Manidae ("spirits") 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),[6] as well as extinct Fayum pangolin.[7]
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][8]
Taxonomy
- Family: Manidae (pangolins)
- Subfamily: Maninae (Gray, 1821) (Asian pangolins)
- Genus: Manis (Linnaeus, 1758) (Asian pangolin)
- (unranked): northern Asian clade
- (unranked): southern Asian clade
- (unranked): African clade
- Subfamily: Phatagininae (Gaubert, 2017) (small African pangolins)
- Genus: Phataginus (Rafinesque, 1821) (African tree pangolin)
- Subfamily: Smutsiinae (Gray, 1873) (large African pangolins)
- Genus: Smutsia (Gray, 1865) (African ground pangolin)
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Incertae sedis
Phylogeny
Phylogenetic position of family Manidae within superfamily Manoidea.[5][8][12][13][6][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
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^ 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"
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^ J. E. Gray. (1821.) "On the natural arrangement of vertebrose animals." The London Medical Repository Monthly Journal and Review 15:296-310
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^ "The CITES Appendices". Convention on International Trade in Endangered Species of Wild Fauna and Flora. CITES. Retrieved 28 January 2019.
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^ 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
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^ 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.
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^ 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
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^ Daniel Gebo, D. Tab Rasmussen (1985.) "The Earliest Fossil Pangolin (Pholidota: Manidae) from Africa" Journal of Mammalogy 66(3):538
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^ 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.
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^ 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
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^ 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
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^ Terhune, C. E.; Gaudin, T.; Curran, S.; Petculescu, A. (2021). "The youngest pangolin (Mammalia, Pholidota) from Europe". Journal of Vertebrate Paleontology. 41 (4): e1990075. doi:10.1080/02724634.2021.1990075. S2CID 245394367.
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^ 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.
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^ 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.
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