Pangolin: Difference between revisions
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{{Short description|Mammals of the order Pholidota}} |
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{{Redirect|Pholidota|the orchid|Pholidota (plant)}} |
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{{other uses}} |
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{{Redirect|Pholidota|the orchid|Pholidota (plant){{!}}''Pholidota'' (plant)}} |
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{{Use dmy dates|date=August 2021}} |
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{{Automatic taxobox |
{{Automatic taxobox |
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|fossil_range = {{Fossil range|47.8|0|earliest=79.47}} Middle [[Eocene]] – present |
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| name = Pangolin |
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|image = Pangolin brought to the Range office, KMTR AJTJ cropped.jpg |
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| fossil_range = {{Fossil range|Paleocene|Present}} |
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|image_caption = [[Indian pangolin]] |
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| image = Pangolin borneo.jpg |
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|display_parents = 2 |
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| image_caption = [[Sunda pangolin]] (''Manis javanica'') |
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|taxon = Pholidota |
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| status = CITES_A1 |
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|authority = [[Max Carl Wilhelm Weber|Weber]], 1904 |
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| status_system = CITES |
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|subdivision_ranks = Subgroups |
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| status_ref = <ref name=CITES/> |
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|subdivision = {{center|[see [[#Classification and phylogeny|classification]]]}} |
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| parent_authority = [[Max Carl Wilhelm Weber|Weber]], 1904 |
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|range_map = Minus ranges.png |
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| taxon = Manidae |
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|range_map_caption = Ranges of living species |
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| authority = [[John Edward Gray|Gray]], 1821 |
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|synonyms = |
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| display_parents = 4 |
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{{collapsible list |bullets=true |title=list of synonyms: |
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| subdivision_ranks = Genera |
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|Afredentata {{small|Szalay & Schrenk, 1994}}<ref>{{cite journal | url=https://www.tandfonline.com/doi/abs/10.1080/02724634.1994.10011592 | doi=10.1080/02724634.1994.10011592 | title=Abstracts of Papers | journal=Journal of Vertebrate Paleontology | date=1994 | volume=14 | pages=1–58 | bibcode=1994JVPal..14S...1. }}</ref> |
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| subdivision = |
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|Lepidota {{small|Lane, 1910}}<ref>Lane, Henry Higgins (1910) [https://archive.org/details/jstor-1636382 "A corrected classification of the edentates."] Science, new ser., vol. 31, pp. 913-914</ref> |
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* ''[[Manis]]'' |
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|Manides {{small|Gervais, 1854}} |
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* ''[[Phataginus]]'' |
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|Maniformes {{small|Zagorodniuk, 2008}}<ref>{{cite journal |last1=Zagorodniuk |first1=I. |year=2008 |title=Scientific names of mammal orders: from descriptive to uniform |url=https://www.academia.edu/27585773 |journal=Visnyk of Lviv University |series=Biology |issue=48 |pages=33–43 }}</ref> |
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* ''[[Smutsia]]'' |
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|Manitheria {{small|Haeckel, 1895}}<ref>{{cite book |first=Ernst |last=Haeckel |title=Systematische Phylogenie: Wirbelthiere |url=https://archive.org/details/systematischephy03haec/page/601 |year=1895 |location=Berlin |publisher=G. Reimer |volume=T.3 |language=de}}</ref> |
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| range_map = Minus ranges.png |
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|Neomanida {{small|Haeckel, 1895}} |
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| range_map_caption = Species ranges |
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|Nomarthra {{small|Cope, 1889}}<ref>{{cite journal |last1=Cope |first1=E. D. |year=1889 |title=The Edentata of North America |journal= American Naturalist |volume=23 |issue=272 |pages= 657–664 |doi=10.1086/274985 |s2cid=83633905 }}</ref> |
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{{leftlegend|#7C1797|''[[Manis crassicaudata]]''|outline=gray}} |
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|Pholidotheria {{small|Haeckel, 1895}} |
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{{leftlegend|#F89B3C|''[[Manis pentadactyla]]''|outline=gray}} |
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|Pholidotiformes {{small|Kinman, 1994}}<ref>Kenneth E. Kinman (1994) "The Kinman System: Toward a Stable Cladisto-Eclectic Classification of Organisms: Living and Extinct, 48 Phyla, 269 Classes, 1,719 Orders", Hays, Kan. (P. O. Box 1377, Hays 67601), 88 pages</ref> |
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{{leftlegend|#59C9FD|''[[Manis javanica]]''|outline=gray}} |
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|Pholidotina {{small|Pearse, 1936}}<ref>{{cite book |last=Pearse |first=Arthur Sperry |year=1936 |title=Zoological names. A list of phyla, classes, and orders, prepared for section F, American Association for the Advancement of Science |url=https://archive.org/details/zoologicalnamesl1936pear/page/24/mode/2up?q=Creodontina |location= |publisher=Duke University Press |page=24}}</ref> |
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{{leftlegend|#FF0800|''[[Manis culionensis]]''|outline=gray}} |
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|Repentia {{small|Newman, 1843}}<ref>Edward Newman (1843) [https://www.biodiversitylibrary.org/item/123076#page/7/mode/1up "The Zoologist: a monthly journal of natural history. (Vol. 1)"], London, J. Van Voorst</ref> |
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{{leftlegend|#9FDA31|''[[Phataginus tricuspis]]''|outline=gray}} |
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|Scutata {{small|Murray, 1866}}<ref>Murray, Andrew (1866) [https://www.biodiversitylibrary.org/item/53245#page/7/mode/1up "The geographic distribution of mammals"], London, Day and Son, Ltd., XVI + 420 pp., 101 maps</ref> |
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{{leftlegend|#FF0090|''[[Phataginus tetradactyla]]''|outline=gray}} |
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|Squamata {{small|Huxley, 1872}}<ref>Huxley, Thomas Henry (1872) [https://www.biodiversitylibrary.org/item/109100#page/7/mode/1up "A manual of the anatomy of vertebrated animals"], New York, d, Appleton and Co., 431 pp.</ref> |
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{{leftlegend|#03C03C|''[[Smutsia gigantea]]''|outline=gray}} |
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|Squamigera {{small|Gill, 1910}}<ref>{{cite journal | url=https://www.science.org/doi/10.1126/science.32.810.56.a | doi=10.1126/science.32.810.56.a | title=Classification of the Edentates | date=1910 | last1=Gill | first1=Theo. | journal=Science | volume=32 | issue=810 | page=56 | pmid=17745887 | s2cid=239573422 }}</ref> |
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{{leftlegend|#1560BD|''[[Smutsia temminckii]]''|outline=gray}} |
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|Squamosa {{small|Haeckel, 1895}} |
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}} |
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}} |
}} |
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[[File:A pangolin in defensive posture, Horniman Museum, London.jpg|thumb|300px|A pangolin in defensive posture, Horniman Museum, London]] |
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'''Pangolins''', sometimes known as '''scaly anteaters''',<ref>{{Cite EB1911 |last1=Thomas |first1=Oldfield |author-link1=Oldfield Thomas |last2=Lydekker |first2=Richard |author-link2=Richard Lydekker |wstitle=Pangolin |short=x}}</ref> are [[mammals]] of the order '''Pholidota''' ({{IPAc-en|f|ɒ|l|ᵻ|ˈ|d|oʊ|t|ə}}). The one [[Neontology|extant]] family, the [[Manidae]], has three genera: ''[[Manis]]'', ''[[Phataginus]]'', and ''[[Smutsia]]''.<ref>{{cite book |last=Schlitter |first= Duane A. |editor-last1=Wilson |editor-first1=D.E. |editor-last2=Reeder |editor-first2= D.M. |year=2005 |title=Mammal Species of the World: A Taxonomic and Geographic Reference |url=https://www.departments.bucknell.edu/biology/resources/msw3/browse.asp?id=13900002 |edition=3rd |location=Baltimore, Maryland, USA |publisher=Johns Hopkins University Press |page=530 |isbn=978-0-8018-8221-0}}</ref> ''Manis'' comprises four species found in Asia, while ''Phataginus'' and ''Smutsia'' include two species each, all found in sub-Saharan Africa.<ref name="web2.utc.edu">{{cite journal |last=Gaudin |first=Timothy |date=28 August 2009 |title=The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis |journal=[[Journal of Mammalian Evolution]] |publisher=[[Springer Science+Business Media]] |location=Heidelberg, Germany |volume=16 |issue=4 |pages=235–305 |doi=10.1007/s10914-009-9119-9 |s2cid=1773698 |url=http://web2.utc.edu/~gvv824/Gaudin%20et%20al%202009.pdf |access-date=14 May 2015 |archive-date=25 September 2015 |archive-url=https://web.archive.org/web/20150925134257/http://web2.utc.edu/~gvv824/Gaudin%20et%20al%202009.pdf |url-status=dead }}</ref> These species range in size from {{convert|30|to|100|cm|abbr=on}}. Several [[extinct]] pangolin species are also known. In September 2023, nine species were reported.<ref name="NYT-20230925">{{cite news |last=Incorvala |first=Darren |title=A Mystery Species Was Discovered in Trafficked Pangolin Scales - Researchers believed there were eight species of the strange mammals. But a ninth was identified genetically, although no one knew it was a separate species in the wild. |url=https://www.nytimes.com/2023/09/25/science/pangolins-new-species-scales.html |date=25 September 2023 |work=[[The New York Times]] |url-status=live |archiveurl=https://archive.today/20230925194503/https://www.nytimes.com/2023/09/25/science/pangolins-new-species-scales.html |archivedate=25 September 2023 |accessdate=26 September 2023 }}</ref> |
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'''Pangolins''' or '''scaly anteaters'''<ref>{{Cite EB1911 |first=Oldfield |last=Thomas |authorlink=Oldfield Thomas |last2=Lydekker |first2=Richard |author2link=Richard Lydekker |wstitle=Pangolin |short=x}}</ref> are mammals of the order '''Pholidota''' (from the Greek word [[wiktionary:φολίς|φολῐ́ς]], "horny scale"). The one [[Neontology|extant]] family, '''Manidae''', has three genera: ''[[Manis]]'', which comprises four species living in Asia; ''[[Phataginus]]'', which comprises two species living in Africa; and ''[[Smutsia]]'', which comprises two species also living in Africa.<ref name="web2.utc.edu">{{cite journal|last1=Gaudin|first1=Timothy|title=The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Tais gay |
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a: A Morphology Based Analysis|journal=Journal of Mammalian Evolution|date=28 August 2009|volume=16|issue=4|pages=235–305|doi=10.1007/s10914-009-9119-9|url=http://web2.utc.edu/~gvv824/Gaudin%20et%20al%202009.pdf|accessdate=14 May 2015}}</ref> These species range in size from {{convert|30|to|100|cm|abbr=on}}. A number of [[extinct]] pangolin species are also known. |
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Pangolins have large, protective [[keratin]] scales covering their skin; they are the only known |
Pangolins have large, protective [[keratin]] scales, similar in material to fingernails and toenails, covering their skin; they are the only known mammals with this feature. Depending on the species, they live in hollow trees or [[burrow]]s. Pangolins are [[nocturnal]], and their diet consists of mainly [[ants]] and [[termites]], which they capture using their long tongues. They tend to be solitary animals, meeting only to mate and produce a litter of one to three offspring, which they raise for about two years. Pangolins superficially resemble [[armadillo]]s, though the two are not closely related; they have merely undergone [[convergent evolution]]. |
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Pangolins are [[pangolin trade|threatened by poaching]] (for their meat and scales) and heavy [[deforestation]] of their natural habitats, and are the most [[ |
Pangolins are [[pangolin trade|threatened by poaching]] (for their meat and scales, which are used in [[traditional medicine]]<ref>{{Cite journal|last1=D'Cruze|first1=Neil|last2=Assou|first2=Délagnon|last3=Coulthard|first3=Emma|last4=Norrey|first4=John|last5=Megson|first5=David|last6=Macdonald|first6=David W.|last7=Harrington|first7=Lauren A.|last8=Ronfot|first8=Delphine|last9=Segniagbeto|first9=Gabriel H.|last10=Auliya|first10=Mark|date=2020-11-05|title=Snake oil and pangolin scales: insights into wild animal use at "Marché des Fétiches" traditional medicine market, Togo|url=https://natureconservation.pensoft.net/article/47879/|journal=[[Nature Conservation]]|language=en|volume=39|pages=45–71|doi=10.3897/natureconservation.39.47879|s2cid=218663851|issn=1314-3301|doi-access=free}}</ref><ref>{{cite journal |title=Chinese Medicine and the Pangolin |journal=Nature |pages=72 |doi=10.1038/141072b0 |date=1 January 1938 |volume=141 |issue=3558 |bibcode=1938Natur.141R..72. |doi-access=free}}</ref>) and heavy [[deforestation]] of their natural habitats, and are the most [[wildlife smuggling|trafficked]] mammals in the world.<ref>{{cite news |last=Goode |first=Emilia |date=31 March 2015 |title=A Struggle to Save the Scaly Pangolin |newspaper=[[The New York Times]] |location=New York City |url=https://www.nytimes.com/2015/03/31/science/a-struggle-to-save-the-scaly-pangolin.html |access-date=1 May 2016}}</ref> {{As of|January 2020}}, there are eight species of pangolin whose [[conservation status]] is listed in the [[Threatened species|threatened]] tier. Three (''[[Manis culionensis]]'', ''[[Manis pentadactyla|M. pentadactyla]]'' and ''[[Manis javanica|M. javanica]]'') are critically endangered, three (''[[Phataginus tricuspis]]'', ''[[Manis crassicaudata]]'' and ''[[Smutsia gigantea]]'') are endangered and two (''[[Phataginus tetradactyla]]'' and ''[[Smutsia temminckii]]'') are vulnerable on the [[Red List of Threatened Species]] of the [[International Union for Conservation of Nature]].<ref name="IUCN Manidae">{{cite web |title=Manidae Family search |website=[[IUCN Red List of Threatened Species]] |publisher=[[IUCN]] |url=https://www.iucnredlist.org/search?taxonomies=101561&searchType=species |access-date=30 January 2020}}</ref> |
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==Etymology== |
== Etymology == |
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The name of order Pholidota comes from [[Ancient Greek]] [[wikt:ϕολιδωτός|ϕολιδωτός]] – "clad in scales"<ref name="RWB">{{cite book |last1=Brown |first1=Roland Wilbur |title=The Composition of Scientific Words |date=1956 |publisher=Smithsonian Institution Press |location=Washington, D.C. |page=604}}</ref> from {{lang|el|φολίς}} ''pholís'' "[[scale (anatomy)|scale]]".<ref>{{cite web|url=https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=foli/s |title=φολίς |last1=Liddell |first1=Henry George |last2=Scott|first2=Robert |work=A Greek-English Lexicon |publisher=Perseus Digital Library |date= 1940 }}</ref> |
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The name ''pangolin'' comes from the [[Malay language|Malay]] word ''pengguling'', meaning "one who rolls up".<ref name=COED>{{cite book |editor= Judy Pearsall | year=2002 |title= Concise Oxford English Dictionary | edition = 10th | publisher= Oxford University Press | page= 1030 |isbn= 978-0-19-860572-0}}</ref> However, the modern name in [[Malay language|Standard Malay]] is ''tenggiling'', whereas in [[Indonesian language|Indonesian]] it is ''trenggiling''. |
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The name "pangolin" comes from the [[Malay language|Malay]] word ''pengguling'' meaning "one who rolls up"<ref name=COED>{{cite book |editor-last=Pearsall |editor-first=Judy |year=2002 |title=Concise Oxford English Dictionary |edition=10th |publisher=[[Oxford University Press]] |location=Oxfordshire, England |isbn=978-0-19-860572-0 |page=1030}}</ref> from ''guling'' or ''giling'' "to roll"; it was used for the [[Sunda pangolin]] (''Manis javanica'').<ref name="Wilkinson">{{Cite encyclopedia|url=https://nla.gov.au/nla.obj-60272783/view?partId=nla.obj-436318167#page/n572/|title=tĕnggiling<!-- "pĕngguling" in page 243 directs to this word --> |dictionary=A Malay-English dictionary (romanised) |last=Wilkinson |first=Richard James |publisher=Salavopoulos & Kinderlis |location=Mytilene, Greece |date=1932 |volume=II |page=567 |via=[[Trove|TROVE]], [[National Library of Australia]]}}</ref> However, the modern name is ''tenggiling''. In [[Javanese language|Javanese]], <!--Please add Javanese script spelling here --> it is ''terenggiling'';<ref name="Wilkinson"/> and in the [[Philippine languages]], it is ''goling'', ''tanggiling'', or ''balintong'' (with the same meaning).<ref name="Vergara">{{cite book |last1=Vergara |first1=Benito S. |last2=Idowu |first2=Panna Melizah H. |last3=Sumangil |first3=Julia H. |last4=Gonzales |first4=Juan Carlos |last5=Dans |first5=Andres |title=Interesting Philippine Animals |publisher=Island Publishing House, Inc. |location=Washington, D.C. |isbn=9718538550 |url=http://scinet.dost.gov.ph/union/UploadFiles/download.php?b=nast_Vergara%20BS%202000%20Interesting%20Philippine%20Animals_423.pdf&f=../Downloads/nast_Vergara%20BS%202000%20Interesting%20Philippine%20Animals_423.pdf&t=application/pdf |access-date=14 December 2019 |archive-date=27 March 2020 |archive-url=https://web.archive.org/web/20200327210150/http://scinet.dost.gov.ph/union/UploadFiles/download.php?b=nast_Vergara%20BS%202000%20Interesting%20Philippine%20Animals_423.pdf&f=..%2FDownloads%2Fnast_Vergara%20BS%202000%20Interesting%20Philippine%20Animals_423.pdf&t=application%2Fpdf |url-status=dead }}</ref> |
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The etymologies of the three generic names ''Manis'' (Linnaeus, 1758), ''Phataginus'' (Rafinesque, 1821), and ''Smutsia'' (Gray, 1865) are sometimes misunderstood. |
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In ancient India, according to [[Claudius Aelianus|Aelian]], it was known as the ''phattáges'' (φαττάγης).<ref>[[Claudius Aelianus|Aelian]], ''On Animals'', 16:6 [[Loeb Classical Library]], translated A. F. Scholfield 1959 vol. 3, pp. 266-268.</ref> |
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[[Carl Linnaeus]] (1758) invented the [[Neo Latin|Neo-Latin]] generic name ''Manis'' apparently as a feminine singular form of the Latin masculine plural [[Manes]], the [[Ancient Rome|Ancient Roman]] name for a type of [[spirit]], after the animal's strange appearance.<ref>{{Cite web|title=ITIS Report Manis Linnaeus, 1758 |url=https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=584905#null |accessdate= 2018-05-28}}</ref> |
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== Description == |
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[[Constantine Rafinesque]] (1821) formed the Neo-Latin generic name ''Phataginus'' from the French term ''phatagin'', adopted by [[Count Buffon]] (1763) after the reported local name ''phatagin'' or ''phatagen'' used in the [[East Indies]]. |
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[[File:Pangolin skeletons.JPG|thumb|Pangolin [[skeleton]]s at the [[Museum of Osteology]] (2009)]] |
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[[File:Pangolin scale histology.jpg|thumb|Schematic drawing of pangolin scale histology]] |
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The physical appearance of a pangolin is marked by large, hardened, overlapping, plate-like scales, which are soft on newborn pangolins, but harden as the animal matures.<ref>{{Cite journal |last1=Li |first1=Hui-Ming |last2=Liu |first2=Ping |last3=Zhang |first3=Xiu-Juan |last4=Li |first4=Lin-Miao |last5=Jiang |first5=Hai-Ying |last6=Yan |first6=Hua |last7=Hou |first7=Fang-Hui |last8=Chen |first8=Jin-Ping|display-authors=4 |date=2020-09-03 |title=Combined proteomics and transcriptomics reveal the genetic basis underlying the differentiation of skin appendages and immunity in pangolin |journal=Scientific Reports |language=en |volume=10 |issue=1 |page=14566 |doi=10.1038/s41598-020-71513-w |pmid=32884035 |issn=2045-2322|pmc=7471334 |bibcode=2020NatSR..1014566L }}</ref> They are made of [[keratin]], the same material from which human [[fingernails]] and [[tetrapod]] [[Claw#Tetrapods|claws]] are made, and are structurally and compositionally very different from the scales of reptiles.<ref name=Spearman2008>{{Cite journal |last=Spearman |first=R.I.C. |title=On the nature of the horny scales of the pangolin |journal=[[Zoological Journal of the Linnean Society]] |publisher=[[Oxford University Press]] |location=Oxfordshire, England |volume=46 |issue=310 |pages=267–273 |year=2008 |doi=10.1111/j.1096-3642.1967.tb00508.x}}</ref> The pangolin's scaled body is comparable in appearance to a [[pine cone]]. It can curl up into a ball when threatened, with its overlapping scales acting as [[Armour (anatomy)|armor]], while it protects its face by tucking it under its tail. The scales are sharp, providing extra defense from predators.<ref name=Wang2016>{{Cite journal |last1=Wang |first1=Bin |last2=Yang |first2=Wen |last3=Sherman |first3=Vincent R. |last4=Meyers |first4=Marc A. |title=Pangolin armor: Overlapping, structure, and mechanical properties of the keratinous scales |journal=Acta Biomaterialia |publisher=[[Elsevier]] |location=Oxfordshire, England |volume=41 |pages=60–74 |year=2016 |doi=10.1016/j.actbio.2016.05.028 |pmid=27221793}}</ref> |
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The British [[naturalist]] [[John Edward Gray]] named ''Smutsia'' for the [[South Africa]]n naturalist [[Johannes Smuts]] (1808–1869),<ref>{{Cite web|title=S2A3 Biographical Database of Southern African Science [Johannes Smuts] |url =http://www.s2a3.org.za/bio/Biograph_final.php?serial=2641|accessdate =2018-05-28}}</ref><ref>{{cite journal |last1= Palmer |first1=T.S. |date=1904|title=Index Generum Mammalium: a List of the Genera and Families of Mammals|url=https://www.biodiversitylibrary.org/item/164195#page/643/mode/1up|journal=North American Fauna|volume=23 |pages=635 |access-date= 2018-05-28}}</ref> the first South African to write a [[treatise]] on mammals in 1832 (in which he described the species ''Manis temminckii''). |
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Pangolins can emit a noxious-smelling chemical from [[gland]]s near the anus, similar to the spray of a [[skunk]].<ref>{{cite web |url=http://pangolins.org/|title= Meet the Pangolin! |year=2015 |publisher=Pangolins.org |archive-url=https://web.archive.org/web/20150222010139/http://pangolins.org/ |archive-date=22 February 2015 |url-status=live}}</ref> They have short legs, with sharp claws which they use for burrowing into [[ant]] and [[termite]] mounds and for climbing.<ref>{{cite web |url=http://animaldiversity.org/accounts/Manis_tricuspis/ |last1=Andrews |first1=James |title=''Manis tricuspis'': tree pangolin |year=2011 |website=Animal Diversity Web |publisher=[[University of Michigan]] |location=Ann Arbor, Michigan |archive-url=https://web.archive.org/web/20141221202454/http://animaldiversity.org/accounts/Manis_tricuspis/ |archive-date=21 December 2014 |url-status=live}}</ref> |
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==Description== |
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{{stack|[[File:Pangolin skeletons.JPG|thumb|Pangolin [[skeleton]]s at the [[Museum of Osteology]] (2009).]]}} |
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The physical appearance of a pangolin is marked by large hardened overlapping plate-like scales, which are soft on newborn pangolins, but harden as the animal matures.<ref name=":2">{{Cite journal|author1=Yu, Jingyu |author2=Jiang, Fulin |author3=Peng, Jianjun |author4=Yin, Xilin |author5=Ma, Xiaohua |date=Oct 2015|title=The First Birth and Survival of Cub in Captivity of Critically Endangered Malayan Pangolin (Mariis javanica).|journal=Agricultural Science & Technology|volume=16|issue=10|doi=|pmid=|via=EBSCO}}</ref> They are made of [[keratin]], the same material from which human [[fingernails]] and [[tetrapod]] [[Claw#Tetrapods|claws]] are made, and are structurally and compositionally very different from the scales of [[reptile]]s.<ref name=Spearman2008>{{Cite journal |last=Spearman |first=R.I.C. |title=On the nature of the horny scales of the pangolin |journal=Zoological Journal of the Linnean Society |volume=46 |issue=310 |pages=267–273 |year=2008 |doi=10.1111/j.1096-3642.1967.tb00508.x}}</ref> The pangolin's scaled body is comparable in appearance to a [[pine cone]]. It can curl up into a ball when threatened, with its overlapping scales acting as [[Armour (anatomy)|armor]], while it protects its face by tucking it under its tail. The scales are sharp, providing extra defense from predators.<ref name=Wang2016>{{Cite journal |last=Wang |first=Bin |title=Pangolin armor: Overlapping, structure, and mechanical properties of the keratinous scales |journal=Acta Biomaterialia |volume=41 |pages=60–74 |year=2016 |doi=10.1016/j.actbio.2016.05.028|pmid=27221793 }}</ref> |
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The tongues of pangolins are extremely long, and like those of the [[giant anteater]] and the [[tube-lipped nectar bat]], the root of the tongue is not attached to the [[hyoid]] bone but is in the [[thorax]] between the [[sternum]] and the [[vertebrate trachea|trachea]].<ref>{{cite journal |last=Chan |first=Lap-Ki |title=Extrinsic Lingual Musculature of Two Pangolins (Pholidota: Manidae) |journal=[[Journal of Mammalogy]] |publisher=[[Oxford University Press]] |location=Oxfordshire, England |volume=76 |issue=2 |pages=472–480 |year=1995 |doi=10.2307/1382356 |jstor=1382356}}</ref> Large pangolins can extend their tongues as much as {{convert|40|cm|in|abbr=on}}, with a diameter of only about {{convert|0.5|cm|in|frac=5|abbr=on}}.<ref name=Mondadori/> |
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Pangolins can emit a noxious-smelling chemical from [[gland]]s near the anus, similar to the spray of a [[skunk]].<ref>{{cite web|url= http://pangolins.org/|title= Meet the Pangolin!|year= 2015|publisher= Pangolins.org|archiveurl= https://web.archive.org/web/20150222010139/http://pangolins.org/|archivedate=2015-02-22|url-status= live}}</ref> They have short legs, with sharp claws which they use for burrowing into [[ant]] and [[termite]] mounds, and climbing.<ref>{{cite web|url= http://animaldiversity.org/accounts/Manis_tricuspis/|title= ''Manis tricuspis'' tree pangolin|year= 2014|publisher= [[University of Michigan]]|archiveurl= https://web.archive.org/web/20141221202454/http://animaldiversity.org/accounts/Manis_tricuspis/|archivedate=2014-12-21|url-status= live}}</ref> |
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The tongues of pangolins are extremely long and – like those of the [[giant anteater]] and the [[tube-lipped nectar bat]] – the root of the tongue is not attached to the [[hyoid]] bone, but is in the [[thorax]] between the [[sternum]] and the [[vertebrate trachea|trachea]].<ref>{{cite journal |last=Chan |first=Lap-Ki | title = Extrinsic Lingual Musculature of Two Pangolins (Pholidota: Manidae) | journal = [[Journal of Mammalogy]]| volume = 76 | issue = 2 | pages = 472–480 | year = 1995 | doi = 10.2307/1382356 | jstor = 1382356}}</ref> Large pangolins can extend their tongues as much as {{Convert|40|cm|in|abbr=on}}, with a diameter of only {{Convert|0.5|cm|abbr=on}}.<ref name=Mondadori/> |
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==Behaviour== |
==Behaviour== |
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[[File:Manis temminckii (29390603130).jpg|thumb|[[Ground pangolin]] in defensive posture]] |
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Most pangolins are [[nocturnal]] animals which use their well-developed sense of smell to find insects. The [[long-tailed pangolin]] is also active by day, while other species of pangolins spend most of the daytime sleeping, curled up into a ball.<ref name=Mondadori>{{cite book | editor = Mondadori, Arnoldo | title = Great Book of the Animal Kingdom | location = New York | publisher = Arch Cape Press | year = 1988 | page = 252|isbn= 978-0517667910}}</ref> |
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[[File:Pangolin defending itself from lions (Gir Forest, Gujarat, India).jpg|thumb|right|[[Indian pangolin]] defending itself against [[Asiatic lion]]s]] |
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Most pangolins are [[nocturnal]] animals<ref>{{cite journal |last1=Wilson |first1=Amelia E. |date=January 1994 |title=Husbandry of pangolins ''Manis'' spp |journal=International Zoo Yearbook |volume=33 |issue=1 |pages=248–251 |doi=10.1111/j.1748-1090.1994.tb03578.x}}</ref> which use their well-developed sense of smell to find insects. The [[long-tailed pangolin]] is also active by day, while other species of pangolins spend most of the daytime sleeping, curled up into a ball ("[[volvation]]").<ref name=Mondadori>{{cite book |editor-first=Arnoldo |editor-last=Mondadori |title=Great Book of the Animal Kingdom |location=New York City |publisher=Arch Cape Press |year=1988 |page=252 |isbn=978-0517667910}}</ref> |
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[[Arboreal]] pangolins live in hollow trees, whereas the ground-dwelling species dig tunnels underground, to a depth of {{Convert|3.5|m|abbr=on}}.<ref name=Mondadori/> |
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Some pangolins walk with their front claws bent under the foot pad, although they use the entire foot pad on their rear limbs. Furthermore, some exhibit a [[Bipedalism|bipedal]] stance for some |
[[Arboreal]] pangolins live in hollow trees, whereas the ground-dwelling species dig tunnels to a depth of {{convert|3.5|m|ft|abbr=on}}.<ref name=Mondadori/> |
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Some pangolins walk with their front claws bent under the foot pad, although they use the entire foot pad on their rear limbs. Furthermore, some exhibit a [[Bipedalism|bipedal]] stance for some behavior, and may walk a few steps bipedally.<ref name="Mohapatra">{{cite journal |first1=Rajesh K. |last1=Mohapatra |first2=Sudarsen |last2=Panda |year=2014 |title=Behavioural descriptions of Indian pangolins (Manis crassicaudata) in captivity. |journal=[[International Journal of Zoology]] |publisher=[[Wiley-Blackwell]] |location=London, England |volume=2014 |pages=1–7 |doi=10.1155/2014/795062 |doi-access=free}}</ref> Pangolins are also good swimmers.<ref name=Mondadori/> |
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===Diet=== |
===Diet=== |
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Pangolins are [[insectivory|insectivorous]]. Most of their diet consists of various species of ants and termites and may be supplemented by other insects, especially larvae. They are somewhat particular and tend to consume only one or two species of insects, even when many species are available. A pangolin can consume {{convert|140|to|200|g|oz|frac=4|abbr=on}} of insects per day.<ref>{{cite web |url=http://bioweb.uwlax.edu/bio203/s2012/grosshue_crai/diet.htm |title=Rollin' With the Pangolin – Diet |first=Craig |last=Grosshuesch |year=2012 |publisher=[[University of Wisconsin–La Crosse]] |location=La Crosse, Wisconsin |archive-url=https://web.archive.org/web/20141223020558/http://bioweb.uwlax.edu/bio203/s2012/grosshue_crai/diet.htm |archive-date=23 December 2014 |url-status=live}}</ref> Pangolins are an important regulator of termite populations in their natural habitats.<ref>{{cite journal |doi=10.7717/peerj.4140 |pmid=29302388 |pmc=5742527 |title=Transcriptomic analysis identifies genes and pathways related to myrmecophagy in the Malayan pangolin (Manis javanica) |journal=[[PeerJ]] |publisher=[[O'Reilly Media]] |location=Corte Madera, California |volume=5 |pages=e4140 |year=2017 |last1=Ma |first1=Jing-E |last2=Li |first2=Lin-Miao |last3=Jiang |first3=Hai-Ying |last4=Zhang |first4=Xiu-Juan |last5=Li |first5=Juan |last6=Li |first6=Guan-Yu |last7=Yuan |first7=Li-Hong |last8=Wu |first8=Jun |last9=Chen |first9=Jin-Ping |doi-access=free }}</ref> |
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[[File:Pangolin defending itself from lions (Gir Forest, Gujarat, India).jpg|thumb|right|[[Indian pangolin]] defending itself against [[Asiatic lion]]s]] |
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Pangolins are [[insectivory|insectivorous]]. Most of their diet consists of various species of ants and termites and may be supplemented by other insects, especially larvae. They are somewhat particular and tend to consume only one or two species of insects, even when many species are available to them. A pangolin can consume {{convert|140|to|200|g|abbr=on}} of insects per day.<ref>{{cite web|url= http://bioweb.uwlax.edu/bio203/s2012/grosshue_crai/diet.htm|title= Rollin' With the Pangolin – Diet|first= Craig|last= Grosshuesch|year= 2012|publisher= [[University of Wisconsin–La Crosse]]|archiveurl= https://web.archive.org/web/20141223020558/http://bioweb.uwlax.edu/bio203/s2012/grosshue_crai/diet.htm|archivedate= 23 December 2014|url-status= live}}</ref> Pangolins are an important regulator of termite populations in their natural habitats.<ref>{{cite journal|doi=10.7717/peerj.4140|pmid=29302388|pmc=5742527|title=Transcriptomic analysis identifies genes and pathways related to myrmecophagy in the Malayan pangolin (Manis javanica)|journal=PeerJ|volume=5|pages=e4140|year=2017|last1=Ma|first1=Jing-E|last2=Li|first2=Lin-Miao|last3=Jiang|first3=Hai-Ying|last4=Zhang|first4=Xiu-Juan|last5=Li|first5=Juan|last6=Li|first6=Guan-Yu|last7=Yuan|first7=Li-Hong|last8=Wu|first8=Jun|last9=Chen|first9=Jin-Ping}}</ref> |
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Pangolins have |
Pangolins have very poor [[Visual perception|vision]]. They also lack teeth. They rely heavily on [[Olfaction|smell]] and [[hearing]], and they have other physical characteristics to help them eat ants and termites. Their skeletal structure is sturdy and they have strong front legs used for tearing into termite mounds.<ref name="Rose-2010">{{Cite book |title=Xenarthra and Pholidota (Armadillos, Anteaters, Sloths and Pangolins) |last1=Rose |first1=K. D. |last2=Gaudin |first2=T. J. |date=2010 |publisher=John Wiley & Sons, Ltd |isbn=978-0470015902 |doi=10.1002/9780470015902.a0001556.pub2|s2cid=82107941 }}</ref> They use their powerful front claws to dig into trees, soil, and vegetation to find prey,<ref>{{Cite journal |author1=Coulson, Ian M |author2=Heath, Martha E |date=December 1997 |title=Foraging behavior and ecology of the Cape pangolin (Manis temminckii) in north-western Zimbabwe |journal=African Journal of Ecology |volume=35 |issue=4 |pages=361–369 |doi=10.1111/j.1365-2028.1997.101-89101.x |bibcode=1997AfJEc..35..361R |via=EBSCO}}</ref> then proceed to use their long tongues to probe inside the insect tunnels and to retrieve their prey. |
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The structure of their tongue and stomach is key to aiding pangolins in obtaining and digesting insects. Their [[saliva]] is sticky,<ref name=" |
The structure of their tongue and stomach is key to aiding pangolins in obtaining and digesting insects. Their [[saliva]] is sticky,<ref name="Rose-2010"/> causing ants and termites to stick to their long tongues when they are hunting through insect tunnels. Without teeth, pangolins cannot also chew;<ref name="Gutteridge2008">{{cite book |first=Lee |last=Gutteridge |title=The South African Bushveld: A Field Guide from the Waterberg |url={{GBurl|id=Dvk4ogj3SxcC|pg=PT36}} |year=2008 |publisher=30° South Publishers |location=Pinetown, South Africa |isbn=978-1-920143-13-8 |page=36}}</ref> but while [[foraging]], they ingest small stones ([[gastroliths]]), which accumulate in their stomachs to help to grind up ants.<ref name="Publishing2015">{{cite book |title=Wildlife of the World |url={{GBurl|id=qqFFCgAAQBAJ|p=215}} |year=2015 |publisher=[[Dorling Kindersley]] |location=London, England |isbn=978-1-4654-4959-7 |page=215}}</ref> This part of their stomach is called the [[gizzard]], and it is also covered in keratinous spines.<ref>{{Cite journal |last1=Davit-Béal |first1=Tiphaine |last2=Tucker |first2=Abigail S. |last3=Sire |first3=Jean-Yves |date=1 April 2009 |title=Loss of teeth and enamel in tetrapods: fossil record, genetic data and morphological adaptations |journal=[[Journal of Anatomy]] |publisher=[[John Wiley & Sons]] |location=New York City |volume=214 |issue=4 |pages=477–501 |doi=10.1111/j.1469-7580.2009.01060.x |pmid=19422426 |pmc=2736120}}</ref> These spines further aid in the grinding up and digestion of the pangolin's prey. |
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Some species, such as the [[tree pangolin]], use their strong, [[prehensile]] tails to hang from tree branches and strip away bark from the trunk, exposing insect nests inside.<ref name="Prothero2016">{{cite book| |
Some species, such as the [[tree pangolin]], use their strong, [[prehensile]] tails to hang from tree branches and strip away bark from the trunk, exposing insect nests inside.<ref name="Prothero2016">{{cite book |first=Donald R. |last=Prothero |title=The Princeton Field Guide to Prehistoric Mammals |url={{GBurl|id=eiftDAAAQBAJ|p=118}} |year=2016 |publisher=[[Princeton University Press]] |location=Princeton, New Jersey |isbn=978-1-4008-8445-2 |page=118}}</ref> |
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===Reproduction=== |
===Reproduction=== |
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[[File:Philippine Pangolin Curled-up by Gregg Yan.jpg|thumb|A [[Philippine pangolin]] pup and its mother, a [[critically endangered]] species [[endemic]] to the [[Palawan]] island group. It is threatened by illegal poaching for the [[pangolin trade]] to China and [[Vietnam]], where it is regarded as a luxury medicinal delicacy.<ref name="Fabro">{{cite news |last1=Fabro |first1=Keith Anthony S. |title=All hope is not lost for vanishing Palawan pangolin |url=https://www.rappler.com/science-nature/environment/232548-all-hope-not-lost-vanishing-palawan-pangolin |access-date=14 December 2019 |work=Rappler |date=10 June 2019}}</ref>]] |
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Pangolins are solitary and meet only to mate. Males are larger than females, weighing up to 40% more. While the mating season is not defined, they typically mate once each year, usually during the summer or autumn. Rather than the males seeking out the females, males mark their location with urine or feces and the females will find them. If there is competition over a female, the males will use their tails as clubs to fight for the opportunity to mate with her.<ref>{{cite web|url= https://bioweb.uwlax.edu/bio203/s2012/grosshue_crai/reproduction.htm|title= Rollin' With the Pangolin - Reproduction|first= Craig|last= Grosshuesch|year= 2012|publisher= University of Wisconsin–La Crosse|archiveurl= https://web.archive.org/web/20141223040747/http://bioweb.uwlax.edu/bio203/s2012/grosshue_crai/reproduction.htm|archivedate= 23 December 2014|url-status= dead}}</ref> |
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Pangolins are solitary and meet only to have sex, with mating typically taking place at night after the male and female pangolin meet near a watering hole. Males are larger than females, weighing up to 40% more. While the mating season is not defined, they typically mate once each year, usually during the summer or autumn. Rather than the males seeking out the females, males [[scent marking|mark their location with urine or feces]] and the females find them. If competition over a female occurs, the males use their tails as clubs to fight for the opportunity to mate with her.<ref>{{cite web |url=https://bioweb.uwlax.edu/bio203/s2012/grosshue_crai/reproduction.htm |title=Rollin' With the Pangolin – Reproduction |first=Craig |last=Grosshuesch |year=2012 |publisher=University of Wisconsin–La Crosse |archive-url=https://web.archive.org/web/20141223040747/http://bioweb.uwlax.edu/bio203/s2012/grosshue_crai/reproduction.htm |archive-date=23 December 2014 |url-status=dead}}</ref> |
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[[Gestation]] periods differ by species, ranging from roughly 70 to 140 days.<ref name=":0">{{Cite journal|last=Gong|first=Shiping|last2=Hua|first2=Liushuai|last3=Wang|first3=Fumin|last4=Li|first4=Weiye|last5=Ge|first5=Yan|last6=Li|first6=Xiaonan|last7=Hou|first7=Fanghui|date=2015-08-06|title=Captive breeding of pangolins: current status, problems and future prospects|url=https://zookeys.pensoft.net/articles.php?id=5419|journal=ZooKeys|language=en|issue=507|pages=99–114|doi=10.3897/zookeys.507.6970|issn=1313-2970|pmc=4490220|pmid=26155072}}</ref> African pangolin females usually give birth to a single offspring at a time, but the Asiatic species may give birth from one to three.<ref name=Mondadori/> Weight at birth is {{convert|80|to|450|g|abbr=on}} and the average length is {{convert|150|mm|abbr=on}}. At the time of birth, the scales are soft and white. After several days, they harden and darken to resemble those of an adult pangolin. During the vulnerable stage, the mother stays with her offspring in the burrow, nursing it, and wraps her body around it if she senses danger. The young cling to the mother's tail as she moves about, although in burrowing species, they remain in the burrow for the first two to four weeks of life. At one month, they first leave the burrow riding on the mother's back. [[Weaning]] takes place around three months of age, at which stage the young begin to eat insects in addition to nursing. At two years of age, the offspring are sexually mature and are abandoned by the mother.<ref name=EoM>{{cite book |last=Dickman |first=Christopher R. |editor-last=MacDonald |editor-first=D. |year=1984 |title=The Encyclopedia of Mammals |publisher=Facts on File |location=New York |pages=780–781 |isbn=978-0-87196-871-5 |url-access=registration |url=https://archive.org/details/encyclopediaofma00mals_0 }}</ref> |
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[[Gestation]] periods differ by species, ranging from roughly 70 to 140 days.<ref name="Hua-2015">{{cite journal |last1=Hua |first1=Liushuai |last2=Gong |first2=Shiping |last3=Wang |first3=Fumin |last4=Li |first4=Weiye |last5=Ge |first5=Yan |last6=Li |first6=Xiaonan |last7=Hou |first7=Fanghui |title=Captive breeding of pangolins: current status, problems and future prospects |journal=ZooKeys |date=8 June 2015 |issue=507 |pages=99–114 |doi=10.3897/zookeys.507.6970 |pmid=26155072 |pmc=4490220|doi-access=free |bibcode=2015ZooK..507...99H }}</ref> African pangolin females usually give birth to a single offspring at a time, but the Asiatic species may give birth to from one to three.<ref name=Mondadori/> Weight at birth is {{convert|80|to|450|g|oz|frac=4|abbr=on}}, and the average length is {{convert|150|mm|in|frac=4|abbr=on}}. At the time of birth, the scales are soft and white. After several days, they harden and darken to resemble those of an adult pangolin. During the vulnerable stage, the mother stays with her offspring in the burrow, nursing it, and wraps her body around it if she senses danger. The young cling to the mother's tail as she moves about, although, in burrowing species, they remain in the burrow for the first 2–4 weeks of life. At one month, they first leave the burrow riding on the mother's back. [[Weaning]] takes place around three months of age, when the young begin to eat insects in addition to nursing. At two years of age, the offspring are sexually mature and are abandoned by the mother.<ref name=EoM>{{cite book |last=Dickman |first=Christopher R. |editor-last=MacDonald |editor-first=D. |year=1984 |title=The Encyclopedia of Mammals |publisher=Facts on File |location=New York |pages=[https://archive.org/details/encyclopediaofma00mals_0/page/780 780–781] |isbn=978-0-87196-871-5 |url-access=registration |url=https://archive.org/details/encyclopediaofma00mals_0/page/780}}</ref> |
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== Classification and phylogeny == |
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{{further|List of pholidotans}} |
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=== Taxonomy === |
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* Order: '''Pholidota''' <small>Weber, 1904</small> |
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** Genus: †''[[Euromanis]]'' <small>Gaudin, Emry & Wible, 2009</small> |
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** Family: [[Extinction|†]][[Eurotamandua|Eurotamanduidae]] <small>Szalay & Schrenk, 1994</small> |
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** Suborder: '''[[Eupholidota]]''' <small>Gaudin, Emry & Wible, 2009</small> |
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*** Superfamily: '''[[Manoidea]]''' <small>Gaudin, Emry & Wible, 2009</small> |
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**** Family: [[Manidae]] <small>Gray, 1821</small> |
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**** Family: †[[Patriomanidae]] <small>Szalay & Schrenk 1998 [''sensu'' Gaudin, Emry & Pogue, 2006]</small> |
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**** ''[[Incertae sedis]]'' |
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***** Genus: †''[[Necromanis]]'' <small>Filhol, 1893</small> |
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*** Superfamily: †'''[[Eomanis|Eomanoidea]]''' <small>Gaudin, Emry & Wible, 2009</small> |
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**** Family: †[[Eomanis|Eomanidae]] <small>Storch, 2003</small> |
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===Phylogeny=== |
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====Among placentals==== |
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{{see also|Pholidotamorpha}} |
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The order Pholidota was long considered to be the [[sister taxon]] to [[Xenarthra]] (neotropical [[anteater]]s, [[sloth]]s, and [[armadillo]]s), but recent genetic evidence indicates their closest living relatives are the [[carnivora]]ns, with which they form a [[clade]], the [[Ferae]].<ref>{{cite journal |name-list-style=vanc |display-authors=2 |last1=Murphy |first1=William J. |last2=Eizirik |first2=Eduardo |last3=O'Brien |first3=Stephen J. |last4=Madsen |first4=Ole |last5=Scally |first5=Mark |last6=Douady |first6=Christophe J. |last7=Teeling |first7=Emma |last8=Ryder |first8=Oliver A. |last9=Stanhope |first9=Michael J. |last10=de Jong |first10=Wilfried W. |last11=Springer |first11=Mark S. |date=14 December 2001 |title=Resolution of the Early Placental Mammal Radiation Using Bayesian Phylogenetics |journal=Science |doi=10.1126/science.1067179 |pmid=11743200 |bibcode=2001Sci...294.2348M |volume=294 |issue=5550 |pages=2348–2351 |s2cid=34367609}}</ref><ref name="AmrineMadsen2003">{{cite journal |author=Amrine-Madsen, H. |author2=Koepfli, K.P. |author3=Wayne, R.K. |author4=Springer, M.S. |year=2003 |title=A new phylogenetic marker, apolipoprotein B, provides compelling evidence for eutherian relationships |journal=Molecular Phylogenetics and Evolution |volume=28 |issue=2 |pages=225–240 |doi=10.1016/S1055-7903(03)00118-0 |pmid=12878460|bibcode=2003MolPE..28..225A }}</ref><ref name="Beck et al 2006">{{cite journal |first1=Robin |last1=Beck |first2=Olaf |last2=Bininda-Emonds |first3=Marcel |last3=Cardillo |first4=Fu-Guo |last4=Liu |first5=Andy |last5=Purvis |title=A higher-level MRP supertree of placental mammals |journal=[[BMC Evolutionary Biology]] |publisher=[[BioMed Central]] |location=London, England |date=2006 |volume=6 |issue=1 |pages=93 |doi=10.1186/1471-2148-6-93 |pmid=17101039 |pmc=1654192 |doi-access=free }}</ref><ref name="Du Toit2014">{{cite journal |last1=Du Toit |first1=Z. |last2=Grobler |first2=J. P. |last3=Kotzé |first3=A. |last4=Jansen |first4=R. |last5=Brettschneider |first5=H. |last6=Dalton |first6=D. L. |title=The complete mitochondrial genome of Temminck's ground pangolin (''Smutsia temminckii''; Smuts, 1832) and phylogenetic position of the Pholidota (Weber, 1904) |journal=Gene |volume=551 |issue=1 |year=2014 |pages=49–54 |doi=10.1016/j.gene.2014.08.040 |pmid=25158133}}</ref> [[Palaeanodonta|Palaeanodonts]] are even closer relatives to pangolins, being classified with pangolins in the clade [[Pholidotamorpha]].<ref name="KondrashovAgadjanian2012">{{cite journal |last1=Kondrashov |first1=Peter |last2=Agadjanian |first2=Alexandre K. |year=2012 |title=A nearly complete skeleton of ''Ernanodon'' (Mammalia, Palaeanodonta) from Mongolia: morphofunctional analysis |journal=[[Journal of Vertebrate Paleontology]] |publisher=[[Taylor & Francis]]|location=Bethesda Maryland |doi=10.1080/02724634.2012.694319 |volume=32 |issue=5 |pages=983–1001 |bibcode=2012JVPal..32..983K |s2cid=86059673}}</ref> The split between carnivorans and pangolins is estimated to have occurred 79.47 [[mega-annum|Ma]] (million years) ago.<ref name="Heighton2023"/> |
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====Among Manidae==== |
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The first dichotomy in the [[phylogeny]] of extant Manidae separates Asian pangolins (''Manis'') from African pangolins (''Smutsia'' and ''Phataginus'').<ref name="Du Toit2014"/> Within the former, ''Manis pentadactyla'' is the [[sister group]] to a clade comprising ''M. crassicaudata'' and ''M. javanica''. Within the latter, a split separates the large terrestrial African pangolins of the genus ''Smutsia'' from the small arboreal African pangolins of the genus ''Phataginus''.<ref name="Gaubert2018">{{Cite journal |last1=Gaubert |first1=Philippe |last2=Antunes |first2=Agostinho |last3=Meng |first3=Hao |last4=Miao |first4=Lin |last5=Peigné |first5=Stéphane |last6=Justy |first6=Fabienne |last7=Njiokou |first7=Flobert |last8=Dufour |first8=Sylvain |last9=Danquah |first9=Emmanuel |last10=Alahakoon |first10=Jayanthi |last11=Verheyen |first11=Erik |date=11 May 2018 |title=The Complete Phylogeny of Pangolins: Scaling Up Resources for the Molecular Tracing of the Most Trafficked Mammals on Earth |journal=Journal of Heredity |language=en |volume=109 |issue=4 |pages=347–359 |doi=10.1093/jhered/esx097 |pmid=29140441 |doi-access=}}</ref> |
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Asian and African pangolins are thought to have diverged about 41.37 Ma ago.<ref name="Heighton2023">Sean P. Heighton, Rémi Allio, Jérôme Murienne, Jordi Salmona, Hao Meng, Céline Scornavacca, Armanda D. S. Bastos, Flobert Njiokou, Darren W. Pietersen, Marie-Ka Tilak, Shu-Jin Luo, Frédéric Delsuc, Philippe Gaubert (2023.) [https://www.biorxiv.org/content/10.1101/2023.02.16.528682v1 "Pangolin genomes offer key insights and resources for the world’s most trafficked wild mammals"]</ref> Moreover, the [[basal (phylogenetics)|basal]] position of ''Manis'' within Pholidota<ref name="Du Toit2014"/><ref name="du Toit2017">{{cite journal |last1=du Toit |first1=Z. |last2=du Plessis |first2= M. |last3=Dalton |first3=D. L. |last4=Jansen |first4=R. |last5=Paul Grobler |first5=J. |last6=Kotzé |first6=A. |title=Mitochondrial genomes of African pangolins and insights into evolutionary patterns and phylogeny of the family Manidae |journal=BMC Genomics |volume=18 |issue=1 |year=2017 |doi=10.1186/s12864-017-4140-5 |pmid=28934931 |doi-access=free |pmc=5609056 |page=746}}</ref> suggests the group originated in Eurasia, consistent with their [[laurasiatheria]]n phylogeny.<ref name="Du Toit2014"/> |
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==Threats== |
==Threats== |
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{{see also|Pangolin trade}} |
{{see also|Pangolin trade}} |
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[[File:Pangolin scale burn in Cameroon. Credit- Kenneth Cameron - USFWS (2) (32575640450).jpg|thumb|Confiscated black market pangolin scales, which are in high demand in [[Chinese traditional medicine|traditional Chinese medicine]],<ref name=cnnchangethelist>{{cite web |publisher=CNN |title=Change the List: The Most Trafficked Mammal You've Never Heard Of |last=Sutter |first=John D. |url=http://www.cnn.com/interactive/2014/04/opinion/sutter-change-the-list-pangolin-trafficking/ |date=April 2014}}</ref> set to be destroyed by authorities in [[Cameroon]] in 2017]] |
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[[File:Coat of Pangolin scales.JPG|thumb|upright|A coat of [[armor]] made of gilded pangolin scales from India, an unusual object, was presented to [[George III of the United Kingdom|George III]] in 1820.]] |
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Pangolins are hunted and eaten in many parts of Africa and are one of the more popular types of [[bush meat]], while local healers use the pangolin as a source of traditional medicine.<ref>{{Cite journal|last=Boakye|first=Maxwell Kwame|last2=Pietersen|first2=Darren William|last3=Kotzé|first3=Antoinette|last4=Dalton|first4=Desiré-Lee|last5=Jansen|first5=Raymond|date=2015-01-20|title=Knowledge and uses of African pangolins as a source of traditional medicine in Ghana|journal=PLOS ONE|volume=10|issue=1|pages=e0117199|doi=10.1371/journal.pone.0117199|issn=1932-6203|pmc=4300090|pmid=25602281|bibcode=2015PLoSO..1017199B}}</ref> They are also in great demand in southern [[China]] and [[Vietnam]] because their meat is considered a delicacy and some believe that pangolin scales have medicinal qualities.<ref>{{cite news |title=Over a million pangolins slaughtered in the last decade |url=http://news.mongabay.com/2014/0729-hance-pangolins-iucn-update.html |newspaper=''[[Mongabay]]'' |date=29 July 2014|accessdate=7 August 2014 |first=Jeremy |last=Hance |archiveurl= https://web.archive.org/web/20141208082730/http://news.mongabay.com/2014/0729-hance-pangolins-iucn-update.html|archivedate= 8 December 2014|url-status= live}}</ref><ref name="javanica">{{cite web |url= http://oldredlist.iucnredlist.org/details/12763/0,|title= Manis javanica|year= 2014|publisher= IUCN Red List|archiveurl= https://web.archive.org/web/20150222182122/http://www.iucnredlist.org/details/12763/0,|archivedate= 22 February 2015|url-status= live}}</ref><ref>{{cite magazine|url=http://news.nationalgeographic.com/2015/12/151220-wildlife-trafficking-crime-blotter/|title=Crime Blotter: Pangolin Scales, Tiger Skins, and More|first=Jani |last=Actman|magazine=National Geographic |date=20 December 2015 |access-date=1 May 2016}}</ref><ref>{{cite magazine|url=http://news.nationalgeographic.com/2015/12/151218-malaysia-endangered-wildlife-tiger-pangolin-crocodile-smuggling/|title=Tiger Eyes, Crocodile Penis: It's What's For Dinner in Malaysia|last=Cruise|first=Adam|date=18 April 2015|access-date=1 May 2016|magazine=National Geographic}}</ref> 100,000 are estimated to be trafficked a year to China and Vietnam,<ref name=":4">{{Cite news|url=https://www.bbc.com/news/world-45784627|title=What's the secret to saving this rare creature?|last=Haenlein|first=Alexandria Reid, Cathy|date=2018-10-10|work=BBC News|access-date=2018-10-10|last2=Keatinge|first2=Tom|language=en-GB}}</ref> amounting to over one million over the past decade.<ref name=":1" /> This makes it the most [[Wildlife trade|trafficked animal]] in the world.<ref name=":1" /><ref>{{Cite news|url=https://www.bbc.com/news/magazine-30833685|title=The world's most-trafficked mammal – and the scaliest|last=Fletcher|first=Martin|date=2015-02-05|work=BBC News|access-date=2018-10-10|language=en-GB}}</ref> This, coupled with [[deforestation]], has led to a large decrease in the numbers of pangolins. Some species, such as ''[[Chinese pangolin|Manis pentadactyla]]'' have become commercially extinct in certain ranges as a result of overhunting.<ref>{{Cite web|url=http://oldredlist.iucnredlist.org/details/12764/0|title=Manis pentadactyla (Chinese Pangolin)|website=www.iucnredlist.org|access-date=2016-09-15}}</ref> In November 2010, pangolins were added to the [[Zoological Society of London]]'s list of evolutionarily distinct and endangered mammals.<ref>{{cite news|last1=Agence France-Presse|title='Asian unicorn' and scaly anteater make endangered list|url=http://www.smh.com.au//breaking-news-world/asian-unicorn-and-scaly-anteater-make-endangered-list-20101119-180sa.html|work=The Sydney Morning Herald|date=November 19, 2010}}</ref> All eight species of pangolin are classified by the [[IUCN]] as threatened with extinction, while two are classified as [[critically endangered]].<ref name="javanica" /><ref>{{cite web |url= http://oldredlist.iucnredlist.org/details/12764/0|title= Manis pentadactyla|year= 2014|publisher= [[IUCN Red List]]|archiveurl= https://web.archive.org/web/20150209200208/http://www.iucnredlist.org/details/12764/0|archivedate= 9 February 2015|url-status= live}}</ref> All pangolin species are currently listed under Appendix I of [[CITES]] which prohibits international trade, except when the product is intended for non-commercial purposes and a permit has been granted. <ref name=CITES>{{cite web |url=https://www.cites.org/eng/app/appendices.php|title= The CITES Appendices |website=Convention on International Trade in Endangered Species of Wild Fauna and Flora |publisher=CITES |accessdate=28 January 2019}}</ref> |
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Pangolins are in high demand in southern China and [[Vietnam]] because their scales are believed to have medicinal properties in [[Traditional Chinese medicine|traditional Chinese]] and [[Traditional Vietnamese medicine|Vietnamese medicine]].<ref>{{Cite journal|last1=Sexton|first1=Rebecca|last2=Nguyen|first2=Trang|last3=Roberts|first3=David L.|date=2021-01-01|title=The Use and Prescription of Pangolin in Traditional Vietnamese Medicine|journal=Tropical Conservation Science|language=en|publication-place=[[University of Kent]], [[WildAid]]|volume=14|pages=1940082920985755|doi=10.1177/1940082920985755|s2cid=233919533|issn=1940-0829|doi-access=free}}</ref> Their meat is also considered a delicacy.<ref>{{cite book |last1=van Uhm |first1=D.P. |title=The Illegal Wildlife Trade: Inside the World of Poachers, Smugglers and Traders (Studies of Organized Crime) |date=2016 |publisher=Springer |location=New York}}</ref><ref>{{cite news |title=Over a million pangolins slaughtered in the last decade |url=http://news.mongabay.com/2014/0729-hance-pangolins-iucn-update.html |newspaper=[[Mongabay]] |date=29 July 2014|access-date=7 August 2014 |first=Jeremy |last=Hance |archive-url= https://web.archive.org/web/20141208082730/http://news.mongabay.com/2014/0729-hance-pangolins-iucn-update.html |archive-date=8 December 2014 |url-status=live}}</ref><ref name="javanica">{{cite iucn |author1=Challender, D. |author2=Willcox, D.H.A. |author3=Panjang, E. |author4=Lim, N. |author5=Nash, H. |author6=Heinrich, S. |author7=Chong, J. |name-list-style=amp |title=''Manis javanica'' |page=e.T12763A123584856 |year=2019 |access-date=30 January 2020}}</ref><ref>{{cite magazine |url=http://news.nationalgeographic.com/2015/12/151220-wildlife-trafficking-crime-blotter/ |archive-url=https://web.archive.org/web/20151222103917/http://news.nationalgeographic.com/2015/12/151220-wildlife-trafficking-crime-blotter/ |url-status=dead |archive-date=22 December 2015 |title=Crime Blotter: Pangolin Scales, Tiger Skins, and More |first=Jani |last=Actman |magazine=National Geographic |date=20 December 2015 |access-date=1 May 2016}}</ref><ref>{{cite magazine |url=http://news.nationalgeographic.com/2015/12/151218-malaysia-endangered-wildlife-tiger-pangolin-crocodile-smuggling/ |archive-url=https://web.archive.org/web/20151220170620/http://news.nationalgeographic.com/2015/12/151218-malaysia-endangered-wildlife-tiger-pangolin-crocodile-smuggling/ |url-status=dead |archive-date=20 December 2015 |title=Tiger Eyes, Crocodile Penis: It's What's For Dinner in Malaysia |last=Cruise |first=Adam |date=18 April 2015 |access-date=1 May 2016 |magazine=National Geographic}}</ref> 100,000 are estimated to be trafficked a year to China and Vietnam,<ref name="Haenlein-2018">{{Cite news |url=https://www.bbc.com/news/world-45784627 |title=What's the secret to saving this rare creature?|last1=Haenlein |first1=Alexandria |last2=Reid |first2=Cathy |last3=Keatinge |first3=Tom |date=10 October 2018 |work=BBC News |access-date=10 October 2018}}</ref> amounting to over one million over the past decade.<ref name="pangolinsg"/><ref>{{Cite journal |last1=Ingram |first1=Daniel J. |last2=Coad |first2=Lauren |last3=Abernethy |first3=Katharine A. |last4=Maisels |first4=Fiona |last5=Stokes |first5=Emma J. |last6=Bobo |first6=Kadiri S. |last7=Breuer |first7=Thomas |last8=Gandiwa |first8=Edson |last9=Ghiurghi |first9=Andrea |last10=Greengrass |first10=Elizabeth |last11=Holmern |first11=Tomas |date=March 2018 |title=Assessing Africa-Wide Pangolin Exploitation by Scaling Local Data: Assessing African pangolin exploitation |journal=Conservation Letters |volume=11 |issue=2 |pages=e12389 |doi=10.1111/conl.12389 |doi-access=free|hdl=1893/25549 |hdl-access=free }}</ref> This makes them the most [[Wildlife trade|trafficked animal]] in the world.<ref name="pangolinsg"/><ref>{{cite news |last=Fletcher |first=Martin |date=5 February 2015 |title=The world's most-trafficked mammal – and the scaliest |work=BBC News |url=https://www.bbc.com/news/magazine-30833685 |access-date=10 October 2018}}</ref> This, coupled with [[deforestation]], has led to a large decrease in the numbers of pangolins. Some species, such as ''[[Chinese pangolin|Manis pentadactyla]]'' have become commercially extinct in certain ranges as a result of overhunting.<ref>{{cite iucn |last1=Challender |first1=D. |last2=Wu |first2=S. |last3=Kaspal |first3=P. |last4=Khatiwada |first4=A. |last5=Ghose |first5=A. |last6=Ching-Min Su |first6=N. |last7=Suwal |first7=Laxmi |year=2019 |title=''Manis pentadactyla'' |page=e.T12764A123585318 |access-date=30 January 2020}}</ref> In November 2010, pangolins were added to the [[Zoological Society of London]]'s list of evolutionarily distinct and endangered mammals.<ref>{{cite news |author=Agence France-Presse |date=19 November 2010 |title='Asian unicorn' and scaly anteater make endangered list |work=The Sydney Morning Herald |url=https://www.smh.com.au//breaking-news-world/asian-unicorn-and-scaly-anteater-make-endangered-list-20101119-180sa.html}}</ref> All eight species of pangolin are assessed as threatened by the [[IUCN]], while three are classified as [[critically endangered]].<ref name="IUCN Manidae"/> All pangolin species are currently listed under Appendix I of [[CITES]] which prohibits international trade, except when the product is intended for non-commercial purposes and a permit has been granted.<ref name=CITES>{{cite web |title=The CITES Appendices |website=Convention on International Trade in Endangered Species of Wild Fauna and Flora |publisher=CITES |url=https://www.cites.org/eng/app/appendices.php |access-date=28 January 2019}}</ref> |
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Though pangolins are protected by an international ban on their trade, populations have suffered from illegal trafficking due to beliefs in [[East Asia]] that their ground-up scales can stimulate [[lactation]] or cure [[cancer]] or [[asthma]].<ref>{{cite news |url= https://www.nytimes.com/2013/03/12/world/asia/no-species-is-safe-from-burgeoning-wildlife-trade.html?_r=0|title= No Species Is Safe From Burgeoning Wildlife Trade|first= Bettina|last= Wassener|date= 12 March 2013|newspaper= [[The New York Times]]|archiveurl= https://web.archive.org/web/20150222182549/http://www.nytimes.com/2013/03/12/world/asia/no-species-is-safe-from-burgeoning-wildlife-trade.html?_r=0|archivedate= 22 February 2015|url-status= live}}</ref> In the past decade, numerous seizures of illegally trafficked pangolin and pangolin meat have taken place in Asia.<ref>{{cite news |url=http://www.cnn.com/interactive/2014/04/opinion/sutter-change-the-list-pangolin-trafficking/ |title=The Most Trafficked Mammal You've Never Heard Of|first= John D.|last= Sutter|publisher=[[CNN]]|date=3 April 2014|archiveurl= https://web.archive.org/web/20150202055005/http://www.cnn.com/interactive/2014/04/opinion/sutter-change-the-list-pangolin-trafficking/|archivedate= 2 February 2015|url-status= live}}</ref><ref>{{cite web |url= http://www.traffic.org/home/2008/3/17/23-tonnes-of-pangolins-seized-in-a-week.html|title= 23 tonnes of pangolins seized in a week|date= 17 March 2008|publisher= Traffic.org|archiveurl= https://web.archive.org/web/20141126034922/http://www.traffic.org/home/2008/3/17/23-tonnes-of-pangolins-seized-in-a-week.html|archivedate= 26 November 2014|url-status= live}}</ref><ref>{{Cite news|url = https://www.theguardian.com/environment/2007/may/26/china.conservation|title = 'Noah's Ark' of 5,000 rare animals found floating off the coast of China|date= 25 May 2007|last = Watts|first = Jonathan|work = [[The Guardian]]|location=London|archiveurl= https://web.archive.org/web/20141003122422/http://www.theguardian.com/environment/2007/may/26/china.conservation|archivedate= 3 October 2014|url-status= live}}</ref><ref>{{Cite news|url = https://www.wsj.com/articles/SB10001424052702303674004577431140183065720#slide/7|title = Asia in Pictures|date = 27 May 2012|work = [[The Wall Street Journal]]|archiveurl = https://web.archive.org/web/20150222184723/http://www.wsj.com/articles/SB10001424052702303674004577431140183065720#slide/7|archivedate = 22 February 2015|url-status = live}}</ref> In one such incident in April 2013, {{convert|10000|kg|ST|abbr=on}} of pangolin meat were seized from a [[China|Chinese]] vessel that ran aground in the [[Philippines]].<ref>{{cite news |title=Chinese vessel on Philippine coral reef caught with illegal pangolin meat |url=https://www.theguardian.com/environment/2013/apr/15/chinese-vessel-philippine-reef-illegal-pangolin-meat |newspaper=[[Associated Press]] |date=15 April 2013 |url-status=live |archiveurl=https://web.archive.org/web/20130416213335/http://www.guardian.co.uk/environment/2013/apr/15/chinese-vessel-philippine-reef-illegal-pangolin-meat |archivedate=2013-04-16 |accessdate=16 April 2013 |location=London |first=Damian |last=Carrington }}</ref><ref>{{cite news |title=Boat Filled With 22,000 Pounds Of Pangolin Hits Endangered Coral Reef |url=http://www.care2.com/causes/boat-filled-with-22000-pounds-of-meat-from-the-protected-pangolin-hits-endangered-coral-reef.html |publisher=Care2 |date=16 April 2013 |url-status=live |archiveurl=https://web.archive.org/web/20130418114310/http://www.care2.com/causes/boat-filled-with-22000-pounds-of-meat-from-the-protected-pangolin-hits-endangered-coral-reef.html |archivedate=2013-04-18 |accessdate=17 April 2013 |location=London |first=Judy |last=Molland }}</ref> In another case in August 2016, an [[Indonesia]]n man was arrested after police raided his home and found over 650 pangolins in freezers on his property.<ref>{{Cite news|url=https://www.bbc.com/news/world-asia-37196635|title=Indonesian man arrested as 650 pangolins found dead in freezers |language=en-GB|access-date=2016-08-27|work=BBC News|date=2016-08-26}}</ref> The same threat is reported in many countries in Africa, especially [[Nigeria]], where the animal is on the verge of extinction due to overexploitation.<ref>The ''[[Media Trust|Daily Trust]]'' (Nigeria), Saturday 18 February 2017</ref> The overexploitation comes from hunting pangolins for game meat and the reduction of their forest habitats due to deforestation caused by timber harvesting.<ref name=":5">{{Cite journal|last=Sodeinde|first=Olufemi A.|date=January 1994|title=Pangolins in south-west Nigeria – current status and prognosis|url=https://www.researchgate.net/publication/231829666|journal=ORYX|volume=28|pages=43–50|via=|doi=10.1017/S0030605300028283}}</ref> The pangolin are hunted as game meat for both medicinal purposes and food consumption. <ref name=":5" /> |
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China had been the main destination country for pangolins until 2018, where it was surpassed by Vietnam. In 2019, Vietnam was reported to have seized the largest volumes of pangolin scales, surpassing Nigeria that year.<ref>{{Cite news|last=Sen|date=February 18, 2020|title=First ivory, now pangolin scales: Vietnam remains wildlife trafficking hotspot|work=[[VnExpress]]|url=https://e.vnexpress.net/news/news/first-ivory-now-pangolin-scales-vietnam-remains-wildlife-trafficking-hotspot-4055838.html}}</ref> |
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Pangolins are also hunted and eaten in Ghana and are one of the more popular types of [[bushmeat]], while local healers use the pangolin as a source of traditional medicine.<ref>{{cite journal |last1=Boakye |first1=Maxwell Kwame |last2=Pietersen |first2=Darren William |last3=Kotzé |first3=Antoinette |last4=Dalton |first4=Desiré-Lee |last5=Jansen |first5=Raymond |date=20 January 2015 |title=Knowledge and uses of African pangolins as a source of traditional medicine in Ghana |journal=PLOS ONE |doi=10.1371/journal.pone.0117199 |pmc=4300090 |pmid=25602281 |bibcode=2015PLoSO..1017199B |volume=10 |issue=1 |pages=e0117199|doi-access=free }}</ref> |
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Though pangolins are protected by an international ban on their trade, populations have suffered from illegal trafficking due to beliefs in East Asia that their ground-up scales can stimulate [[lactation]] or cure cancer or [[asthma]].<ref>{{cite news |last=Wassener |first=Bettina |date=12 March 2013 |title=No Species Is Safe From Burgeoning Wildlife Trade |newspaper=[[The New York Times]] |url=https://www.nytimes.com/2013/03/12/world/asia/no-species-is-safe-from-burgeoning-wildlife-trade.html?_r=0 |url-status=live |archive-url=https://web.archive.org/web/20150222182549/http://www.nytimes.com/2013/03/12/world/asia/no-species-is-safe-from-burgeoning-wildlife-trade.html?_r=0 |archive-date=22 February 2015}}</ref> In the past decade, numerous seizures of illegally trafficked pangolin and pangolin meat have taken place in Asia.<ref>{{cite news |last=Sutter |first=John D. |date=3 April 2014 |title=The Most Trafficked Mammal You've Never Heard Of |publisher=[[CNN]] |url=http://www.cnn.com/interactive/2014/04/opinion/sutter-change-the-list-pangolin-trafficking/ |url-status=live |archive-url=https://web.archive.org/web/20150202055005/http://www.cnn.com/interactive/2014/04/opinion/sutter-change-the-list-pangolin-trafficking/ |archive-date=2 February 2015}}</ref><ref>{{cite web |date=17 March 2008 |title=23 tonnes of pangolins seized in a week |publisher=Traffic.org |url=http://www.traffic.org/home/2008/3/17/23-tonnes-of-pangolins-seized-in-a-week.html |url-status=live |archive-url=https://web.archive.org/web/20141126034922/http://www.traffic.org/home/2008/3/17/23-tonnes-of-pangolins-seized-in-a-week.html |archive-date=26 November 2014}}</ref><ref>{{cite news |last=Watts |first=Jonathan |date=25 May 2007 |title='Noah's Ark' of 5,000 rare animals found floating off the coast of China |work=[[The Guardian]] |location=London |url=https://www.theguardian.com/environment/2007/may/26/china.conservation |url-status=live |archive-url=https://web.archive.org/web/20141003122422/http://www.theguardian.com/environment/2007/may/26/china.conservation |archive-date=3 October 2014}}</ref><ref>{{cite news |date=27 May 2012 |title=Asia in Pictures |work=[[The Wall Street Journal]] |url=https://www.wsj.com/articles/SB10001424052702303674004577431140183065720#slide/7 |url-status=live |archive-url=https://web.archive.org/web/20150222184723/http://www.wsj.com/articles/SB10001424052702303674004577431140183065720#slide/7 |archive-date=22 February 2015}}</ref> In one such incident in April 2013, {{convert|10000|kg|lb|abbr=in}} of pangolin meat were seized from a Chinese vessel that ran aground in the [[Philippines]].<ref>{{cite news |title=Chinese vessel on Philippine coral reef caught with illegal pangolin meat |url=https://www.theguardian.com/environment/2013/apr/15/chinese-vessel-philippine-reef-illegal-pangolin-meat |newspaper=[[Associated Press]] |date=15 April 2013 |url-status=live |archive-url=https://web.archive.org/web/20130416213335/http://www.guardian.co.uk/environment/2013/apr/15/chinese-vessel-philippine-reef-illegal-pangolin-meat |archive-date=16 April 2013 |access-date=16 April 2013 |location=London |first=Damian |last=Carrington}}</ref><ref>{{cite news |last=Molland |first=Judy |date=16 April 2013 |title=Boat Filled With 22,000 Pounds Of Pangolin Hits Endangered Coral Reef |publisher=Care2 |location=London |url=http://www.care2.com/causes/boat-filled-with-22000-pounds-of-meat-from-the-protected-pangolin-hits-endangered-coral-reef.html |access-date=17 April 2013 |url-status=live |archive-url=https://web.archive.org/web/20130418114310/http://www.care2.com/causes/boat-filled-with-22000-pounds-of-meat-from-the-protected-pangolin-hits-endangered-coral-reef.html |archive-date=18 April 2013}}</ref> In another case in August 2016, an Indonesian man was arrested after police raided his home and found over 650 pangolins in freezers on his property.<ref>{{cite news |date=26 August 2016 |title=Indonesian man arrested as 650 pangolins found dead in freezers |work=BBC News |url=https://www.bbc.com/news/world-asia-37196635 |access-date=27 August 2016}}</ref> The same threat is reported in [[Nigeria]], where the animal is on the verge of extinction due to [[overexploitation]].<ref>The ''[[Media Trust|Daily Trust]]'' (Nigeria), Saturday 18 February 2017</ref> The overexploitation comes from hunting pangolins for game meat and the reduction of their forest habitats due to deforestation caused by [[Logging|timber harvesting]].<ref name="Sodeinde-2009">{{cite journal |last1=Sodeinde |first1=Olufemi A. |last2=Adedipe |first2=Segun R. |title=Pangolins in south-west Nigeria – current status and prognosis |journal=Oryx |date=24 April 2009 |volume=28 |issue=1 |pages=43–50 |doi=10.1017/S0030605300028283|doi-access=free }}</ref> The pangolin are hunted as game meat for both medicinal purposes and food consumption.<ref name="Sodeinde-2009"/> |
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== Virology == |
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=== COVID-19 infection === |
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{{See also|Origin of COVID-19}} |
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The [[nucleic acid sequence]] of a specific receptor-binding domain of the [[coronavirus spike protein|spike protein]] belonging to [[coronavirus]]es taken from pangolins was found to be a 99% match with SARS coronavirus 2 ([[SARS-CoV-2]]), the virus which causes [[COVID-19]] and is responsible for the [[COVID-19 pandemic]].<ref name="Cyranoski-2020">{{Cite journal |last=Cyranoski |first=David |date=7 February 2020 |title=Did pangolins spread the China coronavirus to people? |journal=Nature |doi=10.1038/d41586-020-00364-2 |pmid=33547428 |s2cid=212825975}}</ref><ref>{{cite journal |last1=Liu |first1=P. |last2=Chen |first2=W. |last3=Chen |first3=J.-P. |title=Viral Metagenomics Revealed Sendai Virus and Coronavirus Infection of Malayan Pangolins (Manis javanica) |journal=Viruses |year=2019 |volume=11 |issue=11 |page=979 |doi=10.3390/v11110979 |pmid=31652964 |pmc=6893680|doi-access=free }}</ref> Researchers in [[Guangzhou]], China, hypothesized that SARS-CoV-2 had originated in bats, and prior to infecting humans, was circulating among pangolins. The illicit Chinese trade of pangolins for use in [[traditional Chinese medicine]] was suggested as a [[Vector (epidemiology)|vector]] for human transmission.<ref name="Cyranoski-2020"/><ref name="Live">{{cite web |url=https://www.livescience.com/first-case-coronavirus-found.html |title=1st known case of coronavirus traced back to November in China |first1=Jeanna |last1=Bryner |publisher=[[LiveScience]] |date=15 March 2020 |access-date=15 March 2020}}</ref> However, whole-genome comparison found that the pangolin and human [[Template:SARS-CoV-2 related coronavirus|coronaviruses]] share only up to 92% of their RNA.<ref name="Nature-coronavirus-mystery">{{cite journal|last1=Xiao|first1=Kangpeng|last2=Zhai|first2=Junqiong|last3=Feng|first3=Yaoyu|last4=Zhou|first4=Niu|last5=Zhang|first5=Xu|last6=Zou|first6=Jie-Jian|last7=Li|first7=Na|last8=Guo|first8=Yaqiong|last9=Li|first9=Xiaobing|last10=Shen|first10=Xuejuan|last11=Zhang|first11=Zhipeng|date=7 May 2020|title=Isolation of SARS-CoV-2-related coronavirus from Malayan pangolins|journal=Nature|volume=583|issue=7815|pages=286–289|bibcode=2020Natur.583..286X|doi=10.1038/s41586-020-2313-x|pmid=32380510|doi-access=free|last15=Zhang|first22=Fang-Hui|last12=Shu|first12=Fanfan|last13=Huang|first13=Wangyi|first14=Yu|first25=Yongyi|last25=Shen|first24=Lihua|last24=Xiao|first23=Wu|last23=Chen|last22=Hou|first15=Ziding|first21=Qin-Hui|last21=Cai|first20=Wei-Jun|last20=Xie|first19=Mian|last19=Huang|first18=Shi-Ming|last18=Peng|first17=Ya-jiang|last17=Wu|first16=Rui-Ai|last16=Chen|last14=Li}}</ref><ref>{{cite journal |last1=Zhang |first1=Tao |last2=Wu |first2=Qunfu |last3=Zhang |first3=Zhigang |title=Probable Pangolin Origin of SARS-CoV-2 Associated with the COVID-19 Outbreak |journal=Current Biology |date=19 March 2020 |volume=30 |issue=7 |pages=1346–1351.e2 |doi=10.1016/j.cub.2020.03.022 |pmid=32197085 |pmc=7156161|bibcode=2020CBio...30E1346Z }}</ref> Ecologists worried that the early speculation about pangolins being the source may have led to mass slaughters, endangering them further, which was similar to what happened to [[Asian palm civet]]s during the [[SARS]] outbreak.<ref name="Nature-coronavirus-mystery"/><ref>{{cite web |url=https://www.cbsnews.com/news/civet-cat-slaughter-to-fight-sars |title=Civet Cat Slaughter To Fight SARS |publisher=CBSNews |date=11 January 2004 |access-date=28 February 2020}}</ref> It was later proved that the testing which suggested that pangolins were a potential host for the virus was flawed, when [[genetic analysis]] showed that the [[Coronavirus spike protein|spike protein]] and its binding to receptors in pangolins had minimal effect from the virus, and therefore were not likely mechanisms for COVID-19 infections in humans.<ref name="Partin">{{cite journal | last=Partin | first=Clyde | title=Etymologia: ''Sunda pangolin'' | journal=Emerging Infectious Diseases | publisher=Centers for Disease Control and Prevention | volume=27 | issue=7 | year=2021 | issn=1080-6040 | doi=10.3201/eid2707.et2707 |pmc=8237879 | pages=1810| s2cid=235775122 }}</ref> |
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=== ''Pestivirus'' and ''Coltivirus'' === |
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In 2020, two novel RNA viruses distantly related to [[pestivirus]]es and [[coltivirus]]es have been detected in the genomes of dead ''Manis javanica'' and ''Manis pentadactyla''.<ref name="Gao2020">{{Cite journal |last1=Gao |first1=Wen-Hua |last2=Lin |first2=Xian-Dan |last3=Chen |first3=Yan-Mei |last4=Xie |first4=Chun-Gang |last5=Tan |first5=Zhi-Zhou |last6=Zhou |first6=Jia-Jun |last7=Chen |first7=Shuai |last8=Holmes |first8=Edward C |author-link8=Edward C. Holmes |last9=Zhang |first9=Yong-Zhen |author-link9=Zhang Yongzhen |date=1 January 2020 |title=Newly identified viral genomes in pangolins with fatal disease |journal=Virus Evolution |language=en |volume=6 |issue=1 |pages=veaa020 |doi=10.1093/ve/veaa020 |pmid=32296543 |pmc=7151644}}</ref> To refer to both sampling site and hosts, they were named Dongyang pangolin virus (DYPV) and Lishui pangolin virus (LSPV). The DYPV pestivirus was also identified in ''[[Amblyomma javanense]]'' nymph [[tick]]s from a diseased pangolin.<ref name="Gao2020"/> |
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==Folk medicine== |
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Pangolin scales and flesh are used as ingredients for various [[traditional Chinese medicine]] preparations.<ref name="jak">{{cite news |author1=Mariëtte Le Roux |date=25 March 2018 |title=Quackery and superstition: species pay the cost |url=https://www.thejakartapost.com/news/2018/03/25/quackery-and-superstition-species-pay-the-cost.html |access-date=25 August 2020 |work=The Jakarta Post |publisher=[[Agence France-Presse]]}}</ref> While no [[evidence-based medicine|scientific evidence]] exists for the efficacy of those practices, and they have no logical [[mechanism of action]],<ref>{{Cite journal |date=1 January 1938 |title=Chinese Medicine and the Pangolin |journal=Nature |language=en |volume=141 |issue=3558 |pages=72 |doi=10.1038/141072b0 |bibcode=1938Natur.141R..72. |issn=1476-4687 |doi-access=free}}</ref><ref name="Novella2012">{{cite web |author=Steven Novella |date=25 January 2012 |title=What Is Traditional Chinese Medicine? |url=http://www.sciencebasedmedicine.org/what-is-traditional-chinese-medicine/ |url-status=live |archive-url=https://web.archive.org/web/20140415070141/http://www.sciencebasedmedicine.org/what-is-traditional-chinese-medicine/ |archive-date=15 April 2014 |access-date=14 April 2014 |work=[[Science-Based Medicine]]}}</ref><ref name="Jin2005">{{cite book |author=Zhouying Jin |url={{GBurl|id=V7SsFqkHaC4C|pg=PT36}} |title=Global Technological Change: From Hard Technology to Soft Technology |publisher=[[Intellect Books]] |year=2005 |isbn=978-1-84150-124-6 |page=36 |language=en |quote=The vacuum created by China's failure to adequately support a disciplined scientific approach to traditional Chinese medicine has been filled by pseudoscience |access-date=18 February 2016 |archive-url=https://web.archive.org/web/20170320185919/https://books.google.com/books?id=V7SsFqkHaC4C&pg=PT36 |archive-date=20 March 2017 |url-status=live}}</ref> their popularity still drives the [[black market]] for animal body parts, despite concerns about toxicity, transmission of diseases from animals to humans, and species extermination.<ref name=jak/><ref name="Zhang2012">{{cite journal |last1=Zhang |first1=Fang |last2=Kong |first2=Lin-lin |last3=Zhang |first3=Yi-ye |last4=Li |first4=Shu-Chuen |title=Evaluation of Impact on Health-Related Quality of Life and Cost Effectiveness of Traditional Chinese Medicine: A Systematic Review of Randomized Clinical Trials |journal=The Journal of Alternative and Complementary Medicine |volume=18 |issue=12 |year=2012 |pages=1108–20 |issn=1075-5535 |doi=10.1089/acm.2011.0315 |pmid=22924383}}</ref> The ongoing demand for parts as ingredients continues to fuel [[Pangolin trade|pangolin poaching, hunting and trading]].<ref>{{Cite news |last=Boyle |first=Louise |date=30 June 2020 |title='If we don't buy, they don't die': Tackling the global demand that's driving the illegal wildlife trade |work=The Independent |url=https://www.independent.co.uk/environment/china-animal-trade-markets-wild-ivory-pangolins-a9594591.html |archive-url=https://ghostarchive.org/archive/20220524/https://www.independent.co.uk/environment/china-animal-trade-markets-wild-ivory-pangolins-a9594591.html |archive-date=24 May 2022 |url-access=subscription |url-status=live |access-date=21 July 2020}}</ref> |
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The first record of pangolin scales occurs in ''Ben Cao Jinji Zhu'' ("Variorum of Shennong's Classic of Materia Medica", 500 CE), which recommends pangolin scales for protection against ant bites; burning the scales as a cure for people crying hysterically during the night.<ref name="PanMed">{{cite book |last1=Xing |first1=S. |title=Pangolins: Science, Society and Conservation |last2=Bonebrake |first2=T. C. |last3=Cheng |first3=W. |last4=Zhang |first4=M. |last5=Ades |first5=G. |last6=Shaw |first6=D. |last7=Zhou |first7=Y. |date=2019 |publisher=Academic Press |isbn=9780128155073 |editor1-last=Challender |editor1-first=D. |edition=First |page=233 |chapter=Meat and medicine: historic and contemporary use in Asia |access-date=27 March 2020 |editor2-last=Nash |editor2-first=H. |editor3-last=Waterman |editor3-first=C. |chapter-url={{GBurl|id=zey_DwAAQBAJ|p=233}}}}</ref> During the [[Tang dynasty]], a recipe for expelling evil spirits with a formulation of scales, herbs, and minerals appeared in 682, and in 752 CE the idea that pangolin scales could also stimulate milk secretion in lactating women, one of the main uses today, was recommended in the ''Wai Tai Mi Yao'' ("Arcane Essentials from the Imperial Library").<ref name="PanMed" /> In the [[Song dynasty]], the notion of penetrating and clearing blockages was emphasized in the ''Taiping sheng hui fan'' ("Formulas from Benevolent Sages Compiled During the Era of Peace and Tranquility"), compiled by Wang Huaiyin in 992.<ref name="PanMed" /> |
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In the 21st century, the main uses of pangolin scales are [[quackery]] practices based on unproven claims the scales dissolve [[blood clot]]s, promote blood circulation, or help [[lactation|lactating women]] secrete milk.<ref name="jak" /><ref name="PanMed" /> The supposed health effects of pangolin meat and scales claimed by [[traditional Chinese medicine|folk medicine]] practitioners are based on their consumption of ants, long tongues, and protective scales.<ref name="jak" /> |
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The official [[pharmacopoeia]] of the People's Republic of China included Chinese pangolin scales as an ingredient in TCM formulations.<ref name=PanMed/> Pangolins were removed from the pharmacopoeia starting from the first half of 2020.<ref name="ng_maron_2020-09-09">{{cite web |last1=Maron |first1=Dina Fine |url=https://www.nationalgeographic.com/animals/2020/06/pangolins-receive-new-protections-traditional-medicine-in-china/ |archive-url=https://web.archive.org/web/20200610111656/https://www.nationalgeographic.com/animals/2020/06/pangolins-receive-new-protections-traditional-medicine-in-china/ |url-status=dead |archive-date=10 June 2020 |title=Pangolins receive surprising lifeline with new protections in China |date=9 June 2020 |website=National Geographic |access-date=12 June 2020}}</ref> Although pangolin scales have been removed from the list of raw ingredients, the scales are still listed as a key ingredient in various medicines.<ref>{{Cite news |date=2020 |title=Did China really ban the pangolin trade? Not quite, investigators say |website=Mongabay Environmental News |url=https://news.mongabay.com/2020/06/did-china-really-ban-the-pangolin-trade-not-quite-investigators-say/ |access-date=6 July 2020}}</ref> |
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Pangolin parts are also used for medicinal purposes in other Asian countries such as India, Nepal and Pakistan. In some parts of India and Nepal, locals believe that wearing the scales of a pangolin can help prevent pneumonia.<ref>{{Cite book |chapter-url={{GBurl|id=zey_DwAAQBAJ|p=228}} |title=Pangolins: Science, Society and Conservation |publisher=[[Academic Press]] |isbn=978-0-12-815506-6 |publication-date=2019-11-23 |pages=228 |language=en |chapter=Meat and medicine: historic and contemporary use in Asia}}</ref> Pangolin scales have also been used for medicinal purposes in Malaysia, Indonesia and northern Myanmar. Indigenous people in southern [[Palawan]], Philippines, have held the belief that elders could avoid prostate illnesses by wearing belts made with the scales.<ref>{{Cite book |chapter-url={{GBurl|id=zey_DwAAQBAJ|p=231}} |title=Pangolins: Science, Society and Conservation |publisher=[[Academic Press]] |isbn=978-0-12-815506-6 |edition=First |publication-date=2019-11-23 |pages=231 |language=en |chapter=Meat and medicine: historic and contemporary use in Asia}}</ref> |
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==Conservation== |
==Conservation== |
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[[File:Prince of Wales Pangolin Armour.jpg|thumb|upright|left|A coat of [[scale armour|armour made of gilded pangolin scales]] from India, presented in 1875–76 to the then [[Prince of Wales]], the later [[Edward VII]].]] |
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As a result of increasing threats to pangolins, mainly in the form of illegal, international trade in pangolin skin, scales, and meat, these species have received increasing conservation attention in recent years. For example, in 2014, the IUCN re-categorized all eight species of pangolin on its [[IUCN Red List|Red List of Threatened Species]], and each species is now listed as being threatened with extinction.<ref>{{Cite web|title = The IUCN Red List of Threatened Species|url = http://iucnredlist.org/|website = iucnredlist.org|accessdate = 2015-10-17}}</ref> Also, the IUCN SSC Pangolin Specialist Group launched a global action plan to conserve pangolins, dubbed "Scaling up Pangolin Conservation" in July 2014. This action plan aims to improve all aspects of pangolin conservation with an added emphasis on combating poaching and trafficking of the animal, while educating communities in its importance.<ref name=":1">{{Cite web|url=http://www.pangolinsg.org/members-work/action-plan/|title=Action Plan|website=www.pangolinsg.org|access-date=2016-09-15}}</ref> Another suggested approach to fighting pangolin (and general wildlife) trafficking consists in "following the money" rather than "the animal", which aims to disrupt smugglers' profits by interrupting money flows. Financial intelligence gathering could thus become a key tool in protecting these animals, although this opportunity is often overlooked.<ref name=":4" /> In 2018, a Chinese NGO launched the Counting Pangolins movement, calling for joint efforts to save the mammals from [[trafficking]].<ref>Spotlight: Pangolin conservationists call for ban on illegal trade of mammal products http://www.xinhuanet.com/english/2018-11/21/c_137620372.htm</ref><ref>World Pangolin Day: Conservationists demand greater protection to stop extinction http://english.cri.cn/12394/2017/02/18/3124s952140.htm</ref><ref>Xinhua News: How China is combating wildlife trafficking in Africa http://www.xinhuanet.com/english/2018-06/08/c_137240161.htm</ref> Wildlife conservation group [[Traffic (conservation programme)|TRAFFIC]] has indentified 159 smuggling routes used by pangolin traffickers and aims to shut these down. <ref>{{Cite web|url=https://www.traffic.org/what-we-do/species/pangolins/|title=Pangolins – Species we work with at TRAFFIC|website=www.traffic.org|language=en|access-date=2019-01-10}}</ref> |
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As a result of increasing threats to pangolins, mainly in the form of illegal, international trade in pangolin skin, scales, and meat, these species have received increasing conservation attention in recent years.<ref>{{Cite journal|last=Shibaike|first=Takumi|date=2021|title=Small NGOs and Agenda-Setting in Global Conservation Governance: The Case of Pangolin Conservation|journal=Global Environmental Politics|volume=22 |issue=2 |pages=45–69|doi=10.1162/glep_a_00623|s2cid=240588135|issn=1526-3800|doi-access=free}}</ref> {{As of|January 2020}}, the IUCN considered all eight species of pangolin on its [[IUCN Red List|Red List of Threatened Species]] as threatened.<ref name="IUCN Manidae"/> The IUCN SSC Pangolin Specialist Group launched a global action plan to conserve pangolins, dubbed "Scaling up Pangolin Conservation", in July 2014. This action plan aims to improve all aspects of pangolin conservation with an added emphasis on combating poaching and [[trafficking]] of the animal while educating communities on its importance.<ref name="pangolinsg">{{cite web |title=Action Plan |website=www.pangolinsg.org |url=http://www.pangolinsg.org/members-work/action-plan/ |access-date=15 September 2016 |archive-date=10 July 2017 |archive-url=https://web.archive.org/web/20170710234327/http://www.pangolinsg.org/members-work/action-plan/ |url-status=dead }}</ref> Another suggested approach to fighting pangolin (and general wildlife) trafficking consists in "following the money" rather than "the animal", which aims to disrupt smugglers' profits by interrupting money flows. Financial intelligence gathering could thus become a key tool in protecting these animals, although this opportunity is often overlooked.<ref name="Haenlein-2018"/> In 2018, a Chinese NGO launched the Counting Pangolins movement, calling for joint efforts to save the mammals from trafficking.<ref>Xinhua News (21 November 2018) [https://web.archive.org/web/20181123065637/http://www.xinhuanet.com/english/2018-11/21/c_137620372.htm Spotlight: Pangolin conservationists call for ban on illegal trade of mammal products]</ref><ref>China Plus (18 February 2017) [https://web.archive.org/web/20170222062715/http://english.cri.cn/12394/2017/02/18/3124s952140.htm World Pangolin Day: Conservationists demand greater protection to stop extinction]</ref><ref>Xinhua News (8 June 2018) [http://www.xinhuanet.com/english/2018-06/08/c_137240161.htm How China is combating wildlife trafficking in Africa] {{Webarchive|url=https://web.archive.org/web/20201130050244/http://www.xinhuanet.com/english/2018-06/08/c_137240161.htm |date=30 November 2020 }}</ref> Wildlife conservation group [[Traffic (conservation programme)|TRAFFIC]] has identified 159 smuggling routes used by pangolin traffickers and aims to shut these down.<ref>{{cite web |title=Pangolins – Species we work with at TRAFFIC |website=www.traffic.org |url=https://www.traffic.org/what-we-do/species/pangolins/ |access-date=10 January 2019}}</ref> |
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Many attempts have been made to reproduce pangolins in captivity, but due to their reliance on wide-ranging habitats and very particular diets, these attempts are often unsuccessful.<ref name=":0" /> Pangolins have significantly decreased immune responses due to a genetic dysfunction, making them extremely fragile.<ref>{{cite journal|pmc=5052048|year=2016|last1=Choo|first1=S. W.|title=Pangolin genomes and the evolution of mammalian scales and immunity|journal=Genome Research|volume=26|issue=10|pages=1312–1322|last2=Rayko|first2=M.|last3=Tan|first3=T. K.|last4=Hari|first4=R.|last5=Komissarov|first5=A.|last6=Wee|first6=W. Y.|last7=Yurchenko|first7=A. A.|last8=Kliver|first8=S.|last9=Tamazian|first9=G.|last10=Antunes|first10=A.|last11=Wilson|first11=R. K.|last12=Warren|first12=W. C.|last13=Koepfli|first13=K. P.|last14=Minx|first14=P.|last15=Krasheninnikova|first15=K.|last16=Kotze|first16=A.|last17=Dalton|first17=D. L.|last18=Vermaak|first18=E.|last19=Paterson|first19=I. C.|last20=Dobrynin|first20=P.|last21=Sitam|first21=F. T.|last22=Rovie-Ryan|first22=J. J.|last23=Johnson|first23=W. E.|last24=Yusoff|first24=A. M.|last25=Luo|first25=S. J.|last26=Karuppannan|first26=K. V.|last27=Fang|first27=G.|last28=Zheng|first28=D.|last29=Gerstein|first29=M. B.|last30=Lipovich|first30=L.|displayauthors=29|pmid=27510566|doi=10.1101/gr.203521.115}}</ref> They are susceptible to diseases such as [[pneumonia]] and the development of [[Stress ulcer|ulcers]] in captivity, complications which can lead to an early death.<ref name=":0" /> In addition, pangolins rescued from illegal trade often have a higher chance of being infected with parasites such as [[intestinal worms]], further lessening their chance for rehabilitation and reintroduction to the wild.<ref name=":0" /> Recently, researchers have been able to improve artificial pangolin habitats to allow for reproduction of pangolins, providing some hope for future reintroduction of these species into their natural habitats.<ref name=":2" /> Additionally, in countries such as Taiwan the introduction of Wildlife Rehabilitation Centers has greatly improved the survival of Pangolins. These centers have also helped to reveal the causes of death and injury among Taiwan's Pangolin population.<ref>{{Cite journal|last=Pei|first=Curtis Jai-Chyi|last2=Chen|first2=Chen-Chih|last3=Chi|first3=Meng-Jou|last4=Lin|first4=Wen-Chi|last5=Lin|first5=Jing-Shiun|last6=Arora|first6=Bharti|last7=Sun|first7=Nick Ching-Min|date=2019-02-06|title=Mortality and morbidity in wild Taiwanese pangolin (Manis pentadactyla pentadactyla)|journal=PLOS ONE|language=en|volume=14|issue=2|pages=e0198230|doi=10.1371/journal.pone.0198230|issn=1932-6203|pmc=6364958|pmid=30726204|bibcode=2019PLoSO..1498230S}}</ref>The third Saturday in February is promoted as World Pangolin Day by the conservation NPO Annamiticus.<ref>{{cite web|url=https://www.pangolins.org/about-us/|title=World Pangolin Day – About|publisher=Pangolins.org/Annamiticus|date=2011-10-26}}</ref> |
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[[File:Myanmar Illicit Endangered Wildlife Market 04 (cropped).jpg|thumb|Pangolins (in rectangular cages) in an illegal wildlife market in [[Myanmar]]]] |
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In 2017, [[Jackie Chan]] made a public service announcement called ''[[WildAid]]: Jackie Chan & Pangolins (Kung Fu Pangolin)''<ref name=pangolins>{{cite news |
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|url=http://www.chinadaily.com.cn/world/2017-08/23/content_31006476.htm |
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|title=Jackie Chan fights for pangolins |
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|accessdate=2019-03-17 |
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|date=2017-08-23 |
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|newspaper=[[China Daily]] |
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}}</ref> |
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Many attempts have been made to breed pangolins in captivity, but due to their reliance on wide-ranging habitats and very particular diets, these attempts are often unsuccessful.<ref name="Hua-2015"/><ref>{{cite journal |last1=Aitken-Palmer |first1=Copper |last2=deMaar |first2=Thomas W. |last3=Johnson |first3=James G. |last4=Langan |first4=Jennifer |last5=Bergmann |first5=Jonathan |last6=Chinnadurai |first6=Sathya |last7=Guerra |first7=Hector |last8=Carboni |first8=Deborah A. |last9=Adkesson |first9=Michael J. |date=September 2019 |title=Complications Associated with Pregnancy and Parturition in African White-bellied Pangolins (Phataginus Tricuspis) |journal=Journal of Zoo and Wildlife Medicine |doi=10.1638/2019-0019 |volume=50 |issue=3 |pages=678–687 |pmid=33517639 |s2cid=202727948}}</ref> Pangolins have significantly decreased immune responses due to a genetic dysfunction, making them extremely fragile.<ref>{{cite journal |last1=Choo |first1=S. W. |last2=Rayko |first2=M. |last3=Tan |first3=T. K. |last4=Hari |first4=R. |last5=Komissarov |first5=A. |last6=Wee |first6=W. Y. |last7=Yurchenko |first7=A. A. |last8=Kliver |first8=S. |last9=Tamazian |first9=G. |last10=Antunes |first10=A. |last11=Wilson |first11=R. K. |last12=Warren |first12=W. C. |last13=Koepfli |first13=K. P. |last14=Minx |first14=P. |last15=Krasheninnikova |first15=K. |last16=Kotze |first16=A. |last17=Dalton |first17=D. L. |last18=Vermaak |first18=E. |last19=Paterson |first19=I. C. |last20=Dobrynin |first20=P. |last21=Sitam |first21=F. T. |last22=Rovie-Ryan |first22=J. J. |last23=Johnson |first23=W. E. |last24=Yusoff |first24=A. M. |last25=Luo |first25=S. J. |last26=Karuppannan |first26=K. V. |last27=Fang |first27=G. |last28=Zheng |first28=D. |last29=Gerstein |first29=M. B. |last30=Lipovich |first30=L. |display-authors=1 |year=2016 |title=Pangolin genomes and the evolution of mammalian scales and immunity |journal=Genome Research |doi=10.1101/gr.203521.115 |pmid=27510566 |pmc=5052048 |volume=26 |issue=10 |pages=1312–1322}}</ref> They are susceptible to diseases such as [[pneumonia]] and the development of [[Stress ulcer|ulcers]] in captivity, complications that can lead to an early death.<ref name="Hua-2015"/> In addition, pangolins rescued from illegal trade often have a higher chance of being infected with parasites such as [[intestinal worms]], further lessening their chance for rehabilitation and reintroduction to the wild.<ref name="Hua-2015"/> |
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==Taxonomy== |
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[[File:Zoo Leipzig - Chinesisches Ohrenschuppentier.jpeg|thumb|[[Chinese pangolin]]]] |
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The idea of farming pangolins to reduce the number being illegally trafficked is being explored with little success.<ref>{{cite journal |last1=Challender |first1=Daniel W. S. |last2=Sas-Rolfes |first2=Michael't |last3=Ades |first3=Gary W. J. |last4=Chin |first4=Jason S. C. |last5=Ching-Min Sun |first5=Nick |last6=Chong |first6=Ju lian |last7=Connelly |first7=Ellen |last8=Hywood |first8=Lisa |last9=Luz |first9=Sonja |last10=Mohapatra |first10=Rajesh K. |last11=de Ornellas |first11=Paul |date=1 October 2019 |title=Evaluating the feasibility of pangolin farming and its potential conservation impact |journal=Global Ecology and Conservation |doi=10.1016/j.gecco.2019.e00714 |volume=20 |pages=e00714 |doi-access=free|bibcode=2019GEcoC..2000714C |hdl=2263/75894 |hdl-access=free }}</ref> The third Saturday in February is promoted as World Pangolin Day by the conservation NPO Annamiticus.<ref>{{cite web |date=26 October 2011 |title=World Pangolin Day – About |publisher=Pangolins.org/Annamiticus |url=https://www.pangolins.org/about-us/}}</ref> World Pangolin Day has been noted for its effectiveness in generating awareness about pangolins.<ref>{{cite journal |last1=Chua |first1=Marcus A.H. |last2=Tan |first2=Audrey |last3=Carrasco |first3=Luis Roman |title=Species awareness days: Do people care or are we preaching to the choir? |journal=Biological Conservation |date=2021 |volume=255 |pages=109002 |doi=10.1016/j.biocon.2021.109002|bibcode=2021BCons.25509002C |s2cid=233836573 }}</ref> |
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Pangolins were formerly{{When|date=August 2017}} classified with various other orders of ant-eating mammals, most notably [[Xenarthra]], which includes the true [[anteater]]s, [[sloth]]s, and the [[armadillo]]s which pangolins superficially resemble. Newer genetic evidence, however, indicates their closest living relatives are the [[Carnivora]] with which they form the clade [[Ferae]].<ref>{{cite journal |author=Murphy | title = Resolution of the Early Placental Mammal Radiation Using Bayesian Phylogenetics | journal = Science | volume = 294 | issue = 5550 | pages = 2348–2351 | date = 2001-12-14 | doi = 10.1126/science.1067179 | pmid = 11743200 |name-list-format=vanc| author2 = Willian J. | display-authors = 2 | last3 = O'Brien | first3 = SJ | last4 = Madsen | first4 = O | last5 = Scally | first5 = M | last6 = Douady | first6 = CJ | last7 = Teeling | first7 = E | last8 = Ryder | first8 = OA | last9 = Stanhope | first9 = MJ| bibcode = 2001Sci...294.2348M }}</ref><ref name="Beck et al 2006">{{cite journal|last=Beck|first=Robin MD|author2=Bininda-Emonds, Olaf RP |author3=Cardillo, Marcel |author4=Liu, Fu-Guo |author5= Purvis, Andy |title=A higher-level MRP supertree of placental mammals|journal=BMC Evolutionary Biology|year=2006|volume=6|issue=1|pages=93|doi=10.1186/1471-2148-6-93|pmid=17101039|pmc=1654192}}</ref> Some [[palaeontologists]], placing [[Ernanodon]]ta in a separate suborder of Cimolesta near [[Pholidota]],<ref>For example, {{Harvnb|McKenna|Bell|1997| p=222}} in which they included [[Palaeanodonta|palaeanodonts]]. ({{Harvnb|Rose|2006| p=210}})</ref> have classified the pangolins in the order [[Cimolesta]], together with several extinct groups indicated (†) below, though this idea has fallen out of favor since it was determined that cimolestids were not placental mammals.<ref name="rook">{{Cite journal|last1 = Rook|first1 = D.L.|last2 = Hunter|first2 = J.P.|title = Rooting Around the Eutherian Family Tree: the Origin and Relations of the Taeniodonta|year = 2013|journal = Journal of Mammalian Evolution|pages = 1–17|doi = 10.1007/s10914-013-9230-9|volume = 21|df = }}</ref> A 2015 study has supported close affinities between pangolins and the extinct group [[Creodonta]], as well as many former [[cimolesta]]ns (''[[Cimolestes]]'' itself was recovered as a far more basal mammal).<ref>Halliday, Thomas J. D.; Upchurch, Paul; Goswami, Anjali (2015). "Resolving the relationships of Paleocene placental mammals". Biological Reviews: n/a–n/a. {{doi|10.1111/brv.12242}}. {{ISSN|1464-7931}}.</ref> |
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In 2017, [[Jackie Chan]] made a public service announcement called ''[[WildAid]]: Jackie Chan & Pangolins (Kung Fu Pangolin)''.<ref name=pangolins>{{cite news |date=23 August 2017 |title=Jackie Chan fights for pangolins |newspaper=[[China Daily]] |url=http://www.chinadaily.com.cn/world/2017-08/23/content_31006476.htm |access-date=17 March 2019}}</ref> |
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All species of living pangolin had been assigned to the genus ''Manis'' until the late 2000s, when research prompted the splitting of extant pangolins into three genera: ''Manis'', ''Phataginus'', and ''Smutsia''.<ref name="web2.utc.edu"/><ref name="KondrashovAgadjanian2012">{{cite journal|last1=Kondrashov|first1=Peter|last2=Agadjanian|first2=Alexandre K.|title=A nearly complete skeleton of ''Ernanodon'' (Mammalia, Palaeanodonta) from Mongolia: morphofunctional analysis|journal=Journal of Vertebrate Paleontology|volume=32|issue=5|year=2012|pages=983–1001|issn=0272-4634|doi=10.1080/02724634.2012.694319}}</ref> |
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In December 2020, a study found that it is "not too late" to establish conservation efforts for Philippine pangolins (''Manis culionensis''), a species that is only found on the island province of Palawan.<ref>{{Cite web|date=27 January 2021|title=It's not too late – yet – to save the Philippine pangolin, study finds|url=https://news.mongabay.com/2021/01/its-not-too-late-yet-to-save-the-philippine-pangolin-study-finds/|access-date=27 January 2021|website=Mongabay Environmental News|language=en-US}}</ref><ref>{{Cite journal|date=1 December 2020|title=Scaling up local ecological knowledge to prioritise areas for protection: Determining Philippine pangolin distribution, status and threats|journal=Global Ecology and Conservation|language=en|volume=24|pages=e01395|doi=10.1016/j.gecco.2020.e01395|issn=2351-9894|doi-access=free|last1=Archer|first1=Lucy J.|last2=Papworth|first2=Sarah K.|last3=Apale|first3=Charity M.|last4=Corona|first4=Darlyn B.|last5=Gacilos|first5=Josefa T.|last6=Amada|first6=Ronald L.|last7=Waterman|first7=Carly|last8=Turvey|first8=Samuel T.|bibcode=2020GEcoC..2401395A }}</ref> |
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{{cladogram |
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|title=Phylogenetic position of the Pholidota in the context of the order-level cladogram of Boreoeutheria. |
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|caption=The cladogram has been reconstructed from mitochondrial and nuclear DNA and protein characters. |
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|align=center |width=800 |
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|clades={{Laurasiatheria Cladogram}} |
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}} |
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===Taiwan=== |
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[[File:Manis temminckii (29390603130).jpg|thumb|[[Ground pangolin]] in defensive posture]] |
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[[Taiwan]] is one of the few conservation grounds for pangolins in the world after the country enacted the 1989 Wildlife Conservation Act.<ref>{{cite news |title=Taiwan's Path to Pangolin Conservation : How a Mega Pangolin Leather Exporter Transformed into a Conservation Specialist |url=https://www.twreporter.org/a/pangolin-conservation-history-in-taiwan-english |date=23 June 2019 |work=[[The Reporter (Taiwan)]] |access-date=8 February 2020}}</ref> The introduction of Wildlife Rehabilitation Centers in places like Luanshan ([[Yanping Township]]) in [[Taitung County|Taitung]] and [[Xiulin, Hualien|Xiulin]] townships in [[Hualien County|Hualien]] became important communities for protecting pangolins and their habitats and has greatly improved the survival of pangolins. These centers work with local aboriginal tribes and forest police in the National Police Agency to prevent poaching, trafficking, and smuggling of pangolins, especially to black markets in China. These centers have also helped to reveal the causes of death and injury among Taiwan's pangolin population.<ref>{{cite journal |last1=Pei |first1=Curtis Jai-Chyi |last2=Chen |first2=Chen-Chih |last3=Chi |first3=Meng-Jou |last4=Lin |first4=Wen-Chi |last5=Lin |first5=Jing-Shiun |last6=Arora |first6=Bharti |last7=Sun |first7=Nick Ching-Min |date=6 February 2019 |title=Mortality and morbidity in wild Taiwanese pangolin (''Manis pentadactyla pentadactyla'') |journal=PLOS ONE |location=Pingtung, Taiwan |volume=14 |issue=2 |pages=e0198230 |bibcode=2019PLoSO..1498230S |doi=10.1371/journal.pone.0198230 |pmc=6364958 |pmid=30726204 |doi-access=free}}</ref> Today, Taiwan has the highest population density of pangolins in the world.<ref>{{cite news |title=Taiwanese Researchers Collaborate With Locals In Pangolin Conservation |url=https://international.thenewslens.com/feature/pangolin-conservation/125149 |date=25 September 2019 |work=[[The News Lens]] |access-date= 8 February 2020}}</ref> |
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==See also== |
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* '''Order Pholidota''' ''sensu lato'' ([[Pholidotamorpha]])<ref name="mikko">{{cite web|last=Haaramo|first=Mikko|year=2007|title=Manidae– Recent pangolins|url=http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/synapsida/eutheria/leptictida/manidae.html|publisher=Mikko's Phylogeny Archive |accessdate= 23 May 2018}}</ref><ref>{{cite web | url=http://www.paleofile.com/|publisher=[[Paleofile.com]] | title=Taxonomic lists- Mammals| accessdate=23 May 2018}}</ref><ref>{{cite web |title=Palaeanodonta |url=http://fossilworks.org/bridge.pl?action=taxonInfo&taxon_no=43659 |publisher=Paleobiology Database |website=fossilworks.org |access-date=13 March 2017}}</ref><ref name="KondrashovAgadjanian2012"/> |
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* [[Mammal classification]] |
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** Suborder [[Palaeanodonta]] Matthew, 1918 |
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* [[Pholidotamorpha]] |
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*** Genus †''[[Arcticanodon]]'' <small>Rose, Eberle & McKenna, 2004</small> |
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**** Species †''[[Arcticanodon|Arcticanodon dawsonae]]'' <small>Rose, Eberle & McKenna, 2004</small> |
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*** Genus †''[[Melaniella]]'' <small>Fox, 1984</small> |
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**** Species †''[[Melaniella|Melaniella timosa]]'' <small>Fox, 1984</small> |
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*** Genus †''[[Molaetherium]]'' <small>Heissig, 1982</small> |
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*** Genus †''[[Amelotabes]]'' <small>Rose, 1978</small> |
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**** Species †''[[Amelotabes|Amelotabes simpsoni]]'' <small>Rose, 1978</small> |
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*** Family †[[Escavadodontidae]]'' <small>Rose & Lucas, 2000</small> |
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**** Genus †''[[Escavadodon]]'' <small>Rose & Lucas, 2000</small> |
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***** Species †''[[Escavadodon|Escavadodon zygus]]'' <small>Rose & Lucas, 2000</small> |
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*** Family †[[Epoicotheriidae]] <small>Simpson, 1927</small> |
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**** Genus †''[[Alocodontulum]]'' <small>Rose, Bown & Simons, 1978</small> |
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<!--**** Genus †''[[Alocodontulum|Alocodontulum atopum]]'' <small>(Rose, Bown & Simons, 1977) Rose, Bown & Simons, 1978</small> [''[[Alocodon]]'' <small>Rose, Bown & Simons, 1977 non Thulborn, 1973</small>; ''[[Alocodon atopum]]'' <small>Rose, Bown & Simons, 1977</small>; ''[[Tubulodon atopum]]'' <small>(Rose, Bown & Simons, 1977) Rose et al., 1991</small>] --> |
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**** Genus †''[[Auroratherium]]'' <small>Tong & Wamg, 1997</small> |
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***** Species †''[[Auroratherium|Auroratherium sinense]]'' <small>Tong & Wamg, 1997</small> |
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<!--**** Genus †''[[Pentapassalus]]'' <small>Gazin, 1952</small> |
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***** Species †''[[Pentapassalus pearcei|P. pearcei]]'' <small>Gazin, 1952</small> [''[[Tubulodon pearcei]]'' <small>(Gazin, 1952) Rose et al., 1991</small>] |
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***** Species †''[[Pentapassalus woodi|P. woodi]]'' <small>(Guthrie, 1967) Rose, 1978</small> [''[[Palaeanodon woodi]]'' <small>Guthrie, 1967</small>; ''[[Tubulodon woodi]]'' <small>(Guthrie, 1967) Rose et al., 1991</small>]--> |
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**** Genus †''[[Dipassalus]]'' <small>Rose, Krishtalka & Stucky, 1991</small> |
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***** Species †''[[Dipassalus|Dipassalus oryctes]]'' <small>Rose, Krishtalka & Stucky, 1991</small> |
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**** Genus †''[[Tetrapassalus]]'' <small>Simpson, 1959a</small> |
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***** Species †''[[Tetrapassalus mckennai|T. mckennai]]'' <small>Simpson, 1959a</small> |
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***** Species †''[[Tetrapassalus proius|T. proius]]'' <small>West, 1973c</small> |
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**** Genus †''[[Epoicotherium]]'' <small>Simpson, 1927</small> [''[[Xenotherium]]'' <small>Douglass, 1906 non Ameghino, 1904</small>; ''[[Pseudochrysochloris]]'' <small>Turnbull & Reed, 1967</small>] |
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***** Species †''[[Epoicotherium|Epoicotherium unicum]]'' <small>Douglass, 1904</small> [''[[Pseudochrysochloris yoderensis]]'' <small>Turnbull & Reed, 1967</small>] |
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**** Genus †''[[Tubulodon]]'' <small>Jepsen, 1932</small> [''[[Pentapassalus]]'' <small>Gazin, 1952</small>, ''[[Alocodon (Palaeanodonta)|Alocodon]]'' <small>Rose et al., 1977</small>] |
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***** Species †''[[Tubulodon|Tubulodon atopum]]'' <small>Rose et al., 1978</small> |
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***** Species †''[[Tubulodon|Tubulodon pearcei]]'' <small>Gazin, 1952</small> |
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***** Species †''[[Tubulodon|Tubulodon taylori]]'' <small>Jepsen, 1932</small> |
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***** Species †''[[Tubulodon|Tubulodon woodi]]'' <small>Guthrie, 1967</small> |
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**** Genus †''[[Xenocranium]]'' <small>Colbert, 1942</small> |
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***** Species †''[[Xenocranium|Xenocranium pileorivale]]'' <small>Colbert, 1942</small> |
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*** Family †[[Metacheiromyidae]] <small>Wortman, 1903</small> |
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**** Genus †''[[Propalaeanodon]]'' <small>Rose, 1979</small> |
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***** Species †''[[Propalaeanodon|Propalaeanodon schaffi]]'' <small>Rose, 1979</small> |
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**** Genus †''[[Palaeanodon]]'' <small>Matthew, 1918</small> |
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***** Species †''[[Palaeanodon parvulus|P. parvulus]]'' <small>Matthew, 1918</small> |
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***** Species †''[[Palaeanodon ignavus|P. ignavus]]'' <small>Matthew, 1918</small> |
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***** Species †''[[Palaeanodon nievelti|P. nievelti]]'' <small>Gingerich, 1989</small> |
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**** Genus †''[[Brachianodon]]'' <small>Gunnell & Gingerich, 1993</small> |
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***** Species †''[[Brachianodon|Brachianodon westorum]]'' <small>Gunnell & Gingerich, 1993</small> |
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**** Genus †''[[Mylanodon]]'' <small>Secord et al., 2002</small> |
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***** Species †''[[Mylanodon|Mylanodon rosei]]'' <small>Secord et al., 2002</small> |
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**** Genus †''[[Metacheiromys]]'' <small>Wortman, 1903</small> |
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***** Species †''[[Metacheiromys marshi|M. marshi]]'' <small>Wortman, 1903</small> [''[[Metacheiromys tatusia]]'' <small>Osborn, 1904</small>] |
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***** Species †''[[Metacheiromys dasypus|M. dasypus]]'' <small>Osborn, 1904</small> [''[[Metacheiromys osborni]]'' <small>Simpson, 1931</small>] |
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** Pholidota ''sensu stricto'' |
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*** Genus †''[[Argyromanis]]'' <small>Ameghino, 1904</small> |
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**** Species †''[[Argyromanis patagonica]] |
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*** Genus †''[[Orthoarthrus]]'' <small>Ameghino, 1904</small> |
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**** Species †''[[Orthoarthrus mixtus]] |
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*** Genus †''[[Euromanis]]'' <small>(Storch & Martin, 1994) Gaudin, Emry & Wible, 2009</small> [''[[Eomanis krebsi]]'' <small>Storch & Martin, 1994</small>] |
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**** Species †''[[Euromanis krebsi]] |
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*** Genus †''[[Eurotamandua]]'' <small>Storch, 1981</small> |
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**** Species †''[[Eurotamandua joresi]] |
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*** Infraorder [[Eupholidota]] <small>Gaudin, Emry & Wible, 2009</small> |
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**** Family †[[Eomanidae]] <small>Storch, 2003</small> |
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***** Genus †''[[Eomanis]]'' <small>Storch, 1978</small> |
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****** Species †''[[Eomanis waldi]] |
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**** Superfamily [[Manoidea]] |
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***** Family [[Patriomanidae]] <small>Szalay & Schrenk 1998 sensu Gaudin, Emry & Pogue, 2006</small> |
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****** Genus †''[[Cryptomanis]]'' <small>Gaudin, Emry & Pogue, 2006</small> |
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******* Species †''[[Cryptomanis gobiensis]] |
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****** Genus †''[[Patriomanis|Patriomanis]]'' <small>Emry, 1970</small> |
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******* Species †''[[Patriomanis americana]] |
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***** Family '''Manidae''' <small>Gray, 1821</small> |
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****** Genus †''[[Necromanis]]'' <small>Filhol, 1893</small> [''[[Leptomanis]]'' <small>Filhol, 1893</small>; ''[[Necrodasypus]]'' <small>Filhol, 1893</small>; ''[[Teutomanis]]'' <small>Ameghino, 1905</small>; ''[[Galliaetatus]]'' <small>Ameghino, 1905</small>] |
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******* Species †''[[Necromanis franconica|N. franconica]]'' <small>(Quenstedt, 1886)</small> [''[[Lutra franconica]]'' <small>Quenstedt, 1886</small>] |
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******* Species †''[[Necromanis parva|N. parva]]'' <small>Koenigswald, 1969</small> |
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******* Species †''[[Necromanis quercyi|N. quercyi]]'' <small>Filhol, 1893</small> |
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******* Species †''[[Necromanis galliae|N. galliae]]'' |
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****** Subfamily [[Smutsiinae]] <small>Gray, 1873</small> (African pangolins) |
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******* Genus ''[[Phataginus]]'' <small>Rafinesque, 1821</small><ref name="web2.utc.edu"/> [''[[Phataginus]]'' <small>Rafinesque, 1815 (nomen nudum)</small>; ''[[Triglochinopholis]]'' <small>Fitzinger, 1872</small>; ''[[Uromanis]]'' <small>Pocock, 1924</small>] |
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******** [[Tree pangolin]] (''P. tricuspis'' <small>(Rafinesque, 1821) Rafinesque, 1821</small>) |
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******** [[Long-tailed pangolin]] (''P. tetradactyla'' <small>(Linnaeus, 1766)</small>) |
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******* Genus ''[[Smutsia]]'' <small>Gray, 1865</small><ref name="web2.utc.edu"/> (African ground pangolins) |
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******** [[Giant pangolin]] (''S. gigantea'' <small>(Illiger, 1815)</small>) |
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******** [[Ground pangolin]] (''S. temmincki'' <small>(Smuts, 1832)</small>) |
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****** Subfamily [[Maninae]] <small>Gray, 1821</small> |
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******* Genus ''[[Manis]]'' <small>Linnaeus, 1758</small> [''[[Pholidotus]]'' <small>Storr, 1780</small>; ''[[Quaggelo]]'' <small>Frisch, 1775</small>; ''[[Pangolinus]]'' <small>Rafinesque, 1821</small>; ''[[Pangolinus]]'' <small>Rafinesque, 1815 nomen nudum</small>; ''Pangolin'' <small>Gray, 1873</small>; ''[[Phatages]]'' <small>Sundevall, 1843</small>] (Asiatic pangolins) |
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******** †''[[Manis hungarica|M. hungarica]]'' <small>Kormos, 1934</small> |
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******** †''[[Manis lydekkeri|M. lydekkeri]]'' <small>Trouessart</small> |
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******** †''[[Manis paleojavanica|M. paleojavanica]]'' <small>Dubois, 1907</small>) |
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******** Subgenus (''[[Manis (subgenus)|Manis]]'') <small>Linnaeus, 1758</small> |
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********* [[Indian pangolin]] (''M. crassicaudata'' <small>Gray, 1827</small>) |
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********* [[Chinese pangolin]] (''M. pentadactyla'' <small>Linnaeus, 1758</small>) |
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******** Subgenus (''[[Paramanis]]'') <small>Pocock, 1924</small> |
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********* [[Sunda pangolin]] (''M. javanica'' <small>Desmarest, 1822</small>) |
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********* [[Philippine pangolin]] (''M. culionensis'' <small>de Elera, 1895</small>) |
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==References== |
==References== |
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{{ |
{{reflist|2}} |
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==External links== |
==External links== |
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{{Wikispecies|Pholidota}} |
{{Wikispecies|Pholidota}} |
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{{Wiktionary}} |
{{Wiktionary}} |
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* [https://www.pangolinsg.org/ IUCN SSC Pangolin Specialist Group] |
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* [https://www.zsl.org/conservation/threats/illegal-wildlife-trade-crisis/pangolin-conservation ZSL Pangolin Conservation ] |
* [https://www.zsl.org/conservation/threats/illegal-wildlife-trade-crisis/pangolin-conservation ZSL Pangolin Conservation ] |
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* [https://web.archive.org/web/20090417080037/http://www.awf.org/content/wildlife/detail/pangolin Pangolin: Wildlife summary from the African Wildlife Foundation] |
* [https://web.archive.org/web/20090417080037/http://www.awf.org/content/wildlife/detail/pangolin Pangolin: Wildlife summary from the African Wildlife Foundation] |
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* [http://tolweb.org/Pholidota/15954 Tree of Life of Pholidota] |
* [http://tolweb.org/Pholidota/15954 Tree of Life of Pholidota] {{Webarchive|url=https://web.archive.org/web/20210429214815/http://tolweb.org/Pholidota/15954 |date=29 April 2021 }} |
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* [http://video.nationalgeographic.com/video/weirdest-pangolin ''National Geographic'' video of a pangolin] |
* [http://video.nationalgeographic.com/video/weirdest-pangolin ''National Geographic'' video of a pangolin] {{Webarchive|url=https://web.archive.org/web/20181023125211/https://video.nationalgeographic.com/video/weirdest-pangolin |date=23 October 2018 }} |
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* [http://cmsdata.iucn.org/downloads/pangolin_proceedings_final_print__2_.pdf Proceedings of the Workshop on Trade and Conservation of Pangolins Native to South and Southeast Asia] (PDF) |
* [https://web.archive.org/web/20111004225834/http://cmsdata.iucn.org/downloads/pangolin_proceedings_final_print__2_.pdf Proceedings of the Workshop on Trade and Conservation of Pangolins Native to South and Southeast Asia] (PDF) |
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* [http://web2.utc.edu/~gvv824/Gaudin%20et%20al%202009.pdf The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis] (PDF) |
* [http://web2.utc.edu/~gvv824/Gaudin%20et%20al%202009.pdf The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis] {{Webarchive|url=https://web.archive.org/web/20150925134257/http://web2.utc.edu/~gvv824/Gaudin%20et%20al%202009.pdf |date=25 September 2015 }} (PDF) |
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*{{cite AV media|people=Bromley, Victoria (Director/Producer), Young, Nora (Narrator/Host), Diekmann, Maria|date=2018|title=Nature: The World's Most Wanted Animal| |
*{{cite AV media |people=Bromley, Victoria (Director/Producer), Young, Nora (Narrator/Host), Diekmann, Maria |date=2018 |title=Nature: The World's Most Wanted Animal |publisher=PBS |location=United States |url=https://www.pbs.org/video/the-worlds-most-wanted-animal-evunge/}} |
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* [https://www.nytimes.com/2020/03/05/opinion/coronavirus-china-pangolins.html Coronavirus: Revenge of the Pangolins?] ''[[The New York Times]]'', 6 March 2020. |
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{{Mammals}} |
{{Mammals}} |
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{{Pholidotamorpha|state=autocollapse}} |
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{{Pholidota}} |
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{{Taxonbar|from=Q2191516}} |
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{{ |
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[[Category:Critically endangered animals]] |
[[Category:Critically endangered animals]] |
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[[Category:Mammals of Africa]] |
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[[Category:Mammals of Asia]] |
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[[Category:Myrmecophagous mammals]] |
[[Category:Myrmecophagous mammals]] |
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[[Category:Pangolins]] |
[[Category:Pangolins| ]] |
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[[Category:Rolling animals]] |
[[Category:Rolling animals]] |
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[[Category:Extant Paleocene first appearances]] |
[[Category:Extant Paleocene first appearances]] |
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[[Category:Species endangered by human consumption]] |
[[Category:Species endangered by human consumption]] |
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[[Category:Species endangered by human consumption for medicinal or magical purposes]] |
[[Category:Species endangered by human consumption for medicinal or magical purposes]] |
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[[Category:Taxa named by John Edward Gray]] |
Latest revision as of 19:53, 22 November 2024
Pangolin Temporal range: Eocene – present
Middle | |
---|---|
Indian pangolin | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Mirorder: | Ferae |
Clade: | Pholidotamorpha |
Order: | Pholidota Weber, 1904 |
Subgroups | |
[see classification]
| |
Ranges of living species | |
Synonyms | |
list of synonyms:
|
Pangolins, sometimes known as scaly anteaters,[12] are mammals of the order Pholidota (/fɒlɪˈdoʊtə/). The one extant family, the Manidae, has three genera: Manis, Phataginus, and Smutsia.[13] Manis comprises four species found in Asia, while Phataginus and Smutsia include two species each, all found in sub-Saharan Africa.[14] These species range in size from 30 to 100 cm (12 to 39 in). Several extinct pangolin species are also known. In September 2023, nine species were reported.[15]
Pangolins have large, protective keratin scales, similar in material to fingernails and toenails, covering their skin; they are the only known mammals with this feature. Depending on the species, they live in hollow trees or burrows. Pangolins are nocturnal, and their diet consists of mainly ants and termites, which they capture using their long tongues. They tend to be solitary animals, meeting only to mate and produce a litter of one to three offspring, which they raise for about two years. Pangolins superficially resemble armadillos, though the two are not closely related; they have merely undergone convergent evolution.
Pangolins are threatened by poaching (for their meat and scales, which are used in traditional medicine[16][17]) and heavy deforestation of their natural habitats, and are the most trafficked mammals in the world.[18] As of January 2020[update], there are eight species of pangolin whose conservation status is listed in the threatened tier. Three (Manis culionensis, M. pentadactyla and M. javanica) are critically endangered, three (Phataginus tricuspis, Manis crassicaudata and Smutsia gigantea) are endangered and two (Phataginus tetradactyla and Smutsia temminckii) are vulnerable on the Red List of Threatened Species of the International Union for Conservation of Nature.[19]
Etymology
[edit]The name of order Pholidota comes from Ancient Greek ϕολιδωτός – "clad in scales"[20] from φολίς pholís "scale".[21]
The name "pangolin" comes from the Malay word pengguling meaning "one who rolls up"[22] from guling or giling "to roll"; it was used for the Sunda pangolin (Manis javanica).[23] However, the modern name is tenggiling. In Javanese, it is terenggiling;[23] and in the Philippine languages, it is goling, tanggiling, or balintong (with the same meaning).[24]
In ancient India, according to Aelian, it was known as the phattáges (φαττάγης).[25]
Description
[edit]The physical appearance of a pangolin is marked by large, hardened, overlapping, plate-like scales, which are soft on newborn pangolins, but harden as the animal matures.[26] They are made of keratin, the same material from which human fingernails and tetrapod claws are made, and are structurally and compositionally very different from the scales of reptiles.[27] The pangolin's scaled body is comparable in appearance to a pine cone. It can curl up into a ball when threatened, with its overlapping scales acting as armor, while it protects its face by tucking it under its tail. The scales are sharp, providing extra defense from predators.[28]
Pangolins can emit a noxious-smelling chemical from glands near the anus, similar to the spray of a skunk.[29] They have short legs, with sharp claws which they use for burrowing into ant and termite mounds and for climbing.[30]
The tongues of pangolins are extremely long, and like those of the giant anteater and the tube-lipped nectar bat, the root of the tongue is not attached to the hyoid bone but is in the thorax between the sternum and the trachea.[31] Large pangolins can extend their tongues as much as 40 cm (16 in), with a diameter of only about 0.5 cm (1⁄5 in).[32]
Behaviour
[edit]Most pangolins are nocturnal animals[33] which use their well-developed sense of smell to find insects. The long-tailed pangolin is also active by day, while other species of pangolins spend most of the daytime sleeping, curled up into a ball ("volvation").[32]
Arboreal pangolins live in hollow trees, whereas the ground-dwelling species dig tunnels to a depth of 3.5 m (11 ft).[32]
Some pangolins walk with their front claws bent under the foot pad, although they use the entire foot pad on their rear limbs. Furthermore, some exhibit a bipedal stance for some behavior, and may walk a few steps bipedally.[34] Pangolins are also good swimmers.[32]
Diet
[edit]Pangolins are insectivorous. Most of their diet consists of various species of ants and termites and may be supplemented by other insects, especially larvae. They are somewhat particular and tend to consume only one or two species of insects, even when many species are available. A pangolin can consume 140 to 200 g (5 to 7 oz) of insects per day.[35] Pangolins are an important regulator of termite populations in their natural habitats.[36]
Pangolins have very poor vision. They also lack teeth. They rely heavily on smell and hearing, and they have other physical characteristics to help them eat ants and termites. Their skeletal structure is sturdy and they have strong front legs used for tearing into termite mounds.[37] They use their powerful front claws to dig into trees, soil, and vegetation to find prey,[38] then proceed to use their long tongues to probe inside the insect tunnels and to retrieve their prey.
The structure of their tongue and stomach is key to aiding pangolins in obtaining and digesting insects. Their saliva is sticky,[37] causing ants and termites to stick to their long tongues when they are hunting through insect tunnels. Without teeth, pangolins cannot also chew;[39] but while foraging, they ingest small stones (gastroliths), which accumulate in their stomachs to help to grind up ants.[40] This part of their stomach is called the gizzard, and it is also covered in keratinous spines.[41] These spines further aid in the grinding up and digestion of the pangolin's prey.
Some species, such as the tree pangolin, use their strong, prehensile tails to hang from tree branches and strip away bark from the trunk, exposing insect nests inside.[42]
Reproduction
[edit]Pangolins are solitary and meet only to have sex, with mating typically taking place at night after the male and female pangolin meet near a watering hole. Males are larger than females, weighing up to 40% more. While the mating season is not defined, they typically mate once each year, usually during the summer or autumn. Rather than the males seeking out the females, males mark their location with urine or feces and the females find them. If competition over a female occurs, the males use their tails as clubs to fight for the opportunity to mate with her.[44]
Gestation periods differ by species, ranging from roughly 70 to 140 days.[45] African pangolin females usually give birth to a single offspring at a time, but the Asiatic species may give birth to from one to three.[32] Weight at birth is 80 to 450 g (2+3⁄4 to 15+3⁄4 oz), and the average length is 150 mm (6 in). At the time of birth, the scales are soft and white. After several days, they harden and darken to resemble those of an adult pangolin. During the vulnerable stage, the mother stays with her offspring in the burrow, nursing it, and wraps her body around it if she senses danger. The young cling to the mother's tail as she moves about, although, in burrowing species, they remain in the burrow for the first 2–4 weeks of life. At one month, they first leave the burrow riding on the mother's back. Weaning takes place around three months of age, when the young begin to eat insects in addition to nursing. At two years of age, the offspring are sexually mature and are abandoned by the mother.[46]
Classification and phylogeny
[edit]Taxonomy
[edit]- Order: Pholidota Weber, 1904
- Genus: †Euromanis Gaudin, Emry & Wible, 2009
- Family: †Eurotamanduidae Szalay & Schrenk, 1994
- Suborder: Eupholidota Gaudin, Emry & Wible, 2009
- Superfamily: Manoidea Gaudin, Emry & Wible, 2009
- Family: Manidae Gray, 1821
- Family: †Patriomanidae Szalay & Schrenk 1998 [sensu Gaudin, Emry & Pogue, 2006]
- Incertae sedis
- Genus: †Necromanis Filhol, 1893
- Superfamily: †Eomanoidea Gaudin, Emry & Wible, 2009
- Family: †Eomanidae Storch, 2003
- Superfamily: Manoidea Gaudin, Emry & Wible, 2009
Phylogeny
[edit]Among placentals
[edit]The order Pholidota was long considered to be the sister taxon to Xenarthra (neotropical anteaters, sloths, and armadillos), but recent genetic evidence indicates their closest living relatives are the carnivorans, with which they form a clade, the Ferae.[47][48][49][50] Palaeanodonts are even closer relatives to pangolins, being classified with pangolins in the clade Pholidotamorpha.[51] The split between carnivorans and pangolins is estimated to have occurred 79.47 Ma (million years) ago.[52]
Among Manidae
[edit]The first dichotomy in the phylogeny of extant Manidae separates Asian pangolins (Manis) from African pangolins (Smutsia and Phataginus).[50] Within the former, Manis pentadactyla is the sister group to a clade comprising M. crassicaudata and M. javanica. Within the latter, a split separates the large terrestrial African pangolins of the genus Smutsia from the small arboreal African pangolins of the genus Phataginus.[53]
Asian and African pangolins are thought to have diverged about 41.37 Ma ago.[52] Moreover, the basal position of Manis within Pholidota[50][54] suggests the group originated in Eurasia, consistent with their laurasiatherian phylogeny.[50]
Threats
[edit]Pangolins are in high demand in southern China and Vietnam because their scales are believed to have medicinal properties in traditional Chinese and Vietnamese medicine.[56] Their meat is also considered a delicacy.[57][58][59][60][61] 100,000 are estimated to be trafficked a year to China and Vietnam,[62] amounting to over one million over the past decade.[63][64] This makes them the most trafficked animal in the world.[63][65] This, coupled with deforestation, has led to a large decrease in the numbers of pangolins. Some species, such as Manis pentadactyla have become commercially extinct in certain ranges as a result of overhunting.[66] In November 2010, pangolins were added to the Zoological Society of London's list of evolutionarily distinct and endangered mammals.[67] All eight species of pangolin are assessed as threatened by the IUCN, while three are classified as critically endangered.[19] All pangolin species are currently listed under Appendix I of CITES which prohibits international trade, except when the product is intended for non-commercial purposes and a permit has been granted.[68]
China had been the main destination country for pangolins until 2018, where it was surpassed by Vietnam. In 2019, Vietnam was reported to have seized the largest volumes of pangolin scales, surpassing Nigeria that year.[69]
Pangolins are also hunted and eaten in Ghana and are one of the more popular types of bushmeat, while local healers use the pangolin as a source of traditional medicine.[70]
Though pangolins are protected by an international ban on their trade, populations have suffered from illegal trafficking due to beliefs in East Asia that their ground-up scales can stimulate lactation or cure cancer or asthma.[71] In the past decade, numerous seizures of illegally trafficked pangolin and pangolin meat have taken place in Asia.[72][73][74][75] In one such incident in April 2013, 10,000 kg (22,000 pounds) of pangolin meat were seized from a Chinese vessel that ran aground in the Philippines.[76][77] In another case in August 2016, an Indonesian man was arrested after police raided his home and found over 650 pangolins in freezers on his property.[78] The same threat is reported in Nigeria, where the animal is on the verge of extinction due to overexploitation.[79] The overexploitation comes from hunting pangolins for game meat and the reduction of their forest habitats due to deforestation caused by timber harvesting.[80] The pangolin are hunted as game meat for both medicinal purposes and food consumption.[80]
Virology
[edit]COVID-19 infection
[edit]The nucleic acid sequence of a specific receptor-binding domain of the spike protein belonging to coronaviruses taken from pangolins was found to be a 99% match with SARS coronavirus 2 (SARS-CoV-2), the virus which causes COVID-19 and is responsible for the COVID-19 pandemic.[81][82] Researchers in Guangzhou, China, hypothesized that SARS-CoV-2 had originated in bats, and prior to infecting humans, was circulating among pangolins. The illicit Chinese trade of pangolins for use in traditional Chinese medicine was suggested as a vector for human transmission.[81][83] However, whole-genome comparison found that the pangolin and human coronaviruses share only up to 92% of their RNA.[84][85] Ecologists worried that the early speculation about pangolins being the source may have led to mass slaughters, endangering them further, which was similar to what happened to Asian palm civets during the SARS outbreak.[84][86] It was later proved that the testing which suggested that pangolins were a potential host for the virus was flawed, when genetic analysis showed that the spike protein and its binding to receptors in pangolins had minimal effect from the virus, and therefore were not likely mechanisms for COVID-19 infections in humans.[87]
Pestivirus and Coltivirus
[edit]In 2020, two novel RNA viruses distantly related to pestiviruses and coltiviruses have been detected in the genomes of dead Manis javanica and Manis pentadactyla.[88] To refer to both sampling site and hosts, they were named Dongyang pangolin virus (DYPV) and Lishui pangolin virus (LSPV). The DYPV pestivirus was also identified in Amblyomma javanense nymph ticks from a diseased pangolin.[88]
Folk medicine
[edit]Pangolin scales and flesh are used as ingredients for various traditional Chinese medicine preparations.[89] While no scientific evidence exists for the efficacy of those practices, and they have no logical mechanism of action,[90][91][92] their popularity still drives the black market for animal body parts, despite concerns about toxicity, transmission of diseases from animals to humans, and species extermination.[89][93] The ongoing demand for parts as ingredients continues to fuel pangolin poaching, hunting and trading.[94]
The first record of pangolin scales occurs in Ben Cao Jinji Zhu ("Variorum of Shennong's Classic of Materia Medica", 500 CE), which recommends pangolin scales for protection against ant bites; burning the scales as a cure for people crying hysterically during the night.[95] During the Tang dynasty, a recipe for expelling evil spirits with a formulation of scales, herbs, and minerals appeared in 682, and in 752 CE the idea that pangolin scales could also stimulate milk secretion in lactating women, one of the main uses today, was recommended in the Wai Tai Mi Yao ("Arcane Essentials from the Imperial Library").[95] In the Song dynasty, the notion of penetrating and clearing blockages was emphasized in the Taiping sheng hui fan ("Formulas from Benevolent Sages Compiled During the Era of Peace and Tranquility"), compiled by Wang Huaiyin in 992.[95]
In the 21st century, the main uses of pangolin scales are quackery practices based on unproven claims the scales dissolve blood clots, promote blood circulation, or help lactating women secrete milk.[89][95] The supposed health effects of pangolin meat and scales claimed by folk medicine practitioners are based on their consumption of ants, long tongues, and protective scales.[89]
The official pharmacopoeia of the People's Republic of China included Chinese pangolin scales as an ingredient in TCM formulations.[95] Pangolins were removed from the pharmacopoeia starting from the first half of 2020.[96] Although pangolin scales have been removed from the list of raw ingredients, the scales are still listed as a key ingredient in various medicines.[97]
Pangolin parts are also used for medicinal purposes in other Asian countries such as India, Nepal and Pakistan. In some parts of India and Nepal, locals believe that wearing the scales of a pangolin can help prevent pneumonia.[98] Pangolin scales have also been used for medicinal purposes in Malaysia, Indonesia and northern Myanmar. Indigenous people in southern Palawan, Philippines, have held the belief that elders could avoid prostate illnesses by wearing belts made with the scales.[99]
Conservation
[edit]As a result of increasing threats to pangolins, mainly in the form of illegal, international trade in pangolin skin, scales, and meat, these species have received increasing conservation attention in recent years.[100] As of January 2020[update], the IUCN considered all eight species of pangolin on its Red List of Threatened Species as threatened.[19] The IUCN SSC Pangolin Specialist Group launched a global action plan to conserve pangolins, dubbed "Scaling up Pangolin Conservation", in July 2014. This action plan aims to improve all aspects of pangolin conservation with an added emphasis on combating poaching and trafficking of the animal while educating communities on its importance.[63] Another suggested approach to fighting pangolin (and general wildlife) trafficking consists in "following the money" rather than "the animal", which aims to disrupt smugglers' profits by interrupting money flows. Financial intelligence gathering could thus become a key tool in protecting these animals, although this opportunity is often overlooked.[62] In 2018, a Chinese NGO launched the Counting Pangolins movement, calling for joint efforts to save the mammals from trafficking.[101][102][103] Wildlife conservation group TRAFFIC has identified 159 smuggling routes used by pangolin traffickers and aims to shut these down.[104]
Many attempts have been made to breed pangolins in captivity, but due to their reliance on wide-ranging habitats and very particular diets, these attempts are often unsuccessful.[45][105] Pangolins have significantly decreased immune responses due to a genetic dysfunction, making them extremely fragile.[106] They are susceptible to diseases such as pneumonia and the development of ulcers in captivity, complications that can lead to an early death.[45] In addition, pangolins rescued from illegal trade often have a higher chance of being infected with parasites such as intestinal worms, further lessening their chance for rehabilitation and reintroduction to the wild.[45]
The idea of farming pangolins to reduce the number being illegally trafficked is being explored with little success.[107] The third Saturday in February is promoted as World Pangolin Day by the conservation NPO Annamiticus.[108] World Pangolin Day has been noted for its effectiveness in generating awareness about pangolins.[109]
In 2017, Jackie Chan made a public service announcement called WildAid: Jackie Chan & Pangolins (Kung Fu Pangolin).[110]
In December 2020, a study found that it is "not too late" to establish conservation efforts for Philippine pangolins (Manis culionensis), a species that is only found on the island province of Palawan.[111][112]
Taiwan
[edit]Taiwan is one of the few conservation grounds for pangolins in the world after the country enacted the 1989 Wildlife Conservation Act.[113] The introduction of Wildlife Rehabilitation Centers in places like Luanshan (Yanping Township) in Taitung and Xiulin townships in Hualien became important communities for protecting pangolins and their habitats and has greatly improved the survival of pangolins. These centers work with local aboriginal tribes and forest police in the National Police Agency to prevent poaching, trafficking, and smuggling of pangolins, especially to black markets in China. These centers have also helped to reveal the causes of death and injury among Taiwan's pangolin population.[114] Today, Taiwan has the highest population density of pangolins in the world.[115]
See also
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External links
[edit]- ZSL Pangolin Conservation
- Pangolin: Wildlife summary from the African Wildlife Foundation
- Tree of Life of Pholidota Archived 29 April 2021 at the Wayback Machine
- National Geographic video of a pangolin Archived 23 October 2018 at the Wayback Machine
- Proceedings of the Workshop on Trade and Conservation of Pangolins Native to South and Southeast Asia (PDF)
- The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis Archived 25 September 2015 at the Wayback Machine (PDF)
- Bromley, Victoria (Director/Producer), Young, Nora (Narrator/Host), Diekmann, Maria (2018). Nature: The World's Most Wanted Animal. United States: PBS.
- Coronavirus: Revenge of the Pangolins? The New York Times, 6 March 2020.