Hyaenodon

Hyaenodon ("hyena-tooth") is an extinct genus of carnivorous placental mammals from extinct tribe Hyaenodontini within extinct subfamily Hyaenodontinae (in extinct family Hyaenodontidae),[19] that lived in Eurasia and North America from the middle Eocene, throughout the Oligocene, to the early Miocene.

Hyaenodon
Temporal range: Middle Eocene to Early Miocene
Mounted H. sp. skeleton, Science Museum of Minnesota
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: †Hyaenodonta
Superfamily: †Hyaenodontoidea
Family: †Hyaenodontidae
Subfamily: †Hyaenodontinae
Tribe: †Hyaenodontini
Leidy, 1869[1]
Genus: †Hyaenodon
Laizer & Parieu, 1838
Type species
†Hyaenodon leptorhynchus
Laizer and Parieu, 1838
Species
[see classification]
Synonyms[2]
synonyms of genus:
  • Macropterodon (Lavrov, 1999)[3]
  • Megalopterodon (Dashzeveg, 1964)
  • Microhyaenodon (Lavrov, 1999)[3]
  • Neohyaenodon (Thorpe, 1922)
  • Protohyaenodon (Stock, 1933)
  • Pseudopterodon (Schlosser, 1887)
  • Taxotherium (Blainville, 1841)
synonyms of species:
  • H. brachyrhynchus:
    • Canis brachyrhynchus (Blainville, 1841)
    • Hyaenodon cuvieri (Pictet, 1853)[4]
    • Hyaenodon leptorhynchus (Dujardin, 1840)[5]
    • Hyaenodon parisiensis (Laurillard, 1845)[6]
    • Hyaenodon vulpinus (Gervais, 1873)
    • Nasua parisiensis (Blainville, 1841)
    • Pterodon brachyrhynchus (Pomel, 1846)[7]
    • Pterodon cuvieri (Pomel, 1846)
    • Taxotherium parisiense (Blainville, 1841)
  • H. brevirostrus:
    • Hyaenodon brevirostris (Joeckel, 1997)[8]
    • Protohyaenodon brevirostrus (Mellett, 1977)
  • H. chunkhtensis:
    • Microhyaenodon chunkhtensis (Lavrov, 1999)
  • H. crucians:
    • Hyaenodon leptocephalus (Scott, 1888)[9]
    • Hyaenodon minutus (Douglass, 1902)
    • Hyaenodon paucidens (Osborn & Wortman, 1894)[10]
    • Protohyaenodon crucians (Mellett, 1977)
    • Prtohyaenodon crucians (Lavrov, 1999)
    • Pseudopterodon minutus (Douglass, 1902)
  • H. dubius:
    • Hyaenodon aymardi (Filhol, 1882)[11]
  • H. exiguus:
    • Hyaenodon exigus (Lavrov, 1999)
    • Pterodon exiguum (Gervais, 1873)
  • H. filholi:
    • Hyaenodon compressus (Filhol, 1876)
    • Hyaenodon vulpinus (Filhol, 1876)
    • Microhyaenodon filholi (Lavrov, 1999)
    • Pseudopterodon ganodus (Schlosser, 1887)
  • H. gervaisi:
    • Hyaenodon ambiguus (Martin, 1906)
  • H. gigas:
    • Macropterodon zelenovi (Lavrov, 1999)
    • Neohyaenodon gigas (Lavrov, 1999)
  • H. heberti:
    • Hyaenodon arnaudi (Depéret, 1917)[12]
  • H. horridus:
    • Hyaenodon cruentus (Leidy, 1853)
    • Neohyaenodon horridus (Thorpe, 1922)
    • Neohyaenodon semseyi (Kretzoi, 1941)[13]
  • H. incertus:
    • Neohyaenodon incertus (Lavrov, 1999)
    • Pterodon exploratus (Dashzeveg, 1985)
  • H. leptorhynchus:
    • Canis leptorhynchus (Blainville, 1841)
    • Hyaenodon bavaricus (Dehm, 1935)[14]
    • Hyaenodon cayluxi (Filhol, 1876)
    • Hyaenodon martini (Depéret, 1917)
    • Hyaenodon milloquensis (Martin, 1906)
    • Pterodon leptorhynchus (Pomel, 1846)
  • H. macrocephalus:
    • Neohyaenodon macrocephalus (Lavrov, 1999)
  • H. megaloides:
    • Neohyaenodon megaloides (Mellett, 1977)
  • H. microdon:
    • Microhyaenodon microdon (Lavrov, 1999)
    • Protohyaenodon microdon (Mellett, 1977)
  • H. milvinus:
    • Neohyaenodon milvinus (Lavrov, 1999)
  • H. minor:
    • Hyaenodon aimi (Cooper, 1926)[15]
  • H. mongoliensis:
    • Epipterodon mongoliensis (Lavrov, 1999)
    • Megalopterodon mongoliensis (Dashzeveg, 1964)
    • Neohyaenodon mongoliensis (Morlo & Nagel, 2006)[16]
    • Pterodon mongoliensis (Van Valen, 1967)[17]
  • H. montanus:
    • Protohyaenodon montanus
    • Neohyaenodon montanus (Mellett, 1977)
  • H. mustelinus:
    • Protohyaenodon mustelinus (Scott, 1894)
  • H. pervagus:
    • Hyaenodon neimongoliensis (Huang & Zhu, 2002)[18]
  • H. pumilus:
    • Microhyaenodon pumilus (Lavrov, 1999)
  • H. raineyi:
    • Microhyaenodon raineyi (Lavrov, 1999)
    • Protohyaenodon raineyi (Gustafson, 1986)
  • H. requieni:
    • Hyaenodon heberti euzetensis (Depéret, 1917)
    • Pterodon requieni (Gervais, 1846)
  • H. venturae:
    • Hyaenodon exiguus (Stock, 1933)
    • Microhyaenodon venturae (Lavrov, 1999)
    • Protohyaenodon exiguus (Stock, 1933)
    • Protohyaenodon venturae (Mellett, 1977)
  • H. vetus:
    • Neohyaenodon vetus (Mellett, 1977)
    • Pterodon californicus (Stock, 1933)
  • H. yuanchuensis:
    • Hyaenodon yuanchüensis (Young, 1937)

Description
Skull of Hyaenodon horridus
Life reconstruction of H. horridus

Typical of early carnivorous mammals, individuals of Hyaenodon had a very massive skull, but a small brain. The skull is 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.[20]

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.[21] The average weight of adult or subadult H. horridus, the largest North American species, is estimated to about 40 kg (88 lb) and may not have exceeded 60 kg (130 lb). H. gigas, the largest Hyaenodon species, was much larger, being 378 kg (833 lb) and around 10 feet (3.0 m).[20] H. crucians from the early Oligocene of North America is estimated to only 10 to 25 kg (22 to 55 lb). H. microdon and H. mustelinus from the late Eocene of North America were even smaller and weighed probably about 5 kg (11 lb).[22]

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.[23]

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 primitive horses like Mesohippus, Brontotheres, early camels, oreodonts and even primitive rhinos.[20] 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.[24][25]

In North America the last Hyaenodon, in the form of H. brevirostrus, disappeared in the late Oligocene.[26] In Europe, they had already vanished earlier in the Oligocene.[21]

Classification and phylogeny
Reconstruction of Hyaenodon by Heinrich Harder (around 1920.)
Reconstruction of H. horridus and Leptomeryx evansi by W. B. Scott (1913.)

Taxonomy
  • Tribe: †Hyaenodontini (Leidy, 1869)
    • Genus: †Hyaenodon (Laizer & Parieu, 1838)
      • †Hyaenodon brachyrhynchus (Blainville, 1841)[27]
      • †Hyaenodon chunkhtensis (Dashzeveg, 1985)[28]
      • †Hyaenodon dubius (Filhol, 1873)[29]
      • †Hyaenodon eminus (Matthew & Granger, 1925)[30]
      • †Hyaenodon exiguus (Gervais, 1873)[31]
      • †Hyaenodon filholi (Schlosser, 1887)[32]
      • †Hyaenodon gervaisi (Martin, 1906)[33]
      • †Hyaenodon heberti (Europe, 41-33.9 mya) (Filhol, 1876)[34]
      • †Hyaenodon leptorhynchus (Laizer & Parieu, 1838)[35]
      • †Hyaenodon minor (Lange-Badré, 1979)[36]
      • †Hyaenodon pervagus (Matthew & Granger, 1924)[37]
      • †Hyaenodon pumilus (Lavrov, 2019[38]
      • †Hyaenodon requieni (Gervais, 1846)[39]
      • †Hyaenodon rossignoli (Lange-Badré, 1979)
      • †Hyaenodon weilini (China, 23-16,9 mya) (Wang, 2005)[21]
      • †Hyaenodon yuanchuensis (Young, 1937)[40]
      • Subgenus: †Neohyaenodon (paraphyletic subgenus) (Thorpe, 1922)[41]
        • †Hyaenodon gigas (Dashzeveg, 1985)
        • †Hyaenodon horridus (Leidy, 1853)[42]
        • †Hyaenodon incertus (Dashzeveg, 1985)
        • †Hyaenodon macrocephalus (Lavrov, 1999)[3]
        • †Hyaenodon megaloides (Mellett, 1977)[43]
        • †Hyaenodon milvinus (Lavrov, 1999)[3]
        • †Hyaenodon mongoliensis (Dashzeveg, 1964)[44]
        • †Hyaenodon montanus (Douglass, 1902)[45]
        • †Hyaenodon vetus (Stock, 1933)[46]
      • Subgenus: †Protohyaenodon (paraphyletic subgenus) (Stock, 1933)
        • †Hyaenodon brevirostrus (Macdonald, 1970)[47]
        • †Hyaenodon crucians (Leidy, 1853)
        • †Hyaenodon microdon (Mellett, 1977)
        • †Hyaenodon mustelinus (North America, 38-30 mya) (Scott, 1894)[48]
        • †Hyaenodon raineyi (Gustafson, 1986)[49]
        • †Hyaenodon venturae (Mellett, 1977)

Phylogeny

The phylogenetic relationships of genus Hyaenodon are shown in the following cladogram[3][50][51][52][53][54]

 †Hyaenodontidae 

†Paracynohyaenodon

†Quercytherium

†Cynohyaenodon lautricensis

†Cynohyaenodon cayluxi

†Cynohyaenodon ruetimeyeri

†Cynohyaenodon trux

 †Cynohyaenodon

†Cynohyaenodon smithae

†Boritia

 ? 

†Protoproviverra

 ? 

†Praecodens

†Leonhardtina

†Preregidens

†Cartierodon

†Prodissopsalis

 ? 

†Paenoxyaenoides

†Eurotherium theriodis

†Eurotherium mapplethorpei

 †Eurotherium

†Eurotherium matthesi

†Alienetherium

†Matthodon

†Oxyaenoidinae

 ? 

†Neosinopa

 †Hyaenodontinae 
 ? 

†Consobrinus

†Propterodon morrisi

†Propterodon sp. (ZIN no. 34494)

†Propterodon paganensis

†Propterodon tongi

†Propterodon witteri

†Epipterodontini

 †Hyaenodontini 
 †Hyaenodon 

†Hyaenodon chunkhtensis

†Hyaenodon eminus

 †Hyaenodon venturae 

 (†Protohyaenodon) 

†Hyaenodon pumilus

†Hyaenodon filholi

 †Hyaenodon microdon 

 †Hyaenodon raineyi 

 †Hyaenodon mustelinus 

 †Hyaenodon brevirostrus 

 †Hyaenodon crucians 

†Hyaenodon yuanchuensis

†Hyaenodon brachyrhynchus

†Hyaenodon gervaisi

†Hyaenodon requieni

†Hyaenodon minor

†Hyaenodon dubius

†Hyaenodon leptorhynchus

†Hyaenodon exiguus

†Hyaenodon sp. (Ergiliyn Dzo)

†Hyaenodon pervagus

†Hyaenodon rossignoli

†Hyaenodon horridus

†Hyaenodon megaloides

†Hyaenodon montanus

†Hyaenodon vetus

†Hyaenodon macrocephalus

†Hyaenodon milvinus

†Hyaenodon gigas

†Hyaenodon incertus

†Hyaenodon mongoliensis

†Hyaenodon heberti

†Hyaenodon weilini
†Cynohyaenodon/Quercytherium clade
†Leonhardtina clade
†Eurotherium clade
†Matthodon clade
†Oxyaenoides clade
†Propterodon

See also


References
Wikimedia Commons has media related to Hyaenodon.
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