Reconstruction of Embolotherium andrewsi with size comparison to average human, copyright Tim Bertelink.

Belongs within: Euungulata.
Contains: Equoidea, Ceratomorpha, Chalicotheriidae.

The Perissodactyla include the odd-toed hoofed mammals: horses, tapirs and rhinoceroses, as well as their fossil relatives the chalicotheres and brontotheres. Modern perissodactyls are relatively large herbivorous mammals with hind-gut fermenting digestive systems. The Brontotheriidae are a group of rhinoceros-like animals known from the Eocene of North America and Eurasia.

Synapomorphies (from Hooker & Dashzeveg 2004): Upper molar paraconule situated distinctly more mesially than protocone, facet 2A occupying paraconule and part of preprotocrista, notched preprotocrista mesiobuccally directed towards paraconule, lower molar metaconid widely twinned with tips bearing facet 2A; upper molar metaconule distinctly mesial of line drawn between metacone and hypocone, lower molar hypoconulid separated from and distal of crest joining hypoconid and entoconid, M3 with hypocone; upper molar parastyle large; M3 not smaller than M2, M3 larger than M2; upper molars with metaconal fold not joined to metaconule; upper molar metaloph and lower molar hypolophid, especially on M3, oblique.

Perissodactyla [Ancylopoda, Euperissodactyla, Lophodontomorpha, Mesaxonia, Moropomorpha, Selenida, Tapiromorpha]
    |  i. s.: Symborodon C77
    |         Orotherium Marsh 1872 C77
    |           |--O. cristonense Cope 1877 C77
    |           |--O. laevii Cope 1877 C77
    |           `--O. vintanum Marsh 1872 C77
    |         Orientolophus Ting 1993 HD04
    |           |--*O. hengdongensis Ting 1993 HD04
    |           `--O. namadicus (Dashzeveg 1979) [=Homogalax namadicus] HD04
    |         Mesolambdolophus setoni HUG17
    |--Hallensia mathewsi RH14, F02
    `--+--Equoidea RH14
       `--Isectolophidae RH14
            |--Homogalax RH14
            |    |--H. protapirinus [=Systemodon protapirinus] F02
            |    `--H. wutuensis HD04
            `--+--Cardiolophus radinskyi Gingerich 1991 RH14, HD04
               `--+--Ceratomorpha RH14
                  |--Isectolophus RH14
                  |    |--I. annectens RH14
                  |    `--I. latidens RH14
                  |--Chalicotherioidea HD04
                  |    |  i. s.: Kalimantsia Geraards, Spassov & Kovachev 2001 D07
                  |    |           `--K. bulgarica D07
                  |    |--Protomoropus Hooker & Dashzeveg 2004 HD04
                  |    |    `--*P. gabuniai (Dashzeveg 1979) [=Hyracotherium gabuniai, Orientolophus gabuniai] HD04
                  |    `--+--+--Chalicotheriidae RH14
                  |       |  `--Eomoropidae RH14
                  |       |       |--Eomoropus amarorum RH14
                  |       |       `--Litolophus RH14
                  |       |            |--L. gobiensis HUG17
                  |       |            `--L. mongoliensis RH14
                  |       `--Lophiodontidae HD04
                  |            |--Paleomoropus jepseni Radinsky 1964 HD04
                  |            `--+--Lophiaspis maurettei Depéret 1910 HD04
                  |               `--Lophiodon Cuvier 1822 HD04, M02 [incl. Saurornis Fischer 1967 M02]
                  |                    |--L. leptorhynchum RH14
                  |                    |--*Saurornis’ matthesi Fischer 1967 M02
                  |                    `--L. remensis RH14
                  `--+--Lambdotherium [Lambdotheriidae] RH14
                     |    `--L. popoagicum HD04
                     `--Brontotheriidae [Brontotherioidea, Titanotheriidae, Titanotheriomorpha] RH14
                          |  i. s.: Danjiangia HD04
                          |         Dolichorhinus Douglass 1924 D07
                          |           |--D. hyognathus D07
                          |           `--D. intermedius D07
                          |         Nanotitanops shanghuangensis D07
                          |         Aktautitan Mihlbachler, Lucas et al. 2004 D07
                          |           `--A. hippopotamopus D07
                          |         Protitanotherium Lucas 1983 D07
                          |           |--P. curryi D07
                          |           |--P. emarginatum D07
                          |           `--P. superbum D07
                          |         Megacerops Leidy 1870 D07
                          |           |--M. coloradensis D07
                          |           |--M. dispar [=Brontops dispar] D07
                          |           |--M. elatus [=Titanops elatus] D07
                          |           |--M. gigas [=Brontotherium gigas] D07
                          |           |--M. ingens [=Brontotherium ingens] S07
                          |           |--M. kuwagatarhinus D07
                          |           `--M. tyleri D07
                          |         Eotitanops Osborn 1907 RH14, D07
                          |           |--E. borealis D07
                          |           `--E. minimus D07
                          |         Palaeosyops Leidy 1870 RH14, D07
                          |           |--P. fontinalis D07
                          |           |--P. laevidens D07
                          |           |--P. laticeps D07
                          |           |--P. paludosus D07
                          |           `--P. robustus D07
                          |--Rhinotitan [Epimanteoceratinae] WW97
                          |    `--R. mongoliensis WW97
                          |--Embolotherium Osborn 1929 D07 [Embolotheriinae WW97]
                          |    |--E. andrewsi D07
                          |    `--E. grangeri D07
                          `--Metatelmatheriinae WW97
                               |--Desmatotitan WW97
                               |--Hyotitan WW97
                               |--Acrotitan WW97
                               |--Arctotitan WW97
                               |--Qufutitan Wang & Wang 1997 WW97
                               |    `--*Q. zhoui Wang & Wang 1997 WW97
                               `--Metatelmatherium WW97
                                    |--M. cristatum WW97
                                    `--M. ultimum WW97

*Type species of generic name indicated


[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.

[F02] Froehlich, D. J. 2002. Quo vadis eohippus? The systematics and taxonomy of the early Eocene equids (Perissodactyla). Zoological Journal of the Linnean Society 134: 141–256.

[HUG17] Halliday, T. J. D., P. Upchurch & A. Goswami. 2017. Resolving the relationships of Paleocene placental mammals. Biological Reviews 92 (1): 521–550.

[HD04] Hooker, J. J., & D. Dashzeveg. 2004. The origin of chalicotheres (Perissodactyla, Mammalia). Palaeontology 47 (6): 1363–1386.

[M02] Mlíkovský, J. 2002. Cenozoic Birds of the World. Part 1: Europe. Ninox Press: Praha.

[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.

[WW97] Wang Y. & Wang J. 1997. A new brontothere from late Middle Eocene of Qufu, Shandong. Vertebrata PalAsiatica 35 (1): 68–77.

Last updated: 6 August 2018.

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