Thepeony orpaeony (/ˈpiːəni/)[2][3] is anyflowering plant in thegenusPaeonia,[4] the only genus in thefamilyPaeoniaceae. Peonies are native toAsia,Europe, and WesternNorth America.[5] Scientists differ on the number ofspecies that can be distinguished, ranging from 25 to 40,[6][7] although the current consensusdescribes 33 known species.[8] The relationships between the species need to be further clarified.[9]
Most areherbaceousperennial plants 0.25–1 metre (1–3 ft) tall, but some are woodyshrubs 0.25–3.5 metres (1–11 ft) tall. They havecompound, deeply lobed leaves and large, often fragrant flowers, in colors ranging from purple and pink to red, white or yellow, in late spring and early summer. The flowers have a short blooming season, usually lasting for only 7–10 days.
Peonies are popular garden plants in temperate regions. Herbaceous peonies are also sold ascut flowers on a large scale, although they generally are only available in late spring and early summer.[10]
All Paeoniaceae areherbaceousperennials ordeciduous shrubs, with thick storage roots and thin roots for gathering water and minerals. Some species arecaespitose (tufted), because the crown produces adventitious buds, while others havestolons. They have rather large compound leaves withoutglands andstipules, and with anomocyticstomata. In the woody species the new growth emerges from scaly buds on the previous flush or from the crown of the rootstock. The largebisexual flowers are mostly single at the end of the stem. InP. emodi,P. lactiflora,P. veitchii and many of the cultivars these contributed to, few additional flowers develop in the axils of the leaves. Flowers close at night or when the sky is overcast. Each flower is subtended by a number of bracts, that may form a sort ofinvolucre, has 3-7 tough free sepals and mostly 5–8, but occasionally up to 13 free petals. These categories however are intergrading, making it difficult to assign some of them, and the number of these parts may vary. Within are numerous (50–160) freestamens, withanthers fixed at their base to the filaments, and are sagittate in shape, open with longitudinal slitsat the outer side and freepollen grains which have three slits or pores and consist of two cells. Within the circle of stamens is a more or less prominent, lobeddisc, which is presumed not to excretenectar. Within the disk is a varying number (1–15) of separatecarpels, which have a very shortstyle and a decurrentstigma. Each of these develops into a dry fruit (which is called afollicle), which opens with a lengthwise suture and each of which contains one or a few large fleshy seeds. The annual growth is predetermined: if the growing tip of a shoot is removed, no new buds will develop that season.[10][11][12]
The basic chromosome number is five. About half of the species of the sectionPaeonia however is tetraploid (4n=20), particularly many of those in the Mediterranean region. Both allotetraploids and autotetraploids are known, and some diploid species are also of hybrid origin.[11]
The family name "Paeoniaceae" was first used by Friedrich K.L. Rudolphi in 1830, following a suggestion byFriedrich Gottlieb Bartling that same year.[6] The family had been given other names a few years earlier.[14] The composition of the family has varied, but it has always consisted ofPaeonia and one or more genera that are now placed inRanunculales.[7] It has been widely believed thatPaeonia is closest toGlaucidium, and this idea has been followed in some recent works.[6][15]Molecular phylogenetic studies, however, have demonstrated conclusively thatGlaucidium belongs in the family Ranunculaceae, order Ranunculales,[16] but thatPaeonia belongs in the unrelatedorderSaxifragales.[17] The genusPaeonia consists of about 35 species, assigned to three sections:Moutan,Onaepia andPaeoniae. The sectionOnaepia only includesP. brownii andP. californica. The sectionMoutan is divided intoP. delavayi andP. ludlowii, together making up the subsectionDelavayanae, andP. cathayana,P. decomposita,P. jishanensis,P. osti,P. qiui andP. rockii which constitute the subsectionVaginatae.P. suffruticosa is a cultivated hybrid swarm, not a naturally occurring species.[18]
The remainder of the species belongs to the sectionPaeonia, which is characterised by a complicatedreticulate evolution. Only about half of the (sub)species is diploid, the other half tetraploid, while some species both have diploid and tetraploid populations. In addition to the tetraploids, are some diploid species also likely the result of hybridisation, ornothospecies. Known diploid taxa in thePaeonia-section areP. anomala,P. lactiflora,P. veitchii,P. tenuifolia,P. emodi,P. broteri,P. cambedessedesii,P. clusii,P. rhodia,P. daurica subsps.coriifolia,daurica,macrophylla andmlokosewitschii. Tetraploid taxa areP. arietina,P. officinalis,P. parnassica,P. banatica,P. russi,P. peregrina,P. coriacea,P. mascula subsps.hellenica andmascula, andP. daurica subsps.tomentosa andwittmanniana. Species that have both diploid and tetraploid populations includeP. clusii,P. mairei andP. obovata.P. anomala was proven to be a hybrid ofP. lactiflora andP. veitchii, although being a diploid with 10 chromosomes.P. emodi andP. sterniana are diploid hybrids ofP. lactiflora andP. veitchii too, and radically different in appearance.P. russi is the tetraploid hybrid of diploidP. lactiflora andP. mairei, whileP. cambedessedesii is the diploid hybrid ofP. lactiflora, likelyP. mairei, but possibly alsoP. obovata.P. peregrina is the tetraploid hybrid ofP. anomala and eitherP. arietina,P. humilis,P. officinalis,P. parnassica or less likelyP. tenuifolia, or one of their (now extinct) common ancestors.P. banatica is the tetraploid hybrid ofP. mairei and one of this same group.P. broteri,P. coriacea,P. clusii,P. rhodia,P. daurica subsp.mlokosewitschi,P. mascula subsp.hellenica and ssp.mascula, andP. daurica subsp.wittmanniana are all descendants of hybrids ofP. lactiflora andP. obovata.[18][19]
Recent genetic analyses relate themonogeneric family Paeoniaceae to a group of families with woody species in the orderSaxifragales. This results in the following relationship tree.[17] One dissertation suggests the sectionOnaepia branches off earliest, but a later publication of the same author and others suggests theMoutan-section splits off first. Within that sectionP. ludlowii andP. delavayi are more related to each other than to any other species.[20][21]
The genusPaeonia naturally occurs in the temperate and cold areas of the Northern Hemisphere. The sectionMoutan, which includes all woody species, is restricted in the wild to Central and Southern China, including Tibet. The sectionOnaepia consist of two herbaceous species and is present in the West of North-America,P. brownii between southern British Columbia and theSierra Nevada in California and eastward to Wyoming and Utah, whileP. californica is limited to the coastal mountains of Southern and Central California.
The sectionPaeonia, which comprises all other herbaceous species, occurs in a band stretching roughly from Morocco and Spain to Japan. One species of the sectionPaeonia,P. anomala, has by far the largest distribution, which is also north of the distribution of the other species: from theKola peninsula in North-West Russia, toLake Baikal in Siberia and South to theTien Shan Mountains of Kazakhstan. The rest of the section concentrates around the Mediterranean, and in Asia.
The species around the Mediterranean includePaeonia algeriensis that is anendemic of the coastal mountains of Algeria,P. coriacea in theRif Mountains andAndalusia,P. cambessedesii onMajorca,P. russoi onCorsica,Sardinia andSicily,P. corsica on Corsica, Sardinia, theIonian islands and in western Greece,P. clusii subsp.clusii onCrete andKarpathos, and subsp.rhodia onRhodes,P. kesrouanensis in the WesternTaurus Mountains,P. arietina from the Middle Taurus Mountains,P. broteri in Andalucia,P. humilis from Andalucia to theProvence,P. officinalis from the South of France, through Switzerland to the Middle of Italy,P. banatica in western Romania, northern Serbia and Slovenia and in southern Hungary,P. peregrina in Albania, western Bulgaria, northern Greece, western Romania, Serbia, Montenegro and Bosnia, whileP. mascula has a large distribution fromCatalonia and southern France to Israel and Turkey.
Between the two concentrations, the subspecies ofPaeonia daurica occur, with subspeciesvelebitensis in Croatia, anddaurica in the Balkans and Crimea, while the other subspeciescoriifolia,macrophylla,mlokosewitschii,tomentosa andwittmanniana are known from theCaucasus,Kaçkar andAlborz Mountains.
Paeonia emodi occurs in the westernHimalayas between Pakistan and western Nepal,P. sterniana is an endemic of southeastern Tibet,P. veitchii grows in Central China (Qinghai,Ningxia,Gansu,Shaanxi,Shanxi,Sichuan and the eastern rim of Tibet), likeP. mairei (Gansu, Guizhou, Hubei, Shaanxi, Sichuan, and Yunnan), whileP. obovata grows in warm-temperate to cold China, includingManchuria, Korea, Japan, Far Eastern Russia (Primorsky Krai) and onSakhalin, andP. lactiflora occurs in Northern China, including Manchuria, Japan, Korea, Mongolia, Russia (Far East and Siberia).[19]
The species of the sectionPaeonia have adisjunct distribution, with most of the species occurring in the Mediterranean, while many others occur in eastern Asia. Genetic analysis has shown that all Mediterranean species are eitherdiploid or tetraploid hybrids that resulted from thecrossbreeding of species currently limited to eastern Asia. The large distance between the ranges of the parent species and thenothospecies suggest that hybridisation already occurred relatively long ago. It is likely that the parent species occurred in the same region when the hybrids arose, and were later exterminated by successive Pleistocene glaciations, while the nothospecies remained in refugia to the South of Europe. During their retreat,P. lactiflora andP. mairei likely becamesympatric and so produced the Himalayan nothospeciesP. emodi andP. sterniana.[19]
Distribution maps of the species ofPaeonia in Europe and Western Asia
Ancient Chinese texts mention the peony was used for flavoring food. Peonies have been used and cultivated in China since early history. Ornamental cultivars were created from plants cultivated for medicine in China as of the sixth and seventh century. Peonies became particularly popular during theTang dynasty, when they were grown in the imperial gardens. In the tenth century the cultivation of peonies spread through China, and the seat of theSong dynasty,Luoyang, was the centre for its cultivation, a position it still holds today.
A second centre for peony cultivation developed during theQing dynasty in Cáozhōu, now known asHe Ze. Both cities still host annual peony exhibitions and state-funded peony research facilities. Before the tenth century,P. lactiflora was introduced in Japan, and over time many varieties were developed both byself fertilisation andcrossbreeding, particularly during the eighteenth to twentieth centuries (middleEdo to earlyShōwa periods). During the 1940s Toichi Itoh succeeded in crossing tree peonies and herbaceous peonies and so created a new class of so-calledintersectional hybrids. AlthoughP. officinalis and its cultivars were grown in Europe from the fifteenth century on, originally also for medicinal purposes, intensive breeding started only in the nineteenth century whenP. lactiflora was introduced from its native China to Europe. The tree peony was introduced in Europe and planted inKew Gardens in 1789. The main centre of peony breeding in Europe has been in the United Kingdom, and particularly France. Here, breeders likeVictor Lemoine and François Félix Crousse selected many new varieties, mainly withP. lactiflora, such as "Avant Garde" and "Le Printemps". The Netherlands is the largest peony cut flower producing country with about 50 million stems each year, with "Sarah Bernhardt" dominating the sales with over 20 million stems.[10] An emerging source of peonies in mid to late summer is the Alaskan market. Unique growing conditions due to long hours of sunlight create availability from Alaska when other sources have completed harvest.[22]
While the peony takes several years to re-establish itself when moved, it blooms annually for decades once it has done so.[23]
Ants on a bud
Peonies tend to attractants to the flower buds. This is due to the nectar that forms on the outside of the flower buds, and is not required for the plants' own pollination or other growth.[24] The presence of ants is thought to provide some deterrence to other harmful insects though, so the production of ant-attracting nectar is plausibly a functionaladaptation. Ants do not harm the plants.[25]
Peony species come in two distinct growth habits, while hybrid cultivars in addition may occupy an intermediate habit.
herbaceous: During summer, renewal buds develop on the underground stem (the "crown"), particularly at the foot of the current season's annual shoots. These renewal buds come in various sizes. Large buds will grow into stems the following growing season, but smaller buds remain dormant. Theprimordia for the leaves can already be found in June, but the flower only starts differentiating in October, as the annual shoots die down, completing its development in December, when sepals, petals, stamens and pistils are all recognisable.[10]
tree: During the summer, large buds develop at the tip of the annual growth and near its foot. In the autumn, the leaves are shed, and the new stems become woody and are perennial.
Itoh (or "Intersectional"): In 1948 horticulturist Toichi Itoh from Tokyo used pollen from the yellow tree peony "Alice Harding" to fertilize the herbaceousP. lactiflora "Katoden", which resulted in a new category of peonies, the Itoh or intersectional cultivars. These are herbaceous, have leaves like tree peonies, with many large flowers from late spring to early autumn, and good peony wilt resistance. Some of the early Itoh cultivars are "Yellow Crown", "Yellow Dream", "Yellow Emperor" and "Yellow Heaven".[26]
Herbaceous and Itoh peonies are propagated by root division, and sometimes by seed. Tree peonies can be propagated by grafting, division, seed, and from cuttings, although root grafting is most common commercially.[27][28]
Herbaceous peonies such asPaeonia lactiflora, will die back to ground level each autumn. Their stems will reappear the following spring. However tree peonies, such asPaeonia suffruticosa, are shrubbier. They produce permanent woody stems that will lose their leaves in winter but the stem itself remains intact above ground level.[29]
The herb known asPaeonia, in particular the root ofP. lactiflora (Bai Shao, Radix Paeoniae Lactiflorae), has been used frequently intraditional medicines ofKorea,China andJapan. InJapan,Paeonia lactiflora used to be calledebisugusuri ("foreign medicine"). Pronunciation of 牡丹 (peony) in Japan is "botan." Inkampo, the Japanese adaptation ofChinese medicine, its root was used as a treatment for convulsions. It is also cultivated as agarden plant. In JapanPaeonia suffruticosa is called the "King of Flowers" andPaeonia lactiflora is called the "Prime Minister of Flowers."[36]
In China, the fallen petals ofPaeonia lactiflora are parboiled and sweetened as a tea-time delicacy. Peony water, aninfusion of peony petals, was used for drinking in theMiddle Ages. The petals may be added to salads or to punches and lemonades.[37]
In this gold-engravedlacquerware food tray from theSong dynasty (960–1279), the two long-tailed birds represent longevity, and the peony seen at the top centre represents prosperity.Peony, by Chinese artist Wang Qian,Yuan dynasty (1271–1368)Portrait of a peony by Chinese artistYun Shouping, 17th century
The peony is among the longest-used flowers in Eastern culture. Along with theplum blossom, it is a traditional floral symbol of China, where thePaeonia suffruticosa is called 牡丹 (mǔdān). It is also known as 富貴花 (fùguìhuā) "flower of riches and honour" or 花王 (huawang) "king of the flowers", and is used symbolically in Chinese art.[38]
In 1903, theQing dynasty declared the peony as the national flower.[39] Currently, theRepublic of China government inTaiwan designates the plum blossom as thenational flower, while thePeople's Republic of China government has no legally designated national flower. In 1994, the peony was proposed as the national flower after a nationwide poll, but theNational People's Congress failed to ratify the selection. In 2003, another selection process was initiated, but no choice has been made to date.[40]
The ancient Chinese cityLuoyang has a reputation as a cultivation centre for the peonies.[41] ThroughoutChinese history, peonies in Luoyang have been said to be the finest in the country. Dozens of peony exhibitions and shows are still held there annually.[42]
The Greek doctorDioscorides namedaglaophotis, a herb supposedly capable of warding off certain evils, as a member of the peony family.[43]
In theMiddle Ages, peonies were often painted with their ripe seed-capsules, since it was the seeds, not the flowers, which were medically significant.[44] Ancient superstition dictated that great care be taken not to be seen by awoodpecker while picking the plant's fruit, or the bird might peck out one's eyes.[45]
The red flowers of the speciesPaeonia peregrina are important inSerbian folklore. Known asKosovo peonies (Serbian:косовски божур,kosovski božur), they are said to represent the blood of Serbian warriors who died in theBattle of Kosovo.[46]
Mischievous nymphs were said to hide in the petals of the peony, giving it the meaning of Shame or Bashfulness in theLanguage of Flowers.[23]
Peonies are a common subject in tattoos, often used along with koi-fish. The popular use of peonies inJapanese tattoo was inspired by theukiyo-e artistUtagawa Kuniyoshi's illustrations ofSuikoden, a classical Chinese novel. His paintings of warrior-heroes covered in pictorial tattoos included lions, tigers, dragons, koi fish, and peonies, among other symbols. The peony became a masculine motif, associated with a devil-may-care attitude and disregard for consequence.[48]
Famous painters of peonies have includedConrad Gessner (ca. 1550) andAuguste Renoir in 1879.Paeonia officinalis can be found in the altar picture ofMaria im Rosenhag bySchongauer in the former Dominican Church inColmar. The Italian Jesuit, painter and architectGiuseppe Castiglione (1688–1766), who worked at the court of theQianlong Emperor in theQing dynasty, also painted peonies.[44]
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