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== Evolution and phylogeny == [[Image:Rafflesia 80 cm.jpg|thumb|''Rafflesia keithii'' bloom, approximately 80 cm in diameter near Taman Nasional Rafflesia [[Bengkulu]], Indonesia]] [[Image:Rafflesia kerrii flower closeup.jpg|thumb|''Rafflesia kerrii'' flower near [[Langkat]], Indonesia]] [[Image:Rafflesia sp Kinabalu2.jpg|thumb|Three ''Rafflesia pricei'' growing in close proximity near [[Mount Kinabalu]], Borneo]] Comparison of [[mitochondrial DNA]] (mtDNA) sequences of ''Rafflesia'' with other [[angiosperm]] mtDNA indicated this parasite evolved from photosynthetic plants of the order [[Malpighiales]].<ref>{{Cite journal| author-link = Barkman, T. J. | author2= S.-H. Lim | author3=K. Mat Salleh | author4=J. Nais | title = Mitochondrial DNA sequences reveal the photosynthetic relatives of ''Rafflesia'', the world's largest flower | journal = PNAS | volume = 101 | issue = 3 | pages = 787β792 | date = January 20, 2004 | doi = 10.1073/pnas.0305562101 | author = Barkman, T. J. | pmid = 14715901 | pmc = 321759 | doi-access= free }}</ref> Another 2004 study confirmed this result using both mtDNA and nuclear DNA sequences, and showed the three other groups traditionally classified in Rafflesiaceae were unrelated.<ref>{{Cite journal| author-link = Nickrent, D. L. | author2=A. Blarer | author3=Y.-L. Qiu | author4=R. Vidal-Russell | author5=F. E. Anderson | title = Phylogenetic inference in Rafflesiales: the influence of rate heterogeneity and horizontal gene transfer | journal = BMC Evolutionary Biology | volume = 4 | date = October 20, 2004 | doi = 10.1186/1471-2148-4-40 | author = Nickrent, D. L. | pages = 40 | pmid = 15496229 | pmc = 528834 | doi-access=free }}</ref> A 2007 study more specifically found ''Rafflesia'' and its relatives to be embedded within the family [[Euphorbiaceae]] as traditionally [[Circumscription (taxonomy)|circumscribed]], which was surprising, as members of that family typically have very small flowers. According to their analysis, the rate of flower size evolution was more or less constant throughout the family except at the origin of [[Rafflesiaceae]], where the flowers rapidly evolved to become much larger before reverting to the slower rate of change.<ref name=DaviLatvNickWurd07/> To maintain [[Monophyly|monophyletic]] families, in 2016 the [[APG IV system]] separated the family [[Peraceae]] from the Euphorbiaceae.<ref name=APGIV/> A summary cladogram is shown below,<ref name=DaviLatvNickWurd07>{{Citation |last1=Davis |first1=Charles C. |last2=Latvis |first2=Maribeth |last3=Nickrent |first3=Daniel L. |last4=Wurdack |first4=Kenneth J. |last5=Baum |first5=David A. |date=2007-03-30 |title=Floral Gigantism in Rafflesiaceae |journal=Science |volume=315 |issue=5820 |page=1812 |doi=10.1126/science.1135260 |pmid=17218493 |bibcode=2007Sci...315.1812D |s2cid=27620205 |name-list-style=amp |doi-access=free }}</ref> with family placements in the [[APG IV system]].<ref name=APGIV>{{Citation |last=Angiosperm Phylogeny Group |year=2016 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |journal=[[Botanical Journal of the Linnean Society]] |volume=181 |issue=1 |pages=1β20 |doi=10.1111/boj.12385 |doi-access=free }}</ref> {{clade |label1=Euphorbiaceae ''sensu lato'' |1={{clade |1=[[Peraceae]] (3 genera in the study) |2={{clade |label1=Rafflesiaceae |1={{clade |1=''[[Sapria]]'' |2={{clade |1=''[[Rhizanthes]]'' |2=''Rafflesia'' }} }} |2=Euphorbiaceae ''sensu stricto'' (18 genera in the study) }} }} }} A 2010 [[Molecular phylogenetics|molecular phylogenetic]] study of 18 species of ''Rafflesia'' found that they fell into four clear-cut geographically defined groups:<ref name=BendSchuGussNais10>{{Citation |mode=cs1 |last1=Bendiksby |first1=Mika |last2=Schumacher |first2=Trond |last3=Gussarova |first3=Galina |last4=Nais |first4=Jamili |last5=Mat-Salleh |first5=Kamarudin |last6=Sofiyanti |first6=Nery |last7=Madulid |first7=Domingo |last8=Smith |first8=Stephen A. |last9=Barkman |first9=Todd |date=2010-11-01 |title=Elucidating the evolutionary history of the Southeast Asian, holoparasitic, giant-flowered Rafflesiaceae: Pliocene vicariance, morphological convergence and character displacement |journal=Molecular Phylogenetics and Evolution |volume=57 |issue=2 |pages=620β633 |doi=10.1016/j.ympev.2010.08.005 |pmid=20723606 |bibcode=2010MolPE..57..620B |name-list-style=amp }}</ref> {{clade |label1=''Rafflesia'' |1={{clade |label1=Philippines group |1={{clade |1=''[[Rafflesia speciosa]]'' |2={{clade |1=''[[Rafflesia manillana]]'' |2={{clade |1=''[[Rafflesia lobata]]'' |2=''Rafflesia'' sp. }} }} }} |2={{clade |label1=Peninsular Malaysia group |1={{clade |1=''[[Rafflesia kerrii]]'' |2={{clade |1=''[[Rafflesia cantleyi]]'' |2=''[[Rafflesia azlanii]]'' }} }} |2={{clade |label1=Borneo group |1={{clade |1=''[[Rafflesia tuan-mudae]]'' |2={{clade |1=''[[Rafflesia tengku-adlinii]]'' |2={{clade |1=''[[Rafflesia keithii]]'' |2=''[[Rafflesia pricei]]'' }} }} }} |label2=Sumatra + Java group |2={{clade |1={{clade |1=''[[Rafflesia patma]]'' |2=''[[Rafflesia rochussenii]]'' }} |2={{clade |1=''[[Rafflesia zollingeriana]]'' |2={{clade |1=''[[Rafflesia gadutensis]]'' |2={{clade |1=''[[Rafflesia hasseltii]]'' |2={{clade |1=''[[Rafflesia micropylora]]'' |2=''[[Rafflesia arnoldii]]'' }} }} }} }} }} }} }} }} }} However, the clear [[monophyly]] of the four geographical clades does not correspond to any clear difference in appearance. There is no consistency within the clades in the size of flowers, or the presence or absence of white warts; species in different clades resemble one another more than they do some other species within the same clade. [[Homoplasy]] β repeatedly gaining or losing traits β seems to be the rule within ''Rafflesia''.<ref name=BendSchuGussNais10/> === Accepted species === {{div col|colwidth=30em}} {{As of|2020|October}}, [[Plants of the World Online]] accepted the following species:<ref name=POWO_77089854-1>{{cite web |title=''Rafflesia'' R.Br. ex Gray |work=Plants of the World Online |publisher=Royal Botanic Gardens, Kew|url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:77089854-1 |access-date=2020-10-26 }}</ref> *''[[Rafflesia arnoldii]]'' β [[Sumatra]], [[Borneo]] *''[[Rafflesia aurantia]]'' β Philippines ([[Luzon]]) *''[[Rafflesia azlanii]]'' β [[Peninsular Malaysia]] *''[[Rafflesia baletei]]'' β Philippines (Luzon) *''[[Rafflesia banaoana]]'' <small>Malabrigo</small> β Philippines (Luzon); treated as a synonym of ''R. leonardi'' by other sources<ref name=SIUC>{{cite web | title = ''Rafflesia leonardi'' Barcelona & Pelser | url = https://parasiticplants.siu.edu/Rafflesiaceae/Raff.leonardi.page.html | publisher = Parasitic Plant Connection | access-date = 19 January 2022}}</ref> *''[[Rafflesia bengkuluensis]]'' β Sumatra ([[Indonesia]]) *''[[Rafflesia borneensis]]'' <small>Koord.</small> β [[Indonesian Borneo]] (northeast [[Kalimantan]]) *''[[Rafflesia camarinensis]]'' <small>F.B.Valenz., Jaucian-Adan, Agoo & Madulid</small> β the [[Philippines]] *''[[Rafflesia cantleyi]]'' β Peninsular Malaysia *''[[Rafflesia ciliata]]'' <small>Koord.</small> β Indonesian Borneo (northeast [[Kalimantan]]) *''[[Rafflesia consueloae]]'' β Philippines (Luzon) *''[[Rafflesia gadutensis]]'' β Sumatra ([[Indonesia]]) *''[[Rafflesia hasseltii]]'' β Sumatra *''[[Rafflesia horsfieldii]]'' <small>R.Br.</small> β West [[Java]] (Indonesia) *''[[Rafflesia keithii]]'' β Borneo *''[[Rafflesia kemumu]]'' <small>Susatya, Hidayati & Riki</small> β Sumatra (Indonesia) *''[[Rafflesia kerrii]]'' β Thailand, Peninsular Malaysia *''[[Rafflesia lagascae]]'' β Philippines (Luzon) *''[[Rafflesia lawangensis]]'' β Sumatra *''[[Rafflesia leonardi]]'' β Philippines (Luzon) *''[[Rafflesia lobata]]'' β Philippines ([[Panay]]) *''[[Rafflesia manillana]]'' β Philippines ([[Samar]]) *''[[Rafflesia meijeri]]'' <small>Wiriad. & Sari</small> β North Sumatra, Indonesia *''[[Rafflesia micropylora]]'' β Sumatra *''[[Rafflesia mira]]'' β Philippines ([[Mindanao]]) *''[[Rafflesia mixta]]'' β Philippines (Mindanao) *''[[Rafflesia parvimaculata]]'' <small>Sofiyanti, K.Mat-Salleh, Khairil, Zuhailah, Mohd.Ros. & Burs</small> β Peninsular Malaysia *''[[Rafflesia philippensis]]'' β Philippines (Luzon) *''[[Rafflesia pricei]]'' β Borneo *''[[Rafflesia rochussenii]]'' β Java, Sumatra *''[[Rafflesia schadenbergiana]]'' β Philippines (Mindanao) *''[[Rafflesia sharifah-hapsahiae]]'' <small>J.H.Adam, R.Mohamed, Aizat-Juhari & K.L.Wan</small> β Peninsular Malaysia *''[[Rafflesia speciosa]]'' β Philippines ([[Panay]]) *''[[Rafflesia su-meiae]]'' <small>M. Wong, Nais & F.Gan</small> β Peninsular Malaysia *''[[Rafflesia tengku-adlinii]]'' β Borneo ([[Sabah]]) *''[[Rafflesia tiomanensis]]'' <small>Siti-Munirah, Salamah & Razelan</small> β [[Tioman Island|Pulau Tioman]], Peninsular Malaysia<ref>{{cite journal | author1 = Siti-Munirah, Mat Yunoh | author2 = Salamah, Ahmad | author3 = Razelan, Mohd Shah | name-list-style = amp | title = ''Rafflesia tiomanensis'' (Rafflesiaceae), a new species from Pulau Tioman, Pahang, Malaysia | url = https://www.researchgate.net/publication/349862173 | date = March 2021 | journal = Malayan Nature Journal | volume = 73 | issue = 1 | pages = 19β26}}</ref> *''[[Rafflesia tuan-mudae]]'' β Borneo ([[Sarawak]]) *''[[Rafflesia tuanku-halimii]]'' <small>J.H.Adam, Aizat-Juhari, Azilah & K.L.Wan</small> β Peninsular Malaysia *''[[Rafflesia verrucosa]]'' β Philippines (Mindanao) *''[[Rafflesia witkampii]]'' <small>Koord.</small> β Indonesian Borneo (East Kalimantan) *''[[Rafflesia zollingeriana]]'' <small>Koord.</small> β East Java ([[Indonesia]]) {{div col end}} === Other names === *''[[Rafflesia patma]]'' β [[Java]]; considered a [[Synonym (taxonomy)|synonym]] of ''R. horsfieldii'' by Plants of the World Online,<ref name=POWO_77089854-1/> apparently based on the confused writings of a British historian of botany,<ref name=Mabberley1999/> although ''R. horsfieldii'', for which no specimens were ever collected, only a drawing made which was lost centuries ago, is not considered a valid taxon by ''Rafflesia'' experts.<ref name=Meijer1988>{{cite journal |last1=Meijer |first1=W. |last2=Veldkamp |first2=J. F. |date=1988 |title=A revision of ''Rhizanthes'' (Rafflesiaceae) |url=https://repository.naturalis.nl/pub/525536/BLUM1988033002005.pdf |journal=Blumea |volume=33 |issue=2 |pages=329β342 |issn=2212-1676 |access-date=28 October 2020}}</ref> *''[[Rafflesia titan]]'' β A synonym of ''R. arnoldii''; it was in fact validly published earlier, being rushed to publication in Singapore in 1820 by a British botanist who feared that the French, who had actually discovered a species before the British, might deny the glory of the [[species description]] to servants of the British Empire.<ref name=Meijer1997/> In order to retain the honour of naming the species to the famous British scientist Robert Brown, the historian of botany mentioned above chose to pretend a 1821 pre-print Brown sent to a colleague was a valid 'effective publication',<ref name=Mabberley1999/> which has been accepted by the relevant British institutions.<ref name=POWO_77089854-1/><ref name=ipni4>{{cite web |title=''Rafflesia arnoldii'' |url=https://www.ipni.org/n/316069-1 |website=[[International Plant Names Index]] |publisher=The Royal Botanic Gardens, Kew, Harvard University Herbaria & Libraries and Australian National Botanic Gardens |access-date=9 November 2020}}</ref> === Loss of the chloroplast genome === Research published in 2014 revealed that one Philippine ''Rafflesia'' species from the island of [[Luzon]], ''R. lagascae'' (a synonym of [[Rafflesia manillana|''R. manillana'']]), may have lost the genome of its [[chloroplast]] and it is speculated that the loss happened due to the parasitic lifestyle of the plant.<ref name=Molina2014/>
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