Animal Diversity Web

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)

• Biogeographic Regions
• palearctic
  • native
• oriental
  • native

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)

• 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). (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)

• 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). (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). (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)

• 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). (Mah­mood, et al., 2016)

• Parental Investment
• precocial
• male parental care
• female parental care
• pre-hatching/birth
  • provisioning
    • female
  • protecting
    • male
    • female
• pre-weaning/fledging
  • provisioning
    • female
  • protecting
    • male
    • female
• pre-independence
  • protecting
    • female

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.

• 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). (Mah­mood, et al., 2016; Thapa, 2013)

• Other Communication Modes
• pheromones

• Perception Channels
• visual
• tactile
• acoustic
• chemical

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)

• Primary Diet
• carnivore
  • insectivore

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)

• Ecosystem Impact
• creates habitat
• soil aeration

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)

• 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). (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.

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. 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.