White-bellied pangolin

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White-bellied pangolin
White-bellied pangolin (Phataginus tricuspis) at the San Diego Zoo.

White-bellied pangolin ( Phataginus tricuspis ) at the San Diego Zoo .

Systematics
Superordinate : Laurasiatheria
Order : Pholidota
Family : Pangolins (Manidae)
Subfamily : Phatagininae
Genre : Phataginus
Type : White-bellied pangolin
Scientific name
Phataginus tricuspis
( Rafinesque , 1821)

The tree pangolin ( Phataginus tricuspis , Syn. : Manis tricuspis ) is a mammal of the family of pangolins (Manidae). It occurs in western and central Africa and prefers tropical rainforests as a habitat. The animals are predominantly nocturnal and live as solitary animals. With its extremely long tail and small body, the white-bellied pangolin is adapted to life in trees, but it also occurs on the ground and is also a good swimmer. The diet of the pangolin is highly specialized, its main diet consists mainly of termites , it eats ants less often . Excessive hunting for nutritional purposes and for use of individual parts of the body, mainly the scales, in various medicinal customs has resulted in a marked decline in local populations . As a result, the entire population is now viewed as endangered. It was first described scientifically in 1821. The white-bellied pangolin presumably includes several cryptic species .

features

Habitus

The white-bellied pangolin is one of the smallest pangolins along with the closely related long-tailed pangolin ( Phataginus tetradactyla ). It reaches a head-trunk length of 35 to 43 cm, the tail is around 34 to 62 cm long. This makes the tail about one and a half times as long as the rest of the body, but relatively shorter than that of the long-tailed pangolin. The weight varies from 1.6 to 3.0 kg, males are usually larger than females. As with all pangolins, the top of the head, the back and the flanks, the outside of the limbs (but not the forearms) and the tail are covered with horny scales. The scales show a uniform color, which varies from gray-brown to reddish-brown to brown-yellow. The scales are relatively small, longer than wide, and have three points pointing backwards. They form 19 to 22, sometimes also 25, transverse rows on the body. A middle row running lengthways across the back extends to the tail. There it consists of 30 to 33 scales, but breaks off shortly before the tail end and is replaced by a row of 3 to 6 pairs of scales. There are 34 to 37 scales on the sides of the tail. The lower end of the tail is not covered with scales, instead the white-bellied pangolin has a skin pad there. The skin on the unscaled parts of the body has a brownish hue. On the belly it is covered with whitish, thin and long hair, which turns into a brownish color on the legs. The head shows a conical shape, the snout is thicker compared to the long-tailed pangolin, which makes the skull appear more massive. It is only sparsely hairy and has dark spots under the eyes. The eyes are small with dark irises and protrude. Ear bulges are not formed, but a dense tuft of hair surrounds the ear opening. The front legs are slightly shorter than the rear legs. All limbs end in five toes with strong, curved claws. The middle of the front feet is elongated into a large grave claw that is twice as long as the others. The rear foot reaches a length of 4.4 to 5.4 cm.

Skull and skeletal features

Skull of a white-bellied pangolin

The skull of the white-bellied pangolin is between 6 and 8 cm long. The spine is made up of 7 cervical, 13 thoracic, 6 lumbar, 2 sacrum and 41 tail vertebrae, making a total of 69 vertebrae. The tail thus has a little less vertebrae than that of the long-tailed pangolin.

distribution and habitat

Distribution area (green) of the white-bellied pangolin

The range of the white-bellied pangolin covers parts of West and Central Africa . It extends from Guinea to the countries of the Congo Basin to about the northernmost areas of Angola . In the east it can still be found in the southwest of Kenya as well as in the northwest of Tanzania and Zambia . Information about occurrences in the north of Mozambique and Malawi is rather doubtful. The pangolin mainly inhabits tropical rainforests , but it also occurs in mosaic landscapes and more open savannah landscapes , as well as in secondary forests . In some cases it has also been observed in abandoned agricultural areas, for example on abandoned or rarely used oil palm plantations. This suggests that the animals tolerate a certain degree of landscape influence. Over large parts of the distribution area, the white-bellied pangolin appears sympatric with the long-tailed pangolin, which, however, prefers more swampy regions. Observations in Nigeria indicate that the white-bellied pangolin is significantly more common than its close relative. In food-rich areas, the population density can be relatively high. For the Lama Forest Reserve in Benin , 38 white-bellied pangolins were determined over 45 km², which corresponds to 0.84 individuals per square kilometer. No major difference in the frequency of the pangolin species between the primary forests in the center of the protected area and the surrounding old plantations could be determined.

Way of life

Territorial behavior

White-bellied pangolin

Along with the long-tailed pangolin, the white-bellied pangolin is one of the most distinctive tree dwellers among the pangolins, but deviating from this it is much more active at night. Its activity phase begins between 6:00 p.m. and midnight. It lasts about three to four hours in females and up to ten hours in males. The white-bellied pangolin usually rests in tree hollows about 10 to 15 m above the ground. In Benin, animals could preferably be observed in caves of the kapok tree or of Dialium guineense . Sometimes they also dig small caves in the ground 30 to 40 cm deep. The exceptionally long tail of the white-bellied pangolin is a clear adaptation to tree life. It serves as a pronging tail and, as a “fifth limb”, can hold the weight of an animal alone for a long time. In the trees it climbs with the help of its claws and tail, which is often rolled around branches and trunks to support it. The front and rear legs are moved in pairs, creating a typical caterpillar-like gait with a flexing and stretching back. When descending from the tree, an animal moves in a spiral around the trunk. Sometimes it also drops down curled up in a ball, which is seen as a reaction to breaking branches. The white-bellied pangolin uses various modes of locomotion on the ground. The most common is the normal four-footed gait, in which it reaches speeds between 1.0 and 1.5 km / h and the front feet touch the surface of the foot. When running faster, it also falls into a caterpillar-like movement, with the tail then held elevated. Most of the weight is on the hind legs. The pangolin can swim very well. The tail moves undulating like a snake and gives the propulsion. Before swimming, an animal takes in extra air, so that the body diameter can increase by up to 11 cm. This additional air ensures that the body can be kept afloat. After swimming, the air is expelled, which is accompanied by a trumpet-like sound.

The white-bellied pangolin lives mostly solitary and uses action spaces that are 3 to 4 hectares in size for female individuals, but more than seven times the area for males with over 30 hectares. The latter cover up to 1.8 km on their nightly forays, which is relatively little compared to their large area of ​​action. In the action area there are several tree hollows that are used alternately. Male animals change their shelter daily, females less often. The territories of the males can overlap with up to ten of the females. Males tolerate females in their territory, but chase away solitary pups. There is a certain territoriality between male animals; fights are fought with synchronized blows of the front legs. At least in captivity, individual animals were killed. Social communication takes place through the excellent sense of smell . The grazing area is marked with a secretion from the anal glands . Secretions from the perineal glands are used in aggression and sexual contact.

nutrition

The diet of the white-bellied pangolin consists of insects that form colonies , which means that the pangolin lives strictly myrmecophagus . In contrast to the long-tailed pangolin, it prefers termites , especially of the genera Nasutitermes and Microcerotermes , from which it devours the adult individuals and the nymphs . The white-bellied pangolin eats ants less often . Among these, wandering ants such as Dorylus or Mirmicaria predominate , but species of the genera Camponotus , Cataulcus , Crematogaster or Oceophylla also belong to the food spectrum. The animals eat both arboreal and ground-dwelling colonies, but the latter predominate in males and subadult individuals. The food is tracked down with the excellent sense of smell, the white-bellied pangolin often searches under fallen trees or in the leaves. It attacks the nests of the insects from several sides and often breaks them open in several places with the claws of the front feet. It devours the prey with the help of its long, sticky tongue, which can be up to 30 cm long. As a rule, the nests are not completely destroyed, but individual animals return to the same food source several times in a row. Females usually look for nests near the burrow, only a few hundred meters away, and eat for around three to four hours a day. In contrast, males move further away from the burrow and spend five to six hours eating. The white-bellied pangolin can eat between 150 and 200 g of insects every day. After eating, an animal often rolls around on the floor, slips over it, rubs against objects or spreads the scales and scratches its claws to crush insects that have crawled between the scales while defending the nest.

Reproduction

The reproduction takes place all year round. Male animals cross several action areas every day to look for receptive females. The rut begins in females every 3 to 29 days, with an average interval of 9 days. The foreplay consists of ritualized chest-to-chest battles. After that, both partners climb a tree, usually the female clings to the male's tail. During the sexual act that takes place in the tree, the tails of the two animals are intertwined. The gestation period is often given as around 150 days, individual observations are from 112 to 168 days. However, it could also extend over a period of 7 to 9 months. In ten pregnant females examined in captivity, a good half of the birth was problematic and ended with the death of the young and / or the mother. Usually only one young is born. This is about 29 cm long and 100 g in weight. It has open eyes, a pink skin tone, and is hairless except for the eyelids. The young spends the first one to two weeks in a tree hollow, after which it leaves it riding on the root of its mother's tail. The sucking phase ends after around four months and the weight is then around 750 g. With the birth of the next young animal, the older boy leaves the mother animal and goes on a journey with no fixed space for action. Sexual maturity is reached at around eight months, after 15 months the boy weighs over 1 kg and is then fully grown, from this time onwards it lives in its own space of action.

Predators and enemy behavior

The honey badger as predator of the white-bellied pangolin

As the main predator kicking Leopard on. Investigated Kotreste the big cat in Lopé- and Ivindo National Park in Gabon but show only a small part in the wide range of prey. The same applies to the African golden cat , whose feces were examined more closely in the Ituri forest in the northeastern part of the Congo Basin . In addition, the honey badger and jackals chase after the white-bellied pangolin. Some animals are victims of the northern rock python and chimpanzees . Hunting often takes place on the ground. In the case of immediate danger, an animal expels secretions from its glands or rolls up and covers unscaled parts of the body with its tail, otherwise it often takes refuge in a tree.

Parasites

External parasites are ticks of the genus Amblyomma , which usually nest under the scales of the neck and back. In investigations of 26 white-bellied pangolins from different regions of Ghana , the tick genus was found in all individuals, and a member of the genus Haemaphysalis was extremely rare . As internal parasites are especially nematodes and to the coccidia scoring Eimeria known.

Systematics

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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The white-bellied pangolin is a species from the genus Phataginus , which contains another species with the long-tailed pangolin ( Phataginus tetradactyla ). Both species represent the tree-dwelling pangolins of Africa, which, according to molecular genetic analyzes , separated from each other in the Middle Miocene about 13.3 million years ago. Phataginus forms the independent subfamily of the Phatagininae . Opposite it is the subfamily of the Smutsiinae with the genus Smutsia , in which the ground-dwelling pangolins of Africa are united. The Asian pangolins of the subfamily Maninae with the genus Manis are the sister taxons of the two African lines . All three groups together form the family of pangolins (Manidae). The pangolins comprise the currently only member of the order of the Pholidota, so these are recently monotypical . The group is largely related to the predators (Carnivora), but the relationship to each other was only determined and verified through molecular genetic studies.

However, in some other attempts to classify the pangolins, the genus Manis , which here includes the Asian representatives, is regarded as the only recognized genus of the pangolin. All other genera including Phataginus then have the status of subgenera. On the other hand, there is also the opinion that the pangolins are much more fragmented. Here in turn the white-bellied pangolin is the only representative of the genus Phataginus , the closely related long-tailed pangolin is then in the genus Uromanis . The division of the pangolin family into the three genera Manis , Phataginus and Smutsia , which is favored today , was first proposed at the end of the 1990s. Subsequent anatomical and phylogenetic studies supported this view.

Some authors differentiate between two subspecies of the white-bellied pangolin:

The type specimen from M. t. mabirae comes from Uganda , where it was collected by Arthur Loveridge in 1938, and has a few different skull features and an orange to cinnamon-colored peritoneum.

Internal systematics of the white-bellied pangolin according to Gaubert et al. 2016
  Phataginus tricuspis  



 Dahomey Gap


   

 Ghana


   

 West Africa




   

 West Central Africa



   

 Central Africa


   

 Gabon




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Constantine S. Rafinesque-Schmaltz

The division into several subspecies is not generally recognized due to the high variability of the white-bellied pangolin, which is why it is mostly viewed as monotypical. Molecular genetic studies from 2015 reveal large differences between the western and eastern populations, which, according to the authors of the study, may advocate differentiation at species level. This view is supported by the fact that there is a biogeographical barrier between Cameroon and Gabon, the so-called charnière climatique , which separates the two populations. Further analyzes from the following years on over 100 white-bellied pangolin show a much more complex structure. A total of six phylogenetic lines can be identified, each of which is likely to represent distinct species. The breakdown of this widely ramified group began in the transition from the Pliocene to the Pleistocene around 2.7 million years ago. In addition to genetic diversity, there are also differences in skull structure between the individual populations. No fossil finds of the pangolin are known.

The white-bellied pangolin was first scientifically described in 1821 by Constantine S. Rafinesque-Schmaltz under the name Manis tricuspis . Rafinesque gave the type locality as Guinée , but he saw his Manis tricuspis as a synonym for Manis tetradactyla , the long-tailed pangolin, which was named by Linnaeus in 1766 . In the same work, Rafinesque set up the species Manis ceonyx , which today is considered identical to the long-tailed pangolin . For the two species specified by him, Rafinesque established Phataginus as a new subgenus for Manis . For Phataginus he determined differences in the construction of the foot compared to the then known Chinese pangolin ( Manis pentadactyla ) as an Asian representative of Manis , to which Rafinesque certified an allocation to the subgenus Pangolinus . For the white-bellied pangolin, Rafinesque also introduced the colloquial name Phatagin tricuspidé . The specific epithet tricuspis refers to the three tips of the scales.

Threat and protection

The white-bellied pangolin is heavily hunted. Its meat is considered a delicacy and is sold locally to markets where it is offered as bushmeat . In addition, the scales and other parts of the body are used in traditional medical practices in Africa, such as juju , as they are said to have healing powers. The Yoruba in Nigeria use body parts of the white-bellied pangolin for more than 40 diseases, such as the dandruff for stomach problems or diarrhea , the meat for calming down or the eyes for kleptomania . Between April and July 2007 alone, traders in markets in the Nigerian state of Ogun offered 178 white-bellied pangolin for medical purposes, according to research. Studies in the Bombali region of Sierra Leone have shown that a total of 22 different body parts are used to heal a wide variety of diseases, with dandruff by far predominating. The white-bellied pangolin is also increasingly entering international trade, with the greater part being exported to East Asia , where pangolins are used in traditional Chinese medicine . In 2011, at least 100 hides with scales from the white-bellied pangolin were confiscated in Belgium. Because of these three threats, the pangolin species has declined sharply in the past. It is estimated to be extinct in Rwanda . According to reports from hunters, the number of animals in Nigeria was already declining in the 1990s. Since the year 2000 the trade in the white-bellied pangolin or its body parts has been forbidden according to the Washington Convention on Endangered Species (CITES); it is subject to the CITES zero annual export quota . In addition, habitat destruction through deforestation is putting further pressure on local populations, which is particularly the case in West Africa. In some cases, animals are also victims of road accidents. The IUCN therefore lists the species as "endangered" ( vulnerable ). The white-bellied pangolin is present in several protected areas. Further research into the pangolin species' way of life and the effect of hunting pressure on the individual populations are necessary for the conservation of the populations. The development of national protection standards is also necessary, as is the investigation of the routes in international trade.

literature

  • Phillipe 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 (pp. 102–103)
  • Jonathan Kingdon and Michael Hoffmann: Phataginus tricuspis Tree Pangolin (African White-bellied Pangolin). In: Jonathan Kingdom, David Happold, Michael Hoffmann, Thomas Butynski, Meredith Happold and Jan Kalina (eds.): Mammals of Africa Volume V. Carnivores pangolins, equids and rhinoceroses. Bloomsbury, London, 2013, pp. 391-395
  • Ronald M. Nowak: Walker's Mammals of the World . Johns Hopkins University Press, 1999 ISBN 0-8018-5789-9
  • Don E. Wilson, DeeAnn M. Reeder (Eds.): Mammal Species of the World . 3rd edition. The Johns Hopkins University Press, Baltimore 2005, ISBN 0-8018-8221-4 .

Individual evidence

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Web links

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