After McKenna & Bell, 1997, Trajano & Farrarezzi, 1994, Ginsburg & Morales, 1998, and Tedford & Martin, 2001
<==o Ursidae Fisher de Waldheim, 1817 non? Gray, 1825 (bears; karhut) |--o †Ballusia Ginsburg & Morales, 1998 | |-- †B. elmensis (Stehlin, 1917) [Ursavus elmensis Stehlin, 1917] | `-- †B. hareni (Ginsburg, 1989) [Ursavus hareni Ginsburg, 1989] `--+--o Tremarctinae Merriam & Stock, 1925 (silmälasikarhut) | |?-o †Plionarctos Frick, 1926 | | |?- †P. sp. | | |-- †P. edensis Frick, 1926 | | `-- †P. harroldorum Tedford & Martin, 2001 [Ursus sp. cf. U. abstrusus] | `--+--o Tremarctos Gervais, 1855 | | |?- †T. floridanus | | `-- T. ornatus (Spectacled bear; silmälasikarhu) | `--+--o †Arctodus [Arctoidotherium; Arctotherium Bravard, 1857; Dinarctotherium; Pararctotherium; Proarctotherium; Pseudarctotherium; Tremarctotherium] | | |-- †A. pristinus | | |-- †A. simus (Cope) [Arctotherium simum] (short-faced bear; lyhytkuonokarhu) | | `-- †A. angustidens (Kraglievich, 1926) | `--o †Arctotherium Bravard, 1857 | |--o †(Arctotherium) | | |-- †A. (A.) bonariense | | `-- †A. (A.) latidens | `--o †Pararctotherium Ameghino, 1904 [=? Arctodus; Arctotherium] | |?- †A. (P.) enectum (Paula Couto, 1960) | |-- †A. (P.) brasiliensis (Lund, 1838) | `-- †A. (P.) pamparum Ameghino, 1904 `--+--o Ailuropodinae (isopandat) | |?- †Kretzoiarctos beatrix (Abella, Montoya & Morales, 2011) Abella, Alba, Robles, Valenciano, Rotgers, Carmona, Montoya & Morales, 2012 [Agriarctos beatrix] | |--o †Agriarctos Kretzoi, 1942 | | |-- †A. gaali Kretzoi, 1942 | | |-- †A. vighi Kretzoi, 1942 | | `-- †A. depereti | `--+--o †Agriotherium [Hyaenarctos, Lydeccerion] | | |-- †A. insigne | | `-- †A. africanum | `--+--o †Indarctos Pilgrim, 1913 | | |-- †I. salmontanus Pilgrim, 1913 | | |-- †I. nevadensis Macdonald, 1959 | | |-- †I. oregonensis Merriam, 1925 | | |-- †I. zdanskyi Qiu & Tedford, 2003 | | |-- †I. punjabiensis | | |?- †I. laurillardi | | |-- †I. vireti Villalta & Crusafont, 1943 [Agriotherium insignis var. pontiensis Villalta & Crusafont, 1945; Indarctos arctoides vireti Thenius, 1959] | | `--+-- †I. arctoides (Depéret, 1895) | | |-- †I. atticus Weithofer, 1888 [incl. Indarctos bakalovi Kovachev, 1988; Indarctos lagrelii Zdansky, 1924; Indarctos sinensis Zdansky, 1924] | | `-- †I. anthracitis Weithofer, 1888 | `--+--o †Ailurarctos | | |-- †A. yuanmouensis | | `-- †A. lufengensis Qiu & Qi, 1989 | `--o Ailuropoda Milne-Edwards, 1870 [Aelureidopus, Ailuropus] (pandas, pandat) | |-- †A. microta (dwarf panda; kääpiöpanda) | |-- †A. wulingshanensis | `-- A. melanoleuca Milne-Edwards, 1870 (panda; isopanda) `--o Ursinae Fisher de Waldheim, 1817 non? Gray, 1825 (true bears; varsinaiset karhut) |--o †Ursavus Schlosser, 1899 | |-- †U. elmensis Stehlin | `--+-- †U. brevirhinus (Hofmann) | `--+-- †U. primaevus (Gaillard, 1899) | |-- †U. depereti Schlosser, 1902 | |-- †U. ehrenbergi (Brunner, 1942) [Ursus ehrenbergiBrunner, 1942] | |-- †U. sylvestris Qui & Qi, 1990 | `-- †U. isorei Ginsburg & Morales, 1998 `--o Ursus [Euarctos; Helarctos; Melursus; Selenarctos; Thalassarctos] (karhut) |?- †U. minimus Gervais, 1859 [U. boeckhi Schlosser, 1899] (Auvergne bear) |?- †U. wenzensis Stach, 1953 |?- †U. yinanensis Li, 1993 |?- †U. abstrusus Bjork, 1970 |-- U. ursinus Shaw, 1791 [Melursus ursinus (Shaw, 1791) Meyer, 1793] (sloth bear; huulikarhu) |--+-- U. malayanus Raffles, 1821 [Helarctos malayanus (Raffles, 1821) Horsfield, 1825] (Malayan sun bear; malajikarhu) | `--+--o U. thibetanus (Cuvier, 1823) [Selenarctos thibetanus; Ursus torquatus (Blandford, 1888)] (Asiatic black bear; sepelkarhu) | | |-- †U. t. mediterraneus (euroopansepelkarhu) | | |-- †U. t. permjak (uralinsepelkarhu) | | |-- U. t. thibetanus Cuvier, 1823 (Tibetan black bear; tiibetinsepelkarhu) | | |-- U. t. laniger Pocock, 1932 (Himalayan black bear; himalajansepelkarhu) | | |-- U. t. mupinensis Heude, 1901 (Indochinese black bear; indokiinansepelkarhu) | | |-- U. t. ussuricus Heude, 1901 (Ussuri black bear; amurinsepelkarhu) | | |-- †U. t. gedrosianus Blanford, 1877 (belutsistaninkarhu) [extiction status need to be validated] | | |-- U. t. japonicus Schlegel, 1857 (japaninsepelkarhu) | | `-- U. t. formosanus Swinhoe, 1864 (Formosan black bear; formosansepelkarhu) | `--o U. americanus Pallas, 1780 [Ursus americanus Pallas, 1780; Ursus luteolus Griffith, 1821; Ursus amblyceps Baird, 1859; Ursus floridanus Merriam, 1896; Ursus californiensis Miller, 1900; Ursus carlottae Osgood, 1901; Ursus altifrontalis Elliot, 1903; Ursus machetes Elliot, 1903; Ursus kermodei Hornaday, 1905] (black bear; mustakarhu) [16 subsp. (Larivière, 2001)] | |-- U. a. altifrontalis Elliot, 1903 (<>Britt. Kolumbia) | |-- U. a. amblyceps Baird, 1859 | |-- U. a. americanus Pallas, 1780 [U. a. sunborgeri Bangs; U. a. schwenki Shoemaker] (eastern black bear; idänmustakarhu) | |-- U. a. californiensis Miller, 1900 | |-- U. a. carlottae Osgood, 1901. | |-- U. a. cinnamomum Audubon & Bachman, 1854 (kanelikarhu) | |-- U. a. emmonsii Dall, 1895 (jäätikkökarhu) | |-- U. a. eremicus Merriam, 1904 | |-- U. a. floridanus Merriam, 1896 (floridanmustakarhu) | |-- U. a. hamiltoni Cameron, 1957 (Newfoundland black bear; newfoundlandinmustakarhu) | |-- U. a. kermodei Hornaday, 1905 (kermodeskarhu) | |-- U. a. luteolus Griffith, 1821 (louisiananmustakarhu) | |-- U. a. machetes Elliot, 1903 | |-- U. a. perniger Allen, 1910 [U. a. kenaiensis Allen] | |-- U. a. pugnax Swarth, 1911 | `-- U. a. vancouveri Hall, 1928 `--+-- †U. etruscus Cuvier `--+--o †U. spelaeus -group [Spelearctos E. Geoffroy, 1833] (cave bears; luolakarhut) | |-- †U. savini Andrews, 1922 (Savin's bear) | `--+--o †U. deningeri von Reichenau, 1904 (Mid-Pleistocene cave bear) | | |-- †U. d. praekudarensis Baryshnikov, 1998 | | |-- †U. d. kudarensis Baryshnikov, 1998 [U. kudarensis] (Asian cave bear) | | `-- †U. d. deningeri von Reichenau, 1904 | `--+--+-- †U. ingressus Rabeder & Hofreiter, 2004 (Gamssulzen Cave Bear) | | `?-o †U. rossicus Borissiak, 1930 [U. spelaeus rossicus Borissiak, 1930] | | |-- †U. r. rossicus | | `-- †U. r. uralensis [U. uralensis] | `--o †U. spelaeus Rosenmüller, 1794 (cave bear; luolakarhu) | |?- †U. s. kanivetz Vereshchagin, 1973 (Ural Mountain's cave bear) | |?- †U. s. odessanus von Nordmann, 1858 | |-- †U. s. ladinicus Hofreiter, 2005 | `--+-- †U. s. eremus Rabeder & Hofreiter, 2004 | `-- †U. s. spelaeus Rosenmüller, 1794 `--o U. arctos -group |-- U. maritimus Phipps, 1774 (polar bear; jääkarhu) `==o U. arctos Linnaeus, 1758 (brown bear; karhu) [paraphyletic without Ursus maritimus] |--+== U. a. arctos Linnaeus, 1758 (Eurasian brown bear; karhu) [Euraasia] | |-- †U. a. crowtheri (Atlas bear; atlaksenkarhu) | |-- U. a. pruinosus Blyth, 1854 [<Tibet>] | |-- U. a. lasiotus Gray, 1867 [<North China>] | |-- U. a. syriacus Hemprich & Ehrenberg, 1828 | |-- U. a. isabellinus Horsfield, 1826 [<Nepal>] | `-- U. a. collaris Cuvier and Geoffroy, 1824 [<Siberia>] `--+-- U. a. alascensis Merriam, 1896 [<Alaska>] |?- U. maritimus Phipps, 1774 (polar bear; jääkarhu) |-- U. a. beringianus Middendorff, 1853 [<Great Shantar Island, Sea of Okhotsk>] |-- U. a. gyas Merriam, 1902 [<Alaska Peninsula>] |-- U. a. dalli Merriam, 1896 [<NW Alaska>] |-- U. a. sitkensis Merriam, 1896 [<Alaska Panhandle>] |-- U. a. stikeenensis Merriam, 1914 [<northern British Columbia>] |-- U. a. middendorffi Merriam, 1896 (kodiak bear; kodiakinkarhu) |-- U. a. horribilis Ord, 1815 (grizzley; harmaakarhu) |-- †U. a. californicus Merriam, 1896 (Californian grizzley; kalifornianruskeakarhu) `-- †U. a. nelsoni Merriam, 1914 (mexican brown bear; meksikonkarhu)
Alternative phylogenies of Ursidae:
After Abella, Alba, Robles, Valenciano, Rotgers, Carmona, Montoya & Morales, 2012<==o Ursidae |--o Ailuropodinae | |?- †Agriotherium [not treated] | |--o †Indarctini | | `--Indarctos | `--o Ailuropodini | |-- Ailuropoda | `--+-- †Ailurarctos | `--+-- †Agriarctos | `-- †Kretzoiarctos `--+-- Tremarctinae `-- Ursinae Reference(s):
|
After Loreille, Orlando, Patou-Mathis, Philippe, Taberlet & Hänni, 2001
<==o Ursidae
|-- Ailuropoda melanoleuca
`--+-- Tremarctos ornatus
`--+-- Melursus ursinus
`--+--+-- Helarctos malayanus
| `--+-- Ursus thibetanus
| `-- Ursus americanus
`--+-- †Ursus spelaeus
`--+== Ursus arctos
`-- Ursus maritimus
Reference(s):
|
Reference(s):
- Abella, J., Alba, D. M., Robles, J. M., Valenciano, A., Rotgers, C., Carmona, R., Montoya, P. & Morales, J., 2012: Kretzoiarctos gen. nov., the Oldest Member of the Giant Panda Clade.
–Plos One: Vol. 7, #11, pp. e48985 (1-7) [doi: 10.1371/journal.pone.0048985] - Abella, J., Montoya, P. & Morales, J., 2011: A New species of Agriarctos (Ailuropodinae, Ursidae, Carnivora) in the locality of Nombrevilla 2 (Zaragoza, Spain)
[Una nueva especie de Agriarctos (Ailuropodinae, Ursidae, Carnivora) en la localidad de Nombrevilla 2 (Zaragoza, España)].
–Estudios Geológicos: Vol. 67, #2, pp. 187-191 [doi: 10.3989/egeol.40714.182] - Andrews, C. W., 1922: XXII.—Note on a bear (Ursus savini, sp. n.) from the Cromer Forest-bed.
–Annals and Magazine of Natural History: Vol. 9, #50, pp. 204-207 [doi: 10.1080/00222932208632660] - Bocherens, H., Stiller, M., Hobson, K. A., Pacher, M., Rabeder, G., Burns, J. A., Tütken, T. & Hofreiter, M., 2011: Niche partitioning between two sympatric genetically distinct cave bears (Ursus spelaeus and Ursus ingressus) and brown bear (Ursus arctos) from Austria: Isotopic evidence from fossil bones.
–Quaternary International: Vol. 245, #2, pp. 238-248 [doi: http://dx.doi.org/10.1016/j.quaint.2010.12.020] - Carranza-Castañeda, O. & Miller, W. E., 1987: Rediscovered type specimens and other important published Pleistocene mammalian fossils from Central Mexico.
–Journal of Vertebrate Paleontology: Vol. 7, #3, pp. 335-341 - 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. - Crusafont Pairó, M. & Kurtén, B., 1976: Bears and bear-dogs from the Vallesian of the Vallés-Penedés Basin, Spain.
–Acta Zoologica Fennica: Vol. 144, #Separatum, pp. 1-29 - DeMaster, D. P. & Stirling, I., 1981: Ursus maritimus.
–Mammalian Species Archive: Vol. 145, pp. 1-7 [doi: 10.2307/3503828] - Fitzgerald, C. S. & Krausman, P. R., 2002: Helarctos malayanus.
–Mammalian Species Archive: Vol. 696, pp. 1-5 [doi: 10.2307/3504544] - García, N., Santos, E., Arsuaga, J. L. & Carretero, J. M., 2007: Endocranial morphology of the Ursus deningeri von Reichenau, 1904 from the Sima de Los Huesos (Sierra de Atapuerca) Middle Pleistocene site.
–Journal of Vertebrate Paleontology: Vol. 27, #4, pp. 1007-1017 [doi: 10.1671/0272-4634(2007)27[1007:emotud]2.0.co;2] - Geraads, D., Kaya, T. & Mayda, S., 2005: Late Miocene large mammals from Yulafli, Thrace region, Turkey, and their biogeographic implications.
–Acta Palaeontologica Polonica: Vol. 50, #3, pp. 523-544 - Ginsburg, L. & Morales, J., 1998: Les Hemicyoninae (Ursidae, Carnivora, Mammalia) et les formes apparentées du Miocène inférieur et moyet d'Europe occidentale.
[Hemicyoninae (Ursidae, Carnivora, Mammalia) and the related taxa from Early and Middle Miocene of western Europe.]
–Annales de Paléontologie: Vol. 84, #1, pp. 71-123 - Larivière, S., 2001: Ursus americanus.
–Mammalian Species Archive: Vol. 647, pp. 1-11 [doi: 10.2307/3504406] - Lindqvist, C., Schuster, S. C., Sun, Y., Talbot, S. L., Qi, J., Ratan, A., Tomsho, L. P., Kasson, L., Zeyl, E., Aars, J., Miller, W., Ingólfsson, Ó., Bachmann, L. & Wiig, Ø., 2010: Complete mitochondrial genome of a Pleistocene jawbone unveils the origin of polar bear.
–Proceedings of the National Academy of Sciences: Vol. 107, #11, pp. 5053-5057 [doi: 10.1073/pnas.0914266107] - Loreille, O., Orlando, L., Patou-Mathis, M., Philippe, M., Taberlet, P. & Hänni, C., 2001: Ancient DNA analysis reveals divergence of the cave bear, Ursus spelaeus, and brown bear, Ursus arctos, lineages.
–Current Biology: Vol. 11, #3, pp. 200-203 [doi: 10.1016/S0960-9822(01)00046-X] - McKenna, M. C. & Bell, S. K., eds., 1997:Classification of mammals; above the species level.
–Columbia University Press, New York, 1997. - Münzel, S. C., Rivals, F., Pacher, M., Döppes, D., Rabeder, G., Conard, N. J. & Bocherens, H., 2014: Behavioural ecology of Late Pleistocene bears (Ursus spelaeus, Ursus ingressus): Insight from stable isotopes (C, N, O) and tooth microwear.
–Quaternary International: Vol. 339–340, pp. 148-163 [doi: http://dx.doi.org/10.1016/j.quaint.2013.10.020] - Münzel, S. C.; Stiller, M., Pacher, M. & Athen, K., 2010: Genetic Results versus Morphological and Biometrical Analysis – The Case of Ursus spelaeus (sensu lato). –ICAZ 2010 Paris, Session 5-3 Poster: http://alexandriaarchive.org/bonecommons/archive/files/icaz_s5-3_poster_m%C3%BCnzel_etal_2010_ce8d6538d7.pdf
- Nagel, U., 2007: Personal communication (a lecture)
- 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 - Pasitschniak-Arts, M., 1993: Ursus arctos.
–Mammalian Species Archive: Vol. 439, pp. 1-10 [doi: 10.2307/3504138] - Rabeder, G. & Hofreiter, M., 2004: Der neue Stammbaum der alpinen Höhlenbären.
–Die Höhle: Vol. 55, #1-4, pp. 58-77 - Tedford, R. H. & Martin, J., 2001: Plioarctos, a tremarctine bear (Ursidae: Carnivora) from Western North America.
–Journal of Vertebrate Paleontology: Vol. 21, #2, pp. 311-321 - Trajano, E. & Farrarezzi, H., 1994: A fossil bear from northeastern Brazil, with a phylogenetic analysis of the South American extinct Tremarctinae (Ursidae).
–Journal of Vertebrate Paleontology: Vol. 14, #4, pp. 552-561