After Hu, Meng, Li & Wang, 2009, and O'Leary et al., 2013
Alternative phylogeny of Placentalia after Welker et al, 2015
Alternative phylogeny of Placentalia after Hu, Meng, Li, & Wang, 2009
Alternative phylogeny of selected basal eutherians after Wible, Rougier, Novacek & Asher, 2007
Alternative phylogeny of selected basal eutherians after Archibald & Averianov, 2006
Alternative phylogeny of selected basal eutherians after Ji, Luo, Yuan, Wible, Zhang & Georgi, 2002
Alternative molecular phylogeny of Placentalia after Madsen et al., 2001
Alternative molecular phylogeny of Placentalia after Murphy et al., 2001
Alternative molecular phylogeny of Placentalia after Waddell, Okada & Hasegawa 1999
Alternative phylogeny of Eutheria after McKenna & Bell, 1997
<==o Eutheria Gill, 1872 non Huxley, 1880 [Placentaria sensu lato] (istukkanisäkkäät) |-- †Acristatherium yanensis Hu, Meng, Li & Wang, 2009; E.Cret. EAs. |?- †Hovurlestes noyon Lopatin & Averianov, 2018; E.Cret. EAs. |-- †Juramaia sinensis Luo, Yuan, Meng & Ji, 2011; M.Jur. EAs. |-- †Otlestes meiman Nesov, 1985 `--+--+-- †Prokennalestes trofinovi Kielan-Jaworowska & Dashzeveg, 1989 [incl. P. minor Kielan-Jaworowska & Dashzeveg, 1989] | `-- †Eomaia scansoria Ji, Luo, Yuan, Wible, Zhang & Georgi, 2002; E. Cret. EAs. `--+-- †Murtoilestes abramovi (Averianov & Skutschas, 2000) Averianov & Skutschas, 2001 [Prokennalestes abramovi Averianov & Skutschas, 2000] `--+-- †Bobolestes zenge Nessov, 1985; L.Cret. CAs. [Bobotheriidae Nessov, 1989] `--+-- †Montanalestes keeblerorum (Cifelli, 1999) Cifelli, 2000 [Montanalestes keebleri Cifelli, 1999] `--+--+-- †Eozhelestes mangit Nesov, 1997 | `--o †Paranyctoides Fox, 1979 sensu Averianov & Archibald, 2013 [†Paranyctidae (nomen nudum)] [are Zhelestids?] | |-- †P. sternbergi Fox, 1979 [incl.? Paranyctoides maleficus Fox, 1984; Paranyctoides “Wahweap” sp. A (Cifelli, 1990); Paranyctoides “Wahweap” sp. B (Cifelli, 1990); Paranyctoides “Kaiparowits” sp. A (Cifelli, 1990); Paranyctoides “Kaiparowits” sp. B (Cifelli, 1990)] | `-- †P. quadrans (Nessov, 1982) [incl.? Paranyctoides aralensis Nessov, 1993; Marsasia sp. Averianov & Kielan-Jaworowska, 1999] Averianov & Archibald, 2012 `--+-- †Zhelestidae Nessov, 1985 `--+--+--†Bulaklestes kezbe Nessov, 1985 | `--+-- †Uchkudukodon nessovi (McKenna, Kielan-Jaworowska & Meng, 2000) Archibald & Averianov, 2006 [Daulestes nessovi McKenna, Kielan-Jaworowska & Meng, 2000] | `--o †Daulestes Trofimov & Nessov, in Nessov & Trofimov, 1979 [Taslestes Nessov, 1982; Kumlestes Nessov, 1982] [?--> †Palaeoryctidae] | |-- †D. kulbeckensis Trofimov & Nessov, in Nessov & Trofimov, 1979 [Kumlestes olzha Nessov, 1982] | `-- †D. inobservabilis (Nessov, 1982) [Taslestes inobservabilis Nessov, 1982] |--o †Asioryctitheria Novacek, Rougier, Wible, McKenna, Dashzeveg & Horovitz, 1997 [Kennalestoidea Kielan-Jaworowska, 1981] | |--o †Asioryctidae Kielan-Jaworowska, 1981 [incl. Kennalestidae Kielan-Jaworowska, 1981 sensu Archibald & Averianov, 2006; Asioryctinae (Kielan-Jaworowska, 1981) Archibald & Averianov, 2006] | | |-- †Ukhaatherium nessovi Novacek, Rougier, Wible, McKenna, Dashzeveg & Horovitz, 1997 | | `-- †Asioryctes nemegetensis Kielan-Jaworowska, 1975 | `--o †Kennalestidae Kielan-Jaworowska, 1981 | |-- †Kennalestes gobiensis Kielan-Jaworowska, 1969 | `-- †Sailestes quadrans Nessov, 1982 |--o †Didelphodonta McKenna, 1995 [paraphyletic †Cimolesta McKenna, 1995] | `-- †Cimolestidae Marsh, 1889 [taxonomic history] `--+--+?- †Sahnitherium rangapurensis Rana & Wilson, 2003 | `--o †Deccanolestes Prasad & Sahni, 1988 [“Adapisoriculidae” in partim?] | |-- †D. hislopi Prasad & Sahni, 1988 | |-- †D. robustus Prasad, Jaeger, Sahni, Gheerbrant & Khajuria, 1994 | `-- †D. narmadensis Prasad, Verma, Gheerbrant, Goswami, Khosla, Parmar & Sahni, 2010 `--+-- †Zalambdalestidae Gregory & Simpson, 1926 `--o Placentaria Owen, 1837 sensu stricto [Placentalia Owen, 1837; Preptotheria McKenna, 1975, incl. Therictoidea Gregory, 1910; Tokotheria McKenna, 1975] |-- Edentata G. Cuvier, 1798 (edentates; vajaahampaiset) `--o Epitheria McKenna, 1975 |--o Afrotheria Stanhope et al., 1998 | |--o Afroinsectiphillia | | |?-o †Ptolemaiida Simons & Bown, 1995 | | | |-- †Kelba quadeema Savage, 1965; U. Mio. NEAf. [incl. Kenyalutra songhorensis Schmidt-Kittler, 1987; Ndamathaia kubwa Jacobs et al., 1987 sensu Morales et al., 2000] [Kelbidae Cote et al, 2007] [arctocyonid creodont (Savage, 1965); ptolemaiid pantolestan (Van Valen, 1967); Carnivora incertae sedis (McKenna & Bell, 1997); hemigaline viverrid (Morales, Pickford, Salesa & Soria, 2000)] | | | `--o Ptolemaiidae Osborn, 1908 | | | |-- †Qarunavus meyeri Simons & Gingerich, 1974; L.Eoc. NEAf. | | | |--o †Cleopatrodon Bown & Simons, 1987; L.Eoc. NEAf. | | | | |-- †C. ayeshae Bown & Simons, 1987 | | | | `-- †C. major Bown & Simons, 1987 | | | `--o †Ptolemaia Osborn, 1908 [non †Ptolemaia McCutcheon & Wilson, 1961, a fossil rugose coral (Cnidaria: Anthozoa: †Rugosa)]; L.Eoc. NEAf. | | | |-- †P. lyonsi Osborn, 1908 | | | `-- †P. grangeri Bown & Simons, 1987 | | |-- Afrosoricida Stanhope et al., 1998 [Lipotyphla Haeckel, 1866 (in partim)] | | `--+-- †Leptictidae Gill, 1872 [Leptictida McKenna, 1975 (in partim); Proeutheria] | | `-- Macroscelidea Butler, 1956 [Macroscelides Peters, 1864; Macroscelidiformes Kinman, 1994] (norsupäästäiset; sengit; elephant shrews, sengis) | `--+-- Tubulidentata Huxley, 1872 (aardwarks; putkihampaiset) | `--o Paenungulata Simpson, 1945 | |-- Hyracoidea Huxlay, 1869 (hyraxes; tamaanit) | `--+-- †Thomashuxleyia externa Ameghino, 1901, L.Eoc. SA. [Isotemnidae Ameghino, 1897] | `--+-- †Carodnia [Xenungulata; Pyrotheria] | `-- Uranotheria Simpson, 1945 (elephant-like ungulates; norsumaiset ungulaatit) `--o Boreoeutheria |--o Euarchontoglires Murphy & Eizirik, 2001 [Supraprimates] | |-- Archonta Gregory, 1910 (primate-like eutherians; kädellistensukuiset) | `-- Glires Linnaeus, 1758 [Gliriformes Wyss & Meng, 1996] (rodent-like eutherians; jyrsijä-jäniseläimet) `--o Laurasiatheria Waddell et al., 1999 |-- Eulipotyphla [Lipotyphla Haeckel, 1866 (in partim)] (modern insect-eaters; nykyhyönteissyöjät) `--+--o Pholidotamorpha Gaudin et al. 2009 | |== †“Palaeanodonta” Matthew, 1918 [Basal Pholidota sensu McKenna & Bell, 1997] | `-- Pholidota Weber, 1904 (pangolins; muurahaiskävyt) `--+--o Ferae Linnaeus, 1758 | |-- †Creodonta Cope, 1875 (kreodontit; creodonts) | `-- Carnivoramorpha Kalandadze & Rautian, 1992 (modern carnivores and their extinct relatives; nykypetoeläinmäiset nisäkkäät) `--o Ungulata Linnaeus 1766 (in partim) (ungulates; kaviokyntiset) |--+--+-- †Hyopsodontidae Lydekker, 1889 | | `-- †Apheliscidae Matthew, 1918 | `--+-- †Phenacodonta McKenna, 1975 | `--+-- †Didolodus multicuspis Ameghino, 1897 [Enneocodus, Lonchoconus, Paulogervaisia, Cephanodus, Nephacodus, Argyrolambda] | `-- †Litopterna Ameghino, 1889 `--+-- †Mesonychia Van Valen, 1966 [Acreodi] (mesonykit; mesonychians) `--+--o †Rodhocetus Gingerich et al., 1995a | |-- †R. balochistanensis Gingerich, ul Haq, Zalmout, Khan & Malkani, 2001 | `-- †R. kasrani Gingerich et al., 1995a `--o Euungulata |-- Artiodactyla Owen, 1848 [“Paraxonia” Marsh, 1884] (sorkkaeläintensukuiset ungulaatit; artiodactyl-like ungulates; even-toed ungulates) `-- Perissodactyla Owen, 1848 [Mesaxonia Marsh, 1884 sensu Prothero & Schoch, 1989] (kavioeläimet)
Alternative phylogenies of Eutheria:
After Welker et al, 2015 (orders only):<==o Placentaria |--o Afrotheria | |-- Afrosoricida | `--+--+-- Macroscelidea | | `-- Tubulidentata | `--+-- Proboscidea | `--+-- Hyracoidea | `-- Sirenia `--+-- Xenarthra `--+--+-- Primates | `--+-- Scandentia | `--+-- Lagomorpha | `-- Rodentia `--+-- Eulipotyphla `--+-- Chiroptera |-- Carnivora `--+-- Artiodactyla `--+-- †Meridiungulata `-- Perissodactyla Reference(s):
|
After Hu, Meng, Li, & Wang, 2009<==o Eutheria |-- †Acristatherium |-- †Juramaia |-- †Otlestes `--+--+-- †Prokennalestes | `-- †Eomaia `--+-- †Murtoilestes `--+-- †Bobolestes `--+-- †Montanalestes `--+--+-- †Eozhelestes | `-- †Paranyctoides `--+-- †Zhelestidae `--+--+-- †Bulaklestes | `--+-- †Uchkudukodon | `-- †Daulestes |-- †Asioryctitheria |-- †Didelphodonta `--+--+?- †Sahnitherium | `-- †Deccanolestes `--+-- †Zalambdalestidae `--o Placentaria |-- Leptictida `--+--+-- †Purgatorius | `-- Eparctocyona `--+--+-- Carnivoramorpha | `--+-- “Paraxonia” | `--+-- †“Phenacodonta” | `?- Perissodactyla `--+-- Archonta |-- Glires `--+--o Insectivora | `-- Lipotyphla `--+--+-- Macroscelidea | `-- Tubulidentata |-- Edentata `-- Uranotheria Reference(s):
|
Selected basal eutherians after Wible, Rougier, Novacek & Asher, 2007<==o Eutheria |--+-- †Murtoilestes | `--+-- †Prokennalestes | `-- †Eomaia `--+-- †Bobolestes `--+-- †Montanalestes `--+-- †Zhelestidae `--+--+-- †Eozhelestes | `-- †Paranyctoides `--+--+-- †Asioryctitheria | `--o †Cimolestidae [=? Didelphodonta] | |-- †Cimolestes | `--+-- †Batodon | `-- †Maelestes `--+-- †Deccanolestes `--+-- †Zalambdalestidae `--+--+-- †Gypsonictops | `-- †Leptictis `--+--+-- †Purgatorius | `--+-- †Protungulatum | `-- †Oxyprimus `-- Placentalia Reference(s):
|
Selected basal eutherians after Archibald & Averianov, 2006<==o Eutheria |-- †Eomaia `--+-- †Prokennalestes `--+-- †Bobolestes `--+--o †Asioryctitheria | |-- †Uchkudukodon | `--+-- †Daulestes kulbeckensis | |-- †Daulestes inobservabilis | `--+-- †Bulaklestes | `--o †Asioryctidae | |-- †Kennalestes | `--o †Asioryctinae | |-- †Ukhaatherium | `-- †Asioryctes `--+--o †Zhelestidae | |-- †Zhelestes | |-- †Aspanlestes | `--o †Parazhelestes | |-- †P. robustus | `-- †P. mynbulakensis `--+--o †Zalambdalestidae | |-- †Zalambdalestes | `-- †Kulbeckia `?- Placentaria Reference(s):
|
Selected basal eutherians after Ji, Luo, Yuan, Wible, Zhang & Georgi, 2002
<==o Eutheria
|-- †Eomaia
|-- †Prokennalestes
|-- †Murtoilestes
`--+-- †Montanalestes
|--+-- †Asioryctes
| |-- †Kennalestes
| |-- †Ukhaatherium
| `-- †Zalambdalestes
`--+-- †Daulestes
`--+--+-- †Cimolestes
| `-- †Gypsonictops
`--+-- †Aspanlestes
`--+-- †Eoungulatum
`--+-- †Proungulatum
`-- Erinaceus
Reference(s):
|
After Murphy et al, 2001 (orders only):<==o Placentaria |?- †Meridiungulata |--o Afrotheria | |-- Afrosoricida | `--+--+-- †Microhyus | | `-- Macroscelidea | `--o Pseudoungulata | |-- Tubulidentata | `--o Paenungulata | |-- Hyracoidea | `--o Tethytheria | |-- Sirenia | `--o Behemota | |-- †Desmostylia | `-- Proboscidea `--+-- Xenarthra `--o Boreoeutheria |--+--o Glires | | |-- Lagomorpha | | `-- Rodentia | `--o Euarchonta | |-- Scandentia | `--o Primatomorpha | |-- Dermoptera | `-- Primates `--o Laurasiatheria |?- Eulipotyphla `--o Scrotifera |-- Chiroptera `--o Ferungulata |-- Artiodactyla `--o Zooamata |-- Perissodactyla `--o Ferae |-- Pholidota `-- Carnivora Reference(s):
|
After Madsen et al., 2001 (orders only):<==o Placentaria |--o Atlantogenata | |-- Xenarthra | `--+?- †Meridiungulata | `--o Afrotheria | |--+-- Tubulidentata | | `--+-- Afrosoricida | | `--+-- †Microhyus | | `-- Macroscelidea | `--o Paenungulata | |-- Hyracoidea | `--o Tethytheria | |-- Sirenia | `--o Behemota | |?- †Desmostylia | `-- Proboscidea `--o Boreoeutheria |--+-- Scandentia | `--+-- Dermoptera | `--+-- Primates | `--o Glires | |-- Lagomorpha | `-- Rodentia `--o Laurasiatheria |?- Eulipotyphla `--+--o Ferae | |-- Pholidota | `-- Carnivora `--+-- Chiroptera `--+-- Perissodactyla `-- Artiodactyla Reference(s):
|
After Waddell, Okada & Hasegawa 1999 (orders only):<==o Placentaria |--o Atlantogenata | |-- Xenarthra | `--+?- †Meridiungulata | `--o Afrotheria | |--o Afrosoricida | | |-- Tenrecoidea | | `-- Chrysochloroidea | `--+--+-- †Microhyus | | `-- Macroscelidea | `--o Pseudoungulata | |-- Tubulidentata | `--o Paenungulata | |-- Hyracoidea | `--o Tethytheria | |-- Sirenia | `--o Behemota | |-- †Desmostylia | `-- Proboscidea `--+--+--o Glires | | |-- Lagomorpha | | `-- Rodentia | `--o Euarchonta | |-- Scandentia | `--o Primatomorpha | |-- Dermoptera | `-- Primates `--o Laurasiatheria |?- Eulipotyphal `--o Scrotifera |-- Chiroptera `--o Ferungulata |-- Artiodactyla `--o Zooamata |-- Perissodactyla `--o Ferae |-- Pholidota `-- Carnivora Reference(s):
|
After McKenna & Bell, 1997<==o Eutheria |-- †Bobolestes |-- †Otlestes |-- †Murtoilestes |-- †Eomaia `--o Placentaria |-- Edentata `--o Epitheria |?- †Hyotheridium |?- †Deccanolestes |?-o †Asioryctitheria | `--o Asioryctidae | |-- †Asioryctes | `-- †Ukhaatherium |-- Leptictida `--o Preptotheria |--+-- Archonta | `-- Anagalida `--+--o Insectivora | `-- Lipotyphla `?-o Ferungulata |-- Ungulatomorpha `--o Ferae |== Cimolesta `--+-- †Creodonta `-- Carnivoramorpha Reference(s):
|
Basal Eutheria incertae sedis:
†Hyotheridium dobsoni Gregory & Simpson, 1926 †Beleutinus orlovi Bazhanov, 1972 [a zalambdalestid or deltatheroid?] †Cretasorex Nesov & Gureyev, 1981 [identification doubtful, possibly nomen dubium] †Eodesmatodon Zheng & Chi, 1978 †Eutrochodon Roth, 1903 †Helioseus Sudre, 1979 †Horolodectes sunae Scott, Webb & Fox, 2006 †Idiogenomys ozziei Ostrander, 1983 †Neodesmostylus Khomenko, 1927 †Obtususdon hanhuaensis Xu, 1977 †Prozalambdalestes simpsoni †Telacodon laevis Marsh, 1892 [see Leptictida] †Tingamarra porterorum Godthelp, Archer, Cifelli, Hand & Gilkeson, 1992 †Wanotherium xuanchengensis Tang & Yan, 1976 †Veratalpa lugdunensiana Ameghino, 1905 [possibly a rodent, is based on an astragalus (Hutchison, 1974)] †Bibymalagasia MacPhee, 1994 †Plesiorycteropodinae Patterson, 1975 †Plesiorycteropus Filhol, 1895 [Myoryctes Forsyth Major, 1908; Majoria Thomas, 1915] R. Mad. †P. germainepetterae MacPhee, 1994 †P. madagascariensis Filhol, 1895 [Majoria; Myoryctes rapeto]
Reference(s):
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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. & Archibald, J. D., 2013: Variation and taxonomy of Asiamerican eutherian mammal Paranyctoides.
–Canadian Journal of Earth Sciences: Vol. 50, #9, pp. 895-903 [doi: 10.1139/cjes-2013-0051] - Averianov, A. O. & Skutschas, P. P., 2001: A new genus of eutherian mammal from the Early Cretaceous of Transbaikalia, Russia.
–Acta Palaeontologica Polonica: Vol. 46, #3, pp. 431-436 - 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 - Dykes, T., 2002: "Basal" Eutheria. –iNet: Mesozoic mammals – an internet directory: http://home.arcor.de/ktdykes/baseuth.htm
- Hu, Y., Meng, J., Li, C. & Wang, Y., 2009: New basal eutherian mammal from the Early Cretaceous Jehol biota, Liaoning, China.
–Proceedings of the Royal Society of London: Biological series: Vol. 277, #1, pp. 229-236 [doi: 10.1098/rspb.2009.0203] - Ji, Q., Luo, Z.-X., Yuan, C.-X., Wible, J. R., Zhang, J.-P. & Georgi, J. A., 2002: The earliest known eutherian mammal.
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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 - Lopatin, A. V. & Averianov, A. O., 2018: New stem placental mammal from the Early Cretaceous of Mongolia. (in Russian).
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–Nature: Vol. 476, pp. 442-445 [doi: 10.1038/nature10291] - McKenna, M. C. & Bell, S. K., (eds.) 1997: Classification of mammals – above the species level.
–Columbia University Press, New York, 1997, xii-631 - McKenna, M. C., Kielan-Jaworowska, Z. & Meng, J., 2000: Earliest eutherian mammal skull from the Late Cretaceous (Coniacian) of Uzbekistan.
–Acta Paleontologica Polonica: Vol: 45, #1, pp. 1-54 - Montellano-Ballesteros, M., Fox, R. C. & Scott, C. S., 2013: Species composition of the Late Cretaceous eutherian mammal Paranyctoides Fox.
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–Acta Palaeontologica Polonica: Vol. 48, #3, pp. 331-348 - Scott, C. S., Webb, M. W. & Fox, R. C., 2006: Horolodectes sunae, an enigmatic mammal from the Late Paleocene of Alberta, Canada.
–Journal of Paleontology: Vol. 80, #5, pp. 1009-1025