Helogale

Common dwarf mongoose Helogale parvula, copyright Bernard Dupont.

Belongs within: Herpestidae.

Helogale, the dwarf mongooses, is a genus of small, diurnal, social mongooses with a grizzled brown pelage found in eastern and southern Africa.

Helogale
    |--H. brunnula S78
    |--H. dybowskii S78
    |--H. hirtula S78
    |--H. ivori S78
    |--H. macmillani S78
    |--H. mimetra S78
    |--H. parvula S78
    |--H. percivali S78
    |--H. undulata S78
    |--H. varia S78
    |--H. vetula S78
    `--H. victorina S78

*Type species of generic name indicated

REFERENCES

[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).

Connochaetes

Black wildebeest Connochaetes gnou, copyright Bernard Dupont.

Belongs within: Alcephalinae.

Connochaetes, the gnus or wildebeest, is a genus of cattle-like grazing antelopes found in grasslands and open woodland of southern and eastern Africa.

Characters (from Murray 1984): Head massive, both sexes horned. Mane present on neck and shoulders, beard under throat. Tail long, reaching nearly to ground.

Connochaetes [incl. Pultiphagonides Hopwood 1934] G78
    |--+--C. africanus (Hopwood 1934) [=*Pultiphagonides africanus] G78
    |  `--C. gnou G78
    |       |--C. g. gnou G78
    |       |--C. g. antiquus Broom 1913 G78
    |       `--C. g. laticornutus (van Hoepen 1932) G78
    `--C. taurinus G78
         |--C. t. taurinus G78
         |--C. t. cooksoni M84
         |--C. t. mearnsi M84
         |--C. t. olduvaiensis G78
         `--C. t. prognu G78

*Type species of generic name indicated

REFERENCES

[G78] Gentry, A. W. 1978. Bovidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 540–572. Harvard University Press: Cambridge (Massachusetts).

[M84] Murray, M. G. 1984. Grazing antelopes. In: Macdonald, D. (ed.) All the World’s Animals: Hoofed Mammals pp. 120–131. Torstar Books: New York.

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.

Litopterna

Mounted skeleton of Macrauchenia patachonica (larger mount) alongside Phenacodus primaevus, copyright Ryan Somma.

Belongs within: Panameriungulata.

The Litopterna are a group of more or less cursorial herbivores known from the late Palaeocene to the Pleistocene of South America. Distinguishing features of the group include fusion of the mandibular symphysis, and characters of the tarsal bones (Muizon & Cifelli 2000).

<==Litopterna
    |--Neocaliphrium recens FS15
    `--+--Xenorhinotherium bahiense FS15
       `--Macrauchenia Owen 1838 FS15, D07
            `--M. patachonica FS15

Litopterna incertae sedis:
  Miguelsoria parayirunhor (Paula Couto 1952) [=Ernestokokenia parayirunhor] MC00
  Proectocion argentinus MC00
  Requisia vidmari MC00
  Polymorphis MC00
  Cramauchenia MC00
  Proadiantus MC00
  Protolipterna [Protolipternidae] MC00
    `--P. ellipsodontoides Cifelli 1983 MC00
  Thoatherium Ameghino 1887 D07
    `--T. minisculum D07
  Asmithwoodwardia Paula Couto 1952 MC00
    |--A. scotti Paula Couto 1952 MC00
    `--A. subtrigona MC00
  Diadiaphorus Ameghino 1887 D07
    |--D. caniadensis D07
    |--D. majusculus D07
    `--D. zamius D07
  Theosodon Ameghino 1887 D07
    |--T. garretorum D07
    |--T. gracilis D07
    |--T. lydekkeri D07
    `--T. tontanus D07

*Type species of generic name indicated

REFERENCES

[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.

[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.

Hipparion

Mounted skeleton of Hipparion primigenium at the Staatliches Museum für Naturkunde Stuttgart, copyright Ghedoghedo.

Belongs within: Equinae.

Hipparion is a diverse genus of three-toed horses known from the Miocene to the Pleistocene of Eurasia, Africa and North America.

Characters (from Churcher & Richardson 1978): Facial region relatively short and deep; preorbital fossae present or absent; sagittal crest low; bony auditory meatus long and directed dorsolaterally; basicranial region usually with prominent longitudinal crest; occiput with deep fossae above occipital condyles for attachment of nuchal musculature. Coronoid process of dentary high; ascending ramus vertical. Incisors large, cement filled; infundibula typically present on upper and lower I1 and I2; in I₃ infundibulum closed or partly closed. Canines usually present in both sexes, possibly dimorphic. Cheek teeth moderately to very hypsodont, slightly bowed, prismatic. Upper cheek teeth with isolated, more or less elongate protocones, almost always with many plis in fossettes. Lower cheek teeth with well-developed double loop, metaflexid with subequal arms and deep median valley; M₃ usually with tripartite talonid; well-developed ectostylid in milk lower cheek teeth of all forms where known and in permanent lower cheek teeth of more derived forms. Limbs with angled joints, ulna complete. Metapodials moderately to very elongate; metapodials III with traces of II and IV contacts along shafts; metapodials II and IV variably developed, apparently without reduction in size through time; vestigial metacarpal V present. Digits II, III and IV functional but II and IV shorter and thinner than III. Terminal phalanges (ungules) cut by deep median slit.

<==Hipparion de Christol 1832 F99 (see below for synonymy)
    |--H. sitifense Pomel 1897 [=H. sitifensis] CR78
    |    |--H. s. sitifense CR78
    |    `--H. s. gromovae Villalta & Crusafont 1957 CR78
    |--H. turkanense Hooijer & Maglio 1973 CR78
    `--+--H. primigenium (von Meyer 1829) [=Equus primigenius; incl. H. gracile] CR78
       `--+--+--H. baardi Boné & Singer 1965 [=H. (Hipparion) albertense baardi] CR78
          |  `--H. namaquense (Haughton 1932) [=Notohipparion namaquense] CR78
          `--+--H. afarense Eisenmann 1976 CR78
             `--H. libycum Pomel 1897 (see below for synonymy) CR78
                  |--H. l. libycum CR78
                  |--H. l. ethiopicum (Joleaud 1933) [=Libyhipparion ethiopicum] CR78
                  `--H. l. steytleri van Hoepen 1930 [=Libyhipparion steytleri, Stylohipparion steytleri] CR78

Hipparion incertae sedis:
  H. africanum Arambourg 1959 F99
  H. antelopinum Falconer & Cautley 1847 F99
  H. anthonyi D07
  H. brachypus Hensel 1862 F99
  H. campbelli Bernor 1985 F99
  H. catalaunicum Pirlot 1956 F99
  H. chiai DW04
  H. coelophyes DW04
  H. concudense D07
  H. dermatorhinum Sefve 1927 F99
  H. dongxiangense DW04
  H. elegans Gromova 1952 F99
  H. fossatum Sefve 1927 F99
  H. garedzicum Gabuniya 1959 F99
  H. gettyi Bernor 1985 F99
  H. giganteum Gromova 1952 F99
  H. hippidiodus Sefve 1927 [incl. H. platygenys Gromova 1952] F99
  H. houfenense DW04
  H. huangheense D07
  H. licenti Qiu, Huang & Guo 1987 F99
  H. macedonicum Koufos 1984 F99
  H. matthewi Abel 1926 F99
  H. mediterraneum (Roth & Wagner 1855) F99
  H. microtaton Nikolov 1971 F99
  H. mogoicum Zhegallo 1978 F99
  H. mohavense D07
  H. molayanense Zouhri 1992 F99
  H. moldavicum Gromova 1952 F99
  H. nesebricum Bakalov & Nikolov 1962 F99
  H. nicosi Bernor & Tobien 1989 F99
  H. parvum DW04
  H. periafricanum Villalta & Crusafont 1957 F99
  H. platystyle D07
  H. plocodus Sefve 1927 F99
  H. praegiganteum Tarabukin 1967 F99
  H. presulcatum Nikolov 1971 F99
  H. proboscideum Studer 1911 F99
  H. prostylum Gervais 1849 F99
  H. sarmaticum Lungu 1973 F99
  H. schlosseri Antonius 1919 [incl. Hemihipparion dietrichi Wehrli 1941, Hi. dietrichi] F99
  H. shirleyi MacFadden 1984 F99
  H. sinense DW04
  H. tchicoicum Ivanjev 1966 F99
  H. tudorovense Gabuniya 1959 F99
  H. urmiense Gabuniya 1959 F99
  H. verae Gabuniya 1979 [=H. gromovae Gabuniya 1959 non Villalta & Crusafont 1957] F99
  H. weihoense Liu, Li & Zhai 1978 F99

Hipparion de Christol 1832 F99 [incl. Eurygnathohippus van Hoepen 1930 CR78, Libyhipparion Joleaud 1933 CR78, Notohipparion Haughton 1932 CR78, Stylohipparion van Hoeppen 1932 CR78]

Hipparion libycum Pomel 1897 [=H. (Stylohipparion) libycum; incl. H. albertensis Hopwood 1926 (n. d.), H. albertense, Stylohipparion albertense, H. ambiguum Pomel 1897, Eurygnathohippus cornelianus van Hoepen 1930, Stylohipparion hipkini van Hoepen 1932, Hipparion massoesylium Pomel 1897, H. (Hipparion) albertense serengetense Boné & Singer 1965] CR78

*Type species of generic name indicated

REFERENCES

[CR78] Churcher, C. S., & M. L. Richardson. 1978. Equidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 379–422. Harvard University Press: Cambridge (Massachusetts).

[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.

[F99] Forsten, A. 1999. Snout proportions in some Eurasian hipparions (Mammalia, Equidae): taxonomic and functional implications. Geodiversitas 21 (2): 255–278.

Equinae

Mounted skeleton of Pliohippus in the Natural History Museum, Karlsruhe, copyright Ghedoghedo.

Belongs within: Equidae.
Contains: Hipparion, Equus.

The Equinae are a group of horses in which the lateral toes are reduced and non-functional, or lost entirely (Churcher & Richardson 1978).

Characters (from Churcher & Richardson 1978): Skull with postorbital bar. Upper incisors with more or less developed infundibula; in later forms on some or all lower incisors also. Cheek teeth hypsodont, crown height equal to or greater than mesiodistal diameter; cementum in fossettes and in later forms over all of crown; small enamel folds or plis present on lophs, often numerous. Premolars 2 to 4 fully molarised; P1s small or absent. Upper molars with protolophs and metalophs joining ectolophs even in early stages of wear; premolars with transverse lophs potentially remaining free until late stage of wear; fossettes usually closing with wear. Protocone separate or nearly detached from protoloph; elongate, with major diameter oriented mesiodistally. Lower molars with pattern composed of two crescentic lophids joined in middle of tooth; metaconid and metastylid completely fused, mesiodistally elongated, forming double loop in later forms. Ulna greatly reduced in shaft, sometimes incomplete, usually fused to radius. Metapodial III and phalanges relatively elongated; lateral metapodials and digits narrowed, moved posteriorly to limb axis; distal ends of metapodials bent posteriorly. Lateral hoofed phalanges laterally compressed, not extending to plantar surface of III; in latest forms lateral phalanges lost, lateral metapodials reduced to splint bones.

<==Equinae
    |--+--Nannippus Matthew 1926 C96, F99
    |  `--+--Hipparion CR78
    |     `--Neohipparion Gidley 1903 C96, F99
    |          |--N. affine Leidy 1869 F99
    |          `--N. coloradense Osborn 1918 F99
    `--+--Equus CR78
       `--Pliohippus Marsh 1874 C96, C77
            |--P. campestris D07
            |--P. fossulatus D07
            |--P. lullianus D07
            |--P. mexicanus D07
            |--P. mirabilis D07
            |--P. nobilis D07
            |--P. osborni D07
            |--P. pachyops D07
            |--P. pernix D07
            `--P. robustus D07

Equinae incertae sedis:
  Proboscidipparion Sefve 1927 F99
    |--P. houfenense (Teilhard & Young 1931) F99
    |--P. pater Matsumoto 1927 F99
    |--P. rocinantis (Hernandez-Pacheco 1921) F99
    `--P. sinense Sefve 1927 F99
  Pseudhipparion Ameghino 1904 F99
  Dinohippus D07
    |--D. interpolatus D07
    |--D. leardi D07
    |--D. leidyanus D07
    |--D. mexicanus D07
    `--D. spectans D07

*Type species of generic name indicated

REFERENCES

[C96] Campbell, N. A. 1996. Biology 4^th ed. The Benjamin/Cummings Publishing Company, Inc.: Menlo Park (California).

[CR78] Churcher, C. S., & M. L. Richardson. 1978. Equidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 379–422. Harvard University Press: Cambridge (Massachusetts).

[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.

[F99] Forsten, A. 1999. Snout proportions in some Eurasian hipparions (Mammalia, Equidae): taxonomic and functional implications. Geodiversitas 21 (2): 255–278.

Equidae

Mounted skeleton of Merychippus in the Nebraska State Museum of Natural History, copyright James St. John.

Belongs within: Equoidea.
Contains: Equinae, Anchitheriinae.

The Equidae, horses and their fossil relatives, are a group of perissodactyls known from the early Eocene to the present. Most members of the family are distinctly cursorial though the earliest examples were less so.

<==Equidae
    |--Minippus Froehlich 2002 F02
    |    |--*M. index (Cope 1873) [=Hyracotherium index] F02
    |    `--M. jicarillai Froehlich 2002 F02
    `--Arenahippus Froehlich 2002 F02
         |--*A. grangeri (Kitts 1956) [=Hyracotherium angustidens grangeri] F02
         `--+--A. aemulor (Gingerich 1991) [=Hyracotherium aemulor] F02
            `--+--A. pernix (Marsh 1876) [=Hyracotherium pernix, Eohippus pernix; incl. H. seekinsi Morris 1968] F02
               `--+--Xenicohippus Bown & Kihm 1981 F02
                  |    |--X. craspedotum (Cope 1880) [=Hyracotherium craspedotum, Eohippus craspedotus] F02
                  |    `--+--*X. grangeri Bown & Kihm 1981 F02
                  |       `--X. osborni Bown & Kihm 1981 F02
                  `--+--Eohippus Marsh 1876 F02
                     |    `--E. angustidens (Cope 1875) F02 (see below for synonymy)
                     `--+--Pliolophus vulpiceps Owen 1858 [=Hyracotherium vulpiceps] F02
                        `--Protorohippus Wortman 1896 F02
                             |--P. montanum Wortman 1896 F02
                             `--+--*P. venticolum (Cope 1881) (see below for synonymy) F02
                                `--+--Haplohippus McGrew 1953 M76
                                   |--Orohippus Marsh 1874 F02, C77
                                   |    |--O. major F02
                                   |    `--O. pumilus Marsh 1872 C77
                                   `--+--Epihippus Marsh 1878 F02, D07
                                      |    |--E. gracilis D07
                                      |    |--E. parvus RH14
                                      |    `--E. uintensis D07
                                      `--+--Mesohippus Marsh 1876 M76, D07
                                         |    |--M. bairdi OB13
                                         |    |--M. eulophus D07
                                         |    |--M. gridlei D07
                                         |    |--M. intermedius D07
                                         |    |--M. obliquidens D07
                                         |    `--M. validus D07
                                         `--+--Miohippus grandis C96, OB13
                                            `--+--+--Parahippus Leidy 1858 C96, D07
                                               |  |    |--P. leonensis D07
                                               |  |    |--P. mourningi D07
                                               |  |    |--P. pawniensis D07
                                               |  |    `--P. pristinus D07
                                               |  `--+--+--Callipus C96
                                               |     |  `--Equinae M76
                                               |     `--Merychippus Leidy 1857 C96, D07
                                               |          |--M. brevidontus Bode 1934 B76
                                               |          |--M. californicus D07
                                               |          |--M. insignis D07
                                               |          |--M. intermontanus W69
                                               |          |--M. stylodontus W69
                                               |          `--M. sumani W69
                                               `--Anchitheriinae C96

Equidae incertae sedis:
  Hippotherium C77
    |--H. affine C77
    |--H. calamarium Cope 1875 C77
    |--H. gratum C77
    |--H. occidentale C77
    |--H. paniense C77
    `--H. speciosum C77
  Protohippus Leidy 1869 C77
    `--P. sejunctus C77
  Plesiohipparion TH03
  Cormohipparion F99
    |--C. goorisi MacFadden & Skinner 1981 F99
    |--C. occidentale Leidy 1856 F99
    `--C. sphenodus Cope 1885 F99
  Eoequus wilsoni Quinn 1955 F99
  Propachynolophus gaudryi RH14
  Eurohippus parvum RH14

Eohippus angustidens (Cope 1875) F02 [=Orohippus angustidens F02, Hyracotherium angustidens F02; incl. *E. validus Marsh 1876 F02, Orohippus cuspidatus Cope 1875 C77, Hyracotherium cuspidatum F02, H. loevii Cope 1877 F02, Orohippus vasacciensis Cope 1875 C77, H. vasacciense F02, Lophiotherium vasacciense C77]

*Protorohippus venticolum (Cope 1881) [=Eohippus venticolum, Hyracotherium vasacciense venticolum] F02

*Type species of generic name indicated

REFERENCES

[B76] Barnes, L. G. 1976. Outline of eastern North Pacific fossil cetacean assemblages. Systematic Zoology 25 (4): 321–343.

[C96] Campbell, N. A. 1996. Biology 4^th ed. The Benjamin/Cummings Publishing Company, Inc.: Menlo Park (California).

[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.

[F99] Forsten, A. 1999. Snout proportions in some Eurasian hipparions (Mammalia, Equidae): taxonomic and functional implications. Geodiversitas 21 (2): 255–278.

[F02] Froehlich, D. J. 2002. Quo vadis eohippus? The systematics and taxonomy of the early Eocene equids (Perissodactyla). Zoological Journal of the Linnean Society 134: 141–256.

[M76] MacFadden, B. J. 1976. Cladistic analysis of primitive equids, with notes on other perissodactyls. Systematic Zoology 25 (1): 1–14.

[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.

[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.

[TH03] Tedford, R. H., & C. R. Harington. 2003. An Arctic mammal fauna from the Early Pliocene of North America. Nature 425: 388–390.

[W69] Woodburne, M. O. 1969. Systematics, biogeography, and evolution of Cynorca and Dyseohyus (Tayassuidae). Bulletin of the American Museum of Natural History 141 (2): 271–356.

Equoidea

Reconstructed skeleton of Palaeotherium magnum, by Jean Albert Gaudry.

Belongs within: Perissodactyla.
Contains: Equidae.

The Equoidea are a clade of perissodactyls uniting the horses (Equidae) with the fossil Palaeotheriidae from the Eocene and Oligocene of Eurasia.

Synapomorphies (from Hooker & Dashzeveg 2004): Upper molar metaconule large; P³ with trigon broader, protoloph stronger, paraconule (unless subsumed by protoloph) and metaconid much larger and better defined, but smaller than protocone and protoconid respectively; lower M_1–2 distal cingulum lingual of hypoconulid absent.

<==Equoidea [Hippomorpha, Hyracotheriinae] HD04
    |--+--Cymbalophus Hooker 1984 HD04, F02
    |  |    `--*C. cuniculus (Owen 1842) [=Hyracotherium cuniculus] F02
    |  `--Systemodon Cope 1881 F02
    |       |--*S. tapirinus (Cope 1875) (see below for synonymy) F02
    |       `--S. etsagicus (Cope 1884) [=Hyracotherium angustidens etsagicum] F02
    `--+--Sifrhippus Froehlich 2002 HD04, F02
       |    `--*S. sandrae (Gingerich 1989) [=Hyracotherium sandrae] F02
       `--+--Equidae HD04
          `--Palaeotheriidae HD04
               |  i. s.: Anchilophus F02
               |--Hyracotherium Owen 1841 F02
               |    |--*H. leporinum Owen 1841 F02
               |    |--H. agile (Marsh 1873) [=Orohippus agilis] C77
               |    |--H. procyoninum (Cope 1872) [=Helotherium procyoninum] C77
               |    |--H. siderolithicum C77
               |    `--H. sylvaticum C77
               `--+--Propalaeotherium parvulum RH14, S12
                  `--+--+--Plagiolophus annectens RH14
                     |  `--Leptolophus nouleti RH14
                     `--Palaeotherium Cuvier 1804 RH14, H07
                          |--P. castrense D07
                          |--P. crassum D07
                          |--P. curtum D07
                          |--P. lautricense D07
                          |--P. magnum Cuvier 1804 H07
                          |--P. medium D07
                          `--P. ruetimeyeri D07

Systemodon tapirinus (Cope 1875) [=Hyracotherium tapirinum, Orohippus tapirinus; incl. H. cristatum Wortman 1896] F02

*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.

[F02] Froehlich, D. J. 2002. Quo vadis eohippus? The systematics and taxonomy of the early Eocene equids (Perissodactyla). Zoological Journal of the Linnean Society 134: 141–256.

[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.

[HD04] Hooker, J. J., & D. Dashzeveg. 2004. The origin of chalicotheres (Perissodactyla, Mammalia). Palaeontology 47 (6): 1363–1386.

[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.

[S12] Schaal, S. 2012. Messel Pit fossil site—the legacy of the environment and life of the Eocene. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 225–236. Springer.

Chalicotheriidae

Reconstruction of Chalicotherium, from here.

Belongs within: Perissodactyla.

The Chalicotheriidae are a group of perissodactyls known from the Eocene to Pleistocene of Eurasia and North America, with a limited record from Africa. Members of this group possessed feet with large fissured claws rather than hooves and may have been upright browsers in the manner of gorillas or ground sloths.

<==Chalicotheriidae
    |--Borissiakia betpakdalensis Borissiak 1946 [=Phyllotillon betpakdalensis] C75
    |--Nestoritherium C75
    |--Grangeria HD04
    |--Schizotherium HD04
    |    |--S. priscum C75
    |    `--S. turgaicum C75
    |--Phyllotillon B78
    |    |--P. bavaricum (von Koenigswald 1932) B78, C75 [=Metaschizotherium bavaricum B78]
    |    `--P. naricus C75
    |--Ancylotherium B78
    |    |  i. s.: A. hennigi (Dietrich 1942) [=Metaschizotherium hennigi; incl. M. transvaalensis George 1950] B78
    |    |         A. pentelicum B78
    |    `--A. (Metaschizotherium von Koenigswald 1932) C75
    |         `--A. (M.) fraasi (von Koenigswald 1932) [=Metaschizotherium fraasi] C75
    |--Moropus Marsh 1877 HD04, D07
    |    |--M. elatus Marsh 1877 [incl. M. parvus Barbour 1909, M. petersoni Holland 1908] C75
    |    |--M. hollandi D07
    |    |--M. matthewi D07
    |    |--M. merriami D07
    |    `--M. oregonensis D07
    `--Chalicotherium Kaup 1833 D07
         |--C. gigantium D07
         |--C. goldfussi C75
         |--C. grande [=Macrotherium grande] B78
         |--C. pilgrimi B78
         |--C. rusingense Butler 1962 B78
         |--C. salinum D07
         `--C. wetzleri B78

*Type species of generic name indicated

REFERENCES

[B78] Butler, P. M. 1978. Chalicotheriidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 368–370. Harvard University Press: Cambridge (Massachusetts).

[C75] Coombs, M. C. 1975. Sexual dimorphism in chalicotheres (Mammalia, Perissodactyla). Systematic Zoology 24 (1): 55–62.

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

[HD04] Hooker, J. J., & D. Dashzeveg. 2004. The origin of chalicotheres (Perissodactyla, Mammalia). Palaeontology 47 (6): 1363–1386.