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Manispangolins

By Mea Escobedo

Diversity

Pangolins are one of the most highly trafficked mammals in the world, causing all of the extant species to be endangered or critically endangered (Gaubert et al., 2017; Katuwal, Sharma, and Parajuli, 2017; Shepherd et al., 2017). There are four extant species in genus Manis: Manis crassicaudata, Manis culionensis, Manis javanica, and Manis pentadactyla (Cabana et al., 2017; Gaubert et al., 2017; Thapa 2013). (Cabana, et al., 2017; Gaubert, et al., 2017; Katuwal, et al., 2017; Shepherd, et al., 2017; Thapa, 2013)

Manis species are primarily terrestrial creatures. They are myrmecophagous mammals with tough, keratinized dermal scales that cover their body and aid in protection from predators. When threatened, they roll into a ball, leaving only their hardened scaly exterior exposed which deters predation. Manis species have been described as solitary and reclusive in nature (Thapa 2013). (Thapa, 2013)

Geographic Range

Manis species are found throughout the Oriental region and a small portion of the southern Palearctic region (Schlitter, Wilson, and Reeder, 2005). There are no known introduced ranges. (Schlitter, et al., 2005)

Habitat

Manis species have been observed most commonly at elevations of 1,450 to 1,550 m, with some sightings over 2,000 m (Bhandari and Chalise, 2014; Kaspal 2009). They occupy a diverse array of habitats, such as agricultural areas, foothills, forests (bamboo, broadleaf, coniferous, limestone, primary, secondary, subtropical, tropical), grasslands, savannahs, and woodlands (Mahmood, Irshad, and Hussain, 2012; Thapa 2013). Manis species are commonly observed excavating and residing in burrows made of soft, semi-sandy, red and brown soils (Bhandari and Chalise, 2014). (Bhandari and Chalise, 2014; Kaspal, 2009; Mahmood, et al., 2012a; Thapa, 2013)

Systematic and Taxonomic History

Manis species have been synonymously referred to as Manis by Linnaeus in 1758, Pholidotus by Brisson in 1762, Pangolinus by Rafinesque in 1815 and 1821, Phatages by Sundevall in 1843, Phatagin by Gray in 1865, Triglochinopolis by Fitzinger in 1872, and Pangolin by Gray in 1873 (Wilson and Reeder, 2017). (Wilson and Reeder, 2017)

Current evolutionary understanding of relationships within the order Pholidota support only 1 main group: family Manidae. Manidae can be divided into 2 subgroups: Asian pangolins and African pangolins. African Pangolins consist of 2 genera that are closely related to each other (Gaudin, Emry, and Wible, 2009). Manis species constitute the Asian pangolins. (Gaudin, et al., 2009)

Research on synapopmorphies specifically in relation to Manis species does not exist, but research on synapopmorphies within order Pholidota does. Gaudin, Emry, and Wible reported the following as being synapomorphies in their morphological analysis from 2009: transverse width of dorsal surface of lateral cuneiform roughly equal to proximodistal height; fibular facet of astragalus crescent-shaped, with concavity facing proximoplantarly; manual and pedal subungual processes form triangular platform in ventral view, with grooves along either side of subungual processes leading to subungual foramina; obturator foramen small, maximum diameter of acetabulum ≥75 % that of obturator foramen; prominent ischial spine; temporal lines absent; basicranial/basifacial axis reflexed; teeth absent; horizontal ramus of mandible shallow, ≤10% of maximum mandibular length (Gaudin, Emry, and Wible, 2009). (Gaudin, et al., 2009)

  • Synonyms
    • Pholidotus
    • Pangolinus
    • Phatages
    • Phatagin
    • Triglochinopolis
    • Pangolin
  • Synapomorphies
    • Transverse width of dorsal surface of lateral cuneiform roughly equal to proximodistal height
    • Fibular facet of astragalus crescent-shaped, with concavity facing proximoplantarly
    • Manual and pedal subungual processes form triangular platform in ventral view, with grooves along either side of subungual processes leading to subungual foramina
    • Obturator foramen small, maximum diameter of acetabulum ≥75 % that of obturator foramen
    • Prominent ischial spine
    • Temporal lines absent
    • Basicranial/basifacial axis reflexed
    • Teeth absent
    • Horizontal ramus of mandible shallow, ≤10% of maximum mandibular length

Physical Description

Manis species have several adaptations that allow them to successfully hunt insects, such as being edentate, having a long sticky tongue, strong forelimbs, sharp claws, and tough scales (Ashokkumar et al., 2017; Mahmood et al., 2012; Mahmood et al., 2013; Schlitter, Wilson, and Reeder, 2005; Shepherd and Shepherd, 2012). Male and female Manis species are similar barring size. Males are slightly larger than females. Juvenile Manis species resemble adult form. There is no known seasonal variation among Manis species. (Ashokkumar, et al., 2017; Mahmood, et al., 2012b; Mahmood, et al., 2013; Schlitter, et al., 2005; Shepherd and Shepherd, 2012)

Manis crassicaudata are 84 to 122 cm long (including tail) with a tail length of 33 to 47 cm. Manis crassicaudata are 10 to 16 kg (Mahmood et al., 2013; Mahmood et al., 2012). (Mahmood, et al., 2012b; Mahmood, et al., 2013)

Manis culionensis are 58 to 176 cm long (including tail) with a tail length of 28 to 88 cm. Manis culionensis are 1.8 to 2.4 kg (Gaubert and Antunes, 2005). (Gaubert and Antunes, 2005)

Manis javanica are 75 to 121 cm long (including tail) with a tail length of 35 to 56 cm. Manis javanica are up to 10 kg (Schlitter, Wilson, and Reeder, 2005; Shepherd and Shepherd, 2012). (Schlitter, et al., 2005; Shepherd and Shepherd, 2012)

Manis pentadactyla are 65 to 96 cm long (including tail) with a tail length of 25 to 38 cm. Manis javanica are 2 to 7 kg (Shepherd and Shepherd, 2012). (Shepherd and Shepherd, 2012)

  • Sexual Dimorphism
  • male larger

Reproduction

The breeding habits of Manis species have not been studied extensively. Much is unknown in regards to how Manis species find, attract, and defend mates. How mating behaviors affect social structure is also unknown.

Manis crassicaudata have been observed sharing the same burrow with a mate and offspring, even though Manis species are usually solitary. Females give the majority of parental care (Mahmood et al., 2016). (Mahmood, et al., 2016)

Although, information on the reproductive habits of g. Manis species is limited, some studies on Manis crassicaudata and Manis javanica exist.

Manis crassicaudata appear to mate annually, from July to October. On average 1 to 2 offspring are produced after a gestation period of about 80 days (Mahmood et al., 2016). Newborns weigh 235 to 400 g and measure about 30 cm at birth (Mohapatra and Panda, 2014). Manis crassicaudata reach sexual maturity around 2 years (Mahmood et al., 2016). (Mahmood, et al., 2016; Mohapatra and Panda, 2014)

Manis javanica appear to breed year-round; typically producing 1 offspring after a gestation period of about 6 months. Offspring are precocial and juveniles are weaned at about 4 months. Manis javanica reach sexual maturity from 6 months to 1 year (Zhang et al., 2017; Zhang et al., 2015) (Zhang, et al., 2015; Zhang, et al., 2017)

Much is unknown about the degree of parental investment in Manis species.

Female Manis crassicaudata appear to invest more in offspring than male Manis crassicaudata; females have been observed carrying and protecting offspring (Mahmood et al., 2016). (Mahmood, et al., 2016)

Lifespan/Longevity

Information about the lifespan of Manis species is unknown.

Behavior

Manis species are solitary in nature, only coming together to mate and rear young (Thapa 2013; Mahmood et al., 2016). (Mahmood, et al., 2016; Thapa, 2013)

Manis species do not have a known hierarchical structure.

Communication and Perception

Research on communication among the Manis species is scarce. Manis species do not appear to interact with one another often, outside of mating or offspring care (Thapa 2013; Mahmood et al., 2016). (Mahmood, et al., 2016; Thapa, 2013)

Food Habits

Manis species are insectivorous, specifically adept in myrmecophagy. These species have several adaptations that allow them to be successful predators of g. Leptogenys species: long sharp claws and strong forelimbs (Ashokkumar et al., 2017), an elongated tongue for probing, and fibrous hair and keratinized scales that protect from bites and stings (Mahmood et al., 2012; Mahmood et al., 2013; Schlitter, Wilson, and Reeder, 2005; Shepherd and Shepherd, 2012). (Ashokkumar, et al., 2017; Mahmood, et al., 2012b; Mahmood, et al., 2013; Schlitter, et al., 2005; Shepherd and Shepherd, 2012)

Predation

Manis species curl into a ball when threatened (Wang et al., 2016). This protects them from predators since only their hardened, keratinized scales are left exposed. Manis species may also climb into trees as a last resort to avoid predation. (Wang et al., 2016; Challender et al., 2014). Predators of Manis include: Homo sapiens, Lycaon pictus, Panthera pardus, Panthera tigris, and Pythonidae species (Tenaza, 1975; Thapa, 2013). There are no known difference in the way Manis species react to different predators. (Challender, et al., 2014; Tenaza, 1975; Thapa, 2013; Wang, et al., 2016)

  • Known Predators
    • Homo sapiens
    • Lycaon pictus
    • Panthera pardus
    • Panthera tigris
    • Pythonidae

Ecosystem Roles

Manis species fill an ecological role as an insectivore. Their presence helps to keep insect populations under control (Thapa, 2013). (Thapa, 2013)

Manis species also create habitats and aerate soil by digging burrows (Thapa, 2013). (Thapa, 2013)

Manis species serve as a host for several organisms: Amblyomma javanese, Cylicospirura species, and Strongyloides species (Heath and Vanderlip, 1988; Li et al., 2010). (Heath and Vanderlip, 1988; Li, et al., 2010)

Commensal/Parasitic Species
  • Amblyomma javanese
  • Cylicospirura
  • Strongyloides

Economic Importance for Humans: Positive

Manis species help control insect populations (Thapa, 2013) and provide resources for humans such as: their meat, fat, and scales (Mahmood et al., 2012). (Mahmood, et al., 2012b; Thapa, 2013)

Economic Importance for Humans: Negative

There are no known adverse effects of Manis species on humans.

Conservation Status

Manis species are endangered to critically endangered. Populations have been dramatically impacted by illegal hunting and trading of their species. Many people harvest these creatures for their scales, meat, and fat for uses in Chinese medicine, food, and clothing (Mahmood et al., 2012). (Mahmood, et al., 2012b)

  • IUCN Red List [Link]
    Not Evaluated

Other Comments

The only available hypothesis for the etymology of the name Manis comes from the Latin word Manes, meaning spirits. Although, this hypothesis is contradictory to the feminine way Linnaeus appeared to name this genus because Manes is a plural and masculinized word for spirits (ITIS, 2017). (ITIS, 2017)

Manis species have highly impacted Chinese culture. Their scales, meat, and fat is often used in Chinese medicinal practices, as food, and as clothing (Mahmood et al., 2012). (Mahmood, et al., 2012b)

Contributors

Mea Escobedo (author), Colorado State University, Tanya Dewey (editor), University of Michigan-Ann Arbor.

Glossary

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

crepuscular

active at dawn and dusk

drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

ecotourism

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

fossorial

Referring to a burrowing life-style or behavior, specialized for digging or burrowing.

insectivore

An animal that eats mainly insects or spiders.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

male parental care

parental care is carried out by males

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

pet trade

the business of buying and selling animals for people to keep in their homes as pets.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

soil aeration

digs and breaks up soil so air and water can get in

solitary

lives alone

tactile

uses touch to communicate

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

year-round breeding

breeding takes place throughout the year

young precocial

young are relatively well-developed when born

References

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