Ursidae

Spectacled bears Tremarctos ornatus. Photograph from here.


Belongs within: Carnivora.

The Ursidae, bears, are a family of relatively large carnivorans found primarily in Eurasia and the Americas. The living forms are plantigrade and omnivorous (in the case of the giant panda Ailuropoda melanoleuca, primarily herbivorous, while the polar bear Ursus maritimus is primarily carnivorous). Among recent species, the largest is U. maritimus; the smallest is the sun bear Helarctos malayanus. The extinct subfamilies Amphicynodontinae and Hemicyoninae were longer-faced (and often smaller) than modern bears; the Hemicyoninae in particular were superficially similar to dogs.

Synapomorphies (from Flynn et al. 1988): Malleus with tensor tympani insertion process reduced or lost; two upper and three lower molars; upper molars enlarged by expansion of metaconule; protocone and metaconule of upper molars forming longitudinal crest occluding with enlarged talonid cusps of lower molars; M2 developing posteriorly projecting platform occluding with M3; M2 trigonid lacking paraconid. Internal carotid artery folded intracranially within inferior petrosal sinus.

<==Ursidae [Ursoidea]
    |--Amphicynodontinae [Amphicynodontidae] FNT88
    |    |--Amphicynodon teilhardi FF06
    |    |--Pachycynodon gryei T76
    |    `--Parictis parvus MJ11, FF06
    |--Hemicyoninae MJ11
    |    |--Hemicyon DW04
    |    |    |--H. johnhenryi [=Phoberocyon johnhenryi] V91
    |    |    `--H. teilhardi DW04
    |    `--Cephalogale MJ11
    `--+--Tremarctos Gervais 1855 C96, C57 [incl. Nearctos Gray 1876 C57; Tremarctinae]
       |    |--*T. ornatus (Cuvier 1825) (see below for synonymy) C57
       |    `--T. floridanus B84a
       `--Ursinae C96
            |  i. s.: Helarctos malayanus C96, B84a [=Ursus malayanus BP87]
            |         Melursus ursinus C96, B84a
            |         Selenarctos thibetanus C96, B84a [=Ursus thibetanus BP87]
            |           |--S. t. thibetanus I92
            |           `--S. t. japonicus (Schlegel 1857) I92
            |--Ursavus elmensis JC07, B84a
            |--+--Ailurarctos JC07
            |  `--Ailuropoda [Ailuropodidae, Ailuropodinae, Ailuropodini] JC07
            |       |--A. baconi JC07
            |       |--A. melanoleuca (David 1869) H07
            |       |--A. microta HC95
            |       `--A. wulingshanensis JC07
            `--Ursus L84
                 |--U. abstrusus TH03
                 |--U. americanus Pallas 1780 B75
                 |    |--U. a. americanus B84c
                 |    |--U. a. altifrontalis B84c
                 |    |--U. a. carlottae B84c
                 |    |--U. a. cinnamomum B84c
                 |    |--U. a. emmonsii B84c
                 |    |--U. a. eremicus MB86
                 |    |--U. a. hamiltoni B84c
                 |    |--U. a. kermodei B84c
                 |    |--U. a. machetes Elliot 1903 MB86
                 |    `--U. a. vancouveri B84c
                 |--U. arctos B84b
                 |    |--U. a. arctos B84b
                 |    |--U. a. horribilis B84b
                 |    |--U. a. middendorffi B84b
                 |    |--U. a. nelsoni USDI77
                 |    |--U. a. pruinosus USDI77
                 |    `--U. a. yesoensis Lydekker 1897 I92
                 |--U. deningeri [incl. U. dolinensis] ME05
                 |--U. etruscus L84
                 |--U. formosanus Swinhoe 1864 S66
                 |--U. maritimus L84 [=Thalarctos maritimus BP87]
                 |--U. minimus B84a
                 |--U. rodei ME05
                 |--U. spelaeus B84a
                 `--U. syriacus T66

Ursidae incertae sedis:
  Tremarctotherium simum FP64
  Agriotherium S78
    |--A. africanum S78
    `--A. inexpetans DW04
  Arctodus V91
    |--A. pristinus V91
    `--A. simus V91
  Euarctos americanus GM71
  Indarctos atticus FF06
  Amphicticeps FF06
    |--A. dorog FF06
    `--A. shackelfordi FF06

Inorganic: Helarctos malayanus protominilorientalus Okamura 1987 O87
           Thalarctos maritimus minilorientalus Okamura 1987 O87
           Ursus arctos protominilorientalus Okamura 1987 O87

*Tremarctos ornatus (Cuvier 1825) [=Ursus ornatus, Helarctos ornatus, Nearctos ornatus; incl. Ursus frugilegus Tschudi 1844, U. fructilegus, Tremarctos lasallei María 1924, T. ornatus majori Thomas 1902, U. nasutus Sclater 1868, U. ornatus thomasi Hornaday 1911] C57

*Type species of generic name indicated

REFERENCES

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1-184.

[B84a] Bunnell, F. 1984a. The bear family. In All the World’s Animals: Carnivores (D. Macdonald, ed.) pp. 78-79. Torstar Books Inc.: New York.

[B84b] Bunnell, F. 1984b. Grizzly bear. In All the World’s Animals: Carnivores (D. Macdonald, ed.) pp. 80-83. Torstar Books Inc.: New York.

[B84c] Bunnell, F. 1984c. American black bear. In All the World’s Animals: Carnivores (D. Macdonald, ed.) pp. 86-87. Torstar Books Inc.: New York.

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[C57] Cabrera, A. 1957. Catalogo de los mamiferos de America del Sur. I (Metatheria—Unguiculata—Carnivora). Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Instituto Nacional de Investigacion de Las Ciencias Naturales, Ciencias Zoológicas 4 (1): 1-307.

[C96] Campbell, N. A. 1996. Biology (4th ed.) The Benjamin/Cummings Publishing Company, Inc.: Menlo Park (California).

[DW04] Deng T., Wang X., Ni X. & Liu L. 2004. Sequence of the Cenozoic mammalian faunas of the Linxia Basin in Gansu, China. Acta Geologica Sinica (English Edition) 78 (1): 8-14.

[FF06] Finarelli, J. A. & J. J. Flynn. 2006. Ancestral state reconstruction of body size in the Caniformia (Carnivora, Mammalia): the effects of incorporating data from the fossil record. Systematic Biology 55 (2): 301-313.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.

[FNT88] Flynn, J. J., N. A. Neff & R. H. Tedford. 1988. Phylogeny of the Carnivora. In The Phylogeny and Classification of the Tetrapods, vol. 2. Mammals (M. J. Benton, ed.) pp. 73-116. Clarendon Press: Oxford.

[GM71] Goodman, M., & G. W. Moore. 1971. Immunodiffusion systematics of the primates I. The Catarrhini. Systematic Zoology 20 (1): 19-62.

[H07] Hooker, J. J. 2007. Bipedal browsing adaptations of the unusual Late Eocene-earliest Oligocene tylopod Anoplotherium (Artiodactyla, Mammalia). Zoological Journal of the Linnean Society 151 (3): 609-659.

[HC95] Huang W., R. Ciochon, Gu Y., R. Larick, Fang Q., H. Schwarcz, C. Yonge, J. de Vos & W. Rink. 1995. Early Homo and associated artefacts from Asia. Nature 378: 275-278.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[JC07] Jin, C., R. L. Ciochon, W. Dong, R. M. Hunt, Jr, J. Liu, M. Jaeger & Q. Zhu. 2007. The first skull of the earliest giant panda. Proceedings of the National Academy of Sciences of the USA 104 (26): 10932-10937.

[L84] Lentfer, J. W. 1984. Polar bear. In All the World’s Animals: Carnivores (D. Macdonald, ed.) pp. 84-85. Torstar Books Inc.: New York.

[MB86] Matson, J. O. & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1-88.

[MJ11] Meredith, R. W., J. E. Janečka, J. Gatesy, O. A. Ryder, C. A. Fisher, E. C. Teeling, A. Goodbla, E. Eizirik, T. L. L. Simão, T. Stadler, D. L. Rabosky, R. L. Honeycutt, J. J. Flynn, C. M. Ingram, C. Steiner, T. L. Williams, T. J. Robinson, A. Burk-Herrick, M. Westerman, N. A. Ayoub, M. S. Springer & W. J. Murphy. 2011. Impacts of the Cretaceous terrestrial revolution and KPg extinction on mammal diversification. Science 334: 521-524.

[ME05] Moullé, P.-E., A. Echassoux, F. Lacombat, E. Desclaux & S. Bailon. 2005. L’environnement animal des premiers habitants de l’Europe méditerranéenne: les grands mammifères contemporains de l’homme du Vallonnet, données taxonomiques et biostratigraphiques pour la deuxième moitie du Pléistocène inférieur. BAR International Series 1364: 105-113.

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347-573.

[S78] Savage, R. J. G. 1978. Carnivora. In Evolution of African Mammals (V. J. Maglio & H. B. S. Cooke, eds) pp. 249-267. Harvard University Press: Cambridge (Massachusetts).

[S66] Sclater, P. L. 1866. Notice of recent additions to the Society’s menagerie. Proceedings of the Zoological Society of London 1866: 418-419.

[T76] Tedford, R. H. 1976. Relationship of pinnipeds to other carnivores (Mammalia). Systematic Zoology 25 (4): 363-374.

[TH03] Tedford, R. H., & C. R. Harington. 2003. An Arctic mammal fauna from the Early Pliocene of North America. Nature 425: 388-390.

[T66] Tristram, H. B. 1866. Report on the mammals of Palestine. Proceedings of the Zoological Society of London 1866: 84-93.

[USDI77] USDI (United States Department of the Interior). 1977. Endangered and threatened wildlife and plants - republication of list of species. Federal Register 42: 36420-36431.

[V91] Valkenburgh, B. van. 1991. Iterative evolution of hypercarnivory in canids (Mammalia: Carnivora): evolutionary interactions among sympatric predators. Paleobiology 17 (4): 340-362.

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