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)
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)
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)
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)
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)
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)
Information about the lifespan of Manis species is unknown.
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.
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)
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)
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)
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)
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)
There are no known adverse effects of Manis species on humans.
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)
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)
Mea Escobedo (author), Colorado State University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
uses sound to communicate
living in landscapes dominated by human agriculture.
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.
an animal that mainly eats meat
uses smells or other chemicals to communicate
active at dawn and dusk
a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease
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.
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.
parental care is carried out by females
A substance that provides both nutrients and energy to a living thing.
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
Referring to a burrowing life-style or behavior, specialized for digging or burrowing.
An animal that eats mainly insects or spiders.
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).
parental care is carried out by males
Having one mate at a time.
having the capacity to move from one place to another.
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
the area in which the animal is naturally found, the region in which it is endemic.
active during the night
found in the oriental region of the world. In other words, India and southeast Asia.
the business of buying and selling animals for people to keep in their homes as pets.
chemicals released into air or water that are detected by and responded to by other animals of the same species
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.
breeding is confined to a particular season
reproduction that includes combining the genetic contribution of two individuals, a male and a female
digs and breaks up soil so air and water can get in
uses touch to communicate
Living on the ground.
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
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.
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.
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.
uses sight to communicate
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
breeding takes place throughout the year
young are relatively well-developed when born
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. 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.
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.
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. 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., 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., 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.