Scrotifera

Holotype jaw of Mimatuta makpialutae, copyright Yale Peabody Museum of Natural History.

Belongs within: Boreoeutheria.
Contains: Chiroptera, Ferae, Euungulata.

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); these were herbivorous animals with a dentition superficially similar to that of basal perissodactyls. The genera Pleuraspidotherium and Orthaspidotherium are known from France and Belgium whereas Hilalia is known from Turkey. Affinities with Pleuraspidotheriidae have also been suggested for the North American genera Bubogonia and Protoselene (Muizon & Cifelli 2000); however, Protoselene was placed by Halliday et al. (2017) closer to crown euungulates whereas Bubogonia has not yet been subjected to a formal phylogenetic analysis. Other basal North American genera include Desmatoclaenus, previously suggested to be intermediate in morphology between the Arctocyonidae (placed by Halliday et al. as Ferae) and Phenacodontidae (Euungulata). Mimatuta, one of the earliest condylarths, has previously been included in the Periptychidae but possesses a number of primitive dental similarities to contemporaneous taxa such as Protungulatum and Oxyprimus (Muizon & Cifelli 2000). Todralestes variabilis is an insectivorous mammal from the late Palaeocene of Morocco, previously compared to the Pantolestidae.

Scrotifera (see below for synonymy) HUG17
    |--+--Chiroptera HUG17
    |  `--Ferae HUG17
    `--Pan-Euungulata [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 Gazin 1941 HUG17, T90
            |  |    |--D. dianae HUG17
            |  |    |--D. hermaeus HUG17
            |  |    `--D. protogonoides HUG17
            |  `--+--+--Protoselene opisthacus HUG17, MC00
            |     |  `--Todralestes variabilis HUG17, CW07
            |     `--Mimatuta HUG17
            |          |--M. makpialutae HUG17
            |          |--M. minuial van Valen 1978 MC00
            |          `--M. morgoth HUG17
            `--Euungulata OB13

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

Scrotifera [Acreodi, Amblypoda, Arctocyonoidea, Bulbulodentata, Cete, Cetferungulata, Condylarthra, Fereuungulata, Ferungulata, Hyaenodonta, Mesonychoidea, Paraxonia, Pegasoferae, Triisodontinae, Variamana, Zooamata] HUG17

*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.

[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.

[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.

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

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

Last updated: 14 August 2018.