Manidae

Manidae; Temporal range: 41.37–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N middle Eocene - present
	Living species of pangolins
Conservation status
	CITES Appendix I (CITES)
Scientific classification
Domain:	Eukaryota
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] 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

Family: Manidae (pangolins)
- Subfamily: Maninae (Gray, 1821) (Asian pangolins)
  - Genus: Manis (Linnaeus, 1758) (Asian pangolin)
    - (unranked): northern Asian clade
      - Manis pentadactyla (Linnaeus, 1758) (Chinese pangolin)
      - †Manis hungarica (Kormos, 1934)
    - (unranked): southern Asian clade
      - Manis crassicaudata (Gray, 1827) (Indian pangolin)
      - Manis sp. (Scale_H4 & Scale_H8)^[10]^[11]^[1]
      - †Manis lydekkeri (Dubois, 1908)
      - 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) (long-tailed pangolin)
      - Phataginus tricuspis (Rafinesque, 1821) (tree 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)^[12]
- Incertae sedis
  - †Manidae sp. [DPC 3972 & DPC 4364] (Gebo & Rasmussen, 1985) (Fayum pangolin)

Phylogeny

Phylogenetic position of family Manidae within superfamily Manoidea.^[5]^[9]^[13]^[14]^[7]^[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 lydekkeri

	Manis crassicaudata

?	†Manis hungarica

	Manis pentadactyla

?

†Manidае sp. (DPC 3972 & DPC 4364)

Smutsiinae

Phatagininae

Phataginus

	Phataginus tetradactyla

	Phataginus tricuspis

Smutsiinae

Smutsia

	Smutsia gigantea

	†Smutsia olteniensis

	Smutsia temminckii

sensu stricto

sensu lato

sensu stricto

southern Asian clade

northern Asian clade

African clade

(Pholidota sensu lato)

References

^ ^a ^b ^c 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"
^ 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. 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.
^ 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.
^ ^a ^b 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
^ 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. Bibcode:2012JVPal..32..983K. doi:10.1080/02724634.2012.694319. ISSN 0272-4634. S2CID 86059673.
^ Huarong Zhang, Mark P. Miller, Feng Yang, Hon Ki Chan, Philippe Gaubert, Gary Ades, Gunter A. Fischer (2015.) "Molecular tracing of confiscated pangolin scales for conservation and illegal trade monitoring in Southeast Asia", Global Ecology and Conservation, Volume 4, Pages 414-422
^ Jingyang Hu, Christian Roos, Xue Lv, Weimin Kuang, Li Yu (2020.) "Molecular Genetics Supports a Potential Fifth Asian Pangolin Species (Mammalia, Pholidota, Manis)" Zoological Science, 37(6):538-543
^ Terhune, C. E.; Gaudin, T.; Curran, S.; Petculescu, A. (2021). "The youngest pangolin (Mammalia, Pholidota) from Europe". Journal of Vertebrate Paleontology. 41 (4): e1990075. Bibcode:2021JVPal..41E0075T. 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 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.

[Heighton2023-1] 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

[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. 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.

[Gaubert2018-7] 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

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

[KondrashovAgadjanian2012-9] 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. Bibcode:2012JVPal..32..983K. doi:10.1080/02724634.2012.694319. ISSN 0272-4634. S2CID 86059673.

[10] Huarong Zhang, Mark P. Miller, Feng Yang, Hon Ki Chan, Philippe Gaubert, Gary Ades, Gunter A. Fischer (2015.) "Molecular tracing of confiscated pangolin scales for conservation and illegal trade monitoring in Southeast Asia", Global Ecology and Conservation, Volume 4, Pages 414-422

[JingyangHu-11] Jingyang Hu, Christian Roos, Xue Lv, Weimin Kuang, Li Yu (2020.) "Molecular Genetics Supports a Potential Fifth Asian Pangolin Species (Mammalia, Pholidota, Manis)" Zoological Science, 37(6):538-543

[12] Terhune, C. E.; Gaudin, T.; Curran, S.; Petculescu, A. (2021). "The youngest pangolin (Mammalia, Pholidota) from Europe". Journal of Vertebrate Paleontology. 41 (4): e1990075. Bibcode:2021JVPal..41E0075T. doi:10.1080/02724634.2021.1990075. S2CID 245394367.

[Du_Toit2014-13] 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_Toit2017-14] 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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