Pangolins

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Pangolins
Malay pangolin (Manis javanica)

Malay pangolin ( Manis javanica )

Systematics
Class : Mammals (mammalia)
Subclass : Higher mammals (Eutheria)
Superordinate : Laurasiatheria
without rank: Ferae
Order : Pholidota
Family : Pangolins
Scientific name of the  order
Pholidota
Weber , 1904
Scientific name of the  family
Manidae
JE Gray , 1821

The pangolins or pine cones (Manidae) are a family of mammals that also form their own order , the Pholidota . The family consists of three extant genera with eight species, four of which in East , South and Southeast Asia and four in Africa south of the Sahara live. They are insect-eating animals specializing in ants and termites , which are adapted to this diet by means of grave claws , a tubular snout with a toothless jaw and a long tongue . Unique among mammals is their body covering with large, overlapping horn scales. Pangolins live on the ground or in trees, depending on the species, and are usually nocturnal. However, the exact way of life has not been adequately researched. They prefer both forests and partly open landscapes in low-lying areas and medium mountain heights. In case of threat, they can curl up into a ball. The originally Malay word peng-guling also refers to this property , the modification of which pangolin is mainly used in the English and French language areas as a colloquial term for a pangolin .

The current family name Manidae was introduced in 1821. In the early research history of the 19th and early 20th centuries, the pangolins were considered close relatives of the anteaters and armadillos . With the former they share the toothless mouth and the long tongue. Above all, the lack of teeth led to the establishment of a taxon called Edentata, in which all three groups of animals were listed for a long time. Only modern molecular genetic studies showed from the mid-1980s that pangolins are more closely related to predators. The similarities with anteaters and armadillos are based on convergence . The loss of teeth, but also the specialized way of life, also means that pangolins are rarely found in fossil form. The earliest representatives of the Manidae are known from the Pliocene around 5 million years ago, but closely related forms already appeared in the Middle Eocene around 47 million years ago.

All eight pangolin species today are considered to be more or less threatened and are internationally protected. The main causes of the threat are the sale of the meat as an exotic food specialty on the one hand and the use of the scales and other body parts in local ritual customs and traditional Chinese medicine on the other. This means that the pangolins are not only hunted intensively, but also belong to the most frequently illegally trafficked mammals in the world.

features

External physique

Prepared Chinese pangolin ( Manis pentadactyla ): You can see the large grave claws on the front feet and the muscular tail, which can carry the weight of the animal
Indian pangolin ( Manis crassicaudata )

Pangolins have an elongated body with short limbs, a small, pointed head and a long tail. The head-trunk length varies depending on the species, with smaller representatives such as the white-bellied ( Phataginus tricuspis ) and the long-tailed pangolin ( Phataginus tetradactyla ) it is between 25 and 43 cm; the largest species is the giant pangolin ( Smutsia gigantea ) with 67 to 81 cm. The tail becomes again between 25 and 70 cm long. In the tree-dwelling pangolins, the tail exceeds the rest of the body length, in the others it is the same length or shorter. The weight varies between 1.6 and 33 kg, with the males usually being larger than the females. Fossil appeared with Manis palaeojavanica in the Pleistocene of Southeast Asia, a species that reached a total length of around 2.5 m and thus represents the largest known pangolin species.

The pangolin head is small and conical in shape. The eyes are also small and protected by bulging, gland-free lids . Auricles are missing in the African species, in the Asian species there is often only a thickened crest. The nose can be closed by a fold of skin (plica alaris), which is an advantage if the animals stick their snouts in insect burrows to eat.

The eponymous feature is the body covering, which is unique among mammals, and consists of large scales of horn that cover the top of the head, the trunk, the outside of the limbs (in some species without the forearms) and the top and bottom of the tail. Only the face, the stomach and the inside of the limbs are bare and have a gray, coarse skin covered with white, brown or black hair. In African pangolins, the scales on the back of the tip of the tail are arranged irregularly or in pairs, while in Asian pangolins they are always regularly in only one row. On the underside of the tip of the tail, however, arboreal pangolins have a free area covered by horny skin; the armor is closed for those living on the ground. Individual hairs grow between the scales of the back armor only in Asian species.

The limbs appear short and strong and each end in five toes ( pentadactyl ). The front legs show adaptations to a digging way of life in that the middle three fingers are provided with long, curved claws, of which the middle one is again significantly larger. The claws of the first and fifth fingers, on the other hand, are smaller and are not used when digging. The hind legs are stronger and a little longer, the five toes also have claws. In general, the forefoot claws of the pangolins living on the ground are longer and less curved than those of the arboreal ones; the latter, in turn, have significantly longer hind foot claws that support movement in the trees.

Skull features

White-bellied pangolin ( Phataginus tricuspis ) skull

The skull reaches lengths between 6 and 16 cm. It is generally conical in shape with a tubular, slightly narrowing and slightly elongated rostrum towards the front . Since the food is not chewed, the masticatory muscles are regressed, so that only a few bones are formed as muscle attachment points. This makes the skull appear very smooth, making it one of the simplest-built skulls among mammals.

A noticeable characteristic is the not fully developed zygomatic arch , a feature that pangolins share with the anteaters of South America, which also specialize in ants and termites, and is often considered an adaptation to this diet. However, some pangolins, such as the Chinese pangolin, sometimes have closed zygomatic arches. Further general characteristics can be found in the elongated nasal bones and the front bones, which are larger than the parietal bones .

Teeth are completely absent, the lower jaw is only designed as a simple bone brace with weakly developed, backward-pointing and spherically shaped joint ends that leave little room for the movement of the lower jaw. The symphysis of the lower jaw forming a flat surface through which the tongue can slide. However, as a characteristic of all pangolins at the rear end of the symphysis there is a pair of bony, conical, pointed elevations that are similar to a canine tooth .

Scale armor

Individual scales of the scale armor

The scale armor, together with the rest of the skin, makes up about a quarter to a third of the total body weight. It consists of 160 to 290 individual scales, almost half of which are on the tail. They are movable and overlap each other like roof tiles. They are arranged in rows, the number of which varies between 13 and 25 on the trunk. The color of the scales ranges from dark brown to olive green to yellowish. They are triangular to V-shaped; large scales are 7 to 8 cm in length and width. On the surface there are longitudinally directed ripples, and they are also equipped with sharp edges. The largest scales are usually on the back with the tip pointing backwards. When rolled up, the sharp ends protrude similar to a half-opened pine cone . The scale armor protects less against ant bites or termite bites or skin parasites than against injuries caused by larger predators or digging underground.

The scales are keratinized formations of the epidermis that sit on protrusions of the dermis that are bent backwards . Three layers can be distinguished in the cross-section: The upper dorsal plate (back plate) takes up about one sixth of the thickness and consists of flattened, strongly keratinized cells. The intermediate plate, which takes up the greatest amount of space, is formed from less flattened, keratinized cells. The ventral plate (belly plate) forms the underside of the scale and is only a few cells thick. All three plates are formed from different epidermal germinal areas. The absence of filaments indicates that the scales do not correspond to glued hair, as previously assumed. In terms of their structure, they can rather be compared with the fingernails of primates and, like these, grow constantly, which compensates for wear and tear. This also distinguishes them from the scale skin of the scalloped reptiles , which sometimes has to be changed annually.

It is believed that the scale armor was formed early in the development of pangolins - the oldest evidence with Eomanis comes from the Middle Eocene around 47 million years ago, found in the Messel mine in Hesse. Possibly a scaling of the tail formed first, which would be regarded as a homologous development to some representatives of the rodents such as the house mouse or the nutria or the shrews , only later was the body fully armored.

Skeletal features

Skeleton of a long-tailed pangolin ( Phataginus tetradactyla )

The number of vertebrae varies from species to species and ranges from 48 in the steppe pangolin ( Smutsia temminckii ) to over 70 in the long-tailed pangolin. Depending on the type , the spine consists of 7 cervical, 12 to 15 thoracic, 5 to 6 lumbar, 2 to 4 cross and 21 to 50 tail vertebrae. The animals can curl up well because the pelvis is very short, the iliac bone is bent outwards and the lumbar vertebrae are elongated. The tail vertebrae have chevron bones on the underside , which serve as a base for the strong tail muscles, since the tail is wrapped around the body like a shield when it is rolled up. The sword extension at the rear end of the sternum is enlarged up to the pelvic region and serves as a starting point for the complicated tongue muscles.

The humerus in particular is particularly strong for the digging and tree-climbing way of life. It has a very wide elbow joint and - typical for pangolins - a strong crista deltoidea , which surrounds the shaft as a crest of bone and acts as an attachment point for the shoulder muscles. On the thigh bone , the third roll hillock (trochanter tertius), another muscle attachment point on the shaft, is offset far down to the joint ends and is therefore hardly visible. In the case of very primitive Pholidota, this is significantly higher and prominently protruding on the shaft. Another special characteristic are the respective last links of the toes of the fore and hind feet (each phalanx III), which have an elongated shape and deep notches at the end, in which the claws stick.

Internal organs

The worm-shaped tongue , covered with sticky saliva , with which the food is ingested, is very characteristic . In the giant pangolin it can be up to 70 cm long and stretched out up to 25 cm, in the Chinese pangolin it is up to 41 cm long with a diameter of up to 1.1 cm. Your complex musculature consists of longitudinal and radial muscle fibers. In the resting state, the front part of the tongue lies curled up in the mouth, the surface is roughened in the front area by conical papillae, at the tip there are mushroom-shaped taste papillae. Unlike other mammals, the tongue is not connected to the hyoid bone , but rather to the rear part of the sternum via an external muscular system that is partially homologous to the hyoid muscles . The hyoid bone has a different function in pangolins: With it, the insects sticking to the tongue are scraped off at the entrance of the esophagus . The salivary glands are enlarged and extend into the breast and armpit region.

The muscular stomach takes over the mechanical chopping of the insects. It is equipped with keratinized and stratified squamous epithelium , which protects it from the bites and venom of ants and termites. The greatly enlarged porter’s muscles grind the swallowed food and are provided with ossified spikes (pyloric thorns) for better chopping - small stones are also swallowed. The gastric glands are very long and tubular; they form bundles of glands that empty through a central passage to the porter. The entire intestinal tract of the Chinese pangolin reaches a length of 5.2 m and a diameter of around 1 cm. It is coiled in a tubular shape and shows no differences between the small intestine and the large intestine , only in some individuals there is a slight thickening or tortuous formation in the rear area, which may indicate the transition from the small intestine to the large intestine. An appendix is not formed. Pangolins have anal glands whose scented secretions are used for communication and possibly for defense. The females have a two-horned uterus ( uterus bicornis ). Males have a small penis but no scrotum - the testicles are under the skin.

The brain is very simply built and small, it only makes up around 0.2 to 0.5% of the body weight of the Malay pangolin. Only the olfactory bulb is well developed; accordingly, the sense of smell plays an important role in foraging for food and in communicating with conspecifics. According to the structure of the brain - here mainly related to the cerebellum - Asian species are somewhat more primitive than African ones.

distribution and habitat

Distribution of pangolins
Species in Asia
  • Front Indian pangolin
  • Chinese pangolin
  • Malay pangolin
  • Palawan pangolin
    • Species in Africa
  • Steppe pangolin
  • White-bellied pangolin
  • Giant pangolin
  • Long-tailed pangolin

    • Pangolins live in Africa south of the Sahara and in South , Southeast Asia and Southern Asia . In Africa their distribution area extends from Senegal and Sudan to South Africa . In Asia they are distributed from Pakistan and Nepal through India and the Indochinese Peninsula to southern China and from the Malay Peninsula to Borneo and the Philippines . The pangolins thus primarily inhabit tropical regions.

    Their habitat includes a variety of landscape types, such as alluvial and swamp forests , but also rainforests , open savannas and scrublands as well as vegetation areas designed like mosaics. Furthermore, they also tolerate secondary landscapes used by humans such as plantations , garden landscapes and farm areas, which must contain enough protection in the form of trees or rocks and burrows. The animals avoid human settlement areas and farmland and are sensitive to pesticides . The pangolins use the flatlands and highlands; in the Nilgiri Mountains in India, the Indian pangolin ( Manis crassicaudata ) has been detected at altitudes of around 2300 m. In addition to dense underground vegetation, the basic prerequisites for the presence of pangolins are sufficient food sources for ants and termites as well as water.

    Because of the diverse landscapes and specialization in different food groups, the individual ecological niches used rarely overlap in sympatric species . In individual cases, however, there is also an increased formation of niches. The long-tailed pangolin increasingly uses water areas in regions where the white-bellied pangolin appears at the same time. The Chinese pangolin ( Manis pentadactyla ) still lives in northern Vietnam, where the Malay pangolin ( Manis javanica ) is widespread, principally at heights over 600 m. Even with other highly specialized insectivores, such as the African aardvark ( Orycteropus ), there is hardly any overlap in the same used landscapes due to the strong formation of niches.

    Way of life

    Locomotion and social behavior

    The steppe pangolin ( Smutsia temminckii ) is one of the ground-dwelling pangolins.
    The white-bellied pangolin is a tree-climbing pangolin.

    In general, the way of life and social behavior of pangolins has not been well researched. They live mostly solitary, only in the mating season there are brief relationships. The main activity takes place during twilight or at night, but isolated animals can also be observed during the day. The individual individuals are largely tied to their location and use action spaces that are significantly larger for males with 30 to 43 hectares than with females with 3 to 7 hectares. The area of ​​a male animal overlaps several of the females, with some species it can be concluded that there is a certain territoriality because the males actively defend their territory against competitors. The secretion of the anal glands probably plays an important role in communication between conspecifics , as well as in the marking of territories. The ground-dwelling species retreat to rest in earth burrows that they have either dug themselves or taken over from other animals. These burrows are often found in ant or termite mounds , between roots or in depressions that have been created by fallen trees. Generally they are in dense vegetation. Usually the burrows extend several meters below ground and end in a round chamber with a diameter of up to 2 meters. The entrance to the burrow is blocked with mud when the animal is present. Pangolins usually move into a burrow for several days in a row and look for or dig a new one when the food supply has declined. The tree-dwelling species, on the other hand, use tree hollows. Females generally move less far from their burrows than males, and in general the range of action is rather small for both sexes.

    The pangolins move slowly and sedately on the ground, the hand can be placed on the outside edge or the knuckles. It is characteristic of the head swinging back and forth in four-foot gait, with the head moving to the side opposite the guide leg. This swinging is caused by the fact that the shoulder muscles are arranged differently than in unarmored mammals due to the dense covering of the front legs by the scale armor. Individual muscles completely cover the shoulder blade and thus restrict its movements, the swinging head thereby brings about a balance. Ground pangolins, especially the steppe pangolin, can also move on their hind legs and balance the body with their tail, which then does not touch the ground. Generally, pangolins are able to climb trees. The actually tree-dwelling species move like a caterpillar, first with their front legs, then with their back legs, with the back constantly being bent and stretched. The tail serves as a grasping organ and is often wrapped around the trunk or a branch, the scales are splayed out and anchor the animal to the bark. Sometimes it is also used as a fishing rod when a pangolin changes tree or branch. Pangolins are also considered good swimmers, especially the long-tailed pangolin. To do this, they take in extra air so that the body with the heavy scale armor protrudes halfway out of the water and make sideways meandering movements with the tail.

    nutrition

    The diet of pangolins consists primarily of ants and termites ( myrmecophagia ), and they are extremely selective. Only larger species such as the giant pangolin or the Indian pangolin occasionally eat other insects or other invertebrates , such as beetles , cockroaches or worms . The animals eat a comparatively large proportion of biomass , which is 300 to 400 g per day for the smallest representatives and 2 kg for the largest. The food is tracked down with the exceptionally good sense of smell , whereby the ground is constantly searched with the nose, and some animals also dig tunnels to reach nests. The pangolins break open insect burrows or tree bark with their grave claws, and they eat their food with their sticky tongues. The eyes, ears and nostrils can be closed and prevent insects from entering while eating. Insects that get on the body are shaken off. When they eat, they usually rise up on their hind legs. The eating process usually takes several hours, and the structure is not completely destroyed. Usually an animal returns several times in a row over several days to the same ant or termite nest, only when the prey becomes significantly rarer does it look for another.

    Reproduction

    Mother with offspring of the Palawan pangolin ( Maynis culionensis ).

    Little is known about the mating behavior of pangolins. It is most likely not seasonally bound, with the possible exception of the Chinese pangolin. In this species, several males fight very aggressively for mating privilege in late summer or autumn. As a rule, males roam several female territories every night in search of ruthless females, which they recognize by the scent trails. Mating rituals have been observed especially in the white-bellied and long-tailed pangolin. For example, males and females undertake mock competitions with blows chest against chest until the female submits. Usually the female then clings to the male's tail, which pulls it to the mating site. During mating, the tails of the animals are intertwined. Couples then share a resting place for several days. The gestation period in African pangolins is 130–150 days, in Asiatic it may be shorter and should last two to three months. As a rule, the female gives birth to a single newborn, which is very developed in most species. The birth weight is between 70 and 425 g, depending on the species, the birth length between 15 and 30 cm. The newborns have open eyes, the scales are not overlapping and initially soft, they harden in the first days of life. Females have a pair of axillary teats that are used to suckle the young.

    In tree species, the young animal remains in a tree cavity for the first week, only then does it climb onto the tail of the mother animal and can thus be carried until it is weaned. In the ground-dwelling species, the young are born underground and stay there for about two to four weeks until they leave the cave for the first time, clinging to the mother's tail. In the event of danger, it usually slides on its belly and the mother animal covers it with its tail. After two to four weeks, but at the latest after three months, the young animal consumes solid food for the first time; it is often first carried to the food sources on the root of the tail of the mother. After around five months, they separate from their mother and they reach sexual maturity at one to two years of age. The highest known age of a pangolin in human care was just over 19 years.

    defense

    Rolled up steppe pangolin

    Pangolins are rather shy and cautious animals. In the event of a threat, they first try to reach the protective shelter. If they fail, they can curl up into a ball. The muscular tail is placed over the unprotected stomach and face in order to protect these regions from attack. When rolled up, they also erect the scales, the sharp edges of which offer additional protection. Smaller species also interlock the tail scales with those of the neck, so that the animals can hardly be rolled up. In addition, the tail can be used to deliver rapid defensive blows, which are particularly effective thanks to the sharp edges of the scales. There is a report from Indonesia that a pangolin curled up in a ball and rolled down a slope. It covered 30 meters in 10 seconds. Another method of defense is the spraying of a foul-smelling secretion from the anal glands, similar to the skunks . The long claws are not used for defense.

    Systematics

    External system

    Internal systematics of Laurasiatheria according to O'Leary et al. 2013
      Laurasiatheria  

     Eulipotyphla (insect eater)


      Scrotifera  
      Ferae  

     Pholidota (pangolins)


       

     Carnivora (predators, including Pinnipedia (seals))



       

     Chiroptera (bats)


      Euungulata  

     Perissodactyla (odd-toed ungulate)


       

     Cetartiodactyla ( Artiodactyla ( Artiodactyla ( Artiodactyla  ) and  Cetacea  (Whales))






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    The pangolins (Manidae) are the only family from the order of the Pholidota today. According to modern molecular biological studies, the pangolins are the closest living relatives of the predators (Carnivora), both orders together form the higher taxon of the Ferae . They are part of the superordinate order of Laurasiatheria , one of the four main lines of the higher mammals . According to genetic analyzes, the separation of predators from pangolins may have taken place in the Upper Cretaceous more than 80 million years ago or in the early Paleocene just under 65 million years ago.

    In addition to today's pangolins, the Pholidota contain two closely related, albeit extinct, families, the Eomanidae and the Patriomanidae , the latter being regarded as the sister group of the Manidae. The next related large group are the palaeanodonta . This group of mammals, which is also extinct, is known from the Paleocene and Eocene mainly from North America, more rarely from Europe and East Asia, and is divided into three families, the Escavadodontidae , the Epoicotheriidae and the Metacheiromyidae . Possibly, however, Ernanodon from East Asia must also be included in the palaeanodonta. This originally formed the monotypical Ernanodonta, but after analyzes of an almost complete skeleton it is closer to Palaeanodon . The palaeanodonta also showed adaptations to a burrowing and insectivorous way of life, for example strong limbs with large claws and smaller, less specialized teeth with only a thin layer of tooth enamel . Due to the partly primeval features such as trained teeth, they could also represent the forerunners of the Pholidota. The similarity is so strong that the Palaeanodonta and the Pholidota are now grouped together in the parent taxon Pholidotamorpha .

    Internal system

    Internal systematics of the Pholidota according to Gaudin et al. 2009 and Gaudin 2010
      Pholidotamorpha  

     Palaeanodonta


      Pholidota  

     Euromanis


       

     Eurotamandua


       
      Eomanidae  

     Eomanis


      Manoidea 
      Patriomanidae  

     Necromanis


       

     Cryptomanis


       

     Patriomanis




      Manidae  

     Manis


       

     Phataginus


       

     Smutsia






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    Template: Klade / Maintenance / Style

    The pangolin family is now divided into eight species divided into three genera . The genus Manis includes the Asian species as well as some documented fossil forms from the Eurasian region. Smutsia, on the other hand, represents the African pangolins, while Phataginus contains the African tree-dwelling pangolins. This subdivision into several genera can also be substantiated morphologically on the basis of individual skull or foot skeleton features. There is a general difference between Asian and African pangolins in the length of the rostrum , which in the former is markedly more elongated compared to the latter. The anterior zygomatic arch attachment also extends longer in the Asian pangolins than in the African. According to cladistic studies of skeletal features including all fossil forms of the Pholidota, the Asian pangolins are a sister group to the African. The former are viewed as monophyletic , the latter in turn form two separate groups. For this reason, the Asiatic pangolin were placed in the subfamily of the Maninae , while the two African genera represent two subfamilies, the Smutsiinae and the Phatagininae . The anatomically determined basic division of pangolins into an Asian and an African strain, as well as the division of the latter, find their counterparts in the genetic studies that have been increasingly carried out since the 2010s. According to these, the greater diversification of pangolins began with the separation of the Maninae line from that of the African pangolins in the Middle or Upper Eocene a good 46.9 to 37.9 million years ago. The African group, in turn, may have split up during the Oligocene to the transition to the Miocene around 29.6 to 22.9 million years ago.

    Internal systematics of the Manidae according to Gaubert et al. 2018
      Manidae  
      Manis  


     Manis crassicaudata


       

     Manis culionensis


       

     Manis javanica




       

     Manis pentadactyla



       
      Smutsia  

     Smutsia gigantea


       

     Smutsia temminckii



      Phataginus  

     Phataginus tetradactyla


       

     Phataginus tricuspis





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    However, there are also approaches in which the individual species only belong to a single genus, in this case Manis , whereas the other genera only have the status of a subgenus. However, since the genetic investigations, the attempt at structuring has been increasingly rejected. Others, however, differentiated only the Asian ( Manis ) and African forms ( Phataginus ), which was justified morphologically. In addition, the view was expressed that the African tree pangolins can be split into two genera, Phataginus and Uromanis , which would make a total of four genera, a model that was partly followed by the IUCN . In the past, however, there were also suggestions with up to six genres. The suggestion of a division into three genres was first made at the end of the 1990s and was subsequently supported by several cladistic studies.

    Overview of the recent and fossil species of pangolin

    Within the Manidae family, eight recent and three fossil species are now divided into three genera of pangolin:

    • Family: Manidae Gray , 1821
    • Manis (= Pangolinus , Paramanis , Pholidotus ) Linnaeus , 1758
    • Phataginus (= Phatagin , Triglochinopolis , Uromanis ) Rafinesque , 1821

    Genetic data suggests that the white-bellied pangolin may contain several cryptic species . This is also supported by skull morphological findings. Some scientists therefore believe there are more than a dozen species of pangolin. The validity of the extinct species Manis hungarica and Manis lydekkeri is sometimes questioned, as their respective descriptions are based only on isolated phalanges .

    Within the order Pholidota, the following extinct families and genera can also be distinguished:

    Research history

    Taxonomy

    John Edward Gray (1800-1875)

    The name of the pangolin family, Manidae, was introduced by John Edward Gray in 1821 . Gray described the characteristics of the manids as follows:

    Mouth very small: tongue very long, fili-form: body hairy or scaly: claw recurved, acute: tail long: ears short: caecum none: stomach simple

    "Mouth very small: tongue very long, thread-like: body hairy or scaly: claw bent back, pointed: tail long: ears short: appendix missing: stomach simple"

    At that time, however, the description referred to the anteaters of today's pangolins , which Gray incorporated into the Manidae; four years later he separated the anteaters from the pangolins, but regarded both as a subgroup of the armadillos . The generic name Manis was the inspiration for the family name Manidae . This was first used by Linnaeus in his work Systema Naturae in 1758 . With M. pentadactyla he mentioned only one species of which he must have seen a specimen based on the specification of the precise number of toes. The name Manis goes back to the Manen ( Latin manes ), Roman spirits of the dead, and refers to the nocturnal and mostly hidden way of life. The term pangolin , which is often used colloquially in the English- speaking world, is derived from Malay , where peng-guling means something like "roll in" and on the one hand reflects the ability to curl up, on the other hand it emphasizes the ability to be caught by people.

    The Edentata problem - on the systematic position of pangolins

    Originally, the pangolins and some of the groups related to them (palaeanodonta) were closely related to the articulated animals (xenarthra) for anatomical and morphological reasons . The anteaters (Vermilingua) have a comparable toothless, elongated snout with an elongated tongue, a reduced zygomatic arch and grave claws, while the armadillos (Dasypoda) have a similar body armor and in some cases also represent strict insectivors. In contrast to the little diverse pangolins and their predecessors, the Xenarthra represent a very diverse group both today and in their phylogenetic past, which, however, is largely limited to South America . Due to the appearance of the typical xenarthric joints (xenarthral or secondary joints) on the posterior thoracic and lumbar vertebrae, they have long been considered a natural group. In fact, the idea of ​​a taxon Edentata (toothless) was widespread until the 1980s . The edentata were characterized either by the loss of all teeth or by the development of a homodontic dentition with a simultaneous extensive reduction in tooth enamel . As a result, the Edentata differed from most of the higher mammals with their characteristic heterodontic dentition. The taxon Edentata goes back to Étienne Geoffroy Saint-Hilaire and Georges Cuvier , who had introduced the term in 1795 (both used the term "Edentati" used by Félix Vicq d'Azyr as early as 1792), and originally consisted of the pangolins, the Anteaters and the armadillos, three years later Cuvier added the sloths and the African aardvark ( Orycteropus ). Before the establishment of the Edentata, Gottlieb Conrad Christian Storr had already combined the named groups with the exception of the aardvark under the name Mutici in 1780, while Linnaeus had not yet recognized any relationships between these groups. Rather, Linnaeus saw in his work Systema Naturae from 1758 the genus Manis , today's Asian pangolins, in the vicinity of the elephants , but also the anteaters and sloths, while at the same time he placed the armadillos on the side of the insectivores and pigs (the aardvark was not yet known at the time).

    Already towards the end of the 19th century it was recognized that the individual members of the Edentata did not form a natural group. In 1872 Thomas Henry Huxley referred the aardvark to its own order, the Tubulidentata, just like the pangolin. As his favorite name Squamata but already by the Squamata was preoccupied, applies Max Wilhelm Carl Weber's 1904 term coined Pholidota today to refer to the order of pangolins. In 1889 Edward Drinker Cope again established the umbrella term Xenarthra for the South American representatives and thus the actual core group of the Edentata. The composition and systematic position of the Edentata were assessed differently in the further course of the research history. Above all, William Diller Matthew brought the palaeanodonta to the Edentata in 1918. Based on anatomical features, he also recognized the close relationship between the pangolins and the palaeanodonta and viewed the latter as the ancestors of the former. In his study on the classification of mammals from 1945, George Gaylord Simpson separated the Pholidota and thus the pangolins (but not the Palaeanodonta) from the Edentata; he also did not see the two groups in a very close relationship, and he continued to plead for equation the terms Edentata and Xenarthra. Robert J. Emry, in turn, united the palaeanodonta with the pangolin among the Pholidota in 1970, referring to the arguments already put forward by Matthew. In addition, he emphasized, like Simpson before him, the different paleogeographical distribution of the sibling animals and the Pholidota, due to which both groups cannot have a common ancestral ancestor. Around the same period, Malcolm C. McKenna saw the Edentata as a synonym for the Xenarthra and compared it as a sister group to all other higher mammals, which he summarized as Epitheria . He also attested that the Pholidota (and the Tubulidentata) were only distantly related to the secondary articulated animals. However, in the mid-1980s, scientists around Michael J. Novacek renewed the close relationship of the Xenarthra and Pholidota within the Edentata and saw the Edentata in the same position as McKenna before. Other researchers, in turn, regarded the Edentata as synonymous with the Pholidota and the aardvark and excluded the articular animals.

    Also in the mid-1980s, studies based on immunological properties showed for the first time a closer relationship between pangolins and predators , a result that has not been reflected in previous anatomical analyzes. Subsequent investigations were able to confirm this, whereupon in 1998 the Pholidota were united with the Carnivora under the taxon Ferae . As one of the few common morphological features, an ossified tentorium cerebelli between the cerebrum and cerebellum could be identified. Several molecular genetic studies not only underpinned the close relationship between pangolins and predators, but also led to a new division of the higher mammals into four parent groups, with the ferae being given a place in the Laurasiatheria next to the ungulate and unpaired ungulates . The secondary animals, on the other hand, are at the base of the development of the higher mammals, similar to what McKenna had already predicted in the mid-1970s. According to today's understanding, the similarities between scaled and secondary articulated animals are based on convergence and not on relationship and result from the similar ways of life of the two groups. The taxon Edentata is therefore no longer listed today. Occasionally, the term is still used as an informal term for toothless animals, which among mammals include the anteaters and pangolins as well as the baleen whales and the ant urchins .

    Tribal history

    Skeleton of Eurotamandua
    Live reconstruction of Eomanis

    Pangolins fossil remains are generally very rare. The causes are, among other things, the ecology of the animals themselves, such as the preferred wooded habitat, the solitary way of life and the low population density. This is compounded by the early loss of teeth, the most frequently preserved remains of mammals, which are usually indispensable for the exact allocation of fossil finds. As a result, individual skeletal elements are sometimes overlooked, although remains of manids may be present at various sites.

    The Pholidota represent an old order, the oldest undoubted representatives are from the Middle Eocene about 47 million years ago from the Messel mine . This includes several complete skeletons assigned to the genera Eomanis and Eurotamandua . In their physique with their pronounced grave claws and the toothless jaw, these early representatives corresponded very well to today's pangolins. In the case of Eomanis , an approximately 50 cm long animal, the first fossil record of scales was also successful. In tribal history, these two genera stand together with Euromanis , also from Messel, at the basis of the development of the Pholidota and represent the group of " Eomanidae ". In contrast, the Patriomanidae , which are known from the Upper Eocene of North America and East Asia, are much more closely related to today's pangolins . This includes Patriomanis , of which two almost complete partial skeletons and four other individuals from the White River Formation of Wyoming and the Renova Formation of Montana are available; it is the only so far discovered representative in North America. Furthermore, Cryptomanis should be mentioned, which was described using a skullless skeleton from the Shara-Murun formation of Inner Mongolia in China . In Europe later, Necromanis was widespread from the Oligocene to the Middle Miocene . At least three species of this genus are named. The entire find material is distributed among more than a dozen individuals, including an almost complete skeleton from Saulcet in the Allier Basin in France and two partial skeletons from a crevice filling near Petersbuch near Eichstätt in southern Germany. A few remains of bones also come from Solnhofen or Weisenau near Wiesbaden in Germany, from Dolnice near Cheb in the Czech Republic and from El Papiol in Catalonia . Two terminal phalanges of the forefoot and hind foot from the Gebel Qatrani Formation of the Fayyum in Egypt , which show the typical notches for the claws of today's pangolins, belong to a still undescribed representative of the pangolins from the Lower Oligocene of Africa and are estimated to be around 31 million Years dated. According to the fossil record , the Pholidota originated in the northern part of Eurasia, possibly in a more western area, and later reached North America via East Asia. Such an area of ​​origin is also consistent with the current assignment of the Pholidota to the Laurasiatheria, which are generally assigned an origin on more northerly located land masses that originally formed the Laurasia continent . The previously known fossil sites with remains of the earliest pangolins are located around 1000 km north of today's northernmost distribution limit of the recent representatives.

    The Manidae appear for the first time in the Pliocene , but fossil finds are mostly rare here too. A nearly complete skeleton of a large pangolin from the Varswater Formation of Langebaanweg in southwestern South Africa dates to the beginning of the Pliocene . It shows some pathological changes, especially on the foreleg, but clearly resembles the giant pangolin in its limb proportions . The same species is assigned a radius from the Warwire Formation from Lake Albert in Uganda , which is, however, somewhat younger. Both finds belong to animals that were slightly smaller than today's giant pangolin. Manis hungarica , the only European representative, comes from the Upper Pliocene . This was described on the basis of a 5 cm long, but fragmented terminal phalanx of the right middle finger from Villány in southern Hungary and was associated with the Asian pangolins, but the find has since been lost. Other remains of this type have been handed down from the base of a 1.5 m thick silty sand in the valley of Valea Grăunceanului in Romania . These were found together with the primate Paradolichopithecus and the giraffe-like animal Mitilanotherium and are assigned to the transition from the Pliocene to the Lower Pleistocene , the Middle Villafranchian . As early as the beginning of the 20th century, a partial skeleton consisting of 28 bones with partly articulated remains of the skull and the fore and hind legs led to the description of Manis palaeojavanica , which lived in the Middle Pleistocene in Southeast Asia. The find, which only spread over a few square meters, represents an individual of a species to which the largest pangolins known to date belong with a total length of up to 2.5 m; they are about a third larger than the giant pangolin. The species was first detected in the early Middle Pleistocene "Kedung-Brubus-Fauna" on the Indonesian island of Java , from where the remainder of an early human skull originates. The finds are around 800,000 years old, and the fauna of Citarum in the west is similar Javas estimated where a fragment of a right femur of Manis palaeojavanica came from. Further finds of this type in the form of some foot bones are also in the much younger fauna remains of the Niah Caves on Borneo , which are estimated to be around 40,000 years old , where they appeared together with the significantly smaller Malay pangolin. Another fossil form of the Pleistocene was reported with Manis lydekkeri from the Carnul caves near Madras in India . But here there is only a phalanx that differs little from that of today's pangolins. In the Nelson Bay Cave in South Africa, some remains came to light that were originally assigned to the steppe pangolin and which, with an age of 12,000 to 18,000 years, belong to the Upper Pleistocene . But it is questionable whether the finds actually represent a pangolin.

    Pangolins and humans

    Threat and protection

    Jacket made of pangolin, which was given to King George III of England in 1820     . was given
    Destruction of confiscated pangolin scales on     World Pangolin Day , February 17, 2017, in Cameroon
    Chinese pangolin in the Leipzig Zoo

    Due to their way of life, pangolins have a positive ecological benefit in that they regulate the size and spread of ants and termite populations, which is also beneficial to humans. However, the pangolins are subject to strong hunting pressure from humans. In Africa, animal meat is considered a delicacy and is sometimes used as bushmeat . In western Africa in particular, pangolin is one of the most expensive specialties and the animals are usually sold alive in markets. Pangolins are also used in local medicine . Bones and scales are used by the Yoruba and Awori in West Africa, among other things, to relieve stomach problems, but also to increase potency in men and to regulate the menstrual cycle in women. The animals are often said to have magical properties, so that individual body parts are used as talismans or lucky charms for a wide variety of reasons . In Chinese medicine in eastern Asia, the scales also play an important role. They are considered an aphrodisiac and antiseptic agent, have been recommended for the treatment of mental illnesses and nervous disorders and are used in the treatment of poisoning, inflammation, rheumatism , asthma , circulatory disorders and, more recently, against breast cancer. The scales are traded in pieces or ground into powder.

    The pangolin populations are declining. Extensive hunting is primarily responsible for this. All species are under local protection, and they have been listed in Appendix I of the Washington Convention on the Protection of Species (CITES) since 2016, which includes the species with the highest threat status. Since 2000, the CITES zero annual export quota has also been in force , a provision that forbids any international trade in pangolins or their body parts. However, there is a lively black market, some of which operates worldwide, and large quantities of smuggled are discovered again and again. In February and March 2008 alone, a total of 23 t of frozen pangolin were seized in Vietnam and in the same year a further 14 t in Indonesia. With an estimated over one million poached animals between 2004 and 2014, pangolins are the most illegally trafficked mammals in the world, according to the IUCN. In July 2017, Chinese authorities in Shenzhen succeeded in the country's largest ever confiscation of sheds with a total weight of 12 tons. At the beginning of January 2018, customs in the Taiwanese port city of Kaohsiung confiscated a container from Malaysia with 13 tons of frozen pangolins (around 4,000 individuals). The retail value was estimated at NT $ 2,000 per kilogram. International trade in pangolins has been completely banned since January 2017.

    Furthermore, the destruction of the landscape due to deforestation and traffic accidents are threat factors for the individual populations, the exact status of which is difficult to determine due to the hidden way of life. As a result, it is likely that individual local populations have already expired, such as the giant pangolin in Rwanda or the steppe pangolin in the Orange Free State of South Africa, as well as the Indian pangolin in Bangladesh and the Chinese pangolin on the island of Hainan .

    The rearing of pangolins in zoological facilities is also proving to be problematic. Pangolins have been kept in numerous zoos around the world since the 1870s, but usually the animals died after a few years. The causes mostly lay in the unaccepted food. In addition, they hardly produced any offspring. A new food strategy at Taipei Zoo led to success , where a Chinese pangolin was born for the first time in November 1998.

    The IUCN lists the four African species as "vulnerable" ( vulnerable ), which apply from the Asian representatives Palawan pangolin and the leading Indian pangolin as "high risk" ( endangered ) and the Chinese pangolin and the Malayan pangolin as "critically endangered" ( critically endangered ). One of the most important measures is an active observation program to study the exact distribution of the individual pangolin populations ( monitoring ), combined with a more detailed investigation of the ecological needs of the animals in the wild and in zoological facilities. In addition, however, priority is given to training and raising awareness among the local population and government representatives, plus exploring the mechanisms of global trade.

    Pangolins as possible vectors of disease

    Researchers suspect that pangolins could be an intermediate host for the corona virus and the associated COVID-19 pandemic . The matches in the genome of the pangolin-CoV isolated from the Malay pangolin with the SARS-CoV-2, which is active in humans, and the Bat-CoV found in bats as another possible intermediate host are around 91% each.

    literature

    • P. Gaubert: Order Pholidota. In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 2: Hooved Mammals. Lynx Edicions, Barcelona 2011, ISBN 978-84-96553-77-4 , pp. 82-103
    • Timothy J. Gaudin, Robert J. Emry, and John R. Wible: The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis. Journal of Mammalian Evolution 16, 2009, pp. 235-305
    • Ronald M. Nowak: Walker's Mammals of the World. Johns Hopkins University Press, Baltimore, 1999, pp. 1-1936 (1239-1242)

    Web links

    Commons : Pangolins  - Collection of images, videos and audio files

    Individual evidence

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    This article was added to the list of excellent articles in this version on October 27, 2015 .