After Kurtén & Werdelin, 1988, Werdelin & Salounias, 1991, and Werdelin & Lewis, 2000
<==o Hyaenidae Gray, 1829 sensu Werdelin & Salounias, 1991 (hyeenaeläimet; hyaenas) |?- †“Allohyaena” kadici |-- †Protictitherium crassum |-- †“Protictitherium” cingulatum |-- †“Protictitherium” intermedium |-- †“Protictitherium” llopisi |-- †“Protictitherium” punicum |-- †“Protictitherium” gaillardi |-- †“Protictitherium” csakvarensis |-- †“Protictitherium” sumegense `--+--o †Plioviverrops | |?- †P. faventinus | |?- †P. collectus | |-- †P. gervaisi | `--+-- †P. gaudryi | `--+-- †P. guerini | `-- †P. orbigyi |--o Proteles I. Geoffroy Saint-Hilaire, 1824 (Aardwolves; maasudet) [Protelinae Flower, 1869] | |-- †P. amplidentus | `--o P. cristatus (Sparrman, 1783) (Aardwolf; maasusi) | |-- P. c. cristatus (<South Africa>) | `-- P. c. septentrionalis (<East Africa>) `--+-- †Tongxinictis primordialis |-- †Tungurictis spocki Colbert, 1939 `--+--o “Ictitheriinae” | `--o †Ictitherium Wagner, 1848 [Galeotherium; Hyaenalopex; Palhyaena; Thalassictis] | |-- †I. viverrinum Roth & Wagner, 1854 | |-- †I. adroveri Crusafont & Petter, 1969 | |-- †I. arkelaisi Esu & Kotsakis, 1980 [actually a species of Hyaenictitherium (de Bonis et al., 2005)?] | |-- †I. ebu Werdelin | |-- †I. tauricum Borissiak, 1915 | |-- †I. ibericum Meladze, 1967 | |-- †I. ebu Werdelin, 2003 | `--+-- †I. kurteni | `-- †I. pannonicum Kretzoi, 1952 `--o “Hyaeninae” Gray, 1829 (hyeenat) |--o †Thalassictis [monophyly questionable] | |-- †T. robusta | |-- †“T.” certa | |-- †“T.” montadai | |-- †“T.” proava | |-- †“T.” sarmatica [=? Ictitherium sarmaticum Pavlow, 1908] | |-- †“T.” spelaea | |-- †“T.” taurica [=? †Ictitherium tauricum Borissiak, 1915] | |-- †“T.” hipparionum | `-- †“T.” praecurrens `--+-- †Hyaenotherium wongii (Zdansky, 1924) [Hyaenictitherium] `--+-- †Miohyaenotherium bessarabicum Semenov, 1989 [Miohyaena] `--+--o †Hyaenictitherium Kretzoi, 1938 [Hyaenictis] [monophyly questionable] | |-- †H. hyaenoides | |-- †“H.” pilgrimi | |-- †“H.” parvum Khomenko | |-- †“H.” namaquensis (Stromer, 1931) | |-- †“H.” intuberculatum (Ozansoy, 1965) [†Ictitherium intuberculatum Ozansoy, 1965] | |-- †“H.” venator Semenov, 1989 | |-- †“H.” barbarum Geraads, 1997 | `-- †“H.” minimum de Bonis, Peigné, Likius, Mackaye, Vignaud & Brunet, 2005 `--+--o †“Chasmaportheteini” | |--o †Lycyaena | | |-- †L. chaeretis | | |-- †L. dubia | | |-- †L. macrostoma | | `-- †L. crusafonti | `--+--o †Hyaenictis Gaudry, 1861 | | |-- †H. graeca | | |-- †H. almerae | | `-- †H. sp. | `--o †Chasmaporthetes Hay, 1921 [Ailurena Stirton & Christian, 1940; Lycyaena, Euryboas Schaub, 1941] | |?- †C. progressus | |?- †C. melei | |?- †C. kani Caliano & Frailey, 1977 [= Chasmaporthetes lunensis?] [incl. C. progressus?] | |?- †C. johnstoni [Ailurena johnstoni] | |-- †C. exitelus Kurtén & Werdelin, 1988 | `--+-- †C. borissiaki (Khomenko, 1932) sensu Galiano & Frailey, 1977 [Hyaena borissiaki Khomenko, 1932] | `--+--o †C. lunensis (Del Campana, 1914) [Lycyaena lunensis Del Campana, 1914; Lycyaenops lunensis Kretzoi, 1938; Euryboas bielawskyi Schaub, 1941; Chasmaporthetes kani Caliano & Frailey, 1977] | | |-- †C. l. lunensis (Del Campana, 1914) | | `-- †C. l. honanensis (Zdansky, 1924) | |--+-- †C. ossifragus Hay, 1921 | | `-- †C. sp. [Kurtén & Werdelin, 1988] | `--+-- †C. nitidula | `-- †C. australis `--o “Hyaenini” |?- †Metahyaena confector Viranta & Werdelin, 2003 |-- †Palinhyaena reperta Qiu, Huang & Guo, 1979 [incl. Palinhyaena imbricata] `--+-- †Ikelohyaena abronia Werdelin & Solounias, 1991 `--+-- †Belbus beaumonti |-- †Hyaenidae genera et species indet. `--+-- †Leecyaena lycyaenoidea Young & Liu, 1948 |-- †“Leecyaena” bosei `--+-- Parahyaena brunnea (Thunberg, 1820) (Brown hyaena; ruskohyeena) [Hyaena brunnea] `--+-- Hyaena hyaena (Linnaeus, 1758) (Striped hyaena; juovahyeena) [Plesiocrocuta] `--+-- †Pliocrocuta perrieri Kretzoi, 1938 [‘hyena’ Baker, 1835; Hyaena sivalensis Falconer, 1867; Crocuta sivalensis (Falconer, 1867) Lydekker, 1884] `==o †Pachycrocuta Kretzoi, 1938 |?- †Pachycrocuta robusta |?- †Pachycrocuta pyrenica |-- †Pachycrocuta brevirostris |-- †“Pachycrocuta” bellax `--+-- †Adcrocuta eximia (Roth & Wagner, 1854) Kretzoi, 1938 [Hyaena eximia Roth & Wagner, 1854; Hyaena variabilis Zdansky, 1924] `--o Crocuta Kaup, 1828 (spotted hyaenas; täplähyeenat) |-- †C. macrodonta |-- †C. eximia |-- †C. honanensis Zdansky, 1924 [= C. c. honanensis] |-- †C. dietrichi Petter & Howell, 1989 `--o C. crocuta (Erxleben, 1777) [Canis crocuta Erxleben, 1777; Crocuta crocuta Kaup, 1828; Hyaena ultima Matsumoto, 1915; Hyaena ultima Zdansky, 1927; Hyaena ultima Pei, 1934; Crocuta ultima Huang, 1989] (spotted hyaena; täplähyeena) |?- †C. c. honanensis |-- †C. c. ultima |-- †C. c. spelaeus (cave hyaena, Eurasian spotted hyaena; luolahyeena) `-- C. c. crocuta (spotted hyaena; African spotted hyaena; täplähyeena)
Reference(s):
- Carroll, R. L., 1988: Vertebrate paleontology and evolution.
–W. H. Freeman and company, New York, 1988, 698 - Carroll, R. L., 1988: Appendix. 594-648
in Carroll, R. L., 1988: Vertebrate paleontology and evolution.
–W. H. Freeman and company, New York, 1988, 698 - de Bonis, L., Peigné, S., Likius, A., Mackaye, H. T., Vignaud, P. & Brunet, M., 2005: Hyaenictitherium minimum, a new ictithere (Mammalia, Carnivora, Hyaenidae) from the Late Miocene of Toros-Menella, Chad.
–Comptes Rendus de l'Académie des Sciencies, Paris: Palevol: Vol. 4, pp. 671-679 - Kurtén, B. & Werdelin, L., 1988: A review of the genus Chasmaporthetes Hay, 1921 (Carnivora, Hyaenidae).
–Journal of Vertebrate Paleontology: Vol. 8, #1, pp. 46-66 - McKenna, M. C. & Bell, S. K., (eds.) 1997: Classification of mammals – above the species level.
–Columbia University Press, New York, 1997, xii-631 - Nowak, R. M., 1991: Walker's Mammals of the World; Part 1.
–The Johns Hopkins University Press, Baltimore and London, 1991, xlviii-642-lxiii - Nowak, R. M., 1991: Walker's mammals of the World; Part 2.
–The Johns Hopkins University Press, Baltimore and London, 1991, xii-643-1629 - Tseng, Z. J., Jin, C.-Z., Liu, J.-Y., Zheng, L.-T. & Sun, C.-K., 2008: Fossil Hyaenidae (Mammalia: Carnivora) from Huainan, Anhui Province, China.
–Vertebrata PalAsiatica: Vol. 46, #2, pp. 133-146 - Wang, X.-M., 2004: New materials of Tungurictis (Hyaenidae, Carnivora) from Tunggur Formation, Nei Mongol.
–Vertebrata PalAsiatica: Vol. 42, #2, pp. 144-153 - Werdelin, L. & Lewis, M. E., 2000: Carnivora from the South Turkwel hominid site, northern Kenya.
–Journal of Paleontology: Vol. 74, #6, pp. 1173-1180 - Werdelin, L. & Lewis, M. E., 2012: The Taxonomic Identity of the Type Specimen of Crocuta sivalensis (Falconer, 1867).
–Journal of Vertebrate Paleontology: Vol. 32, #6, pp. 1453-1456 [doi: 10.1080/039.032.0607] - Werdelin, L., 2003: Mio-Pliocene Carnivora from Lothagam, Kenya. 261-328.
in Leakey, M. G. & Harris, J. M., (eds.) 2003: Lothagam – The Dawn of Humanity in East Africa.
–Columbia University Press, New York, 2003, 1-678 - Werdelin, L. & Solounias, N., 1991: The Hyaenidae: taxonomy, systematics and evolution.
–Fossils & Strata: #30, 31 May, pp. 1-104