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.

Eulipotyphla

Reconstruction of Nesophontes, copyright Michael R. Long.


Belongs within: Pan-Lipotyphla.
Contains: Talpidae, Erinaceomorpha, Soricidae.

The Eulipotyphla, or 'insectivores', are a group of small mammals including shrews (Soricidae), hedgehogs (Erinaceidae), moles (Talpidae) and others. They are supported as a clade by molecular analyses, albeit one that excludes the African families Tenrecidae (tenrecs) and Chrysochloridae (golden moles) previously regarded as close relatives; these latter groups are instead allied to other Africa-derived mammals in a group called the Afrotheria. A number of fossil taxa previously regarded as related to this group are of uncertain placement as a result of this divide. Future analyses may identify them as true eulipotyphlans, as afrotherians, or as independent extinct groups.

Among the more divergent recent eulipotyphlans are the Solenodontidae (solenodons), moderately large (about a foot long excluding the tail) insectivores found only on the islands of Cuba and Hispaniola. Until recently, the Caribbean islands were also home to the now-extinct species of Nesophontes which were smaller than modern solenodons and possibly more superficially shrew-like. The Parapternodontidae from the early Eocene of Wyoming may represent continental relatives of solenodons (Halliday et al. 2017) but were placed closer to Soricidae by Asher et al. (2002).

Characters (from Butler 1988): Mobile snout, with enlarged olfactory organs, enlarged infraorbital canal, and reduced eyes. Palatine displaced from lacrimal by maxillar wing and confined to orbital floor; jugal reduced or absent, lower border of orbit formed from maxilla. Pubic symphysis reduced or lost. Caecum lost.

Eulipotyphla [Centetoidea, Dormaalioidea, Erinaceota, Lipotyphla, Soricoidea, Soricomorpha] HUG17
    |--+--+--+--Talpidae HUG17
    |  |  |  |--Proscalopidae [Arctoryctinae, Proscalopinae] B88
    |  |  |  |    |--Cryptoryctes Reed 1954 V67
    |  |  |  |    |--Oligoscalops Reed 1961 V67
    |  |  |  |    |--Proscalops Matthew 1901 [incl. Arctoryctes Matthew 1907] V67
    |  |  |  |    `--Mesoscalops Reed 1960 V67
    |  |  |  `--Dimylidae B88
    |  |  |       |--Dimylinae V67
    |  |  |       |    |--Dimylus von Meyer 1846 V67
    |  |  |       |    |    `--D. paradoxus D07
    |  |  |       |    |--Exodaenodus Hürzeler 1944 V67
    |  |  |       |    |--Dimyloides Hürzeler 1944 V67
    |  |  |       |    |--Cordylodon von Meyer 1859 V67
    |  |  |       |    |--Pseudocordylodon Hürzeler 1944 V67
    |  |  |       |    `--Metacordylodon Schlosser 1911 V67
    |  |  |       `--Plesiodimylus Gaillard 1897 [Pseudodimylinae] V67
    |  |  `--+--Wyonycteris HUG17
    |  |     |    |--W. chalix HUG17
    |  |     |    `--W. primitivus HUG17
    |  |     `--+--Erinaceomorpha HUG17
    |  |        `--Litocherinae B88
    |  |             |  i. s.: Cedrocherus B88
    |  |             |--Litocherus HUG17
    |  |             |    |--L. lacunatus HUG17
    |  |             |    |--L. notissimus HUG17
    |  |             |    `--L. zygeus HUG17
    |  |             `--+--Litolestes Jepsen 1930 B88, M60
    |  |                |    |--L. ignotus B88
    |  |                |    `--L. notissimus Simpson 1936 M60
    |  |                `--Eolestes B88
    |  `--Soricidae HUG17
    `--+--Solenodontidae [Solenodontinae, Solenodontini] AM02
       |    |--Clinopternodus Clark 1937 [=Clinodon Scott & Jepsen 1936 non Regan 1920] AM02
       |    `--Solenodon Brandt 1833 FS15, M58 [incl. Antillogale Patterson 1962 V67, Atopogale Cabrera 1925 M58]
       |         |  i. s.: S. arredondoi Morgan & Ottenwalder 1993 AC98
       |         |--*S. paradoxus Brandt 1833 M58, FS15, M58 [=Sorex (Solenodon) paradoxus M58]
       |         `--+--S. cubanus Peters 1864 FS15, M58 [=*Atopogale cubana M58; incl. S. poeyanus Barbour 1944 V67]
       |            `--S. marcanoi FS15
       |--Nesophontes Anthony 1916 FS15, M58 [Nesophontidae, Nesophontoidea]
       |    |--*N. edithae Anthony 1916 M58
       |    |--N. hypomicrus Miller 1929 M58
       |    |--N. longirostris Anthony 1919 M58
       |    |--N. major FS15
       |    |--N. micrus Allen 1917 M58
       |    |--N. paramicrus Miller 1929 M58
       |    |--N. submicrus AC98
       |    `--N. zamicrus Miller 1929 M58
       `--Parapternodontidae AM02
            |--Parapternodus Brown & Schankler 1982 HUG17, AM02
            |    `--*P. antiquus Brown & Schankler 1982 AM02
            `--Koniaryctes Robinson & Kron 1998 AM02
                 `--*K. paulus Robinson & Kron 1998 AM02

Eulipotyphla incertae sedis:
  Iconapternodus Tong 1997 AM02
    `--I. qii Tong 1997 AM02
  Ceciliolemur Weigelt 1933 [incl. Microtarsioides Weigelt 1933] S62
  Pontifactor B88
  Deinogalerix Fruedenthal 1973 D07
    `--D. koeingswaldi D07
  Oligoryctes Hough 1956 [Oligoryctidae] AM02
    |--*O. cameronensis Hough 1956 AM02
    `--O. altitalonidus (Clark 1937) [=Apternodus altitalonidus] AM02
  Plesiosoricidae B88
    |--Butselia Quinet & Misonne 1965 B88, V67 [Butselidae]
    |--Pakilestes B88
    |--Plesiosorex Pomel 1848 V67
    |    |--P. germanicus M60
    |    `--P. soricinoides M60
    |--Entomacodon Matthew 1909 V67
    `--Meterix Hall 1929 V67
         `--M. latidens M60

*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.

[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.

[B88] Butler, P. M. 1988. Phylogeny of the insectivores. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 117–141. Clarendon Press: Oxford.

[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.

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

[M58] McDowell, S. B., Jr. 1958. The Greater Antillean insectivores. Bulletin of the American Museum of Natural History 115 (3): 113–214.

[M60] McKenna, M. C. 1960. The Geolabidinae: a new subfamily of early Cenozoic erinaceoid insectivores. University of California Publications in Geological Sciences 37 (2): 131–164.

[S62] Simons, E. L. 1962. A new Eocene primate genus, Cantius, and a revision of some allied European lemuroids. Bulletin of the British Museum (Natural History): Geology 7 (1): 1–36.

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

Pholidota

Chinese pangolin Manis pentadactyla, copyright Skink Chen.


Belongs within: Pholidotamorpha.

The Pholidota (pangolins) are toothless, insectivorous mammals known from Africa and Asia. Modern species are characterised by a covering of overlapping keratinous scales, and have powerful forelimbs adapted for digging.

Pholidota
    |  i. s.: Euromanis krebsi OB13
    |         Eurotamandua Storch 1981 HUG17, BAA04
    |           `--E. joresi Storch 1981 S03
    |         Argyromanis patagonica A05a
    |--Eomanis Storch 1978 [Eomanidae] S03
    |    |--E. krebsi D07
    |    `--E. waldi Storch 1978 S03
    |--Patriomanidae S03
    |    |--Patriomanis Emry 1970 S03
    |    |    `--*P. americanus Emery 1970 E70
    |    `--Necromanis Filhol 1893 (see below for synonymy) S03
    |         |--‘Leptomanis’ edwardsi Filhol 1894 E70
    |         |--‘Lutra’ franconica Quenstedt 1885 E70 [=Potamotherium franconicum A05b, *Teutomanis franconica A05b]
    |         |--N. parva von Koenigswald 1969 E70
    |         |--‘Teutomanis’ quenstedti Ameghino 1905 [incl. *Galliaetatus schlosseri Ameghino 1905] E70
    |         `--N. quercyi Filhol 1894 E70
    `--Manidae P78
         |--+--Smutsia Gray 1865 FS15, P78
         |  |    |--S. gigantea FS15 [=Manis gigantea M94, Phataginus (Smutsia) giganteus P78]
         |  |    `--S. temminckii FS15 [=Manis temminckii ANG03, Phataginus (Smutsia) temmincki P78]
         |  `--+--Uromanis Pocock 1924 FS15, E70
         |     |    `--U. tetradactyla FS15
         |     `--Phataginus Rafinesque 1821 FS15, P78
         |          |--P. longicaudata P78
         |          `--P. tricuspis P78 [=Manis tricuspis M94]
         `--Manis Linnaeus 1758 FS15, P78 (see below for synonymy)
              |  i. s.: M. aurita P78
              |         M. hungrarica Kormos 1934 E70
              |         M. lydekkeri Dubois 1908 E70
              |         M. tetradactyla [incl. M. longicaudata] E70
              |--+--M. culionensis FS15
              |  `--M. javanica FS15 [=M. (Paramanis) javanica J68]
              `--+--M. paleojavanica Dubois 1907 FS15, E70
                 `--+--*M. pentadactyla Linnaeus 1758 L58, FS15, BAA04
                    `--M. crassicaudata FS15

Manis Linnaeus 1758 FS15, P78 [incl. Pangolinus Rafinesque 1821 E70, Paramanis Pocock 1924 E70, Phatages Sundevall 1843 E70, Pholidotus Brisson 1762 E70]

Necromanis Filhol 1893 [incl. Galliaetatus Ameghino 1905, Leptomanis Filhol 1893, Teutomanis Ameghino 1905] S03

*Type species of generic name indicated

REFERENCES

[A05a] Ameghino, F. 1905a. La perforación astragaliana en el Orycteropus y el origen de los Orycteropidae. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 59–95.

[A05b] Ameghino, F. 1905b. Les édentés fossiles de France et d'Allemagne. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 175–250.

[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.

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

[E70] Emry, R. J. 1970. A North American Oligocene pangolin and other additions to the Pholidota. Bulletin of the American Museum of Natural History 142 (6): 455–510.

[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.

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

[J68] Johnson, M. L. 1968. Application of blood protein electrophoretic studies to problems in mammalian taxonomy. Systematic Zoology 17 (1): 23–30.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[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.

[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.

[P78] Patterson, B. 1978. Pholidota and Tubulidentata. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 268–278. Harvard University Press: Cambridge (Massachusetts).

[S03] Storch, G. 2003. Fossil Old World “edentates” (Mammalia). Senckenbergiana Biologica 83: 51–60.

Ferae

Reconstruction of Chriacus, from Cox (1988).


Belongs within: Scrotifera.
Contains: Arctocyonidae, Pantodonta, Periptychidae, Pholidotamorpha, Didelphodontinae, Creodonta, Mesonychia, Carnivoramorpha.

The Ferae are a clade of mammals supported by molecular data uniting the modern Carnivora with the Pholidota (pangolins). A phylogenetic analysis of Palaeocene mammals by Halliday et al. (2017) identified a number of potential fossil members of this clade when results were constrained to correlate with modern molecular analyses but it remains to be seen whether these associations will be supported by future analyses.

<==Ferae [Ostentoria]
    |  i. s.: Triacodon Marsh 1871 V66
    |--+--+--Arctocyonidae HUG17
    |  |  `--+--+--Bisonalveus Gazin 1956 HUG17, V67
    |  |     |  |    |--B. browni HUG17
    |  |     |  |    `--B. holtzmani HUG17
    |  |     |  `--Pantodonta HUG17
    |  |     `--+--Pentacodontinae [Pentacodontidae] S78
    |  |        |    |--Coriphagus Douglass 1908 V67
    |  |        |    |    `--C. montanus Douglass 1908 S35b
    |  |        |    |--Pentacodon Scott 1892 HUG17, V67
    |  |        |    |    |--P. inversus S35b
    |  |        |    |    `--P. subtrigonus HUG17
    |  |        |    |--Aphronorus Simpson 1935 S78, S35b
    |  |        |    |    |--*A. fraudator Simpson 1935 S35b
    |  |        |    |    `--A. orieli HUG17
    |  |        |    |--Amaramnis Gazin 1962 V67
    |  |        |    |    `--A. gregoryi Gazin 1962 V67
    |  |        |    `--Protentomodon Simpson 1928 V67
    |  |        |         `--P. ursirivalis Simpson 1928 V67
    |  |        `--Periptychidae MC00
    |  `--+--Pholidotamorpha HUG17
    |     `--+--+--Oxyclaenus [Oxyclaenidae, Oxyclaeninae] HUG17
    |        |  |    |--O. antiquus [=Chriacus antiquus] V67
    |        |  |    |--O. cuspidatus V66
    |        |  |    |--O. simplex V66
    |        |  |    `--O. subbituminus HUG17
    |        |  `--+--+--Chriacus Cope 1883 HUG17, D07
    |        |     |  |    |--C. baldwini (Cope 1882) (see below for synonymy) SFY02
    |        |     |  |    |--C. calenancus D07
    |        |     |  |    |--C. gallinae V66
    |        |     |  |    |--C. katrinae D07
    |        |     |  |    |--C. metocemeti D07
    |        |     |  |    |--C. oconostotae HUG17
    |        |     |  |    |--C. pelvidens S35b
    |        |     |  |    |--C. pugnax Simpson 1935 S35b
    |        |     |  |    |--C. punitor D07
    |        |     |  |    `--C. pusillus Simpson 1935 S35b
    |        |     |  `--Thryptacodon Matthew & Granger 1915 HUG17, SFY02 [incl. Metachriacus Simpson 1935 S35b]
    |        |     |       |--T. antiquus Matthew & Granger 1915 SFY02
    |        |     |       |--T. australis Simpson 1935 SFY02
    |        |     |       |--T. orthogonius (Russell 1929) (see below for synonymy) SFY02
    |        |     |       `--T. pseudarctos S35a
    |        |     `--+--Claenodon HUG17 [incl. Neoclaenodon Gidley 1919 S35b]
    |        |        |    |--C. ferox (Cope 1883) S35b
    |        |        |    |--C. latidens (Gidley 1919) S35b
    |        |        |    |--C. montanensis (Gidley 1919) S35b
    |        |        |    |--C. procyonoides V66
    |        |        |    |--C. silberlingi (Gidley 1919) S35b
    |        |        |    `--C. vecordensis Simpson 1935 S35b
    |        |        `--Anacodon ursidens HUG17
    |        `--+--Didelphodontinae HUG17
    |           |--+--Aaptoryctes ivyi HUG17
    |           |  `--Palaeoryctidae [Palaeoryctae, Palaeoryctinae] FN03
    |           |       |--Alveugena carbonensis Eberle 1999 FN03
    |           |       |--Palaeoryctes Matthew 1913 HUG17, M58
    |           |       |    |--*P. puercensis Matthew 1913 M58
    |           |       |    |--P. cruoris D07
    |           |       |    |--P. jepseni HUG17
    |           |       |    |--P. minimus D07
    |           |       |    `--P. punctatus Van Valen 1966 V66
    |           |       `--Pararyctes Van Valen 1966 V67
    |           |            |--*P. pattersoni Van Valen 1966 V66
    |           |            `--P. rutherfordi Scott, Fox & Youzwyshyn 2002 SFY02
    |           `--Creodonta HUG17
    `--+--+--Goniacodon HUG17
       |  |    |--G. hiawathae HUG17
       |  |    `--G. levisanus HUG17
       |  `--+--Eoconodon HUG17
       |     |    |--E. copanus HUG17
       |     |    |--E. gaudrianus V66
       |     |    `--E. heilprinianus HUG17
       |     `--Triisodon O98
       |          |--T. antiquus O98
       |          `--T. quivirensis O98
       `--+--Mesonychia HUG17
          `--Carnivoramorpha HUG17

Chriacus baldwini (Cope 1882) [incl. Metachriacus provocator Simpson 1935, C. truncatus Cope 1884] SFY02

Thryptacodon orthogonius (Russell 1929) SFY02 [=Chriacus orthogonius SFY02; incl. T. belli Gazin 1956 SFY02, T. demari Gazin 1956 SFY02, *Metachriacus punitor Simpson 1935 S35b, SFY02]

*Type species of generic name indicated

REFERENCES

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

[E70] Emry, R. J. 1970. A North American Oligocene pangolin and other additions to the Pholidota. Bulletin of the American Museum of Natural History 142 (6): 455–510.

[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.

[M58] McDowell, S. B., Jr. 1958. The Greater Antillean insectivores. Bulletin of the American Museum of Natural History 115 (3): 113–214.

[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.

[O98] O’Leary, M. A. 1998. Phylogenetic and morphometric reassessment of the dental evidence for a mesonychian and cetacean clade. In: Thewissen, J. G. M. (ed.) The Emergence of Whales pp. 133–161. Plenum Press: New York.

[S78] Savage, R. J. G. 1978. Carnivora. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 249–267. Harvard University Press: Cambridge (Massachusetts).

[SFY02] Scott, C. S., R. C. Fox & G. P. Youzwyshyn. 2002. New earliest Tiffanian (late Paleocene) mammals from Cochrane 2, southwestern Alberta, Canada. Acta Palaeontologica Polonica 47: 691–704.

[S35a] Simpson, G. G. 1935a. The Tiffany fauna, Upper Paleocene. I.–Multituberculata, Marsupialia, Insectivora, and ?Chiroptera. American Museum Novitates 795: 1–19.

[S35b] Simpson, G. G. 1935b. New Paleocene mammals from the Fort Union of Montana. Proceedings of the United States National Museum 83 (2981): 221–244.

[V66] Van Valen, L. 1966. Deltatheridia, a new order of mammals. Bulletin of the American Museum of Natural History 132 (1): 1–126.

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

Hyaenidae

Striped hyaena Hyaena hyaena, copyright Balaji Venkatesh Sivaramakrishnan.


Belongs within: Aeluroidea.

The Hyaenidae, hyaenas, are cursorial carnivores known from Africa and Eurasia, with fossil representatives in North America. Hyaenas have relatively short bodies with conspicuously down-sloping backs, and are conspicuously marked with stripes or spots.

<==Hyaenidae
    |--Crocuta FS15
    |    |--C. crocuta RB84
    |    |    |--C. c. crocuta RB84
    |    |    `--C. c. spelaea RB84
    |    |--C. eximia K54
    |    |--C. honanensis DW04
    |    `--C. variabilis K54
    `--+--Proteles cristatus (Sparrman 1783) FS15, V10
       `--Hyaena FS15
            |--H. abronia S78
            |--H. bellax S78
            |--H. brunnea S78
            |--H. hyaena (Linnaeus 1758) DC02
            |    |--H. h. hyaena USDI77
            |    `--H. h. barbara USDI77
            |--H. priscus ME05
            `--H. striata K54

Hyaenidae incertae sedis:
  Parahyaena brunnea YC04
  Hyaenictis preforfex S78
  Leecyaena forfex [=Hyaenictis forfex] S78
  Progenetta S78
  Miohyaena certa K54
  Chasmaporthetes YC04
    |--C. ossifragus V91
    `--C. progressus DW04
  Euryboas S78
    |--E. namaquensis [=Hyaena namaquensis, Lycyaena namaquensis] S78
    `--E. silberbergi [=Lycyaena silberbergi] S78
  Pachycrocuta ME05
    |--P. brevirostris ME05
    |--P. licenti DW04
    `--P. sinensis DW04
  Percrocuta Schlosser 1903 S78, D07
    |--P. algeriensis [=Hyaena algeriensis] S78
    |--P. australis Hendey 1974 S78
    |--P. gigantea D07
    |--P. hebiensis D07
    |--P. miocenica D07
    |--P. tobieni Crusafont & Aguirre 1971 S78
    `--P. tungurensis D07
  Ictitherium Wagner 1848 (see below for synonymy) K54
    |  i. s.: I. arkesilai D07
    |         I. ibericum D07
    |         I. intuberculatum D07
    |         I. kurteni D07
    |         I. orbignyi K54
    |         I. pannonicum D07
    |--+--I. hipparionum [=Hyaena hipparionum] K54
    |  `--I. wongii K54 [=Hyaenictitherium wongii DW04]
    `--+--I. robustum (Nordmann ex Gervais 1858) (see below for synonymy) K54
       |    |--I. r. robustum K54
       |    `--I. r. tauricum K54
       `--+--I. gaudryi K54
          `--+--I. hyaenoides K54 [=Hyaenictitherium hyaenoides DW04]
             |--I. indicum K54
             `--I. sivalense K54

Inorganic: Proteles cristatus minilorientalus Okamura 1987 O87

Ictitherium Wagner 1848 [incl. Galeotherium Wagner 1840 non Jäger 1839, Hyaenictitherium Kretzoi 1937, Lepthyaena Lydekker 1884, Palhyaena Gervais 1859, Sinictitherium Kretzoi 1937, Thalassictis Nordmann ex Gervais 1850] K54

Ictitherium robustum (Nordmann ex Gervais 1858) [=Thalassictis robusta; incl. T. gracilis Hensel 1862, Ictitherium viverrinum Roth & Wagner 1854] K54

*Type species of generic name indicated

REFERENCES

[DC02] De, J. K., & R. Chakraborty. 2002. Identification of dorsal guard hairs of striped hyena Hyaena hyaena (Linnaeus, 1758) Hyaenidae: Carnivora: Mammalia. Journal of the Bombay Natural History Society 99 (3): 502–506.

[DW04] Deng T., Wang X., Ni X. & Liu L. 2004. Sequence of the Cenozoic mammalian faunas of the Linxia Basin in Gansu, China. Acta Geologica Sinica (English Edition) 78 (1): 8–14.

[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.

[K54] Kurtén, B. 1954. The type collection of Ictitherium robustum (Gervais, ex Nordmann) and the radiation of the ictitheres. Acta Zoologica Fennica 86: 1–26.

[ME05] Moullé, P.-E., A. Echassoux, F. Lacombat, E. Desclaux & S. Bailon. 2005. L’environnement animal des premiers habitants de l’Europe méditerranéenne: les grands mammifères contemporains de l’homme du Vallonnet, données taxonomiques et biostratigraphiques pour la deuxième moitie du Pléistocène inférieur. BAR International Series 1364: 105–113.

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.

[RB84] Richardson, P. K. R., & S. K. Bearder. 1984. The hyaena family. In: Macdonald, D. (ed.) All the World’s Animals: Carnivores pp. 146–151. Torstar Books Inc.: New York.

[S78] Savage, R. J. G. 1978. Carnivora. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 249–267. Harvard University Press: Cambridge (Massachusetts).

[USDI77] USDI (United States Department of the Interior). 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.

[V91] Valkenburgh, B. van. 1991. Iterative evolution of hypercarnivory in canids (Mammalia: Carnivora): evolutionary interactions among sympatric predators. Paleobiology 17 (4): 340–362.

[V10] Vršanský, P. 2010. Cockroach as the earliest eusocial animal. Acta Geologica Sinica (English Edition) 84 (4): 793–808.

[YC04] Yamaguchi, N., A. Cooper, L. Werdelin & D. W. Macdonald. 2004. Evolution of the mane and group-living in the lion (Panthera leo): a review. Journal of Zoology 263: 329–342.

Hyopsodontidae

Partial lower jaw of Litomylus dissentaneus, copyright Yale Peabody Museum of Natural History.


Belongs within: Euungulata.

The Hyopsodontidae are a group of small, generalised mammals known from the Palaeocene and Eocene of Europe and North America. Members of this group have fairly simple incisors and premolars, small canines, and more or less bunodont molars.

See also: Hyopsodontids: little slinkers of the Palaeogene.

Hyopsodontidae
    `--Tricuspiodontinae SFY02
         |--Tricuspiodon Lemoine 1885 V66
         `--Litomylus Simpson 1935 SFY02
              |--*L. dissentaneus Simpson 1935 SFY02
              |--L. grandaletes Scott, Fox & Youzwyshyn 2002 SFY02
              |--L. ishami Gazin 1956 SFY02
              |--L. orthronepius Johnston & Fox 1984 SFY02
              |--L. osceolae Van Valen 1978 SFY02
              `--L. scaphicus Gazin 1956 SFY02

Hyopsodontidae incertae sedis:
  Pascatherium Russell 1964 N85, V67
    `--P. dolloi (Teilhard de Chardin 1927) M60 [=Adapisorex dolloi V67]
  Haplaletes Simpson 1935 S35
    `--*H. disceptatrix Simpson 1935 S35
  Hyopsodus Leidy 1870 D07
    |--*H. paulus C77
    |--H. despiciens D07
    |--H. egressus HUG17
    |--H. gracilis D07
    |--H. lepidus D07
    |--H. loomisi D07
    |--H. marshi D07
    |--H. minisculus D07
    |--H. miticulus (Cope 1874) [=Esthonyx miticulus] C77
    |--H. vicarius C77
    |--H. wardi D07
    `--H. wortmani D07
  Aletodon Gingerich 1977 SFY02
    `--A. quadravus SFY02

*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.

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

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

[N85] Novacek, M. J. 1985. The Sespedectinae, a new subfamily of hedgehog-like insectivores. American Museum Novitates 2822: 1–24.

[SFY02] Scott, C. S., R. C. Fox & G. P. Youzwyshyn. 2002. New earliest Tiffanian (late Paleocene) mammals from Cochrane 2, southwestern Alberta, Canada. Acta Palaeontologica Polonica 47: 691–704.

[S35] Simpson, G. G. 1935. New Paleocene mammals from the Fort Union of Montana. Proceedings of the United States National Museum 83 (2981): 221–244.

[V66] Van Valen, L. 1966. Deltatheridia, a new order of mammals. Bulletin of the American Museum of Natural History 132 (1): 1–126.

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

Arctocyonidae

Reconstruction of Arctocyon by Dmitry Bogdanov.


Belongs within: Ferae.

The Arctocyonidae are a group of probably omnivorous mammals known from the Palaeocene and early Eocene of Europe and North America. They are generalised in morphology, with more or less tritubercular teeth, simple premolars, and moderately long limbs with paraxonic feet and claw-like unguals.

<==Arctocyonidae [Arctocyoninae]
    |--+--Protogonodon pentacus PMF88, S35
    |  `--Deuterogonodon Simpson 1935 PMF88, S35
    |       `--*D. montanus Gidley in Simpson 1935 [=Protogonodon montanus Gidley in Simpson 1935 (n. n.)] S35
    `--+--Mimotricentes PMF88
       `--+--Mentoclaenodon PMF88
          `--+--+--Procynictis V66
             |  `--Arctocyonides ferox PMF88, V66
             `--Arctocyon PMF88
                  |--*A. primaevus de Blainville 1841 HUG17
                  |--A. corrugatus HUG17
                  `--A. ferox PMF88

Arctocyonidae incertae sedis:
  Loxolophus [Loxolophinae] MC00
    |--L. hyattianus HUG17
    |--L. pentacus HUG17
    `--L. priscus HUG17
  Spanoxydon Simpson 1935 S35
    `--*S. latrunculus Simpson 1935 S35
  Carcinodon GH88
  Deltatherium [incl. Lipodectes Cope 1881] V66
    `--D. fundaminis V66
  Prothryptacodon Simpson 1935 PMF88, S35
    `--*P. furens Simpson 1935 S35
  Tricentes PMF88
    |--T. latidens Gidley in Simpson 1935 S35
    `--T. subtrigonus S35
  ‘Tetraclaenodon’ minor (Matthew 1897) T90

*Type species of generic name indicated

REFERENCES

[GH88] Gentry, A. W., & J. J. Hooker. 1988. The phylogeny of the Artiodactyla. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 235–272. Clarendon Press: Oxford.

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

[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.

[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.

[S35] Simpson, G. G. 1935. New Paleocene mammals from the Fort Union of Montana. Proceedings of the United States National Museum 83 (2981): 221–244.

[T90] Thewissen, J. G. M. 1990. Evolution of Paleocene and Eocene Phenacodontidae (Mammalia, Condylarthra). Museum of Paleontology Papers on Paleontology 29: 1–107.

[V66] Van Valen, L. 1966. Deltatheridia, a new order of mammals. Bulletin of the American Museum of Natural History 132 (1): 1–126.