Manis tetradactyla Linnaeus 1766
Creators: name: Don E. Wilson
name: Russell A. Mittermeier
Publisher: name: Lynx Edicions
Year: 2011
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Pholidota
Manidae
Manis
Manis tetradactyla

description: 7.
Long-tailed Pangolin
Manis tetradactyla
French: Pangolin a longue queue / German: Langschwanzschuppentier / Spanish: Pangolin colilargo
Other common names: Black-bellied Pangolin
Taxonomy. Manis tetradactyla Linnaeus, 1766,
West Africa.
Sometimes included in the genus/subgenus Uromanis. Monotypic.
Distribution. W & C African rainforest blocks, from Sierra Leone to Ghana, and from Niger Delta to Congo Basin, as far as C & E DR Congo. Presence in NW Angola (Cabinda) is possible.
Descriptive notes. Head—body 30-40 cm, tail 55-70 cm; weight 2.3-5 kg. A small, golden-scaled and black-bellied pangolin, with a very long tail. One pair of pectoral mammae. Sexual dimorphism unknown. Head conical and rostrum thin and short, giving a V-shape to the head in dorsal view. Nose pad color similar to the dark skin. Auditory orifice without ear pinna. Eyes small and iris dark. Scaled armor with large scales covering the upper face and the whole body, except the belly, the upper foreand hindfeet, and the inner side of legs. Hairs dark brown to black, thin and very long (between 0-5 cm and 1 cm), densely covering all the naked surfaces; no hairs projecting between scales. 13 dorsal scale rows. Width of scales of the distal part of the body similar to that of first rows of post-scapular scales; from base to tip, each scale shows a longitudinal, dark to golden color gradient. Two pairs of post-scapular scales very large, covering the upper forelegs. Tail much longer than head and body. On tail, medio-dorsal row of scales interrupted before tip. Ventral part oftail showing a large, terminal skin pad, due to the absence of two median and two lateral scales. Forelegs slightly shorter than hindlegs. Five short and curved claws on forefeet, with the third claw at least twice as long as the others; walks with wrist folded, with claws perpendicular to ground. The five claws on hindfeet are markedly curved and kept longer than in terrestrial species. Skull 6-7 cm long, V-shaped. Number of caudal vertebrae: 50.
Habitat. Tropical, riparian, and swamp forests, always close to permanent water. Can be considered semi-aquatic. Such habitat specialization may be exacerbated by sympatry with the Common African Pangolin (M. tricuspis). In the Niger Delta, reported from secondary and altered rainforests.
Food and Feeding. Probably strictly myrmecophagous; genera preyed upon are arboreal ants (Crematogaster and Cataulacus). In captivity, does not usually accept termites. Ants are eaten on trees. Tree nests are broken or taken down with forefoot claws, and attacked from all sides, until ants fall down; a tunnel may be dug inside the nests. Nests generally are not destroyed, and thus constitute continuous feeding sources that are visited regularly. Uses olfaction to forage, and extracts ants using the protractile part of the tongue (16-18 cm long). Distal part of the tongue is different from that of M. tricuspus: it is flattened, not pointed, and has an oval cross-section; it has a zone of 4 cm covered with bristles 1 mm long, pointing backwards. The tongue is attached behind the larynx and trachea to a characteristically modified xiphisternum (xiphoid process). Xiphoid cartilages form two elongated bars that extend from the sternum to the pelvic region.
Breeding. Breeding is non-seasonal and apparently continuous. Sexual behavioris extremely elaborate. Preliminaries simulate aggression, chest against chest, followed by female’s submission. The female then clings on the tail of the male (like a young to its mother), and the couple moves to a tree before mating. Tails of male and female are entwined when copulating. Female conceives again within two weeks after birth. Gestation period is around 140 days; placentation is endotheliochorial. A single young is born; stays in a tree hole during the first week, before clinging to mother’s back or tail to get carried. Starts ingesting living prey at about two weeks. Leaves mother at next birth, and wanders for 4-5 months before settling into its own area.
Activity patterns. Solitary, almost strictly arboreal and diurnal. Adults rest in tree hollows and use habitual trails to move; they seem to have a fixed home range, with a series of shelters used in alternation. Younger individuals are wanderers, with no predefined routes; they do not use shelters to rest, but instead simply roll up on branches. If resting in a tree is impossible, an individual may dig a simple, horizontal den in talus. In captivity, observed spending more time above the ground than M. tricuspis; the proportion of arboreal activity reached almost 100% during courtship. Moves slowly. Frequently uses the caterpillar-like walk, with forelegs moving forward and subsequently joined by hindlegs, the pangolin successively arching and stretching its back. This walk is notably used when running. In case of danger, runs away with nose kept close to ground to scent the trail. When running after a congener or following its tracks, rubs its anal glands on supports leaving odoriferous marks. When attacking a congener (especially between males), caterpillar-like walk turns into a series ofjumps forward, the animal standing and pushing on hindlegs; the same behavior has been observed when moving through high grasses. The caterpillar-like walk is also used when moving on 5-20 cm diameter branches: foreand hindlegs are used as clamps, with claws jabbing at branches. Uses the diagonal walk (similarly to M. tricuspis) when moving on terminal, thin branches and creepers, hanging by its claws in a sloth-like manner. Climbing is facilitated by very flexible feet on the vertical axis—butless in torsion (i.e. no supination)—and the sharp, curved claws. Forefoot digits two and three not completely fused so they can spread to hold small branches. The long neck and supple body make the animal reptile-like when using diagonal walk. Can slip into openings 5 cm wide and then enlarge the entrance by powerfully moving the body. Long, prehensile tail with a highly sensitive distal pad, used like a third hindleg. The tail coils tightly thanks to the pangolin’s strong trunk and tail muscles. When climbing large and/or smooth trunks, the tail is curled around the trunk, the animal progressing in a spiral. The scales, large and spread, cling to the uneven surface of the tree. The tip of the tail is also curled around branches and used as a pivot to move from branch to branch ortree to tree. When in danger on the ground, the Long-tailed Pangolin looks for a tree it can climb to an unreachable height. Otherwise, it rolls into a ball, with the tip of the tail folded up onto the scales of the neck, locked into a defensive position.
Movements, Home range and Social organization. The least frequently recorded of African pangolins, although a very specialized mode oflife may be responsible for its apparent low density. Movements of individuals remain unknown. Extremely secretive, well camouflaged in the foliage with its scales that imitate the effect of sunlight on leaves, and endemic to little penetrated habitats. The sole pangolin to be considered semi-aquatic. Frequently travels in streams and inundated areas. Good swimmer, using undulating (snake-like) movements, with tail playing a prominent role for the locomotion in water. Body halfimmersed due to ingestion of extra air while swimming, body diameter increasing by more than 10 cm; breathes regularly at the surface, head position alternating under and above water. After leaving water, emits a trumpet-like sound, emptying extra air from nose and probably anus. When moving, follows trails marked by urine and secretions from anal scent glands; these latter are also expelled to discourage predators. Communication among individuals is mainly olfactory. Perianal glands are involved in marking, sexual, and aggressive behavior, whereas anal glands play a role in marking and repulsive behavior. Olfactory marking permits longdistance regulation of interindividual contacts.
Status and Conservation. CITES Appendix II. Classified as Least Concern on The [IUCN Red List because ofits large distribution, presumably healthy populations, presence in several protected areas, and secretive mode oflife. In addition, its confinement to inaccessible habitats likely spares the species from intense bushmeat activities and forest logging. Nevertheless, requires further monitoring given its poorly known natural history and the existence of local, selective hunting for meat consumption and traditional medicine. In Cameroon, sold on bushmeat markets as a rare item under the name “Pangolin sorcier” (pangolin sorcerer).
Bibliography. Angelici et al. (1999, 2001), Botha & Gaudin (2007), Hatt (1934), Heath & Amachree (1967), Hoffmann (2008d), Jouffroy et al. (1975), Kingdon (1997), Pages (1970, 1972a), Rahm (1956).
description: Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Manidae, pp. 82-103 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on pages 101-102, DOI: 10.5281/zenodo.5720458