Hyainailouridae

Hyainailouridae ("hyena-cats") is a paraphyletic family of extinct predatory mammals from extinct paraphyletic superfamily Hyainailouroidea within extinct order Hyaenodonta. Hyaenodontids arose during the middle Eocene and persisted well into the middle Miocene. Fossils of this group have been found in Asia, Africa, North America and Europe.

Hyainailouridae
Temporal range: middle Eocene to middle Miocene
Comparison of various Early to Middle Miocene hyaenodonts, including the hyainailurids Hyainailouros sulzeri (top) and Megistotherium osteothlastes (center), and teratodontid Dissopsalis pyroclasticus
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: †Hyaenodonta
Superfamily: †Hyainailouroidea
Family: †Hyainailouridae
Pilgrim, 1932[1]
Type genus
†Hyainailouros
(polyphyletic genus)
Biedermann, 1863
Subfamilies
[see classification]
Synonyms
  • Hyaenaelurinae (Pilgrim, 1932)
  • Pterodontidae (Polly, 1996)[2]
  • Pterodontinae (Polly, 1996)

General characteristics

Hyainailourids are characterized by long skulls, slender jaws, slim bodies, and a plantigrade stance. They generally ranged in size from 30 to 140 cm at the shoulder. While some measured as much as 1.4 m high at the shoulder with head-body length up to 3.2 m and weighed up to 1,500 kg, most were in the 5–15 kg range, equivalent to a mid-sized dog. The anatomy of their skulls show that they had a particularly acute sense of smell, while their teeth were adapted for shearing, rather than crushing.

At least one hyainailourid lineage, Apterodontinae, was specialised for aquatic, otter-like habits.[3]

Range

The oldest member of the Hyainailouridae is Orienspterodon from the middle Eocene of China.[4] They were important hypercarnivores in Eurasia and Africa during the Oligocene, but gradually declined, with almost the entire family becoming extinct by the close of the Oligocene. Only Simbakubwa, Megistotherium and several of its sister genera, including Hyainailouros and Sivapterodon, survived into the Miocene.[5] Traditionally this has been attributed to competition with carnivorans, but no formal examination of the correlation between the decline of hyaenodontids and the expansion of carnivorans has been recorded, and the latter may simply have moved into vacant niches after the extinction of hyaenodontid species.

Classification and phylogeny

Relations

Hyainailouridae used to be considered a subfamily of Hyaenodontidae, but cladistic study by Sole et al., (2013, 2015) treats it as a distinct family. Two subfamilies are recognized, Apterodontinae and paraphyletic Hyainailourinae.[6][7]

Taxonomy
  • Family: †Hyainailouridae (paraphyletic family) (Pilgrim, 1932)
    • (unranked): †Maocyon/Orienspterodon clade
      • Genus: †Maocyon (Averianov, 2023)
        • †Maocyon peregrinus (Averianov, 2023)
      • Genus: †Orienspterodon (Egi, 2007)
        • †Orienspterodon dahkoensis (Chow, 1975)
    • Subfamily: †Apterodontinae (Szalay, 1967)
      • Genus: †Apterodon (Fischer, 1880)
        • †Apterodon altidens (Schlosser, 1910)
        • †Apterodon gaudryi (Fischer, 1880)
        • †Apterodon langebadreae (Grohé, 2012)
        • †Apterodon macrognathus (Andrews, 1904)
        • †Apterodon rauenbergensis (Frey, 2010)
        • †Apterodon saghensis (Simons & Gingerich, 1976)
        • †Apterodon sp. [Dur At-Talah escarpment, Libya] (Grohé, 2012)
      • Genus: †Quasiapterodon (Lavrov, 1999)
        • †Quasiapterodon minutus (Schlosser, 1910)
    • Subfamily: †Hyainailourinae (paraphyletic subfamily) (Pilgrim, 1932)
      • Genus: †Megistotherium (Savage, 1973)
        • †Megistotherium osteothlastes (Savage, 1973)
      • Genus: †Mlanyama (Rasmussen & Gutierrez, 2009)
        • †Mlanyama sugu (Rasmussen & Gutierrez, 2009)
      • Genus: †Pakakali (Borths & Stevens, 2017)
        • †Pakakali rukwaensis (Borths & Stevens, 2017)
      • Genus: †Simbakubwa (Borths & Stevens, 2019)
        • †Simbakubwa kutokaafrika (Borths & Stevens, 2019)
      • (unranked): †Akhnatenavus clade
        • Genus: †Akhnatenavus (Holroyd, 1999)
          • †Akhnatenavus leptognathus (Osborn, 1909)
          • †Akhnatenavus nefertiticyon (Borths, 2016)
        • Genus: †Hemipsalodon (Cope, 1885)
          • †Hemipsalodon grandis (Cope, 1885)
          • †Hemipsalodon viejaensis (Gustafson, 1986)
        • Genus: †Ischnognathus (Stovall, 1948)
          • †Ischnognathus savagei (Stovall, 1948)
      • Tribe: †Hyainailourini (polyphyletic tribe) (Ginsburg, 1980)
        • Genus: †Exiguodon (Morales & Pickford, 2017)
          • †Exiguodon pilgrimi (Savage, 1965)
        • Genus: †Falcatodon (Morales & Pickford, 2017)
          • †Falcatodon schlosseri (Holroyd, 1999)
        • Genus: †Hyainailouros (polyphyletic genus) (Biedermann, 1863)
          • †Hyainailouros bugtiensis (Pilgrim, 1912)
          • †Hyainailouros napakensis (Ginsburg, 1980)
          • †Hyainailouros sulzeri (Biedermann, 1863)
        • Genus: †Parapterodon (Lange-Badré, 1979)
          • †Parapterodon lostangensis (Lange-Badré, 1979)
        • Genus: †Sectisodon (Morales & Pickford, 2017)
          • †Sectisodon markgrafi (Holroyd, 1999)
          • †Sectisodon occultus (Morales & Pickford, 2017)
        • Genus: †Sivapterodon (Ginsburg, 1980)
          • †Sivapterodon lahirii (Pilgrim, 1932)
        • Subtribe: †Isohyaenodontina (polyphyletic subtribe) (Lavrov, 1999)
          • Genus: †Isohyaenodon (polyphyletic genus) (Savage, 1965)
            • †Isohyaenodon andrewsi (Savage, 1965)
            • †Isohyaenodon zadoki (Savage, 1965)
        • (unranked): †Pterodon clade
          • Genus: †Kerberos (Solé, 2015)
            • †Kerberos langebadreae (Solé, 2015)
          • Subtribe: †Pterodontina (Lavrov, 1999)
            • Genus: †Pterodon (Blainville, 1839)
              • †Pterodon dasyuroides (Blainville, 1839)
          • Incertae sedis:
            • †"Pterodon" syrtos (Holroyd, 1999)
        • Incertae sedis:
          • †"Pterodon" africanus (Andrews, 1903)
          • †"Pterodon" phiomensis (Osborn, 1909)
          • †"Pterodon" sp. [DPC 5036] (Holroyd, 1999)
      • Tribe: †Leakitheriini (Lavrov, 1999)
        • Genus: †Leakitherium (Savage, 1965)
          • †Leakitherium hiwegi (Savage, 1965)
      • Tribe: †Metapterodontini (Morales & Pickford, 2017)
        • Genus: †Metapterodon (Stromer, 1926)
          • †Metapterodon brachycephalus (Osborn, 1909)
          • †Metapterodon kaiseri (Stromer, 1926)
          • †Metapterodon stromeri (Morales, 1998)
      • Tribe: †Paroxyaenini (Lavrov, 2007)
        • Genus: †Paroxyaena (Martin, 1906)
          • †Paroxyaena galliae (Filhol, 1881)
          • †Paroxyaena pavlovi (Lavrov, 2007)
      • Incertae sedis:
        • †"Pterodon" sp. [BC 15’08] (Pickford, 2008)
        • †Hyainailourinae sp. [GSN AD 100’96] (Morales, 1998)
        • †Hyainailourinae sp. [UON 84-359] (Solé, 2016)
        • †Hyainailourinae sp. A [DPC 6555] (Holroyd, 1999)
        • †Hyainailourinae sp. C [DPC 9243 & DPC 10315] (Holroyd, 1999)
        • †Hyainailourinae sp. D [DPC 6545] (Holroyd, 1999)

Phylogeny

The phylogenetic relationships of family Hyainailouridae are shown in the following cladogram:[3][8][9][10][11][4]

 †Hyaenodonta 

†Eoproviverra

†Boualitomidae

†Arfiidae

†Limnocyonidae

†Sinopidae

†Hyaenodontoidea

†Gazinocyon

†Pyrocyon

†Galecyon

 †Afro‑Arabian clade 

†Parvavorodon

†Indohyaenodontidae

†Glibzegdouia

†Koholiidae

†Tritemnodon

†Teratodontidae

 †Apterodontinae 
 †Apterodon 

†Apterodon altidens

†Apterodon gaudryi

 ? 

†Apterodon rauenbergensis

†Apterodon langebadreae

†Apterodon macrognathus

 ? 

†Apterodon sp. (Dur At-Talah escarpment, Libya)

†Apterodon saghensis

 †Quasiapterodon 

†Quasiapterodon minutus

 †Maocyon 

†Maocyon peregrinus

 †Maocyon/Orienspterodon clade
 †Orienspterodon 

†Orienspterodon dahkoensis

†Hyainailourinae
 †Hemipsalodon 

†Hemipsalodon grandis

†Hemipsalodon viejaensis

 †Ischnognathus 

†Ischnognathus savagei

 ? 
 †Akhnatenavus clade
 †Akhnatenavus 

†Akhnatenavus leptognathus

†Akhnatenavus nefertiticyon

†"Pterodon" sp. (BC 15’08)

†Hyainailourinae sp. (UON 84-359)

†Hyainailourinae sp. C (DPC 9243 & DPC 10315)

†Hyainailourinae sp. D (DPC 6545)

 †Kerberos 

†Kerberos langebadreae

†"Pterodon" syrtos

 †Pterodon clade
 †Pterodontina 
 †Pterodon 

†Pterodon dasyuroides

†Hyainailourinae sp. A (DPC 6555)

†"Pterodon" africanus

 †Parapterodon 

†Parapterodon lostangensis

†"Pterodon" sp. (DPC 5036)

†"Pterodon" phiomensis

†Hyainailourini

 †Paroxyaenini 
 †Paroxyaena 

†Paroxyaena galliae

†Paroxyaena pavlovi

 †Sectisodon 

†Sectisodon markgrafi

†Sectisodon occultus

 †Falcatodon 

†Falcatodon schlosseri

 †Exiguodon 

†Exiguodon pilgrimi

†Isohyaenodon zadoki

†Isohyaenodon  (†Isohyaenodontina)

†Isohyaenodon andrewsi

 †Sivapterodon 

†Sivapterodon lahirii

†Hyainailouros bugtiensis

†Hyainailouros napakensis

†Hyainailouros

†Hyainailouros sulzeri

†Hyainailourinae sp. (GSN AD 100’96)

 †Simbakubwa 

†Simbakubwa kutokaafrika

 †Leakitheriini 
 †Leakitherium 

†Leakitherium hiwegi

 †Megistotherium 

†Megistotherium osteothlastes

 †Mlanyama 

†Mlanyama sugu

 †Metapterodontini 
 †Metapterodon 

†Metapterodon brachycephalus

†Metapterodon kaiseri

†Metapterodon stromeri

 †Pakakali 

†Pakakali rukwaensis

†Prionogalidae

 †Hyainailouridae
†Lahimia clade
†Arfia clade
†Galecyon clade
†Indohyaenodon clade
†Tritemnodon clade

See also

References
  1. Pilgrim G. E. (1932). "The fossil Carnivora of India. Memoirs of the Geological Survey of India". Palaeontologica Indica. 18 (1–232). doi:10.1017/S0016756800096448. Archived from the original on April 13, 2023.
  2. P. D. Polly (1996). "The skeleton of Gazinocyon vulpeculus gen. et. comb nov. and the cladistic relationships of Hyaenodontidae (Eutheria, Mammalia)" (PDF). Journal of Vertebrate Paleontology. 16 (2): 303–319. Archived (PDF) from the original on June 25, 2023.
  3. Laudet, V.; Grohé, C.; Morlo, M.; Chaimanee, Y.; Blondel, C.; Coster, P.; Valentin, X.; Salem, M.; Bilal, A. A.; Jaeger, J. J.; Brunet, M. (2012). "New Apterodontinae (Hyaenodontida) from the Eocene Locality of Dur At-Talah (Libya): Systematic, Paleoecological and Phylogenetical Implications". PLOS ONE. 7 (11): e49054. Bibcode:2012PLoSO...749054G. doi:10.1371/journal.pone.0049054. PMC 3504055. PMID 23185292.
  4. Averianov, Alexander; Obraztsova, Ekaterina; Danilov, Igor; Jin, Jian-Hua (2023). "A new hypercarnivorous hyaenodont from the Eocene of South China". Frontiers in Ecology and Evolution. 11. doi:10.3389/fevo.2023.1076819. ISSN 2296-701X.
  5. Borths, M. R.; Stevens, N. J. (April 2019). "Simbakubwa kutokaafrika, gen. et sp. nov. (Hyainailourinae, Hyaenodonta, 'Creodonta,' Mammalia), a gigantic carnivore from the earliest Miocene of Kenya". Journal of Vertebrate Paleontology. 39: e1570222. Bibcode:2019JVPal..39E0222B. doi:10.1080/02724634.2019.1570222. S2CID 145972918.
  6. Solé, Floréal; Lhuillier, Julie; Adaci, Mohammed; Bensalah, Mustapha; Mahboubi, M’hammed; Tabuce, Rodolphe (2013). "The hyaenodontidans from the Gour Lazib area (?Early Eocene, Algeria): implications concerning the systematics and the origin of the Hyainailourinae and Teratodontinae". Journal of Systematic Palaeontology. 12 (3): 303–322. doi:10.1080/14772019.2013.795196. S2CID 84475034.
  7. Solé, Floréal; Amson, Eli; Borths, Matthew; Vidalenc, Dominique; Morlo, Michael; Bastl, Katharina (2015). "A New Large Hyainailourine from the Bartonian of Europe and Its Bearings on the Evolution and Ecology of Massive Hyaenodonts (Mammalia)". PLOS ONE. 10 (9): e0135698. Bibcode:2015PLoSO..1035698S. doi:10.1371/journal.pone.0135698. PMC 4580617. PMID 26398622.
  8. Borths, Matthew R.; Stevens, Nancy J. (2017). "Deciduous dentition and dental eruption of Hyainailouroidea (Hyaenodonta, "Creodonta," Placentalia, Mammalia)". Palaeontologia Electronica. 20 (3): 55A. doi:10.26879/776.
  9. Matthew R. Borths; Nancy J. Stevens (2019). "Simbakubwa kutokaafrika, gen. et sp. nov. (Hyainailourinae, Hyaenodonta, 'Creodonta,' Mammalia), a gigantic carnivore from the earliest Miocene of Kenya". Journal of Vertebrate Paleontology. 39 (1): e1570222. Bibcode:2019JVPal..39E0222B. doi:10.1080/02724634.2019.1570222. S2CID 145972918.
  10. Floréal Solé; Bernard Marandat; Fabrice Lihoreau (2020). "The hyaenodonts (Mammalia) from the French locality of Aumelas (Hérault), with possible new representatives from the late Ypresian". Geodiversitas. 42 (13): 185–214. doi:10.5252/geodiversitas2020v42a13. S2CID 219585388.
  11. Solé, F.; Morlo, M.; Schaal, T.; Lehmann, T. (2021). "New hyaenodonts (Mammalia) from the late Ypresian locality of Prémontré (France) support a radiation of the hyaenodonts in Europe already at the end of the early Eocene". Geobios. 66–67: 119–141. Bibcode:2021Geobi..66..119S. doi:10.1016/j.geobios.2021.02.004. S2CID 234848856.
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