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Pangolin Temporal range: Paleocene–Present PreЄ Є O S D C P T J K Pg N
Sunda pangolin (Manis javanica)
Conservation status
CITES Appendix I (CITES)[1]
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Scrotifera
Clade:	Fereuungulata
(unranked):	Ferae
Order:	Pholidota Weber, 1904
Family:	Manidae Gray, 1821
Genera
Manis Phataginus Smutsia
Species ranges      Manis crassicaudata      Manis pentadactyla      Manis javanica      Manis culionensis      Phataginus tricuspis      Phataginus tetradactyla      Smutsia gigantea      Smutsia temminckii

A pangolin in defensive posture, Horniman Museum, London

Pangolins or scaly anteaters^[2] are mammals of the order Pholidota (from the Greek word φολῐ́ς, "horny scale"). The one extant family, Manidae, has three genera: Manis, which comprises four species living in Asia; Phataginus, which comprises two species living in Africa; and Smutsia, which comprises two species also living in Africa.^[3] These species range in size from 30 to 100 cm (12 to 39 in). A number of extinct pangolin species are also known.

Pangolins have large, protective keratin scales covering their skin; they are the only known mammals with this feature. They live in hollow trees or burrows, depending on the species. Pangolins are nocturnal, and their diet consists of mainly ants and termites, which they capture using their long tongues. They tend to be solitary animals, meeting only to mate and produce a litter of one to three offspring, which are raised for about two years.

Pangolins are threatened by poaching (for their meat and scales) and heavy deforestation of their natural habitats, and are the most trafficked mammals in the world.^[4] Of the eight species of pangolin, four (Phataginus tetradactyla, P. tricuspis, Smutsia gigantea, and S. temminckii) are listed as vulnerable, two (Manis crassicaudata and M. culionensis) are listed as endangered, and two (M. pentadactyla and M. javanica) are listed as critically endangered on the International Union for Conservation of Nature Red List of Threatened Species.^[5]

Etymology

The name pangolin comes from the Malay word pengguling, meaning "one who rolls up".^[6] However, the modern name in Standard Malay is tenggiling, whereas in Indonesian it is trenggiling.

The etymologies of the three generic names Manis (Linnaeus, 1758), Phataginus (Rafinesque, 1821), and Smutsia (Gray, 1865) are sometimes misunderstood.

Carl Linnaeus (1758) invented the Neo-Latin generic name Manis apparently as a feminine singular form of the Latin masculine plural Manes, the Ancient Roman name for a type of spirit, after the animal's strange appearance.^[7]

Constantine Rafinesque (1821) formed the Neo-Latin generic name Phataginus from the French term phatagin, adopted by Count Buffon (1763) after the reported local name phatagin or phatagen used in the East Indies.

The British naturalist John Edward Gray named Smutsia for the South African naturalist Johannes Smuts (1808—1869),^[8][9] the first South African to do a treatise on mammals in 1832 (in which he described the species Manis temminckii).

Description

Pangolin skeletons at the Museum of Osteology (2009).

The physical appearance of a pangolin is marked by large hardened overlapping plate-like scales, which are soft on newborn pangolins, but harden as the animal matures.^[10] They are made of keratin, the same material from which human fingernails and tetrapod claws are made, and are structurally and compositionally very different from the scales of reptiles.^[11] The pangolin's scaled body is comparable in appearance to a pine cone. It can curl up into a ball when threatened, with its overlapping scales acting as armor, while it protects its face by tucking it under its tail. The scales are sharp, providing extra defense from predators.^[12]

Pangolins can emit a noxious-smelling chemical from glands near the anus, similar to the spray of a skunk.^[13] They have short legs, with sharp claws which they use for burrowing into ant and termite mounds, and climbing.^[14]

The tongues of pangolins are extremely long, like those of the giant anteater and the tube-lipped nectar bat; the root of the tongue is not attached to their hyoid bone, but is in the thorax^[15] between the sternum and the trachea. Large pangolins can extend their tongues as much as 40 cm (400 mm; 16 in), with a diameter of only 0.5 cm (5.0 mm; 0.20 in; 200 thou).^[16]

Behavior

Most pangolins are nocturnal animals that use their well-developed sense of smell to find insects. The long-tailed pangolin is also active by day, while other species of pangolins spend most of the daytime sleeping, curled up into a ball.^[16] They are considered to be shy and secretive creatures.^[17]

Arboreal pangolins live in hollow trees, whereas the ground-dwelling species dig tunnels underground, to a depth of 3.5 m (11 ft).^[16]

Some pangolins walk with their front claws bent under the foot pad, although they use the entire foot pad on their rear limbs. Furthermore, some exhibit a bipedal stance for some behaviour and may walk a few steps bipedally.^[18] Pangolins are also good swimmers.^[16]

Diet

Indian pangolin defending itself against Asiatic lions

Pangolins are insectivorous. Most of their diet consists of various species of ants and termites and may be supplemented by other insects, especially larvae. They are somewhat particular and tend to consume only one or two species of insects, even when many species are available to them. A pangolin can consume 140 to 200 g (4.9 to 7.1 oz) of insects per day.^[19] Pangolins are an important regulator of termite populations in their natural habitats.^[20]

Pangolins have a very poor sense of vision, so they rely heavily on smell and hearing. Pangolins also lack teeth, therefore they have evolved other physical characteristics to help them eat ants and termites. Their skeletal structure is sturdy and they have strong front legs that are useful for tearing into termite mounds.^[21] They use their powerful front claws to dig into trees, ground, and vegetation to find prey,^[22] then proceed to use their long tongues to probe inside the insect tunnels and to retrieve their prey.

The structure of their tongue and stomach is key to aiding pangolins in obtaining and digesting insects. Their saliva is sticky,^[21] causing ants and termites to stick to their long tongues when they are hunting through insect tunnels. Without teeth, pangolins also lack the ability to chew;^[23] however, while foraging, they ingest small stones (gastroliths) which accumulate in their stomachs to help to grind up ants.^[24] This part of their stomach is called the gizzard, and it is also covered in keratinous spines.^[25] These spines further aid in the grinding up and digestion of the pangolin's prey.

Some species, such as the tree pangolin, use their strong, prehensile tails to hang from tree branches and strip away bark from the trunk, exposing insect nests inside.^[26]

Reproduction

Pangolins are solitary and meet only to mate. Males are larger than females, weighing up to 40% more. While the mating season is not defined, they typically mate once each year, usually during the summer or autumn. Rather than the males seeking out the females, males mark their location with urine or feces and the females will find them. If there is competition over a female, the males will use their tails as clubs to fight for the opportunity to mate with her.^[27]

Gestation periods differ by species, ranging from roughly 70 to 140 days.^[28] African pangolin females usually give birth to a single offspring at a time, but the Asiatic species may give birth from one to three.^[16] Weight at birth is 80 to 450 g (2.8 to 15.9 oz) and the average length is 150 mm (5.9 in). At the time of birth, the scales are soft and white. After several days, they harden and darken to resemble those of an adult pangolin. During the vulnerable stage, the mother stays with her offspring in the burrow, nursing it, and wraps her body around it if she senses danger. The young cling to the mother's tail as she moves about, although in burrowing species, they remain in the burrow for the first two to four weeks of life. At one month, they first leave the burrow riding on the mother's back. Weaning takes place around three months of age, at which stage the young begin to eat insects in addition to nursing. At two years of age, the offspring are sexually mature and are abandoned by the mother.^[29]

Threats

A coat of armor made of gilded pangolin scales from India, an unusual object, was presented to George III in 1820

Pangolins are hunted and eaten in many parts of Africa and are one of the more popular types of bush meat, while local healers use the pangolin as a source of traditional medicine.^[30] They are also in great demand in southern China and Vietnam because their meat is considered a delicacy and some believe that pangolin scales have medicinal qualities.^{[31][32][33][34]} 100,000 are estimated to be trafficked a year to China and Vietnam,^[35] amounting to over one million over the past decade.^[36] This makes it the most trafficked animal in the world.^[36][37] This, coupled with deforestation, has led to a large decrease in the numbers of pangolins. Some species, such as Manis pentadactyla have become commercially extinct in certain ranges as a result of overhunting.^[38] In November 2010, pangolins were added to the Zoological Society of London's list of evolutionarily distinct and endangered mammals.^[39] All eight species of pangolin are classified by the IUCN as threatened with extinction, while two are classified as critically endangered.^[32][40] All pangolin species are currently listed under Appendix I of CITES which prohibits international trade, except when the product is intended for non-commercial purposes and a permit has been granted. ^[1]

Though pangolins are protected by an international ban on their trade, populations have suffered from illegal trafficking due to unfounded beliefs in East Asia that their ground-up scales can stimulate lactation or cure cancer or asthma.^[41] In the past decade, numerous seizures of illegally trafficked pangolin and pangolin meat have taken place in Asia.^{[42][43][44][45]} In one such incident in April 2013, 10,000 kg (11 short tons) of pangolin meat were seized from a Chinese vessel that ran aground in the Philippines.^[46][47] In another case in August 2016, an Indonesian man was arrested after police raided his home and found over 650 pangolins in freezers on his property.^[48] The same threat is reported in many countries in Africa, especially Nigeria, where the animal is on the verge of extinction due to overexploitation.^[49]

Conservation

As a result of increasing threats to pangolins, mainly in the form of illegal, international trade in pangolin skin, scales, and meat, these species have received increasing conservation attention in recent years. For example, in 2014, the IUCN recategorised all eight species of pangolin on its Red List of Threatened Species, and each species is now properly listed as being threatened with extinction.^[50] Also, the IUCN SSC Pangolin Specialist Group launched a global action plan to conserve pangolins, dubbed "Scaling up Pangolin Conservation" in July 2014. This action plan aims to improve all aspects of pangolin conservation with an added emphasis on combating poaching and trafficking of the animal, while educating communities in its importance.^[36] Another suggested approach to fighting pangolin (and general wildlife) trafficking consists in "following the money" rather than "the animal", which aims to disrupt smugglers' profits by interrupting money flows. Financial intelligence gathering could thus become a key tool in protecting these animals, although this opportunity is often overlooked.^[35] In 2018, a Chinese NGO launched the Counting Pangolins movement, calling for joint efforts to save the mammals from trafficking.^[51][52][53] Wildlife conservation group TRAFFIC has indentified 159 smuggling routes used by pangolin traffickers and aims to shut these down. ^[54]

Many attempts have been made to reproduce pangolins in captivity, but due to their reliance on wide-ranging habitats and very particular diets, these attempts are often unsuccessful.^[28] Pangolins have significantly decreased immune responses due to a genetic disfunction, making them extremely fragile.^[55] They are susceptible to diseases such as pneumonia and the development of ulcers in captivity, complications which can lead to an early death.^[28] In addition, pangolins rescued from illegal trade often have a higher chance of being infected with parasites such as intestinal worms, further lessening their chance for rehabilitation and reintroduction to the wild.^[28] Recently, researchers have been able to improve artificial pangolin habitats to allow for reproduction of pangolins, providing some hope for future reintroduction of these species into their natural habitats.^[10] The third Saturday in February is promoted as World Pangolin Day by the conservation NPO Annamiticus.^[56]

Taxonomy

Chinese pangolin

Pangolins were formerly^[when?] classified with various other orders of ant-eating mammals, most notably Xenarthra, which includes the true anteaters, sloths, and the armadillos which pangolins superficially resemble. Newer genetic evidence, however, indicates their closest living relatives are the Carnivora with which they form the clade Ferae.^[57][58] Some palaeontologists, placing Ernanodonta in a separate suborder of Cimolesta near Pholidota,^[59] have classified the pangolins in the order Cimolesta, together with several extinct groups indicated (†) below, though this idea has fallen out of favor since it was determined that cimolestids were not placental mammals.^[60] A 2015 study has supported close affinities between pangolins and the extinct group Creodonta, as well as many former Cimolestans (Cimolestes itself was recovered as a far more basal mammal).^[61]

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.^[3][62]

Ground pangolin in defensive posture

• 
  Order Pholidota sensu lato (Pholidotamorpha)^{[63][64][65][62]}
  
  
  • Suborder Palaeanodonta Matthew 1918
    
    
    
    • Genus †Arcticanodon Rose, Eberle & McKenna 2004
      
      • Species †Arcticanodon dawsonae Rose, Eberle & McKenna 2004
        
      
      
    
    
    • Genus †Melaniella Fox 1984
      
      • Species †Melaniella timosa Fox 1984
        
      
      
    
    
    • Genus †Molaetherium Heissig 1982
      
    
    
    • Genus †Amelotabes Rose 1978
      
      • Species †Amelotabes simpsoni Rose 1978
        
      
      
    
    
    • Family †Escavadodontidae Rose & Lucas 2000
      
      • Genus †Escavadodon Rose & Lucas 2000
        
        • Species †Escavadodon zygus Rose & Lucas 2000
          
        
        
      
      
    
    
    • Family †Epoicotheriidae Simpson 1927
      
      
      
      • Genus †Alocodontulum Rose, Bown & Simons 1978
        
      
      
      • Genus †Auroratherium Tong & Wamg 1997
        
        • Species †Auroratherium sinense Tong & Wamg 1997
          
        
        
      
      
      • Genus †Dipassalus Rose, Krishtalka & Stucky 1991
        
        • Genus †Dipassalus oryctes Rose, Krishtalka & Stucky 1991
          
        
        
      
      
      • Genus †Tetrapassalus Simpson 1959a
        
        
        
        • Species †T. mckennai Simpson 1959a
          
        
        
        • Species †T. proius West 1973c
          
        
        
        
        
      
      
      • Genus †Epoicotherium Simpson 1927 [Xenotherium Douglass 1906 non Ameghino 1904; Pseudochrysochloris Turnbull & Reed 1967]
        
        • Species †Epoicotherium unicum Douglass 1904 [Pseudochrysochloris yoderensis Turnbull & Reed 1967]
        
        
      
      
      • Genus †Tubulodon Jepsen 1932 [Pentapassalus Gazin 1952, Alocodon Rose et al. 1977]
        
        
        
        • Species †Tubulodon atopum Rose et al. 1978
          
        
        
        • Species †Tubulodon pearcei Gazin 1952
          
        
        
        • Species †Tubulodon taylori Jepsen 1932
          
        
        
        • Species †Tubulodon woodi Guthrie 1967
          
        
        
        
        
      
      
      • Genus †Xenocranium Colbert 1942
        
        • Species †Xenocranium pileorivale Colbert 1942
          
        
        
      
      
      
      
    
    
    • Family †Metacheiromyidae Wortman 1903
      
      
      
      • Genus †Propalaeanodon Rose 1979
        
        • Species †Propalaeanodon schaffi Rose 1979
          
        
        
      
      
      • Genus †Palaeanodon Matthew 1918
        
        
        
        • Species †P. parvulus Matthew 1918
          
        
        
        • Species †P. ignavus Matthew 1918
          
        
        
        • Species †P. nievelti Gingerich 1989
          
        
        
        
        
      
      
      • Genus †Brachianodon Gunnell & Gingerich 1993
        
        • Species †Brachianodon westorum Gunnell & Gingerich 1993
          
        
        
      
      
      • Genus †Mylanodon Secord et al. 2002
        
        • Species †Mylanodon rosei Secord et al. 2002
          
        
        
      
      
      • Genus †Metacheiromys Wortman 1903
        
        
        
        • Species †M. marshi Wortman 1903 [Metacheiromys tatusia Osborn 1904]
        
        
        • Species †M. dasypus Osborn 1904 [Metacheiromys osborni Simpson 1931]
        
        
        
        
      
      
      
      
    
    
    
    
  
  
  • Pholidota sensu stricto
    
    
    
    • Genus †Argyromanis patagonica Ameghino 1904
      
    
    
    • Genus †Orthoarthrus mixtus Ameghino 1904
      
    
    
    • Genus †Euromanis krebsi (Storch & Martin 1994) Gaudin, Emry & Wible 2009 [Eomanis krebsi Storch and Martin 1994]
    
    
    • Genus †Eurotamandua Storch 1981
      
    
    
    • Suborder Eupholidota Gaudin, Emry & Wible 2009
      
      
      
      • Family †Eomanidae Storch 2003
        
        • Genus †Eomanis waldi Storch 1978
          
        
        
      
      
      • Superfamily Manoidea
        
        
        
        • Family Patriomanidae Szalay & Schrenk 1998 sensu Gaudin, Emry & Pogue 2006
          
          
          
          • Genus †Cryptomanis gobiensis Gaudin, Emry & Pogue 2006
            
          
          
          • Genus †Patriomanis americana Emry 1970
            
          
          
          
          
        
        
        • Family Manidae Gray 1821
          
          
          
          • Genus †Necromanis Filhol 1893 [Leptomanis Filhol 1893; Necrodasypus Filhol 1893; Teutomanis Ameghino 1905; Galliaetatus Ameghino 1905]
            
            
            
            • Species †N. franconica (Quenstedt 1886) [Lutra franconica Quenstedt 1886]
            
            
            • Species †N. parva Koenigswald 1969
              
            
            
            • Species †N. quercyi Filhol 1893
              
            
            
            • Species †N. galliae
              
            
            
            
            
          
          
          • Subfamily Smutsiinae Gray 1873 (African pangolins)
            
            
            
            • Genus Phataginus Rafinesque 1821^[3] [Phataginus Rafinesque 1815 (nomen nudum); Triglochinopholis Fitzinger 1872; Uromanis Pocock 1924]
              
              
              
              • 
                Tree pangolin (P. tricuspis (Rafinesque 1821) Rafinesque 1821)
              
              
              • 
                Long-tailed pangolin (P. tetradactyla (Linnaeus 1766))
              
              
              
              
            
            
            • Genus Smutsia Gray 1865^[3] (African ground pangolins)
              
              
              
              • 
                Giant pangolin (S. gigantea (Illiger 1815))
              
              
              • 
                Ground pangolin (S. temmincki (Smuts 1832))
              
              
              
              
            
            
            
            
          
          
          • Subfamily Maninae Gray 1821
            
            • Genus Manis Linnaeus 1758 [Pholidotus Storr 1780; Quaggelo Frisch 1775; Pangolinus Rafinesque 1821; Pangolinus Rafinesque 1815 nomen nudum; Pangolin Gray 1873; Phatages Sundevall 1843] (Asiatic pangolins)
              
              
              
              • †M. hungarica Kormos 1934
                
              
              
              • †M. lydekkeri Trouessart
                
              
              
              • †Asian giant pangolin (M. paleojavanica Dubois)
              
              
              • Subgenus (Manis) Linnaeus 1758
                
                
                
                • 
                  Indian pangolin (M. crassicaudata Gray 1827)
                
                
                • 
                  Chinese pangolin (M. pentadactyla Linnaeus 1758)
                
                
                
                
              
              
              • Subgenus (Paramanis) Pocock 1924
                
                
                
                • 
                  Sunda pangolin (M. javanica Desmarest 1822)
                
                
                • 
                  Philippine pangolin (M. culionensis Elera 1915)
                
                
                
                
              
              
              
              
            
            
          
          
          
          
        
        
        
        
      
      
      
      
    
    
    
    
  
  
  
  

References

External links

Wikimedia Commons has media related to Pholidota.

Wikispecies has information related to Pholidota

Look up pangolin in Wiktionary, the free dictionary.

• IUCN SSC Pangolin Specialist Group


• 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)