Pangolins, sometimes known asscaly anteaters,[12] aremammals of the orderPholidota (/fɒlɪˈdoʊtə/). The oneextant family, theManidae, has three genera:Manis,Phataginus, andSmutsia.[13]Manis comprises four species found in Asia, whilePhataginus andSmutsia include two species each, all found in sub-Saharan Africa.[14] These species range in size from 30 to 100 cm (12 to 39 in). Severalextinct pangolin species are also known. In September 2023, nine species were reported.[15]
Pangolins have large, protectivekeratin scales, similar in material to fingernails and toenails, covering their skin; they are the only known mammals with this feature. Depending on the species, they live in hollow trees orburrows. Pangolins arenocturnal, and their diet consists of mainlyants andtermites, 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 superficially resemblearmadillos, though the two are not closely related; they have undergoneconvergent evolution.
The name of order Pholidota comes fromAncient GreekΦολιδωτός – "clad in scales"[20] fromφολίςpholís "scale".[21]
The name "pangolin" comes from theMalay wordpengguling meaning "one who rolls up"[22] fromguling orgiling "to roll"; it was used for theSunda pangolin (Manis javanica).[23] However, the modern name istenggiling. InJavanese, it isterenggiling;[23] and in thePhilippine languages, it isgoling,tanggiling, orbalintong (with the same meaning).[24]
In ancient India, according toAelian, it was known as thephattáges (φαττάγης).[25]
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.[26] They are made ofkeratin, the same material from which humanfingernails andtetrapodclaws are made, and are structurally and compositionally very different from the scales of reptiles.[27] The pangolin's scaled body is comparable in appearance to apine cone. It can curl up into a ball when threatened, with its overlapping scales acting asarmor, while it protects its face by tucking it under its tail. The scales are sharp, providing extra defense from predators.[28]
Pangolins can emit a noxious-smelling chemical fromglands near the anus, similar to the spray of askunk.[29] They have short legs, with sharp claws which they use for burrowing intoant andtermite mounds and for climbing.[30]
The tongues of pangolins are extremely long, and like those of thegiant anteater and thetube-lipped nectar bat, the root of the tongue is not attached to thehyoid bone but is in thethorax between thesternum and thetrachea.[31] Large pangolins can extend their tongues as much as 40 cm (16 in), with a diameter of only about 0.5 cm (1⁄5 in).[32]
Most pangolins arenocturnal animals[33] which use their well-developed sense of smell to find insects. Thelong-tailed pangolin is also active by day, while other species of pangolins spend most of the daytime sleeping, curled up into a ball ("volvation").[32]
Arboreal pangolins live in hollow trees, whereas the ground-dwelling species dig tunnels to a depth of 3.5 m (11 ft).[32]
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 abipedal stance for some behavior, and may walk a few steps bipedally.[34] Pangolins are also good swimmers.[32]
Pangolins areinsectivorous. 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 140 to 200 g (5 to 7 oz) of insects per day.[35] Pangolins are an important regulator of termite populations in their natural habitats.[36]
Pangolins have very poorvision. They also lack teeth. They rely heavily onsmell andhearing, 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.[37] They use their powerful front claws to dig into trees, soil, and vegetation to find prey,[38] 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. Theirsaliva is sticky,[37] causing ants and termites to stick to their long tongues when they are hunting through insect tunnels. Without teeth, pangolins cannot also chew;[39] but whileforaging, they ingest small stones (gastroliths), which accumulate in their stomachs to help to grind up ants.[40] This part of their stomach is called thegizzard, and it is also covered in keratinous spines.[41] These spines further aid in the grinding up and digestion of the pangolin's prey.
Some species, such as thetree pangolin, use their strong,prehensile tails to hang from tree branches and strip away bark from the trunk, exposing insect nests inside.[42]
Pangolins are solitary and meet only to have sex, 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, malesmark 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.[44]
Gestation periods differ by species, ranging from roughly 70 to 140 days.[45] 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.[32] Weight at birth is80 to 450 g (2+3⁄4 to15+3⁄4 oz), and the average length is 150 mm (6 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, 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.[46]
The order Pholidota was long considered to be thesister taxon toXenarthra (neotropicalanteaters,sloths, andarmadillos), but recent genetic evidence indicates their closest living relatives are thecarnivorans, with which they form aclade, theFerae.[47][48][49][50]Palaeanodonts are even closer relatives to pangolins, being classified with pangolins in the cladePholidotamorpha.[51] The split between carnivorans and pangolins is estimated to have occurred 79.47Ma (million years) ago.[52]
The first dichotomy in thephylogeny of extant Manidae separates Asian pangolins (Manis) from African pangolins (Smutsia andPhataginus).[50] Within the former,Manis pentadactyla is thesister group to a clade comprisingM. crassicaudata andM. javanica. Within the latter, a split separates the large terrestrial African pangolins of the genusSmutsia from the small arboreal African pangolins of the genusPhataginus.[53]
Asian and African pangolins are thought to have diverged about 41.37 Ma ago.[52] Moreover, thebasal position ofManis within Pholidota[50][54] suggests the group originated in Eurasia, consistent with theirlaurasiatherian phylogeny.[50]
Pangolins are in high demand in southern China andVietnam because their scales are believed to have medicinal properties intraditional Chinese andVietnamese medicine.[56] Their meat is also considered a delicacy.[57][58][59][60][61] 100,000 are estimated to be trafficked a year to China and Vietnam,[62] amounting to over one million over the past decade.[63][64] This makes them the mosttrafficked animal in the world.[63][65] This, coupled withdeforestation, has led to a large decrease in the numbers of pangolins. Some species, such asManis pentadactyla have become commercially extinct in certain ranges as a result of overhunting.[66] In November 2010, pangolins were added to theZoological Society of London's list of evolutionarily distinct and endangered mammals.[67] All eight species of pangolin are assessed as threatened by theIUCN, while three are classified ascritically endangered.[19] All pangolin species are currently listed under Appendix I ofCITES which prohibits international trade, except when the product is intended for non-commercial purposes and a permit has been granted.[68]
China had been the main destination country for pangolins until 2018, where it was surpassed by Vietnam. In 2019, Vietnam was reported to have seized the largest volumes of pangolin scales, surpassing Nigeria that year.[69]
Pangolins are also hunted and eaten in Ghana and are one of the more popular types ofbushmeat, while local healers use the pangolin as a source of traditional medicine.[70]
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 stimulatelactation or cure cancer orasthma.[71] In the past decade, numerous seizures of illegally trafficked pangolin and pangolin meat have taken place in Asia.[72][73][74][75] In one such incident in April 2013, 10,000 kg (22,000 pounds) of pangolin meat were seized from a Chinese vessel that ran aground in thePhilippines.[76][77] 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.[78] The same threat is reported inNigeria, where the animal is on the verge of extinction due tooverexploitation.[79] The overexploitation comes from hunting pangolins for game meat and the reduction of their forest habitats due to deforestation caused bytimber harvesting.[80] The pangolin are hunted as game meat for both medicinal purposes and food consumption.[80]
In recent studies, new discoveries have expanded our understanding of the Giant Ground Pangolin’s distribution and ecological habits. Camera trap surveys proved so much with locating this exclusive species, by targeting burrows, which is a key component of their behavior and habitat (Mattews et al., 2023). Additionally, recent records have confirmed that these species are present in new areas of Kenya, which significantly extended its known range (Sandri et al., 2023).”
Thenucleic acid sequence of a specific receptor-binding domain of thespike protein belonging tocoronaviruses taken from pangolins was found to be a 99% match with SARS coronavirus 2 (SARS-CoV-2), the virus which causesCOVID-19 and is responsible for theCOVID-19 pandemic.[83][84] Researchers inGuangzhou, 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 intraditional Chinese medicine was suggested as avector for human transmission.[83][85] However, whole-genome comparison found that the pangolin and humancoronaviruses share only up to 92% of their RNA.[86][87] 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 toAsian palm civets during theSARS outbreak.[86][88] It was later proved that the testing which suggested that pangolins were a potential host for the virus was flawed, whengenetic analysis showed that thespike 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.[89]
In 2020, two novel RNA viruses distantly related topestiviruses andcoltiviruses have been detected in the genomes of deadManis javanica andManis pentadactyla.[90] 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 inAmblyomma javanense nymphticks from a diseased pangolin.[90]
The first record of pangolin scales occurs inBen 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.[97] During theTang dynasty, 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 theWai Tai Mi Yao ("Arcane Essentials from the Imperial Library").[97] In theSong dynasty, the notion of penetrating and clearing blockages was emphasized in theTaiping sheng hui fan ("Formulas from Benevolent Sages Compiled During the Era of Peace and Tranquility"), compiled by Wang Huaiyin in 992.[97]
In the 21st century, the main uses of pangolin scales arequackery practices based on unproven claims the scales dissolveblood clots, promote blood circulation, or helplactating women secrete milk.[91][97] The supposed health effects of pangolin meat and scales claimed byfolk medicine practitioners are based on their consumption of ants, long tongues, and protective scales.[91]
The officialpharmacopoeia of the People's Republic of China included Chinese pangolin scales as an ingredient in TCM formulations.[97] Pangolins were removed from the pharmacopoeia starting from the first half of 2020.[98] Although pangolin scales have been removed from the list of raw ingredients, the scales are still listed as a key ingredient in various medicines.[99]
Pangolin parts are also used for medicinal purposes in other Asian countries such as India, Nepal and Pakistan. In some parts of India and Nepal, locals believe that wearing the scales of a pangolin can help prevent pneumonia.[100] Pangolin scales have also been used for medicinal purposes in Malaysia, Indonesia and northern Myanmar. Indigenous people in southernPalawan, Philippines, have held the belief that elders could avoid prostate illnesses by wearing belts made with the scales.[101]
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.[102] As of January 2020[update], the IUCN considered all eight species of pangolin on itsRed List of Threatened Species as threatened.[19] 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 andtrafficking of the animal while educating communities on its importance.[63] 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.[62] In 2018, a Chinese NGO launched the Counting Pangolins movement, calling for joint efforts to save the mammals from trafficking.[103][104][105] Wildlife conservation groupTRAFFIC has identified 159 smuggling routes used by pangolin traffickers and aims to shut these down.[106]
Pangolins (in rectangular cages) in an illegal wildlife market inMyanmar
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.[45][107] Pangolins have significantly decreased immune responses due to a genetic dysfunction, making them extremely fragile.[108] They are susceptible to diseases such aspneumonia and the development ofulcers in captivity, complications that can lead to an early death.[45] In addition, pangolins rescued from illegal trade often have a higher chance of being infected with parasites such asintestinal worms, further lessening their chance for rehabilitation and reintroduction to the wild.[45]
The idea of farming pangolins to reduce the number being illegally trafficked is being explored with little success.[109] The third Saturday in February is promoted as World Pangolin Day by the conservation NPO Annamiticus.[110] World Pangolin Day has been noted for its effectiveness in generating awareness about pangolins.[111]
In 2017,Jackie Chan made a public service announcement calledWildAid: Jackie Chan & Pangolins (Kung Fu Pangolin).[112]
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.[113][114]
Taiwan is one of the few conservation grounds for pangolins in the world after the country enacted the 1989 Wildlife Conservation Act.[115] The introduction of Wildlife Rehabilitation Centers in places like Luanshan (Yanping Township) inTaitung andXiulin townships inHualien 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.[116] Today, Taiwan has the highest population density of pangolins in the world.[117]
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