Hippopotamus antiquus: Difference between revisions

Hippopotamus antiquus; Temporal range: Early Pleistocene-Middle Pleistocene 2.1–0.4 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Hippopotamus antiquus in Florence Palaeontology museum
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Artiodactyla
Family:	Hippopotamidae
Genus:	Hippopotamus
Species:	†H. antiquus
Binomial name
	†Hippopotamus antiquus; Desmarest 1822
Synonyms
	Hippopotamus amphibius antiquus; Hippopotamus georgicus; Hippopotamus major; Hippopotamus tiberinus;

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Hippopotamus antiquus is an extinct species of Hippopotamus that ranged across Europe during the Early and Middle Pleistocene.

Chronology

H. antiquus is suggested to be closely related to the African species Hippopotamus gorgops, and may be a descendant of that species.^[2] The oldest records of H. antiquus date to the Early Pleistocene, around 2.1-2 million years ago, which are found in Italy and Greece.^[3] The earliest specimens in the Iberian Peninsula date to around 1.7 million years ago.^[4] H. antiquus first became widespread north of the Alps around 1.1 to 1 million years ago, as evidenced from specimens found dating to this time in France, Germany, the Netherlands and southern Britain.^[5] The youngest remains of the species are from Condeixa in Portugal, suggested to date to approximately 400,000 years ago.^[2] Later records of the genus Hippopotamus in Europe are believed to belong to the modern hippopotamus (Hippopotamus amphibius).^[5] The earliest generally accepted record of H. amphibius in Europe is around 500,000 years old, and it is therefore possible that H. antiquus and H. amphibius coexisted in Europe from 500-400,000 years ago, though this is uncertain.^[6]^[7]

Distribution

H. antiquus ranged across Europe, from the Iberian Peninsula and the Italian Peninsula, to the British Isles to the Rhine River to Greece.^[8]^[9] The easternmost record of the species is from the Caucasus, at the Akhalkalaki site in Georgia.^[6] Remains possibly attributable to the species are also known from the Ubeidiya site in Israel,^[2] though other authors assign these remains to the species H. behemoth.^[6] Their distribution was strongly controlled by temperature, with the species only extending to the northern parts of Europe during warmer interglacial intervals.^[5]

Description

Based on the transverse diameter of its limbs, H. antiquus has been estimated to weigh 3,500–4,200 kilograms (7,700–9,300 lb), more than double the weight of the average specimen of H. amphibius.^[10] In comparison to modern Hippopotamus amphibius, the skull is more slender and elongate, but with a shorter neurocranium.^[11] Hippopotamus antiquus has been suggested to have been more aquatically adapted than Hippopotamus amphibius, with the skull having more elevated eyesockets and the feet having shorter metapodial bones than H. amphibius. An analysis of nitrogen isotopes suggests that H. antiquus preferred aquatic plants, in contrast to modern H. amphibius, which prefers terrestrial grasses.^[5]^[12]

The Cretan dwarf hippopotamus (H. creutzburgi) is believed to have evolved from H. antiquus through the process of insular dwarfism on the island of Crete.^[13] The extinct Cyprus dwarf hippopotamus (H. minor) may also derive from H. antiquus, but this is uncertain.^[14]

Relationship with humans

Remains of the species with cut marks suggestive of butchery by archaic humans have been reported from several sites in Spain, dating to the late Early Pleistocene, including Barranco León (~1.4 Ma), Fuente Nueva 3 (~1.3 Ma), and Vallparadís (~1.0 Ma), At the Marathousa 2 site in Greece, thought to date to the Middle Pleistocene around 500-400,000 years ago, remains of a juvenile Hippopotamus antiquus individual with cut marks were found associated with a lithic artefact. It is unclear whether the remains at these sites were hunted or scavenged. It is unlikely that archaic humans regularly hunted healthy adult hippopotamuses due to their dangerousness.^[15]

References

^ Desmarest, A.G., 1822. Mammalogie ou description des espèces de mammifères. Mme Veuve Agasse imprimeur édit., Paris, 2ème part., pp.277-555.
^ ^a ^b ^c Martino, Roberta; Ríos, Maria Ibanez; Mateus, Octavio; Pandolfi, Luca (December 2022). "Taxonomy, chronology, and dispersal patterns of Western European Quaternary hippopotamuses: New insight from Portuguese fossil material". Quaternary International. 674–675: 121–137. doi:10.1016/j.quaint.2022.12.010. S2CID 255029640.
^ Fidalgo, D.; Rosas, A.; Madurell-Malapeira, J.; Pineda, A.; Huguet, R.; García-Tabernero, A.; Cáceres, I.; Ollé, A.; Vallverdú, J.; Saladie, P. (May 2023). "A review on the Pleistocene occurrences and palaeobiology of Hippopotamus antiquus based on the record from the Barranc de la Boella Section (Francolí Basin, NE Iberia)". Quaternary Science Reviews. 307: 108034. Bibcode:2023QSRv..30708034F. doi:10.1016/j.quascirev.2023.108034. hdl:10261/308555.
^ Fidalgo, Darío; Madurell-Malapeira, Joan; Martino, Roberta; Pandolfi, Luca; Rosas, Antonio (2024-01-10). "An Updated Review of The Quaternary Hippopotamus Fossil Records from the Iberian Peninsula". Quaternary. 7 (1): 4. doi:10.3390/quat7010004. ISSN 2571-550X.
^ ^a ^b ^c ^d Adams, Neil F.; Candy, Ian; Schreve, Danielle C. (2022). "An Early Pleistocene hippopotamus from Westbury Cave, Somerset, England: support for a previously unrecognized temperate interval in the British Quaternary record". Journal of Quaternary Science. 37 (1): 28–41. Bibcode:2022JQS....37...28A. doi:10.1002/jqs.3375. ISSN 1099-1417. S2CID 244179438.
^ ^a ^b ^c Fidalgo, D.; Rosas, A.; Madurell-Malapeira, J.; Pineda, A.; Huguet, R.; García-Tabernero, A.; Cáceres, I.; Ollé, A.; Vallverdú, J.; Saladie, P. (May 2023). "A review on the Pleistocene occurrences and palaeobiology of Hippopotamus antiquus based on the record from the Barranc de la Boella Section (Francolí Basin, NE Iberia)". Quaternary Science Reviews. 307: 108034. Bibcode:2023QSRv..30708034F. doi:10.1016/j.quascirev.2023.108034. hdl:10261/308555.
^ Mecozzi, Beniamino; Iannucci, Alessio; Mancini, Marco; Tentori, Daniel; Cavasinni, Chiara; Conti, Jacopo; Messina, Mattia Yuri; Sarra, Alex; Sardella, Raffaele (2023-11-22). Carnevale, Giorgio (ed.). "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)". PLOS ONE. 18 (11): e0293405. Bibcode:2023PLoSO..1893405M. doi:10.1371/journal.pone.0293405. ISSN 1932-6203. PMC 10664965. PMID 37992018.
^ van Kolfschoten, Th. (2000). "The Eemian mammal fauna of central Europe" (PDF). Netherlands Journal of Geosciences. 79 (2/3): 269–281. Bibcode:2000NJGeo..79..269V. doi:10.1017/S0016774600021752. S2CID 234349144. Archived from the original (PDF) on 2011-07-24.
^ "150 Years of Neanderthal Discoveries; Early Europeans - Continuity & Discontinuity," ed. von Koenigswald, Wighart and Thomas Litt, TERRA NOSTRA 2006/2 University of Bonn, in PDF Archived 2007-02-25 at the Wayback Machine
^ Martínez-Navarro, Bienvenido (2010), Fleagle, John G.; Shea, John J.; Grine, Frederick E.; Baden, Andrea L. (eds.), "Early Pleistocene Faunas of Eurasia and Hominin Dispersals", Out of Africa I, Vertebrate Paleobiology and Paleoanthropology, Dordrecht: Springer Netherlands, pp. 207–224, doi:10.1007/978-90-481-9036-2_13, ISBN 978-90-481-9035-5, retrieved 2023-08-01
^ Martino, R.; Pandolfi, L. (2022-07-03). "The Quaternary Hippopotamus records from Italy". Historical Biology. 34 (7): 1146–1156. Bibcode:2022HBio...34.1146M. doi:10.1080/08912963.2021.1965138. ISSN 0891-2963. S2CID 239713930.
^ Palmqvist, P.; Perez-Claros, J. A.; Janis, C. M.; Figueirido, B.; Torregrosa, V.; Grocke, D. R. (2008-11-01). "Biogeochemical and Ecomorphological Inferences On Prey Selection and Resource Partitioning Among Mammalian Carnivores In An Early Pleistocene Community". PALAIOS. 23 (11): 724–737. Bibcode:2008Palai..23..724P. doi:10.2110/palo.2007.p07-073r. ISSN 0883-1351. S2CID 85577791.
^ A.C. Marra Pleistocene hippopotamuses of Mediterranean islands: looking for ancestors J.A. Alcover, P. Bover (Eds.), Proceedings of the International Symposium “Insular Vertebrate Evolution: The Palaeontological Approach.” Monographias de la Societat d'Historia Natural de !es Balears (2005), pp. 193-204
^ Georgitsis, Michail K.; Liakopoulou, Dionysia Ε.; Theodorou, Georgios E.; Tsiolakis, Efthymios (August 2022). "Functional morphology of the hindlimb of fossilized pygmy hippopotamus from Ayia Napa (Cyprus)". Journal of Morphology. 283 (8): 1048–1079. doi:10.1002/jmor.21488. ISSN 0362-2525. PMID 35708268. S2CID 249709335.
^ Konidaris, George; Tourloukis, Vangelis; Boni, Georgia; Athanassiou, Athanassios; Giusti, Domenico; Thompson, Nicholas; Syrides, George; Panagopoulou, Eleni; Karkanas, Panagiotis; Harvati, Katerina (2023-04-27). "Marathousa 2: A New Middle Pleistocene Locality in the Megalopolis Basin (Greece) With Evidence of Hominin Exploitation of Megafauna (Hippopotamus)". PaleoAnthropology. 2023 (1): 34–55. doi:10.48738/2023.iss1.810. ISSN 1545-0031.

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[1] Desmarest, A.G., 1822. Mammalogie ou description des espèces de mammifères. Mme Veuve Agasse imprimeur édit., Paris, 2ème part., pp.277-555.

[:1-2] Martino, Roberta; Ríos, Maria Ibanez; Mateus, Octavio; Pandolfi, Luca (December 2022). "Taxonomy, chronology, and dispersal patterns of Western European Quaternary hippopotamuses: New insight from Portuguese fossil material". Quaternary International. 674–675: 121–137. doi:10.1016/j.quaint.2022.12.010. S2CID 255029640.

[3] Fidalgo, D.; Rosas, A.; Madurell-Malapeira, J.; Pineda, A.; Huguet, R.; García-Tabernero, A.; Cáceres, I.; Ollé, A.; Vallverdú, J.; Saladie, P. (May 2023). "A review on the Pleistocene occurrences and palaeobiology of Hippopotamus antiquus based on the record from the Barranc de la Boella Section (Francolí Basin, NE Iberia)". Quaternary Science Reviews. 307: 108034. Bibcode:2023QSRv..30708034F. doi:10.1016/j.quascirev.2023.108034. hdl:10261/308555.

[4] Fidalgo, Darío; Madurell-Malapeira, Joan; Martino, Roberta; Pandolfi, Luca; Rosas, Antonio (2024-01-10). "An Updated Review of The Quaternary Hippopotamus Fossil Records from the Iberian Peninsula". Quaternary. 7 (1): 4. doi:10.3390/quat7010004. ISSN 2571-550X.

[:0-5] Adams, Neil F.; Candy, Ian; Schreve, Danielle C. (2022). "An Early Pleistocene hippopotamus from Westbury Cave, Somerset, England: support for a previously unrecognized temperate interval in the British Quaternary record". Journal of Quaternary Science. 37 (1): 28–41. Bibcode:2022JQS....37...28A. doi:10.1002/jqs.3375. ISSN 1099-1417. S2CID 244179438.

[:2-6] Fidalgo, D.; Rosas, A.; Madurell-Malapeira, J.; Pineda, A.; Huguet, R.; García-Tabernero, A.; Cáceres, I.; Ollé, A.; Vallverdú, J.; Saladie, P. (May 2023). "A review on the Pleistocene occurrences and palaeobiology of Hippopotamus antiquus based on the record from the Barranc de la Boella Section (Francolí Basin, NE Iberia)". Quaternary Science Reviews. 307: 108034. Bibcode:2023QSRv..30708034F. doi:10.1016/j.quascirev.2023.108034. hdl:10261/308555.

[7] Mecozzi, Beniamino; Iannucci, Alessio; Mancini, Marco; Tentori, Daniel; Cavasinni, Chiara; Conti, Jacopo; Messina, Mattia Yuri; Sarra, Alex; Sardella, Raffaele (2023-11-22). Carnevale, Giorgio (ed.). "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)". PLOS ONE. 18 (11): e0293405. Bibcode:2023PLoSO..1893405M. doi:10.1371/journal.pone.0293405. ISSN 1932-6203. PMC 10664965. PMID 37992018.

[Kolfschoten2000-8] van Kolfschoten, Th. (2000). "The Eemian mammal fauna of central Europe" (PDF). Netherlands Journal of Geosciences. 79 (2/3): 269–281. Bibcode:2000NJGeo..79..269V. doi:10.1017/S0016774600021752. S2CID 234349144. Archived from the original (PDF) on 2011-07-24.

[Neanderthals-9] "150 Years of Neanderthal Discoveries; Early Europeans - Continuity & Discontinuity," ed. von Koenigswald, Wighart and Thomas Litt, TERRA NOSTRA 2006/2 University of Bonn, in PDF Archived 2007-02-25 at the Wayback Machine

[10] Martínez-Navarro, Bienvenido (2010), Fleagle, John G.; Shea, John J.; Grine, Frederick E.; Baden, Andrea L. (eds.), "Early Pleistocene Faunas of Eurasia and Hominin Dispersals", Out of Africa I, Vertebrate Paleobiology and Paleoanthropology, Dordrecht: Springer Netherlands, pp. 207–224, doi:10.1007/978-90-481-9036-2_13, ISBN 978-90-481-9035-5, retrieved 2023-08-01

[:02-11] Martino, R.; Pandolfi, L. (2022-07-03). "The Quaternary Hippopotamus records from Italy". Historical Biology. 34 (7): 1146–1156. Bibcode:2022HBio...34.1146M. doi:10.1080/08912963.2021.1965138. ISSN 0891-2963. S2CID 239713930.

[12] Palmqvist, P.; Perez-Claros, J. A.; Janis, C. M.; Figueirido, B.; Torregrosa, V.; Grocke, D. R. (2008-11-01). "Biogeochemical and Ecomorphological Inferences On Prey Selection and Resource Partitioning Among Mammalian Carnivores In An Early Pleistocene Community". PALAIOS. 23 (11): 724–737. Bibcode:2008Palai..23..724P. doi:10.2110/palo.2007.p07-073r. ISSN 0883-1351. S2CID 85577791.

[13] A.C. Marra Pleistocene hippopotamuses of Mediterranean islands: looking for ancestors J.A. Alcover, P. Bover (Eds.), Proceedings of the International Symposium “Insular Vertebrate Evolution: The Palaeontological Approach.” Monographias de la Societat d'Historia Natural de !es Balears (2005), pp. 193-204

[14] Georgitsis, Michail K.; Liakopoulou, Dionysia Ε.; Theodorou, Georgios E.; Tsiolakis, Efthymios (August 2022). "Functional morphology of the hindlimb of fossilized pygmy hippopotamus from Ayia Napa (Cyprus)". Journal of Morphology. 283 (8): 1048–1079. doi:10.1002/jmor.21488. ISSN 0362-2525. PMID 35708268. S2CID 249709335.

[15] Konidaris, George; Tourloukis, Vangelis; Boni, Georgia; Athanassiou, Athanassios; Giusti, Domenico; Thompson, Nicholas; Syrides, George; Panagopoulou, Eleni; Karkanas, Panagiotis; Harvati, Katerina (2023-04-27). "Marathousa 2: A New Middle Pleistocene Locality in the Megalopolis Basin (Greece) With Evidence of Hominin Exploitation of Megafauna (Hippopotamus)". PaleoAnthropology. 2023 (1): 34–55. doi:10.48738/2023.iss1.810. ISSN 1545-0031.

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 == Chronology ==
-''H. antiquus'' is suggested to be closely related to the African species ''[[Hippopotamus gorgops]]'', and may be a descendant of that species.<ref name=":1">{{Cite journal |last1=Martino |first1=Roberta |last2=Ríos |first2=Maria Ibanez |last3=Mateus |first3=Octavio |last4=Pandolfi |first4=Luca |date=December 2022 |title=Taxonomy, chronology, and dispersal patterns of Western European Quaternary hippopotamuses: New insight from Portuguese fossil material |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618222003901 |journal=Quaternary International |language=en |pages=121-137|doi=10.1016/j.quaint.2022.12.010}}</ref> The oldest records of ''H. antiquus'' date to the [[Early Pleistocene]], around 2.1-2 million years ago, which are found in Italy and Greece.<ref>{{Cite journal |last1=Fidalgo |first1=D. |last2=Rosas |first2=A. |last3=Madurell-Malapeira |first3=J. |last4=Pineda |first4=A. |last5=Huguet |first5=R. |last6=García-Tabernero |first6=A. |last7=Cáceres |first7=I. |last8=Ollé |first8=A. |last9=Vallverdú |first9=J. |last10=Saladie |first10=P. |date=May 2023 |title=A review on the Pleistocene occurrences and palaeobiology of Hippopotamus antiquus based on the record from the Barranc de la Boella Section (Francolí Basin, NE Iberia) |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379123000823 |journal=Quaternary Science Reviews |language=en |volume=307 |pages=108034 |doi=10.1016/j.quascirev.2023.108034|doi-access=free |hdl=10261/308555 |hdl-access=free }}</ref> The earliest specimens in the Iberian Peninsula date to around 1.7 million years ago.<ref>{{Cite journal |last=Fidalgo |first=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 |url=https://www.mdpi.com/2571-550X/7/1/4 |journal=Quaternary |language=en |volume=7 |issue=1 |pages=4 |doi=10.3390/quat7010004 |issn=2571-550X}}</ref> ''H. antiquus'' first became widespread north of the Alps around 1.1 to 1 million years ago, as evidenced from specimens found dating to this time in France, Germany, the Netherlands and southern Britain.<ref name=":0" /> The youngest remains of the species are from [[Condeixa]] in Portugal, suggested to date to approximately 400,000 years ago.<ref name=":1" /> Later records of the genus ''[[Hippopotamus (genus)|Hippopotamus]]'' in Europe are believed to belong to the modern [[hippopotamus]] (''Hippopotamus amphibius'').<ref name=":0">{{Cite journal|last1=Adams|first1=Neil F.|last2=Candy|first2=Ian|last3=Schreve|first3=Danielle C.|title=An Early Pleistocene hippopotamus from Westbury Cave, Somerset, England: support for a previously unrecognized temperate interval in the British Quaternary record|url=https://onlinelibrary.wiley.com/doi/abs/10.1002/jqs.3375|journal=Journal of Quaternary Science|year=2022|volume=37|pages=28–41|language=en|doi=10.1002/jqs.3375|bibcode=2022JQS....37...28A|s2cid=244179438|issn=1099-1417|doi-access=free}}</ref> The earliest generally accepted record of ''H. amphibius'' in Europe is around 500,000 years old, and it is therefore possible that ''H. antiquus'' and ''H. amphibius'' coexisted in Europe from 500-400,000 years ago, though this is uncertain.<ref name=":2">{{Cite journal |last1=Fidalgo |first1=D. |last2=Rosas |first2=A. |last3=Madurell-Malapeira |first3=J. |last4=Pineda |first4=A. |last5=Huguet |first5=R. |last6=García-Tabernero |first6=A. |last7=Cáceres |first7=I. |last8=Ollé |first8=A. |last9=Vallverdú |first9=J. |last10=Saladie |first10=P. |date=May 2023 |title=A review on the Pleistocene occurrences and palaeobiology of Hippopotamus antiquus based on the record from the Barranc de la Boella Section (Francolí Basin, NE Iberia) |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379123000823 |journal=Quaternary Science Reviews |language=en |volume=307 |pages=108034 |doi=10.1016/j.quascirev.2023.108034|doi-access=free |hdl=10261/308555 |hdl-access=free }}</ref><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 }}</ref>
+''H. antiquus'' is suggested to be closely related to the African species ''[[Hippopotamus gorgops]]'', and may be a descendant of that species.<ref name=":1">{{Cite journal |last1=Martino |first1=Roberta |last2=Ríos |first2=Maria Ibanez |last3=Mateus |first3=Octavio |last4=Pandolfi |first4=Luca |date=December 2022 |title=Taxonomy, chronology, and dispersal patterns of Western European Quaternary hippopotamuses: New insight from Portuguese fossil material |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618222003901 |journal=Quaternary International |volume=674-675 |language=en |pages=121–137|doi=10.1016/j.quaint.2022.12.010|s2cid=255029640 }}</ref> The oldest records of ''H. antiquus'' date to the [[Early Pleistocene]], around 2.1-2 million years ago, which are found in Italy and Greece.<ref>{{Cite journal |last1=Fidalgo |first1=D. |last2=Rosas |first2=A. |last3=Madurell-Malapeira |first3=J. |last4=Pineda |first4=A. |last5=Huguet |first5=R. |last6=García-Tabernero |first6=A. |last7=Cáceres |first7=I. |last8=Ollé |first8=A. |last9=Vallverdú |first9=J. |last10=Saladie |first10=P. |date=May 2023 |title=A review on the Pleistocene occurrences and palaeobiology of Hippopotamus antiquus based on the record from the Barranc de la Boella Section (Francolí Basin, NE Iberia) |journal=Quaternary Science Reviews |language=en |volume=307 |pages=108034 |doi=10.1016/j.quascirev.2023.108034|doi-access=free |bibcode=2023QSRv..30708034F |hdl=10261/308555 |hdl-access=free }}</ref> The earliest specimens in the Iberian Peninsula date to around 1.7 million years ago.<ref>{{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}}</ref> ''H. antiquus'' first became widespread north of the Alps around 1.1 to 1 million years ago, as evidenced from specimens found dating to this time in France, Germany, the Netherlands and southern Britain.<ref name=":0" /> The youngest remains of the species are from [[Condeixa]] in Portugal, suggested to date to approximately 400,000 years ago.<ref name=":1" /> Later records of the genus ''[[Hippopotamus (genus)|Hippopotamus]]'' in Europe are believed to belong to the modern [[hippopotamus]] (''Hippopotamus amphibius'').<ref name=":0">{{Cite journal|last1=Adams|first1=Neil F.|last2=Candy|first2=Ian|last3=Schreve|first3=Danielle 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|language=en|doi=10.1002/jqs.3375|bibcode=2022JQS....37...28A|s2cid=244179438|issn=1099-1417|doi-access=free}}</ref> The earliest generally accepted record of ''H. amphibius'' in Europe is around 500,000 years old, and it is therefore possible that ''H. antiquus'' and ''H. amphibius'' coexisted in Europe from 500-400,000 years ago, though this is uncertain.<ref name=":2">{{Cite journal |last1=Fidalgo |first1=D. |last2=Rosas |first2=A. |last3=Madurell-Malapeira |first3=J. |last4=Pineda |first4=A. |last5=Huguet |first5=R. |last6=García-Tabernero |first6=A. |last7=Cáceres |first7=I. |last8=Ollé |first8=A. |last9=Vallverdú |first9=J. |last10=Saladie |first10=P. |date=May 2023 |title=A review on the Pleistocene occurrences and palaeobiology of Hippopotamus antiquus based on the record from the Barranc de la Boella Section (Francolí Basin, NE Iberia) |journal=Quaternary Science Reviews |language=en |volume=307 |pages=108034 |doi=10.1016/j.quascirev.2023.108034|doi-access=free |bibcode=2023QSRv..30708034F |hdl=10261/308555 |hdl-access=free }}</ref><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>
 == Distribution ==
 [[File:Hippopotamus antiquus 2.JPG|left|thumb|Alternate angle of Florence specimen]]
-''H. antiquus'' ranged across Europe, from the [[Iberian Peninsula]] and the [[Italian Peninsula]], to the [[British Isles]] to the [[Rhine|Rhine River]] to Greece.<ref name="Kolfschoten2000">{{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 |doi=10.1017/S0016774600021752 |s2cid=234349144 |url=http://www.njgonline.nl/publish/articles/000099/article.pdf |archive-url=https://web.archive.org/web/20110724162555/http://www.njgonline.nl/publish/articles/000099/article.pdf |archive-date=2011-07-24 |url-status=dead }}</ref><ref name="Neanderthals">"150 Years of Neanderthal Discoveries; Early Europeans - Continuity & Discontinuity," ed. von Koenigswald, Wighart and Thomas Litt, TERRA NOSTRA 2006/2 [[University of Bonn]], [http://www.neandertal.uni-bonn.de/kongress_bonn/terra%20nostra08-3-8-06.pdf in PDF] {{webarchive|url=https://web.archive.org/web/20070225183257/http://www.neandertal.uni-bonn.de/kongress_bonn/terra%20nostra08-3-8-06.pdf |date=2007-02-25 }}</ref> The easternmost record of the species is from the [[Caucasus]], at the Akhalkalaki site in [[Georgia (country)|Georgia]].<ref name=":2" /> Remains possibly attributable to the species are also known from the [[Ubeidiya prehistoric site|Ubeidiya site]] in Israel,<ref name=":1" /> though other authors assign these remains to the species ''[[Hippopotamus behemoth|H. behemoth]]''.<ref name=":2" /> Their distribution was strongly controlled by temperature, with the species only extending to the northern parts of Europe during warmer [[interglacial]] intervals.<ref name=":0" />
+''H. antiquus'' ranged across Europe, from the [[Iberian Peninsula]] and the [[Italian Peninsula]], to the [[British Isles]] to the [[Rhine|Rhine River]] to Greece.<ref name="Kolfschoten2000">{{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 |doi=10.1017/S0016774600021752 |bibcode=2000NJGeo..79..269V |s2cid=234349144 |url=http://www.njgonline.nl/publish/articles/000099/article.pdf |archive-url=https://web.archive.org/web/20110724162555/http://www.njgonline.nl/publish/articles/000099/article.pdf |archive-date=2011-07-24 |url-status=dead }}</ref><ref name="Neanderthals">"150 Years of Neanderthal Discoveries; Early Europeans - Continuity & Discontinuity," ed. von Koenigswald, Wighart and Thomas Litt, TERRA NOSTRA 2006/2 [[University of Bonn]], [http://www.neandertal.uni-bonn.de/kongress_bonn/terra%20nostra08-3-8-06.pdf in PDF] {{webarchive|url=https://web.archive.org/web/20070225183257/http://www.neandertal.uni-bonn.de/kongress_bonn/terra%20nostra08-3-8-06.pdf |date=2007-02-25 }}</ref> The easternmost record of the species is from the [[Caucasus]], at the Akhalkalaki site in [[Georgia (country)|Georgia]].<ref name=":2" /> Remains possibly attributable to the species are also known from the [[Ubeidiya prehistoric site|Ubeidiya site]] in Israel,<ref name=":1" /> though other authors assign these remains to the species ''[[Hippopotamus behemoth|H. behemoth]]''.<ref name=":2" /> Their distribution was strongly controlled by temperature, with the species only extending to the northern parts of Europe during warmer [[interglacial]] intervals.<ref name=":0" />
 == Description ==
-Based on the transverse diameter of its limbs, ''H. antiquus'' has been estimated to weigh {{Convert|3,500–4,200|kg|lb}}, more than double the weight of the average specimen of ''H. amphibius''.<ref>{{Citation |last=Martínez-Navarro |first=Bienvenido |title=Early Pleistocene Faunas of Eurasia and Hominin Dispersals |date=2010 |url=http://link.springer.com/10.1007/978-90-481-9036-2_13 |work=Out of Africa I |series=Vertebrate Paleobiology and Paleoanthropology |pages=207–224 |editor-last=Fleagle |editor-first=John G. |access-date=2023-08-01 |place=Dordrecht |publisher=Springer Netherlands |doi=10.1007/978-90-481-9036-2_13 |isbn=978-90-481-9035-5 |editor2-last=Shea |editor2-first=John J. |editor3-last=Grine |editor3-first=Frederick E. |editor4-last=Baden |editor4-first=Andrea L.}}</ref> In comparison to modern ''Hippopotamus amphibius'', the skull is more slender and elongate, but with a shorter [[neurocranium]].<ref name=":02">{{Cite journal |last1=Martino |first1=R. |last2=Pandolfi |first2=L. |date=2022-07-03 |title=The Quaternary Hippopotamus records from Italy |journal=Historical Biology |language=en |volume=34 |issue=7 |pages=1146–1156 |doi=10.1080/08912963.2021.1965138 |issn=0891-2963 |s2cid=239713930}}</ref> ''Hippopotamus antiquus'' has been suggested to have been more aquatically adapted than ''Hippopotamus amphibius'', with the skull having more elevated eyesockets and the feet having shorter [[metapodial]] bones than ''H. amphibius.'' An [[Isotopic signature#Nitrogen isotopes|analysis of nitrogen isotopes]] suggests that ''H. antiquus'' preferred aquatic plants, in contrast to modern ''H. amphibius'', which prefers terrestrial grasses.<ref name=":0"/><ref>{{Cite journal|last1=Palmqvist|first1=P.|last2=Perez-Claros|first2=J. A.|last3=Janis|first3=C. M.|last4=Figueirido|first4=B.|last5=Torregrosa|first5=V.|last6=Grocke|first6=D. R.|date=2008-11-01|title=Biogeochemical and Ecomorphological Inferences On Prey Selection and Resource Partitioning Among Mammalian Carnivores In An Early Pleistocene Community|url=https://pubs.geoscienceworld.org/palaios/article/23/11/724-737/145947|journal=PALAIOS|language=en|volume=23|issue=11|pages=724–737|doi=10.2110/palo.2007.p07-073r|bibcode=2008Palai..23..724P|s2cid=85577791|issn=0883-1351}}</ref>
+Based on the transverse diameter of its limbs, ''H. antiquus'' has been estimated to weigh {{Convert|3,500–4,200|kg|lb}}, more than double the weight of the average specimen of ''H. amphibius''.<ref>{{Citation |last=Martínez-Navarro |first=Bienvenido |title=Early Pleistocene Faunas of Eurasia and Hominin Dispersals |date=2010 |url=http://link.springer.com/10.1007/978-90-481-9036-2_13 |work=Out of Africa I |series=Vertebrate Paleobiology and Paleoanthropology |pages=207–224 |editor-last=Fleagle |editor-first=John G. |access-date=2023-08-01 |place=Dordrecht |publisher=Springer Netherlands |doi=10.1007/978-90-481-9036-2_13 |isbn=978-90-481-9035-5 |editor2-last=Shea |editor2-first=John J. |editor3-last=Grine |editor3-first=Frederick E. |editor4-last=Baden |editor4-first=Andrea L.}}</ref> In comparison to modern ''Hippopotamus amphibius'', the skull is more slender and elongate, but with a shorter [[neurocranium]].<ref name=":02">{{Cite journal |last1=Martino |first1=R. |last2=Pandolfi |first2=L. |date=2022-07-03 |title=The Quaternary Hippopotamus records from Italy |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> ''Hippopotamus antiquus'' has been suggested to have been more aquatically adapted than ''Hippopotamus amphibius'', with the skull having more elevated eyesockets and the feet having shorter [[metapodial]] bones than ''H. amphibius.'' An [[Isotopic signature#Nitrogen isotopes|analysis of nitrogen isotopes]] suggests that ''H. antiquus'' preferred aquatic plants, in contrast to modern ''H. amphibius'', which prefers terrestrial grasses.<ref name=":0"/><ref>{{Cite journal|last1=Palmqvist|first1=P.|last2=Perez-Claros|first2=J. A.|last3=Janis|first3=C. M.|last4=Figueirido|first4=B.|last5=Torregrosa|first5=V.|last6=Grocke|first6=D. R.|date=2008-11-01|title=Biogeochemical and Ecomorphological Inferences On Prey Selection and Resource Partitioning Among Mammalian Carnivores In An Early Pleistocene Community|url=https://pubs.geoscienceworld.org/palaios/article/23/11/724-737/145947|journal=PALAIOS|language=en|volume=23|issue=11|pages=724–737|doi=10.2110/palo.2007.p07-073r|bibcode=2008Palai..23..724P|s2cid=85577791|issn=0883-1351}}</ref>
-The [[Cretan dwarf hippopotamus]] (''H. creutzburgi'') is believed to have evolved from ''H. antiquus'' through the process of [[insular dwarfism]] on the island of [[Crete]].<ref>A.C. Marra [http://ibdigital.uib.es/greenstone/sites/localsite/collect/monografiesHistoriaNatural/index/assoc/Monograf/iesSHNB_/2005vol0/12p193.dir/MonografiesSHNB_2005vol012p193.pdf Pleistocene hippopotamuses of Mediterranean islands: looking for ancestors] J.A. Alcover, P. Bover (Eds.), Proceedings of the International Symposium “Insular Vertebrate Evolution: The Palaeontological Approach.” Monographias de la Societat d'Historia Natural de !es Balears (2005), pp. 193-204</ref> The extinct [[Cyprus dwarf hippopotamus]] (''H. minor'') may also derive from ''H. antiquus'', but this is uncertain.<ref>{{Cite journal |last1=Georgitsis |first1=Michail K. |last2=Liakopoulou |first2=Dionysia Ε. |last3=Theodorou |first3=Georgios E. |last4=Tsiolakis |first4=Efthymios |date=August 2022 |title=Functional morphology of the hindlimb of fossilized pygmy hippopotamus from Ayia Napa (Cyprus) |url=https://onlinelibrary.wiley.com/doi/10.1002/jmor.21488 |journal=Journal of Morphology |language=en |volume=283 |issue=8 |pages=1048–1079 |doi=10.1002/jmor.21488 |issn=0362-2525}}</ref>
+The [[Cretan dwarf hippopotamus]] (''H. creutzburgi'') is believed to have evolved from ''H. antiquus'' through the process of [[insular dwarfism]] on the island of [[Crete]].<ref>A.C. Marra [http://ibdigital.uib.es/greenstone/sites/localsite/collect/monografiesHistoriaNatural/index/assoc/Monograf/iesSHNB_/2005vol0/12p193.dir/MonografiesSHNB_2005vol012p193.pdf Pleistocene hippopotamuses of Mediterranean islands: looking for ancestors] J.A. Alcover, P. Bover (Eds.), Proceedings of the International Symposium “Insular Vertebrate Evolution: The Palaeontological Approach.” Monographias de la Societat d'Historia Natural de !es Balears (2005), pp. 193-204</ref> The extinct [[Cyprus dwarf hippopotamus]] (''H. minor'') may also derive from ''H. antiquus'', but this is uncertain.<ref>{{Cite journal |last1=Georgitsis |first1=Michail K. |last2=Liakopoulou |first2=Dionysia Ε. |last3=Theodorou |first3=Georgios E. |last4=Tsiolakis |first4=Efthymios |date=August 2022 |title=Functional morphology of the hindlimb of fossilized pygmy hippopotamus from Ayia Napa (Cyprus) |url=https://onlinelibrary.wiley.com/doi/10.1002/jmor.21488 |journal=Journal of Morphology |language=en |volume=283 |issue=8 |pages=1048–1079 |doi=10.1002/jmor.21488 |pmid=35708268 |s2cid=249709335 |issn=0362-2525}}</ref>
 == Relationship with humans ==