Cypriot pygmy hippopotamus: Difference between revisions

Cypriot pygmy hippopotamus; Temporal range: Pleistocene to Early Holocene, 0.2–0.012 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Composite mounted skeleton of H. minor
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Artiodactyla
Family:	Hippopotamidae
Genus:	Hippopotamus
Species:	†H. minor
Binomial name
	†Hippopotamus minor; Desmarest, 1822
Synonyms
	Phanourios minor Sondaar and Boekschoten, 1972; Hippopotamus minutus Cuvier, 1824;

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The Cypriot pygmy hippopotamus (Hippopotamus minor or Phanourios minor) is an extinct species of dwarf hippopotamus that inhabited the island of Cyprus from the Pleistocene until the early Holocene. One the smallest known hippopotamus species, it was comparable in size to the living pygmy hippopotamus though it was more closely related to the common hippopotamus, with its small body size a result of insular dwarfism. It represented one of only two large terrestrial mammals on Cyprus alongside the Cyprus dwarf elephant. The species became extinct around 12,000 years ago following the arrival of humans on Cyprus, and potential evidence of human hunting has been found at the Aetokremnos rockshelter on the southern coast of the island.

History of discovery and taxonomy

Bones of fossil mammals have been known on Cyprus since at least the 15th century, when Cypriot historian Leontios Machairas reported that bones exposed in the Kyrenia/Pentadactylos mountains in the northern part of Cyprus were believed by locals to be the bones of Maronite Christians who had fled to the island, which they regarded as saints. An account from a later historian, Benedetto Bordone published in 1528, reporting on a similar deposit in the Kyrenia mountains, recounted that locals ground the bones into powder to make a potion they thought could cure many diseases. In 1698, the Dutch traveller Cornelis de Bruijn, remarking on another Kyrenia mountains bone deposit, made several images of bones he found, which he thought were deposited by the Biblical great flood. The remains in one of these images, which he identified as human, is now retrospectively identified as remains of the Cypriot pygmy hippopotamus.^[2]

The earliest scientific description of the species was given by French paleontologist Anselme Gaëtan Desmarest in 1822, who gave the current name Hippopotamus minor. The species Hippopotamus minutus named shortly after by Georges Cuvier in 1824 is now regarded as a junior synonym.^[3] Both authors were unaware of the origin of the specimens which were in the collections of a French museum in Paris, with Desmarest and Cuvier both suggesting that the specimens originated from southern France. Additional remains of the species were collected from Cyprus by British paleontologist Dorothea Bate in 1901, which led Charles Immanuel Forsyth Major to recognise material in the Paris collection as also originating from Cyprus.^[4] The species is now known from over 20 localities across the island.^[5] In 1972, the species was placed in the new genus Phanourios by Paul Yves Sondaar and Gijsbert Jan Boekschoten after Saint Phanourios which local Cypriots associated with its bones.^[6] However this placement has been questioned due to the fact that it is widely agreed that the species descends from a species of the genus Hippopotamus, and other authors have continued to use the combination Hippopotamus minor.^[7]^[8]

Evolution

A partial mitochondrial genome obtained from H. minor suggests that its closest living relative is the common hippopotamus (Hippopotamus amphibius), with an estimated genetic divergence between 1.36 to 1.58 million years ago.^[7] The ancestor of the Cypriot pygmy hippopotamus is uncertain, but is likely either H. amphibius^[7] or the extinct species Hippopotamus antiquus.^[9] The timing of the colonisation is uncertain, though the earliest fossils date to around 219-185,000 years ago, during the late Middle Pleistocene.^[7] Due to Cyprus never having been connected to the mainland, its ancestors must have arrived via crossing the Mediterranean, perhaps as the result of a rare cataclysmic flooding event.^[10] Its small body size is due to insular dwarfism, a common phenomenon on islands.^[7]

Description and ecology

The Cypriot pygmy hippopotamus is the smallest known hippopotamus species, along with the roughly same-sized living African pygmy hippopotamus (Choeropsis liberiensis). The Cypriot pygmy hippopotamus is estimated to have had a body mass of around 130 kilograms (290 lb),^[7] a height of 70 cm (2.3 ft) and a length of 125 cm (4.1 ft),^[2] an over 90% reduction in size from its mainland ancestor.^[8] The cranial cavity containing the brain is proportionally significantly larger relative to skull size than in H. amphibius.^[11] Compared to H. amphibius, the muzzle region of the skull is much shorter, resmbling the condition found in the African pygmy hippopotamus.^[8] Unlike other species of the genus Hippopotamus, the upper fourth premolar has been lost, possibly as a result of the skull shortening.^[8] The teeth of H. minor are more brachydont (less high crowned) than those of H. amphibius, suggesting that H. minor probably occupied a browsing niche, in contrast to the grazing predominant diet of modern Hippopotamus amphibius, though its diet is likely to have varied in correspondence to glacial cycle-induced climatic changes.^[12] Analysis of the limb and hand bones suggests that it was more terrestrial than its living relatives, having a unique form of locomotion distinct from modern hippopotamuses that allowed it to move efficiently on the mountainous and rocky terrain of Cyprus, with changes including the shortening of the distal (closest to foot) part of the legs, and increased robustness of the limb bones, as well as increased rigidity and stability of some of the limb joints.^[13]^[9] It probably habitually moved slowly, moreso than living hippopotamuses,^[13] and was probably incapable of running quickly.^[2]

During the Late Pleistocene, the Cypriot pygmy hippopotamus, along with the similarly sized Cyprus dwarf elephant, were the only large mammals native to the islands, and one of only four native terrestrial mammal species, alongside the still living Cypriot mouse and the extinct genet species Genetta plesictoides,^[14] and had no natural predators.^[15] Remains of the dwarf hippopotamus are abundant at localities where it is found, considerably moreso than the dwarf elephant.^[5]

Extinction

The youngest remains of the species date to the end of the Pleistocene, around 13-12,000 years ago, around the same time as the youngest remains of the dwarf elephant species. These dates roughly coincide with the oldest evidence of human habitation of Cyprus.^[16] Over 200,000 bones of H. minor, representing over 500 individuals, are associated with human artifacts at the Aetokremnos rockshelter on the southern coast of Cyprus, dating to approximately 13-12,000 years Before Present, representing among the youngest records of the species, which is suggested by some authors to provide evidence that the Cypriot pygmy hippopotamus was hunted and driven to extinction by the early human residents of Cyprus.^[17]^[18]^[19]^[20] However, these suggestions have been contested, in part due to a lack of cut marks on the bones that would definitively indicate butchery, with an alternative proposal that bones at Aetokremnos accumulated naturally over hundreds of years, with the human occupation of the site after the bones were initially deposited.^[16]^[21]

A 2024 study estimated that at the time of human arrival, the population of dwarf hippopotamus on Cyprus was around 14,300 individuals. The study suggested that harvesting over 650 dwarf hippos per year would have put the species at risk of extinction, with extinction becoming nearly certain with over 1000 individuals harvested per year (realistically accomplishable with a human population of only a few thousand people likely present on Cyprus during this time). This likely would have resulted in a relatively rapid extinction following the colonisation at Cyprus, with the author estimating a latest possible extinction date (taking into account the Signor-Lipps effect) of around 12,000-11,000 years ago.^[22]

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 van der Geer, Alexandra; Lyras, George; de Vos, John; Dermitzakis, Michael (2010-08-13). "Cyprus". Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands (1 ed.). Wiley. pp. 34–42. doi:10.1002/9781444323986.ch4. ISBN 978-1-4051-9009-1.
^ D.A. Hooijer Notes on some Pontian mammals from Sicily, figured by Seguenza Archives Néerlandaises de Zoologie, 7 (1946), pp. 301-333
^ van der Geer, Alexandra; Lyras, George; de Vos, John; Dermitzakis, Michael (2010-08-13). "Hippopotamuses and Pigs". Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands (1 ed.). Wiley. p. 344. doi:10.1002/9781444323986.ch25. ISBN 978-1-4051-9009-1.
^ ^a ^b Athanassiou, Athanassios; Herridge, Victoria; Reese, David S.; Iliopoulos, George; Roussiakis, Socrates; Mitsopoulou, Vassiliki; Tsiolakis, Efthymios; Theodorou, George (August 2015). "Cranial evidence for the presence of a second endemic elephant species on Cyprus". Quaternary International. 379: 47–57. doi:10.1016/j.quaint.2015.05.065.
^ Boekschoten G.J., Sondaar P.Y. 1972. On the fossil mammalia of Cyprus, I & II. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen (Series B), 75 (4): 306–38.
^ ^a ^b ^c ^d ^e ^f Psonis, Nikolaos; Vassou, Despoina; Nicolaou, Loucas; Roussiakis, Socrates; Iliopoulos, George; Poulakakis, Nikos; Sfenthourakis, Spyros (2022-11-02). "Mitochondrial sequences of the extinct Cypriot pygmy hippopotamus confirm its phylogenetic placement". Zoological Journal of the Linnean Society. 196 (3): 979–989. doi:10.1093/zoolinnean/zlab089. ISSN 0024-4082.
^ ^a ^b ^c ^d van der Geer, Alexandra A. E.; Lyras, George A.; Mitteroecker, Philipp; MacPhee, Ross D. E. (September 2018). "From Jumbo to Dumbo: Cranial Shape Changes in Elephants and Hippos During Phyletic Dwarfing". Evolutionary Biology. 45 (3): 303–317. doi:10.1007/s11692-018-9451-1. ISSN 0071-3260. S2CID 255346580.
^ ^a ^b 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.
^ van der Geer, Alexandra A. E.; Anastasakis, George; Lyras, George A. (April 2015). "If hippopotamuses cannot swim, how did they colonize islands: a reply to Mazza". Lethaia. 48 (2): 147–150. doi:10.1111/let.12095. hdl:10795/3305. ISSN 0024-1164.
^ Lyras, George A. (2018). "Brain Changes during Phyletic Dwarfing in Elephants and Hippos". Brain, Behavior and Evolution. 92 (3–4): 167–181. doi:10.1159/000497268. ISSN 0006-8977.
^ Bethune, Elehna; Kaiser, Thomas M.; Schulz-Kornas, Ellen; Winkler, Daniela E. (November 2019). "Multiproxy dietary trait reconstruction in Pleistocene Hippopotamidae from the Mediterranean islands". Palaeogeography, Palaeoclimatology, Palaeoecology. 533: 109210. doi:10.1016/j.palaeo.2019.05.032. S2CID 181824675.
^ ^a ^b Georgitsis, Michail K.; Liakopoulou, Dionysia Ε; Theodorou, Georgios E. (2022). "Morphofunctional examination of the carpal bones of pygmy hippopotamus from Ayia Napa, Cyprus". The Anatomical Record. 305 (2): 297–320. doi:10.1002/ar.24738. ISSN 1932-8494. PMID 34369097. S2CID 236959026.
^ Athanassiou, Athanassios; van der Geer, Alexandra A.E.; Lyras, George A. (August 2019). "Pleistocene insular Proboscidea of the Eastern Mediterranean: A review and update". Quaternary Science Reviews. 218: 306–321. doi:10.1016/j.quascirev.2019.06.028. S2CID 199107354.
^ Burness, G. P.; Diamond, J.; Flannery, T. (2001-12-04). "Dinosaurs, dragons, and dwarfs: The evolution of maximal body size". Proceedings of the National Academy of Sciences. 98 (25): 14518–14523. Bibcode:2001PNAS...9814518B. doi:10.1073/pnas.251548698. ISSN 0027-8424. JSTOR 3057309. PMC 64714. PMID 11724953.
^ ^a ^b Zazzo, Antoine; Lebon, Matthieu; Quiles, Anita; Reiche, Ina; Vigne, Jean-Denis (2015-08-18). "Direct Dating and Physico-Chemical Analyses Cast Doubts on the Coexistence of Humans and Dwarf Hippos in Cyprus". PLOS ONE. 10 (8). e0134429. Bibcode:2015PLoSO..1034429Z. doi:10.1371/journal.pone.0134429. ISSN 1932-6203. PMC 4540316. PMID 26284623.
^ Swiny, Stuart, ed. (2001). The earliest prehistory of Cyprus from colonization to exploitation (PDF). Monograph Series. Vol. 2. American Schools of Oriental Research. doi:10.2307/1357781. ISBN 0-89757-051-0. JSTOR 1357781. S2CID 161547871.
^ Simmons, A. H. (1999). Faunal extinction in an island society: pygmy hippopotamus hunters of Cyprus. Interdisciplinary Contributions to Archaeology. Kluwer Academic/Plenum Publishers. p. 382. doi:10.1007/b109876. ISBN 978-0306460883. OCLC 41712246.
^ Simmons, A. H.; Mandel, R. D. (December 2007). "Not Such a New Light: A Response to Ammerman and Noller". World Archaeology. 39 (4): 475–482. doi:10.1080/00438240701676169. JSTOR 40026143. S2CID 161791746.
^ Simmons, Alan (June 2023). "From hippos to cattle: The fragile relationship of early cypriots with their animals". Journal of Archaeological Science: Reports. 49: 103957. doi:10.1016/j.jasrep.2023.103957.
^ Nicolaou, Loucas; Iliopoulos, George; Roussiakis, Socrates (December 2020). "Population dynamics on Aetokremnos hippos of Cyprus or have Cypriots ever tasted hippo meat?". Quaternary International. 568: 55–64. Bibcode:2020QuInt.568...55N. doi:10.1016/j.quaint.2020.09.016. S2CID 224982319.
^ Bradshaw, Corey J. A.; Saltré, Frédérik; Crabtree, Stefani A.; Reepmeyer, Christian; Moutsiou, Theodora (September 2024). "Small populations of Palaeolithic humans in Cyprus hunted endemic megafauna to extinction". Proceedings of the Royal Society B: Biological Sciences. 291 (2031). doi:10.1098/rspb.2024.0967. ISSN 0962-8452.

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

[:4-2] van der Geer, Alexandra; Lyras, George; de Vos, John; Dermitzakis, Michael (2010-08-13). "Cyprus". Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands (1 ed.). Wiley. pp. 34–42. doi:10.1002/9781444323986.ch4. ISBN 978-1-4051-9009-1.

[3] D.A. Hooijer Notes on some Pontian mammals from Sicily, figured by Seguenza Archives Néerlandaises de Zoologie, 7 (1946), pp. 301-333

[4] van der Geer, Alexandra; Lyras, George; de Vos, John; Dermitzakis, Michael (2010-08-13). "Hippopotamuses and Pigs". Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands (1 ed.). Wiley. p. 344. doi:10.1002/9781444323986.ch25. ISBN 978-1-4051-9009-1.

[:5-5] Athanassiou, Athanassios; Herridge, Victoria; Reese, David S.; Iliopoulos, George; Roussiakis, Socrates; Mitsopoulou, Vassiliki; Tsiolakis, Efthymios; Theodorou, George (August 2015). "Cranial evidence for the presence of a second endemic elephant species on Cyprus". Quaternary International. 379: 47–57. doi:10.1016/j.quaint.2015.05.065.

[:0-6] Boekschoten G.J., Sondaar P.Y. 1972. On the fossil mammalia of Cyprus, I & II. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen (Series B), 75 (4): 306–38.

[:2-7] ^ ^a ^b ^c ^d ^e ^f Psonis, Nikolaos; Vassou, Despoina; Nicolaou, Loucas; Roussiakis, Socrates; Iliopoulos, George; Poulakakis, Nikos; Sfenthourakis, Spyros (2022-11-02). "Mitochondrial sequences of the extinct Cypriot pygmy hippopotamus confirm its phylogenetic placement". Zoological Journal of the Linnean Society. 196 (3): 979–989. doi:10.1093/zoolinnean/zlab089. ISSN 0024-4082.

[:1-8] van der Geer, Alexandra A. E.; Lyras, George A.; Mitteroecker, Philipp; MacPhee, Ross D. E. (September 2018). "From Jumbo to Dumbo: Cranial Shape Changes in Elephants and Hippos During Phyletic Dwarfing". Evolutionary Biology. 45 (3): 303–317. doi:10.1007/s11692-018-9451-1. ISSN 0071-3260. S2CID 255346580.

[:3-9] 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.

[10] van der Geer, Alexandra A. E.; Anastasakis, George; Lyras, George A. (April 2015). "If hippopotamuses cannot swim, how did they colonize islands: a reply to Mazza". Lethaia. 48 (2): 147–150. doi:10.1111/let.12095. hdl:10795/3305. ISSN 0024-1164.

[11] Lyras, George A. (2018). "Brain Changes during Phyletic Dwarfing in Elephants and Hippos". Brain, Behavior and Evolution. 92 (3–4): 167–181. doi:10.1159/000497268. ISSN 0006-8977.

[12] Bethune, Elehna; Kaiser, Thomas M.; Schulz-Kornas, Ellen; Winkler, Daniela E. (November 2019). "Multiproxy dietary trait reconstruction in Pleistocene Hippopotamidae from the Mediterranean islands". Palaeogeography, Palaeoclimatology, Palaeoecology. 533: 109210. doi:10.1016/j.palaeo.2019.05.032. S2CID 181824675.

[:7-13] Georgitsis, Michail K.; Liakopoulou, Dionysia Ε; Theodorou, Georgios E. (2022). "Morphofunctional examination of the carpal bones of pygmy hippopotamus from Ayia Napa, Cyprus". The Anatomical Record. 305 (2): 297–320. doi:10.1002/ar.24738. ISSN 1932-8494. PMID 34369097. S2CID 236959026.

[14] Athanassiou, Athanassios; van der Geer, Alexandra A.E.; Lyras, George A. (August 2019). "Pleistocene insular Proboscidea of the Eastern Mediterranean: A review and update". Quaternary Science Reviews. 218: 306–321. doi:10.1016/j.quascirev.2019.06.028. S2CID 199107354.

[PNAS_on_Dwards-15] Burness, G. P.; Diamond, J.; Flannery, T. (2001-12-04). "Dinosaurs, dragons, and dwarfs: The evolution of maximal body size". Proceedings of the National Academy of Sciences. 98 (25): 14518–14523. Bibcode:2001PNAS...9814518B. doi:10.1073/pnas.251548698. ISSN 0027-8424. JSTOR 3057309. PMC 64714. PMID 11724953.

[:6-16] Zazzo, Antoine; Lebon, Matthieu; Quiles, Anita; Reiche, Ina; Vigne, Jean-Denis (2015-08-18). "Direct Dating and Physico-Chemical Analyses Cast Doubts on the Coexistence of Humans and Dwarf Hippos in Cyprus". PLOS ONE. 10 (8). e0134429. Bibcode:2015PLoSO..1034429Z. doi:10.1371/journal.pone.0134429. ISSN 1932-6203. PMC 4540316. PMID 26284623.

[17] Swiny, Stuart, ed. (2001). The earliest prehistory of Cyprus from colonization to exploitation (PDF). Monograph Series. Vol. 2. American Schools of Oriental Research. doi:10.2307/1357781. ISBN 0-89757-051-0. JSTOR 1357781. S2CID 161547871.

[Simmons1-18] Simmons, A. H. (1999). Faunal extinction in an island society: pygmy hippopotamus hunters of Cyprus. Interdisciplinary Contributions to Archaeology. Kluwer Academic/Plenum Publishers. p. 382. doi:10.1007/b109876. ISBN 978-0306460883. OCLC 41712246.

[Simmons2-19] Simmons, A. H.; Mandel, R. D. (December 2007). "Not Such a New Light: A Response to Ammerman and Noller". World Archaeology. 39 (4): 475–482. doi:10.1080/00438240701676169. JSTOR 40026143. S2CID 161791746.

[20] Simmons, Alan (June 2023). "From hippos to cattle: The fragile relationship of early cypriots with their animals". Journal of Archaeological Science: Reports. 49: 103957. doi:10.1016/j.jasrep.2023.103957.

[21] Nicolaou, Loucas; Iliopoulos, George; Roussiakis, Socrates (December 2020). "Population dynamics on Aetokremnos hippos of Cyprus or have Cypriots ever tasted hippo meat?". Quaternary International. 568: 55–64. Bibcode:2020QuInt.568...55N. doi:10.1016/j.quaint.2020.09.016. S2CID 224982319.

[22] Bradshaw, Corey J. A.; Saltré, Frédérik; Crabtree, Stefani A.; Reepmeyer, Christian; Moutsiou, Theodora (September 2024). "Small populations of Palaeolithic humans in Cyprus hunted endemic megafauna to extinction". Proceedings of the Royal Society B: Biological Sciences. 291 (2031). doi:10.1098/rspb.2024.0967. ISSN 0962-8452.

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@@ Line 1: / Line 1: @@
 {{Short description|Species of mammal (fossil)}}
 {{speciesbox
-| name = Cyprus dwarf hippo
+| name = Cypriot pygmy hippopotamus
-| fossil_range = [[Pleistocene]] to Early [[Holocene]], {{fossil range|0.781|0.010}}
+| fossil_range = [[Pleistocene]] to Early [[Holocene]], {{fossil range|0.2|0.012}}
 | image = Hippo-Cyprus.JPG
 | image_caption = Composite mounted skeleton of ''H. minor''
 | extinct = yes
 | genus = Hippopotamus
-| parent_authority = [[Carl Linnaeus|Linnaeus]], 1758
+| parent_authority =
 | species = minor
 | authority = [[Anselme Gaëtan Desmarest|Desmarest]], 1822<ref>Desmarest, A.G., 1822. Mammalogie ou description des espèces de mammifères. Mme Veuve Agasse imprimeur édit., Paris, 2ème part., pp.277-555.</ref>
-| synonyms = '''''Phanourios minor''''' <small>Sondaar and Boekschoten, 1972</small>
+| synonyms = *'''''Phanourios minor''''' <small>Sondaar and Boekschoten, 1972</small>
+*'''''Hippopotamus minutus''''' <small>Cuvier, 1824</small>
 }}
-The '''Cyprus dwarf hippopotamus''' or '''Cypriot pygmy hippopotamus''' (''Hippopotamus minor'' or ''Phanourios minor'') is an extinct species of [[Hippopotamidae|hippopotamus]] that inhabited the island of [[Cyprus]] from the [[Pleistocene]] until the early [[Holocene]].
+The '''Cypriot pygmy hippopotamus''' (''Hippopotamus minor'' or ''Phanourios minor'') is an extinct species of dwarf [[Hippopotamidae|hippopotamus]] that inhabited the island of [[Cyprus]] from the [[Pleistocene]] until the early [[Holocene]]. One the smallest known hippopotamus species, it was comparable in size to the living [[pygmy hippopotamus]] though it was more closely related to the [[common hippopotamus]], with its small body size a result of [[insular dwarfism]]. It represented one of only two large terrestrial mammals on Cyprus alongside the [[Cyprus dwarf elephant]]. The species became extinct around 12,000 years ago following the arrival of humans on Cyprus, and potential evidence of human hunting has been found at the [[Aetokremnos]] rockshelter on the southern coast of the island.
+== History of discovery and taxonomy ==
-The {{convert|200|kg|lb|adj=on}} Cyprus dwarf hippo was roughly the same size as the extant [[pygmy hippopotamus]]. Unlike the modern pygmy hippo, the Cyprus dwarf became small through the process of [[insular dwarfism]]. It is estimated to have measured {{cvt|76|cm|ft|sigfig=2}} tall and {{cvt|121|cm|ft|sigfig=2}} long.<ref>{{cite news | first=Menelaos | last=Hadjicostis | title=Dwarf Hippo Fossils Found on Cyprus | date= 6 December 2007 | publisher=The Associated Press | url =http://www.ctvnews.ca/dwarf-hippo-fossils-found-on-cyprus-1.266678 | work =Fox Television | accessdate = 2007-12-06 }}</ref> [[Mitochondrial DNA]] suggests that its closest living relative is the common [[hippopotamus]], (''Hippopotamus amphibius'') with an estimated divergence between 1.36 to 1.58 million years ago.<ref>Mitochondrial sequences of the extinct Cypriot pygmy hippopotamus confirm its phylogenetic placement Nikolaos Psonis, Despoina Vassou, Loucas Nicolaou, Socrates Roussiakis, George Iliopoulos, Nikos Poulakakis, Spyros Sfenthourakis ''Zoological Journal of the Linnean Society'', {{doi|10.1093/zoolinnean/zlab089}} Published: 19 November 2021</ref>
+Bones of fossil mammals have been known on Cyprus since at least the 15th century, when Cypriot historian [[Leontios Machairas]] reported that bones exposed in the [[Kyrenia Mountains|Kyrenia/Pentadactylos mountains]] in the northern part of Cyprus were believed by locals to be the bones of [[Maronite]] Christians who had fled to the island, which they regarded as saints. An account from a later historian, [[Benedetto Bordone]] published in 1528, reporting on a similar deposit in the Kyrenia mountains, recounted that locals ground the bones into powder to make a potion they thought could cure many diseases. In 1698, the Dutch traveller [[Cornelis de Bruijn]], remarking on another Kyrenia mountains bone deposit, made several images of bones he found, which he thought were deposited by the [[Genesis flood narrative|Biblical great flood]]. The remains in one of these images, which he identified as human, is now retrospectively identified as remains of the Cypriot pygmy hippopotamus.<ref name=":4">{{Cite book |last=van der Geer |first=Alexandra |url=https://onlinelibrary.wiley.com/doi/book/10.1002/9781444323986 |title=Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands |last2=Lyras |first2=George |last3=de Vos |first3=John |last4=Dermitzakis |first4=Michael |date=2010-08-13 |publisher=Wiley |isbn=978-1-4051-9009-1 |edition=1 |pages=34–42 |chapter=Cyprus |doi=10.1002/9781444323986.ch4}}</ref>
+The earliest scientific description of the species was given by French paleontologist [[Anselme Gaëtan Desmarest]] in 1822, who gave the current name ''Hippopotamus minor''. The species ''Hippopotamus minutus'' named shortly after by [[Georges Cuvier]] in 1824 is now regarded as a [[junior synonym]].<ref>D.A. Hooijer Notes on some Pontian mammals from Sicily, figured by Seguenza ''Archives Néerlandaises de Zoologie'', 7 (1946), pp. 301-333</ref> Both authors were unaware of the origin of the specimens which were in the collections of a French museum in Paris, with Desmarest and Cuvier both suggesting that the specimens originated from southern France. Additional remains of the species were collected from Cyprus by British paleontologist [[Dorothea Bate]] in 1901, which led [[Charles Immanuel Forsyth Major]] to recognise material in the Paris collection as also originating from Cyprus.<ref>{{Cite book |last=van der Geer |first=Alexandra |url=https://onlinelibrary.wiley.com/doi/book/10.1002/9781444323986 |title=Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands |last2=Lyras |first2=George |last3=de Vos |first3=John |last4=Dermitzakis |first4=Michael |date=2010-08-13 |publisher=Wiley |isbn=978-1-4051-9009-1 |edition=1 |pages=344 |chapter=Hippopotamuses and Pigs |doi=10.1002/9781444323986.ch25}}</ref> The species is now known from over 20 localities across the island.<ref name=":5">{{Cite journal |last=Athanassiou |first=Athanassios |last2=Herridge |first2=Victoria |last3=Reese |first3=David S. |last4=Iliopoulos |first4=George |last5=Roussiakis |first5=Socrates |last6=Mitsopoulou |first6=Vassiliki |last7=Tsiolakis |first7=Efthymios |last8=Theodorou |first8=George |date=August 2015 |title=Cranial evidence for the presence of a second endemic elephant species on Cyprus |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618215005856 |journal=Quaternary International |volume=379 |pages=47–57 |doi=10.1016/j.quaint.2015.05.065}}</ref> In 1972, the species was placed in the new genus '''''Phanourios''''' by Paul Yves Sondaar and Gijsbert Jan Boekschoten after [[Saint Phanourios]] which local Cypriots associated with its bones.<ref name=":0">Boekschoten G.J., Sondaar P.Y. 1972. On the fossil mammalia of Cyprus, I & II. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen (Series B), 75 (4): 306–38.</ref> However this placement has been questioned due to the fact that it is widely agreed that the species descends from a species of the genus ''[[Hippopotamus (genus)|Hippopotamus]],'' and other authors have continued to use the combination ''Hippopotamus minor''.<ref name=":2">{{Cite journal |last=Psonis |first=Nikolaos |last2=Vassou |first2=Despoina |last3=Nicolaou |first3=Loucas |last4=Roussiakis |first4=Socrates |last5=Iliopoulos |first5=George |last6=Poulakakis |first6=Nikos |last7=Sfenthourakis |first7=Spyros |date=2022-11-02 |title=Mitochondrial sequences of the extinct Cypriot pygmy hippopotamus confirm its phylogenetic placement |url=https://academic.oup.com/zoolinnean/article/196/3/979/6432118 |journal=Zoological Journal of the Linnean Society |volume=196 |issue=3 |pages=979–989 |doi=10.1093/zoolinnean/zlab089 |issn=0024-4082}}</ref><ref name=":1">{{Cite journal |last1=van der Geer |first1=Alexandra A. E. |last2=Lyras |first2=George A. |last3=Mitteroecker |first3=Philipp |last4=MacPhee |first4=Ross D. E. |date=September 2018 |title=From Jumbo to Dumbo: Cranial Shape Changes in Elephants and Hippos During Phyletic Dwarfing |url=http://link.springer.com/10.1007/s11692-018-9451-1 |journal=Evolutionary Biology |volume=45 |issue=3 |pages=303–317 |doi=10.1007/s11692-018-9451-1 |issn=0071-3260 |s2cid=255346580}}</ref>
-''H. minor'' is the smallest hippopotamus of all known insular hippopotamuses. The extremely small size of the hippo is in favour of a Middle Pleistocene or perhaps even Early Pleistocene colonization.<ref>Van der Geer A., Lyras G., De Vos J., Dermitzakis M. 2010. [https://books.google.com/books?id=JmSsNuwMAxgC&dq=evolution+of+island+mammals+geer&pg=PP2 Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands]. Wiley-Blackwell.</ref> At the time of its extinction between 11,000 and 9,000 years ago, the Cyprus dwarf hippo, along with the similarly sized [[Cyprus dwarf elephant]], were the only large mammals native to the islands, and one of only four native terrestrial mammal species, alongside the still living [[Cypriot mouse]] and the extinct [[Genet (animal)|genet]] species ''[[Genetta plesictoides]],''<ref>{{Cite journal |last1=Athanassiou |first1=Athanassios |last2=van der Geer |first2=Alexandra A.E. |last3=Lyras |first3=George A. |date=August 2019 |title=Pleistocene insular Proboscidea of the Eastern Mediterranean: A review and update |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119300848 |journal=Quaternary Science Reviews |language=en |volume=218 |pages=306–321 |doi=10.1016/j.quascirev.2019.06.028|s2cid=199107354 }}</ref>  and had no natural predators.<ref name="PNAS on Dwards">{{cite journal
+== Evolution ==
+A partial [[mitochondrial genome]] obtained from ''H. minor'' suggests that its closest living relative is the common [[hippopotamus]] (''Hippopotamus amphibius''), with an estimated [[genetic divergence]] between 1.36 to 1.58 million years ago.<ref name=":2" /> The ancestor of the Cypriot pygmy hippopotamus is uncertain, but is likely either ''H. amphibius''<ref name=":2" /> or the extinct species ''[[Hippopotamus antiquus]]''.<ref name=":3">{{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) |journal=Journal of Morphology |volume=283 |issue=8 |pages=1048–1079 |doi=10.1002/jmor.21488 |issn=0362-2525 |pmid=35708268 |s2cid=249709335}}</ref> The timing of the colonisation is uncertain, though the earliest fossils date to around 219-185,000 years ago, during the late [[Middle Pleistocene]].<ref name=":2" /> Due to Cyprus never having been connected to the mainland, its ancestors must have arrived via crossing the Mediterranean, perhaps as the result of a rare cataclysmic flooding event.<ref>{{Cite journal |last=van der Geer |first=Alexandra A. E. |last2=Anastasakis |first2=George |last3=Lyras |first3=George A. |date=April 2015 |title=If hippopotamuses cannot swim, how did they colonize islands: a reply to Mazza |url=https://www.idunn.no/doi/10.1111/let.12095 |journal=Lethaia |volume=48 |issue=2 |pages=147–150 |doi=10.1111/let.12095 |issn=0024-1164|hdl=10795/3305 |hdl-access=free }}</ref> Its small body size is due to [[insular dwarfism]], a common phenomenon on islands.<ref name=":2" />
+== Description and ecology ==
+[[File:Cypriot pygmy hippopotamus skull and jaw.jpg|right|250px|thumb|Skull and jaw of a Cypriot dwarf hippo]]
+The Cypriot pygmy hippopotamus is the smallest known hippopotamus species, along with the roughly same-sized living African [[pygmy hippopotamus]] (''Choeropsis liberiensis''). The Cypriot pygmy hippopotamus is estimated to have had a body mass of around {{Convert|130|kg|lb}},<ref name=":2" /> a height of {{cvt|70|cm|ft|sigfig=2}} and a length of {{cvt|125|cm|ft|sigfig=2}},<ref name=":4" /> an over 90% reduction in size from its mainland ancestor.<ref name=":1" /> The cranial cavity containing the brain is proportionally significantly larger relative to skull size than in ''H. amphibius''.<ref>{{Cite journal |last=Lyras |first=George A. |date=2018 |title=Brain Changes during Phyletic Dwarfing in Elephants and Hippos |url=https://karger.com/BBE/article/doi/10.1159/000497268 |journal=Brain, Behavior and Evolution |language=en |volume=92 |issue=3-4 |pages=167–181 |doi=10.1159/000497268 |issn=0006-8977}}</ref> Compared to ''H. amphibius'', the muzzle region of the skull is much shorter, resmbling the condition found in the African pygmy hippopotamus.<ref name=":1" />  Unlike other species of the genus ''Hippopotamus,'' the upper fourth premolar has been lost, possibly as a result of the skull shortening.<ref name=":1" /> The teeth of ''H. minor'' are more brachydont (less high crowned) than those of ''H. amphibius'', suggesting that ''H. minor'' probably occupied a [[Browsing (herbivory)|browsing]] niche, in contrast to the grazing predominant diet of modern ''Hippopotamus amphibius'', though its diet is likely to have varied in correspondence to [[glacial cycle]]-induced climatic changes.<ref>{{Cite journal |last1=Bethune |first1=Elehna |last2=Kaiser |first2=Thomas M. |last3=Schulz-Kornas |first3=Ellen |last4=Winkler |first4=Daniela E. |date=November 2019 |title=Multiproxy dietary trait reconstruction in Pleistocene Hippopotamidae from the Mediterranean islands |url=https://zenodo.org/record/3248597 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |volume=533 |pages=109210 |doi=10.1016/j.palaeo.2019.05.032 |s2cid=181824675}}</ref> Analysis of the limb and hand bones suggests that it was more terrestrial than its living relatives, having a unique form of locomotion distinct from modern hippopotamuses that allowed it to move efficiently on the mountainous and rocky terrain of Cyprus, with changes including the shortening of the distal (closest to foot) part of the legs, and increased robustness of the limb bones, as well as increased rigidity and stability of some of the limb joints.<ref name=":7">{{Cite journal |last1=Georgitsis |first1=Michail K. |last2=Liakopoulou |first2=Dionysia Ε |last3=Theodorou |first3=Georgios E. |year=2022 |title=Morphofunctional examination of the carpal bones of pygmy hippopotamus from Ayia Napa, Cyprus |journal=The Anatomical Record |volume=305 |issue=2 |pages=297–320 |doi=10.1002/ar.24738 |issn=1932-8494 |pmid=34369097 |s2cid=236959026 |doi-access=free}}</ref><ref name=":3" /> It probably habitually moved slowly, moreso than living hippopotamuses,<ref name=":7" /> and was probably incapable of running quickly.<ref name=":4" />
+During the [[Late Pleistocene]], the Cypriot pygmy hippopotamus, along with the similarly sized [[Cyprus dwarf elephant]], were the only large mammals native to the islands, and one of only four native terrestrial mammal species, alongside the still living [[Cypriot mouse]] and the extinct [[Genet (animal)|genet]] species ''[[Genetta plesictoides]],''<ref>{{Cite journal |last1=Athanassiou |first1=Athanassios |last2=van der Geer |first2=Alexandra A.E. |last3=Lyras |first3=George A. |date=August 2019 |title=Pleistocene insular Proboscidea of the Eastern Mediterranean: A review and update |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119300848 |journal=Quaternary Science Reviews |volume=218 |pages=306–321 |doi=10.1016/j.quascirev.2019.06.028|s2cid=199107354 }}</ref> and had no natural predators.<ref name="PNAS on Dwards">{{cite journal
   | last = Burness | first = G. P.
   |author2=Diamond, J.
@@ Line 30: / Line 41: @@
   | bibcode = 2001PNAS...9814518B
  | doi-access = free
+ }}</ref> Remains of the dwarf hippopotamus are abundant at localities where it is found, considerably moreso than the dwarf elephant.<ref name=":5" />
- }}</ref> Analysis of the carpal bones and the bones of the hindlimbs suggests that it was more terrestrial than its living relatives, and capable of moving on the rugged terrain of Cyprus.<ref>{{Cite journal |last1=Georgitsis|first1=Michail K.|last2=Liakopoulou|first2=Dionysia Ε|last3=Theodorou |first3=Georgios E.|title=Morphofunctional examination of the carpal bones of pygmy hippopotamus from Ayia Napa, Cyprus |journal=The Anatomical Record|year=2022|volume=305|issue=2|pages=297–320|language=en |doi=10.1002/ar.24738|pmid=34369097|s2cid=236959026|issn=1932-8494}}</ref><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) |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>
+== Extinction ==
-Compared to ''H. amphibius'', the muzzle region of the skull is much shorter, and the skull as a whole resembles that of the [[pygmy hippopotamus]].<ref name=":1" /> The teeth of ''H. minor'' are more brachydont (less high crowned) than those of ''H. amphibius'', suggesting that ''H. minor'' probably occupied a [[Browsing (herbivory)|browsing]] niche, in contrast to the grazing predominant diet of modern ''Hippopotamus amphibius'', though its diet is likely to have varied in correspondence to [[glacial cycle]]-induced climatic changes.<ref>{{Cite journal |last1=Bethune |first1=Elehna |last2=Kaiser |first2=Thomas M. |last3=Schulz-Kornas |first3=Ellen |last4=Winkler |first4=Daniela E. |date=November 2019 |title=Multiproxy dietary trait reconstruction in Pleistocene Hippopotamidae from the Mediterranean islands |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=533 |pages=109210 |doi=10.1016/j.palaeo.2019.05.032|s2cid=181824675 |url=https://zenodo.org/record/3248597 }}</ref>
+The youngest remains of the species date to the end of the Pleistocene, around 13-12,000 years ago, around the same time as the youngest remains of the dwarf elephant species. These dates roughly coincide with the oldest evidence of human habitation of Cyprus.<ref name=":6" /> Over 200,000 bones of ''H. minor,'' representing over 500 individuals, are associated with human artifacts at the [[Aetokremnos]] rockshelter on the southern coast of Cyprus, dating to approximately 13-12,000 years [[Before Present]], representing among the youngest records of the species, which is suggested by some authors to provide evidence that the Cypriot pygmy hippopotamus was hunted and driven to extinction by the early human residents of Cyprus.<ref>{{cite book |title=The earliest prehistory of Cyprus from colonization to exploitation |editor-last=Swiny |editor-first=Stuart |series=Monograph Series |volume=2 |publisher=[[American Schools of Oriental Research]] |year=2001 |isbn=0-89757-051-0 |doi=10.2307/1357781 |jstor=1357781 |s2cid=161547871 |url=https://www.bu.edu/asor/pubs/books-monographs/swiny.pdf}}</ref><ref name="Simmons1">{{Cite book
-Bones of ''H. minor'' are associated with human artifacts at the [[Aetokremnos]] rockshelter on the southern coast of Cyprus, dating to approximately 12,000 years [[Before Present]], which is suggested by some authors to provide evidence that the Cyprus dwarf hippo was encountered and driven to extinction by the early human residents of Cyprus.<ref>{{cite book |title=The earliest prehistory of Cyprus from colonization to exploitation |editor-last=Swiny |editor-first=Stuart |series=Monograph Series |volume=2 |publisher=[[American Schools of Oriental Research]] |year=2001 |isbn=0-89757-051-0 |doi=10.2307/1357781 |jstor=1357781 |s2cid=161547871 |url=https://www.bu.edu/asor/pubs/books-monographs/swiny.pdf}}</ref><ref name = "Simmons1">{{Cite book
   | last = Simmons | first = A. H.
   | title = Faunal extinction in an island society: pygmy hippopotamus hunters of Cyprus | publisher = [[Kluwer Academic/Plenum Publishers]]
@@ Line 40: / Line 50: @@
   | year = 1999 | pages = 382 | url = http://google.com/books?id=hCwYwyEBXEAC&printsec=frontcover
   | doi = 10.1007/b109876 | isbn = 978-0306460883
-  | oclc = 41712246}}</ref><ref name = "Simmons2">{{Cite journal
+  | oclc = 41712246}}</ref><ref name="Simmons2">{{Cite journal
   | last = Simmons | first = A. H. |author2=Mandel, R. D.
   | title = Not Such a New Light: A Response to Ammerman and Noller | journal = [[World Archaeology]]
   | volume = 39 | issue = 4 | pages = 475–482 | date = December 2007
   | jstor = 40026143
-  | doi = 10.1080/00438240701676169| s2cid = 161791746 }}</ref> However, these suggestions have been contested, with an alternative proposal that bones at Aetokremnos accumulated naturally over hundreds of years, with the human occupation of the site after the bones were initially deposited.<ref>{{Cite journal|last1=Zazzo |first1=Antoine|last2=Lebon|first2=Matthieu|last3=Quiles|first3=Anita|last4=Reiche|first4=Ina|last5=Vigne |first5=Jean-Denis|date=2015-08-18 |title=Direct Dating and Physico-Chemical Analyses Cast Doubts on the Coexistence of Humans and Dwarf Hippos in Cyprus|journal=PLOS ONE|language=en|volume=10|issue=8 |at=e0134429|doi=10.1371/journal.pone.0134429|issn=1932-6203|pmc=4540316|pmid=26284623|doi-access=free |bibcode=2015PLoSO..1034429Z}}</ref><ref>{{Cite journal|last1=Nicolaou|first1=Loucas|last2=Iliopoulos |first2=George|last3=Roussiakis|first3=Socrates|date=December 2020 |title=Population dynamics on Aetokremnos hippos of Cyprus or have Cypriots ever tasted hippo meat?|journal=Quaternary International |language=en|volume=568|pages=55–64|doi=10.1016/j.quaint.2020.09.016 |bibcode=2020QuInt.568...55N |s2cid=224982319}}</ref>
+  | doi = 10.1080/00438240701676169| s2cid = 161791746 }}</ref><ref>{{Cite journal |last=Simmons |first=Alan |date=June 2023 |title=From hippos to cattle: The fragile relationship of early cypriots with their animals |url=https://linkinghub.elsevier.com/retrieve/pii/S2352409X23001323 |journal=Journal of Archaeological Science: Reports |volume=49 |pages=103957 |doi=10.1016/j.jasrep.2023.103957}}</ref> However, these suggestions have been contested, in part due to a lack of cut marks on the bones that would definitively indicate butchery, with an alternative proposal that bones at Aetokremnos accumulated naturally over hundreds of years, with the human occupation of the site after the bones were initially deposited.<ref name=":6">{{Cite journal|last1=Zazzo |first1=Antoine|last2=Lebon|first2=Matthieu|last3=Quiles|first3=Anita|last4=Reiche|first4=Ina|last5=Vigne |first5=Jean-Denis|date=2015-08-18 |title=Direct Dating and Physico-Chemical Analyses Cast Doubts on the Coexistence of Humans and Dwarf Hippos in Cyprus|journal=PLOS ONE|volume=10|issue=8 |at=e0134429|doi=10.1371/journal.pone.0134429|issn=1932-6203|pmc=4540316|pmid=26284623|doi-access=free |bibcode=2015PLoSO..1034429Z}}</ref><ref>{{Cite journal|last1=Nicolaou|first1=Loucas|last2=Iliopoulos |first2=George|last3=Roussiakis|first3=Socrates|date=December 2020 |title=Population dynamics on Aetokremnos hippos of Cyprus or have Cypriots ever tasted hippo meat?|journal=Quaternary International |volume=568|pages=55–64|doi=10.1016/j.quaint.2020.09.016 |bibcode=2020QuInt.568...55N |s2cid=224982319}}</ref>
+A 2024 study estimated that at the time of human arrival, the population of dwarf hippopotamus on Cyprus was around 14,300 individuals. The study suggested that harvesting over 650 dwarf hippos per year would have put the species at risk of extinction, with extinction becoming nearly certain with over 1000 individuals harvested per year (realistically accomplishable with a human population of only a few thousand people likely present on Cyprus during this time). This likely would have resulted in a relatively rapid extinction following the colonisation at Cyprus, with the author estimating a latest possible extinction date (taking into account the [[Signor-Lipps effect]]) of around 12,000-11,000 years ago.<ref>{{Cite journal |last=Bradshaw |first=Corey J. A. |last2=Saltré |first2=Frédérik |last3=Crabtree |first3=Stefani A. |last4=Reepmeyer |first4=Christian |last5=Moutsiou |first5=Theodora |date=September 2024 |title=Small populations of Palaeolithic humans in Cyprus hunted endemic megafauna to extinction |url=https://royalsocietypublishing.org/doi/10.1098/rspb.2024.0967 |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=291 |issue=2031 |doi=10.1098/rspb.2024.0967 |issn=0962-8452}}</ref>
-==Taxonomy==
-[[File:Cypriot pygmy hippopotamus skull and jaw.jpg|right|250px|thumb|Skull and jaw of a Cypriot dwarf hippo]]
-Many scientists maintain the name ''Phanourios minor'' for the Cypriot dwarf hippo. This generic name was given by Paul Sondaar and Bert Boekschoten in 1972,<ref name=":0">Boekschoten G.J., Sondaar P.Y. 1972. On the fossil mammalia of Cyprus, I & II. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen (Series B), 75 (4): 306–38.</ref> based on the remains from Agios Georgios, Cyprus. At the site, a chapel had been built into the fossiliferous rocks. The rock strata here are very rich in bone content (bone breccia). For centuries, as already mentioned by Bordone in the 16th century,<ref>Bordone B. 1528. Libro di Benedetto Bordone. Nel qual si ragiona di tutte l'Isole del mondo, con li lor nomi antichi & moderni, historie, favole, & modi del loro vivere. Niccolo Zoppino, Venice. In facsimile, Edizioni Aldine, Modena, 1982.</ref> villagers have gone there to collect these bones, which in their opinion are holy, because they are the petrified remains of Saint Fanourios (see also [[Phanourios (saint)]]), a Greek Orthodox saint who, according to local myth, had fled from Syria to escape his persecutors, but had been stranded on the hostile rocky coast of Cyprus. The collected bones are ground into a powder believed to have medicinal powers. To honour the local tradition and to refer to the site, Sondaar and Boekschoten named their new genus ''Phanourios'', following the Greek spelling. They gave the specific name ''minutus'', but this was later changed to ''minor'' following rules of priority.<ref name=":0" /> Other authors contend that the genus ''[[Hippopotamus (genus)|Hippopotamus]]'' should be maintained for the species, because it descends from other members of the ''Hippopotamus'' genus.<ref name=":1">{{Cite journal |last1=van der Geer |first1=Alexandra A. E. |last2=Lyras |first2=George A. |last3=Mitteroecker |first3=Philipp |last4=MacPhee |first4=Ross D. E. |date=September 2018 |title=From Jumbo to Dumbo: Cranial Shape Changes in Elephants and Hippos During Phyletic Dwarfing |url=http://link.springer.com/10.1007/s11692-018-9451-1 |journal=Evolutionary Biology |language=en |volume=45 |issue=3 |pages=303–317 |doi=10.1007/s11692-018-9451-1 |s2cid=255346580 |issn=0071-3260}}</ref>
 ==See also==
@@ Line 64: / Line 72: @@
 [[Category:Extinct hippopotamuses]]
 [[Category:Mammals described in 1822]]
-[[Category:Pleistocene even-toed ungulates]]
+[[Category:Pleistocene Artiodactyla]]
 [[Category:Extinct mammals of Europe]]
 [[Category:Holocene extinctions]]
 [[Category:Prehistoric Cyprus]]
 [[Category:Fossil taxa described in 1822]]
+[[Category:Mammals in religion]]