List of pholidotans

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

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

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]   Maninae/Manis   Indian pangolin Philippine pangolin Sunda pangolin Chinese pangolin Smutsiinae/Smutsia   Giant pangolin Ground pangolin Phatagininae/Phataginus   Tree pangolin Long-tailed pangolin

Pholidotans

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

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	M. crassicaudata E. Geoffroy, 1803	Indian subcontinent (current range in blue)	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]
Chinese pangolin	M. pentadactyla Linnaeus, 1758 Three subspecies M. p. auritus M. p. pusilla M. p. pentadactyla (Taiwanese pangolin)	Southeast Asia (current range in green)	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] [7]
Philippine pangolin	M. culionensis de Elera, 1895	Southeast Asia (current range in brown)	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]
Sunda pangolin	M. javanica Desmarest, 1822	Southeast Asia (current range in green)	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 habitats, and marine habitats[10] Diet: Primarily consume ants and their larvae, bee pupas, crickets, flies and termites[10]	CR  unknown [10]

Subfamily Phatagininae

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	P. tetradactyla Linnaeus, 1766	Central and West Africa (current range in green-brown)	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]
Tree pangolin	P. tricuspis (Rafinesque, 1821)	Central and West Africa (current range in brown)	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]

Subfamily Smutsiinae

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	S. gigantea Illiger, 1815	Central and East Africa (current range in light brown)	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]
Ground pangolin	S. temminckii (Smuts, 1832)	Southern, East and North Africa (current range in green)	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]

Prehistoric pholidotans

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]		Eocene PreꞒ Ꞓ O S D C P T J K Pg N [1]	Messel Pit in Germany[16]

Superfamily Manoidea

Main article: Manoidea

Genus Necromanis – Filhol, 1894 – 3 species

Species name	Image	Geological range	Biogeography
Necromanis franconica Quenstedt, 1885[17]		Paleogene–Neogene PreꞒ Ꞓ O S D C P T J K Pg N [1]	Europe[15]
Necromanis parva Koenigswald, 1969[18]		Paleogene–Neogene PreꞒ Ꞓ O S D C P T J K Pg N [1]	Europe[15]
Necromanis quercyi Filhol, 1894[19]		Paleogene–Neogene PreꞒ Ꞓ O S D C P T J K Pg N [1]	Europe[15]

Family Patriomanidae

Main article: Patriomanidae

Genus Patriomanis – Emry, 1970 – 1 species

Species name	Image	Geological range	Biogeography
Patriomanis americana Emry, 1970[20]		Eocene PreꞒ Ꞓ O S D C P T J K Pg N [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 PreꞒ Ꞓ O S D C P T J K Pg N [1]	Inner Mongolia, China[21]

Family Eurotamanduidae

Genus Eurotamandua – Storch, 1981 – 1 species

Species name	Image	Geological range	Biogeography
Eurotamandua joresi Storch, 1981[22]		Eocene PreꞒ Ꞓ O S D C P T J K Pg N [23]	Messel Pit in Germany[23]

Family Eomanidae

Main article: Eomanidae

Genus Eomanis – Storch, 1978 – 1 species

Species name	Image	Geological range	Biogeography
Eomanis waldi Storch, 1978[24]		Eocene PreꞒ Ꞓ O S D C P T J K Pg N [1]	Messel Pit in Germany[24]

Family Manidae

Main article: Manidae

Genus Manis – Linnaeus, 1758 – 3[note 1] species

Species name	Image	Geological range	Biogeography
Manis palaeojavanica Dubois, 1907[25]		Pleistocene PreꞒ Ꞓ O S D C P T J K Pg N ↓ [1]	Asia[25]
Manis lydekkeri Dubois, 1908[26]		Pleistocene PreꞒ Ꞓ O S D C P T J K Pg N ↓ [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 PreꞒ Ꞓ O S D C P T J K Pg N ↓ [28]	Europe[28]

See also

• Pholidotamorpha
• Pangolin trade

Notes

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

References

1. Pangolins: Science, Society and Conservation. Academic Press. 23 November 2019. pp. 5–15, 44–46. ISBN 978-0-12-815506-6.
2. 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-US. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. Storch, G.; Martin, T. (1994-01-01). "Eomanis krebsi, ein neues Schuppentier aus dem Mittel-Eozan der Grube Messel bei Darmstadt (Mammalia: Pholidota)". Berliner Geowissenschaftliche Abhandlungen Reihe E Palaeobiologie (in German). 13: 83–97. ISSN 0941-7338.
17. Quenstedt, Fr Aug von (Friedrich August) (1885). Handbuch der petrefaktenkunde (in German). 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 (in German). 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 (in French). Vol. ser.7:t.16 (1894). Paris: Crochard. pp. 129–150.
20. 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. 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-US. Retrieved 2024-11-25.
23. "Abstracts of Papers". Journal of Vertebrate Paleontology. 17 (sup003): 1–93. 4 September 1997. doi:10.1080/02724634.1997.10011028. ISSN 0272-4634.
24. 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. 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. (in Dutch). 24: 449.
26. "Tijdschrift van het Aardrijkskundig Genootschap". Das geologische Alter der Kendeng- oder Trinil-Fauna (in Dutch). 2 (25): 1235–1270. 1 January 1908.
27. Kormos, T. H.; Kormos, T. H. (1934-01-01). "Manis hungarica n sp, das erste Schuppentier aus dem europaischen Oberpliozan". Folia Zoologica et Hydrobiologica (in German). 6 (1): 87–94.
28. 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.

External links

Media related to Manidae at Wikimedia Commons