hyaenodon
{{Short description|Extinct genus of mammals}}
{{Automatic taxobox
| fossil_range = Early Eocene to Early Miocene (Ypresian to Burdigalian) {{fossil range|52.0|16.9}}
| image = Hyaenodon (1).jpg
| image_caption = Mounted H. sp. skeleton, Science Museum of Minnesota
| display_parents = 4
| taxon = Hyaenodon
| authority = Laizer & Parieu, 1838
| type_species = †Hyaenodon leptorhynchus
| type_species_authority = Laizer and Parieu, 1838
| subdivision_ranks = Species
| subdivision = {{center|[see classification]}}
| synonyms = {{collapsible list |bullets=true |title=synonyms of genus:
|Gobipterodon {{small|(Lavrov, 1999)}}A. V. Lavrov (1999.) [https://static.freereferats.ru/_avtoreferats/01000233253.pdf "Adaptive Radiation of Hyaenodontinae (Creodonta, Hyaenodontidae) of Asia."] in 6th Congress of the Theriological Society, Moscow, April 13–16, p. 138 [in Russian].
|Macrohyaenodon {{small|(Lavrov & Emry, 1998)}}
|Macropterodon {{small|(Lavrov, 1999)}}
|Megalopterodon {{small|(Dashzeveg, 1964)}}
|Microhyaenodon {{small|(Lavrov, 1999)}}
|Neohyaenodon {{small|(Thorpe, 1922)}}
|Prohyaenodon {{small|(Lavrov, 1999)}}
|Protohyaenodon {{small|(Stock, 1933)}}
|Pseudopterodon {{small|(Schlosser, 1887)}}
|Taxotherium {{small|(Blainville, 1841)}}
}}
{{collapsible list |bullets=true |title=synonyms of species:
|{{collapsible list |bullets=true |title=H. brachyrhynchus:
|Canis brachyrhynchus {{small|(Blainville, 1841)}}
|Hyaenodon brachyrhenchus {{small|(Lavrov, 1999)}}
|Hyaenodon cuvieri {{small|(Pictet, 1853)}}F. J. Pictet (1853.) [https://www.biodiversitylibrary.org/item/48934#page/9/mode/1up "Traité de Paléontologie."] I (2e edit.):584 p. + atlas 110 pl.
|Hyaenodon leptorhynchus {{small|(Dujardin, 1840)}}F. Dujardin (1840.) [https://gallica.bnf.fr/ark:/12148/bpt6k29690/f134.item "Note sur une tête fossile Hyaenodon trouvée au bord du Tarn, près de Rabastens."] Comptes-Rendus de l'Académie des Sciences de Paris, 10:134-135
|Hyaenodon parisiensis {{small|(Laurillard, 1845)}}Ch. Laurillard (1845.) "Hyaenodon." in: d'Orbigny: "Dictionnaire Universel d'Histoire naturelle 1-6": 767-769, Renard, Martinet édit., Paris
|Hyaenodon vulpinum {{small|(Filhol, 1877)}}
|Hyaenodon vulpinus {{small|(Gervais, 1873)}}
|Nasua parisiensis {{small|(Blainville, 1841)}}
|Pterodon brachyrhynchus {{small|(Pomel, 1846)}}A. Pomel (1846.) [https://www.biodiversitylibrary.org/item/240488#page/405/mode/1up "Note sur le Pterodon, genre voisin des Dasyures dont plusieurs espèces ont été trouvées dans les terrains tertiaires."] Bulletin de la Société Géologique de France, 4(2):385-393
|Pterodon cuvieri {{small|(Pomel, 1846)}}
|Taxotherium parisiense {{small|(Blainville, 1841)}}
}}
|{{collapsible list |bullets=true |title=H. brevirostrus:
|Hyaenodon brevirostris {{small|(Joeckel, 1997)}}R. M. Joeckel, H. W. Bond and G. W. Kabalka (1997). [https://www.jstor.org/stable/4523821 "Internal Anatomy of the Snout and Paranasal Sinuses of Hyaenodon (Mammalia, Creodonta)"] Journal of Vertebrate Paleontology, Vol. 17, No. 2, pp. 440-446
|Protohyaenodon brevirostrus {{small|(Mellett, 1977)}}
}}
|{{collapsible list |bullets=true |title=H. chunkhtensis:
|Microhyaenodon chunkhtensis {{small|(Lavrov, 1999)}}
}}
|{{collapsible list |bullets=true |title=H. crucians:
|Hyaenodon leptocephalus {{small|(Scott, 1888)}}W. B. Scott (1888). [https://www.biodiversitylibrary.org/item/224106#page/171/mode/1up "On some new and little know creodonts."] Journal of the Academy of Natural Sciences of Philadelphia 9:155-185
|Hyaenodon minutus {{small|(Douglass, 1902)}}
|Hyaenodon paucidens {{small|(Osborn & Wortman, 1894)}}H. F. Osborn and J. L. Wortman (1894). [https://digitallibrary.amnh.org/bitstream/handle/2246/1568//v2/dspace/ingest/pdfSource/bul/B006a07.pdf?sequence=1&isAllowed=y "Fossil mammals of the Lower Miocene White River beds. Collection of 1892."] Bulletin of the American Museum of Natural History 6(7):199-228
|Protohyaenodon crucians {{small|(Mellett, 1977)}}
|Prtohyaenodon crucians {{small|(Lavrov, 1999)}}
|Pseudopterodon minutus {{small|(Douglass, 1902)}}
}}
|{{collapsible list |bullets=true |title=H. dubius:
|Hyaenodon aymardi {{small|(Filhol, 1882)}}H. Filhol (1882). "Étude des Mammifères de Ronzon (Haute-Loire)." Ann. Sci. géol., Paris, 12(3):270 p., 25 pl.
}}
|{{collapsible list |bullets=true |title=H. exiguus:
|Hyaenodon exigus {{small|(Lavrov, 1999)}}
|Pterodon exiguum {{small|(Gervais, 1873)}}
}}
|{{collapsible list |bullets=true |title=H. filholi:
|Hyaenodon compressus {{small|(Filhol, 1876)}}
|Hyaenodon vulpinus {{small|(Filhol, 1876)}}
|Microhyaenodon filholi {{small|(Lavrov, 1999)}}
|Pseudopterodon ganodus {{small|(Schlosser, 1887)}}
}}
|{{collapsible list |bullets=true |title=H. gervaisi:
|Hyaenodon ambiguous
|Hyaenodon ambiguus {{small|(Martin, 1906)}}
}}
|{{collapsible list |bullets=true |title=H. gigas:
|Macropterodon zelenovi {{small|(Lavrov, 1999)}}
|Neohyaenodon gigas {{small|(Lavrov, 1999)}}
}}
|{{collapsible list |bullets=true |title=H. heberti:
|Hyaenodon arnaudi {{small|(Depéret, 1917)}}Depéret C. (1917). [https://www.biodiversitylibrary.org/item/112216#page/6/mode/1up "Monographie de la faune de mammiféres fossiles du Ludien inférieur d'Euget-les-Bains (Gard)."] Ann. Univ. Lyon (N. S.), Div. 1, 40, 1-288.
}}
|{{collapsible list |bullets=true |title=H. horridus:
|Hyaenodon cruentus {{small|(Leidy, 1853)}}
|Neohyaenodon horridus {{small|(Thorpe, 1922)}}
|Neohyaenodon semseyi {{small|(Kretzoi, 1941)}}Kretzoi, M. (1941). [http://epa.oszk.hu/01600/01635/00417/pdf/EPA01635_foldtani_kozlony_1941_71_04-06_170-176.pdf "Ausländische Säugetierfossilien der Ungarischen Museen. (1-4)"] Földtani Közlöny, vol. 71, nos. 4-6, pp. 1-6
}}
|{{collapsible list |bullets=true |title=H. incertus:
|Gobipterodon exploratus {{small|(Lavrov, 1999)}}
|Hyaenodon exploratus {{small|(Polly, 1993)}}P. D. Polly (1993.) [https://www.academia.edu/20068607/Hyaenodontidae_Creodonta_Mammalia_from_the_early_Eocene_Four_Mile_Fauna_and_their_biostratigraphic_implications "Hyaenodontidae (Creodonta, Mammalia) from the Early Eocene Four Mile Fauna and their biostratigraphic implications."] PaleoBios, Vol. 14, No. 4, pp. 1-10
|Neohyaenodon incertus {{small|(Lavrov, 1999)}}
|Pterodon exploratus {{small|(Dashzeveg, 1985)}}
}}
|{{collapsible list |bullets=true |title=H. leptorhynchus:
|Canis leptorhynchus {{small|(Blainville, 1841)}}
|Hyaenodon bavaricus {{small|(Dehm, 1935)}}R. Dehm (1935). [http://publikationen.badw.de/de/001933670/pdf/CC%20BY "Über tertiäre Spaltenfüllungen im Fränkischen und Schwäbischen Jura [On Tertiary fissure fillings in the Franconian and Swabian Jura]."] Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung, Neue Folge 29:1-86
|Hyaenodon cayluxi {{small|(Filhol, 1876)}}
|Hyaenodon martini {{small|(Depéret, 1917)}}
|Hyaenodon milloquensis {{small|(Martin, 1906)}}
|Pterodon leptorhynchus {{small|(Pomel, 1846)}}
}}
|{{collapsible list |bullets=true |title=H. macrocephalus:
|Neohyaenodon macrocephalus {{small|(Lavrov, 1999)}}
}}
|{{collapsible list |bullets=true |title=H. megaloides:
|Neohyaenodon megaloides {{small|(Mellett, 1977)}}
}}
|{{collapsible list |bullets=true |title=H. microdon:
|Microhyaenodon microdon {{small|(Lavrov, 1999)}}
|Protohyaenodon microdon {{small|(Mellett, 1977)}}
}}
|{{collapsible list |bullets=true |title=H. milvinus:
|Neohyaenodon milvinus {{small|(Lavrov, 1999)}}
}}
|{{collapsible list |bullets=true |title=H. minor:
|Hyaenodon aimi {{small|(Cooper, 1926)}}Forster Cooper, C. (1926). [https://www.tandfonline.com/doi/abs/10.1080/00222932608633530 "Hyænodon aimi, sp. n., and a note on the occurrence of Anthracotherium minus from the Headon Beds at Hordle."] Annals and Magazine of Natural History (9)18(106):370–373
|Hyaenodon hantonensis {{small|(Lydekker, 1884)}}R. Lydekker (1884.) [https://pdf.zlibcdn.com/dtoken/2271667e518b6f5cb27b077a53a36999/s0016756800185826.pdf "Notes on some fossil Carnivora and Rodentia."] Geol. Mag., London, (3)1:442-445, 2 figs.
}}
|{{collapsible list |bullets=true |title=H. mongoliensis:
|Epipterodon mongoliensis {{small|(Lavrov, 1999)}}
|Megalopterodon mongoliensis {{small|(Dashzeveg, 1964)}}
|Neohyaenodon mongoliensis {{small|(Morlo & Nagel, 2006)}}Michael Morlo, Doris Nagel (2006). [https://www.researchgate.net/publication/248545738_New_remains_of_Hyaenodontidae_Creodonta_Mammalia_from_the_Oligocene_of_Central_Mongolia "New remains of Hyaenodontidae (Creodonta, Mammalia) from the Oligocene of Central Mongolia."] Annales de Paléontologie 92(3):305-321
|Pterodon mongoliensis {{small|(Van Valen, 1967)}}L. Van Valen (1967). [https://digitallibrary.amnh.org/handle/2246/358 "New Paleocene insectivores and insectivore classification."] Bulletin of the American Museum of Natural History 135(5):217-284
}}
|{{collapsible list |bullets=true |title=H. montanus:
|Protohyaenodon montanus
|Neohyaenodon montanus {{small|(Mellett, 1977)}}
}}
|{{collapsible list |bullets=true |title=H. mustelinus:
|Hyaenodon mustilinius
|Prohyaenodon mustelinus {{small|(Lavrov, 1999)}}
|Protohyaenodon mustelinus {{small|(Scott, 1894)}}
}}
|{{collapsible list |bullets=true |title=H. pervagus:
|Hyaenodon neimongoliensis {{small|(Huang & Zhu, 2002)}}{{cite journal |last1=Huang |first1=Xue-Shi |last2=Zhu |first2=Bao-Cheng |title=Creodont (mammalia) remains from the Early Oligocene of Ulantatal, Nei Mongol |journal=Vertebrata PalAsiatica |date=15 March 2002 |volume=40 |issue=1 |pages=17 |url=http://www.vertpala.ac.cn/EN/abstract/abstract856.shtml |access-date=19 November 2023 |language=en |issn=2096-9899}}
}}
|{{collapsible list |bullets=true |title=H. pumilus:
|Microhyaenodon pumilus {{small|(Lavrov, 1999)}}
}}
|{{collapsible list |bullets=true |title=H. raineyi:
|Microhyaenodon raineyi {{small|(Lavrov, 1999)}}
|Protohyaenodon raineyi {{small|(Gustafson, 1986)}}
}}
|{{collapsible list |bullets=true |title=H. requieni:
|Hyaenodon heberti euzetensis {{small|(Depéret, 1917)}}
|Pterodon requieni {{small|(Gervais, 1846)}}
}}
|{{collapsible list |bullets=true |title=H. venturae:
|Hyaenodon exiguus {{small|(Stock, 1933)}}
|Microhyaenodon venturae {{small|(Lavrov, 1999)}}
|Protohyaenodon exiguus {{small|(Stock, 1933)}}
|Protohyaenodon venturae {{small|(Mellett, 1977)}}
}}
|{{collapsible list |bullets=true |title=H. vetus:
|Neohyaenodon vetus {{small|(Mellett, 1977)}}
|Pterodon californicus {{small|(Stock, 1933)}}
}}
|{{collapsible list |bullets=true |title=H. yuanchuensis:
|Hyaenodon yuanchüensis {{small|(Young, 1937)}}
}}
}}
| synonyms_ref = J. Alroy (2002). "Synonymies and reidentifications of North American fossil mammals."
}}
Hyaenodon ("hyena-tooth") is an extinct genus of carnivorous placental mammals from extinct tribe Hyaenodontini within extinct subfamily Hyaenodontinae (in extinct family Hyaenodontidae),Malcolm C. McKenna, Susan K. Bell (1997). [https://books.google.com/books?id=F4jknQEACAAJ "Classification of Mammals: Above the Species Level"], Columbia University Press, New York, 631 pages. {{page|year=|isbn=978-0-231-11013-6|pages=}} that lived in Eurasia, Africa, and North America from the Early Eocene to the Early Miocene.
Classification and phylogeny
File:Hyaenodon Heinrich Harder.jpeg (around 1920)]]
File:Hyaenodon and Leptomeryx.jpg by W. B. Scott (1913)]]
= Taxonomy =
Description
File:Hyaenodon horridus skull.jpg
The skull of Hyaenodon was long with a narrow snout—much larger in relation to the length of the skull than in canine carnivores, for instance. The neck was shorter than the skull, while the body was long and robust and terminated in a long tail. Compared to the larger (but not closely related) Hyainailouros, the dentition of Hyaenodon was geared more towards shearing meat and less towards bone crushing.{{Cite book |last1=Wang |first1=Xiaoming |last2=Tedford |first2=Richard H. |title=Dogs: Their Fossil Relatives and Evolutionary History |location=New York |publisher=Columbia University Press |year=2008 |isbn=978-0-231-13528-3 |page=1|doi=10.7312/wang13528 |jstor=10.7312/wang13528 }}
Some species of this genus were among the largest terrestrial carnivorous mammals of their time; others were only of the size of a marten. Remains of many species are known from North America, Europe, and Asia. H. horridus was the largest North American species. While m1 regressions suggested it could’ve weighed {{cvt|91.8|kg}},{{Cite journal |last=Christison |first=Brigid E |last2=Gaidies |first2=Fred |last3=Pineda-Munoz |first3=Silvia |last4=Evans |first4=Alistair R |last5=Gilbert |first5=Marisa A |last6=Fraser |first6=Danielle |date=2022-01-25 |editor-last=Powell |editor-first=Roger |title=Dietary niches of creodonts and carnivorans of the late Eocene Cypress Hills Formation |url=https://academic.oup.com/jmammal/article/103/1/2/6459704 |journal=Journal of Mammalogy |language=en |volume=103 |issue=1 |pages=2–17 |doi=10.1093/jmammal/gyab123 |issn=0022-2372 |pmc=8789764 |pmid=35087328 |access-date=17 October 2024 |via=Oxford Academic}} regressions based on limb morphology suggests the species instead could’ve been smaller, with adults weighing {{cvt|41.42|kg}} on average and may not have exceeded {{cvt|60|kg}}. H. gigas, the largest Hyaenodon species, was much larger, being {{cvt|378|kg}} and around {{cvt|10|ft|0}}. H. weilini was also very large, being similar in size to H. gigas and H. mongoliensis.
H. crucians from the early Oligocene of North America is estimated to only {{cvt|10|to|25|kg}}. H. microdon and H. mustelinus from the late Eocene of North America were even smaller and weighed probably about {{cvt|5|kg}}.{{cite journal |author=Egi, Naoko |year=2001 |title=Body mass estimates in extinct mammals from limb bone dimensions: the case of North American hyaenodontids. |journal=Palaeontology |volume=44 |number=3 |pages=497–528 |doi=10.1111/1475-4983.00189 |doi-access=free |bibcode=2001Palgy..44..497E |s2cid=128832577}}
Paleobiology
= Predatory behavior =
A 2003 study found that based on elbow morphology H. horridus was a cursorial predator and was the most cursorially Oligocene carnivore within the study.{{Cite journal |last=Andersson |first=Ki |last2=Werdelin |first2=Lars |date=December 2003 |title=The evolution of cursorial carnivores in the Tertiary: Implications of elbow-joint morphology |url=https://www.researchgate.net/publication/8967327_The_evolution_of_cursorial_carnivores_in_the_Tertiary_Implications_of_elbow-joint_morphology |journal=Proceeding of the Royal Society B |volume=270 |doi=10.1098/rsbl.2003.0070}} Furthermore a 2025 study found that based on elbow morphology found that H. crucians and H. horridus were a pounce-pursuit predators.Castellanos, Miguel (2024). [https://huskiecommons.lib.niu.edu/cgi/viewcontent.cgi?article=8874&context=allgraduate-thesesdissertations Hunting Types in North American Eocene and Oligocene Carnivores and Implications for Nimravid Extinction] (Graduate Research Thesis & Disserations) On the other hand, analysis on the bony labyrinth of H. exiguus suggests this species was semi-arboreal and occupied a hyena-like niche.{{Cite journal |last=Pfaff |first=Cathrin |last2=Nagel |first2=Doris |last3=Gunnell |first3=Gregg |last4=Weber |first4=Gerhard W. |last5=Kriwet |first5=Jürgen |last6=Morlo |first6=Michael |last7=Bastl |first7=Katharina |date=25 September 2016 |title=Palaeobiology of Hyaenodon exiguus (Hyaenodonta, Mammalia) based on morphometric analysis of the bony labyrinth |url=https://onlinelibrary.wiley.com/doi/10.1111/joa.12545 |journal=Journal of Anatomy |volume=230 |issue=2 |pages=282-289 |doi=10.1111/joa.12545|pmc=5244453 }} Much like H. horridus, H. eminus, H. gigas, and H. pervagus were recovered as a cursorial predators.{{Cite book |last=Morlo |first=Michael |title=Carnivoran Evolution |last2=Gunnell |first2=Gregg F. |last3=Nagel |first3=Doris |publisher=Cambridge University Press |year=2010 |isbn=9781139193436 |pages=269-310 |language=English |chapter=10 - Ecomorphological analysis of carnivore guilds in the Eocene through Miocene of Laurasia}}
Zigzag Hunter–Schreger bands in the enamel indicate that European and North America Hyaenodon had slightly different niches. Microwear patterns suggests that North American Hyaenodon diet was more similar to lions, on the other hand European Hyaenodon microwear was more similar to spotted hyenas and bone crushing was likely a part of their diet.{{Cite journal |last=Bastl |first=Katharina |last2=Semprebon |first2=Gina |last3=Nagel |first3=Doris |date=1 September 2012 |title=Low‐magnification microwear in Carnivora and dietary diversity in Hyaenodon (Mammalia: Hyaenodontidae) with additional information on its enamel microstructure |url=https://www.sciencedirect.com/science/article/pii/S003101821200301X |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=348-349 |pages=13–20 |doi=10.1016/j.palaeo.2012.05.026 |url-access=subscription |access-date=7 November 2024 |via=Elsevier Science Direct}}
=Tooth eruption=
Studies on juvenile Hyaenodon specimens show that the animal had a very unusual system of tooth replacement. Juveniles took about 3–4 years to complete the final stage of eruption, implying a long adolescent phase. In North American forms, the first upper premolar erupts before the first upper molar, while European forms show an earlier eruption of the first upper molar.{{cite journal |last=Bastl |first=Katharina Anna |year=2013 |title=First evidence of the tooth eruption sequence of the upper jaw in Hyaenodon (Hyaenodontidae, Mammalia) and new information on the ontogenetic development of its dentition |journal=Paläontologische Zeitschrift |volume=88 |issue=4 |pages=481–494 |doi=10.1007/s12542-013-0207-z |s2cid=85304920}}
= Brain anatomy =
While typically assumed that Hyaenodon had a very massive skull, but a small brain. This has been called into question as a recent study found that Hyaenodonta, including Hyaenodon, had encephalization quotient similar to basal and some modern carnivorans.{{Cite journal |last=Dubied |first=Morgane |last2=Solé |first2=Floréal |last3=Mennecart |first3=Bastien |date=29 July 2019 |title=The cranium of Proviverra typica (Mammalia, Hyaenodonta) and its impact on hyaenodont phylogeny and endocranial evolution |url=https://onlinelibrary.wiley.com/doi/10.1111/pala.12437 |journal=Paleontology |volume=62 |issue=6 |pages=983-1001 |doi=10.1111/pala.12437 |doi-access=free|url-access=subscription }} The endocast of Hyaenodon stands out from other Hyaenodontoids as they had relatively high EQ, in addition to relatively gyrencephalic and neocorticalized brains, although the increase in EQ for the genus is still unknown.{{cite journal |last1=Flink |first1=Therese |last2=Cote |first2=Susanne |last3=Rossie |first3=James B. |last4=Kibii |first4=Job M. |last5=Werdelin |first5=Lars |date=March 2021 |title=The neurocranium of Ekweeconfractus amorui gen. et sp. nov. (Hyaenodonta, Mammalia) and the evolution of the brain in some hyaenodontan carnivores |journal=Journal of Vertebrate Paleontology |volume=41 |issue=2 |pages=e1927748 |bibcode=2021JVPal..41E7748F |doi=10.1080/02724634.2021.1927748 |s2cid=237518007 |doi-access=free}}
Paleoecology
The various species of Hyaenodon competed with each other and with other hyaenodont genera (including Sinopa, Dissopsalis and Hyainailurus), and played important roles as predators in ecological communities as late as the Miocene in Asia and preyed on a variety of prey species such as early horses like Mesohippus, Brontotheres, early camels, oreodonts and even early rhinos. Species of Hyaenodon have been shown to have successfully preyed on other large carnivores of their time, including a nimravid ("false sabertooth cat"), according to analysis of tooth puncture marks on a fossil Dinictis skull found in North Dakota.John W. Hoganson and Jeff Person (2010). [https://www.dmr.nd.gov/dmr/sites/www/files/documents/paleontology/pdfs/articles-and-publications/Hoganson--Person-2010-SVP-poster.pdf "Tooth puncture marks on a skull of Dinictis (Nimravidae) from the Oligocene Brule Formation of Northe Dakota attributed to predation by Hyaenodon (Hyaenodontidae)"], North Dakota Geological SurveyJohn W. Hoganson and Jeff Person (2011). [https://www.dmr.nd.gov/ndgs/documents/newsletter/2011Summer/Toothpuncturemarks.pdf "Tooth puncture marks on a 30 million year old Dinictis skull."], Geo News, p. 12-17
In North America the last Hyaenodon, in the form of H. brevirostrus, disappeared in the late Oligocene.{{cite journal |last=Van Valkenburgh |first=Blaire |date=1994 |title=Extinction and replacement among predatory mammals in the North American late Eocene and Oligocene: Tracking a paleoguild over twelve million years |url=https://www.tandfonline.com/doi/abs/10.1080/10292389409380474 |journal=Historical Biology |volume=8 |issue=1–4 |pages=129–150 |doi=10.1080/10292389409380474 |bibcode=1994HBio....8..129V |access-date=12 April 2022|url-access=subscription }} In Europe, they had already vanished earlier in the Oligocene, with the youngest species, H. weilini, persisting in the early Miocene.
References
{{Commons category}}
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{{Pan-Carnivora|H.|state=collapsed}}
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Category:Eocene genus first appearances
Category:Miocene genus extinctions
Category:Cenozoic mammals of Asia
Category:Cenozoic mammals of North America
Category:Cenozoic mammals of Europe