Pangolins, also called scaly anteaters, are mammals of the order Pholidota (). The one extant family, the Manidae, has three genera: Manis, Phataginus, and Smutsia. Manis comprises five species found in Asia, while Phataginus and Smutsia include two species each, all found in sub-Saharan Africa. These species range in size from . Several extinct pangolin species are also known. In September 2023, a ninth species was reported, although it is only known from confiscated scales and has never been observed in the wild.
Pangolins have large, protective keratin scales, covering their skin. Depending on the species, they live in hollow trees or burrows. 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 they raise for about two years.
Pangolins are threatened by poaching (for their meat and scales, which are used in traditional medicine) and heavy deforestation of their natural habitats, and are also the most trafficked mammals in the world. , there are eight species of pangolin whose conservation status is listed in the threatened tier. Three (Manis culionensis, M. pentadactyla and M. javanica) are critically endangered, three (Phataginus tricuspis, Manis crassicaudata and Smutsia gigantea) are endangered and two (Phataginus tetradactyla and Smutsia temminckii) are vulnerable on the Red List of Threatened Species of the International Union for Conservation of Nature.
Etymology The name of order Pholidota comes from Ancient Greek – "clad in scales" from pholís "scale".
The name "pangolin" comes from the Malay word pengguling meaning "one who rolls up" from guling or giling "to roll"; it was used for the Sunda pangolin (Manis javanica). However, the modern name is tenggiling. In Javanese, it is terenggiling; and in the Philippine languages, it is goling, tanggiling, or balintong (with the same meaning).
In ancient India, according to Aelian, it was known as the phattáges (φαττάγης).Aelian, On Animals, 16:6 Loeb Classical Library, translated A. F. Scholfield 1959 vol. 3, pp. 266-268.
Description
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. 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. 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. Despite their appearance, they are not closely related to armadillos, having both evolved scales by convergent evolution.
Pangolins can emit a noxious-smelling chemical 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 ant and termite mounds and for climbing.
The tongues of pangolins are extremely long, and like those of the giant anteater and the tube-lipped nectar bat, the root of the tongue is not attached to the hyoid bone but is in the thorax between the sternum and the trachea. Large pangolins can extend their tongues as much as , with a diameter of only about .
Behaviour
Most pangolins are nocturnal animals which 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 ("volvation").
Arboreal pangolins live in hollow trees, whereas the ground-dwelling species dig tunnels to a depth of .
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 behavior, and may walk a few steps bipedally. Pangolins are also good swimmers.
Diet 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. A pangolin can consume of insects per day. Pangolins are an important regulator of termite populations in their natural habitats.
Pangolins have very poor vision. They also lack teeth. They rely heavily on smell and hearing, and they have other physical characteristics to help them eat ants and termites. Their skeletal structure is sturdy and they have strong front legs used for tearing into termite mounds. They use their powerful front claws to dig into trees, soil, and vegetation to find prey, 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, causing ants and termites to stick to their long tongues when they are hunting through insect tunnels. However, without teeth, pangolins cannot chew; so while foraging, they ingest small stones (gastroliths), which accumulate in their stomachs to help to grind up ants. This part of their stomach is called the gizzard, and it is also covered in keratinous spines. 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.
Reproduction
Pangolins are solitary and meet only to reproduce, with mating typically taking place at night after the male and female pangolin meet near a watering hole. 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 find them. If competition over a female occurs, the males use their tails as clubs to fight for the opportunity to mate with her.
Gestation periods differ by species, ranging from roughly 70 to 140 days. African pangolin females usually give birth to a single offspring at a time, but the Asiatic species may give birth to from one to three. Weight at birth is , and the average length is . 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, when 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.
Classification and phylogeny
Taxonomy Former classification:K. D. Rose and J. D. Archibold (2005) "The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades", Baltimore and London, Johns Hopkins University Press Current classification: Order: Pholidota (Weber, 1904) (pangolins) Family: Manidae (Gray, 1821) (living pangolins) Family: †Eomanidae (Storch, 2003) Family: †Patriomanidae (Szalay & Schrenk, 1998) Suborder: †Palaeanodonta (Matthew, 1918) (stem-pangolins) Family: †Epoicotheriidae (Simpson, 1927) Family: †Escavadodontidae Family: †Metacheiromyidae (Wortman, 1903) Incertae sedis: Genus: †Arcticanodon (Rose, 2004) Genus: †Melaniella (Fox, 1984) Suborder: †Ernanodonta Family: †Ernanodontidae Order: Pholidota (Weber, 1904) (pangolins) Suborder: Eupholidota (Gaudin, Emry & Wible, 2009) (true pangolins) Superfamily: Manoidea (Gaudin, Emry & Wible, 2009) Family: Manidae (Gray, 1821) (living pangolins) Family: †Patriomanidae (Szalay & Schrenk, 1998 [sensu Gaudin, Emry & Pogue, 2006]) Incertae sedis Genus: †Necromanis (Filhol, 1893) Superfamily: †Eomanoidea (Gaudin, Emry & Wible, 2009) Family: †Eomanidae (Storch, 2003) Incertae sedis: †Pholidota sp. (BC 16’08) (Pickford, 2008)
Summary of extant species Common nameBinomial namePopulationStatusTrendNotesImageCryptic pangolinManis mysteriaunknownIdentified via genetic analyses in 2023. Species has not been assessed by IUCN.Philippine pangolinManis culionensisunknownSunda pangolinManis javanicaunknownPopulation in Singapore is estimated to be 1,068 individuals. Thousands of this species are estimated to be illegally exported annually.Chinese pangolinManis pentadactyla45,000-120,000Values given are a very rough estimate given various, disjointed national estimates. Some taxonomists split another species, the Indo Burmese pangolin, from this species; IUCN currently retains it as a subspecies of M. pentadactyla.Indian pangolinManis crassicaudataunknownSpecies is heavily threatened by exportation for illegal wildlife trade.White-bellied pangolinPhataginus tricuspisunknownThe scales of >400,000 individuals were seized from illegal wildlife trade between 2015-2019.Giant ground pangolin (Giant pangolin)Smutsia giganteaunknownBlack-bellied pangolin (Long-tailed pangolin)Phataginus tetradactylaunknownTemminck's ground pangolin (Ground pangolin)Smutsia temminckiiunknownPopulation of this species over its South African range is estimated to be 16,329–24,102 mature individuals.
Phylogeny Among placentals
The order Pholidota was long considered to be the sister taxon to Xenarthra (neotropical anteaters, sloths, and armadillos), but recent genetic evidence indicates their closest living relatives are the carnivorans, with which they form a clade, the Ferae. Palaeanodonts are even closer relatives to pangolins, being classified with pangolins in the clade Pholidotamorpha. The split between carnivoran and pangolin lineages is estimated to have occurred 75.31 Ma (million years) ago.
Cladogram of pangolin phylogeny within order Pholidota Based on Kondrashov & Agadjanian (2012.) study: Based on Rose (2026.) study:
Among Manidae The first dichotomy in the phylogeny of extant Manidae separates Asian pangolins (Manis) from African pangolins (Smutsia and Phataginus). Within the former, Manis pentadactyla is the sister group to a clade comprising M. crassicaudata and M. javanica. Within the latter, a split separates the large terrestrial African pangolins of the genus Smutsia from the small arboreal African pangolins of the genus Phataginus.
Asian and African pangolins are thought to have diverged about 38.05 Ma ago. Moreover, the basal position of Manis within PholidotaSean P. Heighton, Rémi Allio, Jérôme Murienne, Jordi Salmona, Hao Meng, Céline Scornavacca, Armanda D. S. Bastos, Flobert Njiokou, Darren W. Pietersen, Marie-Ka Tilak, Shu-Jin Luo, Frédéric Delsuc, Philippe Gaubert (2023.) "Pangolin genomes offer key insights and resources for the world's most trafficked wild mammals" suggests the group originated in Eurasia, consistent with their laurasiatherian phylogeny.
Cladogram of living pangolins after Wangmo (2025.) study:
Threats