The eight species in the genus Manis are known as pangolins or scaly anteaters and live in Africa and Asia. These are the only members of the family Manidae, which is the only family in the Order Pholidota. The four African species are Manis (Uromanis) tetradactyla, M. (Phataginus) tricuspis, M. (Smutsia) gigantea, and M. (Smutsia) temmincki. The four Asian species are M. (Manis) crassicaudata, M. (Manis) pentadactyla, M. (Paramanis) culionensis, and M. (Paramanis) javanica. The indicated subgenera have sometimes been treated as distinct genera and M. culionensis, from the Philippines, has sometimes been treated as a synonym of M. javanica.
Pangolins have long, muscular tails (head and body length ranges from around 300 to 880 mm; tail length from 350 to 880 mm). All pangolins use their tails as powerful weapons to defend themselves and some pangolins may sometimes walk on just the hind legs, using the heavy tail as a brace. The tail and every outer surface of a pangolin is protected by horny scales, but the face, throat, belly, and inner limbs are naked or covered with ordinary mammalian hair. Three or four hairs are present at the base of each scale in the Asian species, but these hairs are not present in the African pangolins. "Pangolin" is a Malay name meaning "one that rolls up", which describes the defensive strategy employed by all pangolins of curling up into a tight ball, converting themselves into spheres of overlapping armor (one published report described a pangolin [M. javanica] curling itself into a ball and rolling rapidly down a slope!). The tubular skull provides extra protection as a result of its being made of thick, dense bone. All pangolins can travel on the ground, but arboreal species feed and sleep in rainforest trees. The much heavier terrestrial forms have dense massive scales reflecting their greater exposure to more formidable predators. Arboreal pangolins may weigh as little as 2 kg, whereas the terrestrial Giant Pangolin (S. gigantea) can exceed 30 kg.
Pangolin heads are relatively small, consisting largely of a nose with a small mouth. Pangolins have no teeth and cannot chew. All pangolins feed almost exclusively on termites and ants, which are captured with the sticky tongue then swallowed and ground up with tiny pebbles or sand in the hardened gizzard-like stomach. The worm-like tongue is as long as the head and body and when not extended folds back into a throat pocket that visibly bulges and empties as the animal feeds. The tongue is very sticky and associated with enormous salivary glands, requiring frequent drinking. The remarkable structure and attachment of the tongue allow it to travel down a hole in a termite mound, then whip back into the mouth covered in termites. The nostrils and ear openings can be closed and thick lids protect the eyes. All species have sharp claws on their forefeet, which they use to open termite mounds and to rip open hollow branches and trunks (they are reportedly also used by fighting males and possibly in defense). The hind limbs of pangolin species with different body sizes and habits are far more divergent than the forelimbs.
Despite their superficial resembance to the armadillos of the New World, pangolins and armadillos are not closely related. Their resemblance is due to parallel adaptations rather than recent common ancestry of the two groups.
In Africa, the arboreal Long-tailed Pangolin (M. tetradactyla) and Tree Pangolin (M. tricuspis) as well as the Giant Pangolin (M. gigantea) are closely tied to water; the Ground Pangolin (M. temmincki) is more arid-adapted, but only penetrates dry areas on the margins of its range in the Kalahari, Sudanic, and Somali arid regions.
Pangolins inhabit forests and thick bush as well as open or savannah country. Most species are nocturnal, but some are diurnal. Pangolins are mostly silent, although they may make hissing or puffing sounds. Male pangolins apparently defend scent-marked territories that enclose the home ranges of several females. Ground burrows are around 15 to 20 cm in diameter with a depth of several meters and terminating in a circular chamber as much as 2 m in circumference. Burrow entrances are generally closed with dirt when occupied. Giant Pangolin burrows may be as much as 5 m deep and 40 m long.
Young are born in burrows or arboreal hollows and subsequently are transported on their mother's tail. or back. African pangolins typically have just a single offspring at a time, but Asian species may have up to three.
Pangolin populations have suffered greatly from hunting for meat, skins, and scales, which are valued in traditional medicine in both Asia and Africa, as well as from habitat loss and possibly from high sensitivity to insecticides.
(Nowak 1991 and references therein; Kingdon 1997)
- Kingdon, J. 1997. The Kingdon Field Guide to African Mammals. Academic Press, San Diego, CA.
- Nowak, R.M. 1991. Walker's Mammals of the World, 5th edition. Volume 1. Johns Hopkins University Press, Baltimore.
Evolution and Systematics
Scales of pangolins provide flexibility yet strong protection by overlapping like roof shingles.
"Without teeth and without any turn of speed, the pangolin has to be well protected. It has an armour of horny scales that overlap like shingles on a roof. At the slightest danger the animal tucks its head into its stomach and wraps itself into a ball with its muscular tail clasped tight around it. In my experience, there is no way in which a pangolin, once rolled, can be forced to unwind." (Attenborough 1979:228)
Learn more about this functional adaptation.
- Attenborough, David. 1979. Life on Earth. Boston, MA: Little, Brown and Company. 319 p.
Molecular Biology and Genetics
Statistics of barcoding coverage
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A pangolin // (also referred to as a scaly anteater or trenggiling) is a mammal of the order Pholidota. The one extant family, Manidae, has one genus, Manis, which comprises eight species. A number of extinct species are also known. A pangolin has large keratin scales covering its skin, and is the only known mammal with this adaptation. It is found naturally in tropical regions throughout Africa and Asia. The name pangolin comes from the Malay word pengguling, meaning "something that rolls up".
The physical appearance of a pangolin is marked by large, hardened, plate-like scales. The scales, which are soft on newborn pangolins but harden as the animal matures, are made of keratin, the same material of which human fingernails and tetrapod claws are made. The pangolin's scaled body is comparable to a pine cone or globe artichoke. It can curl up into a ball when threatened, with its overlapping scales acting as armour and its face tucked under its tail. The scales are sharp, providing extra defense. The front claws are so long they are unsuited for walking, so the animal walks with its fore paws curled over to protect them. Pangolins can also emit a noxious-smelling acid from glands near the anus, similar to the spray of a skunk. Pangolins, though, are not able to spray this acid like skunks. They have short legs, with sharp claws which they use for burrowing into termite and ant mounds, as well as climbing.
The size of pangolins varies by species, ranging from 30 to 100 centimetres (12 to 39 in). Females are generally smaller than males.
The tongues of pangolins are extremely elongated and extend into the abdominal cavity. By convergent evolution, pangolins, the giant anteater, and the tube-lipped nectar bat all have tongues which are unattached to their hyoid bone and extend past their pharynx deep into the thorax. This extension lies between the sternum and the trachea. Large pangolins can extend their tongues as much as 40 centimetres (16 in), with a diameter of only 0.5 centimetres (0.20 in).
Pangolins are nocturnal animals which use their well-developed sense of smell to find insects. The long-tailed pangolin is also active by day. Other species of pangolins spend most of the daytime sleeping curled up into a ball.
Pangolins lack teeth and the ability to chew. Instead, they tear open anthills or termite mounds with their powerful front claws and probe deep into them with their very long tongues. Pangolins have glands in their chests to lubricate the tongue with sticky, ant-catching saliva.
Gestation is 120–150 days. African pangolin females usually give birth to a single offspring at a time, but the Asiatic species can give birth from one to three. Weight at birth is 80–450 g (3–18 ounces), and the scales are initially soft. 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. Weaning takes place at around three months of age, and pangolins become sexually mature at two years.
Pangolins are hunted and eaten in many parts of Africa, and are one of the more popular types of bush meat. They are also in great demand in China because their meat is considered a delicacy and some Chinese believe pangolin scales have medicinal qualities. This, coupled with deforestation, has led to a large decrease in the numbers of giant pangolins. In November 2010, pangolins were added to the Zoological Society of London's list of genetically distinct and endangered mammals. Two species of pangolin are classified by the IUCN as Endangered species.
Though pangolin are protected by an international ban on their trade, populations have suffered from illegal trafficking due to beliefs in Asia that their ground-up scales can stimulate lactation or cure cancer or asthma. In the past decade there have been numerous seizures of illegally trafficked pangolin and pangolin meat in Asia. In one such incident in 2013, 10,000 kilograms of pangolin meat was seized from a Chinese vessel that ran aground in the Philippines.
The Guardian provided a description of the killing and eating of pangolins: "A Guangdong chef interviewed last year in the Beijing Science and Technology Daily described how to prepare a pangolin: 'We keep them alive in cages until the customer makes an order. Then we hammer them unconscious, cut their throats and drain the blood. It is a slow death. We then boil them to remove the scales. We cut the meat into small pieces and use it to make a number of dishes, including braised meat and soup. Usually the customers take the blood home with them afterwards.'"
Pangolins were classified with various other orders, for example Xenarthra, which includes the ordinary anteaters, sloths, and the similar-looking armadillos. But newer genetic evidence indicates their closest living relatives are the Carnivora with which they form the clade, Ferae. Some palaeontologists have classified the pangolins in the order Cimolesta, together with several extinct groups indicated (†) below.
- ORDER PHOLIDOTA
- Family †Epoicotheriidae
- Family †Metacheiromyidae
- Family Manidae
- Subfamily †Eurotamanduinae
- Genus †Eurotamandua
- Subfamily Maninae
- Genus †Cryptomanis
- Genus †Eomanis
- Genus †Necromanis
- Genus †Patriomanis
- Genus Manis
- Subgenus Manis
- Subgenus Paramanis
- Subgenus Smutsia
- Subgenus Phataginus
- Subgenus Uromanis
- Subfamily †Eurotamanduinae
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- M. javanica, IUCN Red List
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- 23 tonnes of pangolins seized in a week – traffic.org
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- BioMed Central | Full text | A higher-level MRP supertree of placental mammals
- For example, McKenna & Bell 1997, p. 222 placed Ernanodonta in a separate suborder of Cimolesta near Pholidota, in which they included palaeanodonts. (Rose 2006, p. 210)