Atlantogenata

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.

Folivora

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.

Macroscelididae

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.

Pan-Paenungulata

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.

Dasypodidae

Nine-banded armadillos Dasypus novemcinctus, photographed by Charlie Bittner.

Belongs within: Atlantogenata.
Contains: Euphractinae.

The Dasypodidae are the armadillos, a family of omnivorous mammals found in South America and southern North America distinguished by modification of the skin to form an armour of bone and keratin. In the Chlamyphorinae, fairy armadillos, the armour is mostly dorsal and the sides of the body are largely exposed; in other armadillos the armour is more extensive.

Characters (from Nowak 1999): Skull dorsoventrally flattened; lower jaw elongate; tongue long and protrusible; teeth small, ever-growing, peglike, numerous (up to forty in each jaw). Double-layered covering of bone and horn over most of upper surface and sides: shield on top of head; quadrate or polygonal scutes on back and sides, usually arranged into shoulder and hip shields divided by movable rings; irregular horny shields on outer or exposed margins of limbs, which can be withdrawn into space between body and shoulder and hip shields; tail usually encased by bony rings or plates; undersurface of body, inner surfaces of limbs and spaces between scutes with soft, haired skin. Lower bones of legs united proximally and distally. Hands with three to five digits with powerful, curved claws; feet with five digits with claws. Mammae usually located in chest region, abdominal pair occasionally present.

<==Dasypodidae
    |  i. s.: Astegotheriini BAA04
    |           |--Riostegotherium yanei Oliveira & Bergqvist 1998 BAA04
    |           |--Astegotherium Ameghino 1902 BAA04
    |           |--Stegosimpsonia Vizcaíno 1994 BAA04
    |           |--Nanoastegotherium Carlini, Vizcaíno & Scillato-Yané 1997 BAA04
    |           `--Prostegotherium Ameghino 1902 BAA04
    |                `--P. astrifer Ameghino 1902 BAA04
    |--Chlamyphorinae [Chlamyphorini] C57
    |    |--Chlamyphorus Harlan 1825 BR05 (see below for synonymy)
    |    |    `--*C. truncatus Harlan 1825 (see below for synonymy) C57
    |    `--Burmeisteria Gray 1865 C57
    |         `--*B. retusa (Burmeister 1863) (see below for synonymy) C57
    |              |--B. r. retusa C57
    |              `--B. r. clorindae Yepes 1939 C57
    `--+--Dasypus Linnaeus 1758 DSD03, BAA04 (see below for synonymy)
       |    |--*D. novemcinctus Linné 1758 [=Cachicamus novemcinctus, Praopus novemcinctus, Tatusia novemcincta] C57
       |    |    |--D. n. novemcinctus (see below for synonymy) C57
       |    |    |--D. n. aequatorialis Lönnberg 1913 [incl. Tatusia granadiana Gray 1873 (n. d.)] C57
       |    |    |--D. n. mexianae (Hagmann 1908) [=Tatusia novemcincta mexianae] C57
       |    |    `--D. n. mexicanus Peters 1864 MB86
       |    |--D. hybridus (Desmarest 1804) (see below for synonymy) C57
       |    |--D. kappleri Krauss 1862 [=Tatusia kappleri] C57
       |    |    |--D. k. kappleri [incl. D. pentadactylus Peters 1865] C57
       |    |    `--D. k. pastasae (Thomas 1901) (see below for synonymy) C57
       |    |--D. mazzai Yepes 1933 C57
       |    |--D. pilosus (Fitzinger 1856) (see below for synonymy) C57
       |    |--D. sabanicola DSD03
       |    `--D. septemcinctus Linné 1758 (see below for synonymy) C57
       `--+--Tolypeutinae DSD03
          |    |--Tolypeutes Illiger 1811 DSD03, BAA04 (see below for synonymy)
          |    |    |--*T. tricinctus (Linné 1758) (see below for synonymy) C57
          |    |    `--T. matacus (Desmarest 1804) (see below for synonymy) C57
          |    `--Priodontini [Priodontina] DSD03
          |         |--Priodontes Cuvier 1825 (see below for synonymy) C57
          |         |    |--P. giganteus (Geoffroy 1803) (see below for synonymy) C57
          |         |    `--P. maximus Cuvier 1827 BAA04
          |         `--Cabassous McMurtrie 1831 DSD03, BAA04 (see below for synonymy)
          |              |--*C. unicinctus (Linné 1758) (see below for synonymy) C57
          |              |--C. centralis BP87
          |              |--C. chacoensis IT07
          |              |--C. hispidus (Burmeister 1854) (see below for synonymy) C57
          |              |--C. loricatus (Wagner 1855) (see below for synonymy) C57
          |              `--C. tatouay (Desmarest 1819) (see below for synonymy) C57
          `--Euphractinae [Euphractini] DSD03
               |--Zaedyus Ameghino 1889 DSD03, C57 [=Zaedipus (l. c.) C57]
               |    |--Z. pichiy (Desmarest 1804) [=Loricatus pichiy] C57
               |    |    |--Z. p. pichiy (see below for synonymy) C57
               |    |    `--Z. p. caurinus Thomas 1928 C57
               |    `--Z. ciliatus (Fischer 1814) M94, C57 [=Dasypus ciliatus C57]
               `--+--Chaetophractus Fitzinger 1871 DSD03, C57 [incl. Dasyphractus Fitzinger 1871 C57]
                  |    |--*C. villosus (Desmarest 1804) (see below for synonymy) C57
                  |    |--C. nationi (Thomas 1894) [=Dasypus nationi] C57
                  |    `--C. vellerosus (Gray 1865) [=Dasypus vellerosus] C57
                  |         |--C. v vellerosus [incl. Dasyphractus brevirostris Fitzinger 1871] C57
                  |         `--C. v. pannosus (Thomas 1902) [=Dasypus vellerosus pannosus] C57
                  `--Euphractus Wagler 1830 DSD03, C57 (see below for synonymy)
                       `--*E. sexcinctus (Linné 1758) C57 [=Dasypus sexcinctus C57, Chaetophractus sexcinctus ANG03]
                            |--E. s. sexcinctus C57
                            |--E. s. boliviae (Thomas 1907) [=Dasypus sexcinctus boliviae] C57
                            |--E. s. flavimanus (Desmarest 1804) (see below for synonymy) C57
                            |--E. s. setosus (Wied 1826) (see below for synonymy) C57
                            `--E. s. tucumanus (Thomas 1907) (see below for synonymy) C57

*Burmeisteria retusa (Burmeister 1863) [=Chlamydophorus retusus, Calyptophractus retusus, Chlamyphorus retusus] C57

Cabassous McMurtrie 1831 DSD03, BAA04 [incl. Arizostus Gloger 1841 C57, Lysiurus Ameghino 1891 C57, Tatoua Gray 1865 C57, Ziphila Gray 1873 C57]

Cabassous hispidus (Burmeister 1854) [=Dasypus hispidus, Tatoua (Tatoua) hispida; incl. Xenurus latirostris Gray 1873] C57

Cabassous loricatus (Wagner 1855) [=Dasypus gymnurus var. loricatus, Lysiurus loricatus, Xenurus loricatus] C57

Cabassous tatouay (Desmarest 1819) [=Dasypus tatouay, Tatusia tatouay, Xenurus tatouay; incl. Ziphila lugubris Gray 1873] C57

*Cabassous unicinctus (Linné 1758) [=Dasypus unicinctus, Lysiurus unicinctus, Tatoua unicincta, Xenurus (Tatoua) unicinctus; incl. D. dudodecimcinctus Schreber 1774, D. gymnurus Wied 1826, Arizostus gymnurus, Tatusia gymnura, Xenurus gymnurus, Dasypus multicinctus Thumberg 1818, X. nudicaudus Turner 1851, D. octodecimcinctus Erxleben 1777, D. undecimcinctus Illiger 1815 (n. n.), D. verrucosus Wagner 1844, D. xenurus Krauss 1862] C57

*Chaetophractus villosus (Desmarest 1804) [=Loricatus villosus, Dasypus villosus, Euphractus villosus, Tatusia villosa] C57

Chlamyphorus Harlan 1825 BR05 [=Chlamydephorus Lenz 1831 (nom. inv.) non Binney 1879 BR05, Chlamydophorus Wagler 1830 C57]

*Chlamyphorus truncatus Harlan 1825 [=Chlamydophorus truncatus; incl. Chlamydophorus truncatus minor Lahille 1895, Chlamyphorus truncatus minor, Chlamydophorus truncatus ornatus Lahille 1895, Chlamyphorus truncatus patquiensis Yepes 1932, Chlamydophorus truncatus typicus Lahille 1895] C57

Dasypus Linnaeus 1758 DSD03, BAA04 [incl. Cachicama Geoffroy 1835 C57, Cachicamus McMurtrie 1831 C57, Cataphractus Storr 1780 C57, Cryptophractus Fitzinger 1856 C57, Hyperoambon Peters 1864 C57, Loricatus Desmarest 1804 C57, Muletia Gray 1874 C57, Praopus Burmeister 1854 C57, Tatu Blumenbach 1799 C57, Tatus Fermin 1769 C57, Tatusia Lesson 1827 C57, Zonoplites Gloger 1841 C57; Dasypodinae, Dasypodini]

Dasypus hybridus (Desmarest 1804) [=Loricatus hybridus, Cachicama hybridus, Praopus hybridus, Tatu hybridus, Tatusia hybrida, T. hybridus; incl. Dasypus brevicauda Larrañaga 1923] C57

Dasypus kappleri pastasae (Thomas 1901) [=Tatu pastasae; incl. D. kappleri beniensis Lönnberg 1942, D. kappleri peruvianus Lönnberg 1928] C57

*Dasypus novemcinctus novemcinctus Linné 1758 [incl. Tatusia boliviensis Gray 1873, Tatusia brevirostris Gray 1873, Tatusia leptocephala Gray 1873, Tatusia leptorhynchus Gray 1873, Tatusia leptorhinus (l. c.), Dasypus longicaudatus Kerr 1792, D. longicaudus, Praopus longicaudatus, Tatus longicaudatus, Dasypus lundii Fitzinger 1871, Tatus minor Fermin 1769, Loricatus niger Desmarest 1804, Dasypus octocinctus Schreber 1774, D. peba Desmarest 1819, D. (Praopus) peba, Tatusia peba, Tatusia platycercus Hensel 1873, Dasypus serratus Fischer 1814, D. uroceras Lund 1841] C57

Dasypus pilosus (Fitzinger 1856) [=Cryptophractus pilosus, D. (Chryptophractus) pilosus, Tatu pilosa; incl. D. (Praopus) hirsutus Burmeister 1862] C57

Dasypus septemcinctus Linné 1758 [incl. Tatusia megalolepis Cope 1889, D. megalolepe, Tatu megalolepis, Tatusia (Muletia) propalatum Rhoads 1895, Dasypus propalatus] C57

Euphractus Wagler 1830 DSD03, C57 [incl. Encoubertus McMurtrie 1831 C57, Pseudotroctes Gloger 1841 C57, Scleropleura Milne-Edwards 1872 C57]

Euphractus sexcinctus flavimanus (Desmarest 1804) [=Loricatus flavimanus; incl. Dasypus encoubert Desmarest 1820, D. flavipes Fischer 1814, D. sexcinctus gilvipes Lichtenstein 1818, Euphractus sexcinctus gilvipes, D. poyu Larrañaga 1923] C57

Euphractus sexcinctus setosus (Wied 1826) [=Dasypus sexcinctus setosus; incl. Scleropleura bruneti Milne-Edwards 1872, D. mustelinus Fitzinger 1871] C57

Euphractus sexcinctus tucumanus (Thomas 1907) [=Dasypus sexcinctus tucumanus, E. s. tucamanus (l. c.)] C57

Priodontes Cuvier 1825 [=Priodon McMurtrie 1831, Priodonta Gray 1843, Prionodon Gray 1843, Prionodontes Schinz 1845, Prionodos Gray 1865; incl. Cheloniscus Wagler 1830, Polygomphius Gleger 1841] C57

Priodontes giganteus (Geoffroy 1803) [=Dasypus giganteus, Loricatus giganteus, Priodon giganteus; incl. Dasypus gigas Cuvier 1822, Cheloniscus gigas, Priodon gigas, Priodonta gigas, *Priodontes gigas, Prionodontes gigas, Prionodos gigas] C57

Tolypeutes Illiger 1811 DSD03, BAA04 [=Tolypuetes (l. c.) C57; incl. Apar Geoffroy 1847 C57, Apara McMurtrie 1831 C57, Matacus Rafinesque 1815 C57, Sphaerocormus Fitzinger 1871 C57, Tolypoides Neveu-Lemaire & Grandidier 1905 C57; Tolypeutini]

Tolypeutes matacus (Desmarest 1804) [=Loricatus matacus, Tolypeutes mataca, Tolypeutes matacos, Tolypeutes tricinctus matacus, Tolypuetes tricinctus matacos; incl. Dasypus aparoides Bravard in Gervais 1869 (n. n.), Tolypoides bicinctus Grandidier & Neveu-Lemaire 1905, Tolypeutes conurus Geoffroy 1847, Dasypus (Tolypeutes) conurus fossilis Burmeister 1867, Tolypeutes globulus Illiger 1815 (n. n.), Tolypeutes tricinctus muriei Garrod 1878, Tolypuetes tricinctus muriei] C57

*Tolypeutes tricinctus (Linné 1758) [=Dasypus tricinctus, Tolypuetes tricinctus; incl. D. apar Desmarest 1819, Tatusia apar, D. quadricinctus Linné 1758] C57

Zaedyus pichiy pichiy (Desmarest 1804) [incl. Euphractus marginatus Wagler 1830, Dasypus minutus Desmarest 1822, Chaetophractus minutus, Euphractus minutus, Tatusia minuta, *Zaedyus minutus, Dasypus patagonicus Desmarest 1819] C57

*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.

[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.

[BR05] Bouchet, P., J.-P. Rocroi, J. Frýda, B. Hausdorf, W. Ponder, Á. Valdés & A. Warén. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1-2): 1-397.

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[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.

[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.

[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.

[MB86] Matson, J. O. & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1-88.

Nowak, R. M. 1999. Walker's Mammals of the World vol. 2. JHU Press.

Last updated: 21 August 2018.

Proboscidea

Reconstruction of Mammut americanum, copyright James St. John.

Belongs within: Pan-Paenungulata.
Contains: Deinotheriidae, Elephantida.

The Proboscidea include the elephants and their fossil relatives. The earliest proboscideans are known from the early Eocene, and were superficially tapir-like animals. Members of the basal Barytherioidea (including Moeritherium) had bilophodont molars; the molars became trilophodont in the Deinotheriidae and Elephantiformes (Sanders et al. 2004). Modification of the upper second incisors into large tusks is characteristic of the Elephantiformes; basal forms have similarly tusk-like lower first incisors (Tassy & Shoshani 1988).

Characters (from Tassy & Shoshani 1988): Orbitotemporal fossa without processus ascendens of palatine; jugal reduced anteriorly, orbit opening in maxilla; jugal lengthened posteriorly to rear border of glenoid fossa of squamosal; condyloid foramen absent; optic foramen and fissura sphenorbitale on same line under crista orbitotemporalis of orbitotemporal fossa. Metacone on P⁴; upper molars with paracone and metacone transversely thickened and subdivided, paraconule touching precingulum, protocone with postprotocone crest; M^1-2 with distocrista; lower molars with metaconid and entoconid transversely thickened, hypolophid as high as protolophid, hypoconulid developed labially, postentoconulid outlined. Astragalus with enlarged lateral calcaneal facet and protruding tuberculum mediale.

<==Proboscidea
    |  i. s.: Arcanotherium savagei JB10
    |--Barytherioidea [Barypoda] SKR04
    |    |  i. s.: Daouitherium SKR04
    |    |         Barytherium Andrews 1901 [Barytheriidae] H78
    |    |           `--*B. grave Andrews 1901 H78 [=Bradytherium grave DRM86]
    |    |--Phosphatherium Gheerbrant, Sudre & Tassy 1996 RH14, D07
    |    |    `--P. escuilliei D07
    |    `--+--Numidotherium [Numidotheriidae] RH14
    |       |    `--N. koholense TS88
    |       `--Moeritherium Andrews 1901 RH14, CB78 [Moeritheriidae, Moeritherioidea]
    |            |--M. chehbeurameuri JB10
    |            |--M. lyonsi Andrews 1901 [incl. M. ancestrale Petronievics 1923, M. gracile Andrews 1902] CB78
    |            `--M. trigodon Andrews 1904 [incl. M. andrewsi Schlosser 1911] CB78
    `--+--Deinotheriidae TS88
       `--Elephantiformes [Palaeomastodontidae] SW06
            |--Palaeomastodon Andrews 1901 SW06, CM78
            |    `--*P. beadnelli Andrews 1901 (see below for synonymy) CM78
            `--+--Phiomia Andrews & Beadnell 1902 SW06, SKR04 [Phiomiidae]
               |    |--P. major Sanders, Kappelman & Rasmussen 2004 SKR04
               |    |--P. minor (Andrews 1904) S68
               |    |--P. serridens Andrews & Beadnell 1902 (see below for synonymy) CM78
               |    `--P. wintoni (Andrews 1905) S68
               `--Elephantimorpha SW06
                    |  i. s.: Notelephas Owen 1882 LA02
                    |           `--*N. australis Owen 1882 LA02
                    |--Hemimastodon crepusculi TS88
                    |--Eritreum Shoshani, Walter et al. 2006 SW06
                    |    `--*E. melakeghebrekristosi Shoshani, Walter et al. 2006 SW06
                    `--+--Elephantida SW06
                       `--Mammutidae [Mammutida, Mammutoidea] SW06
                            |--Eozygodon morotoensis SW06
                            |--Miomastodon TS88
                            |--Pliomastodon TS88
                            |--Zygolophodon Vacek 1877 SW06, CM78
                            |    |  i. s.: Z. aegyptensis SW06
                            |    |         Z. gobiensis SW06
                            |    |         Z. metachinjiensis SW06
                            |    |         Z. turicensis SW06
                            |    `--Z. (Zygolophodon) CM78
                            |         |--*Z. (Z.) tapiroides CM78
                            |         `--Z. (Z.) borsoni (Hays 1834) [=Mastodon borsoni] CM78
                            `--Mammut Blumenbach 1799 TS88, H07 [incl. Mastodon Cuvier 1806 CM78]
                                 |--M. americanum D07
                                 |--‘Mastodon’ andium C77
                                 |--‘Mastodon’ australis Owen 1844 LA02
                                 |--‘Mastodon’ chapmanii C77
                                 |--‘Mastodon’ humboldtii C77
                                 |--M. obscurus D07 [=Mastodon obscurus C77]
                                 |--‘Mastodon’ ohioticus C77
                                 |--‘Mastodon’ proavus C77
                                 |--‘Mastodon’ productus Cope 1874 C77
                                 |--‘Mastodon’ shepardii C77
                                 `--‘Mastodon’ tapiroides C77

*Palaeomastodon beadnelli Andrews 1901 [incl. P. barroisi Pontier 1907, P. intermedius Matsumoto 1922, P. minor Andrews 1904, P. minus Andrews 1905, P. parvus Andrews 1905, P. wintoni Andrews 1905] CM78

Phiomia serridens Andrews & Beadnell 1902 [=Palaeomastodon (Phiomia) serridens; incl. Ph. osborni Matsumoto 1924] CM78

*Type species of generic name indicated

REFERENCES

[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.

[CB78] Coppens, Y., & M. Beden 1978. Moeritherioidea. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 333–335. Harvard University Press: Cambridge (Massachusetts).

[CM78] Coppens, Y., V. J. Maglio, C. T. Madden & M. Beden. 1978. Proboscidea. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 336–367. Harvard University Press: Cambridge (Massachusetts).

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

[DRM86] Domning, D. P., C. E. Ray & M. C. McKenna. 1986. Two new Oligocene desmostylians and a discussion of tethytherian systematics. Smithsonian Contributions to Paleobiology 59: 1–56.

[H78] Harris, J. M. 1978. Deinotherioidea and Barytherioidea. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 315–332. Harvard University Press: Cambridge (Massachusetts).

[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.

[JB10] Jaeger, J.-J., K. C. Beard, Y. Chaimanee, M. Salem, M. Benammi, O. Hlal, P. Coster, A. A. Bilal, P. Duringer, M. Schuster, X. Valentin, B. Marandat, L. Marivaux, E. Métais, O. Hammuda & M. Brunet. 2010. Late middle Eocene epoch of Libya yields earliest known radiation of African anthropoids. Nature 467: 1095–1098.

[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.

[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.

[SW06] Shoshani, J., R. C. Walter, M. Abraha, S. Berhe, P. Tassy, W. J. Sanders, G. H. Marchant, Y. Libsekal, T. Ghirmai & D. Zinner. 2006. A proboscidean from the late Oligocene of Eritrea, a “missing link” between early Elephantiformes and Elephantimorpha, and biogeographical implications. Proceedings of the National Academy of Sciences of the USA 103 (46): 17296–17301.

[S68] Simons, E. L. 1968. African Oligocene mammals: introduction, history of study, and faunal succession. Peabody Museum of Natural History, Yale University, Bulletin 28: 1–21.

[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.

Last updated: 9 February 2022.