] |url=https://books.google.com/books?id=HP4rAAAAYAAJ}}</ref> |
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"''Een-gonyama Gonyama! Invooboo! Yah-bo! Yah-bo! Invooboo!'' |
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===In zoos=== |
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"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.'"<ref>{{cite web|last=Orans|first=Lewis P.|url=http://www.pinetreeweb.com/bp-dinizulu.htm|title= Scouting in South Africa |
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[[File:Obaysch 1852.jpg|thumb|right|[[Obaysch]] lounging at the [[London Zoo]] in 1852]] |
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1884-1890|publisher=Pinetreeweb.com|date=1997-06-17|archiveurl= https://web.archive.org/web/20140728235654/http://pinetreeweb.com/bp-dinizulu.htm|archivedate= 2014-07-28|deadurl= no|accessdate=2011-03-29}}</ref> |
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Hippos have long been popular zoo animals. The first record of hippos taken into captivity for display is dated to 3500 BC in Hierakonpolis, Egypt.<ref>{{cite journal |author=Rose, M. |year=2010|title=World's first Zoo-Hierakonpolis, Egypt. |journal=Archaeology |volume=63 |issue=1 |pages=25–32}}</ref> The first zoo hippo in modern history was [[Obaysch]], who arrived at the [[London Zoo]] on 25 May 1850, where he attracted up to 10,000 visitors a day and inspired a popular song, the "Hippopotamus Polka".<ref name=Mania>{{Cite journal |author=Root, N. J. |title=Victorian England's Hippomania |journal=[[Natural History (magazine)|Natural History]] |volume=103 |year=1993 |pages=34–39}}</ref> |
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Hippos generally breed well in captivity; birth rates are lower than in the wild, but this can be attributed to zoos' desire to limit births, since hippos are relatively expensive to maintain.<ref name=Mania/><ref>{{cite journal|last1=Snyder|first1=K. D. |title=The common Hippopotamus in the wild and in captivity: Conservation for less charismatic species |journal=Journal of International Wildlife Law & Policy |date=2015 |volume=18 |issue=4 |pages=337–354 |doi=10.1080/13880292.2015.1096162 |s2cid=86295612}}</ref> Starting in 2015, the [[Cincinnati Zoo]] built a US$73 million exhibit to house three adult hippos, featuring a {{cvt|250,000|L}} tank. Modern hippo enclosures also have a complex filtration system for the animal's waste, an underwater viewing area for the visitors, and glass that may be up to {{cvt|9|cm}} thick and capable of holding water under pressures of {{cvt|31|kPa}}.<ref name=reaktion/>{{rp|158–159}} |
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In the U.S., [[United States House of Representatives|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.<ref name="wired2013">{{cite journal| date=2013-12-20| last = Miller | first = Greg | title= The Crazy, Ingenious Plan to Bring Hippopotamus Ranching to America| journal=[[Wired (magazine)|Wired]] | url= http://www.wired.com/2013/12/hippopotamus-ranching/|archiveurl= https://web.archive.org/web/20150117001302/http://www.wired.com/2013/12/hippopotamus-ranching/|archivedate= 2015-01-17|deadurl= no | issn=1059-1028}}</ref><ref name="mooallem2013">{{cite book | last =Mooallem | first = John | title = American Hippopotamus | publisher =The Atavist | year =2013 | location =New York | url = https://periodic.atavist.com/view/theatavist/story/199 | accessdate = March 7, 2014}}</ref> 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.<ref name="mooallem2013"/><ref name="dailybeast2014">{{cite journal| date=August 10, 2014| last = Mooallem | first = Jon | title=Lake Bacon: The Story of The Man Who Wanted Us to Eat Mississippi Hippos| journal=[[The Daily Beast]] | publisher = The Newsweek Daily Beast Company | url =http://www.thedailybeast.com/articles/2014/08/10/the-men-who-wanted-to-every-american-to-eat-wholesome-hippo.html | accessdate= August 13, 2014 | issn=0028-9604}}</ref> The chief collaborators and proponents of Broussard's bill were Major [[Frederick Russell Burnham]] and Captain [[Fritz Joubert Duquesne|Fritz Duquesne]]<ref name="sciam2014">{{cite journal| date=March 27, 2014| last = Eplett| first = Layla | title=The hunger game meat: How hippos early invaded American cuisine| journal=[[Scientific American]] | url=http://blogs.scientificamerican.com/food-matters/2014/03/27/the-hunger-game-meat-how-hippos-nearly-invaded-american-cuisine/ | issn=0036-8733}}</ref><ref name="chances">{{cite book | last =Burnham | first =Frederick Russell | authorlink =Frederick Russell Burnham | title =Taking Chances | publisher =Haynes Corp | year =1944 | location =Los Angeles| url = | doi = | isbn = 1-879356-32-5| pages = 11–23}}</ref> 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".<ref name="sciam2014"/> The "American Hippo Bill" fell just short of being passed.<ref name="mooallem2013"/> |
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===Cultural significance=== |
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[[File:Kalabari masks BM.jpg|thumb|left|upright|Ijaw hippopotamus masks]] |
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The hippopotamus is considered very aggressive and have often been reported to charge and attack boats.<ref>http://www.petethomasoutdoors.com/2015/01/massive-hippo-charges-safari-boat-in-malawi-africa-.html</ref> Small boats can be capsized by hippos and those on board have been killed, and in one case in Niger, a boat was capsized by a hippo and 13 people are killed.<ref>http://www.abc.net.au/news/2014-11-20/hippopotamus-attack-kills-13-in-boat-in-niger/5904646</ref> |
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In [[Egyptian mythology]], the god [[Set (deity)|Set]] takes the form of a red hippopotamus and fights [[Horus]] for control of the land, but is defeated. The goddess [[Tawaret]] is depicted as a pregnant woman with a hippo head, representing fierce maternal love.<ref>{{cite book |last=Sax |first=B. |year=2001 |title=The Mythical Zoo: An Encyclopedia of Animals in World Myth, Legend, and Literature |publisher=ABC-CLIO |page=156 |isbn=9781576076125}}</ref> The [[Ijaw people]] of the [[Niger Delta]] wore masks of aquatic animals like the hippo when practising their [[water spirit]] cults,<ref>{{cite book |author=Segy, L. |year=1976|title=Masks of Black Africa |publisher=Courier Corporation |page=128 |isbn=978-0486231815}}</ref> and hippo ivory was used in the [[divination]] rituals of the Yoruba.<ref>{{cite book |author=Blier, S. P. |year=2015|title=Art and Risk in Ancient Yoruba: Ife History, Power, and Identity, ca. 1300|publisher=Cambridge University Press |page=328 |isbn=978-1107021662}}</ref> Hippo masks were also used in [[Nyau]] funerary rituals of the [[Chewa people|Chewa]] of Southern Africa.<ref name=reaktion/>{{rp|120}} According to [[Robert Baden-Powell, 1st Baron Baden-Powell|Robert Baden-Powell]], [[Zulu people|Zulu]] warriors referred to hippos in war chants.<ref>{{cite web|last=Orans |first=L. P. |title=Scouting in South Africa 1884–1890 |url=http://www.pinetreeweb.com/bp-dinizulu.htm |publisher=Pinetreeweb |date=1997 |archive-url=https://web.archive.org/web/20140728235654/http://pinetreeweb.com/bp-dinizulu.htm |archive-date=28 July 2014 |url-status=dead |access-date=29 March 2011}}</ref> The [[Behemoth]] from the [[Book of Job]], 40:15–24 is thought to be based on the hippo.<ref>{{cite book |editor1=Metzger, B. M. |editor2=Coogan, M. D. |title=The Oxford Companion to the Bible |publisher=Oxford University Press |year=1993 |location=Oxford, UK |page=76 |isbn=978-0-19-504645-8 |url=https://archive.org/details/isbn_9780195046458/page/76}}</ref> |
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Hippos have been the subjects of various [[African folklore|African folktales]]. According to a [[San people|San]] story, when the [[Creator deity|Creator]] assigned each animal its place in nature, the hippos wanted to live in the water, but were refused out of fear they might eat all the fish. After begging and pleading, the hippos were finally allowed to live in the water on the condition they would eat grass instead of fish, and fling their dung so it can be inspected for fish bones. In a Ndebele tale, the hippo originally had long, beautiful hair, but it was set on fire by a jealous hare and the hippo had to jump into a nearby pool. The hippo lost most of his hair and was too embarrassed to leave the water.<ref name=folktale>{{cite book |author1=Greaves, N. |author2=Clement, R. |year=2000|title=When Hippo Was Hairy: And Other Tales from Africa |publisher=Struik|pages=67–71|isbn=978-1-86872-456-7}}</ref> |
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===Hippos in zoos=== |
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[[File:Obaysch 1852.jpg|thumb|[[Obaysch]] lounging at the [[London Zoo]] in 1852]] |
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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".<ref name="Mania">{{Cite journal|author = Root, N. J.|title = Victorian England's Hippomania|journal = [[Natural History (magazine)|Natural History]]|volume = 103|year = 1993|pages = 34–39}}</ref> 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.<ref name="The Hippos"/>{{rp|129}}<ref name="Mania"/> |
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[[File:Hippopotamus-polka-early1850s.jpg|thumb|upright|right|The "Hippopotamus Polka"]] |
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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|Toledo Zoo Hippoquarium]], features a 360,000 gallon pool for hippos.<ref name="Zoos">{{Cite news|title= 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.|work = [[The Atlantic Monthly]]|date = December 1987|author = Melissa Greene}}</ref> 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.<ref>{{cite web|url=http://www.toledozoo.org/plantsanimals/pa_hippoquarium.html|title = Hippoquarium|publisher = [[Toledo Zoo]]|accessdate=2007-03-26| archiveurl = http://web.archive.org/web/20070211115522/http://www.toledozoo.org/plantsanimals/pa_hippoquarium.html| archivedate = February 11, 2007}}</ref> |
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Hippopotamuses were rarely depicted in European art during the [[Renaissance]] and [[Baroque]] periods, due to less access to specimens by Europeans. One notable exception is [[Peter Paul Rubens]]' ''[[The Hippopotamus and Crocodile Hunt]]'' (1615–1616).<ref name=reaktion/>{{rp|122–123}} Ever since [[Obaysch]] inspired the "Hippopotamus Polka", hippos have been popular animals in [[Western culture]] for their rotund appearance, which many consider comical.<ref name="Mania"/> The [[The Walt Disney Company|Disney]] film ''[[Fantasia (1940 film)|Fantasia]]'' featured a ballerina hippo dancing to the opera ''[[La Gioconda (opera)|La Gioconda]]''. The film ''[[Hugo the Hippo]]'' is set in Tanzania and involves the title character trying to escape being slaughtered with the help of local children. The ''[[Madagascar (franchise)|Madagascar]]'' films feature a hippo named [[List of Madagascar (franchise) characters#Gloria|Gloria]].<ref name=reaktion>{{Cite book |last1=Willams|first1=E.|year=2017 |title=Hippopotamus |publisher=[[Reaktion Books]] |isbn=9781780237794 |url=https://books.google.com/books?id=sN5BDwAAQBAJ&pg=PA129}}</ref>{{rp|128–129}} Hippos even inspired a popular board game, [[Hungry Hungry Hippos]].<ref>{{cite book |last1=Robinson|first1=P. T.|last2=Flacke|first2=G. L. |last3=Hentschel |first3=K. M. |year=2017|title=The Pygmy Hippo Story: West Africa's Enigma of the Rainforest |publisher=Oxford University Press |isbn=9780190611859}}</ref> |
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Among the most famous poems about the hippo is "The Hippopotamus" by [[T. S. Eliot]], where he uses the animal to represent the Catholic Church. Hippos are 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 featured in the popular "The Hippopotamous Song" by [[Flanders and Swann]].<ref name=reaktion/>{{rp|128, 136}} |
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===Cultural depictions=== |
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[[File:Hippopotamus-polka-early1850s.jpg|thumb|upright|left|The cover of the Hippopotamus Polka. The unlikely portrayal of dancing hippos was echoed in Disney's ''[[Fantasia (1940 film)|Fantasia]]''.]] |
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A red hippo represented the [[Ancient Egyptian religion|Ancient Egyptian god]] [[Set (mythology)|Set]]; the thigh is the "[[phallic]] leg of Set" symbolic of virility. Set's consort [[Tawaret]] was also seen as part hippo<ref name = "Cooper92">{{cite book |last=Cooper |first=J.C. |title=Symbolic and Mythological Animals |page=129 |year=1992 |publisher= Aquarian Press |location=London |isbn=1-85538-118-4}}</ref> and was a goddess of protection in pregnancy and childbirth, because ancient Egyptians recognized the protective nature of a female hippopotamus toward her young.<ref>{{Cite book|author = Hart, George|title = A Dictionary of Egyptian Gods and Goddesses|publisher = [[Routledge]]|year = 1986|isbn = 0-415-05909-7}}</ref> The [[Ijo people]] wore masks of aquatic animals like the hippo when practicing their water spirit cults.<ref>{{cite book|author=Hamilton, Janice|year=2003|title=Nigeria in Pictures|publisher=Twenty-First Century Books|page=49|isbn=0-8225-0373-5}}</ref> The [[Behemoth]] from the [[Book of Job]], 40:15–24 is thought to be based on a hippo.<ref>{{cite book|editor =Metzeger, Bruce M., Coogan, Michael D. f|title = The Oxford Companion to the Bible|publisher = [[Oxford University Press]]| year = 1993|location = Oxford, UK| page = 76| isbn = 0-19-504645-5}}</ref> |
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==See also== |
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Hippos have been the subjects of various African [[Folklore|folktales]]. According to a [[San people|San]] story; when the [[Creator deity|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.<ref name=folktale>{{cite book|author=Greaves, N.; Clement, R.|year=2000|title=When Hippo Was Hairy: And Other Tales from Africa|publisher=Struik|pages=67–71|isbn=1-86872-456-5}}</ref> 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.<ref name=folktale/> |
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{{Portal|Mammals}} |
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* [[American Hippo bill]] - 1910 bill that proposed the introduction of hippos into Louisiana |
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Ever since [[Obaysch]] inspired the "Hippopotamus Polka", hippos have been popular animals in [[Western culture]] for their rotund appearance that many consider comical.<ref name="Mania"/> Stories of hippos such as [[Huberta (hippopotamus)|Huberta]], which became a celebrity in [[South Africa]] in the 1930s for trekking across the country;<ref>{{Cite book|title = Huberta Goes South, a Record of the Lone Trek of the Celebrated Zululand Hippopotamus|author = Chilvers, H.A.|year = 1931|publisher = Gordon & Gotch|location = London}}</ref> 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.<ref name="NPR">{{Cite news|url = http://www.npr.org/templates/story/story.php?storyId=4754996|title = A hippo and tortoise tale|date =July 17, 2005|accessdate = 2007-06-18|work = [[National Public Radio|NPR]]}}</ref><ref>{{Cite book|title = Owen & Mzee; The True Story of a Remarkable Friendship|author = Hatkoff, Isabella; Hatkoff, Craig and Kahumbu, Paula|publisher = [[Scholastic Press]]|location = [[New York, NY|New York]]|year = 2006|isbn = 0-439-82973-9}}</ref> 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.<ref>{{cite web|title=I Want A Hippopotamus For Christmas Lyrics|url=http://www.christmas-lyrics.org/i-want-a-hippopotamus-for-christmas-lyrics.html|accessdate=2007-12-20|publisher=Christmas-lyrics.org}}</ref> 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]].<ref>{{Cite news|title = Childhood Trauma: Hungry Hungry Hippos|url=http://www.multiplayers.com.au/multi-players-articles/2006/5/2/childhood-trauma-hungry-hungry-hippos/|work = Newcastle Herald (Australia)|date =May 2, 2006}}</ref><ref>{{cite news|title = Fred Kroll, of Trouble and Hungry Hungry Hippos games, dead at 82|url=http://jacksonville.com/tu-online/apnews/stories/080603/D7SO2MJ00.html|agency = Associated Press|date =August 5, 2003}}</ref> |
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* [[Armley Hippo]] fossil ''H. amphibius'' specimen from England |
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[[File:Peter Paul Rubens 083.jpg|thumb|''[[The Hippopotamus and Crocodile Hunt]]'' (1617), by [[Peter Paul Rubens]]]] |
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* [[Allenton hippopotamus]] fossil ''H. amphibius'' specimen from England |
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Hippos have also been popular cartoon characters, where their rotund frames are used for humorous effect. The [[The Walt Disney Company|Disney]] film ''[[Fantasia (1940 film)|Fantasia]]'' featured a ballerina hippopotamus dancing to the opera ''[[La Gioconda (opera)|La Gioconda]]''.<ref name="HippoHaven"/> Other cartoon hippos have included [[Hanna-Barbera]]'s [[Peter Potamus]],<ref>{{cite web |url= http://www.toonopedia.com/potamus.htm|title= Peter Potamus|first= Don|last= Markstein|year= 2008|publisher= [[Don Markstein's Toonopedia]]|archiveurl= http://www.webcitation.org/6Wt404v1A|archivedate= 2015-03-08|deadurl= no}}</ref> the book and TV series ''[[George and Martha]]'', [[Flavio and Marita]] on the ''[[Animaniacs]]'', Pat of the French duo ''[[Pat et Stanley]]'', [[The Backyardigans|''The Backyardigan's'']] Tasha, The [[Moomin|Moomins]], and Gloria and Moto-Moto from the ''[[Madagascar (film series)|Madagascar]]'' franchise. |
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* [[Owen and Mzee]] - hippo and tortoise who bonded |
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* [[Fiona (hippopotamus)|Fiona]] - hippo born 6 weeks early at the [[Cincinnati Zoo and Botanical Garden]] |
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The hippopotamus characters "[[Kinder Happy Hippo|Happy Hippos]]" were created in 1987 by the French designer [[André Roche]]<ref>{{cite web |url= http://citypaper.net/Blogs/WEEKLY-CANDY-Kinder-Happy-Hippos/|title= Weekly Candy: Kinder Happy Hippos|date= 2011-11-22|publisher= ''[[Philadelphia City Paper]]''|archiveurl= http://www.webcitation.org/6Wt3bRsEU|archivedate= 2015-03-08|deadurl= no}}</ref> to be hidden in the "[[Kinder Surprise]] egg" of the Italian chocolate company [[Ferrero SpA]]. The Nintendo Company published Game Boy adventures of them in 2001 and 2007. In the [[chess|game of chess]], the hippopotamus lends its name to the [[Hippopotamus Defense]], an opening system, which is generally considered weak. [[River Horse (sculpture)|''The River Horse'']] is a popular outdoor sculpture at [[George Washington University]] in Washington, D.C. |
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==References== |
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*{{cite web |url=http://awf.org/content/wildlife/detail/hippopotamus |archive-url=https://web.archive.org/web/20101119172410/http://awf.org/content/wildlife/detail/hippopotamus |archive-date=19 November 2010 |title=Hippos: Wildlife summary |publisher=[[African Wildlife Foundation]]}} |
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*{{cite web |url=http://www.ml.duke.edu/projects/hippos/ |title=Hippo Specialist Group of the World Conservation Union |publisher=[[International Union for Conservation of Nature]] |archive-url=https://web.archive.org/web/20141122203937/http://www.ml.duke.edu/projects/hippos/ |archive-date=22 November 2014 |url-status=dead |access-date=4 November 2008 }} |
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*{{cite web |url=http://awf.org/content/wildlife/detail/hippopotamus |archiveurl=http://web.archive.org/web/20101119172410/http://awf.org/content/wildlife/detail/hippopotamus |archivedate=2010-11-19 |title=Hippos: Wildlife summary |publisher=[[African Wildlife Foundation]]}} |
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*{{cite |
*{{cite web |url= http://www.history.com/news/history-lists/11-things-you-may-not-know-about-ancient-egypt|title= 11 Things You May Not Know About Ancient Egypt: King Tut may have been killed by a hippopotamus|date= 12 November 2012|publisher= [[History (U.S. TV channel)|History]]|archive-url= https://web.archive.org/web/20141217213422/http://www.history.com/news/history-lists/11-things-you-may-not-know-about-ancient-egypt|archive-date= 17 December 2014|url-status= live}} |
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*{{cite news|url=http://www.telegraph.co.uk/news/worldnews/southamerica/colombia/6096798/Pablo-Escobars-fugitive-hippos-zoologists-called-in-to-round-up-animals.html|title=Pablo Escobar's fugitive hippos: zoologists called in to round up animals|date=2009-08-27|newspaper= The Daily Telegraph|archiveurl= https://web.archive.org/web/20090910101828/http://www.telegraph.co.uk/news/worldnews/southamerica/colombia/6096798/Pablo-Escobars-fugitive-hippos-zoologists-called-in-to-round-up-animals.html|archivedate= 2009-09-10|deadurl= no}} |
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*{{cite news|url=http://www.bbc.com/news/magazine-27905743|title=Pablo Escobar’s hippos: A growing problem|last=Kremer|first=William|date=2014-06-25|publisher=[[BBC News]]|archiveurl= https://web.archive.org/web/20150225044002/http://www.bbc.com/news/magazine-27905743|archivedate= 2015-02-25|deadurl= no}} |
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*{{cite web |url= http://www.ml.duke.edu/projects/hippos/|title= Hippo Specialist Group of the World Conservation Union|publisher= [[International Union for Conservation of Nature]]|archiveurl= http://www.webcitation.org/6WsequYsT|archivedate= 2015-03-08|deadurl= no}} |
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*{{cite web |url= http://www.history.com/news/history-lists/11-things-you-may-not-know-about-ancient-egypt|title= 11 Things You May Not Know About Ancient Egypt: King Tut may have been killed by a hippopotamus|date= 2012-11-12|publisher= [[History (TV channel)|History]]|archiveurl= https://web.archive.org/web/20141217213422/http://www.history.com/news/history-lists/11-things-you-may-not-know-about-ancient-egypt|archivedate= 2014-12-17|deadurl= no}} |
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[[Category:Hippopotamuses| ]] |
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[[Category:Mammals of Africa]] |
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[[Category:Fauna of Sub-Saharan Africa]] |
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[[Category:Semiaquatic mammals]] |
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[[Category:Fauna of West Africa]] |
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[[Category:Hippopotamuses|*]] |
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Latest revision as of 20:33, 3 January 2025
Hippopotamus Temporal range:
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A hippopotamus in Saadani National Park, Tanzania | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Artiodactyla |
Family: | Hippopotamidae |
Genus: | Hippopotamus |
Species: | H. amphibius
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Binomial name | |
Hippopotamus amphibius | |
Current distribution of the hippopotamus |
The hippopotamus (Hippopotamus amphibius) (/ˌhɪpəˈpɒtəməs/; pl.: hippopotamuses; often shortened to hippo (pl.: hippos), further qualified as the common hippopotamus, Nile hippopotamus and river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extant species in the family Hippopotamidae, the other being the pygmy hippopotamus (Choeropsis liberiensis or Hexaprotodon liberiensis). Its name comes from the ancient Greek for "river horse" (ἱπποπόταμος).
After elephants and rhinoceroses, the hippopotamus is the next largest land mammal. It is also the largest extant land artiodactyl. Despite their physical resemblance to pigs and other terrestrial even-toed ungulates, the closest living relatives of the hippopotamids are cetaceans (whales, dolphins, porpoises, etc.), from which they diverged about 55 million years ago. Hippos are recognisable for their barrel-shaped torsos, wide-opening mouths with large canine tusks, nearly hairless bodies, pillar-like legs, and large size: adults average 1,500 kg (3,300 lb) for bulls (males) and 1,300 kg (2,900 lb) for cows (females). Despite its stocky shape and short legs, it is capable of running 30 km/h (19 mph) over short distances.
Hippos inhabit rivers, lakes, and mangrove swamps. Territorial bulls each preside over a stretch of water and a group of five to thirty cows and calves. Mating and birth both occur in the water. During the day, hippos remain cool by staying in water or mud, emerging at dusk to graze on grasses. While hippos rest near each other in the water, grazing is a solitary activity and hippos typically do not display territorial behaviour on land. Hippos are among the most dangerous animals in the world due to their aggressive and unpredictable nature. They are threatened by habitat loss and poaching for their meat and ivory (canine teeth).
Etymology
The Latin word hippopotamus is derived from the ancient Greek ἱπποπόταμος (hippopótamos), from ἵππος (híppos) 'horse' and ποταμός (potamós) 'river', together meaning 'horse of the river'.[3][4][5] In English, the plural is "hippopotamuses".[6]
Taxonomy and origins
Classification
The modern hippopotamus and the pygmy hippopotamus are the only living members of the family Hippopotamidae. Some taxonomists place hippos and anthracotheres in the superfamily Anthracotheroidea. Hippopotamidae are classified along with other even-toed ungulates in the order Artiodactyla.[7]: 39–40
Five subspecies of hippos have been described based on morphological differences in their skulls as well as differences in geographical range:[7]: 3 [8][9][10]
- H. a. amphibius – (the nominate subspecies) ranges from Gambia east to Ethiopia and then south to Mozambique and historically ranged as far north as Egypt; its skull is distinguished by a moderately reduced preorbital region, a bulging dorsal surface, elongated mandibular symphysis and larger chewing teeth.
- H. a. kiboko – found in Kenya and Somalia; was noted to be smaller and more lightly coloured than other hippos with wider nostrils, somewhat longer snout and more rounded and relatively raised orbits with the space between them being incurved.
- H. a. capensis – found in Zambia and South Africa; distinguished by wider orbits.
- H. a. tschadensis – ranges between Chad and Niger; featured a slightly shorter but broader face, and pronounced, forward-facing orbits.
- H. a. constrictus – ranged from the southern Democratic Republic of Congo to Angola and Namibia; skull characterised by a thicker preorbital region, shorter snout, flatter dorsal surface, reduced mandibular symphysis and smaller chewing teeth.
The suggested subspecies above 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.[7]: 2 A study examining mitochondrial DNA from skin biopsies taken from 13 sampling locations 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 have been tested.[9]
Evolution
Until 1909, naturalists classified hippos together with pigs based on molar patterns. Several lines of evidence, first from blood proteins, then from molecular systematics,[12] DNA[13][14] and the fossil record, show their closest living relatives are cetaceans (whales, dolphins, and porpoises).[15][16] 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.[13][16]
The most recent theory of the origins of Hippopotamidae suggests hippos and whales shared a common semiaquatic ancestor that branched off from other artiodactyls around 60 million years ago.[13][15] This hypothesised ancestral group likely split into two branches again around 54 million years ago.[12]
One branch would evolve into cetaceans, possibly beginning about 52 million years ago, with the protowhale Pakicetus and other early whale ancestors collectively known as Archaeoceti. This group eventually underwent aquatic adaptation into the completely aquatic cetaceans.[16] 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 hippos with comparatively smaller, narrower heads. All branches of the anthracotheres, except that which evolved into Hippopotamidae, became extinct during the Pliocene, leaving no descendants.[15][16]
A rough evolutionary lineage of the hippo can thus be traced from Eocene and Oligocene species: from Anthracotherium and Elomeryx to the Miocene species Merycopotamus and Libycosaurus and finally the very latest anthracotheres in the Pliocene.[17] These groups lived across Eurasia and Africa. The discovery of Epirigenys in East Africa, which was likely a descent of Asian anthracotheres and a sister taxon to Hippopotamidae, suggests that hippo ancestors entered Africa from Asia around 35 million years ago.[18][19] An early hippopotamid is the genus Kenyapotamus, which lived in Africa from 15 to 9 million years ago.[17] Hippopotamid species would spread across Africa and Eurasia, including the modern pygmy hippo. From 7.5 to 1.8 million years ago, a possible ancestor to the modern hippo, Archaeopotamus, lived in Africa and the Middle East.[20] The oldest records of the genus Hippopotamus date to the Pliocene (5.3–2.6 million years ago).[21] The oldest unambiguous records of the modern H. amphibius date to the Middle Pleistocene, though there are possible Early Pleistocene records.[22]
Extinct species
Three species of Malagasy hippopotamus became extinct during the Holocene on Madagascar, the last of them within the past 1,000 years. The Malagasy hippos were smaller than the modern hippo, a likely result of the process of insular dwarfism.[23] Fossil evidence indicates many Malagasy hippos were hunted by humans, a factor in their eventual extinction.[23] Isolated individual Malagasy hippos may have survived in remote pockets; in 1976, villagers described a living animal called the kilopilopitsofy, which may have been a Malagasy hippo.[24]
Hippopotamus gorgops from the Early Pleistocene to the early Middle Pleistocene of Africa and West Asia grew considerably larger than the living hippopotamus, with an estimated body mass of over 4,000 kg (8,800 lb).[21][25][26] Hippopotamus antiquus ranged throughout Europe, extending as far north as Britain during the Early and Middle Pleistocene epochs, before being replaced by the modern H. amphibius in Europe during the latter part of the Middle Pleistocene.[27] The Pleistocene also saw a number of dwarf species evolve on several Mediterranean islands, including Crete (Hippopotamus creutzburgi), Cyprus (the Cyprus dwarf hippopotamus, Hippopotamus minor), Malta (Hippopotamus melitensis), and Sicily (Hippopotamus pentlandi). Of these, the Cyprus dwarf hippo survived until the end of the Pleistocene or early Holocene. Evidence from the archaeological site Aetokremnos continues to cause debate on whether or not the species was driven to extinction or even encountered by humans.[28][29]
Characteristics
The hippopotamus is a megaherbivore and is exceeded in size among land animals only by elephants and some rhinoceros species. The mean adult weight is around 1,480 kg (3,260 lb) for bulls and 1,365 kg (3,009 lb) for cows. Exceptionally large males have been recorded reaching 2,660 kg (5,860 lb).[30] Male hippos appear to continue growing throughout their lives, while females reach maximum weight at around age 25.[31] It is 2.90 to 5.05 m (9 ft 6 in to 16 ft 7 in) long,[32] including a tail of about 35 to 56 cm (14 to 22 in) in length and 1.30 to 1.65 m (4 ft 3 in to 5 ft 5 in) tall at the shoulder,[33][34] with males and females ranging 1.40 to 1.65 m (4 ft 7 in to 5 ft 5 in) and 1.30 to 1.45 m (4 ft 3 in to 4 ft 9 in) tall at the shoulder respectively.[34] The species has a typical head–body length of 3.3–3.45 m (10.8–11.3 ft) and an average standing height of 1.4 m (4 ft 7 in) at the shoulder.[35]
Hippos have barrel-shaped bodies with short tails and legs, and an hourglass-shaped skull with a long snout.[36][7]: 3, 19 Their skeletal structures are graviportal, adapted to carrying their enormous weight,[7]: 8 and their dense bones and low centre of gravity allows them to sink and move along the bottom of the water.[37] Hippopotamuses have small legs (relative to other megafauna) because the water in which they live reduces the weight burden.[38] The toes are webbed and the pelvis rests at an angle of 45 degrees.[7]: 3, 9 Though chubby-looking, hippos have little fat.[7]: 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 is submerged.[39]: 259 The nostrils and ears can close when underwater while nictitating membranes cover the eyes.[7]: 4, 116 The vocal folds of the hippo are more horizontally positioned, much like baleen whales. Underneath are throat tissues, where vibrations are transmitted to produce underwater calls.[40]
The hippo's jaw is powered by huge masseter and digastric muscles which give them large, droopy cheeks.[39]: 259 The jaw hinge allows the animal to open its mouth at almost 180°.[7]: 17 A folded orbicularis oris muscle allows the hippo to attain an extreme gape without tearing any tissue.[41] On the lower jaw, the incisors and canines grow continuously, the former reaching 40 cm (16 in), while the latter can grow to up to 50 cm (20 in). The lower canines are sharpened through contact with the smaller upper canines.[36] The canines and incisors are used mainly for combat instead of feeding. Hippos rely on their flattened, horny lips to grasp and pull grasses which are then ground by the molars.[39]: 259, 263 The hippo is considered to be a pseudoruminant; it has a complex three-chambered stomach, but does not "chew cud".[7]: 22
Hippo skin is 6 cm (2.4 in) thick across much of its body with little hair.[36][39]: 260 The animal is mostly purplish-grey or blue-black, but brownish-pink on the underside and around the eyes and ears.[39]: 260 Their skin secretes a natural, red-coloured sunscreen substance that 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 highly acidic pigments have been identified in the secretions; one red hipposudoric acid and one orange norhipposudoric acid, which inhibit the growth of disease-causing bacteria and their light-absorption profile peaks in the ultraviolet range, creating a sunscreen effect.[42][43] Regardless of diet, all hippos secrete these pigments so food does not appear to be their source; rather, they may be synthesised from precursors such as the amino acid tyrosine.[43] This natural sunscreen cannot prevent the animal's skin from cracking if it stays out of water too long.[44]
The testes of the males do not fully descend and a scrotum is not present. In addition, the penis retracts into the body when not erect. The genitals of the female hippos are unusual in that the vagina is ridged and the vulval vestibule has two large, protruding diverticula. Both of these have an unknown function.[7]: 28–29
A hippo's lifespan is typically 40 to 50 years.[39]: 277 Donna the Hippo was one of the oldest living hippos in captivity. She lived at the Mesker Park Zoo in Evansville, Indiana, in the US[45][46] until her death in 2012 at the age of 61.[47] The oldest hippo ever recorded was called Bertha; she had lived in the Manila Zoo in the Philippines since it first opened in 1959. When she died in 2017, her age was estimated to be 65.[48] The oldest living hippopotamus in captivity is Lu the Hippo, from the Ellie Schiller Homosassa Springs Wildlife State Park. As of 2024, he is 64 years old.[49]
Distribution and status
Hippopotamus amphibius arrived in Europe around 560–460,000 years ago, during the Middle Pleistocene.[50] The distribution of Hippopotamus amphibius in Europe during the Pleistocene was largely confined to Southern Europe, including the Iberian Peninsula,[51] Italy,[52][53] and Greece,[54] but extended into northwestern Europe, including Great Britain (as far north as North Yorkshire), the Netherlands, and western Germany during interglacial periods, such as the Last Interglacial (130–115,000 years ago).[55][56][57] The youngest records of the species in Europe are from the Late Pleistocene of Greece, dating to around 40–30,000 years ago.[54][51]
Archaeological evidence exists of its presence in the Levant, dating to less than 3,000 years ago.[58][59] The species was common in Egypt's Nile region during antiquity, but it has since been driven out. According to Pliny the Elder, in his time, the best location in Egypt for capturing this animal was in the Saite nome;[60] the animal could still be found along the Damietta branch of the Nile after the Arab Conquest in 639. Reports of the slaughter of the last hippo in Natal Province were made at the end of the 19th century.[61] 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.[1]
Genetic evidence suggests common hippos in Africa experienced a marked population expansion during or after the Pleistocene, 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.[9] Hippos are also subject to unregulated hunting and poaching. The species is included in Appendix II of the Convention on International Trade in Endangered Species (CITES) meaning international export/import (including in parts and derivatives) requires CITES documentation to be obtained and presented to border authorities.[1][62]
As of 2017, the IUCN Red List drawn up by the International Union for Conservation of Nature (IUCN) lists the species as vulnerable, with a stable population estimated between 115,000 and 130,000 animals.[1] The hippo population has declined most dramatically in the Democratic Republic of the Congo.[63] By 2005, the population in Virunga National Park had dropped to 800 or 900 from around 29,000 in the mid-1970s.[64] This decline is attributed to the disruptions caused by the Second Congo War.[64] The poachers are believed to be Mai-Mai rebels, underpaid Congolese soldiers, and local militia groups.[64][65] Reasons for poaching include the belief hippos are harmful to society, as well as financial gain.[66] As of 2016, the Virunga hippo population appears to have increased again, possibly due to better protection from park rangers, who have worked with local fishermen.[67] The sale of hippo meat is illegal, but black-market sales are difficult for Virunga National Park officers to track.[65][66] Hippo meat is highly valued in some areas of central Africa and the teeth may be used as a replacement for elephant ivory.[68]
A population of hippos exists in Colombia, descended from captive individuals that escaped from Pablo Escobar's estate after his death in 1993. Their numbers grew to 100 by the 2020s and ecologists believe the population should be eradicated, as they are breeding rapidly and are an increasing menace to humans and the environment. Attempts to control them include sterilisation and culling.[69]
Behaviour and ecology
Hippos are semiaquatic and require enough water to immerse in, while being close to grass.[36] They mostly live in freshwater habitat, but can be found in estuaries.[1] They prefer relatively still waters with gently sloping shores, though male hippos may also be found in very small numbers in more rapid waters with rocky slopes.[39]: 264 Like most herbivores, hippos will consume a variety of plants if presented with them in captivity, but their diet in nature consists almost entirely of grass, with only minimal consumption of aquatic plants.[70] Hippos spend most of the day in water to stay cool and hydrated. Just before night begins, they leave the water to forage on land. A hippo will travel 3–5 km (1.9–3.1 mi) per night, eating around 40 kg (88 lb) of grass. By dawn, they are back in the water.[36]
Despite being semiaquatic, an adult hippo is not a particularly good swimmer, nor can it float. It rarely enters deep water; when it does, the animal moves by bouncing off the bottom. An adult hippo surfaces every four to six minutes, while young need to breathe every two to three minutes.[7]: 3–4 Hippos move on land by trotting, and limb movements do not change between speeds. They can reach an airborne stage (a stage when all limb are off the ground) when they move fast enough. Hippos are reported to reach 30 km/h (19 mph) but this has not been confirmed.[71] They are incapable of jumping but can walk up steep banks.[36] The hippopotamus sleeps with both hemispheres of the brain resting, as in all land mammals, and usually sleeps on land or in water with the nostrils exposed. Despite this, it may be capable of sleeping while submerged, intermittently surfacing to breathe without waking. They appear to transition between different phases of sleep more quickly than other mammals.[72]
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, keeping the land clear of vegetation and depressing the ground. Over prolonged periods, hippos can divert the paths of swamps and channels.[73] By defecating in the water, the animals also appear to pass on microbes from their gut, affecting the biogeochemical cycle.[74] On occasion, hippos have been filmed eating carrion, usually near the water. There are other reports of meat-eating and even cannibalism and predation.[75] Hippos' stomach anatomy lacks adaptions to carnivory and meat-eating is likely caused by lack of nutrients or just an abnormal behaviour.[7]: 82–84
Social life
It is challenging to study the interaction of bulls and cows because hippos are not sexually dimorphic, so cows and young bulls are almost indistinguishable in the field.[76] Hippo pods fluctuate but can contain over 100 hippos. Although they lie close together, adults develop almost no social bonds. Males establish territories in water but not land, and these may range 250–500 m (820–1,640 ft) in lakes and 50–100 m (160–330 ft) in rivers. Territories are abandoned when the water dries up. The bull has breeding access to all the cows in his territory. Younger bachelors are allowed to stay as long as they defer to him. A younger male may challenge the old bull for control of the territory. Within the pods, the hippos tend to segregate by sex and status. Bachelor males lounge near other bachelors, females with other females, and the territorial male is on his own. When hippos emerge from the water to graze, they do so individually.[7]: 4–5, 49–50
Hippos engage in "muck-spreading" which involves defecating while spinning their tails to distribute the faeces over a greater area. Muck-spreading occurs both on land and in water and its function is not well understood. It is unlikely to serve a territorial function, as the animals only establish territories in the water. They may be used as trails between the water and grazing areas.[7]: 5, 51–52 "Yawning" serves as a threat display.[36] When fighting, bulls use their incisors to block each other's attacks and their large canines as offensive weapons.[39]: 259–260 When hippos become over-populated or a habitat shrinks, bulls sometimes attempt infanticide, but this behaviour is not common under normal conditions.[77]
The most common hippo vocalisation is the "wheeze honk", which can travel over long distances in air.[78] This call starts as a high-pitched squeal followed by a deeper, resonant call.[7]: 5 The animals can recognise the calls of other individuals. Hippos are more likely to react to the wheeze honks of strangers than to those they are more familiar with.[78] When threatened or alarmed, they produce exhalations,[36] and fighting bulls will bellow loudly.[7]: 5 Hippos are recorded to produce clicks underwater which may have echolocative properties.[79] They 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 both above and below water.[80]
Reproduction
Cows reach sexual maturity at five to six years of age and have a gestation period of eight months.[81] A study of endocrine systems revealed cows may begin puberty at as early as three or four years.[82] Bulls reach maturity at around 7.5 years. Both conceptions and births are highest during the wet season. Male hippos always have mobile spermatozoa and can breed year-round.[7]: 59–61, 66 After becoming pregnant, a female hippo will typically not begin ovulation again for 17 months.[82] Hippos mate in the water, with the cow remaining under the surface,[7]: 63 her head emerging periodically to draw breath. Cows give birth in seclusion and return within 10 to 14 days. Calves are born on land or shallow water[36] weighing on average 50 kg (110 lb) and at an average length of around 127 cm (50 in). The female lies on her side when nursing, which can occur underwater or on land. The young are carried on their mothers' backs in deep water.[7]: 4, 64
Mother hippos are very protective of their young, not allowing others to get too close.[36] One cow was recorded protecting a calf's carcass after it had died.[83] Calves may be temporarily kept in nurseries, guarded by one or more adults, and will play amongst themselves.[36] 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).[82][77] Calves no longer need to suckle when they are a year old.[7]: 64
Interspecies interactions
Hippos coexist alongside a variety of large predators in their habitats. Nile crocodiles, lions, and spotted hyenas are known to prey on young hippos.[36] Beyond these, adult hippos are not usually preyed upon by other animals due to their aggression and size. Cases where large lion prides have successfully preyed on adult hippos have been reported, but it is generally rare.[84] Lions occasionally prey on adults at Gorongosa National Park and calves are sometimes taken at Virunga.[85] 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 hippos.[86] In turn, very large Nile crocodiles have been observed preying occasionally on calves, "half-grown" hippos, and possibly also adult female hippos. Groups of crocodiles have also been observed finishing off still-living male hippos that were previously injured in mating battles with other males.[87][88]
Hippos occasionally visit cleaning stations in order to be cleaned of parasites by certain species of fishes. They signal their readiness for this service by opening their mouths wide. This is an example of mutualism, in which the hippo benefits from the cleaning while the fish receive food.[89] Hippo defecation creates allochthonous deposits of organic matter along the river beds. These deposits have an unclear ecological function.[70] A 2015 study concluded hippo dung provides nutrients from terrestrial material for fish and aquatic invertebrates,[90] while a 2018 study found that their dung can be toxic to aquatic life in large quantities, due to absorption of dissolved oxygen in water bodies.[91][92]
The parasitic monogenean flatworm Oculotrema hippopotami infests hippopotamus eyes, mainly the nictitating membrane. It is the only monogenean species (which normally live on fish) documented to live on a mammal.[93]
Hippos and humans
Cut marks on bones of H. amphibius found at Bolomor Cave, a site in Spain preserving fossils dating from 230,000 to 120,000 years ago, provides evidence for Neanderthal butchery of hippopotamuses.[94][95] The earliest evidence of modern human interaction with hippos comes from butchery cut marks on hippo bones found at the Bouri Formation and dated to around 160,000 years ago.[96] 4,000–5,000 year art showing hippos being hunted have been found in the Tassili n'Ajjer Mountains of the central Sahara near Djanet.[7]: 1 The ancient Egyptians recognised the hippo as ferocious, and representations on the tombs of nobles show humans hunting them.[97]
The hippo was also known to the Greeks and Romans. The Greek historian Herodotus described the hippo in The Histories (written circa 440 BC) and the Roman naturalist Pliny the Elder wrote about the hippo in his encyclopedia Naturalis Historia (written circa 77 AD).[60][98] The Yoruba people called the hippo erinmi, which means "elephant of the water".[99] Some individual hippos have achieved international fame. Huberta became a celebrity during the Great Depression for trekking a great distance across South Africa.[100][101]: 111–112
Attacks on humans
The hippo is considered to be extremely aggressive and has frequently been reported charging and attacking boats.[102] Small boats can easily be capsized by hippos and passengers can be injured or killed by the animals, or drown in the water. In one 2014 case in Niger, a boat was capsized by a hippo and 13 people were killed.[103] Hippos will often raid farm crops if the opportunity arises, and humans may come into conflict with them on these occasions. These encounters can be fatal to either humans or hippos.[104]
According to the Ptolemaic historian Manetho, the pharaoh Menes was carried off and then killed by a hippopotamus.[105]
In zoos
Hippos have long been popular zoo animals. The first record of hippos taken into captivity for display is dated to 3500 BC in Hierakonpolis, Egypt.[106] The first zoo hippo in modern history was Obaysch, who arrived at the London Zoo on 25 May 1850, where he attracted up to 10,000 visitors a day and inspired a popular song, the "Hippopotamus Polka".[107]
Hippos generally breed well in captivity; birth rates are lower than in the wild, but this can be attributed to zoos' desire to limit births, since hippos are relatively expensive to maintain.[107][108] Starting in 2015, the Cincinnati Zoo built a US$73 million exhibit to house three adult hippos, featuring a 250,000 L (55,000 imp gal; 66,000 US gal) tank. Modern hippo enclosures also have a complex filtration system for the animal's waste, an underwater viewing area for the visitors, and glass that may be up to 9 cm (3.5 in) thick and capable of holding water under pressures of 31 kPa (4.5 psi).[101]: 158–159
Cultural significance
In Egyptian mythology, the god Set takes the form of a red hippopotamus and fights Horus for control of the land, but is defeated. The goddess Tawaret is depicted as a pregnant woman with a hippo head, representing fierce maternal love.[109] The Ijaw people of the Niger Delta wore masks of aquatic animals like the hippo when practising their water spirit cults,[110] and hippo ivory was used in the divination rituals of the Yoruba.[111] Hippo masks were also used in Nyau funerary rituals of the Chewa of Southern Africa.[101]: 120 According to Robert Baden-Powell, Zulu warriors referred to hippos in war chants.[112] The Behemoth from the Book of Job, 40:15–24 is thought to be based on the hippo.[113]
Hippos 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 they might eat all the fish. After begging and pleading, the hippos were finally allowed to live in the water on the condition they would eat grass instead of fish, and fling their dung so it can be inspected for fish bones. In a Ndebele tale, the hippo originally had long, beautiful hair, but it was set on fire by a jealous hare and the hippo had to jump into a nearby pool. The hippo lost most of his hair and was too embarrassed to leave the water.[114]
Hippopotamuses were rarely depicted in European art during the Renaissance and Baroque periods, due to less access to specimens by Europeans. One notable exception is Peter Paul Rubens' The Hippopotamus and Crocodile Hunt (1615–1616).[101]: 122–123 Ever since Obaysch inspired the "Hippopotamus Polka", hippos have been popular animals in Western culture for their rotund appearance, which many consider comical.[107] The Disney film Fantasia featured a ballerina hippo dancing to the opera La Gioconda. The film Hugo the Hippo is set in Tanzania and involves the title character trying to escape being slaughtered with the help of local children. The Madagascar films feature a hippo named Gloria.[101]: 128–129 Hippos even inspired a popular board game, Hungry Hungry Hippos.[115]
Among the most famous poems about the hippo is "The Hippopotamus" by T. S. Eliot, where he uses the animal to represent the Catholic Church. Hippos are 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 featured in the popular "The Hippopotamous Song" by Flanders and Swann.[101]: 128, 136
See also
- American Hippo bill - 1910 bill that proposed the introduction of hippos into Louisiana
- Armley Hippo fossil H. amphibius specimen from England
- Allenton hippopotamus fossil H. amphibius specimen from England
- Owen and Mzee - hippo and tortoise who bonded
- Fiona - hippo born 6 weeks early at the Cincinnati Zoo and Botanical Garden
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External links
- "Hippos: Wildlife summary". African Wildlife Foundation. Archived from the original on 19 November 2010.
- "Hippo Specialist Group of the World Conservation Union". International Union for Conservation of Nature. Archived from the original on 22 November 2014. Retrieved 4 November 2008.
- "11 Things You May Not Know About Ancient Egypt: King Tut may have been killed by a hippopotamus". History. 12 November 2012. Archived from the original on 17 December 2014.