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=== ''M. leprae'' and ''M. lepromatosis'' === [[File:Mycobacterium leprae in Magnification of 2000X.jpg|thumb|upright=1.3|''M. leprae'', one of the causative agents of leprosy: As an [[acid-fast]] bacterium, ''M. leprae'' appears red when a [[Ziehl–Neelsen stain]] is used]] ''[[Mycobacterium leprae|Mycobacterioum leprae]]'' and ''[[Mycobacterium lepromatosis]]'' are the [[Mycobacterium|mycobacteria]] that cause leprosy.<ref name=Rei2019 /> ''M. lepromatosis'' is a relatively newly identified mycobacterium isolated from a fatal case of [[diffuse lepromatous leprosy]] in 2008.<ref name="New2008">{{cite web|url=https://www.sciencedaily.com/releases/2008/11/081124141047.htm|title=New Leprosy Bacterium: Scientists Use Genetic Fingerprint To Nail 'Killing Organism'|date=28 November 2008|work=ScienceDaily|archive-url=https://web.archive.org/web/20100313064458/https://www.sciencedaily.com/releases/2008/11/081124141047.htm|archive-date=13 March 2010|url-status=live|access-date=31 January 2010}}</ref><ref name="Sherris">{{cite book | veditors = Ryan KU, Ray CJ |title=Sherris Medical Microbiology |url=https://archive.org/details/sherrismedicalmi00ryan |url-access=limited |edition= 4th |pages=[https://archive.org/details/sherrismedicalmi00ryan/page/n468 451]–53 |publisher=McGraw Hill |year=2004 |isbn=978-0-8385-8529-0 |oclc=61405904}}</ref> ''M. lepromatosis'' is indistinguishable clinically from ''M. leprae''.<ref>{{Cite web|url=https://internationaltextbookofleprosy.org/chapter/genomic-insights-biology-and-evolution-leprosy-bacilli|title=Genomics Insights into the Biology and Evolution of Leprosy Bacilli|date=11 February 2016|website=International Textbook of Leprosy|access-date=11 February 2019|archive-date=12 February 2019|archive-url=https://web.archive.org/web/20190212070703/https://internationaltextbookofleprosy.org/chapter/genomic-insights-biology-and-evolution-leprosy-bacilli|url-status=live}}</ref> ''M. leprae'' is an [[Aerobic organism|aerobic]], rod-shaped, acid-fast [[Bacteria|bacterium]] with a waxy [[cell envelope]] characteristic of the genus ''[[Mycobacterium]]''.<ref name="Baron">{{cite book |author=McMurray DN |chapter=Mycobacteria and Nocardia |title=Baron's Medical Microbiology |editor1=Baron S |display-editors=etal |edition= 4th |publisher=Univ of Texas Medical Branch |year=1996 |chapter-url=https://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mmed.section.1833 |isbn=978-0-9631172-1-2 |oclc=33838234 |url-status=live |archive-url=https://web.archive.org/web/20090212202626/http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mmed.section.1833 |archive-date=12 February 2009 }}</ref> ''M. leprae'' and ''M. lepromatosis'' are [[Intracellular parasite#Obligate|obligate intracellular pathogens]] and cannot grow or be [[Cell culture|cultured]] outside of host tissues.<ref name="New2008" /><ref name="Bhattacharya-2002">{{cite journal | vauthors = Bhattacharya S, Vijayalakshmi N, Parija SC | title = Uncultivable bacteria: implications and recent trends towards identification | journal = Indian Journal of Medical Microbiology | volume = 20 | issue = 4 | pages = 174–177 | date = October 2002 | pmid = 17657065 | doi = 10.1016/S0255-0857(21)03184-4 | doi-access = free }}</ref> However, they can be grown using research animals such as mice and [[armadillo]]s.<ref>{{Cite web|url=https://www.who.int/lep/in_vitro/en/|archive-url=https://web.archive.org/web/20200809141738/https://www.who.int/lep/in_vitro/en/|url-status=dead|archive-date=9 August 2020|title=WHO {{!}} Microbiology: culture in vitro|publisher=[[World Health Organization]] (WHO)|access-date=22 July 2019}}</ref><ref>{{Cite web|url=https://internationaltextbookofleprosy.org/chapter/armadillos|title=The Armadillo Model for Leprosy|date=11 February 2016|website=International Textbook of Leprosy|access-date=22 July 2019|archive-date=22 July 2019|archive-url=https://web.archive.org/web/20190722225203/https://internationaltextbookofleprosy.org/chapter/armadillos|url-status=live}}</ref> Naturally occurring infections have been reported in nonhuman primates (including the African [[Common chimpanzee|chimpanzee]], the [[sooty mangabey]], and the [[cynomolgus]] macaque), armadillos,<ref>Loughry WJ, Truman RW, McDonough CM, Tilak MK, Garnier S, et al. (2009) "Is leprosy spreading among nine-banded armadillos in the southeastern United States?" ''J Wildl Dis'' 45: 144–52.</ref> and [[red squirrel]]s.<ref name="Meredith2004">{{cite journal | vauthors = Meredith A, Del Pozo J, Smith S, Milne E, Stevenson K, McLuckie J | title = Leprosy in red squirrels in Scotland | journal = The Veterinary Record | volume = 175 | issue = 11 | pages = 285–286 | date = September 2014 | pmid = 25234460 | doi = 10.1136/vr.g5680 | s2cid = 207046489 }}</ref> [[Multilocus sequence typing]] of the armadillo ''M. leprae'' strains suggests that they were of human origin for at most a few hundred years.<ref>Monot M, Honoré N, Garnier T, Araoz R, Coppee JY, et al. (2005). "On the origin of leprosy". ''Science'' 308: 1040–42.</ref> Thus, it is suspected that armadillos first acquired the organism incidentally from early European explorers of the Americas.<ref name="Han-2014" /> This incidental transmission was sustained in the armadillo population, and it may be transmitted back to humans, making leprosy a [[zoonotic disease]] (spread between humans and animals).<ref name="Han-2014">{{cite journal | vauthors = Han XY, Silva FJ | title = On the age of leprosy | journal = PLOS Neglected Tropical Diseases | volume = 8 | issue = 2 | pages = e2544 | date = February 2014 | pmid = 24551248 | pmc = 3923669 | doi = 10.1371/journal.pntd.0002544 | doi-access = free }}</ref> Red squirrels (''Sciurus vulgaris''), a threatened species in Great Britain, were found to carry leprosy in November 2016.<ref>[https://www.science.org/doi/10.1126/science.aah3783 "Red squirrels in the British Isles are infected with leprosy bacilli"] {{Webarchive|url=https://web.archive.org/web/20220612182108/https://www.science.org/doi/10.1126/science.aah3783|date=12 June 2022}}, Dr. Andrej Benjak, Prof Anna Meredith and others. ''Science'', 11 November 2016. [https://www.science.org/doi/10.1126/science.aah3783] . Retrieved 11 November 2016.</ref> It has been suggested that the trade in red squirrel fur, highly prized in the medieval period and intensively traded, may have been responsible for the leprosy epidemic in medieval Europe.<ref name="ScienceDaily" /> A pre-[[Norman England|Norman]] era skull excavated in [[Hoxne|Hoxne, Suffolk]], in 2017 was found to carry DNA from a strain of ''M. leprae'' which closely matched the strain carried by modern red squirrels on [[Brownsea Island]].<ref name="ScienceDaily">{{cite web|url=https://www.sciencedaily.com/releases/2017/10/171025103109.htm|title=Could squirrel fur trade have contributed to England's medieval leprosy outbreak?|website=ScienceDaily|access-date=21 November 2018|archive-date=22 November 2018|archive-url=https://web.archive.org/web/20181122005829/https://www.sciencedaily.com/releases/2017/10/171025103109.htm|url-status=live}}</ref><ref>{{cite journal | vauthors = Inskip S, Taylor GM, Anderson S, Stewart G | title = Leprosy in pre-Norman Suffolk, UK: biomolecular and geochemical analysis of the woman from Hoxne | journal = Journal of Medical Microbiology | volume = 66 | issue = 11 | pages = 1640–1649 | date = November 2017 | pmid = 28984227 | doi = 10.1099/jmm.0.000606 | s2cid = 33997231 | doi-access = free }}</ref>
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