Rhamnus (genus)

''Rhamnus'' is a genus of about 140 accepted species of shrubs or small trees, commonly known as buckthorns, in the family Rhamnaceae. Its species range from tall (rarely to ) and are native mainly in east Asia and North America, but found throughout the temperate and subtropical Northern Hemisphere, and also more locally in the subtropical Southern Hemisphere in parts of Africa and South America. One species, the common buckthorn (''Rhamnus cathartica''), is able to flourish as an invasive plant in parts of Canada and the United States, where it has become naturalized.

Both deciduous and evergreen species occur. The leaves are simple, long, and arranged alternately, in opposite pairs, or almost paired (subopposite). One distinctive character of many buckthorns is the way the veination curves upward towards the tip of the leaf. The plant bears fruits which are black or red berry-like drupes. The name is due to the woody spine on the end of each twig in many species. One species is known to have potential to be used medicinally.

Description

''Rhamnus'' species are shrubs or small to medium-sized trees, with deciduous or rarely evergreen foliage. Branches are unarmed or end in a woody spine. The leaf blades are undivided and pinnately veined. Leaf margins are serrate or rarely entire. ''Rhamnus'' species are generally dioecious, with male and female flowers on separate plants. Most species have yellowish green, small, unisexual or rarely polygamous flowers; which are produced singly or in axillary cymes, cymose racemes, or cymose panicles containing a few flowers. Calyx tube campanulate to cup-shaped, with 4 or 5 ovate-triangular sepals, which are adaxially ± distinctly keeled. Petals 4 or 5 but a few species may lack petals. The petals are shorter than the sepals. Flowers have 4 or 5 stamens which are surrounded by and equal in length the petals or are shorter. The anthers are dorsifixed. The superior ovary is free, rounded, with 2-4 chambers. Fruits are a 2-4 stoned, berrylike drupe, which is obovoid-globose or globose shaped. Seeds are obovoid or oblong-obovoid shaped, unfurrowed or abaxially or laterally margined with a long, narrow, furrow. The seeds have fleshy endosperm.

Species

, Plants of the World Online accepted the following species:

*'' Rhamnus alaternus'' L. – Italian buckthorn
*''Rhamnus alnifolia'' L'Hér. – alderleaf buckthorn, alder-leaved buckthorn
*'' Rhamnus alpina'' L.
*'' Rhamnus arguta'' Maxim. – sharp-tooth buckthorn
*'' Rhamnus arnottiana'' Gardner ex Thwaites
*'' Rhamnus aurea'' Heppeler
*'' Rhamnus baldschuanica'' Grubov
* ''Rhamnus'' × ''bermejoi'' P.Fraga & Rosselló
*'' Rhamnus biglandulosa'' Sessé & Moc.
*'' Rhamnus bodinieri'' H.Lév.
*'' Rhamnus brachypoda'' C.Y.Wu
*'' Rhamnus bungeana'' J.J.Vassil.
*'' Rhamnus calderoniae'' R.Fern.
*''Rhamnus cathartica'' L. – common buckthorn, purging buckthorn (orth. var. ''R. catharticus'')
*'' Rhamnus collettii'' Bhandari & Bhansali
*'' Rhamnus cordata'' Medw.
*'' Rhamnus coriophylla'' Hand.-Mazz.
*'' Rhamnus cornifolia'' Boiss. & Hohen.
*'' Rhamnus costata'' Maxim.
*'' Rhamnus crenulata'' Aiton
*'' Rhamnus crocea'' Nutt. – redberry buckthorn (subsp. ''crocea''), hollyleaf buckthorn (subsp. ''pilosa'')
** ''Rhamnus crocea'' subsp. ''ilicifolia'' (Kellogg) C.B.Wolf – synonym ''Rhamnus ilicifolia''
*'' Rhamnus dalianensis'' S.Y.Li & Z.H.Ning
*'' Rhamnus daliensis'' G.S.Fan & L.L.Deng
*'' Rhamnus darii'' Govaerts
*'' Rhamnus davurica'' Pall. – Dahurian buckthorn
*'' Rhamnus depressa'' Grubov
*'' Rhamnus diffusa'' Clos
*'' Rhamnus disperma'' Ehrenb. ex Boiss.
*'' Rhamnus dolichophylla'' Gontsch.
*'' Rhamnus dumetorum'' C.K.Schneid.
*'' Rhamnus erythroxyloides'' Hoffmanns.
*'' Rhamnus erythroxylum'' Pall.
*'' Rhamnus esquirolii'' H.Lév.
*'' Rhamnus fallax'' Boiss.
*'' Rhamnus flavescens'' Y.L.Chen & P.K.Chou
*'' Rhamnus formosana'' Matsum.
*'' Rhamnus fulvotincta'' F.P.Metcalf
* ''Rhamnus'' × ''gayeri'' Kárpáti ex Soó
*'' Rhamnus gilgiana'' Heppeler
*'' Rhamnus glandulosa'' Aiton
*'' Rhamnus glaucophylla'' Sommier
*'' Rhamnus globosa'' Bunge – Lokao buckthorn
*'' Rhamnus grandiflora'' C.Y.Wu
*'' Rhamnus grubovii'' I.M.Turner
*'' Rhamnus hainanensis'' Merr. & Chun
*'' Rhamnus hemsleyana'' C.K.Schneid.
*'' Rhamnus heterophylla'' Oliv.
*'' Rhamnus hirtella'' Boiss.
*'' Rhamnus hupehensis'' C.K.Schneid.
* ''Rhamnus'' × ''hybrida'' L'Hér.
*'' Rhamnus imeretina'' J.R.Booth ex G.Kirchn.
*'' Rhamnus infectoria'' L.
*'' Rhamnus integrifolia'' DC. –
* ''Rhamnus'' × ''intermedia'' Steud. & Hochst.
*'' Rhamnus ishidae'' Miyabe & Kudô
*'' Rhamnus iteinophylla'' C.K.Schneid.
*'' Rhamnus japonica'' Maxim. – Japanese buckthorn
*'' Rhamnus kanagusukii'' Makino
*'' Rhamnus kayacikii'' Yalt. & P.H.Davis
*'' Rhamnus kurdica'' Boiss. & Hohen.
*'' Rhamnus kwangsiensis'' Y.L.Chen & P.K.Chou
*'' Rhamnus lamprophylla'' C.K.Schneid.
*'' Rhamnus lanceolata'' Pursh – lanceleaf buckthorn
*'' Rhamnus laoshanensis'' D.K.Zang
*'' Rhamnus ledermannii'' Lauterb.
* ''Rhamnus'' × ''lemaniana'' Briq.
*'' Rhamnus leptacantha'' C.K.Schneid.
*'' Rhamnus leptophylla'' C.K.Schneid.
*'' Rhamnus libanotica'' Boiss. – Lebanese buckthorn
*'' Rhamnus liboensis'' Y.F.Deng
*'' Rhamnus liukiuensis'' (E.H.Wilson) Koidz.
*'' Rhamnus lojaconoi'' Raimondo
*'' Rhamnus ludovici-salvatoris'' Chodat – endemic to Mallorca
*'' Rhamnus lycioides'' L. – black hawthorn, European buckthorn, or Mediterranean buckthorn
*'' Rhamnus maximovicziana'' J.J.Vassil.
* ''Rhamnus'' × ''mehreganii'' Alijanpoor & Khodayari
* ''Rhamnus'' × ''mercieri'' Briq.
*'' Rhamnus microcarpa'' Boiss.
*'' Rhamnus mildbraedii'' Engl.
*'' Rhamnus minnanensis'' K.M.Li
*'' Rhamnus mollis'' Merr.
*'' Rhamnus mongolica'' Y.Z.Zhao & L.Q.Zhao
* ''Rhamnus'' × ''mulleyana'' Fritsch
*'' Rhamnus myrtifolia'' Willk.
*'' Rhamnus nakaharae'' (Hayata) Hayata
*'' Rhamnus napalensis'' (Wall.) M.A.Lawson
*'' Rhamnus nigrescens'' Lauterb.
*'' Rhamnus nigricans'' Hand.-Mazz.
*'' Rhamnus ninglangensis'' Y.L.Chen
*'' Rhamnus nitida'' P.H.Davis
*'' Rhamnus oleoides'' L.
*'' Rhamnus orbiculata'' Bornm. – Orjen buckthorn
*'' Rhamnus papuana'' Lauterb.
*'' Rhamnus parvifolia'' Bunge
*'' Rhamnus pentapomica'' R.Parker
*'' Rhamnus persica'' Boiss. – Persian buckthorn (grows in Iran)
*'' Rhamnus persicifolia'' Moris
*'' Rhamnus petiolaris'' Boiss. & Balansa – (endemic to Sri Lanka)
*'' Rhamnus philippinensis'' C.B.Rob.
*'' Rhamnus pichleri'' C.K.Schneid. & Bornm.
*'' Rhamnus pilushanensis'' Y.C.Liu & C.M.Wang
* ''Rhamnus'' × ''pissjaukovae'' O.A.Popova
*''Rhamnus prinoides'' L'Hér. – shiny-leaf buckthorn
*'' Rhamnus procumbens'' Edgew.
*'' Rhamnus prunifolia'' Sm.
*'' Rhamnus pulogensis'' Merr.
*''Rhamnus pumila'' Turra
*'' Rhamnus punctata'' Boiss.
*'' Rhamnus purpurea'' Edgew.
*'' Rhamnus pyrella'' O.Schwarz
*'' Rhamnus qianweiensis'' Z.Y.Zhu
*'' Rhamnus rahiminejadii'' Alijanpoor & Assadi
*'' Rhamnus rhodopea'' Velen.
*'' Rhamnus rosei'' M.C.Johnst. & L.A.Johnst.
*'' Rhamnus rosthornii'' E.Pritz.
*'' Rhamnus rugulosa'' Hemsl.
*'' Rhamnus salixiophylla'' S.S.Ying
*'' Rhamnus sargentiana'' C.K.Schneid.
*'' Rhamnus saxatilis'' Jacq. – rock Buckthorn, Avignon buckthorn, Avignon berry, dyer's buckthorn (syn. ''R. infectoria'', ''R. infectorius'')
*'' Rhamnus schlechteri'' Lauterb.
*'' Rhamnus seravschanica'' (Kom.) Kamelin
*'' Rhamnus serpyllacea'' Greuter & Burdet
*'' Rhamnus serrata'' Humb. & Bonpl. ex Schult. – sawleaf buckthorn
*'' Rhamnus sibthorpiana'' Schult.
*'' Rhamnus smithii'' Greene – Smith's buckthorn
*'' Rhamnus songorica'' Gontsch.
*'' Rhamnus spathulifolia'' Fisch. & C.A.Mey.
*'' Rhamnus staddo'' A.Rich. – (syn. ''R. rhodesicus'')
*'' Rhamnus standleyana'' C.B.Wolf
*'' Rhamnus subapetala'' Merr.
*'' Rhamnus sumatrensis'' Ridl.
*'' Rhamnus sumbawana'' Lauterb.
*'' Rhamnus tangutica'' J.J.Vassil.
*'' Rhamnus taquetii'' (H.Lév. & Vaniot) H.Lév. – Jejudo buckthorn
*'' Rhamnus thymifolia'' Bornm.
*'' Rhamnus tonkinensis'' Pit.
*'' Rhamnus tortuosa'' Sommier & Levier
*'' Rhamnus triquetra'' (Wall.) Brandis
*'' Rhamnus tzekweiensis'' Y.L.Chen & P.K.Chou
*'' Rhamnus utilis'' Decne. – Chinese buckthorn
*'' Rhamnus velutina'' Boiss.
*'' Rhamnus virgata'' Roxb.
*'' Rhamnus wightii'' Wight & Arn. – Wight's buckthorn
*'' Rhamnus wilsonii'' C.K.Schneid.
* ''Rhamnus'' × ''woloszczakii'' Kárpáti
*'' Rhamnus wumingensis'' Y.L.Chen & P.K.Chou
*'' Rhamnus xizangensis'' Y.L.Chen & P.K.Chou
*'' Rhamnus yoshinoi'' Makino

Distribution

''Rhamnus'' has a nearly cosmopolitan distribution, with about 140 species which are native from temperate to tropical regions, the majority of species are from east Asia and North America, with a few species in Europe and Africa.

North American species include ''Rhamnus alnifolia'', alder-leaf buckthorn, occurring across the continent, and '' Rhamnus crocea'', hollyleaf buckthorn, in the west. Though not native to this region, ''Rhamnus cathartica'' can be found in North America.

Buckthorns may be confused with dogwoods, which share the curved leaf venation; indeed, "dogwood" is a local name for '' R. prinoides'' in southern Africa. The two plants are easy to distinguish by slowly pulling a leaf apart; dogwoods will exude thin, white latex strings, while buckthorns will not.

Invasive species

''Rhamnus cathartica'', the common buckthorn, is considered an invasive species in the United States and by many local jurisdictions and state governments, including Minnesota and Wisconsin.

The common buckthorn is well-adapted to spreading in Canada and the U.S. It is an efficient grower that does not need much sunlight and or fertile soil. Its seeds are hardy, as well as being able to grow and spread easily in a variety of environmental conditions. Also other animals prefer to leave buckthorns alone because their leaves are not appetizing and their fruits are toxic to some animals. Overall they are known to have a negative effect on their surrounding environment. For example, the European buckthorn is blamed for increased frog egg mortality from a chemical it releases. Other species, such as the ''Rhamnus alaternus'' also make chemicals that prevent other animals from consuming them.

Ecology

Some species are invasive outside their natural ranges. ''R. cathartica'' was introduced into the United States as a garden shrub and has become an invasive species in many areas there. It is a primary host of the soybean aphid (''Aphis glycines''), a pest for soybean farmers across the US. The aphids use the buckthorn as a host for the winter and then spread to nearby soybean fields in the spring. Italian buckthorn (''R. alaternus''), an evergreen species from the Mediterranean region, has become a serious weed in some parts of New Zealand, especially on Hauraki Gulfe of many Lepidoptera species.

The American species are known to be hosts for the oat fungus ''Puccinia coronata''. In a 1930 study, both kerosene and salt were employed for eradication of ''R. lanceolata'' and both proved to be less expensive than felling these bushes.

Uses

The fruit of most species contain a yellow dye and the seeds are rich in protein. Oils from the seeds are used for making lubricating oil, printing ink, and soap. Many species have been used to make dyes. '' R. utilis'' provides china green, a dye used to give a bright green color to silk and wool. The bark of some species of ''Rhamnus'' afford a beautiful yellow dye. Another species, Avignon buckthorn ('' R. saxatilis'') produces a yellow dye, made from the fruit Persian berry.

Some species may cause demyelinating polyneuropathies.

The purging buckthorn ('' R. cathartica'') is a widespread European native species used in the past as a purgative. It was in mid 17th-century England the only native purgative. It was also known pre-Linnaeus as ''Spina Cervina''. The berries of ''Spina Cervina'' are black and contain a greenish juice, along with four seeds apiece; this serves to distinguish them from those of the black alder and dogberry, which contain only one or two apiece. Its syrup is said to be churlish. William Lewis
"An Experimental History Of The Materia Medica"
/ref> Its toxicity makes this a very risky herbal medicine, and it is no longer in use.Plants for a Future
''Rhamnus cathartica''
/ref>

''Rhamnus prinoides'' is known as ''gesho'' in Ethiopia, where it is used to make a mead called '' tej''.

The species ''Rhamnus alaternus'' shows some promise for medicinal use as well.

See also

* ''Sea buckthorn'' or Hippophae, an unrelated genus of shrubs with a similar common name
* ''Frangula'', a genus formerly included in ''Rhamnus''

References

External links

Scientific chemical research on African ''Rhamnus''

{{Authority control

Rhamnaceae genera
Butterfly food plants
Laxatives
Taxa named by Carl Linnaeus
Dioecious plants