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 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

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

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

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

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
(Pholidotasensu lato)

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] [7]}

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 [8] Diet: Myrmecophagous; eats ants and termites [8]	EN  unknown [8] [8]
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, 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] [9]
Philippine pangolin	M. culionensis de Elera, 1895	Southeast Asia (current range in brown)	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] [11]
Sunda pangolin	M. javanica Desmarest, 1822	Southeast Asia (current range in green)	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] [12]

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 [13] Diet: Feeds on ants and termites [13]	VU  unknown [13] [13]
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 [14] Diet: Feeds on ants and termites [14]	EN  unknown [14] [14]

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 [15] Diet: Eats mainly ants and termites, and sometimes other insects [15]	EN  unknown [15] [15]
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 [16] Diet: Myrmecophagous; preys primarily on ants and termites [16]	VU  16,300–24,000 in South Africa [16] [16]

Prehistoric pholidotans

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

• Pholidotamorpha
• Pangolin trade

References

1. Pangolins: Science, Society and Conservation. Academic Press. 2019-11-23. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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.

External links

Media related to Manidae at Wikimedia Commons