Pan-Paenungulata

Mounted skeleton of Arsinoitherium zitteli in the Natural History Museum, London, copyright Aram Dulyan.

Belongs within: Atlantogenata.
Contains: Sirenia, Hyracoidea, Proboscidea.

The Pan-Paenungulata are a clade of mammals including modern hyraxes, elephants and sirenians, and all fossil taxa more closely related to them than other living mammals. Likely fossil members of this clade include the Desmostylia, aquatic herbivores from the Oligocene and Miocene of the North Pacific, and the Embrithopoda from the Eocene and Oligocene of Africa and Asia.

<==Pan-PaenungulataOB13
    |--Desmostylidae [Cornwalliidae, Cornwalliusidae, Desmostylia, Paleoparadoxidae]TS88
    |    |--Behemotops Domning, Ray & McKenna 1986DRM86
    |    |    |--*B. proteus Domning, Ray & McKenna 1986DRM86
    |    |    `--B. emlongi Domnin, Ray & McKenna 1986DRM86
    |    `--+--Paleoparadoxia Reinhardt 1959DRM86, SM93
    |       |    `--*P. tabataiBDR85, DRM86 [=Cornwallius tabataiDRM86]
    |       `--+--Cornwallius Hay 1923DRM86, SM93
    |          |    `--*C. sookensisDRM86
    |          `--Desmostylus Marsh 1888DRM86, D07 [incl. VanderhoofiusBDR85]
    |               |--D. cymnatius Hannibal 1922R76
    |               `--D. hesperus Marsh 1888R76
    `--Paenungulata [Pan-Tethytheria, Tethytheria, Uranotheria]OB13
         |  i. s.: 0--AstrapotheriaMC00
         |         |    |--Eoastrapostylops Soria & Powell 1981 [Eoastrapostylopidae]SM93
         |         |    |--Trigonostylopidae [Trigonostylopoidea]SM93
         |         |    |    |--Tetragonostylops de Paula Couto 1963SM93
         |         |    |    `--Trigonostylops Ameghino 1897SM93
         |         |    `--AstrapotheriidaeSM93
         |         |         |--Albertogaudrya Ameghino 1901SM93
         |         |         |--Scaglia Simpson 1957SM93
         |         |         |--Monoeidodon Roth 1898SM93
         |         |         `--Astrapotherium Burmeister 1879SM93
         |         |              |--A. grandaD07
         |         |              `--A. magnumD07
         |         `--UintatheriamorphaMC00
         |              |--PyrotheriaPMF88
         |              |    |  i. s.: ColombitheriidaeR06
         |              |    |           |--Proticia Patterson 1977R06, SM93
         |              |    |           `--ColombitheriumR06
         |              |    |--Carodnia Simpson 1935OB13, SM93 [Carodniidae, Xenungulata]
         |              |    |    `--C. vieraiOB13
         |              |    `--+--CarolozitteliaPMF88
         |              |       `--+--GriphodonPMF88
         |              |          `--PyrotheriidaeSM93
         |              |               |--Propyrotherium Ameghino 1901SM93
         |              |               `--Pyrotherium Ameghino 1888PMF88, SM93
         |              |                    |--P. macfaddeniD07
         |              |                    `--P. romeroiD07
         |              `--Dinocerata [Bathyopsinae]MC00
         |                   |  i. s.: Loxolophodon cornutusC77
         |                   |--Prodinoceratidae [Prodinoceratinae]R06
         |                   |    |--BathyopsoidesSM71
         |                   |    `--Prodinoceras Matthew, Granger & Simpson 1929SM93 (see below for synonymy)
         |                   |         |--P. diconicusD07
         |                   |         |--‘Mongolotherium’ efremoviSM71
         |                   |         |--P. martyrD07
         |                   |         |--‘Mongolotherium’ plantigradumSM71
         |                   |         |--‘Probathyopsis’ praecursor Simpson 1929G52
         |                   |         |--P. primigeniumD07
         |                   |         |--P. simplumD07
         |                   |         `--P. turfanensis Chow 1960SM71
         |                   `--Uintatheriidae [Uintatheriinae]SM93
         |                        |  i. s.: Gobiatherium Osborn & Granger 1932SM93
         |                        |--+--Tetheopsis Cope 1885SM71, SM93
         |                        |  |--Bathyopsis fissidens Cope 1881SM71, G52
         |                        |  `--Eobasileus Cope 1872SM71, SM93
         |                        |       |--E. cornutusP96
         |                        |       `--E. pressicornisC77
         |                        `--+--Uintatherium Leidy 1872SM71, D07 [incl. Dinoceras Marsh 1872C77, FT93]
         |                           |    |--U. ancepsH07
         |                           |    |--U. mirabileC77
         |                           |    `--U. robustumD07
         |                           `--ProuintatheriumSM71
         |--Pan-SireniaOB13
         |    |--SireniaFS15
         |    `--ProrastomidaeD78
         |         |--Sirenavus hungaricus Kretzoi 1941D78, S76
         |         |--Prorastomus Owen 1855D07
         |         |    `--P. sirenoidesD01
         |         `--Pezosiren Domning 2001D01
         |              `--*P. portelli Domning 2001D01
         `--+--+--HyracoideaFS15
            |  `--Eritherium azzouzorumRH14, OB13
            `--+--ProboscideaFS15
               `--EmbrithopodaANG03
                    |--Phenacolophus Matthew & Granger 1925 [Phenacolophidae]SM93
                    |    `--P. fallax [incl. Procoryphodon primaevus]SM71
                    `--ArsinoitheriidaeSM93
                         |--Palaeoamasia Ozansoy 1966SM93
                         |--Crivadiatherium Rudalesco et al. 1976SM93
                         |--HypsamasiaR06
                         `--Arsinoitherium Beadnell 1902SKR04
                              |--A. giganteum Sanders, Kappelman & Rasmussen 2004SKR04
                              `--A. zitteli Beadnell 1902T78 [incl. A. andrewsi Lankester 1903SKR04, T78]

Prodinoceras Matthew, Granger & Simpson 1929SM93 [incl. Mongolotherium Flerov 1952R06, SM71, ProbathyopsisR06]

*Type species of generic name indicated

References

[ANG03] Asher, R. J., M. J. Novacek & J. H. Geisler. 2003. Relationships of endemic African mammals and their fossil relatives based on morphological and molecular evidence. Journal of Mammalian Evolution 10 (1–2): 131–194.

[BDR85] Barnes, L. G., D. P. Domning & C. E. Ray. 1985. Status of studies on fossil marine mammals. Marine Mammal Science 1 (1): 15–53.

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i–iv, 1–370.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[D78] Domning, D. P. 1978. Sirenia. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 573–581. Harvard University Press: Cambridge (Massachusetts).

[D01] Domning, D. P. 2001. The earliest known fully quadrupedal sirenian. Nature 413: 625–627.

[DRM86] Domning, D. P., C. E. Ray & M. C. McKenna. 1986. Two new Oligocene desmostylians and a discussion of tethytherian systematics. Smithsonian Contributions to Paleobiology 59: 1–56.

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[FT93] Fensome, R. A., F. J. R. Taylor, G. Norris, W. A. S. Sarjeant, D. I. Wharton & G. L. Williams. 1993. A classification of living and fossil dinoflagellates. Micropaleontology Special Publication 7: i–viii, 1–351.

[G52] Gazin, C. L. 1952. The Lower Eocene Knight Formation of western Wyoming and its mammalian faunas. Smithsonian Miscellaneous Collections 117 (18): 1–82, 11 pls.

[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.

[MC00] Muizon, C. de, & R. L. Cifelli. 2000. The “condylarths” (archaic Ungulata, Mammalia) from the early Palaeocene of Tiupampa (Bolivia): implications on the origin of the South American ungulates. Geodiversitas 22 (1): 47–150.

[OB13] O’Leary, M. A., J. I. Bloch, J. J. Flynn, T. J. Gaudin, A. Giallombardo, N. P. Giannini, S. L. Goldberg, B. P. Kraatz, Z.-X. Luo, J. Meng, X. Ni, M. J. Novacek, F. A. Perini, Z. S. Randall, G. W. Rougier, E. J. Sargis, M. T. Silcox, N. B. Simmons, M. Spaulding, P. M. Velazco, M. Weksler, J. R. Wible & A. L. Cirranello. 2013. The placental mammal ancestor and the post-K–Pg radiation of placentals. Science 339: 662–667.

[P96] Prothero, D. R. 1996. Magnetic stratigraphy and biostratigraphy of the Middle Eocene Uinta Formation, Uinta Basin, Utah. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 3–24. Cambridge University Press.

[PMF88] Prothero, D. R., E. M. Manning & M. Fischer. 1988. The phylogeny of the ungulates. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 201–234. Clarendon Press: Oxford.

[R76] Ray, C. E. 1976. Fossil marine mammals of Oregon. Systematic Zoology 25 (4): 420–436.

[R06] Rose, K. D. 2006. The Beginning of the Age of Mammals. John Hopkins University Press: Baltimore.

[RH14] Rose, K. D., L. T. Holbrook, R. S. Rana, K. Kumar, K. E. Jones, H. E. Ahrens, P. Missiaen, A. Sahni & T. Smith. 2014. Early Eocene fossils suggest that the mammalian order Perissodactyla originated in India. Nature Communications 5: 5570.

[SKR04] Sanders, W. J., J. Kappelman & D. T. Rasmussen. 2004. New large-bodied mammals from the late Oligocene site of Chilga, Ethiopia. Acta Palaeontologica Polonica 49 (3): 365–392.

[S76] Savage, R. J. G. 1976. Review of early Sirenia. Systematic Zoology 25 (4): 344–351.

[SM93] Stucky, R. K., & M. C. McKenna. 1993. Mammalia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 739–771. Chapman & Hall: London.

[SM71] Szalay, F. S., & M. C. McKenna. 1971. Beginning of the age of mammals in Asia: the late Paleocene Gashato fauna, Mongolia. Bulletin of the American Museum of Natural History 144 (4): 269–318.

[T78] Tanner, L. G. 1978. Embrithopoda. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 279–283. Harvard University Press: Cambridge (Massachusetts).

[TS88] Tassy, P., & J. Shoshani. 1988. The Tethytheria: elephants and their relatives. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 283–315. Clarendon Press: Oxford.

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