Rhamnaceae
{{Short description|Family of flowering plants}}
{{cs1 config|name-list-style=vanc}}
{{Automatic taxobox
| name = Rhamnaceae
| fossil_range = {{fossilrange|Late Cretaceous|Recent}}
| image = Rhamnus pumila Atlas Alpenflora.jpg
| image_caption = Rhamnus pumila Turra
| taxon = Rhamnaceae
| authority = Juss.
| type_genus = Rhamnus
| type_genus_authority = L.
| subdivision_ranks = Genera
| subdivision = See text
| synonyms = Frangulaceae DC.
Phylicaceae J.Agardh
| range_map = Rhamnaceae Distribution.svg
| range_map_caption = The range of Rhamnaceae.
}}
The Rhamnaceae are a large family of flowering plants, mostly trees, shrubs, and some vines, commonly called the buckthorn family. Rhamnaceae is included in the order Rosales.{{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|doi-access = free}}
The family contains about 55 genera and 950 species.{{cite journal |vauthors=Christenhusz MJ, Byng JW | 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 | doi-access = free }} 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.{{Cite journal|last1=Jud|first1=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|journal=PLOS ONE|volume=12|issue=5|pages=e0176164|doi=10.1371/journal.pone.0176164|issn=1932-6203|pmc=5425202|pmid=28489895|bibcode=2017PLoSO..1276164J|doi-access=free}}
Leaves of family Rhamnaceae members are simple, i.e., the leaf blades are not divided into smaller leaflets.Flowering Plants of the Santa Monica Mountains, Nancy Dale, 2nd Ed. 2000, p. 166 Leaves can be either alternate or opposite. Stipules are present and modified into spines in many genera. In some (e.g. Paliurus spina-christi and Colletia paradoxa) spectacularly so. Colletia stands out by having two axillary buds instead of one, one developing into a thorn, the other one into a shoot.
File:Ceanothus cuneatus1.jpg]]
File:Helinus integrifolius, blom, Skeerpoort, b.jpg, with five sepals and petals, and a yellow, annular nectary disk. The small, clawed petals embrace the stamens.]]
File:Ziziphus mucronata, blomme, Roodeplaat NR, c.jpg]]
File:Rhamnus saxatilis (Felsen-Kreuzdorn) IMG 7432.JPG]]
The flowers 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 stamens are opposite the petals. The ovary is mostly superior, with 2 or 3 ovules (or one by abortion).
The fruits are mostly berries, fleshy drupes, 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 jujuba) and is a major fruit in China.
The American genus Ceanothus, which has several showy ornamental species, has nitrogen-fixing root nodules.{{cite journal |last1=Kummerow |first1=Jochen |last2=Alexander |first2=James V. |last3=Neel |first3=James W. |last4=Fishbeck |first4=Kathleen |title=Symbiotic Nitrogen Fixation in Ceanothus Roots |journal=American Journal of Botany |date=January 1978 |volume=65 |issue=1 |pages=63–69 |doi= 10.1002/j.1537-2197.1978.tb10836.x|jstor=2442555}}
Economic uses of the Rhamnaceae are chiefly as ornamental plants and as the source of many brilliant green and yellow dyes. The wood of Rhamnus was also the most favoured species to make charcoal for use in gunpowder before the development of modern propellants.
Alphitonia ponderosa (6691195369).jpg|Alphitonia ponderosa
Mountain flower white flower.jpg|Ceanothus
Ceanothus papillosus var roweanus 2.jpg|Ceanothus papillosus var. roweanus at the Regional Parks Botanic Garden, Berkeley, California
Genera
{{Columns-list|colwidth=20em|
- Tribe Ampelozizipheae
- Ampelozizyphus Ducke
- Tribe Bathiorhamneae
- Bathiorhamnus Capuron
- Tribe Colletieae
- Adolphia Meisn.
- Colletia Comm. ex Juss.
- Discaria Hook.
- Kentrothamnus Suess. & Overkott
- Ochetophila Poepp. ex Reissek
- Retanilla (DC.) Brongn.
- Trevoa Miers ex Hook.
- Tribe Doerpfeldieae
- Doerpfeldia Urb.
- Tribe Gouanieae
- Alvimiantha Grey-Wilson
- Crumenaria Mart.
- Gouania Jacq.
- Helinus E.Mey. ex Endl.
- Johnstonalia Tortosa (originally named Johnstonia; by some treated as Gouania)
- Pleuranthodes Weberb. – synonym of Gouania
- Reissekia Endl.
- Tribe Maesopsideae
- Maesopsis Engl.
- Tribe Paliureae
- Condaliopsis {{small|(Weberb.) Suess.}}
- Hovenia Thunb.
- Paliurus Mill.
- Ziziphus Mill.
- Sarcomphalus P.Browne emend. Hauenschild
- Tribe Phyliceae
- Nesiota† Hook.f. Extinct (2003)
- Noltea Rchb.
- Phylica L.
- Trichocephalus Brogn.
- Tribe Pomaderreae
- Blackallia C.A.Gardner
- Cryptandra Sm.
- Papistylus Kellermann, Rye & K.R.Thiele
- Polianthion K.R.Thiele
- Pomaderris Labill.
- Serichonus K.R.Thiele
- Siegfriedia C.A.Gardner
- Spyridium Fenzl
- Stenanthemum Reissek
- Trymalium Fenzl
- Tribe Rhamneae
- Auerodendron Urb.
- Berchemia Neck. ex DC.
- Berchemiella Nakai
- Condalia Cav.
- Frangula Mill.
- Karwinskia Zucc.
- Krugiodendron Urb.
- Phyllogeiton (Weberb.) Herzog
- Pseudoziziphus Hauenschild{{cite journal|jstor=taxon.65.1.47 |title=Phylogenetic relationships within the cosmopolitan buckthorn family (Rhamnaceae) support the resurrection of Sarcomphalus and the description of Pseudoziziphus gen. nov. |journal=Taxon |date=2016 |volume=65 |issue=1 |pages=47–64 |doi=10.12705/651.4 | vauthors = Hauenschild F, Matuszak S, Muellner-Riehl AN, Favre A }}
- Reynosia Griseb.
- Rhamnella Miq.
- Rhamnidium Reissek
- Rhamnus L.
- Sageretia Brongn.
- Scutia (Comm. ex DC.) Brongn.
- Tribe Ventilagineae
- Smythea Seem.
- Ventilago Gaertn.
- Incertae sedis
- Alphitonia Endl. (close to Colubrina)
- Araracuara Fern.Alonso
- Ceanothus L. (close to Pomaderreae)
- Chaydaia Pit. – synonym of Rhamnella
- Colubrina Rich. ex Brongn.
- Conalma {{small|G.L.Nesom}}
- Emmenosperma F.Muell. (close to Colubrina)
- Fenghwaia {{small|G.T.Wang & R.J.Wang}}
- Granitites Rye{{FloraBase|name=Granitites|id= 21733}} (close to Alphitonia)
- Hybosperma Urb. – synonym of Colubrina
- Jaffrea H.C.Hopkins & Pillon
- Lasiodiscus Hook.f.
- Oreorhamnus {{small|Ridl.}}
- Schistocarpaea F.Muell.{{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 |access-date=2011-01-29 |archive-url=https://web.archive.org/web/20150924061906/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?953 |archive-date=2015-09-24 }} (maybe belonging to Colletieae)
}}
Systematics
Modern molecular phylogenetics recommend the following clade-based classification of Rhamnaceae:{{cite journal |vauthors=Sun M, Naeem R, Su JX, Cao ZY, Burleigh JG, Soltis PS, Soltis DE, Chen ZD | 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 | doi = 10.1111/jse.12211| doi-access = free }}
{{Clade| style=line-height:85%;
|1={{clade
|1= Elaeagnaceae (outgroup)
|label2=Rhamnaceae
|2={{clade
|1={{clade
|label1=Ampeloziziphoids
|1={{clade
|1=Ventilago
|2={{clade
|2={{clade
|2=Doerpfeldia
}}
}}
}}
|label2=Rhamnoids
|2={{clade
|1=Maesopsis
|2={{clade
|1={{clade
|1=Scutia
|2={{clade
|1=Rhamnus
|2=Frangula
}}
}}
|2={{clade
|1=Sageretia
|2={{clade
|1=Berchemia
|2={{clade
|1={{clade
|1=Rhamnidium
|2=Rhamnella
}}
|2={{clade
|1=Reynosia
|2={{clade
|2={{clade
|1=Karwinskia
|2=Condalia
}}
}}
}}
}}
}}
}}
}}
}}
}}
|label2=Ziziphoids
|2={{clade
|2={{clade
|1={{clade
|1={{clade
|1=Hovenia
|2={{clade
|1=Ziziphus
|2=Paliurus
}}
}}
|2={{clade
|1={{clade
|1=Gouania
|2=Helinus
}}
|2={{clade
|2={{clade
|1=Crumenaria
|2=Reissekia
}}
}}
}}
}}
|2={{clade
|1={{clade
|1=Lasiodiscus
|2=Colubrina
}}
|2={{clade
|1=Emmenosperma
|2={{clade
|label1=Phyliceae
|1={{clade
|1=Noltea
|2={{clade
|2={{clade
|1=Nesiota
|2=Phylica
}}
}}
}}
|2={{clade
|1={{clade
|1=Granitites
|2=Alphitonia
}}
|label2=Colletieae
|2={{clade
|1={{clade
|1=Adolphia
|2=Trevoa
}}
|2={{clade
|1=Discaria
|3=Colletia
|4=Retanilla
}}
}}
|3={{clade
|1=Ceanothus
|label2=Pomaderreae
|2={{clade
|1=Siegfriedia
|2=Cryptandra
|3={{clade
|1=Stenanthemum
|2={{clade
|1=Trymalium
|2={{clade
|1=Pomaderris
|2=Spyridium
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
Fossil record
The fossil record of the family extends back to the Late Cretaceous, with records from Colombia{{Cite journal |last1=Correa |first1=Edwin |last2=Jaramillo |first2=Carlos |last3=Manchester |first3=Steven |last4=Gutierrez |first4=Mauricio |date=January 2010 |title=A fruit and leaves of Rhamnaceous affinities from the late Cretaceous (Maastrichtian) of Colombia |url=https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.0900093 |journal=American Journal of Botany |language=en |volume=97 |issue=1 |pages=71–79 |doi=10.3732/ajb.0900093 |pmid=21622368 |issn=0002-9122}} and Mexico.{{Cite journal |last1=Calvillo-Canadell |first1=Laura |last2=Cevallos-Ferriz |first2=Sergio R. S. |date=October 2007 |title=Reproductive structures of Rhamnaceae from the Cerro del Pueblo (Late Cretaceous, Coahuila) and Coatzingo (Oligocene, Puebla) Formations, Mexico |url=https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.94.10.1658 |journal=American Journal of Botany |language=en |volume=94 |issue=10 |pages=1658–1669 |doi=10.3732/ajb.94.10.1658 |pmid=21636362 |issn=0002-9122}} Remains from the mid-Cretaceous Burmese amber of Myanmar have been suggested to belong to this family by some authors{{Cite journal |last1=Shi |first1=Chao |last2=Wang |first2=Shuo |last3=Cai |first3=Hao-hong |last4=Zhang |first4=Hong-rui |last5=Long |first5=Xiao-xuan |last6=Tihelka |first6=Erik |last7=Song |first7=Wei-cai |last8=Feng |first8=Qi |last9=Jiang |first9=Ri-xin |last10=Cai |first10=Chen-yang |last11=Lombard |first11=Natasha |date=2022-01-31 |title=Fire-prone Rhamnaceae with South African affinities in Cretaceous Myanmar amber |url=http://dx.doi.org/10.1038/s41477-021-01091-w |journal=Nature Plants |volume=8 |issue=2 |pages=125–135 |doi=10.1038/s41477-021-01091-w |issn=2055-0278 |pmid=35102275 |s2cid=246443363}}{{Cite journal |last1=Oskolski |first1=Alexei A. |last2=Morris |first2=Benjamin B. |last3=Severova |first3=Elena E. |last4=Sokoloff |first4=Dmitry D. |year=2024 |title=Flowers from Myanmar amber confirm the Cretaceous age of Rhamnaceae but not of the extant genus Phylica |journal=Nature Plants |volume=10 |issue=2 |pages=219–222 |doi=10.1038/s41477-023-01591-x |pmid=38278949|bibcode=2024NatPl..10..219O |s2cid=267267981 }}, but this has been doubted by others.{{Cite journal |last1=Beurel |first1=Simon |last2=Bachelier |first2=Julien B. |last3=Schmidt |first3=Alexander R. |last4=Sadowski |first4=Eva-Maria |year=2024 |title=Novel three-dimensional reconstructions of presumed Phylica (Rhamnaceae) from Cretaceous amber suggest Lauralean affinities |journal=Nature Plants |volume=10 |issue=2 |pages=223–227 |doi=10.1038/s41477-023-01592-w |pmid=38278948|bibcode=2024NatPl..10..223B |s2cid=267267851 }} The earliest fossils of modern genera of the family date to the Eocene.{{Cite journal |last1=Jud |first1=Nathan A. |last2=Gandolfo |first2=Maria A. |last3=Iglesias |first3=Ari |last4=Wilf |first4=Peter |date=2017-05-10 |editor-last=Wong |editor-first=William Oki |title=Flowering after disaster: Early Danian buckthorn (Rhamnaceae) flowers and leaves from Patagonia |journal=PLOS ONE |language=en |volume=12 |issue=5 |pages=e0176164 |doi=10.1371/journal.pone.0176164 |doi-access=free |issn=1932-6203 |pmc=5425202 |pmid=28489895|bibcode=2017PLoSO..1276164J }}
References
{{Reflist|2}}
External links
{{Commons category}}
{{Wikispecies}}
- [http://greif.uni-greifswald.de/floragreif/?fam=Rhamnaceae&gen=&spec=&flora_search=taxon Rhamnaceae of Mongolia in FloraGREIF] {{Webarchive|url=https://web.archive.org/web/20160304061655/http://greif.uni-greifswald.de/floragreif/?fam=Rhamnaceae&gen=&spec=&flora_search=taxon |date=2016-03-04 }}
{{Angiosperm families}}
{{Taxonbar|from=Q156083}}
{{Authority control}}