Partial jaws and dentition of Tetraclaenodon puercensis, from Thewissen (1990).

Belongs within: Boreoeutheria.
Contains: Chiroptera, Ferae, Perissodactyla, Hyopsodontidae, Phenacodontidae, Artiodactyla.

The Scrotifera is a clade supported by molecular phylogenetic analyses uniting the modern mammalian orders Chiroptera, Carnivora, Pholidota, Perissodactyla and Artiodactyla. The majority of recent analyses have supported a division of the Scrotifera into three subclades, with the Carnivora and Pholidota forming one (Ferae) and the Perissodactyla and Artiodactyla forming another (Euungulata), but the exact relationships between Chiroptera, Ferae and Euungulata remain disputed. A number of Palaeogene mammalian groups (commonly classed as 'condylarths') are likely to represent early relatives of the Scrotifera. An analysis of Palaeocene mammals by Halliday et al. (2017) supported relationships of a number of primarily herbivorous 'condylarths' (such as Pleuraspidotheriidae, Hyopsodontidae and Phenacodontidae) with the Euungulata, whereas other, more omnivorous 'condylarths' were placed within the Ferae. The European Pleuraspidotheriidae were placed as the most basal members of the Euungulata total group (Pan-Euungulata). A number of North American Palaeocene genera, such as Tetraclaenodon and Ectocion, that had previously been placed in the Phenacodontidae were suggested to form the stem group of the Perissodactyla.

Scrotifera (see below for synonymy) HUG17
    |--+--Chiroptera HUG17
    |  `--Ferae HUG17
    `--Pan-Euungulata [Bulbulodentata, Procreodi] OB13
         |  i. s.: Baioconodon Gazin 1941 PMF88, MC00 [incl. Ragnarok van Valen 1978 MC00]
         |           |--B. middletoni PV07
         |           `--B. nordicum (van Valen 1978) [=Ragnarok nordicum] MC00
         |--Pleuraspidotheriidae [Pleuraspidotheriinae] HUG17
         |    |  i. s.: Bubogonia MC00
         |    |           |--B. bombadil (van Valen 1978) MC00
         |    |           `--B. saskia Johnston & Fox 1984 MC00
         |    |--+--Pleuraspidotherium aumonieri HUG17
         |    |  `--Orthaspidotherium edwardsi HUG17
         |    `--Hilalia HUG17
         |         |--H. robustus HUG17
         |         |--H. saribeya HUG17
         |         `--H. selanneae HUG17
         `--+--+--Desmatoclaenus HUG17
            |  |    |--D. dianae HUG17
            |  |    |--D. hermaeus HUG17
            |  |    `--D. protogonoides HUG17
            |  `--+--Mimatuta HUG17
            |     |    |--M. makpialutae HUG17
            |     |    |--M. minuial van Valen 1978 MC00
            |     |    `--M. morgoth HUG17
            |     `--+--Protoselene opisthacus HUG17, MC00
            |        `--Todralestes variabilis HUG17, CW07
            `--Euungulata [Bellua, Meniscotheriidae, Meniscotheriinae] OB13
                 |  i. s.: Radinskya F02
                 |--+--Lessnessina HUG17
                 |  |    |--L. khushuensis HUG17
                 |  |    |--L. packmani HUG17
                 |  |    |--L. praecipuus HUG17
                 |  |    `--L. thushnensis HUG17
                 |  `--+--Tetraclaenodon HUG17
                 |     |    |--T. pliciferus S35
                 |     |    |--T. puercensis [=Euprotogonia puercensis] PMF88
                 |     |    |--T. superior Simpson 1935 S35
                 |     |    `--T. symbolicus Gidley in Simpson 1935 S35
                 |     `--+--Copecion brachypternus HUG17, MC00
                 |        `--+--Perissodactyla HUG17
                 |           `--Ectocion HUG17
                 |                |--E. collinus Russell 1929 [incl. Meniscotherium semicingulatum Russell 1929] SFY02
                 |                `--E. osbornianus MC00
                 `--+--+--Hyopsodontidae HUG17
                    |  `--Phenacodontidae HUG17
                    `--Artiodactylamorpha OB13
                         |--+--‘Diacodexis’ metsiacus GH88
                         |  `--Leptochoeridae GH88
                         `--Artiodactyla OB13

Scrotifera incertae sedis:
  Eobasileus pressicornis C77
  Tillodontia V66
    |--Anchippodus C77
    |--Tillotherium C77
    |--Trogosus Leidy 1871 D07
    |    |--T. castoroides D07
    |    |--T. fodiens D07
    |    |--T. hyracoides D07
    |    |--T. latidens D07
    |    `--T. vetulus D07
    `--Chungchienia Chow 1963 S03
         |--C. lushia Chow, Wang & Meng 1996 S03
         `--C. sichuanica Chow 1968 S03
  Microclaenodon assurgens O98

Scrotifera [Acreodi, Amblypoda, Arctocyonoidea, Cete, Cetferungulata, Condylarthra, Fereuungulata, Ferungulata, Hyaenodonta, Paraxonia, Triisodontinae, Variamana, Zoomata] HUG17

*Type species of generic name indicated


[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.

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

[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.

[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.

[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.

[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.

[PV07] Prasad, G. V. R., O. Verma, A. Sahni, V. Parmar & A. Khosla. 2007. A Cretaceous hoofed mammal from India. Science 318: 937.

[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.

[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.

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

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

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