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-Paenungulata OB13
    |--Desmostylia TS88
    |    |--Behemotops TS88
    |    |--Cornwallius BDR85
    |    |--Palaeoparadoxia Tokunaga 1939 ANG03, D07
    |    |    `--*P. tabatai BDR85, TS88
    |    `--Desmostylus Marsh 1888 D07 [incl. Vanderhoofius BDR85]
    |         |--D. cymnatius Hannibal 1922 R76
    |         `--D. hesperus Marsh 1888 R76
    `--Paenungulata [Pan-Tethytheria, Tethytheria, Uranotheria] OB13
         |  i. s.: Lammidhania wardi TS88
         |         0--Astrapotheria MC00
         |         |    |--Tetragonostylops MC00
         |         |    `--Astrapotherium Ameghino 1887 D07
         |         |         |--A. granda D07
         |         |         `--A. magnum D07
         |         `--Uintatheriamorpha MC00
         |              |--Pyrotheria PMF88
         |              |    |--Carodnia [Xenungulata] OB13
         |              |    |    `--C. vierai OB13
         |              |    `--+--Carolozittelia PMF88
         |              |       `--+--Griphodon PMF88
         |              |          `--Pyrotherium Ameghino 1889 PMF88, D07
         |              |               |--P. macfaddeni D07
         |              |               `--P. romeroi D07
         |              `--Dinocerata MC00
         |                   |--Loxolophodon cornutus C77
         |                   |--Mongolotherium PMF88
         |                   |--Probathyopsis D07
         |                   |--Uintatherium Leidy 1872 D07 (see below for synonymy)
         |                   |    |--U. anceps H07
         |                   |    |--U. mirabile C77
         |                   |    `--U. robustum D07
         |                   `--Prodinoceras Matthew, Granger & Simpson 1929 D07
         |                        |--P. diconicus D07
         |                        |--P. martyr D07
         |                        |--P. primigenium D07
         |                        |--P. simplum D07
         |                        `--P. turfanensis D07
         |--Pan-Sirenia OB13
         |    |--Sirenia FS15
         |    `--Prorastomidae D78
         |         |--Sirenavus hungaricus Kretzoi 1941 D78, S76
         |         |--Prorastomus Owen 1855 D07
         |         |    `--P. sirenoides D01
         |         `--Pezosiren Domning 2001 D01
         |              `--*P. portelli Domning 2001 D01
         `--+--+--Hyracoidea FS15
            |  `--Eritherium azzouzorum RH14, OB13
            `--+--Proboscidea FS15
               `--Embrithopoda ANG03
                    |--Phenacolophus [Phenacolophidae] PMF88
                    |    `--P. fallax RH14
                    |--Crivadiatherium PMF88
                    |--Palaeoamasia PMF88
                    `--Arsinoitherium Beadnell 1902 [Arsinoitheriidae] SKR04
                         |--A. giganteum Sanders, Kappelman & Rasmussen 2004 SKR04
                         `--A. zitteli Beadnell 1902 T78 [incl. A. andrewsi Lankester 1903 SKR04, T78]

Uintatherium Leidy 1872 D07 [incl. Dinoceras Marsh 1872 C77, FT93; Uintatheriidae PMF88]

*Type species of generic name indicated


[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.

[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.

[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.

[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.

[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.

[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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