Editing Hyena (section)

==Genera of the Hyaenidae (extinct and recent)==
[[File:Crocuta crocuta.jpg|thumb|A [[spotted hyena]] of subfamily Hyaeninae]]
The list follows McKenna and Bell's ''Classification of Mammals'' for prehistoric genera (1997)<ref name="McKenna & Bell">Malcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'', Columbia University Press, New York 1997, 631 Seiten, {{ISBN|0-231-11013-8}}</ref> and Wozencraft (2005) in Wilson and Reeders ''[[Mammal Species of the World]]'' for extant genera.<ref name="MSW3">{{MSW3 Wozencraft|pages=532–548}}</ref> The percrocutids are, in contrast to McKenna and Bell's classification, not included as a subfamily into the Hyaenidae, but as the separate family [[Percrocutidae]], although they are generally grouped as sister-taxa to hyenas.<ref>{{cite journal| year=2013| title=Skull Shape Evolution In Durophagous Carnivorans| journal=Evolution| volume=67| issue=7| pages=1975–93| doi=10.1111/evo.12059 | last1=Figueirido| first1=Borja| last2=Jack Tseng| first2=Zhijie| last3=Martín-Serra| first3=Alberto| pmid=23815654| s2cid=23918004 | doi-access=free| hdl=10630/32909| hdl-access=free}}</ref> However recently, many experts have began reconsidering them as members of Hyaenidae.<ref>{{Cite journal |last=Pérez-Claros |first=Juan A. |date=26 April 2023 |title=An ecomorphological characterization of the percrocutoid hyaenids: a multivariate approach using postcanine dentition |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2023.2197972 |journal=Journal of Vertebrate Paleontology |volume=42 |issue=5}}</ref><ref name="Xiong 2022">{{Cite journal |last=Xiong |first=Wuyang |date=1 May 2022 |title=New species of Percrocuta (Carnivora, Hyaenidae) from the early middle Miocene of Tongxin, China |url=https://doi.org/10.1080/08912963.2022.2067757 |journal=Historical Biology |volume=35 |issue=5 |pages=799–820 |doi=10.1080/08912963.2022.2067757 |issn=0891-2963 |s2cid=248627038 |url-access=subscription}}</ref><ref>{{Cite journal |last=Wuyang |first=Xiong |date=October 2019 |title=Basicranial morphology of Late Miocene Dinocrocuta gigantea (Carnivora: Hyaenidae) from Fugu, Shaanxi |url=https://www.researchgate.net/publication/336578352_Basicranial_morphology_of_Late_Miocene_Dinocrocuta_gigantea_Carnivora_Hyaenidae_from_Fugu_Shaanxi |journal=Vertebrata PalAsiatica |volume=57 |issue=4 |pages=274–307}}</ref><ref>{{Cite journal |last=Barret |first=Paul Z. |last2=Hopkins |first2=Samatha S. B. |date=2024 |title=Mosaic evolution underlies feliform morphological disparity |url=https://royalsocietypublishing.org/doi/10.1098/rspb.2024.0756 |journal=Proc. R. Soc. B |volume=291 |issue=2028}}</ref> Furthermore, the living brown hyena and its closest extinct relatives are not included in the genus ''Pachycrocuta'', but in the genus ''Parahyaena''. However, some research has suggested ''Parahyaena'' may be synonymous with ''Pachycrocuta'', making the brown hyena the only extant member of this genus.<ref>{{cite journal|first1=J. A|last1=Pérez-Claros|year=2024|title= Unravelling the origin of the brown hyena (Parahyena brunnea) and its evolutionary and paleoecological implications for the Pachycrocuta lineage |journal=Palaeontologia Electronica|doi=10.26879/1372 |doi-access=free}}</ref>

* '''Family Hyaenidae'''
** '''Subfamily ''Incertae sedis'''''
*** †''[[Tongxinictis]]''<ref name=CocaOrtega2019>{{cite journal|last1=Coca-Ortega|first1=C.|last2=Pérez-Claros|first2=J.A.|date=January 2019|title=Characterizing ecomorphological patterns in hyenids: a multivariate approach using postcanine dentition|pages=6:e6238|journal=PeerJ|volume=6|doi=10.7717/peerj.6238|pmid=30648005|pmc=6330948|doi-access=free }}</ref> (Middle Miocene of Asia)
** †'''Subfamily Ictitheriinae'''
*** †''[[Herpestides]]'' (Early Miocene of Africa and Eurasia)
*** †''[[Plioviverrops]]'' (including ''Jordanictis'', ''Protoviverrops'', ''Mesoviverrops''; Early Miocene to Early Pliocene of Europe, Late Miocene of Asia)
*** †''[[Ictitherium]]'' (=''Galeotherium''; including ''Lepthyaena'', ''Sinictitherium'', ''Paraictitherium''; Middle Miocene of Africa, Late Miocene to Early Pliocene of Eurasia)
*** †''[[Thalassictis]]'' (including ''Palhyaena'', ''Miohyaena'', ''Hyaenictitherium'', ''Hyaenalopex''; Middle to Late Miocene of Asia, Late Miocene of Africa and Europe)
*** †''[[Hyaenotherium]]'' (Late Miocene to Early Pliocene of Eurasia)
*** †''[[Miohyaenotherium]]''<ref name=Werdelin1996>{{cite journal|last1=Werdelin|first1=Lars|date=March 1996|title=Community-wide character displacement in Miocene hyaenas|journal=Lethaia|volume=29|issue=1|pages=97–106|doi=10.1111/j.1502-3931.1996.tb01843.x|bibcode=1996Letha..29...97W }}</ref>(Late Miocene of Europe)
*** †''[[Lycyaena]]'' (Late Miocene of Eurasia)
*** †''[[Tungurictis]]''<ref name=Wang2020>{{cite journal|last1=Wang|first1=X.|last2=Tsang|first2=Z.J.|display-authors=etal|date=February 2020|title=A new species of Tungurictis Colbert, 1939 (Carnivora, Hyaenidae) from the middle Miocene of Junggar Basin, northwestern China and the early divergence of basal hyaenids in East Asia|journal=Geodiversitas|volume=42|issue=3|pages=29–45|doi=10.5252/geodiversitas2020v42a3|doi-access=free }}</ref> (Middle Miocene of Africa and Eurasia)
*** †''[[Protictitherium]]'' (Middle Miocene of Africa and Asia, Middle to Late Miocene of Europe)
** '''Subfamily Hyaeninae'''
*** †''[[Palinhyaena]]''<ref name=Turner2008>{{cite journal|last1=Turner|first1=A.|last2=Antón|first2=M.|last3=Werdelin|first3=L.|date=September 2008|title=Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe|journal=Geobios|volume=41|issue=5|pages=677–687|doi=10.1016/j.geobios.2008.01.001|bibcode=2008Geobi..41..677T }}</ref> (Late Miocene of Asia)
*** †''[[Ikelohyaena]]''<ref name=Tseng2011>{{cite journal|last1=Tseng|first1=Z.J.|last2=Stynder|first2=D.|date=March 2011|title=Mosaic functionality in a transitional ecomorphology: skull biomechanics in stem Hyaeninae compared to modern South African carnivorans|journal=Biological Journal of the Linnean Society|volume=102|issue=3|pages=540–559|doi=10.1111/j.1095-8312.2010.01602.x|doi-access=free }}</ref> (Early Pliocene of Africa)
*** ''[[Hyaena]]'' (=''Euhyaena'',=''Parahyaena''; including [[striped hyena]], ''Pliohyaena'', ''Pliocrocuta'', ''Anomalopithecus'') Early Pliocene (?Middle Miocene) to Recent of Africa, Late Pliocene (?Late Miocene) to Late Pleistocene of Europe, Late Pliocene to recent in Asia
*** ''[[Parahyaena]]''  (=''Hyaena''; [[brown hyena]] Pliocene to recent of Africa)
*** †''[[Hyaenictis]]''<ref name=Vinuesa2016>{{cite journal|last1=Vinuesa|first1=V.|last2=Madurell-Malapeira|first2=J.|display-authors=etal|date=April 2016|title=A new skull of ''Hyaenictis'' Gaudry, 1861 (Carnivora, Hyaenidae) shows incipient adaptations to durophagy|journal=Journal of Mammalian Evolution|volume=24|issue=2|pages=207–219|doi=10.1007/s10914-016-9334-0|s2cid=23453043|url=http://nrm.diva-portal.org/smash/get/diva2:926393/FULLTEXT02 }}</ref> (Late Miocene of Asia?, Late Miocene of Europe, Early Pliocene (?Early Pleistocene) of Africa)
*** †''[[Leecyaena]]''<ref name=Turner2008/> (Late Miocene and/or Early Pliocene of Asia)
*** †''[[Chasmaporthetes]]'' (=''Ailuriaena''; including ''Lycaenops'', ''Euryboas''; Late Miocene to Early Pleistocene of Eurasia, Early Pliocene to Late Pliocene or Early Pleistocene of Africa, Late Pliocene to Early Pleistocene of North America)
*** †''[[Pachycrocuta]]'' (Pliocene and Pleistocene of Eurasia and Africa)
*** †''[[Adcrocuta]]'' (Late Miocene of Eurasia)
*** ''[[Spotted hyena|Crocuta]]'' (=''Crocotta''; including ''Eucrocuta''; [[spotted hyena]] and [[cave hyena]]. Late Pliocene to recent of Africa, Late Pliocene to Late Pleistocene of Eurasia)
** '''Subfamily Protelinae'''
*** †''[[Gansuyaena]]''<ref>{{cite journal |last1=Tseng |first1=Z. Jack |date=2022 |title=A new aardwolf-line fossil hyena from Middle and Late Miocene deposits of Linxia Basin, Gansu, China |url=https://www.researchgate.net/publication/361162806 |journal=Vertebrata PalAsiatica |volume=60 |issue=2 |pages=81–116 |doi=10.19615/j.cnki.2096-9899.211025 |access-date=}}</ref>
*** ''[[Proteles]]'' (=''Geocyon''; [[aardwolf]]. Pleistocene to Recent of Africa)

===Phylogeny===
The following cladogram illustrates the phylogenetic relationships between extant and extinct hyaenids based on the morphological analysis by Werdelin & Solounias (1991),<ref name="Werdelin1991">{{cite journal|last1=Werdelin|first1=L.|last2=Solounias|first2=Nikos|title=The Hyaenidae: taxonomy, systematics and evolution.|url=http://foreninger.uio.no/ngf/FOS/pdfs/F&S_30.pdf|journal=Fossils and Strata|volume=30|pages=1–104|year=1991|doi=10.18261/8200374815-1991-01 |isbn=8200374815 }}</ref> as updated by Turner et al. (2008).<ref>{{cite journal|title=Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe|first1=Alan|last1=Turner|first2=Mauricio|last2=Antón|first3=Lars|last3=Werdelin|journal=Geobios|volume=41|issue=5|year=2008|pages=677–687|doi=10.1016/j.geobios.2008.01.001|bibcode=2008Geobi..41..677T }}</ref>
{{Clade
 |style=font-size:80%; line-height:85%;background-color:#eeeeff
 |caption=Evolution of hyaenas
 |captionstyle=background-color:#8899ff;padding:10px;font-weight:bold;
 |footer=Phylogenic relationships based on morphological characteristics, after Werdelin & Solounias (1991) and Turner et al (2008)
 |footerstyle=background-color:#eeeeee;text-align:center;
 |1={{clade
  |style=background-color:#ccffcc;
  |label1=
  |1=''[[Protictitherium|Protictitherium crassum]]''
  |2=''"Protictitherium" cingulatum''
  |3=''"Protictitherium" intermedium''
  |4=''"Protictitherium" llopisi''
  |5=''"Protictitherium" punicum''
  |6=''"[[Protictitherium|Protictitherium" gaillardi]]''
  |state7=dotted
  |7=''"Protictitherium" sumegense''
  |state8=dotted
  |8=''"Protictitherium" csakvarense''
  |style9=background-color:#eeeeff;
  |9={{clade
    |label1=
    |1={{clade
        |label1=''[[Plioviverrops]]''
        |style1=background-color:#ccffcc;
        |1={{clade
           |grouplabel1=Civet/mongoose-like insectivore-omnivores 
           |grouplabelstyle1=font-size:12px;font-weight:bold;color:darkgreen;text-align:right;
           |label1=
           |1=''Plioviverrops gervaisi''
           |2={{clade
              |1=''Plioviverrops orbignyi''
              |2={{Clade
                 |label1=
                 |1=''Plioviverrops guerini''
                 |2=''Plioviverrops faventinus''
                 |3=''Plioviverrops gaudryi''
                  }} 
               }} 
            }}
         }}
     |2={{clade
        |label1=
        |1=''Tungurictis spocki''
        |2={{clade
           |label1=
           |1=''[[Thalassictis|Thalassictis robusta]]''
           |2=''"Thalassictis" certa''
           |3=''"Thalassictis" montadai''
           |4=''"Thalassictis" proava''
           |5=''"Thalassictis" sarmatica''
           |6=''"Thalassictis" spelaea''
           |7={{clade
              |label1=
              |1=''Tongxinictis primordialis''
              |2={{clade
                 |label1=
                 |1={{clade
                    |label1=''[[Proteles]]''
                    |style1=background-color:#ddeeff;
                    |1={{clade
                       |label1=
                       |1='''''[[Proteles cristatus]]''''' (aardwolf) [[File:The life of animals (Colored Plate 4) (proteles cristatus).jpg|50 px]]
                       |2=''Proteles amplidentus''
                        }}
                     }}
                 |2={{clade
                    |label1=
                    |1={{clade
                       |label1=''[[Ictitherium]]''
                       |style=background-color:#ddddff;
                       |1={{clade
                          |label1=
                          |1=''[[Ictitherium|Ictitherium viverrinum]]''
                          |2=''Ictitherium ebu''
                          |grouplabel2=Jackal-like hyaenas 
                          |grouplabelstyle2=font-size:12px;font-weight:bold;color:darkcyan;text-align:right;
                          |3={{clade
                             |label1=
                             |1={{clade
                                |label1=
                                |1=''Ictitherium  tauricum''
                                |2=''Ictitherium ibericum''
                                 }}
                             |2={{clade
                                |label1=
                                |1=''Ictitherium kurteni''
                                |2={{clade
                                   |1=''Ictitherium intuberculatum''
                                   |2=''Ictitherium pannonicum''
                                    }}
                                 }} 
                              }} 
                           }} 
                        }}
                    |2={{clade 
                       |label1=
                       |1=''Miohyaenotherium bessarabicum''
                       |2={{clade
                          |1={{clade
                             |label1=''[[Hyaenotherium]]''
                             |1={{clade
                                |1=''Hyaenotherium wongii''
                                |2=''Hyaenictitherium hyaenoides''
                                |3=''"Hyaenictitherium" pilgrimi''
                                |4=''"Hyaenictitherium" parvum''
                                |5=''"Hyaenictitherium" namaquensis''
                                |6=''"Hyaenictitherium" minimum''
                                 }}
                              }}
                          |style2=background-color:#ccccff;
                          |2={{clade
                             |1={{clade
                                |label1=''[[Lycyaena]]''
                                |1={{clade
                                   |grouplabel2=Cursorial hunting hyaenas 
                                   |grouplabelstyle2=font-size:12px;font-weight:bold;color:darkblue;text-align:right;
                                   |1=''Lycyaena chaeretis''
                                   |2=''Lycyaena dubia''
                                   |3={{clade
                                      |1=''Lycyaena macrostoma''
                                      |2=''Lycyaena crusafonti''
                                       }}
                                    }} 
                                 }}
                             |2={{clade
                                |1={{clade
                                   |label1=''[[Hyaenictis]]''
                                   |1={{clade
                                      |1=''Hyaenictis graeca''
                                      |2=''Hyaenictis almerai''
                                      |3=''Hyaenictis hendeyi''
                                       }}
                                    }}
                                 |2={{clade
                                    |1={{clade
                                       |label1=''[[Lycyaenops]]''
                                       |1={{clade
                                          |1=''Lycyaenops rhomboideae''
                                          |2=''Lycyaenops silberbergi''
                                           }}
                                        }}
                                    |2={{clade
                                       |label1=''[[Chasmaporthetes]]''
                                       |sublabel1=(running hyaenas)
                                       |1={{clade
                                          |1={{clade
                                             |1=''Chasmaporthetes exitelus''
                                             |2=''Chasmaporthetes bonisi''
                                              }}
                                          |2={{clade
                                             |1=''Chasmaporthetes borissiaki''
                                             |2={{clade
                                                |1={{clade
                                                   |1=''Chasmaporthetes lunensis''
                                                   |2=''Chasmaporthetes melei''
                                                    }}
                                                |2={{clade
                                                   |1={{clade
                                                      |1=''Chasmaporthetes ossifragus''
                                                      |2=''Chasmaporthetes'' sp. Florida
                                                       }}
                                                   |2={{clade
                                                      |1=''Chasmaporthetes nitidula''
                                                      |2=''Chasmaporthetes australis''
                                                       }} }} }} }} }}
                                       |style2=background-color:#ddccff
                                       |label2=[[Hyaeninae]]
                                       |sublabel2=(bone-cracking hyenas)
                                       |2={HYAENINAE}
                                         }} }} }} }} }} }} }} }} }} }} }} }} }} 
|targetA={HYAENINAE}
|subcladeA={{clade
  |caption=Bone-cracking hyaenas
  |captionstyle=font-size:12px;font-weight:bold;color:darkred;text-align:right;padding:5px 20px 0px 0px;
  |1=''Metahyaena confector''
  |2=''Palinhyaena reperta''
  |3={{clade
     |1=Hyaenid sp. E Langebaar
     |2={{clade
        |1=''Belbus beaumonti''
    <!--|1=''Leecyaena lycyaenoides''
        |2=''"Leecyaena" bosei'' -->
        |style2=background-color:#eeccff;
        |2={{clade
           |1={{clade
              |1={{clade
                 |1=''Hyaena abronia''
                 |2='''''[[Hyaena hyaena]]''''' (striped hyaena) [[File:Hyaena striata - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|50 px]]
                  }}
              |2={{clade
                 |1=''Parahyaena howelli''
                 |2='''''[[Parahyaena brunnea]]''''' (brown hyaena) [[File:Hyaena fusca (white background).jpg|50 px]]
                  }} }}
           |2={{clade
              |1=''Pliocrocuta perrieri''
              |2={{clade
                 |1=''[[Pachycrocuta|Pachycrocuta brevirostris]]'' (giant hyaena) [[File:Pachycrocuta brevirostris restoration.jpg|70 px]]
             <!--|2=''"Pachycrocuta"'' bellax''-->
                 |2={{clade
                    |1=''[[Adcrocuta|Adcrocuta eximia]]'' [[File:Adcrocuta eximia restoration.jpg|60 px]]
                    |2=''Allohyaena kadici''
                    |3={{clade
                       |1='''''[[Crocuta crocuta]]''''' (spotted hyaena) [[File:Spotted hyena (Crocuta crocuta) by Connor Ashbridge.png|50 px]]
                       |2=''Crocuta eturono''
                   <!--|2=''Crocuta sivalensis''
                       |3=''Crocuta dietrichi'' -->
}} }} }} }} }} }} }} }} }}

A more recent molecular analysis agrees on the phylogenetic relationship between the four extant hyaenidae species (Koepfli ''et al'', 2006<ref name="Koepfli2006">{{cite journal|last1=Koepfli|first1=K.-P.|last2=Jenks|first2=S. M.|last3=Eizirik|first3=E.|last4=Zahirpour|first4=T.|last5=Van Valkenburgh|first5=B.|last6=Wayne|first6=R. K.|title=Molecular systematics of the Hyaenidae: Relationships of a relictual lineage resolved by a molecular supermatrix|journal=Molecular Phylogenetics and Evolution|volume=38|issue=3|date=2006|pages=603–620|doi=10.1016/j.ympev.2005.10.017|pmid=16503281|bibcode=2006MolPE..38..603K }}</ref>).

{{clade|style=font-size:90%; line-height:100%;
|label1='''Hyaenidae'''
|1={{clade
  |label1=
  |1={{clade
     |label1=
     |1=''[[Aardwolf|Proteles cristatus]]'' (aardwolf) [[File:The life of animals (Colored Plate 4) (proteles cristatus).jpg|60 px]]
     }}
  |2={{clade
     |label1=
     |1={{clade |label1=|1=''[[Spotted hyena|Crocuta crocuta]]'' (spotted hyena) [[File:Spotted hyena (Crocuta crocuta) by Connor Ashbridge.png|80 px]] }}
     |2={{clade
        |1=''[[Striped hyena|Hyaena hyaena]]'' (striped hyena) [[File:Hyaena striata - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|70 px]]
        |2=''[[Brown hyena|Parahyaena brunnea]]'' (brown hyena) [[File:Hyaena fusca (white background).jpg|65 px]] 
         }} }} }} 
         
}}