Variety of Life

Lower jaw of Pediomys, from here.

Belongs within: Boreosphenida.
Contains: Marsupialia.

The Metatheria are a clade of mammals including the total group of modern marsupials and all fossil taxa more closely related to marsupials than to other mammals. The earliest metatherians are known from the early Late Cretaceous of Eurasia and North America though they only survived into the Cenozoic in South America and Australia.

Synapomorphies (from Bi et al. 2014, for Sulestes + Marsupialia): Posterior vertical shelf of masseteric fossa connected to dentary condyle as thin crest along angular margin of mandible; dentary condyle continuous with semicircular posterior margin of dentary, condyle facing up due to up-turning of posterior-most part of dentary; molar with cristid obliqua present, contact closest to lowest point of protocristid; mesiolingual vertical crest of paraconid on lower molars forming a keel; entoconid near height of hypoconid; paraconid, metaconid, and entoconid on lower molars aligned; functional talonid basin of lower molars as long as wide; hypoconid/protoconid height ratio between 40% and 60%; stylar cuspule "C" (near ectoflexus) present on penultimate upper molar; wear facets 5 and 6 differentiated on labial face of entoconid; five upper incisors present; upper anterior-most incisor anteriorly projecting, separated from following teeth by diastema; body of calcaneus mediolaterally compressed at level of posterior calcaneoastragalar facet; calcaneal tubercle ventrally curved; lacrimal foramen on facial side of lacrimal; optic foramen not separated from sphenorbital fissure; incisive foramina intermediate in size.

<==Metatheria [Boreometatheria, Marsupialiformes, Marsupicarnivora] VG09
    |--Sulestes BW14
    `--+--Marsasia LJ03
       `--+--+--Kokopellia VG09
          |  `--Asiatherium KH14
          `--+--Albertatherium BW14
             `--+--Anchistodelphys LJ03
                `--+--Peradectia [Alphadelphia] CGW04
                   |    |  i. s.: Caroloameghiniidae CGW04
                   |    |--Peradectidae GH99
                   |    |    |--Kasserinotherium tunisiense Crochet 1986 GH99
                   |    |    |--Peradectes Matthew & Granger 1921 S35
                   |    |    |    |--*P. elegans Matthew & Granger 1921 S35
                   |    |    |    `--P. minor OB13
                   |    |    `--Alphadon [Alphadontidae] CGW04
                   |    |         |--A. halleyi G03
                   |    |         |--A. marshi Simpson 1927 FN03
                   |    |         `--A. perexiguus G03
                   |    |--+--Pariadens VG09
                   |    |  `--Didelphodon [Stagodontidae] LJ03
                   |    |       `--D. vorax Marsh 1889 FN03
                   |    `--Pediomys [Pediomyidae] LJ03
                   |         |--P. clemensi G03
                   |         |--P. elegans G03
                   |         |--P. hatcheri SBL05
                   |         |--P. krejcii G03
                   |         `--P. prokrejcii G03
                   `--Notometatheria [Ameridelphia, Sparassodonta] RWN98
                        |--+--Mayulestes [Mayulestidae] WR07
                        |  |    `--M. ferox WR07
                        |  `--+--Marsupialia VG09
                        |     `--+--Pucadelphidae CGW04
                        |        |    |--Pucadelphys andinus WR07
                        |        |    |--Incadelphys CGW04
                        |        |    `--Mizquedelphys CGW04
                        |        `--Andinodelphys LJ03
                        `--Borhyaenoidea M77
                             |  i. s.: Hondadelphys M77 [Hondadelphidae CGW04]
                             |--Thylacosmilus Riggs 1933 D07 [Thylacosmilidae M77]
                             |    |--T. ferox D07
                             |    `--T. lentis D07
                             `--+--Proborhyaenidae [Proborhyaeninae] CGW04
                                |    |--Arminiheringia auceta M77, S32
                                |    `--Proborhyaena M77
                                `--+--Hathliacynidae [Hathliacyninae] M77
                                   |    |--Sipalocyon M77
                                   |    |--Patene M77
                                   |    |--Cladosictis Amighino 1887 M77, D07
                                   |    |    |--C. bardus D07
                                   |    |    |--C. centralis D07
                                   |    |    `--C. patagonica D07
                                   |    `--Allqokirus CGW05
                                   `--Borhyaenidae RWN98
                                        |  i. s.: Pharsophorus lacerans S32
                                        |         Procladosictis M77
                                        |--Prothylacynus [Prothylacyninae] M77
                                        `--Borhyaeninae M77
                                             |--Nemolestes M77
                                             |--Angelocabrerus M77
                                             `--Borhyaena Ameghino 1894 D07
                                                  |--B. excavata D07
                                                  |--B. macrodonta D07
                                                  `--B. tuberata S32

Metatheria incertae sedis:
  Kiruwamaq Goin & Candela 2004 AH07
  Arcantiodelphys Vullo, Gheerbrant et al. 2009 VG09
    `--*A. marchandi Vullo, Gheerbrant et al. 2009 VG09
  Dakotadens VG09
  Iqualadelphis RWN98
  Turgidodon LJ03
    |--T. praesagus G03
    |--T. rhaister G03
    `--T. russelli G03
  Iugomortiferum VG09
  Adelodelphys VG09
  Sinbadelphys VG09

*Type species of generic name indicated

REFERENCES

[AH07] Asher, R. J., I. Horovitz, T. Martin & M. R. Sánchez-Villagra. 2007. Neither a rodent nor a platypus: a reexamination of Necrolestes patagoniensis Ameghino. American Museum Novitates 3546: 1–40.

[BW14] Bi, S., Y. Wang, J. Guan, X. Sheng & J. Meng. 2014. Three new Jurassic euharamiyidan species reinforce early divergence of mammals. Nature 514: 579–584.

[CGW04] Case, J. A., F. J. Goin & M. O. Woodbourne. 2004. “South American” marsupials from the Late Cretaceous of North America and the origin of marsupial cohorts. Journal of Mammalian Evolution 11 (3–4): 223–255.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[FN03] Fox, R. C., & B. G. Naylor. 2003. A Late Cretaceous taeniodont (Eutheria, Mammalia) from Alberta, Canada. Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen 229 (3): 393–420.

[GH99] Gheerbrant, E., & J.-L. Hartenberger. 1999. Nouveau mammifère insectivore (?Lipotyphla, ?Erinaceomorpha) de l’Eocène inférieur de Chambi (Tunisie). Paläontologische Zeitschrift 73 (1–2): 143–156.

[G03] Gordon, C. L. 2003. A first look of estimating body size in dentally conservative marsupials. Journal of Mammalian Evolution 10 (1–2): 1–21.

[LJ03] Luo, Z.-X., Q. Ji, J. R. Wible & C.-X. Yuan. 2003. An Early Cretaceous tribosphenic mammal and metatherian evolution. Science 302: 1934–1940.

[M77] Marshall, L. G. 1977. Cladistic analysis of borhyaenoid, dasyuroid, didelphoid, and thylacinid (Marsupialia: Mammalia) affinity. Systematic Zoology 26 (4): 410–425.

[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.

[RWN98] Rougier, G. W., J. R. Wible & M. J. Novacek. 1998. Implications of Deltatheridium specimens for early marsupial history. Nature 396: 459–463.

[S32] Simpson, G. G. 1932. Skulls and brains of some mammals from the Notostylops beds of Patagonia. American Museum Novitates 578: 1–11.

[S35] Simpson, G. G. 1935. The Tiffany fauna, Upper Paleocene. I.—Multituberculata, Marsupialia, Insectivora, and ?Chiroptera. American Museum Novitates 795: 1–19.

[SBL05] Sullivan, R. M., A. C. Boere & S. G Lucas. 2005. Redescription of the ceratopsid dinosaur Torosaurus utahensis (Gilmore, 1946) and a revision of the genus. Journal of Paleontology 79 (3): 564–582.

[VG09] Vullo, R., E. Gheerbrant, C. De Muizon & D. Néraudeau. 2009. The oldest modern therian mammal from Europe and its bearing on stem marsupial paleobiogeography. Proceedings of the National Academy of Sciences of the USA 106 (47): 19910–19915.

[WR07] Wible, J. R., G. W. Rougier, M. J. Novacek & R. J. Asher. 2007. Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary. Nature 447: 1003–1006.

Southern tamandua Tamandua tetradactyla, copyright Bernard Dupont.

Belongs within: Eutheria.
Contains: Dasypodidae, Folivora, Tubulidentata, Afrosoricida, Macroscelididae, Pan-Paenungulata.

The Atlantogenata are a clade of mammals supported by molecular data uniting the clades Xenarthra and Afrotheria. Xenarthra is the order of mammals containing the anteaters, sloths and armadillos, and is primarily found in South America. The name refers to the distinctive vertebrae of members of the clade. In older references this group was commonly referred to the Edentata, meaning 'toothless', but only the anteaters (Myrmecophagidae) are truly toothless; the sloths and armadillos retain enamelless back teeth. The armadillos and their extinct relatives in the Cingulata are characterised by dorsal armour formed from osteoderms covered in keratinous scales, that may be variously fused to form plates. The sloths and anteaters retain a uniform covering of thick hair. Anteaters have elongate snouts and tongues used for feeding on social insects.

The Afrotheria is a clade of mammals united by molecular analyses, though morphological support for the clade remains elusive. Most phylogenies support division of living afrotherians between the insectivorous taxa—Afrosoricida (golden moles and tenrecs), Macroscelididae (elephant shrews) and Orycteropus afer (aardvark)—and the herbivorous Paenungulata. Possible fossil afrotherians include the Ptolemaiida, a group of insectivorous or omnivorous mammals known from the Oligocene and Miocene of Africa. At least one ptolemaiidan, Kelba quadeemae, was probably about the size of modern coyote (Cote et al. 2007).

<==Atlantogenata [Hicanodonta] HUG17
    |--Xenarthra HUG17
    |    |  i. s.: Proeutatus Ameghino 1891 BAA04
    |    |           `--P. lagena A05a
    |    |         Propaopus Ameghino 1881 BAA04
    |    |--Cingulata [Dasypoda, Dasypodoidea] DSD03
    |    |    |--Stegotheriidae A05c
    |    |    |    |--Stegotherium Ameghino 1887 M94, BAA04
    |    |    |    |    |--S. simplex [=Scotaeops simplex] A05c
    |    |    |    |    |--S. tessellatum M94
    |    |    |    |    `--S. variegatum D07
    |    |    |    |--Necrodasypus galliae Filhol 1894 [=Necromanis galliae] A05c
    |    |    |    `--Pseudostegotherium glangeaudi A05c, A05b
    |    |    |--Dasypodidae BAA04
    |    |    `--Peltephilidae [Peltateloidea] BAA04
    |    |         |--Peltephilus Ameghino 1887 BAA04
    |    |         |    `--P. ferox D07
    |    |         `--Epipeltephilus A05c
    |    `--Pilosa DSD03
    |         |--+--Folivora DSD03
    |         |  `--Pseudoglyptodon chilensis MJ11
    |         `--+--Utaetus Ameghino 1902 HUG17, BAA04
    |            |    |--U. argos A05c
    |            |    |--U. buccatus HUG17
    |            |    `--U. deustus A05c
    |            `--Myrmecophagidae [Vermilingua] DSD03
    |                 |  i. s.: Protamandua rothi OB13
    |                 |--Cyclopes Gray 1821 DSD03, C57 (see below for synonymy)
    |                 |    `--*C. didactylus (Linnaeus 1758) (see below for synonymy) C57
    |                 |         |--C. d. didactylus [incl. Myrmecophaga unicolor Geoffroy in Desmarest 1804] C57
    |                 |         |--C. d. catellus Thomas 1928 C57
    |                 |         |--C. d. dorsalis (Gray 1865) [=Cyclothurus dorsalis] C57
    |                 |         |--C. d. eva Thomas 1902 C57
    |                 |         |--C. d. ida Thomas 1900 [incl. C. didactylus codajazensis Lönnberg 1942] C57
    |                 |         |--C. d. juruanus Lönnberg 1942 C57
    |                 |         |--C. d. melini Lönnberg 1928 C57
    |                 |         `--C. d. mexicanus G69
    |                 `--Myrmecophaginae DC01
    |                      |--Myrmecophaga Linnaeus 1758 BAA04 [incl. Falcifer Rehn 1900 C57]
    |                      |    |--*M. tridactyla Linnaeus 1758 [incl. M. jubata Linné 1766, M. iubata] C57
    |                      |    |    |--M. t. tridactyla C57
    |                      |    |    `--M. t. artata Osgood 1912 [=M. t. artatus] C57
    |                      |    `--M. aculeata G96
    |                      `--Tamandua Gray 1825 DSD03, C57 (see below for synonymy)
    |                           |--*T. tetradactyla (Linnaeus 1758) [=Myrmecophaga tetradactyla] C57
    |                           |    |--T. t. tetradactyla (see below for synonymy) C57
    |                           |    |--T. t. chapadensis Allen 1904 (see below for synonymy) C57
    |                           |    |--T. t. chiriquensis Allen 1904 (see below for synonymy) C57
    |                           |    |--T. t. instabilis Allen 1904 C57
    |                           |    |--T. t. punensis Allen 1916 C57
    |                           |    `--T. t. quicha Thomas 1927 C57
    |                           |--T. longicaudata (Wagner 1844) [=Myrmecophaga longicaudata] C57
    |                           |    |--T. l. longicaudata [=T. tetradactyla longicaudata] C57
    |                           |    `--T. l. mexianae Hagmann 1908 C57
    |                           `--T. mexicana WR07 [=T. tetradactyla mexicana G69]
    `--Afrotheria [Colbertia] HUG17
         |  i. s.: Ptolemaiida CW07
         |           |  i. s.: Qarunavus Simons & Gingerich 1974 CW07
         |           |           `--Q. meyeri Simons & Gingerich 1974 CW07
         |           |--Kelba Savage 1965 [Kelbidae] CW07
         |           |    `--K. quadeemae Savage 1965 [incl. Kenyalutra songhorensis, Ndamathaia kubwa] CW07
         |           `--Ptolemaiidae [Ptolemayidae] CW07
         |                |--Ptolemaia Osborn 1908 CW07
         |                |    |--*P. lyonsi Osborn 1908 CW07
         |                |    `--P. grangeri Bown & Simons 1987 CW07
         |                `--Cleopatrodon Bown & Simons 1987 CW07
         |                     |--C. ayeshae Bown & Simons 1987 CW07
         |                     `--C. robusta Bown & Simons 1987 CW07
         |         Ocepeia daouiensis HUG17
         |--Afroinsectiphilia HUG17
         |    |--+--Tubulidentata FS15
         |    |  `--Plesiorycteropus Filhol 1895 ANG03, M94 (see below for synonymy)
         |    |       |--*P. madagascariensis Filhol 1895 (see below for synonymy) M94
         |    |       `--P. germainepetterae MacPhee 1994 M94
         |    `--Afroinsectivora OB13
         |         |--Afrosoricida OB13
         |         `--+--Macroscelididae OB13
         |            |--+--Pseudictops Matthew, Granger & Simpson 1929 AM05, V67 [Pseudictopidae]
         |            |  `--Anagalidae MHL03
         |            |       |--Anagale Simpson 1931 AM05, V67
         |            |       |    `--A. gobiensis MHL03
         |            |       `--Anagalopsis Bohlin 1951 V67
         |            `--+--Dilambdogale gheerbranti HUG17
         |               `--Widanelfarasia HUG17
         |                    |--W. bowni HUG17
         |                    `--W. rasmusseni HUG17
         `--Pan-Paenungulata OB13

Cyclopes Gray 1821 DSD03, C57 [incl. Cyclothurus Lesson 1842 C57, Myrmidon Wagley 1830 C57; Cyclopedidae]

*Cyclopes didactylus (Linnaeus 1758) [=Myrmecophaga didactyla, Cyclothurus didactyla, Myrmidon didactylus] C57

Plesiorycteropus Filhol 1895 ANG03, M94 [incl. Myoryctes Forsyth Major 1908 (preoc.) M94, Majoria Thomas 1915 M94; Bibymalagasia, Plesiorycteropodidae, Plesiorycteropodinae]

*Plesiorycteropus madagascariensis Filhol 1895 [incl. Hypogeomys boulei Grandidier 1912, Myoryctes rapeto Forsyth Major 1908, Majoria rapeto] M94

Tamandua Gray 1825 DSD03, C57 [=Tamanduus Cuvier 1829 C57; incl. Dryorix Gloger 1841 C57, Uroleptes Wagner 1830 C57, Uroleptess Alston 1882 C57]

Tamandua tetradactyla chapadensis Allen 1904 [incl. T. tetradactyla kriegi Krumbiegel 1940, Myrmecophaga bivittata straminea Cope 1889 (n. d.)] C57

Tamandua tetradactyla chiriquensis Allen 1904 [=T. t. chiriquinus (l. c.); incl. T. t. tambensis Lönnberg 1937] C57

*Tamandua tetradactyla tetradactyla (Linnaeus 1758) [incl. Myrmecophaga bivittata Geoffroy in Desmarest 1804, Tamandua bivittata, M. crispa Rüppell 1845, M. myosura Pallas 1776, M. nigra Geoffroy in Desmarest 1804, Tamandua bivittata var. opistholeuca Gray 1873, T. bivittata var. opithomelas Gray 1873, M. tamandua Cuvier 1817, M. ursina Rapp 1852] C57

*Type species of generic name indicated

REFERENCES

[A05a] Ameghino, F. 1905a. Le perforación astragaliana en Priodontes, Canis (Chrysocyon) y Typotherium. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 1–19.

[A05b] Ameghino, F. 1905b. La perforación astragaliana en el Orycteropus y el origen de los Orycteropidae. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 59–95.

[A05c] Ameghino, F. 1905c. Les édentés fossiles de France et d'Allemagne. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 175–250.

[AM05] Asher, R. J., J. Meng, J. R. Wible, M. C. McKenna, G. W. Rougier, D. Dashzeveg & M. J. Novacek. 2005. Stem Lagomorpha and the antiquity of Glires. Science 307: 1091–1094.

[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.

[BAA04] Bergqvist, L. P., E. A. L. Abrantes & L. dos S. Avilla. 2004. The Xenarthra (Mammalia) of São José de Itaboraí Basin (upper Paleocene, Itaboraian), Rio de Janeiro, Brazil. Geodiversitas 26 (2): 323–337.

[C57] Cabrera, A. 1957. Catalogo de los mamiferos de America del Sur. I (Metatheria—Unguiculata—Carnivora). Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Instituto Nacional de Investigacion de Las Ciencias Naturales, Ciencias Zoológicas 4 (1): 1–307.

[CW07] Cote, S., L. Werdelin, E. R. Seiffert & J. C. Barry. 2007. Additional material of the enigmatic Early Miocene mammal Kelba and its relationship to the order Ptolemaiida. Proceedings of the National Academy of Sciences of the USA 104 (13): 5510–5515.

[DC01] Delsuc, F., F. M. Catzeflis, M. J. Stanhope & E. J. P. Douzery. 2001. The evolution of armadillos, anteaters and sloths depicted by nuclear and mitochondrial phylogenies: implications for the status of the enigmatic fossil Eurotamandua. Proceedings of the Royal Society of London Series B—Biological Sciences 268: 1605–1615.

[DSD03] Delsuc, F., M. J. Stanhope & E. J. P. Douzery. 2003. Molecular systematics of armadillos (Xenarthra, Dasypodidae): contribution of maximum likelihood and Bayesian analyses of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 28: 261–275.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[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.

[G96] Geoffroy, É. L. 1796. Extrait d'un mémoire sur le Myrmecophaga capensis, Gme. Bulletin de la Société Philomatique, a ses Correspondans 50: 1–2.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

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

[M94] MacPhee, R. D. E. 1994. Morphology, adaptations, and relationships of Plesiorycteropus, and a diagnosis of a new order of eutherian mammals. Bulletin of the American Museum of Natural History 220: 1–214.

[MJ11] Meredith, R. W., J. E. Janečka, J. Gatesy, O. A. Ryder, C. A. Fisher, E. C. Teeling, A. Goodbla, E. Eizirik, T. L. L. Simão, T. Stadler, D. L. Rabosky, R. L. Honeycutt, J. J. Flynn, C. M. Ingram, C. Steiner, T. L. Williams, T. J. Robinson, A. Burk-Herrick, M. Westerman, N. A. Ayoub, M. S. Springer & W. J. Murphy. 2011. Impacts of the Cretaceous terrestrial revolution and KPg extinction on mammal diversification. Science 334: 521–524.

[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.

[V67] Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135 (5): 217–284.

[WR07] Wible, J. R., G. W. Rougier, M. J. Novacek & R. J. Asher. 2007. Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary. Nature 447: 1003–1006.

Brown-throated sloth Bradypus variegatus, photographed by Stefan Laube.

Belongs within: Atlantogenata.
Contains: Ortotheriinae.

The Folivora, sloths, are a group of herbivorous mammals known from Central and South America. Living members of the group are arboreal though many fossil members were terrestrial and could reach massive sizes.

<==Folivora [Gravigrada, Phyllophaga, Tardigrada] DSD03
    |--Bradypus Linnaeus 1758 C57 (see below for synonymy)
    |    |  i. s.: B. boliviensis Gray 1871 [=B. tridactylus boliviensis] C57
    |    |         B. infuscatus Wagler 1831 C57
    |    |           |--B. i. infuscatus (see below for synonymy) C57
    |    |           |--B. i. brasiliensis Blainville 1840 (see below for synonymy) C57
    |    |           |--B. i. ephippiger Philippi 1870 [incl. B. ignavus Goldman 1913, B. violeta Thomas 1917] C57
    |    |           |--B. i. gorgon Thomas 1926 C57
    |    |           `--B. i. marmoratus (Gray 1849) (see below for synonymy) C57
    |    |--B. torquatus Illiger 1811 FS15, C57 (see below for synonymy)
    |    `--+--*B. tridactylus Linnaeus 1758 C57, FS15, C57 (see below for synonymy)
    |       `--+--B. pygmaeus FS15
    |          `--B. variegatus Schinz 1823 FS15, C57 [=B. (Scaeopus) variegatus C57]
    `--Megalonychidae FS15
         |  i. s.: Eucholoeops ingens S32
         |         Schismotherium S32
         |         Hapaloides S32
         |         Hapalops Ameghino 1887 S32, BAA04
         |           |--H. cadens D07
         |           |--H. elongatus OB13
         |           |--H. indifferens D07
         |           `--H. longicers D07
         |         Proschismotherium S32
         |           |--P. oppositum S32
         |           `--P. scarritti Simpson 1932 S32
         |--+--+--Nothrotheriinae MM95
         |  |  |    |--Pronothrotherium MM95
         |  |  |    `--+--Nothropus MM95
         |  |  |       `--+--+--Nothrotheriops shastense MM95, FS15
         |  |  |          |  `--Nothrotherium maquinense FS15
         |  |  |          `--Thalassocnus Muizon & McDonald 1995 MM95
         |  |  |               |--*T. natans Muizon & McDonald 1995 MM95
         |  |  |               |--T. antiquus D07
         |  |  |               |--T. carolomartini N10
         |  |  |               |--T. littoralis D07
         |  |  |               `--T. yaucensis N10
         |  |  `--+--Eremotherium laurillardi FS15
         |  |     `--Megatherium Cuvier 1796 FS15, H07 [Megatheriidae]
         |  |          |--M. americanum H07
         |  |          |--M. lundii O05
         |  |          `--M. tarijense FS15
         |  `--+--+--Paramylodon harlani FS15
         |     |  `--+--Mylodon darwinii FS15
         |     |     `--+--Lestodon armatus FS15
         |     |        `--Glossotherium Owen 1842 FS15, D07
         |     |             `--G. robustum D07
         |     `--+--Scelidotherium leptocephalum FS15
         |        `--+--Catonyx cuvieri FS15
         |           `--+--Valgipes deformis FS15
         |              `--Scelidodon chiliensis FS15
         `--+--+--Choloepus Illiger 1811 FS15, C57 [=Chaelopus Gray in Griffith 1827 C57, Cholopus Sclater 1872 C57]
            |  |    |--*C. didactylus (Linnaeus 1758) (see below for synonymy) C57
            |  |    `--C. hoffmanni Peters 1858 C57
            |  |         |--C. h. hoffmanni [incl. C. didactylus var. columbianus Gray 1871] C57
            |  |         |--C. h. agustinus Allen 1913 C57
            |  |         |--C. h. capitalis Allen 1913 [incl. C. andinus Allen 1913] C57
            |  |         |--C. h. florenciae Allen 1913 [incl. C. napensis Lönnberg 1922] C57
            |  |         |--C. h. juruanus Lönnberg 1942 C57
            |  |         `--C. h. pallescens Lönnberg 1928 [incl. C. h. peruvianus Menegaux 1906 (n. d.)] C57
            |  `--Ortotheriinae FS15
            `--+--Megalonyx Jefferson 1799 FS15, H07 [Megalonychidae]
               |    `--M. jeffersoni H07
               `--+--+--Diabolotherium nordenskioldi FS15
                  |  `--Acratocnus Anthony 1916 FS15, AC98
                  |       |--*A. odontrigonus Anthony 1916 AC98
                  |       `--A. ye FS15
                  `--Megalocninae PC56
                       |--Megalocnus Leidy 1868 [=Megalochnus Ameghino 1881; incl. Myomorphus Pomel 1868] PC56
                       |    |--*M. rodens Leidy 1868 PC56 (see below for synonymy)
                       |    `--M. zile FS15
                       |--Neomesocnus Arredondo 1961 AC98
                       |    `--*N. brevirrostris Arredondo 1961 AC98
                       |--Parocnus Miller 1929 AC98
                       |    |--*P. serus Miller 1929 AC98
                       |    `--P. browni FS15
                       `--Mesocnus Matthew 1931 AC98
                            |--*M. browni Matthew 1931 AC98
                            `--M. torrei Matthew 1931 AC98

Bradypus Linnaeus 1758 C57 [incl. Acheus Cuvier 1825 C57, Arctopithecus Gray 1849 non Virey 1819 C57, Eubradypus Lönnberg 1942 C57, Hemibradypus Anthony 1906 C57, Ignavus Blumenbach 1779 C57, Neobradypus Lönnberg 1942 C57, Scaeopus Peters 1864 C57; Bradypodidae DSD03]

Bradypus infuscatus brasiliensis Blainville 1840 [=B. braziliensis, B. tridactylus brasiliensis; incl. Arctopithecus blainvilli Gray 1849, B. dorsalis Fitzinger 1871, B. miritibae Lönnberg 1942, B. pallidus Wagner 1844] C57

Bradypus infuscatus infuscatus Wagler 1831 [incl. B. infuscatus var. brachydactylus Wagner 1844, B. marmoratus codajazensis Lönnberg 1942, B. macrodon Thomas 1917, B. speculiger Fitzinger 1871, B. trivittatus Cornalia in Osculati 1850] C57

Bradypus infuscatus marmoratus (Gray 1849) [=Arctopithecus marmoratus; incl. A. problematicus Gray 1849, A. flaccidus var. smithii Gray 1869, B. (Eubradypus) tocantinus Lönnberg 1942, B. unicolor Fitzinger 1871] C57

Bradypus torquatus Illiger 1811 FS15, C57 [=B. (*Scaeopus) torquatus C57; incl. B. affinis Gray 1849 C57, B. crinitus Gray 1849 C57, Hemibradypus mareyi Anthony 1906 C57]

*Bradypus tridactylus Linnaeus 1758 C57, FS15, C57 [=Arctopithecus tridactylus C57; incl. Acheus ai Lesson 1827 C57, B. ai C57, B. cristatus Temminck in Fitzinger 1871 C57, B. cuculliger Wagler 1831 C57, B. tridactylus guianensis Blainville 1940 C57, B. gularis Rüppell 1845 C57, Arctopithecus gularis C57]

*Choloepus didactylus (Linnaeus 1758) [=Bradypus didactylus, Cholopus didactylus; incl. Choloepus brasiliensis Fitzinger 1871, B. curi Link 1795, Choloepus guianensis Fitzinger 1871, B. unau Link 1795] C57

*Megalocnus rodens Leidy 1868 PC56 [incl. M. casimbae Matthew 1959 AC98, M. intermedius Mayo 1969 AC98, M. junius Matthew 1959 AC98, M. ursulus Matthew 1959 AC98]

*Type species of generic name indicated

REFERENCES

[AC98] Alcover, J. A., X. Campillo, M. Macias & A. Sans. 1998. Mammal species of the world: additional data on insular mammals. American Museum Novitates 3248: 1–29.

[BAA04] Bergqvist, L. P., E. A. L. Abrantes & L. dos S. Avilla. 2004. The Xenarthra (Mammalia) of São José de Itaboraí Basin (upper Paleocene, Itaboraian), Rio de Janeiro, Brazil. Geodiversitas 26 (2): 323–337.

[C57] Cabrera, A. 1957. Catalogo de los mamiferos de America del Sur. I (Metatheria—Unguiculata—Carnivora). Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Instituto Nacional de Investigacion de Las Ciencias Naturales, Ciencias Zoológicas 4 (1): 1–307.

[DSD03] Delsuc, F., M. J. Stanhope & E. J. P. Douzery. 2003. Molecular systematics of armadillos (Xenarthra, Dasypodidae): contribution of maximum likelihood and Bayesian analyses of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 28: 261–275.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[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.

[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.

[MM95] Muizon, C. de, & H. G. McDonald. 1995. An aquatic sloth from the Pliocene of Peru. Nature 375: 224–227.

[N10] Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: Bideford (UK).

[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.

[O05] Outes, F. F. 1905. Sobre un instrumento paleolítico de Luján (Provincia de Buenos Aires). Anales del Museo Nacional de Buenos Aires, serie 3, 6: 169–173.

[PC56] Paula Couto, C de. 1956. On two mounted skeletons of Megalocnus rodens. Journal of Mammalogy 37 (3): 423–427.

[S32] Simpson, G. G. 1932. Some new or little-known mammals from the Colpodon beds of Patagonia. American Museum Novitates 575: 1–12.

Bushveld elephant shrew Elephantulus intufi, copyright Yathin.

Belongs within: Atlantogenata.

The Macroscelididae, elephant shrews, are a group of small, cursorial insectivorous mammals known from Africa. Members of this group have long, mobile snouts that provide the inspiration for their vernacular name.

<==Macroscelididae [Dipogales, Macroscelidea, Macroscelidoidea]
    |  i. s.: Chambius kasserinensis Hartenberger 1986 GH99
    |         Miorhynchocyon OB13
    |--Mylomygale Broom 1948 [Mylomygalinae] V67
    |    `--M. spiersi Broom 1948 B78
    |--Myohyracinae [Myohyracidae] M78
    |    |--Protypotheroides Stromer 1922 M78
    |    |    `--P. beetzi Stromer 1922 B78
    |    `--Myohyrax Andrews 1914 M78
    |         |--*M. oswaldi Andrews 1914 M78 [incl. M. doederleini Stromer 1926 B78]
    |         `--M. osborni D07
    |--Rhynchocyon Peters 1847 V67 [incl. Rhinonax Thomas 1918 V67; Rhynchocyoninae B78, Rhynchocyonini]
    |    |  i. s.: R. clarki Butler & Hopwood 1957 B78
    |    |         R. rusingae Butler 1969 B78
    |    |         R. stuhlmanni JP84
    |    |--+--R. chrysopygus FS15
    |    |  `--R. udzungwensis FS15
    |    `--+--R. cirnei FS15
    |       `--R. petersi FS15
    `--Macroscelidinae [Macroscelidini] B78
         |  i. s.: Palaeothentoides Stromer 1932 V67
         |           `--P. africanus Stromer 1932 B78
         |         Metolbodotes Schlosser 1910 [=Metoldobotes] V67
         |           `--M. stromeri Schlosser 1910 B78
         |         Nasilio Thomas & Schwann 1906 V67
         |--+--Macroscelides Smith 1829 FS15, V67 (see below for synonymy)
         |  |    `--M. proboscideus ANG03
         |  |         |--M. p. proboscideus B78
         |  |         `--M. p. vagans Butler & Greenwood 1976 B78
         |  `--+--‘Elephantulus’ rozeti FS15
         |     `--Petrodromus Peters 1846 FS15, V67 [incl. Cercoctenus Hollister 1916 V67, Mesoctenus Thomas 1918 V67]
         |          |--P. sultan GM71
         |          `--P. tetradactylus IT07
         `--Elephantulus Thomas & Schwann 1906 FS15, V67 [incl. Elephantomys Broom 1937]
              |  i. s.: E. antiquus Broom 1948 B78
              |         E. broomi [incl. Elephantomys langi Broom 1937] B78
              |         E. vandami B78
              |--+--E. revoili FS15
              |  `--+--E. fuscipes FS15 [=Nasilio fuscipes B78]
              |     `--E. fuscus FS15 [=Nasilio fuscus B78]
              |          |--E. f. fuscus B78
              |          `--E. f. leakeyi (Butler & Greenwood 1965) B78
              `--+--+--E. myurus FS15
                 |  `--+--E. edwardii FS15
                 |     `--E. pilicaudus FS15
                 `--+--+--*E. rupestris B78, FS15
                    |  `--E. intufi FS15 [=Elephantomys intufi B78]
                    `--+--E. brachyrhynchus FS15 [=Nasilio brachyrhynchus B78]
                       `--E. rufescens FS15

Macroscelides Smith 1829 FS15, V67 [incl. Diposorex Blainville 1838 V67, Eumerus Geoffroy Saint-Hailaire 1829 V67, Macroscelis Fischer 1830 V67, Rhinomys Lichtenstein 1831 V67]

*Type species of generic name indicated

REFERENCES

[AM02] Asher, R. J., M. C. McKenna, R. J. Emry, A. R. Tabrum & D. G. Kron. 2002. Morphology and relationships of Apternodus and other extinct, zalambdodont, placental mammals. Bulletin of the American Museum of Natural History 273: 1–117.

[B78] Butler, P. M. 1978. Insectivora and Chiroptera. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 56–68. Harvard University Press: Cambridge (Massachusetts).

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[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.

[GH99] Gheerbrant, E. & J.-L. Hartenberger. 1999. Nouveau mammifère insectivore (?Lipotyphla, ?Erinaceomorpha) de l’Eocène inférieur de Chambi (Tunisie). Paläontologische Zeitschrift 73 (1–2): 143–156.

[GM71] Goodman, M., & G. W. Moore. 1971. Immunodiffusion systematics of the primates I. The Catarrhini. Systematic Zoology 20 (1): 19–62.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[JP84] Jolicoeur, P., P. Pirlot, G. Baron & H. Stephan. 1984. Brain structure and correlation patterns in Insectivora, Chiroptera, and Primates. Systematic Zoology 33 (1): 14–29.

[M78] Meyer, G. E. 1978. Hyracoidea. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 284–314. Harvard University Press: Cambridge (Massachusetts).

[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.

[V67] Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135 (5): 217–284.

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

REFERENCES

[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.

Reconstructed skull of Uchkudukodon nessovi, from McKenna et al. (2000).

Belongs within: Eutheria.

The Asioryctitheria are a group of small, probably insectivorous mammals known from the Late Cretaceous of Asia. Monophyly of this group is uncertain: Wible et al.'s (2007) phylogenetic analysis found them to be paraphyletic to modern placentals whereas Halliday et al. (2017) found a clade uniting Asioryctitheria with the contemporary Zalambdalestidae.

<==Asioryctitheria WR07
    |--+--+--Bulaklestes kezbe HUG17, WR07
    |  |  `--Uchkudukodon nessovi HUG17, OB13
    |  `--Daulestes Trofimov & Nessov in Nessov & Trofimov 1979 HUG17, FN03
    |       |--D. inobservabilis WR07
    |       |--D. kulbeckensis WR07
    |       `--D. nessovi AAE01
    `--+--+--Kennalestes Kielan-Jaworowska 1969 HUG17, FN03
       |  |    `--K. gobiensis WR07
       |  `--+--Ukhaatherium Novacek 1997 HUG17, D07
       |     |    `--U. nessovi OB13
       |     `--Asioryctes Kielan-Jaworowska 1975 HUG17, FN03
       |          `--A. nemegtensis WR07
       `--Zalambdalestidae WR07
            |--Kulbeckia HUG17
            |    |--K. kansaica HUG17
            |    `--K. kulbecke WR07
            `--+--Lainodon orueetxebarriai HUG17, WR07
               |--Zhangolestes jilinensis HUG17
               |--Barunlestes butleri HUG17, WR07
               `--+--Alymlestes kielanae HUG17
                  `--Zalambdalestes Gregory & Simpson 1926 HUG17, D07
                       |--Z. grangeri HUG17
                       `--Z. lechei WR07

*Type species of generic name indicated

REFERENCES

[AAE01] Archibald, J. D., A. O. Averianov & E. G. Ekdale. 2001. Late Cretaceous relatives of rabbits, rodents, and other extant eutherian mammals. Nature 414: 62–65.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[FN03] Fox, R. C., & B. G. Naylor. 2003. A Late Cretaceous taeniodont (Eutheria, Mammalia) from Alberta, Canada. Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen 229 (3): 393–420.

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

[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.

[WR07] Wible, J. R., G. W. Rougier, M. J. Novacek & R. J. Asher. 2007. Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary. Nature 447: 1003–1006.

Tiger quolls Dasyurus maculatus, copyright J.-L. Klein & M.-L. Hubert.

Belongs within: Dasyuridae.

The Dasyurinae is a group of carnivorous marsupials including the quolls (marsupial cats), mulgaras and related taxa. Members of this group include the largest living carnivorous marsupials.

Dasyurinae
    |--+--Dasykaluta rosamondae CB-E04
    |  `--Parantechinus Tate 1947 CB-E04, R64
    |       `--*P. apicalis [=Phascogale apicalis] R64
    |--+--‘Pseudantechinus’ ningbing CB-E04
    |  `--Myoictis Gray 1858 CB-E04, LA02
    |       `--*M. melas (Müller 1840) LA02
    `--+--Pseudantechinus Tate 1947 CB-E04, R64
       |    |--*P. macdonnellensis [=Phascogale macdonnellensis] R64
       |    |--P. roryi CWS05
       |    `--P. woolleyae CB-E04
       |--Dasycercus Peters 1875 CB-E04, LA02
       |    |--*D. cristicauda (Krefft 1867) LA02
       |    |--D. blythi TL70
       |    |--D. byrnei CB-E04
       |    |--D. hillieri LA02
       |    `--D. worboysi Dawson, Muirhead & Wroe 1999 LA02
       `--+--‘Antechinus’ bilarni Johnson 1954 R64 [=Parantechinus bilarni CB-E04]
          `--+--+--Neophascogale lorentzi CB-E04
             |  `--Phascolosorex CB-E04
             |       |--P. doriae CB-E04
             |       `--P. dorsalis CB-E04
             `--+--Sarcophilus Cuvier 1837 CB-E04, LA02 [Sarcophilini]
                |    |--*S. laniarius (Owen 1838) LA02
                |    |--S. harrisii (Boitard 1841) K92 [incl. S. ursinus S61]
                |    `--S. moornaensis Crabb 1982 LA02
                `--Dasyurus Geoffroy 1796 CB-E04, LA02
                     |  i. s.: D. bowlingi A14
                     |--D. hallucatus CB-E04 [=Satanellus hallucatus TL70]
                     `--+--+--*D. maculatus (Kerr 1792) LA02, CB-E04, LA02 [=Viverra maculata G91]
                        |  `--D. dunmalli Bartholomai 1971 LA02
                        `--+--D. viverrinus (Shaw 1800) CB-E04, R64 (see below for synonymy)
                           `--+--D. albopunctatus CB-E04
                              `--+--D. geoffroii Gould 1841 CB-E04, BD-D09
                                 `--D. spartacus CB-E04

Dasyurinae incertae sedis:
  Dasyurops D02
  Dasyurinus D02
  Archerium Wroe & Mackness 2000 LA02
    `--*A. chinchillaensis Wroe & Mackness 2000 LA02
  Dasyuroides Spencer 1896 LA02
    |--*D. byrnei Spencer 1896 LA02
    `--D. achilpatna Archer 1982 LA02
  Ganbulanyi Wroe 1998 LA02
    `--*G. djadjinguli Wroe 1998 LA02
  Glaucodon Stirton 1957 LA02
    `--*G. ballaratensis Stirton 1957 LA02

Dasyurus viverrinus (Shaw 1800) CB-E04, R64 [=Didelphis viverrina R64; incl. Mustela quoll Zimmermann 1777 (nom. inv.) R64, Dasyurus quoll R64]

*Type species of generic name indicated

REFERENCES

[A14] Anderson, W. 1914. Notes on the occurrence of the sand-rock containing bones of extinct species of marsupials (emu, kangaroo, wombat, etc.) on King Island, Bass Strait, Tasmania. Records of the Australian Museum 10 (9): 275-283.

[BD-D09] Beveridge, I. & M.-C. Durette-Desset. 2009. A new species of the nematode genus Copemania (Nematoda: Trichostrongylida), parasitic in the western quoll, Dasyurus geoffroii and short-nosed bandicoot, Isoodon obesulus from south-western Western Australia. Records of the Western Australian Museum 25 (3): 345-349.

[CB-E04] Cardillo, M., O. R. P. Bininda-Emonds, E. Boakes & A. Purvis. 2004. A species-level phylogenetic supertree of marsupials. Journal of Zoology 264: 11-31.

[CWS05] Cooper, N. K., P. C. Withers & T. Stewart. 2005. Description of sternal glands in Western Australian Sminthopsis and Ningaui (Marsupialia: Dasyuridae). Journal of the Royal Society of Western Australia 88 (2): 45-49.

[D02] Dyck, S. van. 2002. Morphology-based revision of Murexia and Antechinus (Marsupialia: Dasyuridae). Memoirs of the Queensland Museum 48: 239-330.

[G91] Groves, C. P. 1991. A Theory of Human and Primate Evolution, revised ed. Clarendon Press: Oxford.

[K92] Klompen, J. S. H. 1992. Phylogenetic relationships in the mite family Sarcoptidae (Acari: Astigmata). Miscellaneous Publications, Museum of Zoology, University of Michigan 180: i-vi, 1-154.

[LA02] Long, J., M. Archer, T. Flannery & S. Hand. 2002. Prehistoric Mammals of Australia and New Guinea: One Hundred Million Years of Evolution. University of New South Wales Press: Sydney.

[R64] Ride, W. D. L. 1964. A list of mammals described from Australia between the years 1933 and 1963 (comprising newly proposed names and additions to the Australian faunal list). Australian Mammal Society Bulletin 7 (Suppl.): 1-15.

[S61] Sharman, G. B. 1961. The mitotic chromosomes of marsupials and their bearing on taxonomy and phylogeny. Australian Journal of Zoology 9 (1): 38-60.

[TL70] Turnbull, W. D. & E. L. Lundelius, Jr. 1970. The Hamilton fauna: a Late Pliocene mammalian fauna from the Grange Burn, Victoria, Australia. Fieldiana Geology 19: 1-163.