Manidae

Manidae Temporal range: 41.37–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N middle Eocene - present[1]
Living species of pangolins
Conservation status
CITES Appendix I [3]
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Pholidotamorpha
Order:	Pholidota
Suborder:	Eupholidota
Superfamily:	Manoidea
Family:	Manidae Gray, 1821[2]
Type genus
Manis Linnaeus, 1758
Genera
[see classification]
Species ranges:   Manis crassicaudata   Manis pentadactyla   Manis javanica   Manis culionensis   Phataginus tricuspis   Phataginus tetradactyla   Smutsia gigantea   Smutsia temminckii
Synonyms
list of synonyms: Manida (Kalandadze & Rautian, 1992) Manididae (Gray, 1865) Manina (Gray, 1825)[4] Manisia (Rafinesque, 1815) Perilepia (Rafinesque, 1815)

Manidae ("spirits") is the only extant family of pangolins.^[5] This family comprises three genera (Manis from subfamily Maninae, Phataginus from subfamily Phatagininae, and Smutsia from subfamily Smutsiinae),^[6][7] as well as the extinct Fayum pangolin.^[8]

Classification and phylogeny

History of classification

All species of living pangolin had been assigned to the genus Manis until the late 2000s, when research prompted the splitting of extant pangolins into three genera: Manis, Phataginus, and Smutsia.^[5][9]

Taxonomy

Former classification (McKenna & Bell, 1997):[10]

Current classification:

Family: Manidae (Gray, 1821) (pangolins) Subfamily: Maninae (Gray, 1821) (Asian pangolins) Genus: Manis (Linnaeus, 1758) (Asian pangolin) †Manis hungarica (Kormos, 1934) (Hungarian pangolin) †Manis lydekkeri (Dubois, 1908) (Lydekker's pangolin) †Manis palaeojavanica (Dubois, 1907) (giant Asian pangolin) Subgenus: Manis (Linnaeus, 1758) Manis crassicaudata (Gray, 1827) (Indian pangolin) Manis pentadactyla (Linnaeus, 1758) (Chinese pangolin) Subgenus: Paramanis (Pocock, 1924) (Southeast Asian pangolin) Manis javanica (Desmarest, 1822) (Sunda pangolin) Manis culionensis (de Elera, 1895) (Philippine pangolin) Genus: †Eomanis (Storch, 1978) †Eomanis krebsi (Storch & Martin, 1994) †Eomanis waldi (Storch, 1978) Genus: †Necromanis (Filhol, 1894) †Necromanis franconica (Quenstedt, 1885) †Necromanis parva (Koenigswald, 1969) †Necromanis quercyi (Filhol, 1894) Genus: †Patriomanis (Emry, 1970) †Patriomanis americana (Emry, 1970) Subfamily: Smutsiinae (Gray, 1873) (African pangolins) Tribe: Smutsiini (Gray, 1873) (large African pangolins) Genus: Smutsia (Gray, 1865) (African ground pangolin) Smutsia gigantea (Illiger, 1815) (giant pangolin) Smutsia temmincki (Smuts, 1832) (ground pangolin) Tribe: Uromanini (McKenna & Bell, 1997) (small African pangolins) Genus: Phataginus (Rafinesque, 1821) Phataginus tricuspis (Rafinesque, 1821) (white-bellied pangolin) Genus: Uromanis (Pocock, 1924) Phataginus tetradactyla (Linnaeus, 1766) (black-bellied pangolin) Genus: †Eurotamandua (Storch, 1981) †Eurotamandua joresi (Storch, 1981) Incertae sedis: †Manidae sp. [DPC 3972 & DPC 4364] (Gebo & Rasmussen, 1985) (Fayum pangolin)

Family: Manidae (Gray, 1821) (pangolins) Subfamily: Maninae (Gray, 1821) (Asian pangolins) Genus: Manis (Linnaeus, 1758) (Asian pangolin) †Manis hungarica (Kormos, 1934) (Hungarian pangolin) (unranked): northern Asian clade Manis indoburmanica (Wangmo, 2025) (Indo Burmese pangolin) Manis pentadactyla (Linnaeus, 1758) (Chinese pangolin) (unranked): southern Asian clade Manis crassicaudata (Gray, 1827) (Indian pangolin) Manis mysteria (Tong-Tong, 2023) (cryptic pangolin) †Manis lydekkeri (Dubois, 1908) (Lydekker's pangolin) Subgenus: Paramanis (Pocock, 1924) (Southeast Asian pangolin) Manis javanica (Desmarest, 1822) (Sunda pangolin) Manis culionensis (de Elera, 1895) (Philippine pangolin) †Manis palaeojavanica (Dubois, 1907) (giant Asian pangolin) (unranked): African clade Subfamily: Phatagininae (Gaubert, 2017) (small African pangolins) Genus: Phataginus (Rafinesque, 1821) (African tree pangolin) Phataginus tetradactyla (Linnaeus, 1766) (black-bellied pangolin) Phataginus tricuspis (Rafinesque, 1821) (white-bellied pangolin) Subfamily: Smutsiinae (Gray, 1873) (large African pangolins) Genus: Smutsia (Gray, 1865) (African ground pangolin) Smutsia gigantea (Illiger, 1815) (giant pangolin) Smutsia temmincki (Smuts, 1832) (ground pangolin) †Smutsia olteniensis (Terhune, 2021) (Oltenian pangolin) Incertae sedis †Manidae sp. [DPC 3972 & DPC 4364] (Gebo & Rasmussen, 1985) (Fayum pangolin)

Phylogeny

Phylogenetic position of family Manidae within order Pholidota based on Kondrashov & Agadjanian (2012.) study:^[9]

	Eulipotyphla  Erinaceus  Carnivora  Nandinia  Pholidotamorpha  †Palaeanodonta  Pholidota  †Euromanis †Afredentata  Eupholidota  †Eomanoidea  Manoidea  †Patriomanidae ? †Necromanis Manidae

Phylogeny within family Manidae based on Wangmo (2025.) study:^[11]

Manidae	Maninae   Manis   (Paramanis)  Manis culionensis Manis javanica Manis mysteria Manis crassicaudata Manis indoburmanica Manis pentadactyla  Phatagininae   Phataginus  Phataginus tetradactyla Phataginus tricuspis  Smutsiinae   Smutsia  Smutsia gigantea Smutsia temminckii

See also

• Mammal classification
• Manoidea

References

1. Sean P. Heighton, Rémi Allio, Jérôme Murienne, Jordi Salmona, Hao Meng, Céline Scornavacca, Armanda D. S. Bastos, Flobert Njiokou, Darren W. Pietersen, Marie-Ka Tilak, Shu-Jin Luo, Frédéric Delsuc, Philippe Gaubert (2023.) "Pangolin genomes offer key insights and resources for the world's most trafficked wild mammals"
2. J. E. Gray. (1821.) "On the natural arrangement of vertebrose animals." The London Medical Repository Monthly Journal and Review 15:296-310
3. "The CITES Appendices". Convention on International Trade in Endangered Species of Wild Fauna and Flora. CITES. Retrieved 28 January 2019.
4. J. E. Gray. (1825.) "An outline of an attempt at the disposition of Mammalia into Tribes and Families, with a list of genera apparently appertaining to each Tribe." Annals of Philosophy, new series 10:337-344
5. Gaudin, Timothy (2009). "The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis" (PDF). Journal of Mammalian Evolution. 16 (4). Heidelberg, Germany: Springer Science+Business Media: 235–305. doi:10.1007/s10914-009-9119-9. S2CID 1773698. Archived from the original (PDF) on 2015-09-25. Retrieved 2021-01-20.
6. Schlitter, Duane A. (2005). Wilson, D.E.; Reeder, D.M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore, Maryland, USA: Johns Hopkins University Press. p. 530. ISBN 978-0-8018-8221-0.
7. Philippe Gaubert; Agostinho Antunes; Hao Meng; Lin Miao; Stéphane Peigné; Fabienne Justy; Flobert Njiokou; Sylvain Dufour; Emmanuel Danquah; Jayanthi Alahakoon; Erik Verheyen (11 May 2018). "The Complete Phylogeny of Pangolins: Scaling Up Resources for the Molecular Tracing of the Most Trafficked Mammals on Earth". Journal of Heredity. 109 (4): 347–359. doi:10.1093/jhered/esx097. PMID 29140441.
8. Daniel Gebo, D. Tab Rasmussen (1985.) "The Earliest Fossil Pangolin (Pholidota: Manidae) from Africa" Journal of Mammalogy 66(3):538
9. Kondrashov, Peter; Agadjanian, Alexandre K. (2012). "A nearly complete skeleton of Ernanodon (Mammalia, Palaeanodonta) from Mongolia: morphofunctional analysis". Journal of Vertebrate Paleontology. 32 (5): 983–1001. doi:10.1080/02724634.2012.694319. ISSN 0272-4634. S2CID 86059673.
10. McKenna, M. C.; Bell, S. K. (1997). Classification of Mammals: Above the Species Level. Columbia University Press. p. 631. ISBN 978-0-231-52853-5. Retrieved 2024-09-16.
11. Wangmo, Lenrik Konchok; Ghosh, Avijit; Dolker, Stanzin; Joshi, Bheem Dutt; Sharma, Lalit Kumar; Thakur, Mukesh (2025-01-07). "Indo-Burmese pangolin (Manis indoburmanica): a novel phylogenetic species of pangolin evolved in Asia". Mammalian Biology. 105 (5): 691–698. doi:10.1007/s42991-024-00475-7. ISSN 1616-5047.

External links

• ZSL Pangolin Conservation
• Pangolin: Wildlife summary from the African Wildlife Foundation
• Tree of Life of Pholidota
• National Geographic video of a pangolin
• Proceedings of the Workshop on Trade and Conservation of Pangolins Native to South and Southeast Asia (PDF)
• The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis Archived 2015-09-25 at the Wayback Machine (PDF)
• Bromley, Victoria (Director/Producer), Young, Nora (Narrator/Host), Diekmann, Maria (2018). Nature: The World's Most Wanted Animal. United States: PBS.
• Coronavirus: Revenge of the Pangolins? The New York Times, March 6, 2020.