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The peony or paeony is a flowering plant in the genus ''Paeonia'' , the only genus in the family Paeoniaceae . Peonies are native to Asia, Europe and Western
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
. Scientists differ on the number of species that can be distinguished, ranging from 25 to 40, although the current consensus is 33 known species. The relationships between the species need to be further clarified. Most are
herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...
perennial plants tall, but some are woody
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...
s tall. They have compound, 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 only 7–10 days. Peonies are popular garden plants in temperate regions. Herbaceous peonies are also sold as cut flowers on a large scale, although generally only available in late spring and early summer.


Description


Morphology

All Paeoniaceae are
herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...
perennials or deciduous shrubs, with thick storage roots and thin roots for gathering water and minerals. Some species are caespitose (tufted), because the crown produces adventitious buds, while others have
stolon In biology, stolons (from Latin '' stolō'', genitive ''stolōnis'' – "branch"), also known as runners, are horizontal connections between organisms. They may be part of the organism, or of its skeleton; typically, animal stolons are external s ...
s. They have rather large compound leaves without
gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...
s and stipules, and with anomocytic
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
ta. In the woody species the new growth emerges from scaly buds on the previous flush or from the crown of the rootstock. The large
bisexual flowers Plant reproductive morphology is the study of the physical form and structure (the morphology) of those parts of plants directly or indirectly concerned with sexual reproduction. Among all living organisms, flowers, which are the reproductive st ...
are mostly single at the end of the stem. In ''P. 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 of involucre, 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) free
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
s, with
anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
s fixed at their base to the filaments, and are sagittate in shape, open with longitudal slits at the outer side and free
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
grains which have three slits or pores and consist of two cells. Within the circle of stamens is a more or less prominent, lobed disc, which is presumed not to excrete
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
. Within the disk is a varying number (1-15) of separate carpels, which have a very short style and a decurrent stigma. Each of these develops into a dry fruit (which is called a follicle), 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. File:Paeonia-suffruticosa-buds.jpg, ''Paeonia suffruticosa'',
buds File:Paeonia.jpg, young growth of
a tree peony File:Paeonia vetchii feuille.jpg, ''Paeonia veitchii'',
leaf File:Paeonia 2015-04-16 183.jpg, ''Paeonia tenuifolia'',
leaves and flower buds File:Paeonia suffruticosa2.jpg, ''Paeonia suffruticosa'', showing the disk that encloses the carpels File:Paeonia wittmanniana Fruits BOGA.jpg, ''Paeonia wittmanniana'',
ripe follicles with seeds File:Peony anomil.jpg, ''Paeonia anomala'',
seeds


Phytochemistry

Over 262 compounds have been obtained so far from the plants of Paeoniaceae. These include monoterpenoid
glucoside A glucoside is a glycoside that is derived from glucose. Glucosides are common in plants, but rare in animals. Glucose is produced when a glucoside is hydrolysed by purely chemical means, or decomposed by fermentation or enzymes. The name was o ...
s,
flavonoid Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...
s, tannins, stilbenoids, triterpenoids,
steroid A steroid is a biologically active organic compound with four rings arranged in a specific molecular configuration. Steroids have two principal biological functions: as important components of cell membranes that alter membrane fluidity; and a ...
s, paeonols, and phenols. '' In vitro'' biological activities include antioxidant, antitumor, antipathogenic, immunomodulative, cardiovascular-system-protective activities and central-nervous-system activities. Paeoniaceae are dependent on C3 carbon fixation. They contain ellagic acid, myricetin, ethereal oils and
flavone Flavone is an organic compound with the formula . A white solid, flavone is a derivative of chromone with a phenyl (Ph) substituent adjacent to the ether group. The compound is of little direct practical importance, but susbstituted derivatives, t ...
s, as well as crystals of
calcium oxalate Calcium oxalate (in archaic terminology, oxalate of lime) is a calcium salt of oxalic acid with the chemical formula . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydrate ...
. The wax tubules that are formed primarily consist of palmitone (the
ketone In organic chemistry, a ketone is a functional group with the structure R–C(=O)–R', where R and R' can be a variety of carbon-containing substituents. Ketones contain a carbonyl group –C(=O)– (which contains a carbon-oxygen double bo ...
of palmitic acid).


Genome

The basic chromosome number is five. About half of the species of the section ''Paeonia'' 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.


Taxonomy

The family name "Paeoniaceae" was first used by Friedrich K.L. Rudolphi in 1830, following a suggestion by Friedrich Gottlieb Bartling that same year. The family had been given other names a few years earlier.James L. Reveal. 2008 onward. "A Checklist of Family and Suprafamilial Names for Extant Vascular Plants." At: Home page of James L. Reveal and C. Rose Broome. (see ''External links'' below). The composition of the family has varied, but it has always consisted of ''Paeonia'' and one or more genera that are now placed in Ranunculales. It has been widely believed that ''Paeonia'' is closest to '' Glaucidium'', and this idea has been followed in some recent works.David J. Mabberley. 2008. ''Mabberley's Plant-Book''.Cambridge University Press: UK.
Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies, however, have demonstrated conclusively that ''Glaucidium'' belongs in the family Ranunculaceae, order Ranunculales, but that ''Paeonia'' belongs in the unrelated
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
Saxifragales. The genus ''Paeonia'' consists of about 35 species, assigned to three sections: ''Moutan'', ''Onaepia'' and ''Paeoniae''. The section ''Onaepia'' only includes ''P. brownii'' and ''P. californicum''. The section ''Moutan'' is divided into ''P. delavayi'' and ''P. ludlowii'', together making up the subsection ''Delavayanae'', and ''P. catayana'', ''P. decomposita'', ''P. jishanensis'', ''P. osti'', ''P. qiui'' and ''P. rockii'' which constitute the subsection ''Vaginatae''. ''
P. suffruticosa P. is an abbreviation or acronym that may refer to: * Page (paper), where the abbreviation comes from Latin ''pagina'' * Paris Herbarium, at the ''Muséum national d'histoire naturelle'' * ''Pani'' (Polish), translating as Mrs. * The ''Pacific Repo ...
'' is a cultivated hybrid swarm, not a naturally occurring species. The remainder of the species belongs to the section ''Paeonia'', which is characterised by a complicated reticulate 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, or nothospecies. Known diploid taxa in the ''Paeonia''-section are ''P. anomala'', ''P. lactiflora'', ''P. veitchii'', ''P. tenuifolia'', ''P. emodi'', ''P. broteri'', ''P. cambedessedesii'', ''P. clusii'', ''P. rhodia'', ''P. daurica'' subsps. ''coriifolia'', ''daurica'', ''macrophylla'' and ''mlokosewitschii''. Tetraploid taxa are ''P. arietina'', ''P. officinalis'', ''P. parnassica'', ''P. banatica'', ''P. russi'', ''P. peregrina'', ''P. coriacea'', ''P. mascula'' subsps. ''hellenica'' and ''mascula'', and ''P. daurica'' subsps. ''tomentosa'' and ''wittmanniana''. Species that have both diploid and tetraploid populations include ''P. clusii'', ''P. mairei'' and ''P. obovata''. ''P. anomala'' was proven to be a hybrid of ''P. lactiflora'' and ''P. veitchii'', although being a diploid with 10 chromosomes. ''P. emodi'' and ''P. sterniana'' are diploid hybrids of ''P. lactiflora'' and ''P. veitchii'' too, and radically different in appearance. ''P. russi'' is the tetraploid hybrid of diploid ''P. lactiflora'' and ''P. mairei'', while ''P. cambedessedesii'' is the diploid hybrid of ''P. lactiflora'', likely ''P. mairei'', but possibly also ''P. obovata''. ''P. peregrina'' is the tetraploid hybrid of ''P. anomala'' and either ''P. arietina'', ''P. humilis'', ''P. officinalis'', ''P. parnassica'' or less likely ''P. tenuifolia'', or one of their (now extinct) common ancestors. ''P. banatica'' is the tetraploid hybrid of ''P. 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'', and ''P. daurica'' subsp. ''wittmanniana'' are all descendants of hybrids of ''P. lactiflora'' and ''P. obovata''.


Phylogeny

Recent genetic analyses relate the monogeneric family Paeoniaceae to a group of families with woody species in the order Saxifragales. This results in the following relationship tree. One dissertation suggests the section ''Onaepia'' branches off earliest, but a later publication of the same author and others suggests the ''Moutan''-section splits off first. Within that section ''P. ludlowii'' and ''P. delavayi'' are more related to each other than to any other species.


Species

*
Herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...
species (about 30 species) **'' Paeonia algeriensis'' **''
Paeonia anomala ''Paeonia anomala'' is a species of herbaceous perennial peony. This plant is ½-1 m high, with a thick irregular taproot and thin side roots. The deeply incised leaves have leaflets which are themselves divided in fine segments. It flowers ...
'' **'' Paeonia arietina'' **''
Paeonia broteri ''Paeonia broteri'' is a perennial, herbaceous species of peony. It is an endemic species of the Iberian Peninsula. It bears rose-pink highly fragrant flowers about 12 cm wide and glossy green leaves. It reaches up to in height. Descrip ...
'' **'' Paeonia brownii'' (Brown's peony) **''
Paeonia californica ''Paeonia californica'' is a perennial herbaceous plant of 35–70 cm high, that retreats underground in summer, and reoccurs with the arrival of the winter rains. It has lobed leaves, elliptic (cup-shaped) drooping flowers with dark maroon- ...
'' (California peony or wild peony) **'' Paeonia cambessedesii'' (Majorcan peony) **''
Paeonia clusii ''Paeonia clusii'' is a relatively low (25–50 cm) species of herbaceous peony with scented, white or pink flowers of up to 12 cm in diameter. In the wild, the species can only be found on the islands of Crete and Karpathos (subsp. ''c ...
'' ***subsp. ''clusii'' ***subsp. ''rhodia'' **''
Paeonia coriacea ''Paeonia coriacea'', also known as the Corsican peony or Andalusian peony, is a species of flowering plant within the family Paeoniaceae. Description ''P. coriacea'' is a herbaceous, perennial species. Plants possess stems that range in height ...
'' **''
Paeonia corsica ''Paeonia corsica'' is a perennial herbaceous plant of high that belongs to the peonies. It naturally occurs on Corsica, Sardinia, on the Ionian islands and in western Greece. It has hairless green to purple stems, and the lower leaves consist m ...
'' **'' Paeonia daurica'' ***subsp. ''coriifolia'' ***subsp. ''daurica'' ***subsp. '' macrophylla'' ***subsp. '' mlokosewitschii'' ***subsp. ''tomentosa'' ***subsp. ''velebitensis'' ***subsp. '' wittmanniana'' **'' Paeonia emodi'' **''
Paeonia intermedia ''Paeonia intermedia'' is a species of flowering plant in the peony family Paeoniaceae, native to Central Asia (except Turkmenistan), the Altai Mountains, and Xinjiang in China. A perennial herb reaching , it is found on scrubby and grassy slopes ...
'' **'' Paeonia kesrouanensis'' ( Keserwan peony) **''
Paeonia lactiflora ''Paeonia lactiflora'' (Chinese peony, Chinese herbaceous peony, or common garden peony) is a species of herbaceous perennial flowering plant in the family Paeoniaceae, native to central and eastern Asia from eastern Tibet across northern China t ...
'' ( Chinese or common garden peony) **'' Paeonia mairei'' **'' Paeonia mascula'' (
Balkan The Balkans ( ), also known as the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throughout the who ...
, wild or male peony) **''
Paeonia obovata ''Paeonia obovata'' is a perennial herbaceous species of peony growing 30–70 cm high. It has white, pink or purple-red flowers and its lower leaves consist of no more than nine leaflets or segments. In English it is sometimes called wood ...
'' ***subsp. ''willmottiae'' **'' Paeonia officinalis'' ( European or common peony, type species) **'' Paeonia parnassica'' ( Greek peony) **''
Paeonia peregrina ''Paeonia peregrina'' is a species of flowering plant in the peony family ''Paeoniaceae'', native to Southeastern Europe and Turkey. It is an erect, herbaceous perennial with 9-lobed, deeply divided leaves. Single, glossy red flowers, in diamete ...
'' **'' Paeonia sterniana'' **'' Paeonia tenuifolia'' (Steppe peony) **''
Paeonia veitchii ''Paeonia veitchii'' is a species of herbaceous perennial peony. The vernacular name in China is 川赤芍 (chuan chi shao). This species is ½-1 m high, has a thick irregular taproot and thin side roots, and deeply incised leaves, with leaf ...
'' (Veitch's peony) * Woody species (about 8 species) **'' Paeonia decomposita'' **'' Paeonia delavayi'' (Delavay's tree peony) **'' Paeonia jishanensis'' (Jishan peony) **'' Paeonia ludlowii'' (Ludlow's tree peony) **'' Paeonia ostii'' (Osti's peony) **'' Paeonia qiui'' (Qiu's peony) **'' Paeonia rockii'' (Rock's peony or tree peony;
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
'' Paeonia suffruticosa'' subsp. ''rockii'' (Chinese tree peony, known as "moutan (moutan peony)" in China)) File:Paeonia anomala riu orkhon.jpg, ''Paeonia anomala'' File:Peonía o Rosa alabardera (Peonia broteroi) (3814459852).jpg, ''Paeonia broteri'' File:Paeoniabrownii3.jpg, ''Paeonia brownii'' File:Paeonia californica 2320679478.jpg, ''Paeonia californica'' File:Paeonia cambessedesii - Copenhagen Botanical Garden - DSC07470.JPG, ''Paeonia cambessedesii'' File:Paeonia cambessedesii 2c.JPG, ''Paeonia cambessedesii'' File:Paeonia daurica ssp mlokosewitschii.jpg, ''Paeonia daurica mlokosewitschii'' File:Paeonia delavayi feuille.jpg, ''Paeonia delavayi'' File:Paeonia emodi - Kew Gardens.jpg, ''Paeonia emodi'' File:Paeonia anomala var. intermedia 2016-04-19 7716.JPG, ''Paeonia intermedia'' File:Paeonia ludlowii.jpg, ''Paeonia ludlowii'' File:Paeonia mairei.jpg, ''Paeonia mairei'' File:Paeonia mascula 02.jpg, ''Paeonia mascula'' File:Paeonia mascula subsp. russoi 3.jpg, ''Paeonia russoi '' File:Paeonia japonica.JPG, ''Paeonia obovata japonica'' File:Rugova Mountains.jpg, ''Paeonia officinalis'' File:Paeonia ostii.jpg, ''Paeonia ostii'' File:Paeonia rockii 2011 G2.jpg, ''Paeonia rockii'' File:Воронець 09.jpg, ''Paeonia tenuifolia''


Etymology

The peony is named after Paeon (also spelled Paean), a student of Asclepius, the Greek god of medicine and healing. In Greek mythology, when Asclepius became jealous of his pupil, Zeus saved Paeon from the wrath of Asclepius by turning him into the peony flower.


Distribution

The genus ''Paeonia'' naturally occurs in the temperate and cold areas of the Northern Hemisphere. The section ''
Moutan Tree peony is the vernacular name for the section ''Moutan'' of the plant genus '' Paeonia'', or one of the species or cultivars belonging to this section. It consists of shrubs that have perennial aerial woody stems. Other peonies do not have ...
'', which includes all woody species, is restricted in the wild to Central and Southern China, including Tibet. The section ''Onaepia'' consist of two herbaceous species and is present in the West of North-America, '' P. brownii'' between southern British Columbia and the Sierra Nevada in California and eastward to Wyoming and Utah, while '' P. californica'' is limited to the coastal mountains of Southern and Central California. The section ''Paeonia'', which comprises all other herbaceous species, occurs in a band stretching roughly from Morocco and Spain to Japan. One species of the section ''Paeonia'', '' P. anomala'', has by far the largest distribution, which is also north of the distribution of the other species: from the Kola peninsula in North-West Russia, to
Lake Baikal Lake Baikal (, russian: Oзеро Байкал, Ozero Baykal ); mn, Байгал нуур, Baigal nuur) is a rift lake in Russia. It is situated in southern Siberia, between the federal subjects of Irkutsk Oblast to the northwest and the Repu ...
in Siberia and South to the Tien Shan Mountains of Kazakhstan. The rest of the section concentrates around the Mediterranean, and in Asia. The species around the Mediterranean include '' Paeonia algeriensis'' that is an endemic of the coastal mountains of Algeria, '' P. coriacea'' in the Rif Mountains and Andalusia, '' P. cambessedesii'' on
Majorca Mallorca, or Majorca, is the largest island in the Balearic Islands, which are part of Spain and located in the Mediterranean. The capital of the island, Palma, is also the capital of the autonomous community of the Balearic Islands. The Bal ...
, '' P. russoi'' on
Corsica Corsica ( , Upper , Southern ; it, Corsica; ; french: Corse ; lij, Còrsega; sc, Còssiga) is an island in the Mediterranean Sea and one of the 18 regions of France. It is the fourth-largest island in the Mediterranean and lies southeast of ...
, Sardinia and Sicilly, '' P. corsica'' on Corsica, Sardinia, the Ionian islands and in western Greece'', P. clusii'' subsp. ''clusii'' on Crete and Karpathos, and subsp. ''rhodia'' on Rhodes, '' P. kesrouanensis'' in the Western Taurus Mountains, '' P. arietina'' from the Middle Taurus Mountains, '' P. broteri'' in Andalucia, '' P. humilis'' from Andalucia to the Provence, '' 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, while '' P. mascula'' has a large distribution from Catalonia and southern France to Israel and Turkey. Between the two concentrations, the subspecies of '' Paeonia daurica'' occur, with subspecies ''velebitensis'' in Croatia, and ''daurica'' in the Balkans and Crimea, while the other subspecies ''coriifolia'', ''macrophylla'', ''mlokosewitschii'', ''tomentosa'' and ''wittmanniana'' are known from the Caucasus, Kaçkar and Alborz Mountains. '' Paeonia emodi'' occurs in the western Himalayas between Pakistan and western Nepal, '' P. sterniana'' is an endemic of southeastern Tibet, '' P. veitchii'' grows in Central China ( Qinghai, Ningxia,
Gansu Gansu (, ; alternately romanized as Kansu) is a province in Northwest China. Its capital and largest city is Lanzhou, in the southeast part of the province. The seventh-largest administrative district by area at , Gansu lies between the Tibet ...
, Shaanxi,
Shanxi Shanxi (; ; formerly romanised as Shansi) is a landlocked province of the People's Republic of China and is part of the North China region. The capital and largest city of the province is Taiyuan, while its next most populated prefecture-lev ...
, Sichuan and the eastern rim of Tibet), like '' P. mairei'' (Gansu, Guizhou, Hubei, Shaanxi, Sichuan, and Yunnan), while '' P. obovata'' grows in warm-temperate to cold China, including Manchuria, Korea, Japan, Far Eastern Russia ( Primorsky Krai) and on Sakhalin, and '' P. lactiflora'' occurs in Northern China, including Manchuria, Japan, Korea, Mongolia, Russia (Far East and Siberia).


Distributional history

The species of the section Paeonia have a
disjunct distribution In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...
, 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 either
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
or tetraploid hybrids that resulted from the crossbreeding of species currently limited to eastern Asia. The large distance between the ranges of the parent species and the nothospecies 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'' and ''P. mairei'' likely became
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...
and so produced the Himalayan nothospecies ''P. emodi'' and ''P. sterniana''.


Cultivation

Ancient Chinese texts mention the peony was used for flavoring food. Confucius (551–479 BC) is quoted to have said: "I eat nothing without its sauce. I enjoy it very much, because of its flavor." 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 the Tang dynasty, when they were grown in the imperial gardens. In the tenth century the cultivation of peonies spread through China, and the seat of the Song dynasty, Luoyang, was the centre for its cultivation, a position it still holds today. A second centre for peony cultivation developed during the Qing dynasty in Cáozhōu, now known as He 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 by self fertilisation and crossbreeding, particularly during the eighteenth to twentieth centuries (middle
Edo Edo ( ja, , , "bay-entrance" or "estuary"), also romanized as Jedo, Yedo or Yeddo, is the former name of Tokyo. Edo, formerly a ''jōkamachi'' (castle town) centered on Edo Castle located in Musashi Province, became the ''de facto'' capital of ...
to early Shōwa periods). During the 1940s Toichi Itoh succeeded in crossing tree peonies and herbaceous peonies and so created a new class of so-called intersectional hybrids. Although ''P. 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 when ''P. lactiflora'' was introduced from its native China to Europe. The tree peony was introduced in Europe and planted in Kew Gardens in 1789. The main centre of peony breeding in Europe has been in the United Kingdom, and particularly France. Here, breeders like
Victor Lemoine Pierre Louis Victor Lemoine (October 21, 1823 in Delme, Moselle - December 11, 1911) was a celebrated and prolific French flower breeder who, among other accomplishments, created many of today's lilac varieties. As a result of his accomplishment ...
and François Félix Crousse selected many new varieties, mainly with ''P. 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. 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.


Plant growth habits

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. The primordia 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. * 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 herbaceous ''P. 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".


Flower types

Six types of flower are generally distinguished in cultivars of herbaceous peonies. * single: a single or double row of broad petals encircle fertile stamens, carpels visible. * Japanese: a single or double row of broad petals encircle somewhat broadened
staminode In botany, a staminode is an often rudimentary, sterile or abortive stamen, which means that it does not produce pollen.Jackson, Benjamin, Daydon; ''A Glossary of Botanic Terms with their Derivation and Accent''; Published by Gerald Duckworth & Co. ...
s, may carry pollen along the edges, carpels visible. * anemone: a single or double row of broad petals encircle narrow incurved petal-like staminodes; fertile stamens are absent, carpels visible. * semi-double: a single or double row of broad petals encircles further broad petals intermingled with stamens. * bomb: a single row of broad petals encircles a shorter dense pompon of narrower petals. * double: the flower consists of many broad petals only, including those which likely are altered stamens and carpels. File:Paeonia Claire de Lune1a.UME.JPG, ''Paeonia ×arendsii'' 'Claire de Lune',
single flowered File:Paeonia 'Walter Mains'.jpg, ''Paeonia'' 'Walter Mains',
Japanese flowered File:BowlOfBeauty1b.UME.JPG, ''Paeonia lactiflora'' 'Bowl Of Beauty',
anemone flowered File:Paeonia lactiflora 'James Kelway'.JPG, ''Paeonia lactiflora'' 'James Kelway',
semi-double flowered File:Paeonia 'Ruth Clay' 14.JPG, ''Paeonia'' 'Ruth Clay',
bomb flowered File:Paeonia Da Fu Gui.jpg, ''Paeonia lactiflora'' 'Da Fu Gui',
double flowered


Propagation

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. Herbaceous peonies such as ''Paeonia lactiflora'', will die back to ground level each autumn. Their stems will reappear the following spring. However tree peonies, such as ''Paeonia suffruticosa'', are shrubbier. They produce permanent woody stems that will lose their leaves in winter but the stem itself remains intact above ground level.


Hybrid cultivars

The following hybrids and cultivars have gained the Royal Horticultural Society's
Award of Garden Merit The Award of Garden Merit (AGM) is a long-established annual award for plants by the British Royal Horticultural Society (RHS). It is based on assessment of the plants' performance under UK growing conditions. History The Award of Garden Merit ...
:- *'Bartzella' *'Coral Charm' *''Paeonia'' × ''festiva'' 'Rubra Plena' *''Paeonia'' × ''lemoinei'' 'High Noon'


Uses

The herb known as Paeonia, in particular the root of ''P. lactiflora'' (Bai Shao, Radix Paeoniae Lactiflorae), has been used frequently in traditional medicines of Korea,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
and
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
. In
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
, ''
Paeonia lactiflora ''Paeonia lactiflora'' (Chinese peony, Chinese herbaceous peony, or common garden peony) is a species of herbaceous perennial flowering plant in the family Paeoniaceae, native to central and eastern Asia from eastern Tibet across northern China t ...
'' used to be called ''ebisugusuri'' ("foreign medicine"). Pronunciation of 牡丹 (peony) in Japan is "botan." In '' kampo'', the Japanese adaptation of Chinese medicine, its root was used as a treatment for convulsions. It is also cultivated as a garden plant. In Japan '' Paeonia suffruticosa'' is called the "King of Flowers" and ''
Paeonia lactiflora ''Paeonia lactiflora'' (Chinese peony, Chinese herbaceous peony, or common garden peony) is a species of herbaceous perennial flowering plant in the family Paeoniaceae, native to central and eastern Asia from eastern Tibet across northern China t ...
'' is called the "Prime Minister of Flowers." In China, the fallen petals of ''Paeonia lactiflora'' are parboiled and sweetened as a tea-time delicacy. Peony water, an infusion of peony petals, was used for drinking in the Middle Ages. The petals may be added to salads or to punches and lemonades. Peonies are also extensively grown as ornamental plants for their very large, often scented flowers.


Culture

Located in Cuigou Village, Chaoyang Town, Mengjin District, Luoyang, Henan, China. The first generation of Kao, wild peony, Ranunculaceae, and medicinal genus, was founded in 1800 meters in Baiyun Mountain, Song County, Henan. Transplanted by Cui Yueqi, the owner of the house, in May 1958, after 27 years of labor and land, the plant chapter of "Henan Geography Knowledge Expo" was filmed by China Education Television in April 2017. The height of the tree is 1.7 meters, the diameter of the canopy is more than two meters, and the number of flowers will reach 350 in the future. The peony is among the longest-used flowers in Eastern culture. Along with the plum blossom, it is a traditional floral symbol of China, where the '' Paeonia 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. In 1903, the Qing dynasty declared the peony as the national flower. Currently, the
Republic of China Taiwan, officially the Republic of China (ROC), is a country in East Asia, at the junction of the East and South China Seas in the northwestern Pacific Ocean, with the People's Republic of China (PRC) to the northwest, Japan to the northeast ...
government in Taiwan designates the plum blossom as the
national flower In a number of countries, plants have been chosen as symbols to represent specific geographic areas. Some countries have a country-wide floral emblem; others in addition have symbols representing subdivisions. Different processes have been used to ...
, while the People'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 the National People's Congress failed to ratify the selection. In 2003, another selection process was initiated, but no choice has been made to date. The ancient Chinese city Luoyang has a reputation as a cultivation centre for the peonies. Throughout
Chinese history The earliest known written records of the history of China date from as early as 1250 BC, from the Shang dynasty (c. 1600–1046 BC), during the reign of king Wu Ding. Ancient historical texts such as the ''Book of Documents'' (early chapter ...
, peonies in Luoyang have been said to be the finest in the country. Dozens of peony exhibitions and shows are still held there annually. In the Middle Ages, peonies were often painted with their ripe seed-capsules, since it was the seeds, not the flowers, which were medically significant. Ancient superstition dictated that great care be taken not to be seen by a woodpecker while picking the plant's fruit, or the bird might peck out one's eyes. The red flowers of the species ''
Paeonia peregrina ''Paeonia peregrina'' is a species of flowering plant in the peony family ''Paeoniaceae'', native to Southeastern Europe and Turkey. It is an erect, herbaceous perennial with 9-lobed, deeply divided leaves. Single, glossy red flowers, in diamete ...
'' are important in Serbian folklore. Known as Kosovo peonies ( sr, косовски божур, ), they are said to represent the blood of Serbian warriors who died in the
Battle of Kosovo The Battle of Kosovo ( tr, Kosova Savaşı; sr, Косовска битка) took place on 15 June 1389 between an army led by the Serbian Prince Lazar Hrebeljanović and an invading army of the Ottoman Empire under the command of Sultan ...
. In 1957, the Indiana General Assembly passed a law to make the peony the
state flower This is a list of U.S. state, federal district, and territory flowers. See also *List of U.S. state trees * Lists of U.S. state insignia References External linksList of state flowers {{USStateLists * U.S. state flowers Flowers ...
of Indiana, a title which it holds to this day. It replaced the zinnia, which had been the state flower since 1931. Mischievous nymphs were said to hide in the petals of the Peony, giving it the meaning of Shame or Bashfulness in the Language of Flowers. While the peony takes several years to re-establish itself when moved, it blooms annually for decades once it has done so. Peonies tend to attract ants 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. 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 functional adaptation. Ants do not harm the plants. Peonies are a common subject in tattoos, often used along with koi-fish. The popular use of peonies in Japanese tattoo was inspired by the ukiyo-e artist Utagawa Kuniyoshi's illustrations of ''
Suikoden is a series of role-playing video games created by Yoshitaka Murayama. The games are loosely based on the classical Chinese novel ''Water Margin'', whose title is rendered as in Japanese. Each individual game centers on themes of politics, co ...
'', 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. Famous painters of peonies have included Conrad Gessner (ca. 1550) and
Auguste Renoir Pierre-Auguste Renoir (; 25 February 1841 – 3 December 1919) was a French artist who was a leading painter in the development of the Impressionist style. As a celebrator of beauty and especially feminine sensuality, it has been said that " ...
in 1879. ''Paeonia officinalis'' can be found in the altar picture of ''Maria im Rosenhag'' by Schongauer in the former Dominican Church in
Colmar Colmar (, ; Alsatian: ' ; German during 1871–1918 and 1940–1945: ') is a city and commune in the Haut-Rhin department and Grand Est region of north-eastern France. The third-largest commune in Alsace (after Strasbourg and Mulhouse), it is ...
. The Italian Jesuit, painter and architect Giuseppe Castiglione (1688-1766), who worked at the court of the
Qianlong Emperor The Qianlong Emperor (25 September 17117 February 1799), also known by his temple name Emperor Gaozong of Qing, born Hongli, was the fifth Emperor of the Qing dynasty and the fourth Qing emperor to rule over China proper, reigning from 1735 t ...
in the Qing dynasty, painted peonies.


References


Bibliography

*


External links


Family and Suprafamilial Names
At
James L. RevealFlora Europaea: ''Paeonia''Ornamental Plants from Russia: ''Paeonia''The Peony Society (UK)
(defunct as of 2106)
Canadian Peony SocietyU.S. Peony SocietyCarsten Burkhardt's Open Source Peony ProjectGerman Peony Group
{{Authority control Garden plants Monogeneric plant families Taxa named by Carl Linnaeus