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Four Manid species in a collage; from top-left, clockwise: ground pangolin, tree pangolin, Philippine pangolin and Sunda pangolin
Manidae species of different genera; from top-left, clockwise: ground pangolin (Smutsia temminckii), tree pangolin (Phataginus tricuspis), Philippine pangolin (Manis culionensis), and Sunda pangolin (Manis javanica)

Pholidota is an order of placental mammals in the Pholidotamorpha clade. A member of this order is called a pholidotan or a pangolin. They are distributed throughout Africa, the Indian subcontinent and southeast Asia, and are usually found in habitats such as savannas, forests, grasslands and shrublands. Pholidotans range in size from the giant pangolin, at 30 kg (66 lb) and 68 cm (27 in) in length, to the tree pangolin, at only 2.3 kg (5.1 lb) and 34 cm (13 in) in length. They have large, hardened, keratin scales which cover the entirety of the body surface, and long claws which they use for digging or climbing trees. Most species are nocturnal and feed primarily on ants and termites, though other insects may also be taken. Despite only a few pholidotans having population estimates, all eight species are classified as threatened by the IUCN Red List, with three being classified as Critically endangered, largely due to intensive poaching of their keratin scales.[1][2]

The eight extant species of Pholidota are split into three genera within three subfamilies: Maninae (Asian pangolin), Phatagininae (African tree pangolin) and Smutsiinae (African ground pangolin). Manidae is the only extant family within the Pholidota order, as the other three families—Eurotamandua, Patriomanidae, and Eomanis—have gone entirely extinct. As such, around a dozen or more extinct species have been described, though the exact categorization and number is not fixed due to ongoing research.[1]

Conventions

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IUCN Red List categories
Conservation status
 EX Extinct (0 species)
 EW Extinct in the wild (0 species)
 CR Critically Endangered (3 species)
 EN Endangered (3 species)
 VU Vulnerable (2 species)
 NT Near threatened (0 species)
 LC Least concern (0 species)
Other categories
 DD Data deficient (0 species)
 NE Not evaluated (0 species)

Conservation status codes listed follow the International Union for Conservation of Nature (IUCN) Red List of Threatened Species. Range maps are provided where possible; if unavailable, a description of the species will instead be provided. All extinct species and subspecies listed alongside extant species went extinct in prehistoric times, and are indicated by dagger symbol "†". Population figures are rounded to the nearest hundred.

Classification

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The Pholidota order is composed of eight extant species belonging to three genera in three subfamilies. This does not include extinct species or hybrid animals.

Phylogenetic tree of the Pholidotamorpha clade.[3][4][5][6]

 Pholidotamorpha 
southern Asian clade
northern Asian clade
African clade
 (Pholidota sensu lato) 

Pholidotans

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The following classification is based on the taxonomy described by the reference work Mammal Species of the World (2005), with augmentation by generally accepted proposals made since using molecular phylogenetic analysis, as supported by the IUCN Red List.[1][7]

Subfamily Maninae

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Main article: Maninae
Genus Manis – Linnaeus, 1758 – four species
Common name Scientific name and subspecies Range Size and ecology IUCN status and estimated population
Indian pangolin

Indian pangolin walking in dirt

M. crassicaudata
E. Geoffroy, 1803 		Indian subcontinent (current range in blue)
Map of range 		Size: 51–75 cm (20–30 in) long, with a 33 to 47 cm (13 to 19 in) long tail
and a weight of 10–16 kg (22–35 lb)[2]

Habitat: Savanna, forest, grassland and shrubland[8]

Diet: Myrmecophagous; eats ants and termites[8] 		 EN 


unknown[8] Population decreasing[8]

Chinese pangolin

Chinese pangolin in green grass

M. pentadactyla
Linnaeus, 1758

Three subspecies
 Southeast Asia (current range in green)
Map of range 		Size: 40–58 cm (16–23 in) long, plus 25–38 cm (9.8–15.0 in) tail
2.5–7 kg (5.5–15.4 lb)[2]

Habitat: Forest, shrubland and grassland[9]

Diet: Eats insects such as ants and termites[9] 		 CR 


10,000[10] Population declining[9]

Philippine pangolin

A Phillipine pangolin female and her young in a muddy area near some green grass

M. culionensis
de Elera, 1895 		Southeast Asia (current range in brown)
Map of range 		Size: 45–54 cm (18–21 in) long, plus 39–50 cm (15–20 in) tail
2.5–8 kg (5.5–17.6 lb)[2]

Habitat: Forest, shrubland, and artificial habitats[11]

Diet: Preys exclusively on ant and termite species[11] 		 CR 


unknown[11] Population decreasing[11]

Sunda pangolin

A Sunda pangolin amongst fallen tree branches

M. javanica
Desmarest, 1822 		Southeast Asia (current range in green)
Map of range 		Size: 40–65 cm (16–26 in) long, plus 35–58 cm (14–23 in) tail
3–10 kg (6.6–22.0 lb)[2]

Habitat: Forests, shrublands, artificial terrestrial and aquatic habits, and marine habitats[12]

Diet: Primarily consume ants and their larvae, bee pupas, crickets, flies and termites[12] 		 CR 


unknown[12] population decreasing[12]

Subfamily Phatagininae

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Main article: Phatagininae
Genus Phataginus – Linnaeus, 1766 – two species
Common name Scientific name and subspecies Range Size and ecology IUCN status and estimated population
Long-tailed pangolin

A bright gold and black pangolin on a very thin green branch

P. tetradactyla
Linnaeus, 1766 		Central and West Africa (current range in green-brown)
Map of range 		Size: 30–40 cm (12–16 in) long, with a 55–70 cm (22–28 in) long tail and a weight of 2–3.5 kg (4.4–7.7 lb)[2]

Habitat: Forest, savanna, and artificial habitats[13]

Diet: Feeds on ants and termites[13] 		 VU 


unknown[13] Population declining[13]

Tree pangolin

A tree pangolin hanging near vertically on a thin branch that is surrounded by green leaves

P. tricuspis
(Rafinesque, 1821) 		Central and West Africa (current range in brown)
Map of range 		Size: 25–43 cm (9.8–16.9 in) long, with a 35–62 cm (14–24 in) long tail and a weight of 1.6–3 kg (3.5–6.6 lb)[2]

Habitat: Forest, savanna, and artificial habitats[14]

Diet: Feeds on ants and termites[14] 		 EN 


unknown[14] Population declining[14]

Subfamily Smutsiinae

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Main article: Smutsiinae
Genus Smutsia – Gray, 1865 – two species
Common name Scientific name and subspecies Range Size and ecology IUCN status and estimated population
Giant pangolin

A giant pangolin exhibited in the Natural History Museum of London

S. gigantea
Illiger, 1815 		Central and East Africa (current range in light brown)
Map of range 		Size: 67–81 cm (26–32 in) long, with a 58–68 cm (23–27 in) long tail and a weight of 30 kg (66 lb)[2]

Habitat: Forest and savanna[15]

Diet: Eats mainly ants and termites, and sometimes other insects[15] 		 EN 


unknown[15] Population declining[15]

Ground pangolin

A ground pangolin walking with its forelegs slightly above the earth's surface in a horizontal manner rather than a vertical one

S. temminckii
(Smuts, 1832) 		Southern, East and North Africa (current range in green)
Map of range 		Size: 45–55 cm (18–22 in) long, with a 40–52 cm (16–20 in) long tail and a weight of 5–20 kg (11–44 lb)[2]

Habitat: Forest, savanna and grassland[16]

Diet: Myrmecophagous; preys primarily on ants and termites[16] 		 VU 


16,300–24,000 in South Africa[16] Population declining[16]

Prehistoric pholidotans

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In addition to extant species, multiple extinct species have been described and classified into Pholidota. They are placed into three extinct families: Eomanis, Patriomanidae and Eurotamandua, as well as the extant family Manidae. However, the genus Necromanis is placed as incertae sedis within the pholidotid superfamily Manoidea, together with the families Manidae and Patriomanidae. The species listed here are mainly based off the 2019 reference work Pangolins: Science, Society and Conservation, unless otherwise cited. Where available, the temporal range of each species will be given in millions of years before the present time (mya). This list only includes extinct species that went extinct during prehistoric times.[17][1]

Image Binomial name Genus Superfamily or family Temporal range
Euromanis krebsi Euromanis None ~45 mya[1]
Eurotamandua joresi Eurotamandua Eurotamanduidae 47.8–46.3 mya[18]
Image unavailable Cryptomanis gobiensis Cryptomanis Patriomanidae ~40 mya[1]
Patriomanis americana Patriomanis Patriomanidae ~35–37 mya[1]
Eomanis waldi Eomanis Eomanidae ~45 mya[1]
Image unavailable Manis hungarica Manis Manidae Unknown[19]
Image unavailable Manis lydekkeri Manis Manidae Pleistocene[1]
Manis palaeojavanica Manis Manidae Pleistocene[1]
Image unavailable Smutsia olteniensis Smutsia Manidae 2.2–1.9 mya[20]
Necromanis franconica Necromanis Manoidea ~28–14 mya[1]
Image unavailable Necromanis parva Necromanis Manoidea ~28–14 mya[1]
Image unavailable Necromanis quercyi Necromanis Manoidea ~28–14 mya[1]

See also

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References

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  1. ^ a b c d e f g h i j k l m Pangolins: Science, Society and Conservation. Academic Press. 2019-11-23. pp. 5–15, 44–46. ISBN 978-0-12-815506-6.
  2. ^ a b c d e f g h i Chernasky, Amy; Motis, Anna; Burgin, Connor, eds. (2023). All the Mammals of the World. Lynx Nature Books. pp. 638–639. ISBN 978-84-16728-66-4.
  3. ^ 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.
  4. ^ 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.
  5. ^ 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.
  6. ^ 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.
  7. ^ Gardner, Alfred L. (2005). Wilson, Don E.; Reeder, DeeAnn M. (eds.). Mammal Species of the World. Vol. 1 (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0.
  8. ^ a b c d Mahmood, T.; Challender, D.; Khatiwada, A.; Andleeb, S.; Perera, P.; Trageser, S.; Ghose, A.; Mohapatra, R.K. (2019). "Manis crassicaudata". IUCN Red List of Threatened Species. 2019: e.T12761A123583998. doi:10.2305/IUCN.UK.2019-3.RLTS.T12761A123583998.en. Retrieved 17 January 2022.
  9. ^ a b c Challender, D.; Wu, S.; Kaspal, P.; Khatiwada, A.; Ghose, A.; Ching-Min Su, N. & Laxmi Suwal, T. (2020) [errata version of 2019 assessment]. "Manis pentadactyla". IUCN Red List of Threatened Species. 2019: e.T12764A168392151. doi:10.2305/IUCN.UK.2019-3.RLTS.T12764A168392151.en. Retrieved 17 January 2022.
  10. ^ Zhang, Fuhua; Wu, Shibao; Cen, Peng (2022-01-01). "The past, present and future of the pangolin in Mainland China". Global Ecology and Conservation. 33: e01995. Bibcode:2022GEcoC..3301995Z. doi:10.1016/j.gecco.2021.e01995. ISSN 2351-9894.
  11. ^ a b c d Schoppe, S.; Katsis, L.; Lagrada, L. (2019). "Manis culionensis". IUCN Red List of Threatened Species. 2019: e.T136497A123586862. doi:10.2305/IUCN.UK.2019-3.RLTS.T136497A123586862.en. Retrieved 19 November 2021.
  12. ^ a b c d Challender, D.; Willcox, D.H.A.; Panjang, E.; Lim, N.; Nash, H.; Heinrich, S.; Chong, J. (2019). "Manis javanica". IUCN Red List of Threatened Species. 2019: e.T12763A123584856. doi:10.2305/IUCN.UK.2019-3.RLTS.T12763A123584856.en. Retrieved 17 January 2022.
  13. ^ a b c d Ingram, D.J.; Shirley, M.H.; Pietersen, D.; Godwill Ichu, I.; Sodeinde, O.; Moumbolou, C.; Hoffmann, M.; Gudehus, M.; Challender, D. (2019). "Phataginus tetradactyla". IUCN Red List of Threatened Species. 2019: e.T12766A123586126. doi:10.2305/IUCN.UK.2019-3.RLTS.T12766A123586126.en. Retrieved 19 November 2021.
  14. ^ a b c d Pietersen, D.; Moumbolou, C.; Ingram, D.J.; Soewu, D.; Jansen, R.; Sodeinde, O.; Keboy Mov Linkey Iflankoy, C.; Challender, D.; Shirley, M.H. (2019). "Phataginus tricuspis". IUCN Red List of Threatened Species. 2019: e.T12767A123586469. doi:10.2305/IUCN.UK.2019-3.RLTS.T12767A123586469.en. Retrieved 19 November 2021.
  15. ^ a b c d Nixon, S.; Pietersen, D.; Challender, D.; Hoffmann, M.; Godwill Ichu, I.; Bruce, T.; Ingram, D. J.; Matthews, N. & Shirley, M. H. (2019). "Smutsia gigantea". IUCN Red List of Threatened Species. 2019: e.T12762A123584478. doi:10.2305/IUCN.UK.2019-3.RLTS.T12762A123584478.en. Retrieved 17 January 2022.
  16. ^ a b c d Pietersen, D.; Jansen, R.; Connelly, E. (2019). "Smutsia temminckii". IUCN Red List of Threatened Species. 2019: e.T12765A123585768. doi:10.2305/IUCN.UK.2019-3.RLTS.T12765A123585768.en. Retrieved 19 November 2021.
  17. ^ Gaudin, J; Emry, J; Wible, R (2014). "The phylogeny of living and extinct pangolins (Mammalia, Pholidota) and associated taxa: a morphology based analysis (project)". MorphoBank datasets. doi:10.7934/p291. Retrieved 2024-11-10.
  18. ^ "Abstracts of Papers". Journal of Vertebrate Paleontology. 17 (sup003): 1–93. 1997-09-04. Bibcode:1997JVPal..17S...1.. doi:10.1080/02724634.1997.10011028. ISSN 0272-4634.
  19. ^ Botha, Jennifer; Gaudin, Timothy (2007-06-12). "An early pliocene pangolin (Mammalia; Pholidota) from Langebaanweg, South Africa". Journal of Vertebrate Paleontology. 27 (2): 484–491. doi:10.1671/0272-4634(2007)27[484:aepppf]2.0.co;2. ISSN 0272-4634.
  20. ^ Terhune, Claire E.; Gaudin, Timothy; Curran, Sabrina; Petculescu, Alexandru (2021-07-04). "The youngest pangolin (Mammalia, Pholidota) from Europe". Journal of Vertebrate Paleontology. 41 (4). Bibcode:2021JVPal..41E0075T. doi:10.1080/02724634.2021.1990075. ISSN 0272-4634.
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Media related to Manidae at Wikimedia Commons

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