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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Cladus: Synapsida
Cladus: Eupelycosauria
Cladus: Sphenacodontia
Cladus: Sphenacodontoidea
Cladus: Therapsida
Cladus: Theriodontia
Cladus: Cynodontia
Cladus: Eucynodontia
Cladus: Probainognathia
Cladus: Prozostrodontia
Cladus: Mammaliaformes
Classis: Mammalia
Subclassis: Trechnotheria
Infraclassis: Zatheria
Supercohors: Theria
Cohors: Eutheria
Infraclassis: Placentalia
Cladus: Boreoeutheria
Superordo: Laurasiatheria
Cladus: Ferae
Ordo: Pholidota

Familia: Manidae
Genus: ManisPhataginusSmutsia

Name

Manidae Gray, 1821: 305
References
Primary references

Gray, J.E. 1821. On the Natural Arrangement of Vertebrose Animals. London Medical Repository 15: 296–310. HathiTrust. Reference page.

Additional references

Gaudin, T.J., Emry, R.J. & Wible, J.R. 2009. The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis. Journal of Mammalian Evolution 16(4): 235–305. DOI: 10.1007/s10914-009-9119-9 ResearchGate Reference page.

Links

Manidae in Mammal Species of the World.
Wilson, Don E. & Reeder, DeeAnn M. (Editors) 2005. Mammal Species of the World – A Taxonomic and Geographic Reference. Third edition. ISBN 0-8018-8221-4.

Vernacular names
Deutsch: Schuppentiere
Nordfriisk: Skolepdiarten
magyar: Tobzoskafélék
한국어: 천산갑과(穿山甲科)
Nederlands: Schubdier
polski: Łuskowcowate
português: Manídeos
ไทย: ลิ่น, นิ่ม

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][7] as well as 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)
See also

Mammal classification
Manoidea

References

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
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.
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, 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
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.
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. 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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