Eutheria
Placental Mammals
This tree diagram shows the relationships between several groups of organisms.
The root of the current tree connects the organisms featured in this tree to their containing group and the rest of the Tree of Life. The basal branching point in the tree represents the ancestor of the other groups in the tree. This ancestor diversified over time into several descendent subgroups, which are represented as internal nodes and terminal taxa to the right.
You can click on the root to travel down the Tree of Life all the way to the root of all Life, and you can click on the names of descendent subgroups to travel up the Tree of Life all the way to individual species.
For more information on ToL tree formatting, please see Interpreting the Tree or Classification. To learn more about phylogenetic trees, please visit our Phylogenetic Biology pages.
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Adkins, R. M. and R. L. Honeycutt. 1991. Molecular phylogeny of the superorder Archonta. Proceedings of the National Academy of Sciences (USA) 88:10317-10321.
Allard, M. W., B. E. McNiff, M. M. Miyamoto. 1996. Support for interordinal eutherian relationships with an emphasis on primates and their Archontan relatives. Molecular Phylogenetics and Evolution 5:78-88.
Amrine-Madsen, H., K.-P. Koepfli, R. K. Wayne and M. S. Springer. 2003. A new phylogenetic marker, apolipoprotein B, provides compelling evidence for eutherian relationships. Molecular Phylogenetics and Evolution 28(2):225-240.
Archibald, J. D. 1996. Fossil evidence for a Late Cretaceous origin of 'hoofed' mammals. Science 272:1150-1153.
Archibald, J. D., A. O. Averianov, and E. G. Ekdale. 2001. Late Cretaceous relatives of rabbits, rodents, and other extant eutherian mammals. Nature 414:62?65.
Arnason, U., A. Gullberg, S. Gretarsdottir, B. Ursing, and A. Janke. 2000. The mitochondrial genome of the sperm whale and a new molecular reference for estimating eutherian divergence dates. Journal of Molecular Evolution 50:569-578.
Arnason, U., A. Gullberg, and A. Janke. 1997. Phylogenetic analyses of mitochondrial DNA suggest a sister group relationship between Xenarthra (Edentata) and Ferungulates. Molecular Biology and Evolution 14:762-768.
Arnason, U., A. Gullberg, and A. Janke. 1999. The mitochondrial DNA molecule of the aardvark, Orycteropus afer, and the position of the Tubulidentata in the eutherian tree. Proceedings of the Royal Society of London Series B 266: 339-345.
Asher, R. J., J. Meng, J. R. Wible, M. C. McKenna, G. W. Rougier, D. Dashzeveg, and M. J. Novacek. 2005. Stem Lagomorpha and the antiquity of Glires. Science 307(5712):1091-1094.
Cao, Y., J. Adachi, A. Janke, S. Pääbo, and M. Hasegawa. 1994. Phylogenetic relationships among eutherian orders estimated from inferred sequences of mitochondrial proteins - instability of a tree based on a single gene. Journal of Molecular Evolution 39:519-527.
Cao, Y., A. Janke, P. J. Waddell, M. Westerman, O. Takenaka, S. Murata, N. Okada, S. Pääbo, and M. Hasegawa. 1998. Conflict among individual mitochondrial proteins in resolving the phylogeny of eutherian orders. Journal of Molecular Evolution 47:307-322.
Cao, Y., N. Okada, and M. Hasegawa. 1997. Phylogenetic position of guinea pigs revisited. Molecular Biology and Evolution 14:461-464.
Carter, A. M. 2001. Evolution of the placenta and fetal membranes seen in the light of molecular phylogenetics. Placenta 22:800-807.
Corbet,G.B. and J. E. Hill. 1992. The Mammals of the Indomalayan Region : A Systematic Review. Oxford University Press, Oxford.
Corneli, P. S. and R. H. Ward. 2000. Mitochondrial genes and mammalian phylogenies: Increasing the reliability of branch length estimation. Molecular Biology and Evolution 17:224-234.
Corneli, P. S. 2002. Complete mitochondrial genomes and eutherian evolution. Journal of Mammalian Evolution 9:281-305.
Estes, R. D. 1991. The Behavior Guide to African Mammals. University of California Press, Berkeley.
Debry, R. W. 1999. Maximum likelihood analysis of gene-based and structure-based process partitions, using mammalian mitochondrial genomes. Systematic Biology 48:286-299.
de Jong, W. W., M. A. M. van Dijka, C. Pouxa, G. Kappéa, T. van Rheedea, and O. Madsen. Indels in protein-coding sequences of Euarchontoglires constrain the rooting of the eutherian tree. Molecular Phylogenetics and Evolution 28(2):328-340.
D'Erchia, A. M., C. Gissi, G. Pesole, C. Saccone, and U. Arnason. 1996. The guinea-pig is not a rodent. Nature 381:597-600.
Douzery, E. and F. M. Cazeflis. 1995. Molecular evolution of the mitochondrial 12S rRNA in Ungulata (Mammalia). Journal of Molecular Evolution 41:622-636.
Easteal, S. 1999. Molecular evidence far the early divergence of placental mammals. BioEssays 21:1052-1058.
Emerson, G. L., C. W. Kilpatrick, B. E. McNiff, J. Ottenwalder, and M. W. Allard. 1999. Phylogenetic relationships of the order insectivora based on complete 12S rRNA sequences from mitochondria. Cladistics 15:221-230.
Foote, M., J. P. Hunter, C. M. Janis, and J. J. Sepkoski. 1999. Evolutionary and preservational constraints on origins of biologic groups: Divergence times of eutherian mammals. Science 283:1310-1314.
Frye, M. S. and S. B. Hedges. 1995. Monophyly of the order Rodentia inferred from mitochondrial DNA sequences of the genes for 12S ribosomal RNA, 16S ribosomal RNA, and transfer RNA valine. Molecular Biology and Evolution 12:168-176.
Gatesy, J. 1997. More DNA support for a Cetacea/Hippopotamidae clade: the blood-clotting protein gene gamma-fibrinogen. Molecular Biology and Evolution 14:537-543.
Gatesy, J., M. Milinkovitch, V. Waddell, and M. Stanhope. 1999. Stability of cladistic relationships between Cetacea and higher-level Artiodactyl taxa. Systematic Biology 48:6-20.
Gatesy, J., P. O'Grady, and R. H. Baker. 1999. Corroboration among data sets in simultaneous analysis: Hidden support for phylogenetic relationships among higher level artiodactyl taxa. Cladistics 15:271-313.
Gatesy, J. and M. A. O'Leary. 2001. Deciphering whale origins with molecules and fossils. Trends in Ecology and Evolution 16:562-570.
Geisler, J.H. and M.D. Uhen. 2003. Morphological support for a close relationship between hippos and whales. Journal of Vertebrate Paleontology 23:991?996.
Graur, D. 1993. Toward a molecular resolution of the ordinal phylogeny of the eutherian mammals. FEBS Letters 325:152-159.
Graur, D., L. Duret, and M. Gouy. 1996. Phylogenetic position of the order Lagomorpha (rabbits, hares and allies). Nature 379:333-335.
Graur, D., M. Gouy, and L. Duret. 1997. Evolutionary affinities of the order Perissodactyla and the phylogenetic status of the superordinal taxa Ungulata and Altungulata. Molecular Phylogenetics and Evolution 7:195-200.
Graur, D., W. A. Hide, and W.-H. Li. 1991. Is the guinea pig a rodent? Nature 351:649-652.
Graur, D. and D. Higgins. 1994. Molecular evidence for the inclusion of cetaceans within the order Artiodactyla. Molecular Biology and Evolution 11:357-364.
Halanych, K. M. 1998. Lagomorphs misplaced by more characters and fewer taxa. Systematic Biology 47:138-146.
Hasegawa, M., Y. Cao, J. Adachi, and T. Yano. 1992. Rodent polyphyly. Nature 355:595.
Hauf, J., P. J. Waddell, N. Chalwatzis, U. Joger, and F. K. Zimmermann. 1999. The complete mitochondrial genome sequence of the African elephant (Loxodonta africana), phylogenetic relationships of Proboscidea to other mammals, and D-loop heteroplasmy. Zoology - Analysis of Complex Systems 102:184-195.
Honeycutt, R. L. and R. M. Adkins. 1993. Higher-level systematics of eutherian mammals–an assessment of molecular characters and phylogenetic hypotheses. Annual Review of Ecology and Systematics 24: 279-305.
Huchon D., O. Madsen, M.J.J.B. Sibbald, K. Ament, M. J. Stanhope, F. Catzeflis, W. W. De Jong, and E. J. P. Douzery. 2002. Rodent phylogeny and a timescale for the evolution of glires: Evidence from an extensive taxon sampling using three nuclear genes. Molecular Biology and Evolution 19(7):1053-1065.
Hudelot, C., V. Gowri-Shankar, H. Jow, M. Rattray, and P. G. Higgs. 2003. RNA-based phylogenetic methods: application to mammalian mitochondrial RNA sequences. Molecular Phylogenetics and Evolution 28:241-252.
Irwin, D. M. and U. Arnason. 1994. Cytochrome b gene of marine mammals: phylogeny and evolution. Journal of Mammalian Evolution 2:37-55.
Ji, Q., Z.-X. Luo, C.-X. Yuan, J. R. Wible, J.-P. Zhang, and J. A. Georgi. 2002. The earliest known eutherian mammal. Nature 416:816-822.
Kingdon, J. 1997. The Kingdon Field Guide to African Mammals. Academic Press, London.
Kleineidam, R. G., G. Pesole, H. J. Breukelman, J. J. Beintema, R. A. Kastelein. 1999. Inclusion of cetaceans within the order Artiodactyla based on phylogenetic analysis of pancreatic ribonuclease genes. Journal of Molecular Evolution 48:360-368.
Kupfermann, H., Y. Satta, N. Takahata, H. Tich, and J. Klein. 1999. Evolution of Mhc-DRB introns: Implications for the origin of primates. Journal of Molecular Evolution 48:663-674.
Lavergne, A., E. Douzery, T. Stichler, F. M. Catzeflis, and M. S. Springer. 1996. Interordinal mammalian relationships: Evidence for paenungulate monophyly is provided by complete mitochondrial 125 rRNA sequences. Molecular Phylogenetics and Evolution 6:245-258.
Li, W., M. Gouy, P. M. Sharp, C. O'HUigin, and Y. Yang. 1990. Molecular Phylogeny of Rodentia, Lagomorpha, Primates, Artiodactyla, and Carnivora and Molecular Clocks. Proceedings of the National Academy of Sciences (USA) 87:6703-6707.
Liu, F. G. R. and M. M. Miyamoto. 1999. Phylogenetic assessment of molecular and morphological data for eutherian mammals. Systematic Biology 48:54-64.
Liu, F. G. R., M. M. Miyamoto, N. P. Freire, P. Q. Ong, M. R. Tennant, T. S. Young, and K. F. Gugel. 2001. Molecular and morphological supertrees for eutherian (placental) mammals. Science 291:1786-1789.
Luckett, W. P. and J.-L. Hartenberger. 1993. Monophyly or polyphyly of the order Rodentia: possible conflict between morphological and molecular interpretations. Journal of Mammalian Evolution 1:127-147.
Luckett, W. P. and N. Hong. 1998. Phylogenetic relationships between the orders Artiodactyla and Cetacea: a combined assessment of morphological and molecular evidence. Journal of Mammalian Evolution 5:127-182.
Madsen, O., P. M. T. Deen, G. Pesole, C. Saccone, and W. W. deJong. 1997. Molecular evolution of mammalian aquaporin-2: Further evidence that elephant shrew and aardvark join the paenungulate clade. Molecular Biology and Evolution 14:363-371.
Madsen, O., M. Scally, C. J, Douady, D. J. Kao, R. W. Debry, R. Adkins, H. M. Ambrine, M. J. Stanhope, W. W. DeJong, and M. S. Springer. 2001. Parallel adaptive radiations in two major clades of placental mammals. Nature 409 :610-614.
McNiff, B.E. and M. W. Allard. 1998. A test of Archonta monophyly and the phylogenetic utility of the mitochondrial gene 12S rRNA. American Journal of Physical Anthropology 107:225-241.
Milinkovitch, M. C. and J. G. M. Thewissen. 1997. Even-toed fingerprints on whale ancestry. Nature 388:622-624.
Misawa, K. and A. Janke. 2003. Revisiting the Glires concept––phylogenetic analysis of nuclear sequences. Molecular Phylogenetics and Evolution 28(2):320-327.
Miyamoto, M. M. 1996. A congruence study of molecular and morphological data for Eutherian mammals. Molecular Phylogenetics and Evolution 6:373-390.
Montgelard, C., F. M. Catzeflis, and E. Douzery. 1997. Phylogenetic relationships of artiodactyls and cetaceans as deduced from the comparison of cytochrome b and 12S rRNA mitochondrial sequences. Molecular Biology and Evolution 14:550-559.
Mouchaty, S. K., A. Gullberg, A. Janke, and U. Arnason. 2000. The phylogenetic position of the Talpidae within eutheria based on analysis of complete mitochondrial sequences. Molecular Biology and Evolution 17:60-67.
Murata, Y., M. Nikaido, T. Sasaki, Y. Cao, Y. Fukumoto, M. Hasegawa and N. Okada. 2003. Afrotherian phylogeny as inferred from complete mitochondrial genomes. Molecular Phylogenetics and Evolution 28(2):253-260.
Murphy, W. J., E. Eizirik, S. J. O'Brien, O. Madsen, M. Scally, C. J. Douady, E. Teeling, O. A. Ryder, M. J. Stanhope, W. W. de Jong, and M. S. Springer. 2001. Resolution of the early placental mammal radiation using Bayesian phylogenetics. Science 294:2348-2351.
Murphy, W. J., E. Eizirik, W. E. Johnson, Y. P. Zhang, O. A. Ryder, and S. J. O'Brien. 2001. Molecular phylogenetics and the origins of placental mammals. Nature 409:614-618.
Nikaido, M., A. P. Rooney, and N. Okada. 1999. Phylogenetic relationships among cetartiodactyls based on insertions of short and long interspersed elements: hippopotamuses are the closest extant relatives of whales. Proceedings of the National Academy of Sciences (USA) 96 :10261?10266.
Nishihara, H., Y. Satta, M. Nikaido, J. G. M. Thewissen, M. J. Stanhope, and N. Okada. 2005. A retroposon analysis of afrotherian phylogeny. Molecular Biology and Evolution 22(9):1823-1833.
Novacek, M. J. 1992. Fossils, topologies, missing data, and the higher level phylogeny of eutherian mammals. Systematic Biology 41:58-73.
Novacek, M. J. 1996. Where do rabbits and kin fit in? Nature 379:299-300.
Novacek, M. J., G. W. Rougier, J. R. Wible, M. C. McKenna, D. Dashzeveg, and I. Horovitz. 1997. Epipubic bones in eutherian mammals from the late Cretaceous of Mongolia. Nature 389:483-486.
O'Leary, M. A. and J. H. Geisler. 1999. The position of Cetacea within Mammalia: Phylogenetic analysis of morphological data from extinct and extant taxa. Systematic Biology 48:455-490.
O'Leary, M. A. 1999. Parsimony analysis of total evidence from extinct and extant taxa and the cetacean-artiodactyl question (Mammalia, Ungulata). Cladistics 15:315-330.
O'Leary M. A. 2001. The phylogenetic position of cetaceans: Further combined data analyses, comparisons with the stratigraphic record and a discussion of character optimization. American Zoologist 41: 487-506.
Onuma, M., T. Kusakabe, and S. Kusakabe. 1998. Phylogenetic positions of Insectivora in Eutheria inferred from mitochondrial cytochrome c oxidase subunit II gene. Zoological Science 15:139-145.
Ozawa, T., S. Hayashi, and V. M. Mikhelson. 1997. Phylogenetic position of Mammoth and Steller's sea cow within Tethytheria demonstrated by mitochondrial DNA sequences. Journal of Molecular Evolution 44:406-413.
Philippe, H. 1997. Rodent monophyly: Pitfalls of molecular phylogenies. Journal of Molecular Evolution 45:712-715.
Philippe, H. and E. Douzery. 1994. The pitfalls of molecular phylogeny based on four species, as illustrated by the Cetacea/Artiodactyla relationships. Journal of Mammalian Evolution 2:133-152.
Porter, C. A., M. Goodman, M. J. Stanhope. 1996. Evidence on mammalian phylogeny from sequences of exon 28 of the von Willebrand factor gene. Molecular Phylogenetics and Evolution 5:89-101.
Prothero, D.R and R.M. Schoch. 1993. Horns, Tusks, Hooves, and Flippers: The Evolution of Hoofed Mammals and Their Relatives. Princeton University Press, Princeton, New Jersey.
Pumo, D. E., P. S. Finamore, W. R. Franek, C. J. Phillips, S. Tarzami, and D. Balzarano. 1998. Complete mitochondrial genome of a neotropical fruit bat, Artibeus jamaicensis, and a new hypothesis of the relationships of bats to other eutherian mammals. Journal of Molecular Evolution 47:709-717.
Reynolds, J. E. and S. A. Rommel. 1999. Biology of Marine Mammals. Smithsonian Institution Press, Wahington, D.C.
Reyes, A., C. Gissi, G. Pesole, F. M. Catzeflis, and C. Saccone. 2000. Where do rodents fit? Evidence from the complete mitochondrial genome of Sciurus vulgaris. Molecular Biology and Evolution 17:979-983.
Reyes, A., G. Pesole, and C. Saccone. 1998. Complete mitochondrial DNA sequence of the fat dormouse, Glis glis: Further evidence of rodent paraphyly. Molecular Biology and Evolution 15:499-505.
Rose, K. D. 1996. On the origin of the order Artiodactyla. Proceedings of the National Academy of Sciences (USA) 93:1705-1709.
Rose, K. D. and J. D. Archibald, eds. 2005. The Rise of Placental Mammals. Origins and Relationships of the Major Clades. Johns Hopkins University Press.
Schmitz, J. and H. Zischler. 2003. A novel family of tRNA-derived SINEs in the colugo and two new retrotransposable markers separating dermopterans from primates. Molecular Phylogenetics and Evolution 28:341-349.
Shimamura, M., H. Yasue, K. Ohshima, H. Abe, H. Kato, T. Kishiro, M. Goto, I. Munechika, and N. Okada. 1997. Molecular evidence from retroposons that whales form a clade within even-toed ungulates. Nature 388:666-670.
Springer, M. S. , H. M. Amrine, A. Burk, and M. J. Stanhope. 1999. Additional support for Afrotheria and Paenungulata, the performance of mitochondrial versus nuclear genes, and the impact of data partitions with heterogeneous base composition. Systematic Biology 48:65-75.
Springer, M. S. , G. C. Cleven, O. Madsen, W. W. deJong, V. G. Waddell, H. M. Amrine, and M. J. Stanhope. 1997. Endemic African mammals shake the phylogenetic tree. Nature 388:61-64.
Springer, M. S. and W. W. deJong. 2001. Which mammalian supertree to bark up? Science 291:1709-1711.
Stanhope, M.J., O. Madsen, V. G. Waddell, G. C. Cleven, W. W. de Jong, and M. S. Springer. 1998. Highly congruent molecular support for a diverse superordinal clade of endemic African mammals. Molecular Phylogenetics and Evolution 9:501-508.
Stanhope, M. J., M. R. Smith, V. G. Waddell, C. A. Porter, M. S. Shivji, and M. Goodman. 1996. Mammalian evolution and the interphotoreceptor retinoid binding protein (IRBP) gene: Convincing evidence for several superordinal clades. Journal of Molecular Evolution 43:83-92.
Stanhope, M. J., V. G. Waddell, O. Madsen, W. de Jong, S. B. Hedges, G. C. Cleven, D. Kao, and M. S. Springer. 1998. Molecular evidence for multiple origins of Insectivora and for a new order of endemic African insectivore mammals. Proceedings of the National Academy of Sciences (USA) 95:9967-9972.
Sullivan, J. and D. L. Swofford. 1997. Are guinea pigs rodents? The importance of adequate models in molecular phylogenetics. Journal of Mammalian Evolution 4:77-86.
Szalay, F. S., M. J. Novacek, and M. C. McKenna (eds.) 1993. Mammal Phylogeny. Volume 2. Placentals. Springer Verlag, New York.
Thewissen, J.G.M. and S.I. Madar. 1999. Ankle morphology of the earliest Cetaceans and its implications for the phylogenetic relations among Ungulates. Systematic Biology 48:21-30.
van Dijk, M. A. M., E. Paradis, F. Catzeflis, and W. W. De Jong. 1999. The virtues of gaps: Xenarthran (Edentate) monophyly supported by a unique deletion in alphaA-crystallin. Systematic Biology 48: 94-106.
van Dijk, M. A. M., O. Madsen, F. Catzeflis, M. J. Stanhope, W. W. de Jong, and M. Pagel. 2001. Protein sequence signatures support the African clade of mammals. Proceedings of the National Academy of Sciences (USA) 98:188-193.
Waddell, P.J., Y. Cao, J. Hauf, and M. Hasegawa. 1999. Using novel phylogenetic methods to evaluate mammalian mtDNA, including amino acid-invariant sites-LogDet plus site stripping, to detect internal conflicts in the data, with special reference to the positions of hedgehog, armadillo, and elephant. Systematic Biology 48:31-53.
Waddell, P.J., Y. Cao, M. Hasegawam and D. P. Mindell. 1999. Assessing the Cretaceous superordinal divergence times within birds and placental mammals by using whole mitochondrial protein sequences and an extended statistical framework. Systematic Biology 48:119-137.
Waddell, P.J., N. Okada, and M. Hasegawa. 1999. Towards resolving the interordinal relationships of placental mammals. Systematic Biology 48:1-5.
Waddell, P. J. and S. Shelley. 2003. Evaluating placental inter-ordinal phylogenies with novel sequences including RAG1, -fibrinogen, ND6, and mt-tRNA, plus MCMC-driven nucleotide, amino acid, and codon models. Molecular Phylogenetics and Evolution 28(2):197-224.
Xu, X. F., A. Janke, and U. Arnason. 1996. The complete mitochondrial DNA sequence of the greater Indian rhinoceros, Rhinoceros unicornis, and the phylogenetic relationship among Carnivora, Perissodactyla, and Artiodactyla (plus Cetacea). Molecular Biology and Evolution 13:1167-1173.
Information on the Internet
- African Mammals Databank. A GIS-based databank on the distribution and conservation of all the big and medium-sized mammals over the whole African continent.
- ZOO in the wild. A virtual zoo with information on African and European mammals.
- African Wildlife Foundation
- Mammals of Africa. NatureExplorer.com.
- Les grands Mammifères du Niokolo-Badiar. A guide to West-African mammals, en français.
- Aubrey Manning Gallery Collection of ungulates, rodents, carnivores and primates.
- The Ultimate Ungulate Page. Your Guide to the World's Hoofed Mammal Species.
- New Model May Resolve Paleontology Family Feud. Information about research on the origin of placental mammals. UniSci Science and Research News.
- Fossil Eutheria:
- Cretaceous Critters. Two 80 million-year-old fossil mammals, including a new species, shed light on early reproduction. Information about Ukhaatherium nessovi and a zalambdalestid. American Museum of Natural History.
- Introduction to the Desmostylia. UCMP Berkeley.
- Introduction to the Embrithopoda. UCMP Berkeley.