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|image_caption = Comparison of various Early to Middle Miocene hyaenodonts, including the hyainailurids ''[[Hyainailouros|Hyainailouros sulzeri]]'' (top) and ''[[Megistotherium|Megistotherium osteothlastes]]'' (center), and teratodontid ''[[Dissopsalis|Dissopsalis pyroclasticus]]''
|image_caption = Comparison of various Early to Middle Miocene hyaenodonts, including the hyainailurids ''[[Hyainailouros|Hyainailouros sulzeri]]'' (top) and ''[[Megistotherium|Megistotherium osteothlastes]]'' (center), and teratodontid ''[[Dissopsalis|Dissopsalis pyroclasticus]]''
|taxon = Hyainailouridae
|taxon = Hyainailouridae
|authority = Pilgrim, 1932<ref>Pilgrim G. E. (1932.) [https://www.cambridge.org/core/journals/geological-magazine/article/abs/fossil-carnivora-of-india-by-guy-e-pilgrim-dsc-palaeontologia-indica-new-series-vol-xviii-pp-iii-232-calcutta-government-of-india-1932-price-22s-6d/86B306399D654CC4D032CA9D670354F7 "The fossil Carnivora of India."] Memoirs of the Geological Survey of India. Palaeontologica Indica, 18:1-232.</ref>
|authority = Pilgrim, 1932<ref>{{cite journal|author=Pilgrim G. E.|date=1932|url=https://www.cambridge.org/core/journals/geological-magazine/article/abs/fossil-carnivora-of-india-by-guy-e-pilgrim-dsc-palaeontologia-indica-new-series-vol-xviii-pp-iii-232-calcutta-government-of-india-1932-price-22s-6d/86B306399D654CC4D032CA9D670354F7|title=The fossil Carnivora of India. Memoirs of the Geological Survey of India.|journal=Palaeontologica Indica|volume=18|issue=1-232|doi=10.1017/S0016756800096448|archive-url=https://web.archive.org/web/20230413194629/https://www.cambridge.org/core/journals/geological-magazine/article/abs/fossil-carnivora-of-india-by-guy-e-pilgrim-dsc-palaeontologia-indica-new-series-vol-xviii-pp-iii-232-calcutta-government-of-india-1932-price-22s-6d/86B306399D654CC4D032CA9D670354F7|archive-date=April 13, 2023|url-status=live}}</ref>
|type_genus = †''[[Hyainailouros]]''<br><sup>([[polyphyletic]] genus)</sup>
|type_genus = †''[[Hyainailouros]]''<br><sup>([[polyphyletic]] genus)</sup>
|type_genus_authority = Biedermann, 1863
|type_genus_authority = Biedermann, 1863
|subdivision_ranks = Subfamilies
|subdivision_ranks = Subfamilies
|subdivision =
|subdivision = {{center|[see [[#Taxonomy|classification]]]}}
* †[[Apterodontinae]]
* †[[Hyainailourinae]]<br><sup>([[paraphyletic]] subfamily)</sup>
|synonyms =
|synonyms =
* Hyaenaelurinae {{small|(Pilgrim, 1932)}}
* Hyaenaelurinae {{small|(Pilgrim, 1932)}}
* Pterodontidae {{small|(Polly, 1996)}}<ref>P. D. Polly (1996.) [https://pollylab.indiana.edu/doc/papers/Polly,%201996,%20Gazinocyon.pdf "The skeleton of ''Gazinocyon vulpeculus'' gen. et. comb nov. and the cladistic relationships of Hyaenodontidae (Eutheria, Mammalia)."] Journal of Vertebrate Paleontology 16(2):303-319</ref>
* Pterodontidae {{small|(Polly, 1996)}}<ref>{{cite journal|author=P. D. Polly|date=1996|url=https://pollylab.indiana.edu/doc/papers/Polly,%201996,%20Gazinocyon.pdf|title=The skeleton of ''Gazinocyon vulpeculus'' gen. et. comb nov. and the cladistic relationships of Hyaenodontidae (Eutheria, Mammalia)|journal=Journal of Vertebrate Paleontology|volume=16|issue=2|pages=303-319|archive-url=https://web.archive.org/web/20230625215529/https://pollylab.indiana.edu/doc/papers/Polly,%201996,%20Gazinocyon.pdf|archive-date=June 25, 2023|url-status=live}}</ref>
* Pterodontinae {{small|(Polly, 1996)}}
* Pterodontinae {{small|(Polly, 1996)}}
}}
}}


'''Hyainailouridae''' ("[[hyena]]-[[Felidae|cats]]") is a [[paraphyletic]] family of extinct predatory [[mammal]]s 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''' ("[[hyena]]-[[Felidae|cats]]") is a family of extinct predatory [[mammal]]s within the 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]].


==General characteristics==
==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&nbsp;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&nbsp;kg, most were in the 5–15&nbsp;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.
Hyainailourids are characterized by long skulls, slender jaws, slim bodies, and a [[plantigrade]] stance. They generally ranged in size from 30 to 140&nbsp;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&nbsp;kg, most were in the 5–15&nbsp;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.<ref name="ReferenceA">{{cite journal |last1=Laudet |first1=V. |last2=Grohé |first2=C. |last3=Morlo |first3=M. |last4=Chaimanee |first4=Y. |last5=Blondel |first5=C. |last6=Coster |first6=P. |last7=Valentin |first7=X. |last8=Salem |first8=M. |last9=Bilal |first9=A. A. |last10=Jaeger |first10=J. J. |last11=Brunet |first11=M. |title=New Apterodontinae (Hyaenodontida) from the Eocene Locality of Dur At-Talah (Libya): Systematic, Paleoecological and Phylogenetical Implications |journal=PLOS ONE |volume=7 |issue=11 |year=2012 |pages=e49054 |doi=10.1371/journal.pone.0049054 |pmid=23185292 |pmc=3504055 |bibcode=2012PLoSO...749054G |doi-access=free}}</ref>
At least one hyainailourid lineage, [[Apterodontinae]], was specialised for aquatic, [[otter]]-like habits.<ref name="ReferenceA"/>


==Range==
==Range==
They were important [[Hypercarnivore|hypercarnivores]] in Eurasia and Africa during the [[Oligocene]], but gradually declined, with almost the entire family becoming extinct by the close of the Oligocene.{{cn|date=May 2019}} Only ''[[Megistotherium]]'' and several of its sister genera, including ''[[Hyainailouros]]'' and ''[[Sivapterodon]]'', survived into the Miocene.<ref>{{cite journal |last1=Borths |first1=M. R. |last2=Stevens |first2=N. J. |date=April 2019 |title=''Simbakubwa kutokaafrika'', gen. et sp. nov. (Hyainailourinae, Hyaenodonta, 'Creodonta,' Mammalia), a gigantic carnivore from the earliest Miocene of Kenya |journal=Journal of Vertebrate Paleontology |volume=39 |pages=e1570222 |doi=10.1080/02724634.2019.1570222 |s2cid=145972918|url=https://figshare.com/articles/journal_contribution/_i_Simbakubwa_kutokaafrika_i_gen_et_sp_nov_Hyainailourinae_Hyaenodonta_Creodonta_Mammalia_a_gigantic_carnivore_from_the_earliest_Miocene_of_Kenya/8009951 }}</ref> 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.
The oldest member of the Hyainailouridae is ''[[Orienspterodon]]'' from the middle Eocene of China.<ref name="Maocyon">{{Cite journal |last1=Averianov |first1=Alexander |last2=Obraztsova |first2=Ekaterina |last3=Danilov |first3=Igor |last4=Jin |first4=Jian-Hua |date=2023 |title=A new hypercarnivorous hyaenodont from the Eocene of South China |journal=Frontiers in Ecology and Evolution |volume=11 |doi=10.3389/fevo.2023.1076819 |issn=2296-701X |doi-access=free }}</ref> They were important [[Hypercarnivore|hypercarnivores]] in Eurasia and Africa during the [[Oligocene]], but gradually declined, with almost the entire family becoming extinct by the close of the Oligocene.{{cn|date=May 2019}} Only ''[[Simbakubwa]]'', ''[[Megistotherium]]'' and several of its sister genera, including ''[[Hyainailouros]]'' and ''[[Sivapterodon]]'', survived into the Miocene.<ref>{{cite journal |last1=Borths |first1=M. R. |last2=Stevens |first2=N. J. |date=April 2019 |title=''Simbakubwa kutokaafrika'', gen. et sp. nov. (Hyainailourinae, Hyaenodonta, 'Creodonta,' Mammalia), a gigantic carnivore from the earliest Miocene of Kenya |journal=Journal of Vertebrate Paleontology |volume=39 |pages=e1570222 |doi=10.1080/02724634.2019.1570222 |bibcode=2019JVPal..39E0222B |s2cid=145972918|url=https://figshare.com/articles/journal_contribution/8009951 }}</ref> Traditionally this has been attributed to competition with [[carnivora]]ns, 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==
==Classification and phylogeny==
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===Taxonomy===
===Taxonomy===
* Family: †'''Hyainailouridae''' <sup>([[Paraphyly|paraphyletic]] family)</sup> {{small|(Pilgrim, 1932)}}
* Family: †'''Hyainailouridae''' {{small|(Pilgrim, 1932)}}
** (unranked): †'''''Maocyon''/''Orienspterodon'' clade'''
*** Genus: †''[[Maocyon]]'' {{small|(Averianov, 2023)}}
**** †''Maocyon peregrinus'' {{small|(Averianov, 2023)}}
*** Genus: †''[[Orienspterodon]]'' {{small|(Egi, 2007)}}
**** †''Orienspterodon dahkoensis'' {{small|(Chow, 1975)}}
**** †''Orienspterodon mianchiensis'' {{small|(Sheng, Bi & Li, 2024)}}
** Subfamily: †'''[[Apterodontinae]]''' {{small|(Szalay, 1967)}}
** Subfamily: †'''[[Apterodontinae]]''' {{small|(Szalay, 1967)}}
*** Genus: †''[[Apterodon]]'' {{small|(Fischer, 1880)}}
*** Genus: †''[[Apterodon]]'' {{small|(Fischer, 1880)}}
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**** †''Apterodon rauenbergensis'' {{small|(Frey, 2010)}}
**** †''Apterodon rauenbergensis'' {{small|(Frey, 2010)}}
**** †''Apterodon saghensis'' {{small|(Simons & Gingerich, 1976)}}
**** †''Apterodon saghensis'' {{small|(Simons & Gingerich, 1976)}}
**** †''Apterodon'' sp. [Dur At-Talah escarpment, Libya] {{small|(Grohé, 2012)}}
*** Genus: †''[[Quasiapterodon]]'' {{small|(Lavrov, 1999)}}
*** Genus: †''[[Quasiapterodon]]'' {{small|(Lavrov, 1999)}}
**** †''Quasiapterodon minutus'' {{small|(Schlosser, 1910)}}
**** †''Quasiapterodon minutus'' {{small|(Schlosser, 1910)}}
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*** Genus: †''[[Mlanyama]]'' {{small|(Rasmussen & Gutierrez, 2009)}}
*** Genus: †''[[Mlanyama]]'' {{small|(Rasmussen & Gutierrez, 2009)}}
**** †''Mlanyama sugu'' {{small|(Rasmussen & Gutierrez, 2009)}}
**** †''Mlanyama sugu'' {{small|(Rasmussen & Gutierrez, 2009)}}
*** Genus: †''[[Orienspterodon]]'' {{small|(Egi, 2007)}}
**** †''Orienspterodon dahkoensis'' {{small|(Chow, 1975)}}
*** Genus: †''[[Pakakali]]'' {{small|(Borths & Stevens, 2017)}}
*** Genus: †''[[Pakakali]]'' {{small|(Borths & Stevens, 2017)}}
**** †''Pakakali rukwaensis'' {{small|(Borths & Stevens, 2017)}}
**** †''Pakakali rukwaensis'' {{small|(Borths & Stevens, 2017)}}
Line 65: Line 66:
**** Genus: †''[[Ischnognathus]]'' {{small|(Stovall, 1948)}}
**** Genus: †''[[Ischnognathus]]'' {{small|(Stovall, 1948)}}
***** †''Ischnognathus savagei'' {{small|(Stovall, 1948)}}
***** †''Ischnognathus savagei'' {{small|(Stovall, 1948)}}
*** Tribe: †'''[[Hyainailourini]]''' <sup>([[Polyphyly|polyphyletic]] tribe)</sup> {{small|(Ginsburg, 1980)}}
*** Tribe: †'''[[Hyainailourini]]''' {{small|(Ginsburg, 1980)}}
**** Genus: †''[[Exiguodon]]'' {{small|(Morales & Pickford, 2017)}}
**** Genus: †''[[Exiguodon]]'' {{small|(Morales & Pickford, 2017)}}
***** †''Exiguodon pilgrimi'' {{small|(Savage, 1965)}}
***** †''Exiguodon pilgrimi'' {{small|(Savage, 1965)}}
**** Genus: †''[[Falcatodon]]'' {{small|(Morales & Pickford, 2017)}}
**** Genus: †''[[Falcatodon]]'' {{small|(Morales & Pickford, 2017)}}
***** †''Falcatodon schlosseri'' {{small|(Holroyd, 1999)}}
***** †''Falcatodon schlosseri'' {{small|(Holroyd, 1999)}}
**** Genus: †''[[Hyainailouros]]'' <sup>(polyphyletic genus)</sup> {{small|(Biedermann, 1863)}}
**** Genus: †''[[Hyainailouros]]'' {{small|(Biedermann, 1863)}}
***** †''Hyainailouros bugtiensis'' {{small|(Pilgrim, 1912)}}
***** †''Hyainailouros bugtiensis'' {{small|(Pilgrim, 1912)}}
***** †''Hyainailouros napakensis'' {{small|(Ginsburg, 1980)}}
***** †''Hyainailouros napakensis'' {{small|(Ginsburg, 1980)}}
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**** Genus: †''[[Sivapterodon]]'' {{small|(Ginsburg, 1980)}}
**** Genus: †''[[Sivapterodon]]'' {{small|(Ginsburg, 1980)}}
***** †''Sivapterodon lahirii'' {{small|(Pilgrim, 1932)}}
***** †''Sivapterodon lahirii'' {{small|(Pilgrim, 1932)}}
**** Subtribe: †'''[[Isohyaenodon]]tina''' <sup>(polyphyletic subtribe)</sup> {{small|(Lavrov, 1999)}}
**** Subtribe: †'''[[Isohyaenodon]]tina''' {{small|(Lavrov, 1999)}}
***** Genus: †''[[Isohyaenodon]]'' <sup>(polyphyletic genus)</sup> {{small|(Savage, 1965)}}
***** Genus: †''[[Isohyaenodon]]'' {{small|(Savage, 1965)}}
****** †''Isohyaenodon andrewsi'' {{small|(Savage, 1965)}}
****** †''Isohyaenodon andrewsi'' {{small|(Savage, 1965)}}
****** †''Isohyaenodon zadoki'' {{small|(Savage, 1965)}}
****** †''Isohyaenodon zadoki'' {{small|(Savage, 1965)}}
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***** †''[[Pterodon africanus|"Pterodon" africanus]]'' {{small|(Andrews, 1903)}}
***** †''[[Pterodon africanus|"Pterodon" africanus]]'' {{small|(Andrews, 1903)}}
***** †''[[Pterodon phiomensis|"Pterodon" phiomensis]]'' {{small|(Osborn, 1909)}}
***** †''[[Pterodon phiomensis|"Pterodon" phiomensis]]'' {{small|(Osborn, 1909)}}
***** †[[Pterodon sp. (DPC 5036)|''"Pterodon"'' sp. [''DPC 5036'']]] {{small|(Holroyd, 1999)}}
*** Tribe: †'''[[Leakitherium|Leakitheriini]]''' {{small|(Lavrov, 1999)}}
*** Tribe: †'''[[Leakitherium|Leakitheriini]]''' {{small|(Lavrov, 1999)}}
**** Genus: †''[[Leakitherium]]'' {{small|(Savage, 1965)}}
**** Genus: †''[[Leakitherium]]'' {{small|(Savage, 1965)}}
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***** †''Paroxyaena galliae'' {{small|(Filhol, 1881)}}
***** †''Paroxyaena galliae'' {{small|(Filhol, 1881)}}
***** †''Paroxyaena pavlovi'' {{small|(Lavrov, 2007)}}
***** †''Paroxyaena pavlovi'' {{small|(Lavrov, 2007)}}
*** ''[[Incertae sedis]]'':
**** †[[Pterodon sp. (BC 15’08)|''"Pterodon"'' sp. [''BC 15’08'']]] {{small|(Pickford, 2008)}}
**** †[[Hyainailourinae sp. (GSN AD 100’96)|Hyainailourinae sp. [''GSN AD 100’96'']]] {{small|(Morales, 1998)}}
**** †[[Hyainailourinae sp. (UON 84-359)|Hyainailourinae sp. [''UON 84-359'']]] {{small|(Solé, 2016)}}
**** †[[Hyainailourinae sp. A (DPC 6555)|Hyainailourinae sp. A [''DPC 6555'']]] {{small|(Holroyd, 1999)}}
**** †[[Hyainailourinae sp. C (DPC 9243 & DPC 10315)|Hyainailourinae sp. C [''DPC 9243'' & ''DPC 10315'']]] {{small|(Holroyd, 1999)}}
**** †[[Hyainailourinae sp. D (DPC 6545)|Hyainailourinae sp. D [''DPC 6545'']]] {{small|(Holroyd, 1999)}}

===Phylogeny===
The phylogenetic relationships of family Hyainailouridae are shown in the following cladogram:<ref name="ReferenceA">{{cite journal |last1=Laudet |first1=V. |last2=Grohé |first2=C. |last3=Morlo |first3=M. |last4=Chaimanee |first4=Y. |last5=Blondel |first5=C. |last6=Coster |first6=P. |last7=Valentin |first7=X. |last8=Salem |first8=M. |last9=Bilal |first9=A. A. |last10=Jaeger |first10=J. J. |last11=Brunet |first11=M. |title=New Apterodontinae (Hyaenodontida) from the Eocene Locality of Dur At-Talah (Libya): Systematic, Paleoecological and Phylogenetical Implications |journal=PLOS ONE |volume=7 |issue=11 |year=2012 |pages=e49054 |doi=10.1371/journal.pone.0049054 |pmid=23185292 |pmc=3504055 |bibcode=2012PLoSO...749054G |doi-access=free}}</ref><ref name="Matthew2017">{{Cite journal |last1=Borths |first1=Matthew R. |last2=Stevens |first2=Nancy J. |date=2017 |title=Deciduous dentition and dental eruption of Hyainailouroidea (Hyaenodonta, "Creodonta," Placentalia, Mammalia) |url=http://palaeo-electronica.org/content/2017/2056-deciduous-hyaenodont-teeth |journal=Palaeontologia Electronica |volume=20 |issue=3 |pages=55A |doi=10.26879/776 |via= |doi-access=free}}</ref><ref name="Simbakubwa">{{Cite journal |author1=Matthew R. Borths |author2=Nancy J. Stevens |year=2019 |title=''Simbakubwa kutokaafrika'', gen. et sp. nov. (Hyainailourinae, Hyaenodonta, 'Creodonta,' Mammalia), a gigantic carnivore from the earliest Miocene of Kenya |journal=Journal of Vertebrate Paleontology |volume=39 |issue=1 |pages=e1570222 |doi=10.1080/02724634.2019.1570222|s2cid=145972918 |url=https://figshare.com/articles/journal_contribution/_i_Simbakubwa_kutokaafrika_i_gen_et_sp_nov_Hyainailourinae_Hyaenodonta_Creodonta_Mammalia_a_gigantic_carnivore_from_the_earliest_Miocene_of_Kenya/8009951 }}</ref><ref name="Geodiversitas4213">{{Cite journal |author1=Floréal Solé |author2=Bernard Marandat |author3=Fabrice Lihoreau |year=2020 |title=The hyaenodonts (Mammalia) from the French locality of Aumelas (Hérault), with possible new representatives from the late Ypresian |journal=Geodiversitas |volume=42 |issue=13 |pages=185–214 |doi=10.5252/geodiversitas2020v42a13 |s2cid=219585388 |url=http://sciencepress.mnhn.fr/en/periodiques/geodiversitas/42/13|doi-access=free }}</ref><ref name="hyaenodontsGeobios">{{Cite journal |last1=Solé |first1=F. |last2=Morlo |first2=M. |last3=Schaal |first3=T. |last4=Lehmann |first4=T. |title=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 |year=2021 |journal=Geobios |volume=66-67 |pages=119–141 |doi=10.1016/j.geobios.2021.02.004|s2cid=234848856 }}</ref><ref name="Maocyon">{{Cite journal |last=Averianov |first=Alexander |last2=Obraztsova |first2=Ekaterina |last3=Danilov |first3=Igor |last4=Jin |first4=Jian-Hua |date=2023 |title=A new hypercarnivorous hyaenodont from the Eocene of South China |url=https://www.frontiersin.org/articles/10.3389/fevo.2023.1076819 |journal=Frontiers in Ecology and Evolution |volume=11 |doi=10.3389/fevo.2023.1076819/full |issn=2296-701X}}</ref>
{{clade |style=font-size:85%;line-height:100%;
|grouplabel1={{clade labels
|label1=†'''''Lahimia'' clade''' |top1=2%
|label2=†'''''Arfia'' clade''' |top2=4%
|label3=†'''''Galecyon'' clade''' |top3=12%
|label4=†'''''Indohyaenodon'' clade''' |top4=16%
|label5=†'''''Tritemnodon'' clade''' |top5=20%
|label6=†'''[[Hyainailouroidea]]''' |top6=72%
}}
|label1=&nbsp;†'''[[Hyaenodonta]]'''&nbsp;
|1={{clade
|1=†''[[Eoproviverra]]''
|2={{clade
|grouplabel1x=†'''''Lahimia'' clade'''
|1=†'''[[Boualitomidae]]''' |barbegin1=purple |barend1=purple
|2={{clade
|1={{clade
|1={{clade
|grouplabel1x=†'''''Arfia'' clade'''
|1=†'''[[Arfia|Arfiidae]]''' |barbegin1=teal |barend1=teal
|2=†'''[[Limnocyonidae]]'''
}}
|2=†'''[[Sinopidae]]'''
}}
|2=†'''[[Hyaenodontoidea]]'''
|3={{clade
|1={{clade
|1=†''[[Gazinocyon]]''
|2=†''[[Pyrocyon]]''
}}
|grouplabel2x=†'''''Galecyon'' clade'''
|2=†''[[Galecyon]]'' |barbegin2=blue |barend2=blue
|label3=&nbsp;†'''Afro-Arabian clade'''&nbsp;
|3={{clade
|1=†''[[Parvavorodon]]''
|2={{clade
|grouplabel1x=†'''''Indohyaenodon'' clade''' |barbegin1=yellow |barend1=yellow
|1={{clade
|1=†'''[[Indohyaenodontidae]]'''
|2=†''[[Glibzegdouia]]''
}}
|2={{clade
|thickness=2 |thickness1=1 |thickness2=1 |color=green |color1=black |color2=black
|1=†'''[[Koholia|Koholiidae]]'''
|grouplabel2x=†'''''Tritemnodon'' clade''' |barbegin2=red |barend2=red
|2=†''[[Tritemnodon]]''
|3=†'''[[Teratodontidae]]''' |barbegin3=green
|barend4=green |grouplabel4x=†'''[[Hyainailouroidea]]'''
|4={{clade
|thickness=2 |color=green |style1=background-color:#74C365 |grouplabel3=†'''Hyainailouridae'''
|1={{clade
|thickness=2 |label1=&nbsp;†'''[[Apterodontinae]]'''&nbsp; |color=green
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Apterodon]]''&nbsp; |color=green
|1={{clade
|thickness=2 |color=green
|1={{clade
|thickness=2 |color=green
|1={{clade
|thickness=2 |color=green
|1={{clade
|thickness=2 |color=green
|1=†''Apterodon altidens'' |barbegin1=brown
|2=†''Apterodon gaudryi'' |bar2=brown
|label3=&nbsp;'''?'''&nbsp; |state3=dotted
|3=†''Apterodon rauenbergensis'' |bar3=brown
}}
|2=†''Apterodon langebadreae'' |bar2=brown
}}
|2=†''Apterodon macrognathus'' |bar2=brown
}}
|label2=&nbsp;'''?'''&nbsp; |state2=dotted
|2=†''Apterodon'' sp. (Dur At-Talah escarpment, Libya) |bar2=brown
|3=†''Apterodon saghensis'' |bar3=brown
}}
|label2=&nbsp;†''[[Quasiapterodon]]''&nbsp;
|2=†''Quasiapterodon minutus'' |bar2=brown
}} }}
|style2=background-color:#74C365 |bar2=brown
|2={{clade
|thickness=2 |label1=&nbsp;†''[[Maocyon]]''&nbsp; |color=green |grouplabel1=†'''''Maocyon''/''Orienspterodon'' clade'''
|1=†''Maocyon peregrinus'' |barbegin1=gray
|label2=&nbsp;†''[[Orienspterodon]]''&nbsp;
|2=†''Orienspterodon dahkoensis'' |barend2=gray
}}
|3={{clade
|thickness=2 |color=green |style1=background-color:#96C8A2 |bar1=brown
|1={{clade
|thickness=2 |color=green |grouplabel1=†'''''Akhnatenavus'' clade''' |caption=†'''[[Hyainailourinae]]''' |captionstyle=background-color:#96C8A2;text-align:center;font-weight:bold;padding:2px 10px 1px 1px;
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Hemipsalodon]]''&nbsp; |color=green |barbegin1=magenta
|1={{clade
|thickness=2 |color=green
|1=†''Hemipsalodon grandis''
|2=†''Hemipsalodon viejaensis''
}}
|label2=&nbsp;†''[[Ischnognathus]]''&nbsp; |sublabel2=&nbsp;'''?'''&nbsp; |state2=dotted |bar2=magenta
|2=†''Ischnognathus savagei''
}}
|2={{clade
|thickness=2 |label1=&nbsp;†''[[Akhnatenavus]]''&nbsp; |color=green
|1={{clade
|thickness=2 |color=green
|1=†''Akhnatenavus leptognathus'' |bar1=magenta
|2=†''Akhnatenavus nefertiticyon'' |barend2=magenta
}} }} }}
|style2=background-color:#96C8A2 |bar2=brown
|2=†[[Pterodon sp. (BC 15’08)|''"Pterodon"'' sp. (''BC 15’08'')]]
|style3=background-color:#96C8A2 |bar3=brown
|3=†[[Hyainailourinae sp. (UON 84-359)|Hyainailourinae sp. (''UON 84-359'')]]
|style4=background-color:#96C8A2 |bar4=brown
|4=†[[Hyainailourinae sp. C (DPC 9243 & DPC 10315)|Hyainailourinae sp. C (''DPC 9243'' & ''DPC 10315'')]]
|style5=background-color:#96C8A2 |bar5=brown
|5=†[[Hyainailourinae sp. D (DPC 6545)|Hyainailourinae sp. D (''DPC 6545'')]]
|style6=background-color:#96C8A2 |bar6=brown
|6={{clade
|thickness=2 |color=green |style=background-color:#ACE1AF |grouplabel1=†'''''Pterodon'' clade'''
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Kerberos (mammal)|Kerberos]]''&nbsp; |color=green
|1=†''Kerberos langebadreae'' |barbegin1=black
|2=†''[[Pterodon syrtos|"Pterodon" syrtos]]'' |bar2=black
}}
|2={{clade
|thickness=2 |label1=&nbsp;†'''[[Pterodon (mammal)|Pterodontina]]'''&nbsp; |color=green
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Pterodon (mammal)|Pterodon]]''&nbsp; |color=green
|1=†''Pterodon dasyuroides'' |barend1=black
}} }} }}
|7={{clade
|thickness=2 |color=green |style1=background-color:#96C8A2 |bar1=brown
|1={{clade
|thickness=2 |color=green
|1={{clade
|thickness=2 |color=green
|1=†[[Hyainailourinae sp. A (DPC 6555)|Hyainailourinae sp. A (''DPC 6555'')]]
|style2=background-color:#ACE1AF
|2=†''[[Pterodon africanus|"Pterodon" africanus]]''
|label3=&nbsp;†''[[Parapterodon]]''&nbsp; |style3=background-color:#ACE1AF
|3=†''Parapterodon lostangensis''
}}
|style2=background-color:#ACE1AF
|2=†[[Pterodon sp. (DPC 5036)|''"Pterodon"'' sp. (''DPC 5036'')]]
|style3=background-color:#ACE1AF
|3=†''[[Pterodon phiomensis|"Pterodon" phiomensis]]''
|footer=†'''[[Hyainailourini]]''' |footerstyle=background-color:#ACE1AF;text-align:center;font-weight:bold;padding:2px 10px 1px 1px;
}}
|style2=background-color:#96C8A2 |bar2=brown
|2={{clade
|thickness=2 |label1=&nbsp;†'''[[Paroxyaena|Paroxyaenini]]'''&nbsp; |color=green
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Paroxyaena]]''&nbsp; |color=green
|1={{clade
|thickness=2 |color=green
|1=†''Paroxyaena galliae''
|2=†''Paroxyaena pavlovi''
}} }} }}
|3={{clade
|thickness=2 |color=green |style1=background-color:#96C8A2 |bar1=brown
|1={{clade
|thickness=2 |style1=background-color:#ACE1AF |color=green
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Sectisodon]]''&nbsp; |color=green
|1={{clade
|thickness=2 |color=green
|1=†''Sectisodon markgrafi''
|2=†''Sectisodon occultus''
}} }}
|style2=background-color:#ACE1AF
|2={{clade
|thickness=2 |label1=&nbsp;†''[[Falcatodon]]''&nbsp; |color=green
|1=†''Falcatodon schlosseri''
}}
|style3=background-color:#ACE1AF
|3={{clade
|thickness=2 |label1=&nbsp;†''[[Exiguodon]]''&nbsp; |color=green
|1=†''Exiguodon pilgrimi''
|2={{Uuline|†''[[Isohyaenodon|Isohyaenodon zadoki]]''}}
}}
|4={{clade
|thickness=2 |color=green |style1=background-color:#ACE1AF
|1={{clade
|thickness=2 |color=green |footer={{Uline|c=lime|†''[[Hyainailouros]]''}} |footerstyle=background-color:#ACE1AF;text-align:center;font-weight:bold;padding:2px 10px 1px 1px;
|1={{clade
|thickness=2 |caption={{Uuline|†''[[Isohyaenodon]]''&nbsp;&nbsp;(†[[Isohyaenodon]]tina)}} |captionstyle=background-color:#ACE1AF;text-align:center;font-weight:bold;padding:2px 10px 1px 1px; |color=green
|1={{Uuline|†''[[Isohyaenodon|Isohyaenodon andrewsi]]''}}
|2={{clade
|thickness=2 |label1=&nbsp;†''[[Sivapterodon]]''&nbsp; |color=green
|1=†''Sivapterodon lahirii''
|2={{Uline|c=lime|†''[[Hyainailouros|Hyainailouros bugtiensis]]''}}
}} }}
|2={{Uline|c=lime|†''[[Hyainailouros|Hyainailouros napakensis]]''}}
}}
|2={{clade
|thickness=2 |color=green |style1=background-color:#ACE1AF
|1={{Uline|c=lime|†''[[Hyainailouros|Hyainailouros sulzeri]]''}}
|2=†[[Hyainailourinae sp. (GSN AD 100’96)|Hyainailourinae sp. (''GSN AD 100’96'')]]
}}
|3={{clade
|thickness=2 |label1=&nbsp;†''[[Simbakubwa]]''&nbsp; |color=green
|1=†''Simbakubwa kutokaafrika''
}} }}
|5={{clade
|thickness=2 |label1=&nbsp;†'''[[Leakitherium|Leakitheriini]]'''&nbsp; |color=green
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Leakitherium]]''&nbsp; |color=green
|1=†''Leakitherium hiwegi''
}}
|label2=&nbsp;†''[[Megistotherium]]''&nbsp;
|2=†''Megistotherium osteothlastes''
}}
|6={{clade
|thickness=2 |label1=&nbsp;†''[[Mlanyama]]''&nbsp; |color=green
|1=†''Mlanyama sugu''
|label2=&nbsp;†'''[[Metapterodon]]tini'''&nbsp;
|2={{clade
|thickness=2 |label1=&nbsp;†''[[Metapterodon]]''&nbsp; |color=green
|1={{clade
|thickness=2 |color=green
|1=†''Metapterodon kaiseri''
|2=†''Metapterodon stromeri''
}} }} }} }}
|style2=background-color:#96C8A2
|2={{clade
|thickness=2 |label1=&nbsp;†''[[Pakakali]]''&nbsp; |color=green
|1=†''Pakakali rukwaensis'' |barend1=brown
}}
|3=†'''[[Prionogalidae]]'''
}} }} }} }} }} }} }} }} }} }} }} }}

==See also==
* [[Mammal classification]]
* [[Hyainailouroidea]]


==References==
==References==

Latest revision as of 18:16, 29 October 2024

Hyainailouridae
Temporal range: 47.8–11.3 Ma 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
Synonyms
  • Hyaenaelurinae (Pilgrim, 1932)
  • Pterodontidae (Polly, 1996)[2]
  • Pterodontinae (Polly, 1996)

Hyainailouridae ("hyena-cats") is a family of extinct predatory mammals within the 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.

General characteristics

[edit]

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

[edit]

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.[citation needed] 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

[edit]

Relations

[edit]

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

[edit]
  • Family: †Hyainailouridae (Pilgrim, 1932)
    • (unranked): †Maocyon/Orienspterodon clade
      • Genus: †Maocyon (Averianov, 2023)
        • Maocyon peregrinus (Averianov, 2023)
      • Genus: †Orienspterodon (Egi, 2007)
        • Orienspterodon dahkoensis (Chow, 1975)
        • Orienspterodon mianchiensis (Sheng, Bi & Li, 2024)
    • 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)
      • 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 (Ginsburg, 1980)
        • Genus: †Exiguodon (Morales & Pickford, 2017)
          • Exiguodon pilgrimi (Savage, 1965)
        • Genus: †Falcatodon (Morales & Pickford, 2017)
          • Falcatodon schlosseri (Holroyd, 1999)
        • Genus: †Hyainailouros (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 (Lavrov, 1999)
          • Genus: †Isohyaenodon (Savage, 1965)
            • Isohyaenodon andrewsi (Savage, 1965)
            • Isohyaenodon zadoki (Savage, 1965)
        • (unranked): †Pterodon clade
        • Incertae sedis:
      • 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)

References

[edit]
  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.