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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 their skin, 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 assigned to the family Manidae which is split into three genera within three subfamilies: Maninae (Asian pangolins), Phatagininae (African tree pangolins) and Smutsiinae (African ground pangolins). A ninth species, Manis mysteria, was proposed in 2023, though its status as valid taxa remains uncertain. Around a dozen extinct prehistoric pholidotan species have been described; however, due to ongoing research the exact categorization and number is not fixed.[1][3]

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)

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.

  • Family Manidae
    • Subfamily Maninae
      • Genus: Manis (Asian pangolins): four species
    • Subfamily Phatagininae
      • Genus: Phataginus (African tree pangolins): two species
    • Subfamily Smutsiinae
      • Genus: Smutsia (African ground pangolins): two species
Manidae[4]  

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][5]

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[6]

Diet: Myrmecophagous; eats ants and termites[6] 		 EN 


unknown[6] Population decreasing[6]

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, with a 25–38 cm (9.8–15.0 in) long tail and a weight of 2.5–7 kg (5.5–15.4 lb)[2]

Habitat: Forest, shrubland and grassland[7]

Diet: Eats insects such as ants and termites[7] 		 CR 


10,000[8] Population declining[7]

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, with a 39–50 cm (15–20 in) long tail and a weight of 2.5–8 kg (5.5–17.6 lb)[2]

Habitat: Forest, shrubland, and artificial habitats[9]

Diet: Preys exclusively on ant and termite species[9] 		 CR 


unknown[9] Population decreasing[9]

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, with a 35–58 cm (14–23 in) long tail and a weight of 3–10 kg (6.6–22.0 lb)[2]

Habitat: Forests, shrublands, artificial terrestrial and aquatic habits, and marine habitats[10]

Diet: Primarily consume ants and their larvae, bee pupas, crickets, flies and termites[10] 		 CR 


unknown[10] population decreasing[10]

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[11]

Diet: Feeds on ants and termites[11] 		 VU 


unknown[11] Population declining[11]

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[12]

Diet: Feeds on ants and termites[12] 		 EN 


unknown[12] Population declining[12]

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[13]

Diet: Eats mainly ants and termites, and sometimes other insects[13] 		 EN 


unknown[13] Population declining[13]

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[14]

Diet: Myrmecophagous; preys primarily on ants and termites[14] 		 VU 


16,300–24,000 in South Africa[14] Population declining[14]

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: Eomanidae, Patriomanidae and Eurotamandua, as well as the extant family Manidae. The genus Necromanis on the other hand, is placed as incertae sedis within the pholidotid superfamily Manoidea, together with the families Manidae and Patriomanidae. Euromanis, a genus described in 2009, is not placed under any family and is instead assigned to the Pholidota order.[15][1] The extinct 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 will only contain taxa that went extinct during prehistoric times.

Genus Euromanis – Gaudin, 2009 – 1 species
Species name Image Geological range Biogeography
Euromanis krebsi

Storch & Martin, 1994[16]

Euromanis krebsi bones Eocene [1] Messel Pit in Germany[16]

Superfamily Manoidea

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Main article: Manoidea
Genus Necromanis – Filhol, 1894 – 3 species
Species name Image Geological range Biogeography
Necromanis franconica

Quenstedt, 1885[17]

Paleogene–Neogene [1] Europe[15]
Necromanis parva

Koenigswald, 1969[18]

Paleogene–Neogene [1] Europe[15]
Necromanis quercyi

Filhol, 1894[19]

Paleogene–Neogene [1] Europe[15]

Family Patriomanidae

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Main article: Patriomanidae
Genus Patriomanis – Emry, 1970 – 1 species
Species name Image Geological range Biogeography
Patriomanis americana

Emry, 1970[20]

Life reconstruction of Patriomanis americana

Eocene [1] North America[20]
Genus Cryptomanis – Gaudin, Emry, and Pogue, 2006 – 1 species
Species name Image Geological range Biogeography
Cryptomanis gobiensis

Gaudin, Emry, and Pogue, 2006[21]

Eocene [1] Inner Mongolia, China[21]

Family Eurotamanduidae

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Genus Eurotamandua – Storch, 1981 – 1 species
Species name Image Geological range Biogeography
Eurotamandua joresi

Storch, 1981[22]

Life reconstruction of Eurotamandua joresi Eocene [23] Messel Pit in Germany[23]

Family Eomanidae

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Main article: Eomanidae
Genus Eomanis – Storch, 1978 – 1 species
Species name Image Geological range Biogeography
Eomanis waldi

Storch, 1978[24]

Life reconstruction of Eomanis waldi Eocene [1] Messel Pit in Germany[24]

Family Manidae

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Main article: Manidae
Genus Manis – Linnaeus, 1758 – 3[note 1] species
Species name Image Geological range Biogeography
Manis palaeojavanica

Dubois, 1907[25]

Pleistocene
↓
[1] 		Asia[25]
Manis lydekkeri

Dubois, 1908[26]

Pleistocene
↓
[1] 		Unknown
Manis hungarica

Kormos, 1934[27]

Unknown Unknown
Genus Smutsia – Gray, 1865 – 1[note 2] species
Species name Image Geological range Biogeography
Smutsia olteniensis

Terhune, Gaudin, Curran & Petculescu, 2021[28]

Pleistocene
↓
[28] 		Europe[28]

See also

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Notes

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  1. ^ This only includes extinct Manis species
  2. ^ This only includes extinct Smutsia species

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. 23 November 2019. 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. ^ Anderson, Natali (2 October 2023). "DNA Study Reveals Cryptic New Species of Pangolin | Sci.News". Sci.News: Breaking Science News. Retrieved 2024-11-25.
  4. ^ Gaubert, Philippe; Antunes, Agostinho; Meng, Hao; Miao, Lin; Peigné, Stéphane; Justy, Fabienne; Njiokou, Flobert; Dufour, Sylvain; Danquah, Emmanuel; Alahakoon, Jayanthi; Verheyen, Erik (11 May 2018). "The Complete Phylogeny of Pangolins: Scaling Up Resources for the Molecular Tracing of the Most Trafficked Mammals on Earth". Journal of Heredity. 109 (4): 347–359. doi:10.1093/jhered/esx097. PMID 29140441.
  5. ^ 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.
  6. ^ 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.
  7. ^ 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.
  8. ^ Zhang, Fuhua; Wu, Shibao; Cen, Peng (1 January 2022). "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.
  9. ^ 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.
  10. ^ 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.
  11. ^ 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.
  12. ^ 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.
  13. ^ 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.
  14. ^ 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.
  15. ^ a b c d Gaudin, Timothy J.; Emry, Robert J.; Wible, John R. (1 December 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. ISSN 1573-7055.
  16. ^ a b Storch, G. and Martin, T., 1994. Eomanis krebsi, ein neues Schuppentier aus dem Mittel-Eozän der Grube Messel bei Darmstadt (Mammalia: Pholidota). Berliner geowissenschaftliche Abhandlungen E, 13, pp.83-97.
  17. ^ Quenstedt, Fr Aug von (Friedrich August) (1885). Handbuch der petrefaktenkunde. Oxford University. Tübingen, H. Laupp. p. 1239.
  18. ^ Koenigswald, W.V. (1969). Die Maniden (Pholidota, Mamm.) des europäischen Tertiärs. Mitteilungen der Bayerischen Staatssammlung für Paläontologie und historische Geologie 9. pp. 61–71.
  19. ^ Audouin, Jean Victor; Audouin, Jean Victor; Bouvier, E.-L.; Grassé, Pierre-Paul; Milne-Edwards, H.; Milne-Edwards, Alphonse; Perrier, Edmond (1834). Annales des sciences naturelles. Zoologie. Vol. ser.7:t.16 (1894). Paris: Crochard. pp. 129–150.
  20. ^ a b Emry, R. J. (1970). "A North American Oligocene pangolin and other additions to the Pholidota". Bulletin of the American Museum of Natural History. 142 (6): 1–278.
  21. ^ a b Gaudin, Timothy J.; Emry, Robert J.; Pogue, Brandon (30 March 2006). "A new genus and species of pangolin (Mammalia, Pholidota) from the late Eocene of Inner Mongolia, China". Journal of Vertebrate Paleontology. 26 (1): 146–159. doi:10.1671/0272-4634(2006)26[146:ANGASO]2.0.CO;2. ISSN 0272-4634.
  22. ^ Bechly, Günter (14 October 2022). "Eurotamandua: Anteater or Not Even Close?". Evolution News. Retrieved 2024-11-25.
  23. ^ a b "Abstracts of Papers". Journal of Vertebrate Paleontology. 17 (sup003): 1–93. 4 September 1997. doi:10.1080/02724634.1997.10011028. ISSN 0272-4634.
  24. ^ a b Difouo Fopa, Ghislain (2023). Influence of habitat on pangolins’ (Pholidota: Manidae) feeding behavior, potential and effective diet composition in two forest-savanna mosaic zones of Cameroon (Thesis thesis). Université de Yaoundé I.
  25. ^ a b E, Dubois (1907). "Eenige van Nederlanschen kant verkregen uitkomsten met betrekking tot de kennis der Kendeng-fauna (Fauna van Trinil)". Tijdschr. K. Nederlansch. Aardrijkskd. Genoot. 24: 449.
  26. ^ "Tijdschrift van het Aardrijkskundig Genootschap, 1908, 01-01-1908". Das geologische Alter der Kendeng- oder Trinil-Fauna. 2 (25): 1235–1270. 1 January 1908.
  27. ^ Kormos, T. 1934. Manis hungarica n. s., das erste Schuppentier aus dem europäischen Oberpliozän. [./Folia_zoologica_et_hydrobiologica?action=edit&redlink=1 Folia zoologica et hydrobiologica] 6: 87–94.
  28. ^ a b c Terhune, Claire E.; Gaudin, Timothy; Curran, Sabrina; Petculescu, Alexandru (4 July 2021). "The youngest pangolin (Mammalia, Pholidota) from Europe". Journal of Vertebrate Paleontology. 41 (4). doi:10.1080/02724634.2021.1990075. ISSN 0272-4634.
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Media related to Manidae at Wikimedia Commons

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