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.

Camelidae

Dromedary camel Camelus dromedarius, copyright Jjron.

Belongs within: Artiodactyla.

The Camelidae are a group of long-necked herbivorous mammals with living species native to Africa, Asia and South America. Members of this group have a three-chambered ruminating stomach.

Characters (from Animal Diversity Web): Large, ranging in weight from 35 to 650 kg. Head small; neck long, gracile; legs long, slender. Upper lip deeply cleft. Fore legs plantigrade; toes splayed. Skull with rostrum elongate; sagittal crest well developed; postorbital bar complete. Adult dental formula 1/3, 1/1, 2-3/1-2, 3/3; juveniles with three pairs of upper incisors but adults with only one canine-like pair; lower incisors spatulate, projecting forward; canines medium-sized, hooked; cheek teeth selenodont. Wide diastema separating incisors and cheek teeth. Legs with ulna and fibula much reduced, present mainly as splint on radius or tibia. Third and fourth metapodials fusing to form cannon bone proximally only, splaying apart distally. First two phalanges of each digit flattened and expanded; last reduced, bearing a nail. Middle (second) phalanges imbedded in broad pad making up sole of foot.

<==Camelidae
    |--Camelus Linnaeus 1758 FS15, L58
    |    |--C. bactrianus Linnaeus 1758 GC-BG04
    |    |--C. dromedarius Linnaeus 1758 L58
    |    |--C. ferus Przewalski 1878 GC-BG04
    |    |--C. glama Linnaeus 1758 L58
    |    |--C. pacos Linnaeus 1758 L58
    |    |--C. sivalensis 78
    |    `--C. thomasi Pomel 1893 G78
    `--+--Palaeolama FS15
       |    |--P. weddeli FS15
       |    `--+--P. major FS15
       |       `--P. mirifica FS15
       `--+--Camelops hesternus FS15
          `--+--Hemiauchenia FS15
             |    |--H. macrocephala FS15
             |    `--H. paradoxa FS15
             `--+--Vicugna FS15
                |    |--V. pacos (Linnaeus 1758) GC-BG04
                |    `--V. vicugna (Molina 1782) GC-BG04
                `--Lama FS15
                     |--L. glama (Linnaeus 1758) GC-BG04
                     |--L. guanicoe (Müller 1776) GC-BG04
                     `--L. pacos F84

Camelidae incertae sedis:
  Auchenia lama C77
  Alticamelus M66
  Protolabis Cope 1876 C77
    `--P. heterodontus C77
  Pliauchenia Cope 1875 C77
    |--P. humphresiana Cope 1875 C77
    `--P. vulcanorum Cope 1875 C77
  Procamelus Leidy 1858 C77
    |--P. angustidens C77
    |--P. coartatus D07
    |--P. fissidens Cope 1876 C77
    |--P. gazini D07
    |--P. gracilis Leidy 1858 C77
    |--P. grandis D07
    |--P. leptocolon D07
    |--P. occidentalis Leidy 1858 C77
    `--P. robustus C77
  Protomeryx Leidy 1856 (n. d.) M66
    `--P. halli M66
  Pseudolabis [Pseudolabidinae] M66
    `--P. dakotensis Matthew 1904 M66
  0--Rakomylus M66
  `--Stenomylus Peterson 1908 M66, D07
       |--S. crassipes D07
       |--S. gracilis D07
       |--S. hitchcocki D07
       `--S. keelinensis D07
  0--Gentilicamelus Loomis 1936 [=Gomphotherium Cope 1886 nec Burmeister 1837 nec Filhol 1884] M66
  |    `--*G. sternbergi (Cope 1879) (see below for synonymy) M66
  |--Paratylopus M66
  |    |--P. matthewi (Lull 1921) [=Paralabis matthewi] M66
  |    `--P. primaevus (Matthew 1904) M66
  `--+--Oxydactylus M66
     |    |  i. s.: *O. longipes Peterson 1904 M66
     |    |         O. brachyodontus M66
     |    |         O. exilis M66
     |    |         O. lagota M66
     |    |         O. wyomingensis (Loomis 1936) [=Gentilicamelus wyomingensis] M66
     |    |--O. cedrensis (Matthew 1901) P98, M66 (see below for synonymy)
     |    `--+--O. campestris M66
     |       `--+--O. cameloides (Wortman 1898) [=Paratylopus cameloides] M66
     |          `--O. wortmani (Lull 1921) [=Paratylopus wortmani] M66
     `--+--Miotylopus Schlaikjer 1935 M66
        |    |--*M. brachygnathus Schlaikjer 1935 M66 [=M. bathygnathus P98]
        |    |--M. gibbi (Loomis 1911) [=Oxydactylus gibbi] M66
        |    |--M. leonardi (Loomis 1911) [=Protomeryx leonardi, Gentilicamelus leonardi] P98
        |    `--M. taylori Prothero 1996 P98
        `--+--Nothokemas White 1947 [Nothokemadidae] M66
           |    `--N. floridanus (Simpson 1932) (see below for synonymy) M66
           `--Floridatragulus White 1940 [incl. Hypermekops White 1942] M66
                |--F. barbouri White 1947 M66
                `--F. dolichanthereus White 1940 [incl. Hypermekops olseni White 1942] M66
  Dyseotylopus Stock 1935 M66
    `--D. migrans Stock 1935 M66, P98

*Gentilicamelus sternbergi (Cope 1879) [=*Gomphotherium sternbergi, Miolabis (Paratylopus) sternbergi, Protomeryx sternbergi] M66

Nothokemas floridanus (Simpson 1932) [=Oxydactylus floridanus; incl. Paratylopus grandis White 1940] M66

Oxydactylus cedrensis (Matthew 1901) P98, M66 [=Protomeryx cedrensis M66; incl. P. campester Matthew 1901 non O. campestris M66]

*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.

[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.

[F84] Franklin, W. L. 1984. Camels and llamas. In: Macdonald, D. (ed.) All the World’s Animals: Hoofed Mammals pp. 72–75. Torstar Books: New York.

[G78] Gentry, A. W. 1978. Tragulidae and Camelidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 536–539. Harvard University Press: Cambridge (Massachusetts).

[GC-BG04] Gentry, A., J. Clutton-Brock & C. P. Groves. 2004. The naming of wild animal species and their domestic derivatives. Journal of Archaeological Science 31: 645–651.

[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.

[M66] McKenna, M. C. 1966. Synopsis of Whitneyan and Arikareean camelid phylogeny. American Museum Novitates 2253: 1–11.

[P98] Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

Basilosauridae

Mounted skeleton of Dorudon atrox in the Senckenberg Museum in Frankfurt am Main, copyright Eva Kröcher.

Belongs within: Cetaceamorpha.
Contains: Mysticeti, Odontoceti.

The Basilosauridae are a group of stem-cetaceans known from the upper Eocene (Barnes et al. 1985). This group is probably paraphyletic to the crown Cetacea.

Basilosauridae [Dorudontidae, Hydrarchidae, Prozeuglodontidae, Stegorhinidae, Zeuglodontidae]
    |  i. s.: Pontogeneus Leidy 1852 G03
    |           `--P. brachyspondylus (Müller 1849) [=Zeuglodon brachyspondylus] BM78
    |         Ancalecetus Gingerich & Uhen 1996 G03
    |         Basiloterus Gingerich et al. 1997 G03
    |         Chrysocetus Uhen & Gingerich 2001 G03
    |         Zeuglodon cetoides A05
    |--+--Basilosaurinae [Zeuglodontinae] BDR85
    |  |    |--Platyosphys Kellogg 1936 BM78, WS76
    |  |    |    `--P. paulsonii (Brandt 1873) BDR85
    |  |    |--Prozeuglodon Andrews 1906 BM78
    |  |    |    `--*P. atrox Andrews 1906 G92 [incl. Zeuglodon intermedius Dart 1923 G92, Dorudon intermedius BM78]
    |  |    `--Basilosaurus Harlan 1843 SOG09, D07
    |  |         |--B. cetoides (Harlan 1845) SB02
    |  |         |--B. hussaini D07
    |  |         `--B. isis (Beadnell in Andrews 1904) G92 (see below for synonymy)
    |  `--Cetacea [Autoceta] SOG09
    |       |--Mysticeti SOG09
    |       `--Odontoceti SOG09
    `--Dorudontinae BDR85
         |--Zygorhiza True 1908 BM78, G03
         |    `--Z. kochii (Reichenbach 1847) SB02
         |--Saghacetus Gingerich 1992 G92
         |    `--*S. osiris (Dames 1894) G92 (see below for synonymy)
         `--Dorudon Gibbes 1845 SOG09, G92 [=Durodon (l. c.) G92; incl. Doryodon Cope 1868 G92]
              |--*D. serratus Gibbes 1845 G92
              |--D. atrox G03
              `--D. stromeri Kellogg 1936 SB02 [=Prozeuglodon stromeri G92]

Basilosaurus isis (Beadnell in Andrews 1904) G92 [=Zeuglodon isis G92, Protocetus isis BM78, Prozeuglodon isis BM78]

*Saghacetus osiris (Dames 1894) G92 [=Zeuglodon osiris BM78, Z. (Dorudon) osiris BM78; incl. Z. elliotsmithii Dart 1923 G92, Dorudon elliotsmithii BM78, Z. sensitivus Dart 1923 G92, D. sensitivus BM78, Z. zitteli Stromer 1903 G92, D. zitteli BM78, Protocetus zitteli BM78]

*Type species of generic name indicated

REFERENCES

[A05] Ameghino, F. 1905. Les édentés fossiles de France et d'Allemagne. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 175–250.

[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.

[BM78] Barnes, L. G., & E. Mitchell. 1978. Cetacea. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 582–602. Harvard University Press: Cambridge (Massachusetts).

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

[G92] Gingerich, P. D. 1992. Marine mammals (Cetacea and Sirenia) from the Eocene of Gebel Mokattam and Fayum, Egypt: stratigraphy, age, and paleoenvironments. University of Michigan Papers on Paleontology 30: i–ix, 1–84.

[G03] Gingerich, P. D. 2003. Land-to-sea transition in early whales: evolution of Eocene Archaeoceti (Cetacea) in relation to skeletal proportions and locomotion of living semiaquatic mammals. Paleobiology 29 (3): 429–454.

[SB02] Sanders, A. E., & L. G. Barnes. 2002. Paleontology of the Late Oligocene Ashley and Chandler Bridge Formations of South Carolina, 3: Eomysticetidae, a new family of primitive mysticetes (Mammalia: Cetacea). Smithsonian Contributions to Paleobiology 93: 313–356.

[SOG09] Spaulding, M., M. A. O'Leary & J. Gatesy. 2009. Relationships of Cetacea (Artiodactyla) among mammals: increased taxon sampling alters interpretation of key fossils and character evolution. PLoS One 4 (9): e7062.

[WS76] Whitmore, F. C., Jr & A. E. Sanders. 1976. Review of the Oligocene Cetacea. Systematic Zoology 25 (4): 304–320.

Suinae

Central European boar Sus scrofa scrofa, copyright Valentin Panzirsch.

Belongs within: Suina.

The Suinae are a group of pigs including all living members of the family. They are united by elongation of the skull, complication of the cheek teeth, and the development of canine tusks (Cooke & Wilkinson 1978).

Suinae
    |  i. s.: Microstonyx Pilgrim 1926 CW78
    |           `--M. major DW04
    |         Dicoryphochoerus CW78
    |           |--D. haydeni CW78
    |           |--D. robustus CW78
    |           `--D. titan CW78
    |--Babyrousa FS15
    |    |--B. togeanensis FS15
    |    `--+--B. babyrussa FS15
    |       `--B. celebensis FS15
    `--+--+--Porcula salvania FS15
       |  `--Sus Linnaeus 1758 FS15, CW78
       |       |  i. s.: S. babyrussa Linnaeus 1758 L58
       |       |         S. heureni IT07
       |       |         S. minor Depéret 1890 P04
       |       |         S. porcus Linnaeus 1758 L58
       |       |         S. riukiuanus Kuroda 1924 I92
       |       |         S. salvanius IT07
       |       |         S. strozzii ME05
       |       |         S. taivanus S66
       |       |         S. timoriensis IT07
       |       |         S. vittatus M89
       |       |--*S. scrofa Linnaeus 1758 CW78, FS15, CW78 (see below for synonymy)
       |       |    |--S. s. scrofa K92
       |       |    |--S. s. domesticus Erxleben 1777 K92, GC-BG04
       |       |    `--S. s. leucomystax Temminck 1842 M76
       |       `--+--+--+--S. bucculentus FS15
       |          |  |  `--S. celebensis FS15
       |          |  `--+--S. cebifrons FS15
       |          |     `--S. philippensis FS15
       |          `--+--+--S. ahoenobarbus FS15
       |             |  `--S. barbatus FS15
       |             `--+--S. oliveri FS15
       |                `--+--S. brachgnathus FS15
       |                   `--S. verrucosus FS15
       `--+--Potamochoerus Gray 1854 FS15, CW78
          |    |--*P. porcus (Linnaeus 1758) CW78
          |    |--P. larvatus FS15
          |    `--P. penicillatus B66
          `--+--Kolpochoerus van Hoepen & van Hoepen 1932 FS15, CW78 (see below for synonymy)
             |    |  i. s.: ‘Hylochoerus’ grabhami Hopwood 1929 (n. d.) [=Mesochoerus grabhami] CW78
             |    |         K. major (Hopwood 1934) (n. d.) FS15, CW78 (see below for synonymy)
             |    |         K. phacochoeroides (Thomas 1884) [=Sus phacochoeroides] CW78
             |    |--K. afarensis Cooke 1978 CW78
             |    `--+--K. limnetes (Hopwood 1926) (see below for synonymy) CW78
             |       |--+--K. paiceae (Broom 1931) (see below for synonymy) CW78
             |       |  `--K. olduvaiensis (Leakey 1942) CW78
             |       `--Hylochoerus Thomas 1904 CW78
             |            |--*H. meinertzhageni Thomas 1904 CW78
             |            `--H. antiquus Leakey 1958 CW78
             `--+--Potamochoeroides Dale 1948 CW78
                |    `--P. shawi (Dale 1948) [=Pronotochoerus shawi; incl. *P. hypsodon Dale 1948] CW78
                `--+--Phacochoerus Cuvier 1817 (see below for synonymy) CW78
                   |    |  i. s.: P. kabuae Whitworth 1965 CW78
                   |    |--P. modestus (van Hoepen & van Hoepen 1932) (see below for synonymy) CW78
                   |    `--+--*P. aethiopicus (Pallas 1767) (see below for synonymy) CW78
                   |       `--P. africanus (Gmelin 1788) (see below for synonymy) CW78
                   |            |--P. a. africanus CW78
                   |            `--P. a. helmei Dreyer & Lyle 1931 (see below for synonymy) CW78
                   `--Metridiochoerus Hopwood 1926 FS15, CW78 [incl. Pronotochoerus Leakey 1943 CW78]
                        |  i. s.: M. compactus (Leakey 1958) D07, CW78 [=Notochoerus compactus CW78]
                        |         M. modestus D07
                        |         ‘Notochoerus’ serengetensis Dietrich 1942 CW78
                        |--M. jacksoni (Leakey 1943) [=*Pronotochoerus jacksoni; incl. M. pygmaeus Leakey 1958] CW78
                        |--M. nyanzae (Leakey 1958) [=Pronotochoerus nyanzae] CW78
                        `--+--*M. andrewsi Hopwood 1926 (see below for synonymy) CW78
                           `--Stylochoerus van Hoepen & van Hoepen 1932 (see below for synonymy) CW78
                                `--S. compactus van Hoepen & van Hoepen 1932 (see below for synonymy) CW78

Inorganic: Babirousa babyrussa minilorientalis Okamura 1987 O87
           Sus scrofa minilorientalis Okamura 1987 O87

Kolpochoerus van Hoepen & van Hoepen 1932 FS15, CW78 [incl. Ectopotamochoerus Leakey 1965 CW78, Mesochoerus Shaw & Cooke 1941 CW78, Omochoerus Arambourg 1942 CW78, Promesochoerus Leakey 1965 CW78]

Kolpochoerus limnetes (Hopwood 1926) [=Sus limnetes, Mesochoerus limnetes; incl. *Ectopotamochoerus dubius Leakey 1965, M. heseloni Leakey 1943, Potamochoerus intermedius Leakey 1965, Omochoerus maroccanus Ennouchi 1954, *Promesochoerus mukiri Leakey 1965, *O. pachygnathus Arambourg 1943, Sus waylandi] CW78

Kolpochoerus major (Hopwood 1934) (n. d.) FS15, CW78 [=Koiropotamus majus CW78, Potamochoerus majus CW78]

Kolpochoerus paiceae (Broom 1931) [=Notochoerus paiceae; incl. Mesochoerus lategani Singer & Keen 1955, *Kolpochoerus sinuosus van Hoepen & van Hoepen 1932] CW78

*Metridiochoerus andrewsi Hopwood 1926 [incl. Notochoerus dietrichi Hopwood 1934, N. hopwoodi Leakey 1958, N. meadowsi Broom 1928, Kolpochoerus meadowsi, Phacochoerus meadowsi, Tapinochoerus meadowsi] CW78

Phacochoerus Cuvier 1817 [incl. Potamochoerops Ewer 1956, Tapinochoerus van Hoepen & van Hoepen 1932; Phacochoerinae] CW78

*Phacochoerus aethiopicus (Pallas 1767) [=Aper aethiopicus; incl. P. dreyeri, P. meiringi, P. venteri Dreyer & Lyle 1931] CW78

Phacochoerus africanus (Gmelin 1788) [=Sus africanus; incl. P. barbarus Pomel 1897, P. congolensis van Straelen 1924, P. mauritanicus Pomel 1897, P. stenobunus Pia 1930] CW78

Phacochoerus africanus helmei Dreyer & Lyle 1931 [incl. P. laticolumnatus van Hoepen & van Hoepen 1932] CW78

Phacochoerus modestus (van Hoepen & van Hoepen 1932) [=*Tapinochoerus modestus; incl. Ph. antiquus Broom 1948, *Potamochoerops antiquus, Notochoerus broomi Shaw & Cooke 1941, Tapinochoerus minutus Leakey 1958] CW78

Stylochoerus van Hoepen & van Hoepen 1932 [incl. Afrochoerus Leakey 1942, Orthostonyx Leakey 1958] CW78

Stylochoerus compactus van Hoepen & van Hoepen 1932 [incl. Phacochoerus altidens, Orthostonyx brachyops Leakey 1958, P. complectidens, Synaptochoerus hieroglyphicus, *Afrochoerus nicoli Leakey 1942] CW78

*Sus scrofa Linnaeus 1758 CW78, FS15, CW78 [incl. S. algeriensis Pomel 1896 CW78, S. barbarus Pomel 1896 (preoc.) CW78]

*Type species of generic name indicated

REFERENCES

[B66] Blyth, E. 1866. A note on African buffalos. Proceedings of the Zoological Society of London 1866: 371–373.

[CW78] Cooke, H. B. S., & A. F. Wilkinson. 1978. Suidae and Tayassuidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 435–482. 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.

[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.

[GC-BG04] Gentry, A., J. Clutton-Brock & C. P. Groves. 2004. The naming of wild animal species and their domestic derivatives. Journal of Archaeological Science 31: 645–651.

[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.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[K92] Krylov, M. V. 1992. The origin of heteroxeny in Sporozoa. Parazitologiya 26 (5): 361–368.

[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.

[M76] Masui, M. 1976. Nihon no Doobutsu. Kogakukan: Tokyo.

[M89] Modigliani, E. 1889. Appunti intorno ai mammiferi dell’isola Nias. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 238–245.

[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.

[P04] Popov, V. V. 2004. Pliocene small mammals (Mammalia, Lipotyphla, Chiroptera, Lagomorpha, Rodentia) from Muselievo (north Bulgaria). Geodiversitas 26 (3): 403–491.

[S66] Sclater, P. L. 1866. Notice of recent additions to the Society’s menagerie. Proceedings of the Zoological Society of London 1866: 418–419.

Ruminantiamorpha

Reconstruction of Brachyodus aequatorialis, copyright WillemSvdMerwe.

Belongs within: Artiodactyla.
Contains: Protoceratidae, Tragulidae, Pecora.

The Ruminantiamorpha are a clade of artiodactyls including all species closer to modern ruminants than to other living artiodactyls. Living ruminants are characterised by the possession of a multi-chambered stomach in which food is broken down by bacteria living in a large fermentation chamber, the rumen.

See also: Horns and guts.

Synapomorphies (from Janis & Scott 1988): Tarsus with fused cuboid and navicular; upper incisors reduced or lost; incisiform lower canines.

<==Ruminantiamorpha SOG09
    |  i. s.: Bothriogenys Schmidt 1913 SOG09, B78
    |           |--*B. fraasi (Schmidt 1913) [=Brachyodus (*Bothriogenys) fraasi] B78
    |           |--B. africanus (Andrews 1899) B78
    |           |--B. andrewsi Schmidt 1913 B78
    |           |--B. gorringei (Andrews & Beadnell 1902) B78
    |           |--B. parvus (Andrews 1906) B78
    |           `--B. rugulosus Schmidt 1913 B78
    |--+--Anthracotherium BLB05
    |  `--+--Brachyodus BLB05
    |     |    |--B. aequatorialis MacInnes 1951 BLB05, B78 [=Masritherium aequitorialis B78]
    |     |    |--B. andrewsi Schmidt 1913 S68
    |     |    |--B. gorringei (Andrews & Beadnell 1902) S68
    |     |    |--B. parvulus (Andrews 1906) S68
    |     |    `--B. rugulosus Schmidt 1913 S68
    |     `--Elomeryx Marsh 1894 SOG09, D07
    |          |--E. armatus BLB05
    |          |--E. crispus OB13
    |          `--E. woodi D07
    `--+--+--Libycosaurus petrocchii Bonarelli 1947 SOG09, B78 [=Merycopotamus petrocchii B78]
       |  `--+--Leptoreodon TC07
       |     `--+--Heteromeryx TC07
       |        `--Protoceratidae SOG09
       `--Ruminantia [Tragulina] SOG09
            |  i. s.: Russa equina M89
            |         Cervulus M89
            |           |--C. crinifrons TD89
            |           |--C. feae Thomas & Doria 1889 TD89
            |           |--C. moschatus TD89
            |           |--C. muntjac M89
            |           |--C. reevesii D59
            |           `--C. vaginalis D59
            |         Indomeryx JS88
            |         Pseudoceras JS88
            |         Pseudomeryx JS88
            |         Archaeomeryx Matthew & Granger 1925 D07 [Archaeomerycidae HD03]
            |           `--A. optatus D07
            |         Hexameryx White 1941 D07
            |           |--H. elmori D07
            |           `--H. simpsoni D07
            |--Tragulidae JS88
            `--+--Bachitherium [Bachitheriidae] JS88
               `--+--Pecora JS88
                  `--Lophiomerycidae JS88
                       |--Lophiomeryx JS88
                       |--Cryptomeryx JS88
                       `--Iberomeryx JS88

*Type species of generic name indicated

REFERENCES

[B78] Black, C. C. 1978. Anthracotheriidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 423–434. Harvard University Press: Cambridge (Massachusetts).

[BLB05] Boisserie, J.-R., F. Lihoreau & M. Brunet. 2005. The position of Hippopotamidae within Cetartiodactyla. Proceedings of the National Academy of Sciences of the USA 102 (5): 1537–1541.

[D59] Darwin, C. 1859. The Origin of Species by Means of Natural Selection, or the preservation of favoured races in the struggle for life 1^st ed. John Murray: London.

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

[HD03] Hassanin, A., & E. J. P. Douzery. 2003. Molecular and morphological phylogenies of Ruminantia and the alternative position of the Moschidae. Systematic Biology 52 (2): 206–228.

[JS88] Janis, C. M., & K. M. Scott. 1988. The phylogeny of the Ruminantia (Artiodactyla, Mammalia). In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 273–282. Clarendon Press: Oxford.

[M89] Modigliani, E. 1889. Appunti intorno ai mammiferi dell’isola Nias. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 238–245.

[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.

[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.

[SOG09] Spaulding, M., M. A. O'Leary & J. Gatesy. 2009. Relationships of Cetacea (Artiodactyla) among mammals: increased taxon sampling alters interpretation of key fossils and character evolution. PLoS One 4 (9): e7062.

[TC07] Thewissen, J. G. M., L. N. Cooper, M. T. Clementz, S. Bajpai & B. N. Tiwari. 2007. Whales originated from aquatic artiodactyls in the Eocene epoch of India. Nature 450: 1190–1194.

[TD89] Thomas, O., & G. Doria. 1889. Diagnosi di una nuova specie del genere Cervulus raccolta da L. Fea nel Tenasserim. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 92.

Artiodactyla

Mounted skeleton of Cainotherium at the Natural History Museum, Basel, copyright Ghedoghedo.

Belongs within: Euungulata.
Contains: Choeropotamoidea, Anoplotherioidea, Oreodonta, Camelidae, Suina, Ruminantiamorpha, Cetancodontamorpha.

The Artiodactyla are a clade of mammals containing the even-toed hoofed mammals, as well as the cetaceans which are nested well within this group. Most recent studies place the cetaceans closest to the Hippopotamidae among living artiodactyls, with the two in turn often sister to the Ruminantia. Previously, the ruminants had been thought more closely related to the Camelidae due to shared characters such as stomach divided into three or four chambers and rear teeth with crescent-shaped cusps. However, modern molecular analyses mostly place camelids as the sister group to all other living artiodactyls.

The Artiodactyla have an extensive fossil record going back to the Eocene, but the affinities of many basal taxa are currently uncertain in the light of the aforementioned changes in our understanding of artiodactyl phylogeny. The Cainotheriidae are small, rabbit-sized animals known from the late Eocene to the middle Miocene of Europe that may be distantly related to the camels. The Dichobuninae and Hyperdichoduninae are other groups of small artiodactyls (or possibly stem-artiodactyls) from the Eocene of Europe. The Dichobuninae have a bulbous dentition and moderately elongated snouts. The Hyperdichobuninae have molarised, crescentiform premolars and molars with a tendency to broaden the distal side of the latter.

See also: Where do you put your camels?

Synapomorphies (from Spaulding et al. 2009, for Artiodactylamorpha): Sustentacular facet wide; contact of distal astragalus with cuboid present; ectal facet primary orientation lateral; astragalus with lateral edges of proximal and distal trochlea aligned; astragalar head arc to dorsoplantar wide, ~200 degrees; astragalar neck as wide as tibial trochlea.

Artiodactyla (see below for synonymy) SOG09
    |--Camelidamorpha SOG09
    |    |--Oreodonta SOG09
    |    `--+--+--Camelidae TC07
    |       |  |--Oromerycidae GH88
    |       |  |--Eotylops TC07
    |       |  `--Poebrotherium Leidy 1848 SOG09, D07
    |       |       |--P. labiatum HUG17
    |       |       `--P. wilsoni D07
    |       `--Cainotheriidae GH88
    |            |--Robiacina GH88
    |            `--Cainotherium Bravard 1835 SOG09, D07
    |                 |--C. commune D07
    |                 |--C. geoffroyi D07
    |                 `--C. laticurvatum D07
    `--+--Suina SOG09
       `--Cetruminantia [Cetruminantiamorpha] SOG09
            |--Ruminantiamorpha SOG09
            `--Cetancodontamorpha SOG09

Artiodactyla incertae sedis:
  Heptacodon GH01
  Siamotragulus CJ03
  Merychyus Leidy 1858 C77
    |--M. elegans C77
    |--M. major C77
    `--M. medius Leidy 1858 C77
  Wasatchia GH88
  Chorlakkia GH88
  Lantianius GH88
  Texodon GH88
  Bunomerycidae GH88
    |--Bunomeryx Wortman 1898 TC07, GH88
    |--Hylomeryx Peterson 1919 GH88
    |--Mytonomeryx Gazin 1955 GH88
    `--Pentacemylus Peterson 1931 GH88
  Arretotherium MS56
    |--A. acridens Douglass 1901 MS56
    `--A. fricki Macdonald & Schultz 1956 MS56
  Masritherium Fourtau 1918 B78
    `--*M. depereti Fourtau 1918 B78
  Hyoboops Trouessart 1904 B78
    |--*H. palaeindicus (Lydekker 1883) B78
    |--H. africanus (Andrews 1914) [=Merycops africanus] B78
    `--H. moneyi (Fourtau 1918) [=Brachyodus moneyi] B78
  Anthracothema S68
  Choeropotamoidea BLB05
  Gervachoerus TW01
  Aumelasia Sudre 1980 SL00
  Protodichobune Lemoine 1878 SL00
    `--*P. oweni Lemoine 1878 SL00
  Messelobunodon GH88
  Dichobuninae GH88
    |--Dichobune Cuvier 1822 GH88
    |--Metriotherium Filhol 1882 GH88
    `--Meniscodon Rütimeyer 1888 GH88
         `--M. europaeus (Ruetimeyer 1888) [=Phenacodus europaeus] T90
  Sarcolemur Cope 1875 C77 [incl. Antiacodon Marsh 1872 (n. n.) GH88, C77]
    |--*S. furcatus C77
    |--S. crassus (Cope 1875) [=Antiacodon crassus] C77
    |--S. mentalis (Cope 1875) [=Antiacodon mentalis] C77
    `--S. pygmaeus C77
  Hyperdichobuninae GH88
    |--Hyperdichobune Stehlin 1910 GH88
    `--Mouillacitherium Filhol 1882 GH88
  Brachycrus Merriam 1919 D07
    |--B. buwaldi W69
    |--B. laticeps D07
    `--B. siouense D07
  Protylopus Wortman 1898 D07
    `--P. petersoni D07
  Aepycamelus Leidy 1886 D07
    |--A. alexandrae D07
    |--A. bradyi D07
    |--A. elrodi D07
    |--A. giraffinus D07
    |--A. major D07
    |--A. priscus D07
    |--A. proceras D07
    |--A. robustus D07
    `--A. stocki D07
  Titanotylopus Barbour & Schultz 1934 D07
    |--T. nebraskensis D07
    `--T. spatulus D07
  Leptotragulidae GH88
  Amphimeryx [Amphimerycidae] TC07
  Anoplotherioidea TC07
  Anthracokeryx ulnifer SOG09
  Microbunodon minimum BLB05
  Gobiohyus SOG09
    |--G. orientalis Matthew & Granger 1925 GH88, SL00
    |--G. pressidens HUG17
    `--G. robustus HUG17
  ‘Proviverra’ americanus Scott 1892 V65

Artiodactyla [Ancodonta, Anthracotheriidae, Anthracotherioidea, Bunodontia, Cameloidea, Cetartiodactyla, Dichobunidae, Dichobunoidea, Helohyidae, Merycotheria, Neoselenodontia, Palaeodonta, Selenodontia, Suiformes, Tylopoda, Whippomorpha] SOG09

*Type species of generic name indicated

REFERENCES

[B78] Black, C. C. 1978. Anthracotheriidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 423–434. Harvard University Press: Cambridge (Massachusetts).

[BLB05] Boisserie, J.-R., F. Lihoreau & M. Brunet. 2005. The position of Hippopotamidae within Cetartiodactyla. Proceedings of the National Academy of Sciences of the USA 102 (5): 1537–1541.

[CJ03] Chaimanee, Y., D. Jolly, M. Benammi, P. Tafforeau, D. Duzer, I. Moussa & J.-J. Jaeger. 2003. A Middle Miocene hominoid from Thailand and orangutan origins. Nature 422: 61–65.

[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.

[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.

[GH01] Gingerich, P. D., M. ul Haq, I. S. Zalmout, I. H. Khan & M. S. Malkani. 2001. Origin of whales from early artiodactyls: hands and feet of Eocene Protocetidae from Pakistan. Science 293: 2239–2242.

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

[MS56] Macdonald, J. R., & C. B. Schultz. 1956. Arretotherium fricki, a new Miocene anthracothere from Nebraska. Bulletin of the University of Nebraska State Museum 4 (3): 53–67.

[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.

[SOG09] Spaulding, M., M. A. O'Leary & J. Gatesy. 2009. Relationships of Cetacea (Artiodactyla) among mammals: increased taxon sampling alters interpretation of key fossils and character evolution. PLoS One 4 (9): e7062.

[SL00] Sudre, J., & G. Lecomte. 2000. Relations et position systématique du genre Cuisitherium Sudre et al., 1983, le plus dérivé des artiodactyles de l’Éocène inféreur d’Europe. Geodiversitas 22 (3): 415–432.

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

[TC07] Thewissen, J. G. M., L. N. Cooper, M. T. Clementz, S. Bajpai & B. N. Tiwari. 2007. Whales originated from aquatic artiodactyls in the Eocene epoch of India. Nature 450: 1190–1194.

[TW01] Thewissen, J. G. M., E. M. Williams, L. J. Roe & S. T. Hussain. 2001. Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls. Nature 413: 277–281.

[V65] Van Valen, L. 1965. Some European Proviverrini (Mammalia, Deltatheridia). Palaeontology 8 (4): 638–665.

[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.

Last updated: 31 July 2018.