Euphorbia

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Euphorbia
Euphorbia February 2008-2.jpg
Euphorbia serrata
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Malpighiales
Family: Euphorbiaceae
Subfamily: Euphorbioideae
Tribe: Euphorbieae
Subtribe: Euphorbiinae
Griseb.
Genus: Euphorbia
L.
Type species
Euphorbia antiquorum
L.
Subgenera

Chamaesyce
Esula
Euphorbia
Rhizanthium
and see below

Contents

Diversity
c.2008 species
Synonyms [1]
List
    • AdenopetalumKlotzsch & Garcke
    • AdenorimaRaf.
    • AgalomaRaf.
    • AklemaRaf.
    • AlectoroctonumSchltdl.
    • AllobiaRaf.
    • AnisophyllumHaw.
    • AnthacanthaLem.
    • AplarinaRaf.
    • ArthrothamnusKlotzsch & Garcke
    • AthymalusNeck.
    • ChamaesyceGray
    • CharaciasGray
    • ChylogalaFourr.
    • CrepidariaHaw.
    • CtenadenaProkh.
    • CubanthusMillsp.
    • CyathophoraRaf.
    • CystidospermumProkh.
    • DactylanthesHaw.
    • DematraRaf.
    • DesmonemaRaf.
    • DiadenariaKlotzsch & Garcke
    • DichyliumBritton
    • DiplocyathiumHeinr.Schmidt
    • DitritraRaf.
    • ElaeophorbiaStapf
    • EndadeniumL.C.Leach
    • EndoisilaRaf.
    • EpurgaFourr.
    • Esula(Pers.) Haw.
    • EumecanthusKlotzsch & Garcke
    • EuphorbiastrumKlotzsch & Garcke
    • EuphorbiodendronMillsp.
    • EuphorbionSt.-Lag.
    • EuphorbiumHill
    • GalarhoeusHaw.
    • GalorhoeusEndl.
    • HexadeniaKlotzsch & Garcke
    • KanopikonRaf.
    • KeraselmaNeck. ex Juss.
    • KobiosisRaf.
    • LacanthisRaf.
    • LepadenaRaf.
    • LeptopusKlotzsch & Garcke
    • LophobiosRaf.
    • LortiaRendle
    • Lyciopsis(Boiss.) Schweinf.
    • MeduseaHaw.
    • MonadeniumPax
    • MurtekiasRaf.
    • NisomenesRaf.
    • PeccanaRaf.
    • PedilanthusNeck. ex Poit.
    • PetalomaRaf. ex Boiss.
    • PleuradenaRaf.
    • PoinsettiaGraham
    • PythiusRaf.
    • SclerocyathiumProkh.
    • StenadeniumPax
    • SterigmantheKlotzsch & Garcke
    • SynadeniumBoiss.
    • TirucaliaRaf.
    • TithymalodesLudw. ex Kuntze
    • TithymaloidesOrtega
    • TithymalopsisKlotzsch & Garcke
    • TithymalusGaertn.
    • TithymalusSég.
    • TithymalusMill.
    • TorfasadisRaf.
    • TreisiaHaw.
    • TrichosterigmaKlotzsch & Garcke
    • TumalisRaf.
    • VallarisRaf.
    • VentenatiaTratt.
    • XamesikeRaf.
    • ZaliteaRaf.
    • ZygophyllidiumSmall
Euphorbia as a small tree: Euphorbia dendroides Euphorbia dendroides3-Corsica~2011 04 30.jpg
Euphorbia as a small tree: Euphorbia dendroides

Euphorbia is a very large and diverse genus of flowering plants, commonly called spurge, in the family Euphorbiaceae. "Euphorbia" is sometimes used in ordinary English to collectively refer to all members of Euphorbiaceae (in deference to the type genus), not just to members of the genus. [2]

Euphorbias range from tiny annual plants to large and long-lived trees. [3] with perhaps the tallest being Euphorbia ampliphylla at 30 m (98 ft) or more. [4] [5] The genus has roughly 2,000 members, [6] [7] making it one of the largest genera of flowering plants. [8] [9] It also has one of the largest ranges of chromosome counts, along with Rumex and Senecio . [8] Euphorbia antiquorum is the type species for the genus Euphorbia. [10] It was first described by Carl Linnaeus in 1753 in Species Plantarum .

Some euphorbias are widely available commercially, such as poinsettias at Christmas. Some are commonly cultivated as ornamentals, or collected and highly valued for the aesthetic appearance of their unique floral structures, such as the crown of thorns plant ( Euphorbia milii ). Succulent euphorbias from the deserts of Southern Africa and Madagascar have evolved physical characteristics and forms similar to cacti of North and South America, so they are often incorrectly referred to as cacti. [11] Some are used as ornamentals in landscaping, because of beautiful or striking overall forms, and drought and heat tolerance. [7] [3]

Euphorbia all share the feature of having a poisonous, latex-like sap and unique floral structures. [7] When viewed as a whole, the head of flowers looks like a single flower (a pseudanthium). [7] It has a unique kind of pseudanthium, called a cyathium, where each flower in the head is reduced to its barest essential part needed for sexual reproduction. [7] The individual flowers are either male or female, with the male flowers reduced to only the stamen, and the females to the pistil. [7] These flowers have no sepals, petals, or other parts that are typical of flowers in other kinds of plants. [7] Structures supporting the flower head and other structures underneath have evolved to attract pollinators with nectar, and with shapes and colors that function in a way petals and other flower parts do in other flowers. It is the only genus of plants that has all three kinds of photosynthesis, CAM, C3 and C4. [7]

Etymology

The common name "spurge" derives from the Middle English/Old French espurge ("to purge"), due to the use of the plant's sap as a purgative. The botanical name Euphorbia derives from Euphorbos, the Greek physician of King Juba II of Numidia and Mauretania (52–50 BC – 23 AD), who married the daughter of Anthony and Cleopatra. [12] Juba was a prolific writer on various subjects, including natural history. Euphorbos wrote that one of the cactus-like euphorbias (now called Euphorbia obtusifolia ssp. regis-jubae ) was used as a powerful laxative. [12] In 12 BC, Juba named this plant after his physician Euphorbos, as Augustus Caesar had dedicated a statue to the brother of Euphorbos, Antonius Musa, who was the personal physician of Augustus. [12] In 1753, botanist and taxonomist Carl Linnaeus assigned the name Euphorbia to the entire genus in the physician's honor. [13]

Description

The plants are annual, biennial or perennial herbs, woody shrubs, or trees with a caustic, poisonous milky latex. The roots are fine or thick and fleshy or tuberous. Many species are more or less succulent, thorny, or unarmed. The main stem and mostly also the side arms of the succulent species are thick and fleshy, and often winged, 15–91 cm (6–36 in) tall. The succulent trees and large shrubs are mostly confined to southern and eastern Africa, but Euphorbia neutra is native to the Caatinga of Brazil and Euphorbia royleana is found in the Himalayan foothills. [14] The deciduous[ citation needed ] leaves may be opposite, alternate, or in whorls. In succulent species, the leaves are mostly small and short-lived. The stipules are mostly small, partly transformed into spines or glands, or missing.

Inflorescence and fruit

Euphorbia false-flower EUPHORBIA.JPG
Euphorbia false-flower

Like all members of the family Euphorbiaceae, spurges have unisexual flowers.

In Euphorbia, flowers occur in a head, called the cyathium (plural cyathia). Each male or female flower in the cyathium head has only its essential sexual part, in males the stamen, and in females the pistil. The flowers do not have sepals, petals, or nectar to attract pollinators, although other nonflower parts of the plant have an appearance and nectar glands with similar roles. Euphorbias are the only plants known to have this kind of flower head. [15]

Nectar glands and nectar that attract pollinators are held in the involucre, a cup-like part below and supporting the cyathium head. The "involucre" in the genus Euphorbia is not to be confused with the "involucre" in family Asteraceae members, which is a collection of bracts called phyllaries, which surround and encase the unopened flower head, then support the receptacle under it after the flower head opens.

The involucre is above and supported by bract-like modified leaf structures (usually in pairs)[ citation needed ] called cyathophylls', or cyathial leaves. The cyathophyll often has a superficial appearance of being petals of a flower.

Euphorbia flowers are tiny, and the variation attracting different pollinators, with different forms and colors occurs, in the cyathium, involucre, cyathophyll, or additional parts such as glands that attached to these.

The collection of many flowers may be shaped and arranged to appear collectively as a single individual flower, sometimes called a pseudanthium in the Asteraceae, and also in Euphorbia.

The majority of species are monoecious (bearing male and female flowers on the same plant), although some are dioecious with male and female flowers occurring on different plants. It is not unusual for the central cyathia of a cyme to be purely male, and for lateral cyathia to carry both sexes. Sometimes, young plants or those growing under unfavorable conditions are male only, and only produce female flowers in the cyathia with maturity or as growing conditions improve.[ citation needed ]

The female flowers reduced to a single pistil usually split into three parts, often with two stigmas at each tip.[ citation needed ] Male flowers often have anthers in twos.[ citation needed ] Nectar glands usually occur in fives, [16] may be as few as one, [16] and may be fused into a "U" shape. [15] The cyathophylls often occur in twos, are leaf-like, and may be showy and brightly coloured and attractive to pollinators, or be reduced to barely visible tiny scales.[ citation needed ]

The fruits are three- or rarely two-compartment capsules, sometimes fleshy, but almost always ripening to a woody container that then splits open, sometimes explosively. The seeds are four-angled, oval, or spherical, and some species have a caruncle.[ citation needed ]

Xerophytes and succulents

In the genus Euphorbia, succulence in the species has often evolved divergently and to differing degrees. Sometimes, it is difficult to decide, and is a question of interpretation, whether or not a species is really succulent or "only" xerophytic. In some cases, especially with geophytes, plants closely related to the succulents are normal herbs. About 850 species are succulent in the strictest sense. If one includes slightly succulent and xerophytic species, this figure rises to about 1000, representing about 45% of all Euphorbia species.

Irritants

The milky sap of spurges (called "latex") evolved as a deterrent to herbivores. It is white, and transparent when dry, except in E. abdelkuri , where it is yellow. The pressurized sap seeps from the slightest wound and congeals after a few minutes in air. The skin-irritating and caustic effects are largely caused by varying amounts of diterpenes. Triterpenes such as betulin and corresponding esters are other major components of the latex. [17] In contact with mucous membranes (eyes, nose, mouth), the latex can produce extremely painful inflammation. The sap has also been known to cause mild to extreme Keratouveitis, which affects vision. [18] Therefore, spurges should be handled with caution and kept away from children and pets. Wearing eye protection while working in close contact with Euphorbia is advised. [18] Latex on skin should be washed off immediately and thoroughly. Congealed latex is insoluble in water, but can be removed with an emulsifier such as milk or soap. A physician should be consulted if inflammation occurs, as severe eye damage including permanent blindness may result from exposure to the sap. [19]

The poisonous qualities were well known: in the Ethiopian Kebra Nagast , the serpent king Arwe is killed with juice from the Euphorbia. [20]

Uses

Detail of poinsettia flowers and immature fruits Pointsetta flowers.jpg
Detail of poinsettia flowers and immature fruits
An old Euphorbia hybrid Euphorbia hybrid 11.jpg
An old Euphorbia hybrid
Euphorbia obesa Euphorbia obesa 222.JPG
Euphorbia obesa

Several spurges are grown as garden plants, among them poinsettia (E. pulcherrima) and the succulent E. trigona . E. pekinensis (Chinese : 大戟 ; pinyin :dàjǐ) is used in traditional Chinese medicine, where it is regarded as one of the 50 fundamental herbs. Several Euphorbia species are used as food plants by the larvae of some Lepidoptera (butterflies and moths), like the spurge hawkmoths ( Hyles euphorbiae and Hyles tithymali ), as well as the giant leopard moth.

Ingenol mebutate, a drug used to treat actinic keratosis, is a diterpenoid found in Euphorbia peplus .

Euphorbias are often used as hedging plants in many parts of Africa. [21]

Misidentification as cacti

Euphorbia milii Euphorbia milii - flower view01.jpg
Euphorbia milii

Among laypeople, Euphorbia species are among the plant taxa most commonly confused with cacti, especially the stem succulents. [22] Euphorbias secrete a sticky, milky-white fluid with latex, but cacti do not. [22] Individual flowers of euphorbias are usually tiny and nondescript (although structures around the individual flowers may not be), without petals and sepals, unlike cacti, which often have fantastically showy flowers. [22] Euphorbias from desert habitats with growth forms similar to cacti have thorns, which are different from the spines of cacti. [22]

Systematics and taxonomy

The present taxon "Euphorbia" corresponds to its own former subtribe, the Euphorbiinae.[ citation needed ] It has over 2000 species. [7] Morphological description using the presence of a cyathium (see section above) is consistent with nuclear and chloroplast DNA sequence data in testing of about 10% of its members. This testing supports inclusion of formerly other genera as being best placed in this single genus, including Chamaesyce, Monadenium, Pedilanthus, and poinsettia (E. pulcherrima).

Genetic tests have shown that similar flower head structures or forms within the genus, might not mean close ancestry within the genus. The genetic data show that within the genus, convergent evolution of inflorescence structures may be from ancestral subunits that are not related. So using morphology within the genus becomes problematic for further subgeneric grouping. As stated on the Euphorbia Planetary Biodiversity Inventory project webpage: [7]

Previous morphologically based delimitations of subgenera or sections within the genus should not be taken at face value. The genus is in fact rife with striking examples of morphological convergence in cyathial and vegetative features, which justifies a global approach to studying the genus to obtain a proper phylogenetic understanding of the whole group.... The bottom line is that a number of clades have been placed inside or outside of Euphorbia at different times... few of the subgeneric circumscriptions hold up under DNA sequence analysis.

According to a 2002 publication on studies of DNA sequence data, [23] [24] [25] most of the smaller "satellite genera" around the huge genus Euphorbia nest deep within the latter. Consequently, these taxa, namely the never generally accepted genus Chamaesyce, as well as the smaller genera Cubanthus , [26] Elaeophorbia, Endadenium, Monadenium, Synadenium, and Pedilanthus were transferred to Euphorbia. The entire subtribe Euphorbiinae now consists solely of the genus Euphorbia.

Selected species

See List of Euphorbia species for complete list.

Hybrids

Euphorbia has been extensively hybridised for garden use, with many cultivars available commercially. Moreover, some hybrid plants have been found growing in the wild, for instance E. × martini Rouy, [32] a cross of E. amygdaloides × E. characias subsp. characias, found in southern France.

Subgenera

Simplified diagram of relations in subtribe Euphorbiinae, 2006 Euphorbiinae tree.png
Simplified diagram of relations in subtribe Euphorbiinae, 2006

The genus Euphorbia is one of the largest and most complex genera of flowering plants, and several botanists have made unsuccessful attempts to subdivide the genus into numerous smaller genera. According to the recent phylogenetic studies, [23] [24] [25] Euphorbia can be divided into four subgenera, each containing several sections and groups. Of these, subgenus Esula is the most basal. The subgenera Chamaesyce and Euphorbia are probably sister taxa, but very closely related to subgenus Rhizanthium. Extensive xeromorph adaptations in all probability evolved several times; [33] it is not known if the common ancestor of the cactus-like Rhizanthium and Euphorbia lineages had been xeromorphic—in which case a more normal morphology would have re-evolved namely in Chamaesyce—or whether extensive xeromorphism is entirely polyphyletic even to the level of the subgenera.

See also

Related Research Articles

<span class="mw-page-title-main">Poinsettia</span> Species of flowering plant in the spurge family Euphorbiaceae

The poinsettia is a commercially important flowering plant species of the diverse spurge family Euphorbiaceae. Indigenous to Mexico and Central America, the poinsettia was first described by Europeans in 1834. It is particularly well known for its red and green foliage and is widely used in Christmas floral displays. It derives its common English name from Joel Roberts Poinsett, the first United States minister to Mexico, who is credited with introducing the plant to the US in the 1820s. Poinsettias are shrubs or small trees, with heights of 0.6 to 4 m. Though often stated to be highly toxic, the poinsettia is not dangerous to pets or children. Exposure to the plant, even consumption, most often results in no effect, though it can cause nausea, vomiting, or diarrhea.

<i>Euphorbia heterophylla</i> Species of plant in the spurge family, Euphorbiaceae, native to tropical and subtropical America

Euphorbia heterophylla, also known under the common names of Mexican fireplant, painted euphorbia, Japanese poinsettia, paintedleaf, painted spurge and milkweed, is a plant belonging to the Euphorbiaceae or spurge family.

<i>Chamaesyce</i> Genus of flowering plants

Chamaesyce is a genus of plants in the family Euphorbiaceae. Recent phylogenetic studies have shown that Chamaesyce is deeply nested within the broader Euphorbia. Specifically, Chamaesyce is very closely related to plants like Euphorbia pulcherrima, the popular poinsettia. Currently, all species have now been reclassified as species of Euphorbia. Specifically, this group now belongs to Euphorbia subgenus Chamaesyce section Anisophyllum. Taxonomically speaking, Chamaesyce is considered a synonym of Euphorbia.

<i>Euphorbia tithymaloides</i> Species of plant

Euphorbia tithymaloides is a perennial succulent spurge. An erect shrub, the plant is also known by the scientific name Pedilanthus tithymaloides. However, the genus Pedilanthus has been submerged into the genus Euphorbia, and is more correctly known by its new name.

<i>Euphorbia milii</i> Species of plant

Euphorbia milii, the crown of thorns, Christ plant, or Christ's thorn, is a species of flowering plant in the spurge family Euphorbiaceae, native to Madagascar. The species name commemorates Baron Milius, once Governor of Réunion, who introduced the species to France in 1821.

<i>Euphorbia maculata</i> Species of plant

Euphorbia maculata, known as spotted spurge, prostrate spurge, milk purslane, or spotted sandmat, is a fast-growing annual plant in the family Euphorbiaceae. While it is native to North America, where it is a common garden and lawn weed in the United States, it has become a common introduced species throughout the world, including Europe, Japan, Korea, Australia, and New Zealand.

<span class="mw-page-title-main">Cyathium</span> A form of inflorescence of the type found on the poinsettia

A cyathium is one of the specialised pseudanthia forming the inflorescence of plants in the genus Euphorbia (Euphorbiaceae). A cyathium consists of:

<i>Astrophytum asterias</i> Species of cactus

Astrophytum asterias is a species of cactus in the genus Astrophytum, and is native to small parts of Texas in the United States and Mexico. Common names include sand dollar cactus, sea urchin cactus, star cactus and star peyote.

<i>Euphorbia misera</i> Species of flowering plant

Euphorbia misera is a semi-succulent shrub in the genus Euphorbia commonly known as the cliff spurge or coast spurge. A drought-deciduous shrub, it is typically found as a gnarled, straggly plant occupying seashore bluffs, hills and deserts. Like other members of its genus, it has a milky sap, which can be found exuding out of the light gray bark when damaged. The alternately-arranged leaves are round and folded in the middle, with small hairs on them. The "flowers" can be found blooming year-round, and are colored maroon or yellow in the center with 5 white to light-yellow petal-like appendages attached outside. This species is native to the Baja California peninsula and Sonora in Mexico, and the coast of southern California in the United States, where it is a rare species. It is threatened in some localities by the development of its coastal habitat, which tends to be prime locations for high-end residential and commercial developments.

<i>Euphorbia canariensis</i> Species of flowering plant in the spurge family Euphorbiaceae

Euphorbia canariensis, commonly known as the Canary Island spurge, Hercules club or in Spanish cardón, is a succulent member of the genus Euphorbia and family Euphorbiaceae endemic to the Canary Islands. It is the plant symbol of the island of Gran Canaria.

<span class="mw-page-title-main">Euphorbiaceae</span> Family of Eudicot flowering plants

Euphorbiaceae, the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of the type genus of the family. Most spurges, such as Euphorbia paralias, are herbs, but some, especially in the tropics, are shrubs or trees, such as Hevea brasiliensis. Some, such as Euphorbia canariensis, are succulent and resemble cacti because of convergent evolution. This family has a cosmopolitan global distribution. The greatest diversity of species is in the tropics; however, the Euphorbiaceae also have many species in nontropical areas of all continents except Antarctica.

<i>Euphorbia micromera</i> Species of plant in the family Euphorbiaceae

Euphorbia micromera is a species of flowering plant in the family Euphorbiaceae. It is known by the common name Sonoran sandmat. It is native to the southwestern United States from California to Texas, and northern Mexico, where it grows in sandy soils in desert and other dry habitat. It is an annual herb forming a small mat of slender stems. The hairy to hairless leaves are oblong in shape and just a few millimeters long. The tiny inflorescence is a cyathium less than a millimeter wide. It lacks the appendages that many similar species have in their cyathia. It has only a central female flower and 2 to 5 male flowers surrounded by round red nectar glands. The fruit is a minute round capsule.

<i>Euphorbia revoluta</i> Species of flowering plant

Euphorbia revoluta is a species of euphorb known by the common name threadstem sandmat. It is native to Mexico and the southwestern United States from California to the Rocky Mountains. It is an annual herb producing thin, erect stems with pairs of linear leaves, each leaf up to 2.6 centimeters long. The inflorescence is a cyathium with rounded nectar glands surrounding one female flower and several male flowers. There are sometimes white petal-like appendages as well. The Navajo used this plant as a skin lotion.

<i>Euphorbia balsamifera</i> Species of flowering plant in the spurge family Euphorbiaceae

Euphorbia balsamifera is a flowering plant in the spurge family Euphorbiaceae. It is distributed in the Canary Islands and the western Sahara. It is the vegetable symbol of the island of Lanzarote. Euphorbia adenensis has been treated as a subspecies of this species.

<span class="mw-page-title-main">Huntington Desert Garden</span>

The Huntington Desert Garden is part of The Huntington Library, Art Collections and Botanical Gardens in San Marino, California. The Desert Garden is one of the world's largest and oldest collections of cacti, succulents and other desert plants, collected from throughout the world. It contains plants from extreme environments, many of which were acquired by Henry E. Huntington and William Hertrich in trips taken to several countries in North, Central and South America. One of the Huntington's most botanically important gardens, the Desert Garden brought together a group of plants largely unknown and unappreciated in the beginning of the 1900s. Containing a broad category of xerophytes, the Desert Garden grew to preeminence and remains today among the world's finest, with more than 5,000 species in the 10 acre garden.

<i>Euphorbia celastroides</i> Species of plant in the family Euphorbiaceae

Euphorbia celastroides is a flowering plant in the family Euphorbiaceae. It is referred to by the common name 'akoko by Hawaiians, and is a species of spurge closely related to the poinsettia. This species develops into a round-shape shrub. This species is endemic to the Hawaiian Islands.

<i>Euphorbia royleana</i> Species of plant in the family Euphorbiaceae

Euphorbia royleana is a species of flowering plant in the family Euphorbiaceae. It is also known as Sullu spurge, and Royle's spurge. It is a succulent and almost cactus like in appearance although unrelated. It grows right across the Himalaya mountains from Pakistan, India, Bhutan, Myanmar, Nepal to western China, It prefers dry and rocky slopes between 1000 and 1500 meters, but has been found up to 2000 meters. Flowering and fruiting is in spring to early summer (March–July) and seeding is in June–October. It is used as a hedging plant in northern India and has medicinal uses.

<i>Euphorbia missurica</i> Species of flowering plant

Euphorbia missurica, commonly called prairie sandmat, or Missouri spurge, is a species of flowering plant in the spurge family (Euphorbiaceae). It is native to North America, where it is found primarily in area of the Great Plains. Its natural habitat is in dry, often calcareous areas, including glades, bluffs, and open woodlands.

<i>Euphorbia bicolor</i> Species of plant

Euphorbia bicolor, commonly known as snow on the prairie, is a species of flowering plant in the genus Euphorbia, native to the southern United States. It grows 1-4 feet tall, has green and white alternate leaves, and is monoecious with unisexual flowers. It grows in hard clay soils of prairies, rangelands, and edges of forests.

<i>Euphorbia nivulia</i> Species of succulent flowering plant in the family Euphorbiaceae

Euphorbia nivulia is a subtropical succulent species of flowering plant in the family Euphorbiaceae. It is found in the Indian subcontinent and is commonly known as the leafy milk hedge, holy milk hedge or dog's tongue.

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Further reading