Difference between revisions of "Rhamnaceae"

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{{taxobox
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#redirect [[:Category:Rhamnaceae]]
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|name = Buckthorn family
 
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|image = Rhamnus pumila Atlas Alpenflora.jpg
 
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|image_caption = ''[[Rhamnus pumila]]'' <small>Turra</small>
 
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|regnum = [[Plant]]ae
 
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|unranked_divisio = [[Angiosperms]]
 
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|unranked_classis = [[Eudicots]]
 
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|unranked_ordo = [[Rosids]]
 
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|ordo = [[Rosales]]<ref>{{cite journal|author = Walter S. Judd and Richard G. Olmstead|year = 2004|title = A survey of tricolpate (eudicot) phylogenetic relationships|journal = American Journal of Botany| volume = 91|pages = 1627–1644|doi = 10.3732/ajb.91.10.1627|pmid=21652313|issue = 10}} ([http://www.amjbot.org/cgi/content/full/91/10/1627 full text] )</ref>
 
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|familia = '''Rhamnaceae'''
 
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|familia_authority = [[Antoine Laurent de Jussieu|Juss.]]
 
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|type_genus = ''[[Rhamnus (plant)|Rhamnus]]''
 
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|type_genus_authority = [[Carl Linnaeus|L.]]
 
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|subdivision_ranks = Genera
 
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|subdivision = See text.
 
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|synonyms = Frangulaceae <small>DC.</small><br/>
 
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Phylicaceae <small>J.Agardh</small><br/>
 
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Ziziphaceae <small>Adans. ex Post & Kuntze</small><ref name="GRIN">{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?953 |title=Family: ''Rhamnaceae''  Juss., nom. cons. |work=Germplasm Resources Information Network |publisher=United States Department of Agriculture |date=2003-01-17 |accessdate=2011-01-29}}</ref>
 
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| range_map = Rhamnaceae Distribution.svg
 
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| range_map_caption = The range of Rhamnaceae.
 
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}}
 
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The '''Rhamnaceae''' are a large [[Family (biology)|family]] of [[flowering plant]]s, mostly trees, [[shrub]]s, and some vines, commonly called the '''buckthorn family'''.<ref name=FPSMM/>
 
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The family contains about 55 genera and 950 species.<ref name="Christenhusz-Byng2016">{{cite journal | author = Christenhusz, M. J. M., and Byng, J. W. | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | publisher = Magnolia Press }}</ref> The Rhamnaceae have a worldwide distribution, but are more common in the subtropical and tropical regions. The earliest fossil evidence of Rhamnaceae is from the Late Cretaceous. Fossil flowers have been collected from the Upper Cretaceous of Mexico and the Paleocene of Argentina.<ref>{{Cite journal|last=Jud|first=Nathan A.|last2=Gandolfo|first2=Maria A.|last3=Iglesias|first3=Ari|last4=Wilf|first4=Peter|date=2017-05-10|title=Flowering after disaster: Early Danian buckthorn (Rhamnaceae) flowers and leaves from Patagonia|url=http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0176164|journal=PLOS ONE|volume=12|issue=5|pages=e0176164|doi=10.1371/journal.pone.0176164|issn=1932-6203|pmc=5425202|pmid=28489895}}</ref>
 
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Leaves of Rhamnaceae family members are [[Simple leaf|simple]], i.e., the leaf blades are not divided into smaller leaflets.<ref name=FPSMM>Flowering Plants of the Santa Monica Mountains, Nancy Dale, 2nd Ed. 2000, p. 166</ref> Leaves can be either alternate and spiraling, or [[opposite leaves|opposite]]. Stipules are present. These leaves are modified into spines in many genera, in some (e.g. ''[[Paliurus spina-christi]]'' and ''[[Colletia]] cruciata'') spectacularly so. ''Colletia'' stands out by having two [[axil]]lary buds instead of one, one developing into a thorn, the other one into a shoot.
 
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[[File:Ceanothus cuneatus1.jpg|thumb|right|Flowers of ''[[Ceanothus cuneatus]]'']]
 
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[[File:Helinus integrifolius, blom, Skeerpoort, b.jpg|thumb|right|Bisexual flower of ''[[Helinus]]'', with five sepals and petals, and a yellow, annular nectary disk. The small, clawed petals embrace the stamens.]]
 
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[[File:Ziziphus mucronata, blomme, Roodeplaat NR, c.jpg|thumb|right|Flowers of ''[[Ziziphus mucronata]]'']]
 
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[[File:Rhamnus saxatilis (Felsen-Kreuzdorn) IMG 7432.JPG|thumb|right|Flowers of ''[[Rhamnus saxatilis]]'']]
 
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The [[flower]]s are radially symmetrical. There are 5 (sometimes 4) separate sepals and 5 (sometimes 4 or none) separate petals. The petals may be white, yellowish, greenish, pink or blue, and are small and inconspicuous in most genera, though in some (e.g. ''Ceanothus'') the dense clusters of flowers are conspicuous. The 5 or 4 [[stamen]]s are opposite the petals.<ref name=FPSMM/> The [[Ovary (plants)|ovary]] is superior, with 2 or 3 [[ovule]]s (or one by abortion).
 
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The [[fruit]]s are mostly berries, fleshy [[drupe]]s, or nuts. Some are adapted to wind carriage, but most are dispersed by mammals and birds. Chinese jujube is the fruit of the [[jujube]] tree (''Ziziphus zizyphus'') and is a major fruit in China.
 
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The American genus ''[[Ceanothus]]'', which has several showy [[Ornamental plant|ornamental]] species, has [[Nitrogen fixation#Non-leguminous nitrogen-fixing plants|nitrogen-fixing]] [[root nodule]]s.
 
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Economic uses of the Rhamnaceae are chiefly as ornamental plants and as the source of many brilliant green and yellow [[dye]]s. The [[wood]] of ''[[Rhamnus (genus)|Rhamnus]]'' was also the most favoured species to make [[charcoal]] for use in [[gunpowder]] before the development of modern propellants.
 
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== Genera ==
 
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{|
 
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|- valign=top
 
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|
 
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;Tribe '''[[Ampelozizyphus|Ampelozizipheae]]'''
 
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*''[[Ampelozizyphus]]'' <small>Ducke</small>
 
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;Tribe '''[[Bathiorhamneae]]'''
 
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*''[[Bathiorhamnus]]'' <small>Capuron</small>
 
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;Tribe '''[[Colletieae]]'''
 
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*''[[Adolphia]]'' <small>Meisn.</small>
 
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*''[[Colletia]]'' <small>Comm. ex Juss.</small>
 
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*''[[Discaria]]'' <small>Hook.</small>
 
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*''[[Kentrothamnus]]'' <small>Suess. & Overkott</small>
 
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*''[[Ochetophila]]'' <small>Poepp. ex Reissek</small>
 
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*''[[Retanilla]]'' <small>(DC.) Brongn.</small>
 
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*''[[Trevoa]]'' <small>Miers ex Hook.</small>
 
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;Tribe '''[[Doerpfeldieae]]'''
 
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*''[[Doerpfeldia]]'' <small>Urb.</small>
 
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;Tribe '''[[Gouanieae]]'''
 
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*''[[Alvimiantha]]'' <small>Grey-Wilson</small>
 
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*''[[Crumenaria]]'' <small>Mart.</small>
 
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*''[[Gouania]]'' <small>Jacq.</small>
 
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*''[[Helinus]]'' <small>E.Mey. ex Endl.</small>
 
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*''[[Johnstonalia]]'' <small>Tortosa</small> (originally named ''Johnstonia''; by some treated as ''[[Gouania]]'')
 
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*''[[Pleuranthodes]]'' <small>Weberb.</small>
 
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*''[[Reissekia]]'' <small>Endl.</small>
 
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;Tribe '''[[Maesopsideae]]'''
 
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*''[[Maesopsis]]'' <small>Engl.</small>
 
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;Tribe '''[[Paliureae]]'''
 
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*''[[Hovenia]]'' <small>Thunb.</small>
 
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*''[[Paliurus]]'' <small>Mill.</small>
 
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*''[[Ziziphus]]'' <small>Mill.</small>
 
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*''[[Sarcomphalus]]'' <small>P.Browne emend. Hauenschild</small>
 
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;Tribe '''[[Phyliceae]]'''
 
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*''[[Saint Helena Olive|Nesiota]]''† <small>Hook.f.</small>
 
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*''[[Noltea]]'' <small>Rchb.</small>
 
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*''[[Phylica]]'' <small>L.</small>
 
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*''[[Trichocephalus]]'' <small>Brogn.</small>
 
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;Tribe '''[[Pomaderreae]]'''
 
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*''[[Blackallia]]'' <small>C.A.Gardner</small>
 
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*''[[Cryptandra]]'' <small>Sm.</small>
 
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*''[[Papistylus]]'' <small>Kellermann, Rye & K.R.Thiele</small>
 
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*''[[Polianthon]]'' <small>K.R.Thiele</small>
 
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*''[[Pomaderris]]'' <small>Labill.</small>
 
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*''[[Serichonus]]'' <small>K.R.Thiele</small>
 
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*''[[Siegfriedia]]'' <small>C.A.Gardner</small>
 
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*''[[Spyridium]]'' <small>Fenzl</small>
 
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*''[[Stenanthemum]]'' <small>Reissek</small>
 
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*''[[Trymalium]]'' <small>Fenzl</small>
 
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|
 
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;Tribe '''[[Rhamneae]]'''
 
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*''[[Auerodendron]]'' <small>Urb.</small>
 
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*''[[Berchemia]]'' <small>Neck. ex DC.</small>
 
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*''[[Berchemiella]]'' <small>Nakai</small>
 
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*''[[Condalia]]'' <small>Cav.</small>
 
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*''[[Frangula]]'' <small>Mill.</small>
 
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*''[[Karwinskia]]'' <small>Zucc.</small>
 
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*''[[Krugiodendron]]'' <small>Urb.</small>
 
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*''[[Reynosia]]'' <small>Griseb.</small>
 
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*''[[Rhamnella]]'' <small>Miq.</small>
 
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*''[[Rhamnidium]]'' <small>Reissek</small>
 
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*''[[Rhamnus (plant)|Rhamnus]]'' <small>L.</small>
 
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*''[[Sageretia]]'' <small>Brongn.</small>
 
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*''[[Scutia]]'' <small>(Comm. ex DC.) Brongn.</small>
 
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;Tribe '''[[Ventilagineae]]'''
 
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*''[[Smythea]]'' <small>Seem.</small>
 
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*''[[Ventilago]]'' <small>Gaertn.</small>
 
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;'''[[Incertae sedis]]'''
 
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*''[[Alphitonia]]'' <small>[[Stephan Ladislaus Endlicher|Endl.]]</small> (close to ''[[Colubrina]]'')
 
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*''[[Araracuara]]'' <small>Fern.Alonso</small>
 
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*''[[Ceanothus]]'' <small>L.</small> (close to Pomaderreae)
 
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*''[[Chaydaia]]'' <small>Pit.</small> (by some treated as ''[[Rhamnella]]'')
 
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*''[[Colubrina]]'' <small>Rich. ex Brongn.</small>
 
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*''[[Emmenosperma]]'' <small>F.Muell.</small> (close to ''[[Colubrina]]'')
 
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*''[[Granitites]]'' <small>Rye</small><ref>{{FloraBase|name=''Granitites''|id= 21733}}</ref> (close to ''[[Alphitonia]]'')
 
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*''[[Hybosperma]]'' <small>Urb.</small> (by some treated as ''[[Colubrina]]'')
 
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*''[[Jaffrea]]'' <small>H.C.Hopkins & Pillon</small>
 
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*''[[Lasiodiscus]]'' <small>Hook.f.</small>
 
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*''[[Schistocarpaea]]'' <small>F.Muell.</small><ref name="GRINGenera">{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?953 |title=GRIN Genera of ''Rhamnaceae'' |work=Germplasm Resources Information Network |publisher=United |accessdate=2011-01-29}}</ref> (maybe belonging to Colletieae)
 
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*''[[Talguenea]]'' <small>Miers ex Endl.</small> (by some treated as ''[[Trevoa]]'')
 
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|}
 
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==Systematics==
 
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Modern [[molecular phylogenetics]] recommend the following clade-based classification of Rhamnaceae:<ref>{{cite journal | authors = Sun M, Naeem R, Su J-X, Cao Z-Y, J. Burleigh G, Soltis PS, Soltis DE, Chen Z-D. | year = 2016 | title = Phylogeny of the Rosidae: A dense taxon sampling analysis | journal = Journal of Systematics and Evolution | volume = 54 | issue = 4 | pages = 363–391 | url = http://onlinelibrary.wiley.com/wol1/doi/10.1111/jse.12211/abstract | doi = 10.1111/jse.12211}}</ref>
 
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{{Clade| style=line-height:75%;
 
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|1={{clade
 
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  |1=&nbsp;[[Elaeagnaceae]]&nbsp;([[Outgroup (cladistics)|outgroup]])
 
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  |label2=Rhamnaceae
 
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  |2={{clade
 
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    |1={{clade
 
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      |label1=[[Ampeloziziphoids]]
 
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      |1={{clade
 
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        |1=''[[Ventilago]]''
 
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        |2={{clade
 
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          |1=''[[Bathiorhamnus]]''
 
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          |2={{clade
 
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            |1=''[[Ampelozizyphus]]''
 
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            |2=''[[Doerpfeldia]]''
 
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            }}
 
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          }}
 
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        }}
 
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      |label2=[[Rhamnoids]]
 
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      |2={{clade
 
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        |1=''[[Maesopsis]]''
 
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        |2={{clade
 
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          |1={{clade
 
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            |1=''[[Scutia]]''
 
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            |2={{clade
 
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              |1=''[[Rhamnus (plant)|Rhamnus]]''
 
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              |2=''[[Frangula]]''
 
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              }}
 
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            }}
 
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          |2={{clade
 
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            |1=''[[Sageretia]]''
 
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            |2={{clade
 
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              |1=''[[Berchemia]]''
 
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              |2={{clade
 
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                |1={{clade
 
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                  |1=''[[Rhamnidium]]''
 
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                  |2=''[[Rhamnella]]''
 
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                  }}
 
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                |2={{clade
 
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                  |1=''[[Reynosia]]''
 
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                  |2={{clade
 
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                    |1=''[[Krugiodendron]]''
 
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                    |2={{clade
 
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                      |1=''[[Karwinskia]]''
 
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                      |2=''[[Condalia]]''
 
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                      }}
 
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                    }}
 
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                  }}
 
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                }}
 
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              }}
 
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            }}
 
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          }}
 
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        }}
 
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      }}
 
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    |label2=[[Ziziphoids]]
 
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    |2={{clade
 
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      |1=''[[Schistocarpeia]]''
 
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      |2={{clade
 
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        |1={{clade
 
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          |1={{clade
 
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            |1=''[[Hovenia]]''
 
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            |2={{clade
 
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              |1=''[[Ziziphus]]''
 
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              |2=''[[Paliurus]]''
 
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              }}
 
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            }}
 
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          |2={{clade
 
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            |1={{clade
 
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              |1=''[[Gouania]]''
 
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              |2=''[[Helinus]]''
 
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              }}
 
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            |2={{clade
 
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              |1=''[[Pleuranthodes]]''
 
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              |2={{clade
 
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                |1=''[[Crumenaria]]''
 
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                |2=''[[Reissekia]]''
 
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                }}
 
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              }}
 
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            }}
 
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          }}
 
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        |2={{clade
 
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          |1={{clade
 
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            |1=''[[Lasiodiscus]]''
 
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            |2=''[[Colubrina]]''
 
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            }}
 
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          |2={{clade
 
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            |1=''[[Emmenosperma]]''
 
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            |2={{clade
 
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              |1={{clade
 
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                |1=''[[Noltea]]''
 
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                |2={{clade
 
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                  |1=''[[Trichocephalus]]''
 
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                  |2={{clade
 
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                    |1=''[[Nesiota]]''
 
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                    |2=''[[Phylica]]''
 
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                    }}
 
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                  }}
 
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                }}
 
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              |2={{clade
 
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                |1={{clade
 
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                  |1=''[[Granitites]]''
 
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                  |2=''[[Alphitonia]]''
 
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                  }}
 
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                |label2=[[Colletieae]]
 
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                |2={{clade
 
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                  |1={{clade
 
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                    |1=''[[Adolphia]]''
 
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                    |2={{clade
 
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                      |1=''[[Trevoa]]''
 
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                      |2=''[[Talguenea]]''
 
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                      }}
 
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                    }}
 
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                  |2={{clade
 
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                    |1=''[[Discaria]]''
 
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                    |2=''[[Kentrothamnus]]''
 
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                    |3=''[[Colletia]]''
 
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                    |4=''[[Retanilla]]''
 
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                    }}
 
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                  }}
 
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                |3={{clade
 
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                  |1=''[[Ceanothus]]''
 
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                  |label2=[[Pomaderreae]]
 
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                  |2={{clade
 
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                    |1=''[[Siegfriedia]]''
 
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                    |2=''[[Cryptandra]]''
 
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                    |3={{clade
 
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                      |1=''[[Stenanthemum]]''
 
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                      |2={{clade
 
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                        |1=''[[Trymalium]]''
 
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                        |2={{clade
 
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                          |1=''[[Pomaderris]]''
 
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                          |2=''[[Spyridium]]''
 
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                          }}
 
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                        }}
 
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                      }}
 
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                    }}
 
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                  }}
 
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                }}
 
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              }}
 
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            }}
 
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          }}
 
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        }}
 
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      }}
 
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    }}
 
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  }}
 
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}}
 
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==References==
 
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{{Reflist|2}}
 
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== External links ==
 
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{{commons category}}
 
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{{wikispecies}}
 
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*[http://greif.uni-greifswald.de/floragreif/?fam=Rhamnaceae&gen=&spec=&flora_search=taxon Rhamnaceae of Mongolia in FloraGREIF]
 
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{{taxonbar}}
 
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[[Category:Rhamnaceae]]
 
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[[Category:Rosid families]]
 

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