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Cavalier-Smith 2020</small> * "Izemaplasma" <small>(sic) Skennerton et al. 2016</small> * "Paramycetes" <small>Sabin 1941</small> }} '''Mollicutes''' is a class of [[bacteria]]<ref>[http://www.britannica.com/EBchecked/topic/388373/Mollicutes] "Mollicutes", from Encyclopædia Britannica Online. Retrieved 15 Oct. 2011.</ref> distinguished by the absence of a [[cell wall]]. The word "Mollicutes" is derived from the Latin ''{{wikt-lang|la|mollis}}'' (meaning "soft" or "pliable"), and ''{{wikt-lang|la|cutis}}'' (meaning "skin"). Individuals are very small, typically only 0.2–0.3 [[micrometre|μm]] (200–300 [[nanometer|nm]]) in size and have a very small [[genome size]]. They vary in form, although most have [[sterol]]s that make the cell membrane somewhat more rigid. Many are able to move about through [[bacterial gliding|gliding]], but members of the genus ''[[Spiroplasma]]'' are helical and move by twisting. The best-known genus in the Mollicutes is ''[[Mycoplasma]]''. Colonies show the typical "fried-egg" appearance. Mollicutes are parasites of various animals and plants, living on or in the host's cells. Many cause diseases in humans, attaching to cells in the respiratory or urogenital tracts, particularly species of ''Mycoplasma'' and ''[[Ureaplasma urealyticum|Ureaplasma]]''. [[Phytoplasma]] and ''Spiroplasma'' are plant pathogens associated with insect vectors. Whereas formerly the [[trivial name]] "mycoplasma" (plural: "mycoplasmas") has commonly denoted any member of the class Mollicutes, it now in scientific classification refers exclusively to a member of the genus ''Mycoplasma''. ==Origin and development to parasitic life== Analysis of the genomes of Mollicutes ("mycoplasmas") gives solid support for the hypothesis that Mollicutes have developed from Gram-positive bacteria by a process of reductive evolution. By adopting a parasitic mode of life with use of nutrients from their hosts, many Mollicutes were able to reduce their genetic material considerably. On the other hand, they lost the genes for many assimilative processes. Thus, ''Mycoplasma'' possibly became the smallest self-replicating organism in nature. ''[[Mycoplasma genitalium]]'', with 580,000 base pairs, has an especially small genome size. Some phytoplasmas also have a very small genome size. The genera with the smallest genome are considered to be phylogenetically the most "recent" mollicutes. To maintain their parasitic mode of life the Mollicutes have developed rather sophisticated mechanisms to colonize their hosts and resist the host immune system.<ref>Shmuel Razin, David Yogev and Yehudith Naot [http://mmbr.asm.org/cgi/content/full/62/4/1094 ''Molecular Biology and Pathogenicity of Mycoplasmas'']. Micr. and Molec. Biology Reviews, December '''1998''', pp. 1094–1156, Vol. 62, No. 4</ref> ===Proliferation=== Mollicutes (formerly also called pleuropneumonia-like organisms, PPLO) as well as [[L-form bacteria]] (formerly also called L-phase bacteria), both lacking cell walls, do not proliferate by binary fission but by a budding mechanism. In 1954, this mode of proliferation has been shown by continual observations of live cells using phase-contrast microscopy.<ref>{{Cite journal |last1=Kandler |first1=Gertraud |last2=Kandler |first2=Otto |author-link2=Otto Kandler |date=1954 |others=(Article in English available) |title=Untersuchungen über die Morphologie und die Vermehrung der pleuropneumonie-ähnlichen Organismen und der L-Phase der Bakterien. I. Lichtmikroskopische Untersuchungen |trans-title=Studies on morphology and multiplication of pleuropneumonia-like organisms and on bacterial L-phase, I. Light microscopy (now mycoplasmas and L-form bacteria) |url=https://badw.de/fileadmin/members/K/1501/1954-Kandler_Kandler-PPLO-I-English2015-03-11.pdf |journal=Archiv für Mikrobiologie |language=German |volume=21 |issue=2 |pages=178–201 |doi=10.1007/BF01816378 |pmid=14350641|s2cid=21257985 }}</ref> Previously, Mollicutes were sometimes considered stable L-form bacteria, but phylogenetic analysis has identified them as bacteria that have lost their cell walls in the course of evolution.<ref name="woese 1980" /> ==History of the classification== The classification of the Mollicutes has always been difficult. The individuals are tiny, and being parasites, they have to be cultivated on special media. Until now, many species could not be isolated at all. In the beginning, whether they were fungi, viruses, or bacteria was not clear. Also, the resemblance to [[L-form bacteria|L-forms]] was confusing. At first, all members of the class Mollicutes were generally named "mycoplasma" or pleuropneumonia-like organism (PPLO). Mollicutes other than some members of genus ''Mycoplasma'' were still unidentified. The first species of ''Mycoplasma''/Mollicutes, that could be isolated was ''[[Mycoplasma mycoides]]''. This bacterium was [[Microbiological culture|cultivated]] by Nocard and Roux in 1898.<ref name=Hayflick>{{cite journal|author=Hayflick L. & Chanock, R.M.|title=Mycoplasma Species of Man|journal=Bacteriological Reviews|volume=29|issue=2|pages=185–221|year=1965|doi=10.1128/MMBR.29.2.185-221.1965|pmid=14304038 |pmc=441270 |url = http://mmbr.asm.org/cgi/reprint/29/2/185.pdf|doi-access=free}}</ref> In 1956, D.G. Edward and E.A. Freundt made a first proposal for classifying and naming PPLOs. They left undecided, however, whether they belong to the bacteria ([[prokaryote]]s, in 1956 called "Schizomycetes") or to the [[eukaryote]]s. As [[type species]] (name-giving species) of the PPLOs/mycoplasmas, Edward and Freundt proposed ''Mycoplasma mycoides'', being the causative organism of bovine pleuropneumonia and referring to the pleuropneumonia-like organisms. Until then, ''Mycoplasma mycoides'' was known as ''Asterococcus mycoides'', but later that name was not recognized as valid. In their publication of 1956, they described 15 species of ''Mycoplasma''.<ref>{{cite journal|author=Edward DG, Freundt EA|date=February 1956|title=The classification and nomenclature of organisms of the pleuropneumonia group|journal=J. Gen. Microbiol.|volume=14|issue=1|pages=197–207|doi=10.1099/00221287-14-1-197|pmid=13306904|doi-access=free}}pdf [http://mic.sgmjournals.org/cgi/reprint/14/1/197.pdf]</ref> In 1967 the class Mollicutes, containing the order [[Mycoplasmatales]], was proposed by the Subcommittee on Taxonomy of the Mycoplasmata.<ref name="Edward">{{cite journal|last=Edward|first=D.G.|author2=F.A. Freundt|date=July 1967|title=Proposal for Mollicutes as name of the class established for the order Mycoplasmatales|journal=Int J Syst Bacteriol|volume=17|issue=3|pages=267–268|doi=10.1099/00207713-17-3-267|doi-access=free}}</ref> Now, the name ''[[Mycoplasma]]'' should exclusively be used for members of the '''genus''' ''Mycoplasma'', rather than the use as a trivial name for any Mollicute. As the trivial name has been used in literature for a long time, this is yet not always the case. ===Three divisions of the Bacteria=== {{main|Bacterial taxonomy}} Traditionally, the taxonomy of bacteria was based on similarities and differences in [[Morphology (biology)|morphology]] (Linnaean taxonomy). In 1962, R.G.E. Murray proposed to divide the "kingdom" (now [[Domain (biology)|domain]]<ref>{{Cite journal |last1=Woese |first1=Carl R. |author-link=Carl Woese |last2=Kandler |first2=Otto |author-link2=Otto Kandler |last3=Wheelis |first3=Mark L. |date=1990 |title=Towards a natural system of organisms: proposal for the domains Archaea, Bacteria, and Eucarya |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=87 |issue=12 |pages=4576–9 |bibcode=1990PNAS...87.4576W |doi=10.1073/pnas.87.12.4576 |pmc=54159 |pmid=2112744|doi-access=free }}</ref>) Bacteria into three divisions (= phyla) on the basis of the cell wall types: # [[Gram-negative bacteria|Gram-negative]] [[Gracilicutes]], with a thin cell wall and little [[peptidoglycan]]; # [[Gram-positive bacteria|Gram-positive]] [[Firmicutes]], with a thicker cell wall and more peptidoglycan (the name was later changed in "Firm'''i'''cutes"), and # the "Mollicutes", without a cell wall.<ref>N.E. Gibbons & R.G.E. Murray [http://ijs.sgmjournals.org/cgi/reprint/28/1/1.pdf ''Proposals Concerning the Higher Taxa of Bacteria'']{{Dead link|date=April 2020 |bot=InternetArchiveBot |fix-attempted=yes }} Int J Syst Bacteriol Vol. 28 (1) Jan. 1978, pp. 1–6.</ref> ==Phylogeny== [[File:Phylogenetic position of Mollicutes among bacteria.jpg|right|thumb|300px| Phylogenetic position of Mollicutes among bacteria, using 16S rRNA sequences.<ref name="Oshima2013">{{Cite journal| pmid = 23966988| pmc = 3743221| year = 2013| last1 = Oshima| first1 = K| title = Genomic and evolutionary aspects of phytoplasmas| journal = Frontiers in Microbiology| volume = 4| pages = 230| last2 = Maejima| first2 = K| last3 = Namba| first3 = S| doi = 10.3389/fmicb.2013.00230| doi-access = free}}</ref>]] For classification and nomenclature of Mollicutes, there are special rules, which are maintained by the ''International Committee on Systematics of Prokaryotes (ICSP) Subcommittee on the Taxonomy of Mollicutes'' (formerly the ''International Committee on Systematic Bacteriology (ICSB) Subcommittee on taxonomy of Mycoplasmatales'').<ref name="standards">[http://ijs.sgmjournals.org/cgi/content/full/57/11/2703 ''Revised minimal standards for description of new species of the class Mollicutes (division Tenericutes)''] {{Webarchive|url=https://web.archive.org/web/20100710160820/http://ijs.sgmjournals.org/cgi/content/full/57/11/2703 |date=2010-07-10 }} PDF [http://ijs.sgmjournals.org/cgi/reprint/45/3/605.pdf]{{Dead link|date=April 2020 |bot=InternetArchiveBot |fix-attempted=yes }}; Daniel R. Brown, Robert F. Whitcomb and Janet M. Bradbury (2007) ''Int J Syst Evol Microbiol'' 57 2703–2719; {{doi|10.1099/ijs.0.64722-0}}</ref> Traditionally, Mollicutes [[taxonomy (biology)|taxonomy]] has been based on [[serology]] and [[phenotype|phenotypic characteristics]]. However, most modern classifications are based on DNA or RNA sequences, especially [[16S ribosomal RNA|16S rRNA]] sequences<ref name="woese 1980" /> (see Figure). ===The phylum for Mollicutes=== The results of Mollicutes [[phylogenetics|phylogenetic analyses]] have been controversial. Some taxonomists place them in Bacillota, others in Mycoplasmatota. [[Carl Woese|Woese]] et al. suggested that the Mollicutes might have been derived from different branches of bacteria. They concluded, that the Mollicutes are not a phylogenetically coherent group and therefore do not form a distinct higher level taxon. Instead, they cluster within Gram-positive bacteria of the phylum Bacillota.<ref name="woese 1980">C.R. Woese, J. Maniloff and L.B. Zablen [http://www.pnas.org/content/77/1/494.full.pdf ''Phylogenetic analysis of the mycoplasmas''] Proc. Natl. Acad. Sci. USA Vol. 77, No. 1, pp. 494–498, January 1980</ref> The results of [[Molecular phylogenetics|molecular phylogenetic analyses]] have been partly dependent on the chosen molecular marker, like [[Ribosomal RNA|rRNA]], [[elongation factor]] or another protein.<ref name="ludwig2005">Ludwig, W. and Schleifer, K.H. [https://books.google.com/books?hl=nl&lr=&id=LA8JHPM3-zsC&pg=PA70 ''Molecular phylogeny of bacteria based on comparative sequence analysis of conserved genes''] Microbial phylogeny and evolution, 2005, p. 70–98.</ref> Phylogenetic trees based on [[phosphoglycerate kinase]] (Pgk) amino acid sequences' indicated a monophyletic origin for the Mollicutes within the Bacillota.<ref name="Wolf">{{cite journal|last1=Wolf|first1=Matthias|year=2004|title=Phylogeny of Firmicutes with special reference to Mycoplasma (Mollicutes) as inferred from phosphoglycerate kinase amino acid sequence data|journal=Int J Syst Evol Microbiol|volume=54|pages=871–875|doi=10.1099/ijs.0.02868-0|pmid=15143038|last2=Müller|first2=T|last3=Dandekar|first3=T|last4=Pollack|first4=JD|issue=Pt 3|display-authors=etal|doi-access=free}} DOI 10.1099/ijs.0.02868-0</ref> An early edition of ''[[Bergey's Manual of Systematic Bacteriology]]'' placed class Mollicutes within phylum [[Bacillota]],<ref name="bergeys">[http://www.bergeys.org/outlines/bergeysoutline_5_2004.pdf Taxonomic Outline of the Prokaryotes. Bergey's Manual of Systematic Bacteriology.] {{webarchive|url=https://web.archive.org/web/20091229160912/http://www.bergeys.org/outlines/bergeysoutline_5_2004.pdf |date=2009-12-29 }} Garrity, Bell & Lilburn; Second Edition, Release 5.0 (2004); pp. 140–204. {{doi|10.1007/bergeysoutline200405}}.</ref><ref>K. Johansson and B. Pettersson, ''Taxonomy of Mollicutes'' [https://books.google.com/books?id=WuMVqDapIbEC&pg=PA10 books.google.com] (2002)</ref> whereas in the announced 2nd edition, they are moved to a separate phylum Tenericutes (later renamed [[Mycoplasmatota]]).<ref name="taxbrowser2">{{cite web|url = https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=544448|title = Tenericutes|access-date = 2008-09-11|work = Taxonomy Browser|publisher = NCBI}}</ref><ref name="Ludwig2009">Wolfgang Ludwig, Karl-Heinz Schleifer and William B. Whitman (In press, release in '''2009'''). "Revised road map to the phylum Firmicutes". ''In'' P. De Vos ''et al.'' (eds.) ''Bergey's Manual of Systematic Bacteriology'', 2nd ed., vol. 3 (The Firmicutes). Springer-Verlag, New York. {{ISBN|0-387-95041-9}}; [http://www.bergeys.org/outlines/Bergeys_Vol_3_Outline.pdf PDF] {{webarchive|url=https://web.archive.org/web/20100921022410/http://bergeys.org/outlines/Bergeys_Vol_3_Outline.pdf |date=2010-09-21 }}</ref><ref>[[cell wall|''Validation of the Publication of New Names and New Combinations ...'']] Int. J. Syst. Bacteriol.; July 1984, pp. 355–357</ref> The change is motivated by "their unique phenotypic properties, in particular the lack of rigid cell walls, and the general low support by alternative markers".<ref name="ludwig2005"/> In the ''Taxonomic Outline of Bacteria and Archaea'' (TOBA Release 7.7), March 2007, the Mollicutes are a class in the phylum Bacillota.<ref name="TOBA">Garrity, et al. [http://www.taxonomicoutline.org/ ''The Taxonomic Outline of Bacteria and Archaea''] {{Webarchive|url=https://web.archive.org/web/20130502073028/http://www.taxonomicoutline.org/ |date=2013-05-02 }} (TOBA Release 7.7); March 2007. {{cite journal |url=http://www.taxonomicoutline.org/index.php/toba/article/view/185/217.pdf |title=Taxonomic Outline of Bacteria and Archaea |journal=Taxonomic Outline of Bacteria and Archaea |access-date=2009-08-13 |url-status=dead |archive-url=https://web.archive.org/web/20111007150532/http://www.taxonomicoutline.org/index.php/toba/article/view/185/217.pdf |archive-date=2011-10-07 }}</ref> The currently accepted taxonomy is based on the [[List of Prokaryotic names with Standing in Nomenclature]] (LPSN)<ref>{{cite web | author=J.P. Euzéby | url= https://lpsn.dsmz.de/class/erysipelotrichia | title=Erysipelotrichia | publisher=[[List of Prokaryotic names with Standing in Nomenclature]] (LPSN) | access-date=2016-10-21}}</ref> and [[National Center for Biotechnology Information]] (NCBI)<ref>{{cite web |author = Sayers| url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Undef&id=526524&lvl=9&lin |title=Erysipelotrichia |access-date=2013-03-20 |publisher=[[National Center for Biotechnology Information]] (NCBI) taxonomy database |display-authors=etal}}</ref> {| class="wikitable" |- ! colspan=1 | 16S rRNA based [[The All-Species Living Tree Project|LTP]]_08_2023<ref name=LTP>{{cite web|title=The LTP |url=https://imedea.uib-csic.es/mmg/ltp/#LTP| access-date=20 November 2023}}</ref><ref>{{cite web|title=LTP_all tree in newick format|url=https://imedea.uib-csic.es/mmg/ltp/wp-content/uploads/ltp/LTP_all_08_2023.ntree |access-date=20 November 2023}}</ref><ref>{{cite web|title=LTP_08_2023 Release Notes| url=https://imedea.uib-csic.es/mmg/ltp/wp-content/uploads/ltp/LTP_08_2023_release_notes.pdf |access-date=20 November 2023}}</ref> ! colspan=1 | 120 marker proteins based [[Genome Taxonomy Database|GTDB]] 08-RS214<ref name="about">{{cite web |title=GTDB release 08-RS214 |url=https://gtdb.ecogenomic.org/about#4%7C |website=[[Genome Taxonomy Database]]|access-date=10 May 2023}}</ref><ref name="tree">{{cite web |title=bac120_r214.sp_label |url=https://data.gtdb.ecogenomic.org/releases/release214/214.0/auxillary_files/bac120_r214.sp_labels.tree |website=[[Genome Taxonomy Database]]|access-date=10 May 2023}}</ref><ref name="taxon_history">{{cite web |title=Taxon History |url=https://gtdb.ecogenomic.org/taxon_history/ |website=[[Genome Taxonomy Database]]|access-date=10 May 2023}}</ref> |- | style="vertical-align:top| {{Clade | style=font-size:90%;line-height:80% |1={{clade |1={{clade |label1=[[Bacilli|Bacilliia]] |1={{clade |1=other |2={{clade |1={{clade |1={{clade |label1="Lottiidibacillaceae" |1=''[[Lottiidibacillus]]'' }} |2={{clade |1={{clade |label1=Culicoidibacteraceae |1=''[[Culicoidibacter]]'' }} |2={{clade |label1=[[Haloplasmataceae]] |1=''[[Haloplasma]]'' ♦ |label2=[[Turicibacteraceae]] |2=''[[Turicibacter]]'' }} }} }} |2=other }} }} }} |2={{clade |label1=[[Fusobacteriota]] |1={{clade |label1=[[Fusobacteria|Fusobacteriia]] |1=[[Fusobacteriales]] }} |label2=[[Mycoplasmatota]] |2={{clade |1={{clade |1=''[[Asteroleplasma]]'' ♦ |2=[[Coprobacillaceae]] }} |2={{clade |1={{clade |label1=[[Erysipelotrichales]] |1=[[Erysipelotrichaceae]] |label2=[[Acholeplasmatales]] |sublabel2=/[[Anaeroplasmatales]] |2=♦ [[Acholeplasmataceae]] <br>(incl. [[Anaeroplasmataceae]])<br/> }} |2={{clade |label1=Mollicutes ♦ |1={{clade |1=[[Metamycoplasmataceae]] |2={{clade |label1=[[Mycoplasmoidales]] |1={{clade |1=''[[Spiroplasma]]'' species-group 3 |2=[[Mycoplasmoidaceae]] }} |label2=[[Mycoplasmatales]] |2={{clade |1=''[[Spiroplasma]]'' {[[Spiroplasmataceae]]} |2={{clade |1=''[[Spiroplasma]]'' species-group 2 |2={{clade |1="''[[Edwardiiplasma]]''" |2={{clade |1=[[Entomoplasmataceae]] |2=[[Mycoplasmataceae]] }} }} }} }} }} }} }} }} }} }} }} }} | {{Clade | style=font-size:90%;line-height:80% |label1=[[Bacilli|Bacilliia]] |1={{clade |1=other |2={{clade |1={{clade |1="Staphylococcales" |2={{clade |label1="Erysipelotrichidae" |1={{clade |1={{clade |label1=Culicoidibacterales |1={{clade |label1=Culicoidibacteraceae |1=''[[Culicoidibacter]]'' }} }} |2={{clade |1={{clade |label1=[[Haloplasmatales]] |1={{clade |label1=[[Turicibacteraceae]] |1=''[[Turicibacter]]'' |label2=[[Haloplasmataceae]] |2=''[[Haloplasma]]'' ♦ }} }} |2={{clade |1={{clade |1={{clade |label1="Bathyoplasmales" |1={{clade |label1="Bathyoplasmaceae" |1="''Ca.'' [[Bathoplasma]]" [MAG-NZ] }} }} |2={{clade |label1="Izemoplasmatales" |1={{clade |1="Hujiaoplasmataceae" |2="Izemoplasmataceae" }} |label2=[[Acholeplasmatales]] ♦ |sublabel2=/[[Anaeroplasmatales]]) |2={{clade |1=[[Anaeroplasmataceae]] |2={{clade |1=UBA5453 |2=[[Acholeplasmataceae]] }} }} }} }} |2={{clade |1={{clade |label1="Aphodocolales" |1="Aphodocolaceae" [RF39] }} |2={{clade |1={{clade |label1=RFN20 |1={{clade |1="Caccosomataceae" |2={{clade |1="Enterosomataceae" |2="Enteromonadaceae" }} }} |label2=[[Erysipelotrichales]] |2={{clade |1=[[Coprobacillaceae]] |2=[[Erysipelotrichaceae]] }} }} |label2=[[Mycoplasmatales]] ♦ |sublabel2=[[Entomoplasmatales]]) |2={{clade |1={{clade |1=''[[Spiroplasma]]'' species-group 3 {VBWQ01} |2=[[Mycoplasmataceae]] (incl. [[Entomoplasmataceae]] & [[Spiroplasmataceae]]) }} |2={{clade |1={{clade |1="''Ca.'' [[Spiroplasma holothuricola]]" {MT37} |2=[[Mycoplasmoidaceae]] }} |2={{clade |1="''Ca.'' [[Hepatoplasma]]" {"Hepatoplasmataceae"} |2=[[Metamycoplasmataceae]] }} }} }} }} }} }} }} }} }} }} |2=[[Lactobacillales]] }} }} }} |} ♦ Paraphyletic Mollicutes ==See also== * [[List of bacterial orders]] * [[List of bacteria genera]] ==References== {{Reflist|2}} ==External links== {{MeshName|Mollicutes}} {{Bacteria classification}} {{Taxonbar|from=Q131796}} [[Category:Mollicutes| ]] [[Category:Bacteria classes]] </textarea><div class="templatesUsed"><div class="mw-templatesUsedExplanation"><p><span id="templatesused">Pages transcluded onto the current version of this page<span class="posteditwindowhelplinks"> (<a href="/wiki/Help:Transclusion" title="Help:Transclusion">help</a>)</span>:</span> </p></div><ul> <li><a href="/wiki/Template:;" title="Template:;">Template:;</a> (<a href="/w/index.php?title=Template:;&action=edit" title="Template:;">view source</a>) (template editor protected)</li><li><a href="/wiki/Template:Anglicise_rank" title="Template:Anglicise rank">Template:Anglicise 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title="Module:Disambiguation/templates">view source</a>) (protected)</li><li><a href="/wiki/Module:Exponential_search" title="Module:Exponential search">Module:Exponential search</a> (<a href="/w/index.php?title=Module:Exponential_search&action=edit" title="Module:Exponential search">view source</a>) (template editor protected)</li><li><a href="/wiki/Module:Format_link" title="Module:Format link">Module:Format link</a> (<a href="/w/index.php?title=Module:Format_link&action=edit" title="Module:Format link">view source</a>) (template editor protected)</li><li><a href="/wiki/Module:GetParameters" title="Module:GetParameters">Module:GetParameters</a> (<a href="/w/index.php?title=Module:GetParameters&action=edit" title="Module:GetParameters">view source</a>) (protected)</li><li><a href="/wiki/Module:Hatnote" title="Module:Hatnote">Module:Hatnote</a> (<a href="/w/index.php?title=Module:Hatnote&action=edit" title="Module:Hatnote">view source</a>) (template editor protected)</li><li><a 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