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.

    |  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


[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.

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