Manidae

Manidae; Temporal range: 37.9–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Late Eocene - present
	Living speacies of pangolins
Conservation status
	CITES Appendix I (CITES)
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
	Subfamily: Maninae Manis; ; Subfamily: Phatagininae Phataginus; ; Subfamily: Smutsiinae Smutsia; ; Incertae sedis †; ;
	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 is the only extant family of pangolins from superfamily Manoidea.^[5] This family comprises three genera (Manis from subfamily Maninae, Phataginus from subfamily Phatagininae, and Smutsia from subfamily Smutsiinae),^[1] as well as extinct Fayum pangolin.^[6]

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]^[7]

Taxonomy[]

Family: Manidae (pangolins)
- Subfamily: Maninae (Gray, 1821)
  - Genus: Manis (Linnaeus, 1758) (Asiatic pangolins)
    - Manis crassicaudata (Gray, 1827) (Indian pangolin)
    - Manis pentadactyla (Linnaeus, 1758) (Chinese pangolin)
    - † (Kormos, 1934)
    - † (Dubois, 1908)
    - Subgenus: Paramanis (Pocock, 1924)
      - Manis javanica (Desmarest, 1822) (Sunda pangolin)
      - Manis culionensis (de Elera, 1895) (Philippine pangolin)
      - †Manis palaeojavanica (Dubois, 1907) (giant asian pangolin)
- Subfamily: Phatagininae (Gaubert, 2017) (small African pangolins)
  - Genus: Phataginus (Rafinesque, 1821) (African tree pangolins)
    - Phataginus tetradactyla (Linnaeus, 1766) (long-tailed pangolin)
    - Phataginus tricuspis (Rafinesque, 1821) (tree pangolin)
- Subfamily: Smutsiinae (Gray, 1873) (large African pangolins)
  - Genus: Smutsia (Gray, 1865) (African ground pangolins)
    - Smutsia gigantea (Illiger, 1815) (giant pangolin)
    - Smutsia temmincki (Smuts, 1832) (ground pangolin)
    - † (Terhune, 2021)^[8]
- Incertae sedis
  - †Manidae sp. [DPC 3972 and DPC 4364] (Gebo & Rasmussen, 1985) (Fayum pangolin)

Phylogeny[]

Phylogenetic position of family Manidae within superfamily Manoidea.^[5]^[7]^[9]^[10]^[1]

Pholidotamorpha

†Palaeanodonta

Pholidota

†Euromanis

?

†Pholidota sp. (BC 16’08)

†Eurotamanduidae

Eupholidota

†Eomanoidea

Manoidea

†Patriomanidae

?

†Necromanis

Manidae

Maninae

Manis

(Paramanis)

	Manis culionensis

	Manis javanica

?	†Manis palaeojavanica

?

Manis sp. (Scale_H4 & Scale_H8)

?	†

	Manis crassicaudata

?	†

	Manis pentadactyla

?

†

Smutsiinae

Phatagininae

Phataginus

	Phataginus tetradactyla

	Phataginus tricuspis

Smutsiinae

Smutsia

	Smutsia gigantea

	†

	Smutsia temminckii

sensu stricto

sensu lato

sensu stricto

(Pholidota sensu lato)

References[]

^ ^a ^b ^c Philippe Gaubert, Agostinho Antunes, Hao Meng, Lin Miao, Stéphane Peigné, Fabienne Justy, Flobert Njiokou, Sylvain Dufour, Emmanuel Danquah, Jayanthi Alahakoon, Erik Verheyen, William T Stanley, Stephen J O’Brien, Warren E Johnson, Shu-Jin Luo (2018) "The Complete Phylogeny of Pangolins: Scaling Up Resources for the Molecular Tracing of the Most Trafficked Mammals on Earth" Journal of Heredity, Volume 109, Issue 4, Pages 347–359
^ J. E. Gray. (1821.) "On the natural arrangement of vertebrose animals." The London Medical Repository Monthly Journal and Review 15:296-310
^ "The CITES Appendices". Convention on International Trade in Endangered Species of Wild Fauna and Flora. CITES. Retrieved 28 January 2019.
^ 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
^ ^a ^b ^c Gaudin, Timothy (2009). "The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis" (PDF). Journal of Mammalian Evolution. Heidelberg, Germany: Springer Science+Business Media. 16 (4): 235–305. doi:10.1007/s10914-009-9119-9. S2CID 1773698.
^ Daniel Gebo, D. Tab Rasmussen (1985.) "The Earliest Fossil Pangolin (Pholidota: Manidae) from Africa" Journal of Mammalogy 66(3):538
^ ^a ^b 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.
^ Terhune, C. E.; Gaudin, T.; Curran, S.; Petculescu, A. (2021). "The youngest pangolin (Mammalia, Pholidota) from Europe". Journal of Vertebrate Paleontology. in press: e1990075. doi:10.1080/02724634.2021.1990075. S2CID 245394367.
^ Du Toit, Z.; Grobler, J. P.; Kotzé, A.; Jansen, R.; Brettschneider, H.; Dalton, D. L. (2014). "The complete mitochondrial genome of Temminck's ground pangolin (Smutsia temminckii; Smuts, 1832) and phylogenetic position of the Pholidota (Weber, 1904)". Gene. 551 (1): 49–54. doi:10.1016/j.gene.2014.08.040. PMID 25158133.
^ du Toit, Z.; du Plessis, M.; Dalton, D. L.; Jansen, R.; Paul Grobler, J.; Kotzé, A. (2017). "Mitochondrial genomes of African pangolins and insights into evolutionary patterns and phylogeny of the family Manidae". BMC Genomics. 18 (1): 746. doi:10.1186/s12864-017-4140-5. PMC 5609056. PMID 28934931.

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 (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.

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[Gaubert2018-1] Philippe Gaubert, Agostinho Antunes, Hao Meng, Lin Miao, Stéphane Peigné, Fabienne Justy, Flobert Njiokou, Sylvain Dufour, Emmanuel Danquah, Jayanthi Alahakoon, Erik Verheyen, William T Stanley, Stephen J O’Brien, Warren E Johnson, Shu-Jin Luo (2018) "The Complete Phylogeny of Pangolins: Scaling Up Resources for the Molecular Tracing of the Most Trafficked Mammals on Earth" Journal of Heredity, Volume 109, Issue 4, Pages 347–359

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

[CITES-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

[web2.utc.edu-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. Heidelberg, Germany: Springer Science+Business Media. 16 (4): 235–305. doi:10.1007/s10914-009-9119-9. S2CID 1773698.

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

[KondrashovAgadjanian2012-7] 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.

[8] Terhune, C. E.; Gaudin, T.; Curran, S.; Petculescu, A. (2021). "The youngest pangolin (Mammalia, Pholidota) from Europe". Journal of Vertebrate Paleontology. in press: e1990075. doi:10.1080/02724634.2021.1990075. S2CID 245394367.

[Du_Toit2014-9] Du Toit, Z.; Grobler, J. P.; Kotzé, A.; Jansen, R.; Brettschneider, H.; Dalton, D. L. (2014). "The complete mitochondrial genome of Temminck's ground pangolin (Smutsia temminckii; Smuts, 1832) and phylogenetic position of the Pholidota (Weber, 1904)". Gene. 551 (1): 49–54. doi:10.1016/j.gene.2014.08.040. PMID 25158133.

[du_Toit2017-10] u Toit, Z.; du Plessis, M.; Dalton, D. L.; Jansen, R.; Paul Grobler, J.; Kotzé, A. (2017). "Mitochondrial genomes of African pangolins and insights into evolutionary patterns and phylogeny of the family Manidae". BMC Genomics. 18 (1): 746. doi:10.1186/s12864-017-4140-5. PMC 5609056. PMID 28934931.

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Manidae Temporal range: 37.9–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Late Eocene - present^[1]

Living speacies of pangolins
Conservation status
CITES Appendix I (CITES)^[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
Subfamily: Maninae Manis Subfamily: Phatagininae Phataginus Subfamily: Smutsiinae Smutsia Incertae sedis †

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)