Behavior

Some species of pangolins, the larger ones, are classified as terrestrial (ground-dwelling) (Manis gigantea, Manis temminckii, Manis crassicaudata, and Manis javanica) while others, the smaller species, are classified as arboreal (tree-climbing) (Manis pentadactyla, Manis tetradactyla, and Manis tricuspis). Whether classified as terrestrial or arboreal, some species can dwell both on the ground and in trees.

All species of pangolins are nocturnal except for the predominantly diurnal Manis tetradactyla. Because pangolins are active mostly at night, they have poor vision. Their ability to

Pangolins may use their prehensile tails to anchor themselves in a tree while they break open termites' tree nests. (Photo by A. Blank/ Photo Researchers, Inc. Reproduced by permission.)

hear is only average, and is used little in intraspecific relationships. They make no well-distinguished sounds; with the only noticeable utterances being hisses and snorts. However, large olfactory lobes allow their sense of smell to be exceptionally sharp, playing the major role in finding food and, in all likelihood, playing a major role in intraspecific communication. They have very few expressive behaviors, the most frequent being the closing of the eyes and the sticking out of the tongue. Pangolins are generally shy, solitary, and unsociable creatures who only rarely associate with a conspecific.

Panglolins are slow and deliberate movers. Most are good climbers, and some also swim. The terrestrial species will rest and sleep during the day in burrows that they have previously dug (or those dug by other animals), while the arboreal species find haven in hollow trees. For additional protection from small- and medium-sized enemies, they roll up (basically) into a ball when sleeping. They do not actively defend a fixed territory from neighboring animals of the same species. However, they do repeatedly mark selected trees, branches, and rocks with secretions from their anal glands and urine, which announces to neighboring and trespassing animals that the area is already occupied. The terrestrial species regularly mark their territories with droppings, and arboreal species may do so as well.

All species can move quickly when alarmed. If unable to reach shelter, they will often curl into a tight ball with their scaled limbs and tail protecting the soft under-parts. When in this position the sharp-edged scales are erected as protective armor. Movements of the scales and twitches of the tail apparently deter predators. They do not appear to use their fore-claws in defense. The animals will also expel urine and posterior gland secretion in defense of their immediate safety. Often a stream of urine is released if a pangolin is unrolled by force. Feces may also be released, which is often combined with secretion from the anal glands.

Pangolins are plantigrade; that is, they walk on the soles of their feet with heels touching the ground. When they walk on all four limbs their toes are doubled under and the tips of the long nails are placed on the ground. The large digging claws of the forefeet are held inward by terrestrial species, which walk on the outer sole. The arboreal species bend the claw downward when traveling on the ground. In walking on the ground, primarily the hind limbs are used. They stop periodically and raise themselves up, supporting themselves on

A pangolin searches for insects in the Democratic Republic of the Congo (Zaire). (Photo by K & K Ammann/Photo Researchers, Inc. Reproduced by permission.)
Ground pangolins (Manis temminckii) live in central Africa to northern South Africa. (Photo by Nigel J. Dennis/Photo Researchers, Inc. Reproduced by permission.)

their hind legs and tail. In this position, with the front of the body erect and the head protruding, they smell or look out for possible enemies. At a fast pace, the tail is lifted from the ground.

The giant pangolin (M. gigantea), ground pangolin (M. ter-mminckii), long-tailed pangolin (M. tetradactyla), and tree pangolin (M. tricuspis) are all good swimmers, holding only the head above water while swimming with a "doggy paddle" motion. They are able to cross narrow rivers easily, but tire on longer journeys.

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