Manidae

Manidae; Temporal range: 40.5–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Pal. Eocene Oligo. Miocene P P middle Eocene - present
	Living species of pangolins
Conservation status
	CITES Appendix I
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Pholidotamorpha
Order:	Pholidota
Suborder:	Eupholidota
Superfamily:	Manoidea
Family:	Manidae; Gray, 1821
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) ; Manisia (Rafinesque, 1815) ; Perilepia (Rafinesque, 1815) ;

Manidae ("spirits") is the only extant family of pangolins.^[5]^[6]^[7] This family comprises three genera (Manis from subfamily Maninae, Phataginus from subfamily Phatagininae, and Smutsia from subfamily Smutsiinae),^[8]^[9] as well as the extinct Fayum pangolin.^[10]

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]^[6]

Taxonomy

Former classification (McKenna & Bell, 1997):^[11]	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

Cladogram of phylogeny within family Manidae

Position within order Pholidota based on Rose (2026.) study:^[12]

Within family based on Wangmo (2025.) study:^[13]

outhgroup

Pholidotamorpha

	†Afredentata

	†Palaeanodonta

†Euromanis

Pholidota

Eupholidota

†Eomanoidea

Manoidea

	†Patriomanidae

?	†Necromanis

Manidae

Maninae

Manis

Smutsiinae

Phatagininae	Phataginus

Smutsiinae	Smutsia
sensu stricto

sensu lato

sensu stricto

(Pholidota [sensu lato])

outhgroup

Manidae

Maninae

Manis

(Paramanis)

	Manis culionensis

	Manis javanica

Manis mysteria

Manis crassicaudata

	Manis indoburmanica

	Manis pentadactyla

Smutsiinae

Phatagininae

Phataginus

	Phataginus tetradactyla

	Phataginus tricuspis

Smutsiinae

Smutsia

	Smutsia gigantea

	Smutsia temminckii

sensu stricto

sensu lato

southern Asian clade

northern Asian clade

African clade

References

↑ Koju, Narayan Prasad; Zeng, Zeling; Zhang, Guihua; Yao, Zhicheng; Huang, Xia; Wang, Xiaoyun; Hawkins, Melissa T. R.; Hinckley, Arlo; Flores, Mary Faith C.; Guo, Ce; Li, Jun; Maharjan, Devendra; Byanjankar, Saraswoti; Huang, Lianghua; Yu, Wenhua (2026-07-01). "Revalidation of Manis aurita based on integrative genomic and morphological evidence". Communications Biology. 9 (1). doi:10.1038/s42003-026-10314-9. ISSN 2399-3642. PMC 13323723. PMID 42387092.
↑ "The CITES Appendices". Convention on International Trade in Endangered Species of Wild Fauna and Flora. CITES. Retrieved 28 January 2019.
↑ J. E. Gray. (1821.) "On the natural arrangement of vertebrose animals." The London Medical Repository Monthly Journal and Review 15:296-310
↑ 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
1 2 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.
1 2 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.
↑ 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"
↑ 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.
↑ 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.
↑ Daniel Gebo, D. Tab Rasmussen (1985.) "The Earliest Fossil Pangolin (Pholidota: Manidae) from Africa" Journal of Mammalogy 66(3):538
↑ 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.
↑ Rose, K. D.; Rabenstein, R.; Habersetzer, J.; Gaudin, T. J. (2026). "Anatomy of Eurotamandua joresi (Mammalia) from Grube Messel, Germany, based on computed tomography, and implications for its relationships. Part I: Background and forelimb". Palaeobiodiversity and Palaeoenvironments. doi:10.1007/s12549-025-00686-x.
↑ 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.

[Koju2026-1] Koju, Narayan Prasad; Zeng, Zeling; Zhang, Guihua; Yao, Zhicheng; Huang, Xia; Wang, Xiaoyun; Hawkins, Melissa T. R.; Hinckley, Arlo; Flores, Mary Faith C.; Guo, Ce; Li, Jun; Maharjan, Devendra; Byanjankar, Saraswoti; Huang, Lianghua; Yu, Wenhua (2026-07-01). "Revalidation of Manis aurita based on integrative genomic and morphological evidence". Communications Biology. 9 (1). doi:10.1038/s42003-026-10314-9. ISSN 2399-3642. PMC 13323723. PMID 42387092.

[2] "The CITES Appendices". Convention on International Trade in Endangered Species of Wild Fauna and Flora. CITES. Retrieved 28 January 2019.

[3] J. E. Gray. (1821.) "On the natural arrangement of vertebrose animals." The London Medical Repository Monthly Journal and Review 15:296-310

[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

[web2.utc.edu-5] 1 2 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.

[KondrashovAgadjanian2012-6] 1 2 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.

[Heighton2023-7] 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"

[8] 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.

[Gaubert2018-9] 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.

[10] Daniel Gebo, D. Tab Rasmussen (1985.) "The Earliest Fossil Pangolin (Pholidota: Manidae) from Africa" Journal of Mammalogy 66(3):538

[McKenna-11] 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.

[Rose2026-12] Rose, K. D.; Rabenstein, R.; Habersetzer, J.; Gaudin, T. J. (2026). "Anatomy of Eurotamandua joresi (Mammalia) from Grube Messel, Germany, based on computed tomography, and implications for its relationships. Part I: Background and forelimb". Palaeobiodiversity and Palaeoenvironments. doi:10.1007/s12549-025-00686-x.

[Wangmo2025-13] 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.

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Manidae

Classification and phylogeny

History of classification

Taxonomy

Phylogeny

See also

References

External links