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).
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).
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.
- Biogeographic Regions
- palearctic
- oriental
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).
- Habitat Regions
- tropical
- terrestrial
- Terrestrial Biomes
- savanna or grassland
- forest
- rainforest
- mountains
- Other Habitat Features
- agricultural
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).
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.
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).
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.
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).
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).
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).
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).
- Other Physical Features
- endothermic
- homoiothermic
- bilateral symmetry
- 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).
- Mating System
- monogamous
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).
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)
- Key Reproductive Features
- iteroparous
- seasonal breeding
- year-round breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- viviparous
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).
- Parental Investment
- precocial
- male parental care
- female parental care
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
-
pre-independence
-
protecting
- female
-
protecting
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).
Manis species do not have a known hierarchical structure.
- Key Behaviors
- terricolous
- fossorial
- nocturnal
- crepuscular
- motile
- solitary
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).
- Other Communication Modes
- pheromones
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).
- Primary Diet
- carnivore
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.
Ecosystem Roles
Manis
species fill an ecological role as an insectivore. Their presence helps to keep insect
populations under control (Thapa, 2013).
Manis
species also create habitats and aerate soil by digging burrows (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).
- Ecosystem Impact
- creates habitat
- soil aeration
- 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).
- Positive Impacts
- pet trade
- food
- body parts are source of valuable material
- ecotourism
- source of medicine or drug
- research and education
- controls pest population
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).
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).
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).
Additional Links
Contributors
Mea Escobedo (author), Colorado State University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- 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.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- 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.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- agricultural
-
living in landscapes dominated by human agriculture.
- 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.
- 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.
- monogamous
-
Having one mate at a time.
- 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).
- seasonal breeding
-
breeding is confined to a particular season
- year-round breeding
-
breeding takes place throughout the year
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- young precocial
-
young are relatively well-developed when born
- male parental care
-
parental care is carried out by males
- female parental care
-
parental care is carried out by females
- fossorial
-
Referring to a burrowing life-style or behavior, specialized for digging or burrowing.
- nocturnal
-
active during the night
- crepuscular
-
active at dawn and dusk
- motile
-
having the capacity to move from one place to another.
- solitary
-
lives alone
- pheromones
-
chemicals released into air or water that are detected by and responded to by other animals of the same species
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- soil aeration
-
digs and breaks up soil so air and water can get in
- pet trade
-
the business of buying and selling animals for people to keep in their homes as pets.
- food
-
A substance that provides both nutrients and energy to a living thing.
- 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.
- drug
-
a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease
- carnivore
-
an animal that mainly eats meat
- insectivore
-
An animal that eats mainly insects or spiders.
References
Ashokkumar, M., D. Valsarajan, M. Suresh, A. Kaimal, G. Chandy. 2017. Stomach contents of the Indian Pangolin Manis crassicaudata (Mammalia: Pholidota: Manidae) in tropical forests of southern India. Journal of Threatened Taxa , 9/5: 10246-10248. Accessed January 29, 2018 at http://threatenedtaxa.org/index.php/JoTT/article/view/2873 .
Bhandari, N., M. Chalise. 2014. Habitat and Distribution of Chinese Pangolin (Manis Pentadactyla Linnaeus, 1758) in Nagarjun Forest of Shivapuri Nagarjun National Park, Nepal. Nepalese Journal of Zoology , 2/1: 18-25. Accessed February 11, 2018 at https://www.researchgate.net/profile/Mukesh_Chalise/publication/278406616_Habitat_and_Distribution_of_Chinese_Pangolin_Manis_Pentadactyla_Linnaeus_1758_in_Nagarjun_Forest_of_Shivapuri_Nagarjun_National_Park_Nepal/links/558050f808aed40dd8cd2492/Habitat-and-Distribution-of-Chinese-Pangolin-Manis-Pentadactyla-Linnaeus-1758-in-Nagarjun-Forest-of-Shivapuri-Nagarjun-National-Park-Nepal.pdf .
Cabana, F., A. Plowman, T. Nguyen, S. Chin, S. Wu, H. Lo, H. Watabe, F. Yamamoto. 2017. Feeding Asian pangolins: An assessment of current diets fed in institutions world wide. Zoo Biology , 36/4: 298-305. Accessed January 29, 2018 at http://onlinelibrary.wiley.com.ezproxy2.library.colostate.edu/doi/10.1002/zoo.21375/full .
Challender, D., T. Nyguyen, C. Shepherd, K. Krishnasamy, A. Wang, B. Lee, E. Panjang, L. Fletcher, S. Heng, J. Seah, A. Olsson, A. Nguyen, Q. Nguyen, Y. Chung. 2014. Manis javanica. The IUCN Red List of Threatened Species 2014: e.T12763A45222303 , NA: NA. Accessed March 05, 2018 at http://www.iucnredlist.org/details/12763/0 .
Feiler, 1998. The Philippine pangolin, Manis culionensis ELERA, 1915, an almost overlooked species (Mammalia: Pholidota: Manidae). Zoologische Abhandlungen , 50/4-12: 161-164.
Gaubert, P., A. Antunes, H. Meng, L. Miao, S. Peigné, F. Justy, F. Njiokou, S. Dufour, E. Danquah, J. Alahakoon, E. Verheyen, W. Stanley, S. O'Brien, W. Johnson, S. Luo. 2017. The complete phylogeny of pangolins: Scaling up resources for the molecular tracing of the most trafficked mammals on Earth. The Journal of Heredity , 87/1: 1-13. Accessed January 29, 2018 at https://www.ncbi.nlm.nih.gov/pubmed/29140441 .
Gaubert, P., A. Antunes. 2005. ASSESSING THE TAXONOMIC STATUS OF THE PALAWAN PANGOLIN MANIS CULIONENSIS (PHOLIDOTA) USING DISCRETE MORPHOLOGICAL CHARACTERS. Journal of Mammalogy , 86/6: 1068-1074. Accessed February 19, 2018 at https://watermark.silverchair.com/86-6-1068.pdf?token=AQECAHi208BE49Ooan9kkhW_Ercy7Dm3ZL_9Cf3qfKAc485ysgAAAdowggHWBgkqhkiG9w0BBwagggHHMIIBwwIBADCCAbwGCSqGSIb3DQEHATAeBglghkgBZQMEAS4wEQQM1cW0CFuSNTUQMxo1AgEQgIIBjb6C4eZzPpvwCewtpW5ra1eFAuGN6ALtUtBpPNcRvt1nv9-6M3Wb65xoCgJWRr8Dn2IyhQww29olJ8k9hikElpmwxlzZaMemotcDyqYeQcGQDE8RBJrUK2xXOk37qiZtgIhfSaVlz1hxBBh7yCimJMdi0l5-vD2y05EJ414nK2ZNs1eaGN3QgTtn-q9DYFFKQDItVd9bIrr430YZRDScz74at0b3uJjHjJX082NSPpWJwOn698XU-pEAL6DDoy6E64p1JCkymUay6nApj0QA6fsin7iPyYJVUpDrJrlEOhv4HkuSCYeOLxcwKzuyHIdx6h4Ug6S7CvohbB0GpNEvRy49pOnwHKLr0Pt9seTLq4OtxNYR_DZqstzeYFSPP4WbKsYteA3b3ATcPodr1ur7By6LkZ_PgDzd7Hb69dILo-ktR8-XGxmGm9AGLGvm_jP6QbnC_QdnmLfV5Ye_B52gTXwpzS6K94Tai93iG7vxNiflmjVOdPH2vc5ca2dkNCGF13Jzi_feyED_DBHe448 .
Gaudin, T., R. Emry, J. Wible. 2009. The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis. Journal of Mammalian Evolution , 16/4: 235-305. Accessed March 26, 2018 at https://www.researchgate.net/publication/225550717_The_Phylogeny_of_Living_and_Extinct_Pangolins_Mammalia_Pholidota_and_Associated_Taxa_A_Morphology_Based_Analysis .
Heath, M., S. Vanderlip. 1988. Biology, husbandry, and veterinary care of captive Chinese pangolins (Manis pentadactyla). Zoo Biology , 7/4: 293-312. Accessed March 05, 2018 at http://onlinelibrary.wiley.com/doi/10.1002/zoo.1430070402/full .
ITIS, 2017. "Manis Linnaeus, 1758" (On-line). Integrated Taxonomic Information System. Accessed March 26, 2018 at https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=584905#null .
Kamalakannan, M. 2017. CHARACTERIZATION OF BRISTLE HAIR OF THE INDIAN PANGOLIN MANIS CRASSICAUDATA E. GEOFFROY, 1803 AND CHINESE PANGOLIN MANIS PENTADACTYLA LINNAEUS, 1758 (MANIDAE: PHOLIDATA). Journal of Experimental Zoology India , 20/2: 1033-1036.
Kaspal, P. 2009. Saving the Pangolins: Ethno zoology and Pangolin conservation awareness in Human dominated Landscapes. Himalayan Nature , NA: 1-19. Accessed February 11, 2018 at https://www.rufford.org/files/40.09.08%20Preliminary%20Report.pdf .
Katuwal, H., H. Sharma, K. Parajuli. 2017. Anthropogenic impacts on the critically endangered Chinese pangolin (Manis pentadactyla) in Nepal. Journal of Mammalogy , 98/6: 1667-1673. Accessed January 29, 2018 at https://academic-oup-com.ezproxy2.library.colostate.edu/jmammal/article/98/6/1667/4160377 .
Li, Y., S. Chao, Z. FuHua, W. ShiBao, M. GuangZhi. 2010. Age structure and parasites of Malayan pangolin (Manis javanica).. Journal of Economic Animal , 14/1: 22-25. Accessed March 05, 2018 at https://www.cabdirect.org/cabdirect/abstract/20103112417 .
Mahmood, T., R. Hussain, N. Irshad, F. Akrim, M. Nadeem. 2012. Illegal Mass Killing of Indian Pangolin (Manis crassicaudata) in Potohar Region, Pakistan. Pakistan Journal of Zoology , 44/5: 1457-1461. Accessed March 05, 2018 at http://www.zsp.com.pk/pdf44/1457-1461_41_%20SHORT%20COMMUNICATIONS_.pdf .
Mahmood, T., N. Irshad, R. Hussain, F. Akrim, I. Hussain, M. Anwar, M. Rais, M. Nadeem. 2016. Breeding habits of the Indian pangolin (Manis crassicaudata) in Potohar Plateau, Pakistan. Mammalia , 80/2: 231-234. Accessed January 29, 2018 at https://www-degruyter-com.ezproxy2.library.colostate.edu/view/j/mamm.2016.80.issue-2/mammalia-2014-0153/mammalia-2014-0153.xml .
Mahmood, T., N. Irshad, R. Hussain. 2012. Habitat Preference and Population Estimates of Indian Pangolin (Manis crassicaudata) in District Chakwal of Potohar Plateau, Pakistan. Russian Journal of Ecology , 45/1: 70-75. Accessed February 11, 2018 at https://link.springer.com/article/10.1134%2FS1067413614010081 .
Mahmood, T., K. Jabeen, I. Hussain, A. Kayani. 2013. Plant Species Association, Burrow Characteristics and the Diet of the Indian Pangolin. Pakistan Journal of Zoology , 45/6: 1533-1539. Accessed March 05, 2018 at http://zsp.com.pk/pdf45/1533-1539%20_8_%20PJZ-1146-12%2026-10-13%20Page%20Proof%20final%20_23-10-13_%20MS%20No..pdf .
Mohapatra, R., S. Panda. 2014. Behavioural Descriptions of Indian Pangolins (Manis crassicaudata) in Captivity. International Journal of Zoology , NA: 1-7. Accessed February 19, 2018 at https://www.hindawi.com/journals/ijz/2014/795062/ .
Schlitter, D., D. Wilson, D. Reeder. 2005. Mammal Species of the World: A Taxonomic and Geographic Reference, Volume 2 . Baltimore, Maryland: Johns Hopkins University Press.
Shepherd, C., E. Connelly, L. Hywood, P. Cassey. 2017. Taking a stand against illegal wildlife trade: the Zimbabwean approach to pangolin conservation. Oryx , 51/2: 280-285. Accessed January 29, 2018 at https://www-cambridge-org.ezproxy2.library.colostate.edu/core/services/aop-cambridge-core/content/view/3128970E18D11C84BA7395665FC293D6/S0030605316000119a.pdf/div-class-title-taking-a-stand-against-illegal-wildlife-trade-the-zimbabwean-approach-to-pangolin-conservation-div.pdf .
Shepherd, C., L. Shepherd. 2012. A Naturalist's Guide to the Mammals of Southeast Asia . Wiltshire, UK: John Beaufoy Publishing.
Tenaza, R. 1975. Pangolins Rolling Away from Predation Risks. Journal of Mammalogy , 56/1: 257. Accessed March 05, 2018 at https://academic.oup.com/jmammal/article-abstract/56/1/257/839099 .
Thapa, P. 2013. An Overview of Chinese Pangolin (Manis pentadactyla): Its General Biology, Status, Distribution and Conservation Threats in Nepal. The Initiation , NA: 164-170. Accessed February 06, 2018 at https://www.nepjol.info/index.php/INIT/article/view/10267/8346 .
Wang, B., W. Yang, V. Sherman, M. Meyers. 2016. Pangolin armor: Overlapping, structure, and mechanical properties of the keratinous scales. Acta Biomaterialia , 41: 60-74. Accessed March 05, 2018 at https://www.sciencedirect.com/science/article/pii/S174270611630246X?via%3Dihub .
Wilson, D., D. Reeder. 2017. "Genus Manis" (On-line). Mammal Species of the World: 3rd Edition. Accessed March 26, 2018 at https://www.departments.bucknell.edu/biology/resources/msw3/browse.asp?s=y&id=13900003 .
Zhang, F., S. Wu, L. Yang, L. Zhang, R. Sun, S. Li. 2015. Reproductive parameters of the Sunda pangolin, Manis javanica. Folia Zoologica , 64/2: 129-135. Accessed January 29, 2018 at http://www.bioone.org.ezproxy2.library.colostate.edu/doi/pdf/10.25225/fozo.v64.i2.a6.2015 .
Zhang, F., J. Yu, S. Wu, S. Li, C. Zou, Q. Wang, R. Sun. 2017. Keeping and breeding the rescued Sunda pangolins (Manis javanica) in captivity. Zoo Biology , 36/6: 387-396. Accessed January 29, 2018 at http://onlinelibrary.wiley.com.ezproxy2.library.colostate.edu/doi/10.1002/zoo.21388/full .