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A '''black panther''' is the [[Melanism|melanistic]] colour variant of the [[leopard]] (''Panthera pardus'') and [[jaguar]] (''Panthera onca''). Black panthers of both species have excess black [[pigment]]s, but their typical [[Rosette (zoology)|rosettes]] are also present. They have been documented mostly in [[tropical forest]]s, with black leopards in Africa and Asia, and black jaguars in South America. Melanism is caused by a [[recessive allele]] in the leopard, and by a [[dominant allele]] in the jaguar.
A '''black panther''' is the [[Melanism|melanistic]] colour variant of the [[leopard]] (''Panthera pardus'') and the [[jaguar]] (''Panthera onca''). Black panthers of both species have excess black [[pigment]]s, but their typical [[Rosette (zoology)|rosettes]] are also present. They have been documented mostly in [[tropical forest]]s, with black leopards in Africa and Asia, and black jaguars in South America. Melanism is caused by a [[recessive allele]] in the leopard, and by a [[dominant allele]] in the jaguar.


==Leopard==
==Leopard==
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|image1=Black Panther - India.jpg |caption1=A melanistic [[Indian leopard]] in [[Nagarhole National Park]], [[Karnataka]]
|image1=Black Panther - India.jpg |caption1=A melanistic [[Indian leopard]] in [[Nagarhole National Park]], [[Karnataka]]
|image2=Blackleopard.JPG |caption2=A melanistic leopard in Out of Africa Wildlife Park, [[Camp Verde, Arizona]]
|image2=Blackleopard.JPG |caption2=A melanistic leopard in Out of Africa Wildlife Park, [[Camp Verde, Arizona]]
|image3=Black leopard.JPG |caption3=Markings on a female black leopard at the Rhino and Lion Nature Reserve, [[Kromdraai]]}}
|image3=Black leopard.JPG |caption3=Particularly prominent [[Rosette (zoology)|rosette]]s on a female black leopard at the Rhino and Lion Nature Reserve, [[Kromdraai]]}}
In 1788, [[Jean-Claude Delamétherie]] described a black leopard that was kept in the [[Tower of London]] and had been brought from [[Bengal]].<ref>{{cite journal |last1=Delamétherie |first1=J.-C. |year=1788 |title=Description d'une Panthère noire |trans-title=Description of a black Panther |journal=Observations et Mémoires sur la Physique, sur l'Histoire Naturelle et sur les Arts et Métiers, etc. |volume=33 |page=45 |language=fr |url=https://archive.org/details/observationsetm33pari/page/45}}</ref> In 1794, [[Friedrich Albrecht Anton Meyer]] proposed the [[scientific name]] ''Felis fusca'' for this cat, the [[Indian leopard]] (''P. p. fusca'').<ref name=Meyer1794>{{cite book |last=Meyer, F. A. A. |year=1794 |title=Zoologische Annalen |volume=Erster Band |location=Weimar |publisher=Im Verlage des Industrie-Comptoirs |pages=394–396 |language=de |chapter=Über de la Metheries schwarzen Panther [About de la Metheries black Panther] |chapter-url=https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394 |access-date=14 January 2020 |archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505145120/https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394#v=onepage&q&f=false |url-status=live }}</ref><ref name=Pocock1930b>{{cite journal |author=Pocock, R. I. |year=1930 |title=The Panthers and Ounces of Asia |journal=Journal of the Bombay Natural History Society |volume=34 |issue=1 |pages=307–336 |url=https://archive.org/details/journalofbomb34121930bomb/page/306/mode/2up}}</ref>
In 1788, [[Jean-Claude Delamétherie]] described a black leopard that was kept in the [[Tower of London]] and had been brought from [[Bengal]].<ref>{{cite journal |last1=Delamétherie |first1=J.-C. |year=1788 |title=Description d'une Panthère noire |trans-title=Description of a black Panther |journal=Observations et Mémoires sur la Physique, sur l'Histoire Naturelle et sur les Arts et Métiers, etc. |volume=33 |page=45 |language=fr |url=https://archive.org/details/observationsetm33pari/page/45}}</ref> In 1794, [[Friedrich Albrecht Anton Meyer]] proposed the [[scientific name]] ''Felis fusca'' for this cat, the [[Indian leopard]] (''P. p. fusca'').<ref name=Meyer1794>{{cite book |last=Meyer, F. A. A. |year=1794 |title=Zoologische Annalen |volume=Erster Band |location=Weimar |publisher=Im Verlage des Industrie-Comptoirs |pages=394–396 |language=de |chapter=Über de la Metheries schwarzen Panther [About de la Metheries black Panther] |chapter-url=https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394 |access-date=14 January 2020 |archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505145120/https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394#v=onepage&q&f=false |url-status=live }}</ref><ref name=Pocock1930b>{{cite journal |author=Pocock, R. I. |year=1930 |title=The Panthers and Ounces of Asia |journal=Journal of the Bombay Natural History Society |volume=34 |issue=1 |pages=307–336 |url=https://archive.org/details/journalofbomb34121930bomb/page/306/mode/2up}}</ref>
In 1809, [[Georges Cuvier]] described a black leopard kept in the [[Ménagerie du Jardin des plantes]] that had been brought from [[Java]]. Cuvier proposed the name ''Felis melas'', the [[Javan leopard]] (''P. p. melas'').<ref name=Pocock1930b/><ref>{{cite journal |author=Cuvier, G. |year=1809 |title=Recherches sur les espėces vivantes de grands chats, pour servir de preuves et d'éclaircissement au chapitre sur les carnassiers fossils |journal=Annales du Muséum National d'Histoire Naturelle |volume=XIV |pages=136–164 |url=https://archive.org/details/annalesdumusum14mus/page/152/mode/2up}}</ref> By the late 19th century, the occurrence of black and spotted leopard cubs in the same [[Litter (zoology)|litter]] had been repeatedly recorded in [[India]]. Black leopards were thought to be more common in [[Travancore]] and in the hills of [[southern India]] than in other parts of the country.<ref>{{cite book |author=Blanford, W. T. |year=1888 |title=The Fauna of British India, including Ceylon and Burma. Mammalia: Volume 1 |publisher=Taylor and Francis |location=London |pages=67–71 |chapter=''Felis pardus''. The Leopard or Panther |chapter-url=https://archive.org/details/mammalia00blaniala/page/66/mode/2up}}</ref> Black leopards were also frequently encountered in southern [[Myanmar]].<ref>{{cite book |title=Burma, its people and productions; or, Notes on the fauna, flora, and minerals of Tenasserim, Pegu, and Burma |author=Mason, F. |year=1882 |place=Hertford |publisher=Chief Commissioner of British Burma |edition=Rewritten and enlarged by W. Theobald |volume=1. Geology, mineralogy, and zoology |chapter=''F. pardus'', L. The leopard |page=472 |chapter-url=https://archive.org/details/burmaitspeoplepr01maso/page/472/mode/2up}}</ref>
In 1809, [[Georges Cuvier]] described a black leopard kept in the [[Ménagerie du Jardin des plantes]] that had been brought from [[Java]]. Cuvier proposed the name ''Felis melas'', the [[Javan leopard]] (''P. p. melas'').<ref name=Pocock1930b/><ref>{{cite journal |author=Cuvier, G. |year=1809 |title=Recherches sur les espėces vivantes de grands chats, pour servir de preuves et d'éclaircissement au chapitre sur les carnassiers fossils |journal=Annales du Muséum National d'Histoire Naturelle |volume=XIV |pages=136–164 |url=https://archive.org/details/annalesdumusum14mus/page/152/mode/2up}}</ref> By the late 19th century, the occurrence of black and spotted leopard cubs in the same [[Litter (zoology)|litter]] had been repeatedly recorded in [[India]]. Black leopards were thought to be more common in [[Travancore]] and in the hills of [[southern India]] than in other parts of the country.<ref>{{cite book |author=Blanford, W. T. |year=1888 |title=The Fauna of British India, including Ceylon and Burma. Mammalia: Volume 1 |publisher=Taylor and Francis |location=London |pages=67–71 |chapter=''Felis pardus''. The Leopard or Panther |chapter-url=https://archive.org/details/mammalia00blaniala/page/66/mode/2up}}</ref> Black leopards were also frequently encountered in southern [[Myanmar]].<ref>{{cite book |title=Burma, its people and productions; or, Notes on the fauna, flora, and minerals of Tenasserim, Pegu, and Burma |author=Mason, F. |year=1882 |place=Hertford |publisher=Chief Commissioner of British Burma |edition=Rewritten and enlarged by W. Theobald |volume=1. Geology, mineralogy, and zoology |chapter=''F. pardus'', L. The leopard |page=472 |chapter-url=https://archive.org/details/burmaitspeoplepr01maso/page/472/mode/2up}}</ref>
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At least one black leopard was photographed in mixed [[deciduous]] forest in Thailand's [[Kaeng Krachan National Park]] during a one-year-long camera trapping survey from 2003 to 2004.<ref>{{cite journal |author1=Ngoprasert, D. |name-list-style=amp |author2=Lynam, A. J. |author3=Gale, G. A. |year=2007 |title=Human disturbance affects habitat use and behaviour of Asiatic leopard "Panthera pardus" in Kaeng Krachan National Park, Thailand |journal=Oryx |volume=41 |issue=3 |pages=343–351 |doi=10.1017/S0030605307001102 |doi-access=free}}</ref> In 2009, black leopards were photographed more often than spotted leopards in [[Kui Buri National Park]].<ref>{{cite report |author1=Steinmetz, R. |name-list-style=amp |author2=Seuaturien, N. |author3=Chutipong, W. |author4=Poonnil, B. |year=2009 |title=The ecology and conservation of tigers and their prey in Kuiburi National Park, Thailand |publisher=WWF Thailand, Department of National Parks, Wildlife and Plant Conservation |location=Bangkok |url=https://wwfasia.awsassets.panda.org/downloads/kuiburi_tiger_project_annual_report__aug_31_2009.pdf |archive-url=https://web.archive.org/web/20210502194409/https://wwfasia.awsassets.panda.org/downloads/kuiburi_tiger_project_annual_report__aug_31_2009.pdf |archive-date=2021-05-02 |url-status=live}}</ref>
At least one black leopard was photographed in mixed [[deciduous]] forest in Thailand's [[Kaeng Krachan National Park]] during a one-year-long camera trapping survey from 2003 to 2004.<ref>{{cite journal |author1=Ngoprasert, D. |name-list-style=amp |author2=Lynam, A. J. |author3=Gale, G. A. |year=2007 |title=Human disturbance affects habitat use and behaviour of Asiatic leopard "Panthera pardus" in Kaeng Krachan National Park, Thailand |journal=Oryx |volume=41 |issue=3 |pages=343–351 |doi=10.1017/S0030605307001102 |doi-access=free}}</ref> In 2009, black leopards were photographed more often than spotted leopards in [[Kui Buri National Park]].<ref>{{cite report |author1=Steinmetz, R. |name-list-style=amp |author2=Seuaturien, N. |author3=Chutipong, W. |author4=Poonnil, B. |year=2009 |title=The ecology and conservation of tigers and their prey in Kuiburi National Park, Thailand |publisher=WWF Thailand, Department of National Parks, Wildlife and Plant Conservation |location=Bangkok |url=https://wwfasia.awsassets.panda.org/downloads/kuiburi_tiger_project_annual_report__aug_31_2009.pdf |archive-url=https://web.archive.org/web/20210502194409/https://wwfasia.awsassets.panda.org/downloads/kuiburi_tiger_project_annual_report__aug_31_2009.pdf |archive-date=2021-05-02 |url-status=live}}</ref>
Most leopards recorded at 16 sites south of the [[Kra Isthmus]] between 1996 and 2009 were black, indicating a near-[[Fixed allele|fixation]] of melanism in Peninsular Malaysia.<ref>{{cite journal |last=Kawanishi |first=K. |author2=Sunquist, M. E. |author3=Eizirik, E. |author4=Lynam, A. J. |author5=Ngoprasert, D. |author6=Shahruddin, W. N. W. |author7=Rayan, D. M. |author8=Sharma, D. S. K. |author9=Steinmetz, R. |name-list-style=amp |title=Near fixation of melanism in panthers of the Malay Peninsula |journal=Journal of Zoology |year=2010 |volume=282 |issue=3 |pages=201–206 |doi=10.1111/j.1469-7998.2010.00731.x}}</ref> In 2019, a black individual was photographed outside a protected area in [[Jeli District]].<ref>{{cite journal |author1=Hambali, K. |name-list-style=amp |author2=Fazli, N. F. M. |author3=Amir, A. |author4=Fauzi, N. |author5=Hassin, N. H. |author6=Abas, M. A. |author7=Karim, M. F. A. |author8=Sow, A. Y. |year=2021 |title=The discovery of a melanistc Leopard ''Panthera pardus delacouri'' (Linnaeus, 1758) (Mammalia: Carnivora: Felidae) at Bukit Kudung in Jeli, Kelantan, Peninsular Malaysia: conservation and ecotourism |journal=Journal of Threatened Taxa |volume=13 |issue=1 |pages=17513–17516 |doi=10.11609/jott.6060.13.1.17513-17516 |doi-access=free}}</ref>
Most leopards recorded at 16 sites south of the [[Kra Isthmus]] between 1996 and 2009 were black, indicating a near-[[Fixed allele|fixation]] of melanism in Peninsular Malaysia.<ref>{{cite journal |last=Kawanishi |first=K. |author2=Sunquist, M. E. |author3=Eizirik, E. |author4=Lynam, A. J. |author5=Ngoprasert, D. |author6=Shahruddin, W. N. W. |author7=Rayan, D. M. |author8=Sharma, D. S. K. |author9=Steinmetz, R. |name-list-style=amp |title=Near fixation of melanism in panthers of the Malay Peninsula |journal=Journal of Zoology |year=2010 |volume=282 |issue=3 |pages=201–206 |doi=10.1111/j.1469-7998.2010.00731.x}}</ref> In 2019, a black individual was photographed outside a protected area in [[Jeli District]].<ref>{{cite journal |author1=Hambali, K. |name-list-style=amp |author2=Fazli, N. F. M. |author3=Amir, A. |author4=Fauzi, N. |author5=Hassin, N. H. |author6=Abas, M. A. |author7=Karim, M. F. A. |author8=Sow, A. Y. |year=2021 |title=The discovery of a melanistc Leopard ''Panthera pardus delacouri'' (Linnaeus, 1758) (Mammalia: Carnivora: Felidae) at Bukit Kudung in Jeli, Kelantan, Peninsular Malaysia: conservation and ecotourism |journal=Journal of Threatened Taxa |volume=13 |issue=1 |pages=17513–17516 |doi=10.11609/jott.6060.13.1.17513-17516 |doi-access=free}}</ref>
Both black and spotted leopards were recorded in [[Gunung Gede Pangrango National Park]] in [[West Java]] between 2005 and 2017.<ref>{{cite journal |author1=Ario, A. |name-list-style=amp |author2=Supian |author3=Hidayat, E. |author4=Hidayatullah, R. |author5=Rustiadi, A. |author6=Gunawan, A. |author7=Triprajawan, T. |author8=Sopian, I. |author9=Zatnika, R. R. |author10=Yusup, D. M. |author11=Hindrayani, W. |author12=Mulyanto, A. |author13=Iskandar, D. |year=2018 |title=Population dynamics and ecology of Javan leopard, ''Panthera pardus melas'', in Gunung Gede Pangrango National Park, West Java |journal=Journal of Indonesian Natural History |volume=6 |issue=1 |pages=6–13 |url=http://jinh.fmipa.unand.ac.id/index.php/jinh/article/view/106/75 |access-date=15 May 2021 |archive-date=27 September 2021 |archive-url=https://web.archive.org/web/20210927124141/http://jinh.fmipa.unand.ac.id/index.php/jinh/article/view/106/75 |url-status=live }}</ref>
Both black and spotted leopards were recorded in [[Gunung Gede Pangrango National Park]] in [[West Java]] between 2005 and 2017.<ref>{{cite journal |author1=Ario, A. |name-list-style=amp |author2=Supian |author3=Hidayat, E. |author4=Hidayatullah, R. |author5=Rustiadi, A. |author6=Gunawan, A. |author7=Triprajawan, T. |author8=Sopian, I. |author9=Zatnika, R. R. |author10=Yusup, D. M. |author11=Hindrayani, W. |author12=Mulyanto, A. |author13=Iskandar, D. |year=2018 |title=Population dynamics and ecology of Javan leopard, ''Panthera pardus melas'', in Gunung Gede Pangrango National Park, West Java |journal=Journal of Indonesian Natural History |volume=6 |issue=1 |pages=6–13 |url=http://jinh.fmipa.unand.ac.id/index.php/jinh/article/view/106/75 |access-date=15 May 2021 |archive-date=27 September 2021 |archive-url=https://web.archive.org/web/20210927124141/http://jinh.fmipa.unand.ac.id/index.php/jinh/article/view/106/75 |url-status=live}}</ref>


Frequency of melanism appears to be approximately 11% over the leopard's range. Data on the distribution of leopard populations indicates that melanism occurs in five subspecies in the wild: the Indian leopard, Javan leopard, African leopard, [[Indochinese leopard]] (''P. p. delacouri'') and [[Sri Lankan leopard]] (''P. p. kotiya''). Based on records from camera traps, melanistic leopards occur foremost in [[tropical and subtropical moist broadleaf forests]].<ref name=daSilva17>{{cite journal |author1=Da Silva L. G. |author2=Kawanishi, K. |first=K. |author3=Henschel P. |author4=Kittle, A. |author5=Sanei, A. |author6=Reebin, A. |author7=Miquelle, D. |author8=Stein, A. B. |author9=Watson, A. |author10=Kekule, L. B. |author11=Machado, R. B. |author12=Eizirik, E. |name-list-style=amp |year=2017 |title=Mapping black panthers: Macroecological modeling of melanism in leopards (''Panthera pardus'') |journal=PLOS ONE |volume=12 |issue=4 |pages=e0170378 |doi=10.1371/journal.pone.0170378 |pmid=28379961 |pmc=5381760 |bibcode=2017PLoSO..1270378D |doi-access=free}}</ref>
Frequency of melanism appears to be approximately 11% over the leopard's range. Data on the distribution of leopard populations indicates that melanism occurs in five subspecies in the wild: the Indian leopard, Javan leopard, African leopard, [[Indochinese leopard]] (''P. p. delacouri'') and [[Sri Lankan leopard]] (''P. p. kotiya''). Based on records from camera traps, melanistic leopards occur foremost in [[tropical and subtropical moist broadleaf forests]].<ref name=daSilva17>{{cite journal |author1=Da Silva, L. G. |author2=Kawanishi, K. |first=K. |author3=Henschel P. |author4=Kittle, A. |author5=Sanei, A. |author6=Reebin, A. |author7=Miquelle, D. |author8=Stein, A. B. |author9=Watson, A. |author10=Kekule, L. B. |author11=Machado, R. B. |author12=Eizirik, E. |name-list-style=amp |year=2017 |title=Mapping black panthers: Macroecological modeling of melanism in leopards (''Panthera pardus'') |journal=PLOS ONE |volume=12 |issue=4 |pages=e0170378 |doi=10.1371/journal.pone.0170378 |pmid=28379961 |pmc=5381760 |bibcode=2017PLoSO..1270378D |doi-access=free}}</ref>


Melanism in the leopard is conferred by a [[recessive allele]].<ref>{{cite journal |author1=Robinson, R. |s2cid=5446868 |year=1970 |title=Inheritance of the black form of the leopard ''Panthera pardus'' |journal=Genetica |volume=41 |issue=1 |pages=190–197 |doi=10.1007/BF00958904 |pmid=5480762}}</ref>
Melanism in the leopard is conferred by a [[recessive allele]].<ref>{{cite journal |author1=Robinson, R. |s2cid=5446868 |year=1970 |title=Inheritance of the black form of the leopard ''Panthera pardus'' |journal=Genetica |volume=41 |issue=1 |pages=190–197 |doi=10.1007/BF00958904 |pmid=5480762}}</ref>
It is thought that melanism confers a [[Natural selection#Fitness|selective advantage]] under certain conditions since it is more common in regions of dense forest, where light levels are lower. Preliminary studies also suggest that melanism might be linked to beneficial mutations in the [[immune system]].<ref>{{cite journal |author=Sunquist, F. |date=2007 |title=Malaysian Mystery Leopards |journal=[[National Wildlife Magazine]] |volume=45 |issue=1 |url=https://www.nwf.org/en/Magazines/National-Wildlife/2007/Malasian-Mystery |access-date=6 January 2020 |archive-date=13 August 2022 |archive-url=https://web.archive.org/web/20220813093112/https://www.nwf.org/en/Magazines/National-Wildlife/2007/Malasian-Mystery |url-status=live }}</ref>
It is thought that melanism confers a [[Natural selection#Fitness|selective advantage]] under certain conditions since it is more common in regions of dense forest, where light levels are lower. Preliminary studies also suggest that melanism might be linked to beneficial mutations in the [[immune system]].<ref>{{cite journal |author=Sunquist, F. |date=2007 |title=Malaysian Mystery Leopards |journal=[[National Wildlife Magazine]] |volume=45 |issue=1 |url=https://www.nwf.org/en/Magazines/National-Wildlife/2007/Malasian-Mystery |access-date=6 January 2020 |archive-date=13 August 2022 |archive-url=https://web.archive.org/web/20220813093112/https://www.nwf.org/en/Magazines/National-Wildlife/2007/Malasian-Mystery |url-status=live}}</ref>
The typical spots and rosettes are present but hidden due to the excess melanin.<ref name=Schneider_al12>{{cite journal |author1=Schneider, A. |author2=David, V. A. |author3=Johnson, W. E. |author4=O'Brien, S. J. |author5=Barsh, G. S. |author6=Menotti-Raymond, M. |author7=Eizirik, E. |name-list-style=amp |year=2012 |title=How the leopard hides its spots: ASIP mutations and melanism in wild cats |journal=PLOS ONE |volume=7 |issue=12 |pages=e50386 |doi=10.1371/journal.pone.0050386|pmid=23251368 |pmc=3520955 |bibcode=2012PLoSO...750386S |doi-access=free}}</ref>
The typical spots and rosettes are present but hidden due to the excess melanin.<ref name=Schneider_al12>{{cite journal |author1=Schneider, A. |author2=David, V. A. |author3=Johnson, W. E. |author4=O'Brien, S. J. |author5=Barsh, G. S. |author6=Menotti-Raymond, M. |author7=Eizirik, E. |name-list-style=amp |year=2012 |title=How the leopard hides its spots: ASIP mutations and melanism in wild cats |journal=PLOS ONE |volume=7 |issue=12 |pages=e50386 |doi=10.1371/journal.pone.0050386|pmid=23251368 |pmc=3520955 |bibcode=2012PLoSO...750386S |doi-access=free}}</ref>


The [[Taxonomy (biology)|taxonomic]] status of [[captive animal|captive]] black leopards and the extent of [[Hybridisation (biology)|hybridization]] between the Javan leopard and other [[Leopard#Living subspecies|leopard subspecies]] is uncertain. Therefore, coordinated breeding programs for black leopards do not exist in European and North American zoos.<ref>{{cite journal |author=Gippoliti, S. |author2=Meijaard, E. |name-list-style=amp |year=2007 |title=Taxonomic uniqueness of the Javan Leopard; an opportunity for zoos to save it |journal=Contributions to Zoology |volume=76 |issue=1 |pages=55–57 |url=http://www.repository.naturalis.nl/record/217425 |doi=10.1163/18759866-07601005 |s2cid=55715897 |access-date=4 September 2019 |archive-date=8 October 2019 |archive-url=https://web.archive.org/web/20191008151849/https://www.repository.naturalis.nl/record/217425 |url-status=live }}</ref> Black leopards occupy space needed for breeding endangered leopard subspecies and are not included within the North American [[Species Survival Plan]].<ref>{{cite book |author=Richardson, D. M. |year=2001 |chapter=A simple analysis of leopard (''Panthera pardus'') space within EAZA collections |pages=391–392 |title=EAZA Yearbook 1999/2000 |editor1-last=Hiddinga, B. |editor2-last=Brouwer, K. |publisher=EAZA Executive Office |location=Amsterdam}}</ref><ref>{{cite book |author=Swanson, B. |author2=Fletchall, N. |author3=Shoemaker, A. |name-list-style=amp |year=2003 |title=Felid Taxon Advisory Group North American Regional Collection Plan 2003–2005 |publisher=Disney's Animal Kingdom |location=Bay Lake}}</ref>
The [[Taxonomy (biology)|taxonomic]] status of [[captive animal|captive]] black leopards and the extent of [[Hybridisation (biology)|hybridization]] between the Javan leopard and other [[Leopard#Living subspecies|leopard subspecies]] is uncertain. Therefore, coordinated breeding programs for black leopards do not exist in European and North American zoos.<ref>{{cite journal |author=Gippoliti, S. |author2=Meijaard, E. |name-list-style=amp |year=2007 |title=Taxonomic uniqueness of the Javan Leopard; an opportunity for zoos to save it |journal=Contributions to Zoology |volume=76 |issue=1 |pages=55–57 |url=http://www.repository.naturalis.nl/record/217425 |doi=10.1163/18759866-07601005 |s2cid=55715897 |access-date=4 September 2019 |archive-date=8 October 2019 |archive-url=https://web.archive.org/web/20191008151849/https://www.repository.naturalis.nl/record/217425 |url-status=live }}</ref> Black leopards occupy space needed for breeding endangered leopard subspecies and are not included within the North American [[Species Survival Plan]].<ref>{{cite book |author=Richardson, D. M. |year=2001 |chapter=A simple analysis of leopard (''Panthera pardus'') space within EAZA collections |pages=391–392 |title=EAZA Yearbook 1999/2000 |editor1-last=Hiddinga, B. |editor2-last=Brouwer, K. |publisher=EAZA Executive Office |location=Amsterdam}}</ref><ref>{{cite book |author=Swanson, B. |author2=Fletchall, N. |author3=Shoemaker, A. |name-list-style=amp |year=2003 |title=Felid Taxon Advisory Group North American Regional Collection Plan 2003–2005 |publisher=Disney's Animal Kingdom |location=Bay Lake}}</ref>
A black [[Amur leopard]] (''P. p. orientalis'') was exhibited at the [[San Diego Zoo]] in 2017.<ref>{{cite web |first=D. |last=Parham |title=Black Beauty |date=2017 |url=https://zoonooz.sandiegozoo.org/zoonooz/black-beauty/ |website=Zoonooz |access-date=24 October 2018 |archive-date=24 October 2018 |archive-url=https://web.archive.org/web/20181024152518/https://zoonooz.sandiegozoo.org/zoonooz/black-beauty/ |url-status=live }}</ref>
A black [[Amur leopard]] (''P. p. orientalis'') was exhibited at the [[San Diego Zoo]] in 2017.<ref>{{cite web |first=D. |last=Parham |title=Black Beauty |date=2017 |url=https://zoonooz.sandiegozoo.org/zoonooz/black-beauty/ |website=Zoonooz |access-date=24 October 2018 |archive-date=24 October 2018 |archive-url=https://web.archive.org/web/20181024152518/https://zoonooz.sandiegozoo.org/zoonooz/black-beauty/ |url-status=live}}</ref>


A pseudo-melanistic leopard has a normal background color, but the spots are more densely packed than normal, and merge to obscure the golden-brown background color. Any spots on the flanks and limbs that have not merged into the mass of swirls and stripes are unusually small and discrete, rather than forming rosettes. The face and underparts are paler and dappled, like those of ordinary spotted leopards.<ref>{{cite book |last=Gamble |first=C. |title=Leopards: Natural History & Conservation |year=2004 |publisher=Voyageur Press |location=Stillwater, MN |isbn=978-0896586567}}</ref>
A pseudo-melanistic leopard has a normal background color, but the spots are more densely packed than normal, and merge to obscure the golden-brown background color. Any spots on the flanks and limbs that have not merged into the mass of swirls and stripes are unusually small and discrete, rather than forming rosettes. The face and underparts are paler and dappled, like those of ordinary spotted leopards.<ref>{{cite book |last=Gamble |first=C. |title=Leopards: Natural History & Conservation |year=2004 |publisher=Voyageur Press |location=Stillwater, MN |isbn=978-0896586567}}</ref>
Line 47: Line 47:
In eastern Panama, black jaguars were repeatedly photographed in the [[Mamoní River]] Valley between 2016 and 2018, mostly in [[primary forest]].<ref>{{cite journal |author1=Yacelga, M. |name-list-style=amp |author2=Craighead, K. |year=2019 |title=Melanistic jaguars in Panama |journal=Cat News |issue=70 |pages=39–41 |url=https://www.academia.edu/41977728 |access-date=14 April 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021257/https://www.academia.edu/41977728 |url-status=live }}</ref>
In eastern Panama, black jaguars were repeatedly photographed in the [[Mamoní River]] Valley between 2016 and 2018, mostly in [[primary forest]].<ref>{{cite journal |author1=Yacelga, M. |name-list-style=amp |author2=Craighead, K. |year=2019 |title=Melanistic jaguars in Panama |journal=Cat News |issue=70 |pages=39–41 |url=https://www.academia.edu/41977728 |access-date=14 April 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021257/https://www.academia.edu/41977728 |url-status=live }}</ref>
Melanism in the jaguar is caused by deletions in the [[melanocortin 1 receptor]] gene and conferred by a [[dominant allele]].<ref name=Eizirik_al03>{{cite journal |author1=Eizirik, E. |author2=Yuhki, N. |author3=Johnson, W. E. |author4=Menotti-Raymond, M. |author5=Hannah, S. S. |author6=O'Brien, S. J. |s2cid=19021807 |name-list-style=amp |title=Molecular Genetics and Evolution of Melanism in the Cat Family |journal=Current Biology |year=2003 |volume=13 |issue=5 |pages=448–453 |doi=10.1016/S0960-9822(03)00128-3 |pmid=12620197|doi-access=free }}</ref> Five black jaguars were monitored by researchers in the [[Mamirauá Sustainable Development Reserve]] in the Brazilian State of [[Amazonas (Brazilian state)|Amazonas]] between 2003 and 2018. The researchers studied their behaviour in [[Várzea forest]].<ref>{{Cite web |last=Publicidade |first=W5 |title=Onça-preta é monitorada na Amazônia por pesquisadores do Instituto Mamirauá |url=https://www.mamiraua.org.br/noticias/onca-preta-e-monitorada-na-amazonia-por-pesquisadores-do-instituto-mamiraua |access-date=2022-08-20 |website=Instituto de Desenvolvimento Sustentável Mamirauá |language=pt-br |archive-date=20 August 2022 |archive-url=https://web.archive.org/web/20220820190257/https://www.mamiraua.org.br/noticias/onca-preta-e-monitorada-na-amazonia-por-pesquisadores-do-instituto-mamiraua |url-status=live }}</ref>
Melanism in the jaguar is caused by deletions in the [[melanocortin 1 receptor]] gene and conferred by a [[dominant allele]].<ref name=Eizirik_al03>{{cite journal |author1=Eizirik, E. |author2=Yuhki, N. |author3=Johnson, W. E. |author4=Menotti-Raymond, M. |author5=Hannah, S. S. |author6=O'Brien, S. J. |s2cid=19021807 |name-list-style=amp |title=Molecular Genetics and Evolution of Melanism in the Cat Family |journal=Current Biology |year=2003 |volume=13 |issue=5 |pages=448–453 |doi=10.1016/S0960-9822(03)00128-3 |pmid=12620197|doi-access=free }}</ref> Five black jaguars were monitored by researchers in the [[Mamirauá Sustainable Development Reserve]] in the Brazilian State of [[Amazonas (Brazilian state)|Amazonas]] between 2003 and 2018. The researchers studied their behaviour in [[Várzea forest]].<ref>{{Cite web |last=Publicidade |first=W5 |title=Onça-preta é monitorada na Amazônia por pesquisadores do Instituto Mamirauá |url=https://www.mamiraua.org.br/noticias/onca-preta-e-monitorada-na-amazonia-por-pesquisadores-do-instituto-mamiraua |access-date=2022-08-20 |website=Instituto de Desenvolvimento Sustentável Mamirauá |language=pt-br |archive-date=20 August 2022 |archive-url=https://web.archive.org/web/20220820190257/https://www.mamiraua.org.br/noticias/onca-preta-e-monitorada-na-amazonia-por-pesquisadores-do-instituto-mamiraua |url-status=live }}</ref>

==Cougar==
There is no authenticated case of a truly melanistic [[cougar]]. No specimen has been photographed or killed in the wild, nor has it ever been bred in captivity. Unconfirmed sightings known as the "North American black panther" are currently attributed to errors in species identification by non-experts, and by the [[mimesis|mimetic]] exaggeration of size.<ref>{{cite journal |last1=Holbert|first1=C. |date=2002 |title=Stranded in the Wasteland: Literary Allusion in The Sharpest Sight |journal=Studies in American Indian Literatures |volume=14 |issue=1 |page=3|jstor=20737121 }}</ref>{{better source needed|date=July 2021}}{{clarify|reason=species misidentification of *what*? exaggeration of size of *what*? cougars are bigger than leopards, but jaguars are bigger than cougars. if a black panther were to be in North America, and it wasn't a cougar, twould be a jaguar - and thus bigger thana cougar|date=July 2021}}


==See also==
==See also==

Revision as of 21:17, 26 August 2024

A black panther is the melanistic colour variant of the leopard (Panthera pardus) and the jaguar (Panthera onca). Black panthers of both species have excess black pigments, but their typical rosettes are also present. They have been documented mostly in tropical forests, with black leopards in Africa and Asia, and black jaguars in South America. Melanism is caused by a recessive allele in the leopard, and by a dominant allele in the jaguar.

Leopard

A melanistic leopard in Out of Africa Wildlife Park, Camp Verde, Arizona
Particularly prominent rosettes on a female black leopard at the Rhino and Lion Nature Reserve, Kromdraai

In 1788, Jean-Claude Delamétherie described a black leopard that was kept in the Tower of London and had been brought from Bengal.[1] In 1794, Friedrich Albrecht Anton Meyer proposed the scientific name Felis fusca for this cat, the Indian leopard (P. p. fusca).[2][3] In 1809, Georges Cuvier described a black leopard kept in the Ménagerie du Jardin des plantes that had been brought from Java. Cuvier proposed the name Felis melas, the Javan leopard (P. p. melas).[3][4] By the late 19th century, the occurrence of black and spotted leopard cubs in the same litter had been repeatedly recorded in India. Black leopards were thought to be more common in Travancore and in the hills of southern India than in other parts of the country.[5] Black leopards were also frequently encountered in southern Myanmar.[6] By 1929, the Natural History Museum, London had skins of black leopards collected in South Africa, Nepal, Assam and Kanara in India.[7] Black leopards were thought to be common on the Malay Peninsula and on Java.[8]

A black African leopard (P. p. pardus) was sighted in the alpine zone of Mount Kenya in the winter of 1989–1990.[9] In Kenya's Laikipia County, a black leopard was photographed by a camera trap in 2007; in 2018, a female subadult black leopard was repeatedly recorded together with a spotted leopard about 50 km (31 mi) farther east in a grassland.[10]

In India's Western Ghats, black leopards were sighted and photographed in 2010 and 2012 in the Kas Plateau Reserved Forest, and in Bhadra Wildlife Sanctuary in 2012.[11] In 2015, a dead black leopard was found on a highway near Satara in Maharashtra.[12] In May 2012, a black leopard was photographed at an elevation of 4,300 m (14,100 ft) in Nepal's Kanchenjunga Conservation Area.[13]

At least one black leopard was photographed in mixed deciduous forest in Thailand's Kaeng Krachan National Park during a one-year-long camera trapping survey from 2003 to 2004.[14] In 2009, black leopards were photographed more often than spotted leopards in Kui Buri National Park.[15] Most leopards recorded at 16 sites south of the Kra Isthmus between 1996 and 2009 were black, indicating a near-fixation of melanism in Peninsular Malaysia.[16] In 2019, a black individual was photographed outside a protected area in Jeli District.[17] Both black and spotted leopards were recorded in Gunung Gede Pangrango National Park in West Java between 2005 and 2017.[18]

Frequency of melanism appears to be approximately 11% over the leopard's range. Data on the distribution of leopard populations indicates that melanism occurs in five subspecies in the wild: the Indian leopard, Javan leopard, African leopard, Indochinese leopard (P. p. delacouri) and Sri Lankan leopard (P. p. kotiya). Based on records from camera traps, melanistic leopards occur foremost in tropical and subtropical moist broadleaf forests.[19]

Melanism in the leopard is conferred by a recessive allele.[20] It is thought that melanism confers a selective advantage under certain conditions since it is more common in regions of dense forest, where light levels are lower. Preliminary studies also suggest that melanism might be linked to beneficial mutations in the immune system.[21] The typical spots and rosettes are present but hidden due to the excess melanin.[22]

The taxonomic status of captive black leopards and the extent of hybridization between the Javan leopard and other leopard subspecies is uncertain. Therefore, coordinated breeding programs for black leopards do not exist in European and North American zoos.[23] Black leopards occupy space needed for breeding endangered leopard subspecies and are not included within the North American Species Survival Plan.[24][25] A black Amur leopard (P. p. orientalis) was exhibited at the San Diego Zoo in 2017.[26]

A pseudo-melanistic leopard has a normal background color, but the spots are more densely packed than normal, and merge to obscure the golden-brown background color. Any spots on the flanks and limbs that have not merged into the mass of swirls and stripes are unusually small and discrete, rather than forming rosettes. The face and underparts are paler and dappled, like those of ordinary spotted leopards.[27]

Jaguar

A melanistic jaguar
A melanistic jaguar at the Henry Doorly Zoo

In 1801, Félix de Azara described a black jaguar observed by local people near the Paraná River in Paraguay.[28] In 2004, a female black jaguar was recorded in Mexico's Sierra Madre Occidental.[29] In 2009, a black jaguar was photographed by a camera trap for the first time in Costa Rica's Alberto Manuel Brenes Biological Reserve.[30] In Barbilla National Park, black jaguars were recorded in 2013.[31] In the mountains of the Cordillera de Talamanca, 104 records of jaguars were obtained between 2010 and 2019; 26 of them showed melanistic jaguars.[32] In eastern Panama, black jaguars were repeatedly photographed in the Mamoní River Valley between 2016 and 2018, mostly in primary forest.[33] Melanism in the jaguar is caused by deletions in the melanocortin 1 receptor gene and conferred by a dominant allele.[34] Five black jaguars were monitored by researchers in the Mamirauá Sustainable Development Reserve in the Brazilian State of Amazonas between 2003 and 2018. The researchers studied their behaviour in Várzea forest.[35]

See also

References

  1. ^ Delamétherie, J.-C. (1788). "Description d'une Panthère noire" [Description of a black Panther]. Observations et Mémoires sur la Physique, sur l'Histoire Naturelle et sur les Arts et Métiers, etc. (in French). 33: 45.
  2. ^ Meyer, F. A. A. (1794). "Über de la Metheries schwarzen Panther [About de la Metheries black Panther]". Zoologische Annalen (in German). Vol. Erster Band. Weimar: Im Verlage des Industrie-Comptoirs. pp. 394–396. Archived from the original on 5 May 2024. Retrieved 14 January 2020.
  3. ^ a b Pocock, R. I. (1930). "The Panthers and Ounces of Asia". Journal of the Bombay Natural History Society. 34 (1): 307–336.
  4. ^ Cuvier, G. (1809). "Recherches sur les espėces vivantes de grands chats, pour servir de preuves et d'éclaircissement au chapitre sur les carnassiers fossils". Annales du Muséum National d'Histoire Naturelle. XIV: 136–164.
  5. ^ Blanford, W. T. (1888). "Felis pardus. The Leopard or Panther". The Fauna of British India, including Ceylon and Burma. Mammalia: Volume 1. London: Taylor and Francis. pp. 67–71.
  6. ^ Mason, F. (1882). "F. pardus, L. The leopard". Burma, its people and productions; or, Notes on the fauna, flora, and minerals of Tenasserim, Pegu, and Burma. Vol. 1. Geology, mineralogy, and zoology (Rewritten and enlarged by W. Theobald ed.). Hertford: Chief Commissioner of British Burma. p. 472.
  7. ^ Pocock, R. I. (1929). "Black panthers – an inquiry". Journal of the Bombay Natural History Society. 33 (3): 693–694.
  8. ^ Pocock, R. I. (1930). "The Panthers and Ounces of Asia". Journal of the Bombay Natural History Society. 34 (1): 65–82.
  9. ^ Young, T. P. & Evans, M. E. (1993). "Alpine vertebrates of Mount Kenya, with particular notes on the rock hyrax". Journal of the East African Natural History Society. 82 (202): 54–79.
  10. ^ Pilfold, N. W.; Letoluai, A.; Ruppert, K.; Glikman, J. A.; Stacy-Dawes, J.; O'Connor, D. & Owen, M. (2019). "Confirmation of black leopard (Panthera pardus pardus) living in Laikipia County, Kenya". African Journal of Ecology. 57 (2): 270–273. doi:10.1111/aje.12586. S2CID 92543492.
  11. ^ Sayyed, A. & Mahabal, A. (2013). "Records of the melanistic Leopard Panthera pardus (Linnaeus) from Western Ghats area of Maharashtra and Karnataka, India". Journal of the Bombay Natural History Society. 110 (2): 151.
  12. ^ Sayyed, A. & Mahabal, A. (2015). "Second record of melanistic leopard Panthera pardus (Linnaeus) from Satara, Maharashtra: a case of roadkill". Zoo's Print. 30 (5): 29.
  13. ^ Thapa, K.; Pradhan, N. M. B.; Barker, J.; Dahal, M.; Bhandari, A. R.; Gurung, G. S.; Rai, D. P.; Thapa, G. J.; Shrestha, S.; Singh, G. R. (2013). "High elevation record of a leopard cat in the Kangchenjunga Conservation Area, Nepal". Cat News (58): 26–27.
  14. ^ Ngoprasert, D.; Lynam, A. J. & Gale, G. A. (2007). "Human disturbance affects habitat use and behaviour of Asiatic leopard "Panthera pardus" in Kaeng Krachan National Park, Thailand". Oryx. 41 (3): 343–351. doi:10.1017/S0030605307001102.
  15. ^ Steinmetz, R.; Seuaturien, N.; Chutipong, W. & Poonnil, B. (2009). The ecology and conservation of tigers and their prey in Kuiburi National Park, Thailand (PDF) (Report). Bangkok: WWF Thailand, Department of National Parks, Wildlife and Plant Conservation. Archived (PDF) from the original on 2 May 2021.
  16. ^ Kawanishi, K.; Sunquist, M. E.; Eizirik, E.; Lynam, A. J.; Ngoprasert, D.; Shahruddin, W. N. W.; Rayan, D. M.; Sharma, D. S. K. & Steinmetz, R. (2010). "Near fixation of melanism in panthers of the Malay Peninsula". Journal of Zoology. 282 (3): 201–206. doi:10.1111/j.1469-7998.2010.00731.x.
  17. ^ Hambali, K.; Fazli, N. F. M.; Amir, A.; Fauzi, N.; Hassin, N. H.; Abas, M. A.; Karim, M. F. A. & Sow, A. Y. (2021). "The discovery of a melanistc Leopard Panthera pardus delacouri (Linnaeus, 1758) (Mammalia: Carnivora: Felidae) at Bukit Kudung in Jeli, Kelantan, Peninsular Malaysia: conservation and ecotourism". Journal of Threatened Taxa. 13 (1): 17513–17516. doi:10.11609/jott.6060.13.1.17513-17516.
  18. ^ Ario, A.; Supian; Hidayat, E.; Hidayatullah, R.; Rustiadi, A.; Gunawan, A.; Triprajawan, T.; Sopian, I.; Zatnika, R. R.; Yusup, D. M.; Hindrayani, W.; Mulyanto, A. & Iskandar, D. (2018). "Population dynamics and ecology of Javan leopard, Panthera pardus melas, in Gunung Gede Pangrango National Park, West Java". Journal of Indonesian Natural History. 6 (1): 6–13. Archived from the original on 27 September 2021. Retrieved 15 May 2021.
  19. ^ Da Silva, L. G., K.; Kawanishi, K.; Henschel P.; Kittle, A.; Sanei, A.; Reebin, A.; Miquelle, D.; Stein, A. B.; Watson, A.; Kekule, L. B.; Machado, R. B. & Eizirik, E. (2017). "Mapping black panthers: Macroecological modeling of melanism in leopards (Panthera pardus)". PLOS ONE. 12 (4): e0170378. Bibcode:2017PLoSO..1270378D. doi:10.1371/journal.pone.0170378. PMC 5381760. PMID 28379961.
  20. ^ Robinson, R. (1970). "Inheritance of the black form of the leopard Panthera pardus". Genetica. 41 (1): 190–197. doi:10.1007/BF00958904. PMID 5480762. S2CID 5446868.
  21. ^ Sunquist, F. (2007). "Malaysian Mystery Leopards". National Wildlife Magazine. 45 (1). Archived from the original on 13 August 2022. Retrieved 6 January 2020.
  22. ^ Schneider, A.; David, V. A.; Johnson, W. E.; O'Brien, S. J.; Barsh, G. S.; Menotti-Raymond, M. & Eizirik, E. (2012). "How the leopard hides its spots: ASIP mutations and melanism in wild cats". PLOS ONE. 7 (12): e50386. Bibcode:2012PLoSO...750386S. doi:10.1371/journal.pone.0050386. PMC 3520955. PMID 23251368.
  23. ^ Gippoliti, S. & Meijaard, E. (2007). "Taxonomic uniqueness of the Javan Leopard; an opportunity for zoos to save it". Contributions to Zoology. 76 (1): 55–57. doi:10.1163/18759866-07601005. S2CID 55715897. Archived from the original on 8 October 2019. Retrieved 4 September 2019.
  24. ^ Richardson, D. M. (2001). "A simple analysis of leopard (Panthera pardus) space within EAZA collections". In Hiddinga, B.; Brouwer, K. (eds.). EAZA Yearbook 1999/2000. Amsterdam: EAZA Executive Office. pp. 391–392.
  25. ^ Swanson, B.; Fletchall, N. & Shoemaker, A. (2003). Felid Taxon Advisory Group North American Regional Collection Plan 2003–2005. Bay Lake: Disney's Animal Kingdom.
  26. ^ Parham, D. (2017). "Black Beauty". Zoonooz. Archived from the original on 24 October 2018. Retrieved 24 October 2018.
  27. ^ Gamble, C. (2004). Leopards: Natural History & Conservation. Stillwater, MN: Voyageur Press. ISBN 978-0896586567.
  28. ^ Azara, F. de (1801). "L'Yagouarété". Essais sur l'histoire naturelle des quadrupedes de la province du Paraguay. Vol. 1. Paris: Charles Pougens. pp. 114–132.
  29. ^ Dinets, V. & Polechla, P. J. (2005). "First documentation of melanism in the jaguar (Panthera onca) from northern Mexico". Cat News. 42: 18. Archived from the original on 26 September 2006.
  30. ^ Núñez, M. C. & Jiménez, E. C. (2009). "A new record of a black jaguar, Panthera onca (Carnivora: Felidae) in Costa Rica". Brenesia. 71: 67–68. Archived from the original on 29 November 2021. Retrieved 12 April 2021.
  31. ^ Sáenz-Bolaños, C.; Montalvo, V.; Fuller, T. K. & Carrillo, E. (2015). "Records of black jaguars at Parque Nacional Barbilla, Costa Rica". Cat News (62): 38–39.
  32. ^ Mooring, M. S.; Eppert, A. A. & Botts, R. T. (2020). "Natural Selection of Melanism in Costa Rican Jaguar and Oncilla: A Test of Gloger's Rule and the Temporal Segregation Hypothesis". Tropical Conservation Science. 13: 1–15. doi:10.1177/1940082920910364.
  33. ^ Yacelga, M. & Craighead, K. (2019). "Melanistic jaguars in Panama". Cat News (70): 39–41. Archived from the original on 29 November 2021. Retrieved 14 April 2021.
  34. ^ Eizirik, E.; Yuhki, N.; Johnson, W. E.; Menotti-Raymond, M.; Hannah, S. S. & O'Brien, S. J. (2003). "Molecular Genetics and Evolution of Melanism in the Cat Family". Current Biology. 13 (5): 448–453. doi:10.1016/S0960-9822(03)00128-3. PMID 12620197. S2CID 19021807.
  35. ^ Publicidade, W5. "Onça-preta é monitorada na Amazônia por pesquisadores do Instituto Mamirauá". Instituto de Desenvolvimento Sustentável Mamirauá (in Brazilian Portuguese). Archived from the original on 20 August 2022. Retrieved 20 August 2022.{{cite web}}: CS1 maint: numeric names: authors list (link)