After Archibald, 1996, McKenna & Bell, 1997, Janis, Archibald, Cifelli, Lucas, Schaff, Schoch & Williamson, 1998, Averianov, 2000 and de Muizon & Cifelli, 2000
<==o Ungulatomorpha Archibald, 1996 |?- †Zhelestidae genera et species indet. [UA 8699] [U. Cret. Mad.] [Averianov, Archibald & Martin, 2003] |?- †Kumlestes olzha Nesov, 1985 [paraphyletic Zhelestidae Nesov, 1985] |?- †Kumsuperus avus Nesov, 1984 [paraphyletic Zhelestidae Nesov, 1985] |?-o †Sorlestes Nesov, 1985 [paraphyletic Zhelestidae Nesov, 1985] | |-- †S. budan Nesov, 1985 | `-- †S. kara Nesov, 1993 |?- †Taslestes inobservabilis Nesov, 1982 [paraphyletic Zhelestidae Nesov, 1985] |-- †Aspanlestes aptap Nesov, 1985 [paraphyletic Zhelestidae Nesov, 1985] `--+-- †Avitotherium utahensis Cifelli, 1990d [paraphyletic Zhelestidae Nesov, 1985] |?-o †Gallolestes Lillegraven, 1976 | |-- †G. pachymandibularis Lillegraven, 1976 | `-- †G. agujaensis Cifelli, 1994 |?- †Lainodon orueetxebarriai Gheerbrant & Astibia, 1994 |?-o †Labes Sigè, 1992, in Pol, Buscalioni, Carballeira, Francés, López Martinez, Marandat, Moratella, Sanz, Sigé, Villatte, 1992 | |-- †L. quintanillensis Sigé, 1992, in Pol et al., 1992 | `-- †L. garimondi Sigé, 1992, in Pol et al., 1992 |?- †Valentinella vitrollense Tabuce, Vianey-Liaud & Garcia, 2004 `--+-- †Alostera saskatchewanensis Fox, 1989 [incl.? Paranyctoides megaceros Lillegraven & McKenna 1986] [paraphyletic Zhelestidae Nesov, 1985] `--+--o †"Zhelestinae" | |--o †Zhelestes Nesov, 1985 [paraphyletic Zhelestidae Nesov, 1985] | | |-- †Z. temirkazyk Nesov, 1985 | | `-- †Z. bezelgen Nesov, 1987 | `--o †Parazhelestes Nesov, 1993 [paraphyletic Zhelestidae Nesov, 1985] | |-- †P. robustus Nesov, 1993 [paraphyletic Zhelestidae Nesov, 1985] | `-- †P. minor Nesov, Archibald & Kielan-Jaworowska, 1998 [paraphyletic Zhelestidae Nesov, 1985] `--+-- †Eoungulatum kudukensis Nesov et al., 1998 [??] [paraphyletic Zhelestidae] `--o Ungulata Linnaeus 1766 (ungulates; kaviokyntiset) |?- †Kharmerungulatum vanvaleni Prasad, Verma, Sahni, Parmar & Khosla, 2007; U.Cret. SAs. (India) |?-o †Hilalia Maas, Thewissen, Sen, Kazanci & Kappelman, 2001 | |-- †H. sezerorum Maas, Thewissen, Sen, Kazanci & Kappelman, 2001 | `--+-- †H. saribeya Maas, Thewissen, Sen, Kazanci & Kappelman, 2001 [type of Hilalia] | `--+-- †H. selanneae Maas, Thewissen, Sen, Kazanci & Kappelman, 2001 | `-- †H. robusta Maas, Thewissen, Sen, Kazanci & Kappelman, 2001 |?- Tubulidentata (putkihampaiset, aardwarks) |?- †Dinocerata (uintatheriids) |-- Altungulata [Paenungulata] (horse- and elephant-like ungulates; hevos- ja norsumaiset ungulaatit) `--o Eparctocyona |== †Procreodi sensu McKenna & Bell, 1997 [paraphyletic assemblage of basal ungulates??] `--+-- †Bulbulodentata Archibald, 1998 [Condylartha sensu McKenna & Bell, 1997, incl. †Panameriungulata de Muizon, & Cifelli, 2000 and Meridiungulata] (etelä-amerikkalaiset ungulaatit; South American ungulates) `--+-- †Arctostylopida sensu Cifelli and Schaff, 1998 `-- "Paraxonia" (artiodactyl-like ungulates; sorkkaeläintensukuiset ungulaatit)
Note:
Taxonomy and phylogeny of basal ungulates is quite a mess. A large number of scientist have studied these animals over the years, but there is still no consensus about the relationships of main ungulate-groups.
I have here mainly followed McKenna & Bell (1997), but it's almost certain that some basal orders they recognized, such as Procreodi and Condylartha are paraphyletic. Actually all early Tertiary ungulates were once placed in Condylartha (Carroll, 1988).
One of the fun aspects when doing taxonomic research of early ungulates, are the rather funny names given by Leigh Van Valen. OK, I'm a Tolkien-fan too, but really... Did he have to...?
Reference(s):
- Alroy, J., 2002: North American fossil mammal systematics database.
–iNet: North American fossil mammal systematics database: http://www.nceas.ucsb.edu/~alroy/nafmsd.html - Archibald, J. D., 1996: Fossil evidence for a Late Cretaceous origin of "hoofed" mammals.
–Science: Vol. 272, 24 May, pp. 1150-1153 - Averianov, A. O., 2000: Mammals from the Mesozoic of Kirgizstan, Uzbekistan, Kazakhstan and Tadzhikistan. 627-652
in Benton, M. J., Shishkin, M. A., Unvin, D. M. & Kurochkin, E. N., (eds.) 2000: The Age of Dinosaurs in Russia and Mongolia.
–Cambridge University Press, Cambridge, 2000 xxxix-696 - Averianov, A. O., Archibald, J. D. & Martin, T., 2003: Placental nature of the alleged marsupial from the Cretaceous of Madagascar.
–Acta Palaeontologica Polonica: Vol. 48, #1, pp. 149-151 - 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. - Cifelli, R. L., 1990: Cretaceous mammals of Southern Utah. IV. Eutherian mammals from the Wahweap (Aquilan) and Kaiparowits (Judithian) Formations.
–Journal of Vertebrate Paleontology: Vol. 10, #3, pp. 346-360 - Cifelli, R. L. & Schaff, C. R., 1998: Arctostylopida. 332-336.
in Janis, C. M., Scott, K. M. & Jacobs, L. L., 1998: Evolution of Tertiary mammals of North America.
–Cambridge University Press, New York. 1998 - Gheerbrant, E., Sudre, J., Iarochene, M. & Moumni, A., 2001: First ascertained african "condylarth" mammals (primitive ungulates: cf. Bulbulodentata and cf. Phenacodonta) from the earlies Ypresian of the Ouled Abdoun Basin, Marocco.
–Journal of Vertebrate Paleontology: Vol. 21, #1, pp. 107-118 - Gingerich, P. D., Abbas, S. G. & Arif, M., 1997: Early Eocene Quettacyon parachai (Condylarthra) from the Ghazij Formation of Baluchistan (Pakistan): Oldest Cenozoic land mammal from South Asia.
–Journal of Vertebrate Paleontology: Vol. 17, #4, pp. 629-637 - Gingerich, P. D., Arif, M., Khan, I. H., Clyde, W. C. & Bloch, J. I., 1999: Machocyon abbasi, a new Early Eocene quettacyonid (Mammalia, Condylartha) from the Middle Ghazij Formation of Mach and Daghari coal fields, Baluchistan (Pakistan).
–Contributions from the Museum of Paleontology, The University of Michigan: Vol. 30, #9, pp. 233-250 - Luo, Z.-X., 1991: Variability of dental morphology and the relationships of the earliest arctocyonid species.
–Journal of Vertebrate Paleontology: Vol. 11, #4, pp. 452-471 - Maas, M. C., Thewissen, J. G. M., Sen, S., Kazanci, N. & Kappelman, J., 2001: Enigmatic new ungulates from the early Middle Eocene of central Anatolia, Turkey.
–Journal of Vertebrate Paleontology: Vol. 21, #3, pp. 578-590 - McKenna, M. C. & Bell, S. K., (eds.) 1997: Classification of mammals – above the species level.
–Columbia University Press, New York, 1997, xii-631 - de Muizon, C. & Cifelli, R. L., 2000: The "condylarths" (archaic Ungulata, Mammalia) from the early Palaeocene of Tiupampa (Bolivia): implications on the origin of the South American ungulates.
–Geodiversitas: Vol. 22, #1, pp. 47-150 - Janis, C. M., Archibald, J. D., Cifelli, R. L., Lucas, S. L., Schaff, C. R., Schoch, R. M. & Williamson, T. E., 1998: Archaic ungulates and ungulate like mammals. 247-259
in Janis, C. M., Scott, K. M. & Jacobs, L. L., 1998: Evolution of Tertiary mammals of North America.
–Cambridge University Press, New York. 1998 - Prasad, G. V. R., Verma, O., Sahni, A., Parmar, V. & Khosla, A., 2007: A Cretaceous hoofed mammal from India.
–Science: Vol. 318, pp. 937 [doi: 10.1126/science.1149267] - Tabuce, R., Vianey-Liaud, M. & Garcia, G., 2004: A eutherian mammal in the latest Cretaceous of Vitrolles, southern France.
–Acta Palaeontologica Polonica: Vol. 49, #3, pp. 347-356