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Hippidion

Hippidion (meaning little horse) is an extinct genus of equine that lived in South America from the Late Pliocene to the end of the Late Pleistocene (Lujanian), between 2.5 million and 11,000 years ago. They were one of two lineages of equines native to South America during the Pleistocene epoch, alongside Equus (Amerhippus) neogeus.

Hippidion
Temporal range: Late Pliocene-Holocene (Uquian-Lujanian)
~2.5–0.011 Ma
H. principale skeleton
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Equidae
Subfamily: Equinae
Tribe: Equini
Genus: Hippidion
Owen, 1869
Species
  • H. principale (Lund, 1846) (type)
  • H. saldiasi Roth, 1899
  • H. devillei (Gervais, 1855)
Synonyms
  • Hipphaplous Ameghino 1882
  • Hipphaplus Ameghino 1882
  • Onohippidion Filhol 1888
  • Onohippidium Moreno 1891
  • Parahipparion Ameghino 1904

Taxonomy edit

Evolution edit

Although early ancient DNA analysis studies suggested a close relationship with the wild horse, Equus ferus,[1][2] this was later shown to be incorrect, with more complete sequences finding Hippidion as an outgroup to all living equines[3] and less closely related to living equines than the North American "New World stilt legged horse", Haringtonhippus francisci. Cladogram shown below:[4]

Equini

Hippidion

Haringtonhippus

Genus Equus

Equus ferus (wild horse)

Equus ovodovi

Subgenus Asinus (asses)

Subgenus Hippotigris (zebras)

Hippidion is part of a distinct lineage of equines belonging to the tribe Equini that are suggested to have diverged from the ancestors of living equines of the genus Equus at least 6 million years ago.[4] The earliest members of the Hippidion lineage are known from the Late Miocene of North America.[5][6] Hippidion migrated into the South American continent at the Pliocene-Pleistocene transition, around 2.5 million years ago as part of the Great American Biotic Interchange[1][2] with the hippidiform lineage becoming extinct in North America during the Early Pleistocene.[5] It is disputed as to whether Hippidion is an exclusively South American genus or whether remains from North America are attributable to it.[7][6]

Hippidion is traditionally thought to have 3 species, H. principale, H. saldiasi and H. devillei, however, in a 2015 DNA analysis, the single sampled H. principale specimen was found to be nested with H. saldiasi, while H. devillei was found to be clearly genetically distinct.[3]

Description edit

 
Skeleton in Natural History Museum, London
 
Reconstruction of skull musculature with proposed prehensile lip

Specimens of Hippidion saldiasi have been estimated to weigh in the range of 252.7–326.4 kilograms (557–720 lb), while specimens of H. devillei have been estimated to weigh in the region of 227.3–366.5 kilograms (501–808 lb). H. principale was somewhat larger, with an estimated mass range of 252.7–468.6 kilograms (557–1,033 lb).[8] The skull of Hippidion is noted for its very long nasal notch separating the elongate nasal bone from the rest of the skull.[9] This structure is suggested to correspond to the presence of a prehensile upper lip.[10]

Paleobiology edit

 
Restoration of a Hippidion browsing with prehensile lips.

Isotopic evidence from δ13C suggests that Hippidion consumed plants of either C3 or mixed C3 and C4 type carbon fixation.[11] The δ13C values are consistent with the occupation of woodland and wooded grassland habitats.[12] H. principale is suggested to have been a mixed feeder (both browsing and grazing), as opposed to the grazing diet of the contemporary Equus neogeus.[13]

Distribution edit

Fossils of Hippidion have been found across South America. Hippidion principale is primarily known from the Pampas grasslands of Argentina, but its range may have extended to Peru and Ecuador. Hippidion devillei is also known from Argentinan Pampas, but is also found in the Peruvian central Andes, with remains possibly found in Venezuela. Hippidion saldiasi was native to the Southern Cone, including Patagonia and the southern Andes.[8]

Extinction edit

Hippidion became extinct alongside the other South American equines at the end of the Late Pleistocene, between 12,000 and 10,000 years Before Present (BP) as part of the Quaternary extinction event, which resulted in the extinction of most large animals in both North and South America. Climatic modelling suggests that the preferred habitat for species of Hippidion declined after the Holocene transition, but the decline is not enough to explain the extinction.[14] At the Arroyo Seco 2 site in the Argentinan Pampas (14,782–11,142 cal yr BP), and Piedra Museo site in Santa Cruz Province, Patagonia (two separate layers dating to 12,463–10,457 cal yr BP and 15,517–12,352 cal yr BP), remains of Hippidion are associated with human artifacts, including Fishtail projectile points, and bear cut marks, indicating that they were hunted by recently arrived humans, which may have played a role in their extinction.[15]

References edit

  1. ^ a b Weinstock, J.; et al. (2005). "Evolution, systematics, and phylogeography of Pleistocene horses in the New World: a molecular perspective". PLOS Biology. 3 (8): e241. doi:10.1371/journal.pbio.0030241. PMC 1159165. PMID 15974804.
  2. ^ a b Orlando, L.; et al. (2008). "Ancient DNA Clarifies the Evolutionary History of American Late Pleistocene Equids". Journal of Molecular Evolution. 66 (5): 533–538. Bibcode:2008JMolE..66..533O. doi:10.1007/s00239-008-9100-x. PMID 18398561. S2CID 19069554.
  3. ^ a b Der Sarkissian, Clio; Vilstrup, Julia T.; Schubert, Mikkel; Seguin-Orlando, Andaine; Eme, David; Weinstock, Jacobo; Alberdi, Maria Teresa; Martin, Fabiana; Lopez, Patricio M.; Prado, Jose L.; Prieto, Alfredo (March 2015). "Mitochondrial genomes reveal the extinct Hippidion as an outgroup to all living equids". Biology Letters. 11 (3): 20141058. doi:10.1098/rsbl.2014.1058. ISSN 1744-9561. PMC 4387498. PMID 25762573.
  4. ^ a b Heintzman, P.D.; Zazula, G.D.; MacPhee, R.D.E; Scott, E.; Cahill, J.A.; McHorse, B.K.; Kapp, J.D.; Stiller, M.; Wooller, M.J.; Orlando, L.; Southon, J.; Froese, D.G.; Shapiro, B. (2017). "A new genus of horse from Pleistocene North America". eLife. 6. doi:10.7554/eLife.29944. PMC 5705217. PMID 29182148.
  5. ^ a b Santos Avilla, Leonardo Dos; Bernardes, Camila; Mothé, Dimila (2015-05-04). "A new genus for Onohippidium galushai Macfadden and Skinner, 1979 (Mammalia, Equidae), from the late Hemphillian of North America". Journal of Vertebrate Paleontology. 35 (3): e925909. doi:10.1080/02724634.2014.925909. ISSN 0272-4634.
  6. ^ a b MacFadden, B. J., and M. Skinner. 1979. Diversification and biogeography of the one-toed horses Onohippidium and Hippidion. Yale Peabody Museum of Natural History Postilla 175:1–9.
  7. ^ Prado, José; Alberdi, María (2023). "Diversity of the fossil equidsfrom South America". Publicación Electrónica de la Asociación Paleontológica Argentina. doi:10.5710/PEAPA.06.06.2022.415.
  8. ^ a b Labarca, Rafael; Caro, Francisco J.; Villavicencio, Natalia A.; Capriles, José M.; Briones, Esteban; Latorre, Claudio; Santoro, Calogero M. (2020-12-10). "A partially complete skeleton of Hippidion saldiasi Roth, 1899 (Mammalia: Perissodactyla) from the late Pleistocene of the high Andes in northern Chile". Journal of Vertebrate Paleontology. 40 (6): e1862132. Bibcode:2020JVPal..40E2132L. doi:10.1080/02724634.2020.1862132. ISSN 0272-4634. S2CID 233706381.
  9. ^ Prado, José Luis; Alberdi, María Teresa (2017), "Nomenclatures and Taxonomy", Fossil Horses of South America, Cham: Springer International Publishing, pp. 7–59, doi:10.1007/978-3-319-55877-6_2, ISBN 978-3-319-55876-9, retrieved 2024-01-19
  10. ^ Bernardes, Camila; Sicuro, Fernando L.; Avilla, Leonardo S.; Pinheiro, André E.P. (2012). "Rostral reconstruction of South American hippidiforms (Mammalia, Perissodactyla, Equidae): New anatomical and ecomorphological inferences". Acta Palaeontologica Polonica. doi:10.4202/app.2011.0107.
  11. ^ Prado, José L; Sánchez, Begoña; Alberdi, María T (December 2011). "Ancient feeding ecology inferred from stable isotopic evidence from fossil horses in South America over the past 3 Ma". BMC Ecology. 11 (1): 15. Bibcode:2011BMCE...11...15P. doi:10.1186/1472-6785-11-15. ISSN 1472-6785. PMC 3129290. PMID 21672241.
  12. ^ Prado, José Luis; Alberdi, María Teresa (2017), "Ancient Feeding Ecology and Niche Differentiation of Pleistocene Horses", Fossil Horses of South America, Cham: Springer International Publishing, pp. 101–118, doi:10.1007/978-3-319-55877-6_6, ISBN 978-3-319-55876-9, retrieved 2024-01-20
  13. ^ Morosi, Elizabeth; Ubilla, Martin (December 2019). "Feeding and environmental studies on late Pleistocene horses in mid-latitudes of South America (northern Uruguay)". Quaternary Science Reviews. 225: 106025. Bibcode:2019QSRv..22506025M. doi:10.1016/j.quascirev.2019.106025. S2CID 210240458.
  14. ^ Villavicencio, Natalia A.; Corcoran, Derek; Marquet, Pablo A. (2019-06-27). "Assessing the Causes Behind the Late Quaternary Extinction of Horses in South America Using Species Distribution Models". Frontiers in Ecology and Evolution. 7: 226. doi:10.3389/fevo.2019.00226. ISSN 2296-701X.
  15. ^ Bampi, Hugo; Barberi, Maira; Lima-Ribeiro, Matheus S. (December 2022). "Megafauna kill sites in South America: A critical review". Quaternary Science Reviews. 298: 107851. Bibcode:2022QSRv..29807851B. doi:10.1016/j.quascirev.2022.107851. S2CID 253876769.

hippidion, meaning, little, horse, extinct, genus, equine, that, lived, south, america, from, late, pliocene, late, pleistocene, lujanian, between, million, years, they, were, lineages, equines, native, south, america, during, pleistocene, epoch, alongside, eq. Hippidion meaning little horse is an extinct genus of equine that lived in South America from the Late Pliocene to the end of the Late Pleistocene Lujanian between 2 5 million and 11 000 years ago They were one of two lineages of equines native to South America during the Pleistocene epoch alongside Equus Amerhippus neogeus HippidionTemporal range Late Pliocene Holocene Uquian Lujanian 2 5 0 011 Ma PreꞒ Ꞓ O S D C P T J K Pg N H principale skeletonScientific classificationDomain EukaryotaKingdom AnimaliaPhylum ChordataClass MammaliaOrder PerissodactylaFamily EquidaeSubfamily EquinaeTribe EquiniGenus HippidionOwen 1869SpeciesH principale Lund 1846 type H saldiasi Roth 1899 H devillei Gervais 1855 SynonymsHipphaplous Ameghino 1882 Hipphaplus Ameghino 1882 Onohippidion Filhol 1888 Onohippidium Moreno 1891 Parahipparion Ameghino 1904 Contents 1 Taxonomy 1 1 Evolution 2 Description 3 Paleobiology 4 Distribution 5 Extinction 6 ReferencesTaxonomy editEvolution edit Although early ancient DNA analysis studies suggested a close relationship with the wild horse Equus ferus 1 2 this was later shown to be incorrect with more complete sequences finding Hippidion as an outgroup to all living equines 3 and less closely related to living equines than the North American New World stilt legged horse Haringtonhippus francisci Cladogram shown below 4 Equini Hippidion HaringtonhippusGenus Equus Equus ferus wild horse Equus ovodoviSubgenus Asinus asses Subgenus Hippotigris zebras Hippidion is part of a distinct lineage of equines belonging to the tribe Equini that are suggested to have diverged from the ancestors of living equines of the genus Equus at least 6 million years ago 4 The earliest members of the Hippidion lineage are known from the Late Miocene of North America 5 6 Hippidion migrated into the South American continent at the Pliocene Pleistocene transition around 2 5 million years ago as part of the Great American Biotic Interchange 1 2 with the hippidiform lineage becoming extinct in North America during the Early Pleistocene 5 It is disputed as to whether Hippidion is an exclusively South American genus or whether remains from North America are attributable to it 7 6 Hippidion is traditionally thought to have 3 species H principale H saldiasi and H devillei however in a 2015 DNA analysis the single sampled H principale specimen was found to be nested with H saldiasi while H devillei was found to be clearly genetically distinct 3 Description edit nbsp Skeleton in Natural History Museum London nbsp Reconstruction of skull musculature with proposed prehensile lipSpecimens of Hippidion saldiasi have been estimated to weigh in the range of 252 7 326 4 kilograms 557 720 lb while specimens of H devillei have been estimated to weigh in the region of 227 3 366 5 kilograms 501 808 lb H principale was somewhat larger with an estimated mass range of 252 7 468 6 kilograms 557 1 033 lb 8 The skull of Hippidion is noted for its very long nasal notch separating the elongate nasal bone from the rest of the skull 9 This structure is suggested to correspond to the presence of a prehensile upper lip 10 Paleobiology edit nbsp Restoration of a Hippidion browsing with prehensile lips Isotopic evidence from d13C suggests that Hippidion consumed plants of either C3 or mixed C3 and C4 type carbon fixation 11 The d13C values are consistent with the occupation of woodland and wooded grassland habitats 12 H principale is suggested to have been a mixed feeder both browsing and grazing as opposed to the grazing diet of the contemporary Equus neogeus 13 Distribution editFossils of Hippidion have been found across South America Hippidion principale is primarily known from the Pampas grasslands of Argentina but its range may have extended to Peru and Ecuador Hippidion devillei is also known from Argentinan Pampas but is also found in the Peruvian central Andes with remains possibly found in Venezuela Hippidion saldiasi was native to the Southern Cone including Patagonia and the southern Andes 8 Extinction editHippidion became extinct alongside the other South American equines at the end of the Late Pleistocene between 12 000 and 10 000 years Before Present BP as part of the Quaternary extinction event which resulted in the extinction of most large animals in both North and South America Climatic modelling suggests that the preferred habitat for species of Hippidion declined after the Holocene transition but the decline is not enough to explain the extinction 14 At the Arroyo Seco 2 site in the Argentinan Pampas 14 782 11 142 cal yr BP and Piedra Museo site in Santa Cruz Province Patagonia two separate layers dating to 12 463 10 457 cal yr BP and 15 517 12 352 cal yr BP remains of Hippidion are associated with human artifacts including Fishtail projectile points and bear cut marks indicating that they were hunted by recently arrived humans which may have played a role in their extinction 15 References edit a b Weinstock J et al 2005 Evolution systematics and phylogeography of Pleistocene horses in the New World a molecular perspective PLOS Biology 3 8 e241 doi 10 1371 journal pbio 0030241 PMC 1159165 PMID 15974804 a b Orlando L et al 2008 Ancient DNA Clarifies the Evolutionary History of American Late Pleistocene Equids Journal of Molecular Evolution 66 5 533 538 Bibcode 2008JMolE 66 533O doi 10 1007 s00239 008 9100 x PMID 18398561 S2CID 19069554 a b Der Sarkissian Clio Vilstrup Julia T Schubert Mikkel Seguin Orlando Andaine Eme David Weinstock Jacobo Alberdi Maria Teresa Martin Fabiana Lopez Patricio M Prado Jose L Prieto Alfredo March 2015 Mitochondrial genomes reveal the extinct Hippidion as an outgroup to all living equids Biology Letters 11 3 20141058 doi 10 1098 rsbl 2014 1058 ISSN 1744 9561 PMC 4387498 PMID 25762573 a b Heintzman P D Zazula G D MacPhee R D E Scott E Cahill J A McHorse B K Kapp J D Stiller M Wooller M J Orlando L Southon J Froese D G Shapiro B 2017 A new genus of horse from Pleistocene North America eLife 6 doi 10 7554 eLife 29944 PMC 5705217 PMID 29182148 a b Santos Avilla Leonardo Dos Bernardes Camila Mothe Dimila 2015 05 04 A new genus for Onohippidium galushai Macfadden and Skinner 1979 Mammalia Equidae from the late Hemphillian of North America Journal of Vertebrate Paleontology 35 3 e925909 doi 10 1080 02724634 2014 925909 ISSN 0272 4634 a b MacFadden B J and M Skinner 1979 Diversification and biogeography of the one toed horses Onohippidium and Hippidion Yale Peabody Museum of Natural History Postilla 175 1 9 Prado Jose Alberdi Maria 2023 Diversity of the fossil equidsfrom South America Publicacion Electronica de la Asociacion Paleontologica Argentina doi 10 5710 PEAPA 06 06 2022 415 a b Labarca Rafael Caro Francisco J Villavicencio Natalia A Capriles Jose M Briones Esteban Latorre Claudio Santoro Calogero M 2020 12 10 A partially complete skeleton of Hippidion saldiasi Roth 1899 Mammalia Perissodactyla from the late Pleistocene of the high Andes in northern Chile Journal of Vertebrate Paleontology 40 6 e1862132 Bibcode 2020JVPal 40E2132L doi 10 1080 02724634 2020 1862132 ISSN 0272 4634 S2CID 233706381 Prado Jose Luis Alberdi Maria Teresa 2017 Nomenclatures and Taxonomy Fossil Horses of South America Cham Springer International Publishing pp 7 59 doi 10 1007 978 3 319 55877 6 2 ISBN 978 3 319 55876 9 retrieved 2024 01 19 Bernardes Camila Sicuro Fernando L Avilla Leonardo S Pinheiro Andre E P 2012 Rostral reconstruction of South American hippidiforms Mammalia Perissodactyla Equidae New anatomical and ecomorphological inferences Acta Palaeontologica Polonica doi 10 4202 app 2011 0107 Prado Jose L Sanchez Begona Alberdi Maria T December 2011 Ancient feeding ecology inferred from stable isotopic evidence from fossil horses in South America over the past 3 Ma BMC Ecology 11 1 15 Bibcode 2011BMCE 11 15P doi 10 1186 1472 6785 11 15 ISSN 1472 6785 PMC 3129290 PMID 21672241 Prado Jose Luis Alberdi Maria Teresa 2017 Ancient Feeding Ecology and Niche Differentiation of Pleistocene Horses Fossil Horses of South America Cham Springer International Publishing pp 101 118 doi 10 1007 978 3 319 55877 6 6 ISBN 978 3 319 55876 9 retrieved 2024 01 20 Morosi Elizabeth Ubilla Martin December 2019 Feeding and environmental studies on late Pleistocene horses in mid latitudes of South America northern Uruguay Quaternary Science Reviews 225 106025 Bibcode 2019QSRv 22506025M doi 10 1016 j quascirev 2019 106025 S2CID 210240458 Villavicencio Natalia A Corcoran Derek Marquet Pablo A 2019 06 27 Assessing the Causes Behind the Late Quaternary Extinction of Horses in South America Using Species Distribution Models Frontiers in Ecology and Evolution 7 226 doi 10 3389 fevo 2019 00226 ISSN 2296 701X Bampi Hugo Barberi Maira Lima Ribeiro Matheus S December 2022 Megafauna kill sites in South America A critical review Quaternary Science Reviews 298 107851 Bibcode 2022QSRv 29807851B doi 10 1016 j quascirev 2022 107851 S2CID 253876769 Retrieved from https en 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