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)
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.
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimens with Sequences:155
Specimens with Barcodes:145
Species With Barcodes:6
|This article's lead section may not adequately summarize key points of its contents. (February 2015)|
The 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. These species range in size from 30 to 100 cm (12 to 39 in). A number of extinct species are also known. The name pangolin comes from the Malay word "pengguling", meaning "something that rolls up". It is found naturally in tropical regions throughout Africa and Asia.
Pangolins have large, protective keratin scales covering their skin. The pangolin is the only known mammal with this adaptation. 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, specially adapted 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 hunting (for their meat and armor) and heavy deforestation of their natural habitats. They are the most trafficked mammal in the world. All eight pangolin species are listed as threatened on the IUCN Red List of Threatened Species.
The physical appearance of a pangolin is marked by large, hardened, overlapping 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. 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 that are not attached 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).
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.
Pangolins are insectivorous. The bulk 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 will consume an average of 140 to 200 g (4.9 to 7.1 oz) of insects per day.
Pangolins have a very poor sense of vision, and therefore rely heavily on smell and hearing. After locating their prey, they tear open the anthills or termite mounds with their powerful front claws. Their front claws are so large that their anterior feet are not useful for walking. The animal uses its long tail to counterbalance its torso as it walks on its two hind legs. After tearing open the ant or termite mound, it uses its long tongue to probe inside the insect tunnels and retrieve its prey. They have glands in their chests to lubricate the tongue with sticky, ant-catching saliva. The tongue extends all the way into a cavity of the abdomen and is longer than the pangolin's entire body length. Pangolins lack teeth and, therefore, the ability to chew, however, they ingest small stones while foraging, which accumulate in the muscular stomach and help to grind up ants.
Pangolins are solitary and meet only to mate. Males are larger than females, weighing up to 50% more. While there is no defined mating season, they typically mate once each year, usually during the summer or autumn months. 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.
Gestation lasts for approximately 120–150 days. African pangolin females usually give birth to a single offspring at a time, but the Asiatic species may give birth from one to three. Weight at birth is 80 to 450 g (2.8 to 15.9 oz) and the average length is 6 inches (150 mm). 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 will wrap 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 at approximately 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.
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 and Vietnam because their meat is considered a delicacy and some 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. All eight species of pangolin are classified by the IUCN as threatened to extinction, while two are classified as critically endangered.
Though pangolin are protected by an international ban on their trade, populations have suffered from illegal trafficking due to unfounded 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 during 2013, 10,000 kilograms of pangolin meat was seized from a Chinese vessel that ran aground in the Philippines.
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 (International Union for the Conservation of Nature) re-categorised all eight species of pangolin on its Red List of Threatened Species (www.iucnredlist.org), and each species is now threatened with extinction. Also, the IUCN SSC Pangolin Specialist Group (www.pangolinsg.org) launched a global action plan to conserve pangolins, dubbed 'Scaling up Pangolin Conservation' in July 2014.
Pangolins were formerly classified with various other orders, for example Xenarthra, which includes the ordinary anteaters, sloths, and the similar-looking armadillos. Newer genetic evidence, however, indicates their closest living relatives are the Carnivora with which they form the clade Ferae. Some palaeontologists placed Ernanodonta in a separate suborder of Cimolesta near Pholidota, have classified the pangolins in the order Cimolesta, together with several extinct groups indicated (†) below, though this idea has fallen out of favor since cimolestids have been determined to have not been placental mammals.
- 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
- Tree pangolin (M. tricuspis)
- Subgenus Uromanis
- Long-tailed pangolin (M. tetradactyla)
- Subfamily †Eurotamanduinae
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