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Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (). The one
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
family, the
Manidae Manidae is the only extant family of pangolins from superfamily Manoidea. This family comprises three genera ('' Manis'' from subfamily Maninae, ''Phataginus'' from subfamily Phatagininae, and ''Smutsia'' from subfamily Smutsiinae), as well as ex ...
, has three genera: ''
Manis ''Manis'' is a genus of South Asian and East Asian pangolins, the Asiatic pangolins, from subfamily Maninae, within family Manidae. Etymology Carl Linnaeus (1758) invented the Neo-Latin generic name ''Manis'' apparently as a feminine singular ...
'', ''
Phataginus African tree pangolin (''Phataginus'') is a genus of African pangolins from subfamily small African pangolins (Phatagininae), within family Manidae. Its members are the more arboreal of the African pangolins. The number of illegally trafficked ...
'', and ''
Smutsia ''Smutsia'' is a genus of African pangolins, better known as the African ground pangolins, from subfamily Smutsiinae, within family Manidae. It was formerly considered a subgenus of '' Manis''. Its members are the more terrestrial of the African ...
''. ''Manis'' comprises of four species found in Asia, while ''Phataginus'' and ''Smutsia'' include two species each, all found in sub-Saharan Africa. These species range in size from . A number of
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
pangolin species are also known. Pangolins have large, protective
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
scales, similar in material to fingernails and toenails, covering their skin; they are the only known mammals with this feature. They live in hollow trees or
burrow An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of sh ...
s, depending on the species. Pangolins are
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
, and their diet consists of mainly ants and
termites Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
, 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 Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
) and heavy
deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated d ...
of their natural habitats, and they are the most
trafficked ''Trafficked'' is a 2017 American thriller drama film directed by Will Wallace and starring Ashley Judd, Sean Patrick Flanery and Anne Archer. Plot In California, Sara is eighteen and has to leave her foster home; she is offered training to be a ...
mammals in the world. , there are eight species of pangolin whose
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservation ...
is listed in the
threatened Threatened species are any species (including animals, plants and fungi) which are vulnerable to endangerment in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of ''critical depensat ...
tier. Three (''
Manis culionensis The Philippine pangolin or Palawan pangolin (''Manis culionensis''), also locally known as ''balintong'', is a pangolin species endemic to the Palawan province of the Philippines. Its habitat includes primary and secondary forests, as well as sur ...
'', '' M. pentadactyla'' and '' M. javanica'') are critically endangered, three (''
Phataginus tricuspis The tree pangolin (''Phataginus tricuspis'') is one of eight extant species of pangolins ("scaly anteaters"), and is native to equatorial Africa. Also known as the white-bellied pangolin or three-cusped pangolin, it is the most common of the A ...
'', ''
Manis crassicaudata The Indian pangolin (''Manis crassicaudata''), also called thick-tailed pangolin and scaly anteater is a pangolin native to the Indian subcontinent. Like other pangolins, it has large, overlapping scales on its body which act as armour. The colou ...
'' and '' Smutsia gigantea'') are endangered and two (''
Phataginus tetradactyla The long-tailed pangolin (''Phataginus tetradactyla''), also called the African black-bellied pangolin, or ''ipi'', is a diurnal, Wiktionary:arboreal#Adjective, arboreal pangolin species belonging to the family Manidae, in the order Pholidota. T ...
'' and ''
Smutsia temminckii The ground pangolin (''Smutsia temminckii''), also known as Temminck's pangolin, Cape pangolin or steppe pangolin, is one of four species of pangolins which can be found in Africa, and the only one in southern and eastern Africa. The animal was n ...
'') are vulnerable on the
Red List of Threatened Species The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biologi ...
of the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
.


Etymology

The name "pangolin" comes from the
Malay Malay may refer to: Languages * Malay language or Bahasa Melayu, a major Austronesian language spoken in Indonesia, Malaysia, Brunei and Singapore ** History of the Malay language, the Malay language from the 4th to the 14th century ** Indonesi ...
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 The Philippine languages or Philippinic are a proposed group by R. David Paul Zorc (1986) and Robert Blust (1991; 2005; 2019) that include all the languages of the Philippines and northern Sulawesi, Indonesia—except Sama–Bajaw (languages ...
, it is ''goling'', ''tanggiling'', or ''balintong'' (with the same meaning). In ancient India, according to
Aelian Aelian or Aelianus may refer to: * Aelianus Tacticus, Greek military writer of the 2nd century, who lived in Rome * Casperius Aelianus, Praetorian Prefect, executed by Trajan * Claudius Aelianus, Roman writer, teacher and historian of the 3rd centu ...
, it was known as the ''phattáge'' (φαττάγης).


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 Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
, the same material from which human fingernails and
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
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. Pangolins can emit a noxious-smelling chemical from
gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...
s near the anus, similar to the spray of a
skunk Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or ginge ...
. They have short legs, with sharp claws which they use for burrowing into ant and
termite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
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 The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
between the sternum and the
trachea The trachea, also known as the windpipe, is a Cartilage, cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends ...
. Large pangolins can extend their tongues as much as , with a diameter of only about .


Behavior

Most pangolins are
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
animals which use their well-developed sense of smell to find insects. The
long-tailed pangolin The long-tailed pangolin (''Phataginus tetradactyla''), also called the African black-bellied pangolin, or ''ipi'', is a diurnal, arboreal pangolin species belonging to the family Manidae, in the order Pholidota. They feed on ants rather tha ...
is also active by day, while other species of pangolins spend most of the daytime sleeping, curled up into a ball ("
volvation Volvation (from Latin ''volvere'' "roll", and the suffix ''-(a)tion''; sometimes called enrollment or conglobation), is a defensive behavior in certain animals, in which the animal rolls its own body into a ball, presenting only the hardest parts ...
"). 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 behaviour, and may walk a few steps bipedally. Pangolins are also good swimmers.


Diet

Pangolins are
insectivorous A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores were ...
. 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 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 Smell may refer to; * Odor, airborne molecules perceived as a scent or aroma * Sense of smell, the scent also known scientifically as olfaction * "Smells" (''Bottom''), an episode of ''Bottom'' * The Smell, a music venue in Los Angeles, Californ ...
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 that are 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 Saliva (commonly referred to as spit) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which DNA can be ...
is sticky, 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; but while foraging, they ingest small stones (
gastroliths A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In oth ...
), 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 The tree pangolin (''Phataginus tricuspis'') is one of eight extant species of pangolins ("scaly anteaters"), and is native to equatorial Africa. Also known as the white-bellied pangolin or three-cusped pangolin, it is the most common of the ...
, use their strong,
prehensile Prehensility is the quality of an appendage or organ (anatomy), organ that has Adaptation (biology), adapted for grasping or holding. The word is derived from the Latin term ''prehendere'', meaning "to grasp". The ability to grasp is likely der ...
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 mate, 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 Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregna ...
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 2–4 weeks of life. At one month, they first leave the burrow riding on the mother's back. Weaning takes place around 3 months of age, when the young begin to eat insects in addition to nursing. At 2 years of age, the offspring are sexually mature and are abandoned by the mother.


Classification and phylogeny


Taxonomy

The order name Pholidota comes from
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
ϕολιδωτός – "clad in scales" from ''pholís'' "
scale Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number ...
". * Order: Pholidota (Weber, 1904) (pangolins) ** Genus:
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species). It is one of the modern descendan ...
'' Euromanis'' (Gaudin, Emry & Wible, 2009) *** †''Euromanis krebsi'' (Storch & Martin, 1994) ** Family: † Eurotamanduidae (Szalay & Schrenk, 1994) *** Genus: †'' Eurotamandua'' (Storch, 1981) **** †''Eurotamandua joresi'' (Storch, 1981) ** '' Incertae sedis'' *** †Pholidota sp. (BC 16’08) ** Suborder:
Eupholidota Eupholidota ("true pangolins") is a suborder of pangolins that includes two superfamilies: extant Manoidea and extinct Eomanoidea. Taxonomy * Suborder: Eupholidota (true pangolins) ** Superfamily: Manoidea *** Family: Manidae (pangolins) *** ...
(Gaudin, Emry & Wible, 2009) (true pangolins) *** Superfamily: † Eomanoidea (Gaudin, Emry & Wible, 2009) **** Family: † Eomanidae (Storch, 2003) ***** Genus: †'' Eomanis'' (Storch, 1978) ****** †''Eomanis waldi'' (Storch, 1978) *** Superfamily:
Manoidea Manoidea is a superfamily of pangolins from suborder Eupholidota that includes extant family Manidae, extinct family Patriomanidae and extinct genus ''Necromanis''. Taxonomy * Superfamily: Manoidea ** Family: Manidae (pangolins) ** Family: ...
(Gaudin, Emry & Wible, 2009) **** Family:
Manidae Manidae is the only extant family of pangolins from superfamily Manoidea. This family comprises three genera ('' Manis'' from subfamily Maninae, ''Phataginus'' from subfamily Phatagininae, and ''Smutsia'' from subfamily Smutsiinae), as well as ex ...
(Gray, 1821) (pangolins) ***** Subfamily: Maninae (Gray, 1821) (Asiatic pangolins) ****** Genus: ''
Manis ''Manis'' is a genus of South Asian and East Asian pangolins, the Asiatic pangolins, from subfamily Maninae, within family Manidae. Etymology Carl Linnaeus (1758) invented the Neo-Latin generic name ''Manis'' apparently as a feminine singular ...
'' (Linnaeus, 1758) (Asiatic pangolins) ******* ''
Manis crassicaudata The Indian pangolin (''Manis crassicaudata''), also called thick-tailed pangolin and scaly anteater is a pangolin native to the Indian subcontinent. Like other pangolins, it has large, overlapping scales on its body which act as armour. The colou ...
'' (Gray, 1827) (Indian pangolin) ******* ''
Manis pentadactyla The Chinese pangolin (''Manis pentadactyla'') is a pangolin native to the northern Indian subcontinent, northern parts of Southeast Asia and southern China. It has been listed as Critically Endangered on the IUCN Red List since 2014, as the wil ...
'' (Linnaeus, 1758) (Chinese pangolin) ******* ''Manis'' sp. (Scale_H4 & Scale_H8) ******* †'' Manis hungarica'' (Kormos, 1934) ******* †'' Manis lydekkeri'' (Dubois, 1908) ******* Subgenus: ''
Paramanis ''Manis'' is a genus of South Asian and East Asian pangolins, the Asiatic pangolins, from subfamily Maninae, within family Manidae. Etymology Carl Linnaeus (1758) invented the Neo-Latin generic name ''Manis'' apparently as a feminine sing ...
'' (Pocock, 1924) ******** ''
Manis javanica The Sunda pangolin (''Manis javanica''), also known as the Malayan or Javan pangolin, is a species of pangolin. It is found throughout Southeast Asia, including Myanmar, Thailand, Cambodia, Laos, Malaysia, Singapore, Vietnam, and the islands of ...
'' (Desmarest, 1822) (Sunda pangolin) ******** ''
Manis culionensis The Philippine pangolin or Palawan pangolin (''Manis culionensis''), also locally known as ''balintong'', is a pangolin species endemic to the Palawan province of the Philippines. Its habitat includes primary and secondary forests, as well as sur ...
'' (de Elera, 1895) (Philippine pangolin) ******** †'' Manis palaeojavanica'' (Dubois, 1907) (Giant asian pangolin) ***** Subfamily: Phatagininae (Gaubert, 2017) (small African pangolins) ****** Genus: ''
Phataginus African tree pangolin (''Phataginus'') is a genus of African pangolins from subfamily small African pangolins (Phatagininae), within family Manidae. Its members are the more arboreal of the African pangolins. The number of illegally trafficked ...
'' (Rafinesque, 1821) (African tree pangolins) ******* ''
Phataginus tetradactyla The long-tailed pangolin (''Phataginus tetradactyla''), also called the African black-bellied pangolin, or ''ipi'', is a diurnal, Wiktionary:arboreal#Adjective, arboreal pangolin species belonging to the family Manidae, in the order Pholidota. T ...
'' (Linnaeus, 1766) (Long-tailed pangolin) ******* ''
Phataginus tricuspis The tree pangolin (''Phataginus tricuspis'') is one of eight extant species of pangolins ("scaly anteaters"), and is native to equatorial Africa. Also known as the white-bellied pangolin or three-cusped pangolin, it is the most common of the A ...
'' (Rafinesque, 1821) (Tree pangolin) ***** Subfamily: Smutsiinae (Gray, 1873) (large African pangolins) ****** Genus: ''
Smutsia ''Smutsia'' is a genus of African pangolins, better known as the African ground pangolins, from subfamily Smutsiinae, within family Manidae. It was formerly considered a subgenus of '' Manis''. Its members are the more terrestrial of the African ...
'' (Gray, 1865) (African ground pangolins) ******* '' Smutsia gigantea'' (Illiger, 1815) (Giant pangolin) ******* '' Smutsia temmincki'' (Smuts, 1832) (Ground pangolin) ******* †'' Smutsia olteniensis'' (Terhune, 2021) ***** '' Incertae sedis'' ****** †
Fayum pangolin Faiyum ( ar, الفيوم ' , borrowed from cop,  ̀Ⲫⲓⲟⲙ or Ⲫⲓⲱⲙ ' from egy, pꜣ ym "the Sea, Lake") is a city in Middle Egypt. Located southwest of Cairo, in the Faiyum Oasis, it is the capital of the modern Faiyum ...
(Manidae sp. 'DPC 3972'' & ''DPC 4364''(Gebo & Rasmussen, 1985)) **** Family: †
Patriomanidae Patriomanidae ("fathers of pangolins") is a extinct family of pangolins from superfamily Manoidea that includes two extinct genera ''Patriomanis'' and '' Cryptomanis''. Taxonomy * Family: †Patriomanidae ** Genus: †'' Cryptomanis'' *** ...
(Szalay & Schrenk 1998) 'sensu'' Gaudin, Emry & Pogue, 2006/small> ***** Genus: †'' Cryptomanis'' (Gaudin, Emry & Pogue, 2006) ****** †''Cryptomanis gobiensis'' (Gaudin, Emry & Pogue, 2006) ***** Genus: †''
Patriomanis ''Patriomanis'' ("father of pangolins") is an extinct genus of pangolin from extinct family Patriomanidae. It lived during the Eocene of North America and it currently represents the only pangolin known from the Western Hemisphere. The genus co ...
'' (Emry, 1970) ****** †''Patriomanis americana'' (Emry, 1970) **** '' Incertae sedis'' ***** Genus: †''
Necromanis ''Necromanis'' ("extinct pangolin") is an extinct genus of pangolin from superfamily Manoidea. It lived during the Oligocene and Miocene of Europe. It was originally placed within family Manidae, but was eventually removed from it as more f ...
'' (Filhol, 1893) ****** †''Necromanis franconica'' (Quenstedt, 1886) ****** †''Necromanis parva'' (Koenigswald, 1969) ****** †''Necromanis quercyi'' (Filhol, 1893)


Phylogeny


Among placentals

The order Pholidota was considered to be the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to Xenarthra (neotropical
anteater Anteater is a common name for the four extant mammal species of the suborder Vermilingua (meaning "worm tongue") commonly known for eating ants and termites. The individual species have other names in English and other languages. Together with ...
s,
sloth Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their li ...
s, and
armadillo Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, along wi ...
s), but recent genetic evidence indicates their closest living relatives are the
carnivora Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...
ns, with which they form a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
termed either
Ferae Ferae ( , , "wild beasts") is a mirorder of placental mammalsMalcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'' in Columbia University Press, New York (1997), 631 Seiten. that groups together clades Pan-Ca ...
or Ostentoria. Fossil groups like the
creodont Creodonta ("meat teeth") is a former order of extinct carnivorous placental mammals that lived from the early Paleocene to the late Miocene epochs in North America, Europe, Asia and Africa. Originally thought to be a single group of animals ance ...
s and palaeanodonts are even closer relatives to pangolins (the latter group being classified with pangolins in the clade Pholidotamorpha). The split between carnivorans and pangolins is estimated to have occurred 79–87 Ma (million years) ago.


Among Manidae

The first dichotomy in the
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
of extant Manidae separates Asian pangolins (''Manis'') from African pangolins (''Smutsia'' and ''Phataginus''). Within the former, ''Manis pentadactyla'' is the
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
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 38–47 Ma ago. Moreover, the
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
position of ''Manis'' within Pholidota suggests the group originated in Eurasia, consistent with their laurasiatherian phylogeny.


Threats

Pangolins are in high demand for
traditional Chinese A tradition is a belief or behavior (folk custom) passed down within a group or society with symbolic meaning or special significance with origins in the past. A component of cultural expressions and folklore, common examples include holidays or ...
and Vietnamese medicine in southern China and
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
because their scales are believed to have medicinal properties. Their meat is also considered a delicacy. 100,000 are estimated to be trafficked a year to China and Vietnam, amounting to over one million over the past decade. This makes them the most trafficked animal in the world. This, coupled with
deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated d ...
, has led to a large decrease in the numbers of pangolins. Some species, such as ''
Manis pentadactyla The Chinese pangolin (''Manis pentadactyla'') is a pangolin native to the northern Indian subcontinent, northern parts of Southeast Asia and southern China. It has been listed as Critically Endangered on the IUCN Red List since 2014, as the wil ...
'' have become commercially extinct in certain ranges as a result of overhunting. In November 2010, pangolins were added to the
Zoological Society of London The Zoological Society of London (ZSL) is a charity devoted to the worldwide conservation of animals and their habitats. It was founded in 1826. Since 1828, it has maintained the London Zoo, and since 1931 Whipsnade Park. History On 29 ...
's list of evolutionarily distinct and endangered mammals. All eight species of pangolin are assessed as threatened by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
, while three are classified as critically endangered. All pangolin species are currently listed under Appendix I of
CITES CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
which prohibits international trade, except when the product is intended for non-commercial purposes and a permit has been granted. China had been the main destination country for pangolins until 2018, when it was reportedly surpassed by Vietnam. In 2019, Vietnam was reported to have seized the largest volumes of pangolin scales, surpassing Nigeria that year. Pangolins are also hunted and eaten in Ghana and are one of the more popular types of
bushmeat Bushmeat is meat from wildlife species that are hunted for human consumption, most often referring to the meat of game in Africa. Bushmeat represents a primary source of animal protein and a cash-earning commodity for inhabitants of humid tropi ...
, while local healers use the pangolin as a source of traditional medicine. Though pangolins are protected by an international ban on their trade, populations have suffered from illegal trafficking due to beliefs in East Asia that their ground-up scales can stimulate
lactation Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The proces ...
or cure cancer or
asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, cou ...
. In the past decade, numerous seizures of illegally trafficked pangolin and pangolin meat have taken place in Asia. In one such incident in April 2013, of pangolin meat were seized from a Chinese vessel that ran aground in the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
. 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. The same threat is reported in
Nigeria Nigeria ( ), , ig, Naìjíríyà, yo, Nàìjíríà, pcm, Naijá , ff, Naajeeriya, kcg, Naijeriya officially the Federal Republic of Nigeria, is a country in West Africa. It is situated between the Sahel to the north and the Gulf o ...
, where the animal is on the verge of extinction due to
overexploitation Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term app ...
. The overexploitation comes from hunting pangolins for game meat and the reduction of their forest habitats due to deforestation caused by
timber harvesting Logging is the process of cutting, processing, and moving trees to a location for transport. It may include skidding, on-site processing, and loading of trees or logs onto trucks or skeleton cars. Logging is the beginning of a supply chain ...
. The pangolin are hunted as game meat for both medicinal purposes and food consumption.


Virology


COVID-19 infection

The
nucleic acid sequence A nucleic acid sequence is a succession of Nucleobase, bases signified by a series of a set of five different letters that indicate the order of nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. By convention, sequence ...
of a specific receptor-binding domain of the spike protein belonging to
coronavirus Coronaviruses are a group of related RNA viruses that cause diseases in mammals and birds. In humans and birds, they cause respiratory tract infections that can range from mild to lethal. Mild illnesses in humans include some cases of the com ...
es taken from pangolins was found to be a 99% match with SARS coronavirus 2 (
SARS-CoV-2 Severe acute respiratory syndrome coronavirus 2 (SARS‑CoV‑2) is a strain of coronavirus that causes COVID-19 (coronavirus disease 2019), the respiratory illness responsible for the ongoing COVID-19 pandemic. The virus previously had a ...
), the virus which causes
COVID-19 Coronavirus disease 2019 (COVID-19) is a contagious disease caused by a virus, the severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2). The first known case was COVID-19 pandemic in Hubei, identified in Wuhan, China, in December ...
and is responsible for the
COVID-19 pandemic The COVID-19 pandemic, also known as the coronavirus pandemic, is an ongoing global pandemic of coronavirus disease 2019 (COVID-19) caused by severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2). The novel virus was first identif ...
. Researchers in
Guangzhou Guangzhou (, ; ; or ; ), also known as Canton () and alternatively romanized as Kwongchow or Kwangchow, is the capital and largest city of Guangdong province in southern China. Located on the Pearl River about north-northwest of Hong Kon ...
, China, hypothesized that SARS-CoV-2 had originated in bats, and prior to infecting humans, was circulating among pangolins. The illicit Chinese trade of pangolins for use in
traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of action ...
was suggested as a vector for human transmission. However, whole-genome comparison found that the pangolin and human
coronaviruses Coronaviruses are a group of related RNA viruses that cause diseases in mammals and birds. In humans and birds, they cause respiratory tract infections that can range from mild to lethal. Mild illnesses in humans include some cases of the com ...
share only up to 92% of their RNA. Ecologists worried that the early speculation about pangolins being the source may have led to mass slaughters, endangering them further, which was similar to what happened to Asian palm civets during the SARS outbreak. It was later proved that the testing which suggested that pangolins were a potential host for the virus was flawed, when
genetic analysis Genetic analysis is the overall process of studying and researching in fields of science that involve genetics and molecular biology. There are a number of applications that are developed from this research, and these are also considered parts of ...
showed that the spike protein and its binding to receptors in pangolins had minimal effect from the virus, and therefore were not likely mechanisms for COVID-19 infections in humans.


''Pestivirus'' and ''Coltivirus''

In 2020, two novel RNA viruses distantly related to
pestivirus ''Pestivirus'' is a genus of viruses, in the family ''Flaviviridae''. Viruses in the genus ''Pestivirus'' infect mammals, including members of the family Bovidae (which includes cattle, sheep, and goats) and the family Suidae (which includes var ...
es and
coltivirus Coltivirus is a genus of viruses (belonging to the ''Reoviridae'' family) that infects vertebrates and invertebrates. It includes the causative agent of Colorado tick fever. Colorado tick fever virus can cause a fever, chills, headache, photopho ...
es have been detected in the genomes of dead ''Manis javanica'' and ''Manis pentadactyla''. To refer to both sampling site and hosts, they were named Dongyang pangolin virus (DYPV) and Lishui pangolin virus (LSPV). The DYPV pestivirus was also identified in ''
Amblyomma javanense ''Amblyomma javanense'' is a hard-bodied tick of the genus ''Amblyomma''. It is found in Vietnam, Indonesia, Malaysia, India, Sri Lanka, Philippines and Thailand Thailand ( ), historically known as Siam () and officially the Kingdom ...
'' nymph
tick Ticks (order Ixodida) are parasitic arachnids that are part of the mite superorder Parasitiformes. Adult ticks are approximately 3 to 5 mm in length depending on age, sex, species, and "fullness". Ticks are external parasites, living by ...
s from a diseased pangolin.


Folk medicine

Pangolin scales and flesh are used as ingredients for various
traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of action ...
preparations. While no
scientific evidence Scientific evidence is evidence that serves to either support or counter a scientific theory or hypothesis, although scientists also use evidence in other ways, such as when applying theories to practical problems. "Discussions about empirical ev ...
exists for the efficacy of those practices, and they have no logical mechanism of action, their popularity still drives the
black market A black market, underground economy, or shadow economy is a clandestine market or series of transactions that has some aspect of illegality or is characterized by noncompliance with an institutional set of rules. If the rule defines the se ...
for animal body parts, despite concerns about toxicity, transmission of diseases from animals to humans, and species extermination. The ongoing demand for parts as ingredients continues to fuel pangolin poaching, hunting and trading. In the 21st century, the main uses of pangolin scales are
quackery Quackery, often synonymous with health fraud, is the promotion of fraudulent or ignorant medical practices. A quack is a "fraudulent or ignorant pretender to medical skill" or "a person who pretends, professionally or publicly, to have skill, ...
practices based on unproven claims the scales dissolve
blood clot A thrombus (plural thrombi), colloquially called a blood clot, is the final product of the blood coagulation step in hemostasis. There are two components to a thrombus: aggregated platelets and red blood cells that form a plug, and a mesh of c ...
s, promote blood circulation, or help lactating women secrete milk. The supposed health effects of pangolin meat and scales claimed by folk medicine practitioners and quacks are based on their consumption of ants, long tongues, and protective scales. The Chinese name ''chuan shan jia'' () "penetrating-the-mountain scales") emphasizes the idea of penetration or passing through even massive obstructions such as mountains, plus the distinctive scales which embody penetration and protection. The official
pharmacopoeia A pharmacopoeia, pharmacopeia, or pharmacopoea (from the obsolete typography ''pharmacopœia'', meaning "drug-making"), in its modern technical sense, is a book containing directions for the identification of compound medicines, and published by ...
of the People's Republic of China included Chinese pangolin scales as an ingredient in TCM formulations. Pangolins were removed from the pharmacopoeia starting from the first half of 2020. Although pangolin scales have been removed from the list of raw ingredients, the scales are still listed as a key ingredient in various medicines. The first record of pangolin scales occurs in ''Ben Cao Jinji Zhu'' ("Variorum of Shennong's Classic of Materia Medica", 500 CE), which recommends pangolin scales for protection against ant bites; burning the scales as a cure for people crying hysterically during the night. During the
Tang dynasty The Tang dynasty (, ; zh, t= ), or Tang Empire, was an Dynasties in Chinese history, imperial dynasty of China that ruled from 618 to 907 AD, with an Zhou dynasty (690–705), interregnum between 690 and 705. It was preceded by the Sui dyn ...
, a recipe for expelling evil spirits with a formulation of scales, herbs, and minerals appeared in 682, and in 752 CE the idea that pangolin scales could also stimulate milk secretion in lactating women, one of the main uses today, was recommended in the ''Wai Tai Mi Yao'' ("Arcane Essentials from the Imperial Library"). In the
Song dynasty The Song dynasty (; ; 960–1279) was an imperial dynasty of China that began in 960 and lasted until 1279. The dynasty was founded by Emperor Taizu of Song following his usurpation of the throne of the Later Zhou. The Song conquered the rest ...
, the notion of penetrating and clearing blockages was emphasized in the ''Taiping sheng hui fan'' ("Formulas from Benevolent Sages Compiled During the Era of Peace and Tranquility"), compiled by Wang Huaiyin in 992.


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. , the IUCN considered all eight species of pangolin on its
Red List of Threatened Species The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biologi ...
as threatened. 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 on its importance. 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. In 2018, a Chinese NGO launched the Counting Pangolins movement, calling for joint efforts to save the mammals from trafficking. Wildlife conservation group
TRAFFIC Traffic comprises pedestrians, vehicles, ridden or herded animals, trains, and other conveyances that use public ways (roads) for travel and transportation. Traffic laws govern and regulate traffic, while rules of the road include traffic ...
has identified 159 smuggling routes used by pangolin traffickers and aims to shut these down. Many attempts have been made to breed pangolins in captivity, but due to their reliance on wide-ranging habitats and very particular diets, these attempts are often unsuccessful. Pangolins have significantly decreased immune responses due to a genetic dysfunction, making them extremely fragile. They are susceptible to diseases such as
pneumonia Pneumonia is an inflammatory condition of the lung primarily affecting the small air sacs known as alveoli. Symptoms typically include some combination of productive or dry cough, chest pain, fever, and difficulty breathing. The severity ...
and the development of ulcers in captivity, complications that can lead to an early death. 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. The idea of farming pangolins to reduce the number being illegally trafficked is being explored with little success. The third Saturday in February is promoted as World Pangolin Day by the conservation NPO Annamiticus. World Pangolin Day has been noted for its effectiveness in generating awareness about pangolins. In 2017,
Jackie Chan Fang Shilong (born 7 April 1954), known professionally in English as Jackie Chan and in Chinese as Cheng Long ( zh, c=成龍, j=Sing4 Lung4; "becoming the dragon"), is a Hong Kong actor, filmmaker, martial artist, and stuntman known for ...
made a public service announcement called '' WildAid: Jackie Chan & Pangolins (Kung Fu Pangolin)''. In December 2020, a study found that it is "not too late" to establish conservation efforts for Philippine pangolins (''Manis culionensis''), a species that is only found on the island province of Palawan.


Taiwan

Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia, at the junction of the East and South China Seas in the northwestern Pacific Ocean, with the People's Republic of China (PRC) to the northwest, Japan to the nort ...
is one of the few conservation grounds for pangolins in the world after the country enacted the 1989 Wildlife Conservation Act. The introduction of Wildlife Rehabilitation Centers in places like Luanshan (
Yanping Township Yanping Township () is a mountain indigenous township in Taitung County, Taiwan. The main population is the Bunun people of the Taiwanese aborigines. Administrative divisions The township comprises five villages: Hongye, Luanshan, Taoyuan, W ...
) in Taitung and Xiulin townships in Hualien became important communities for protecting pangolins and their habitats and has greatly improved the survival of pangolins. These centers work with local aboriginal tribes and forest police in the National Police Agency to prevent poaching, trafficking, and smuggling of pangolins, especially to black markets in China. These centers have also helped to reveal the causes of death and injury among Taiwan's pangolin population. Today, Taiwan has the highest population density of pangolins in the world.


References


External links


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) *

''
The New York Times ''The New York Times'' (''the Times'', ''NYT'', or the Gray Lady) is a daily newspaper based in New York City with a worldwide readership reported in 2020 to comprise a declining 840,000 paid print subscribers, and a growing 6 million paid ...
'', 6 March 2020. {{Authority control Critically endangered animals Myrmecophagous mammals Rolling animals Extant Paleocene first appearances Species endangered by human consumption Species endangered by human consumption for medicinal or magical purposes Taxa named by John Edward Gray