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The peony or paeony is a
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''Paeonia'' , the only genus in the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Paeoniaceae . Peonies are native to
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
,
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
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 In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
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 plant A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
s 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 Compound may refer to: Architecture and built environments * Compound (enclosure), a cluster of buildings having a shared purpose, usually inside a fence or wall ** Compound (fortification), a version of the above fortified with defensive struct ...
, 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 Cut flowers are flowers or flower buds (often with some stem and leaf) that have been cut from the plant bearing it. It is usually removed from the plant for decorative use. Typical uses are in vase displays, wreaths and garlands. Many gardene ...
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 ...
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
s or
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
shrubs, with thick storage roots and thin roots for gathering water and minerals. Some species are
caespitose This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...
(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
stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...
s, 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 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 In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
, 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
carpel Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...
s, which have a very short
style Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...
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 Monoterpenes are a class of terpenes that consist of two isoprene units and have the molecular formula C10H16. Monoterpenes may be linear (acyclic) or contain rings (monocyclic and bicyclic). Modified terpenes, such as those containing oxygen funct ...
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,
tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' (from Anglo-Norman ''tanner'', ...
s,
stilbenoid Stilbenoids are hydroxylated derivatives of stilbene. They have a C6–C2–C6 structure. In biochemical terms, they belong to the family of phenylpropanoids and share most of their biosynthesis pathway with chalcones. Most stilbenoids are prod ...
s,
triterpenoid Triterpenes are a class of chemical compounds composed of three terpene units with the molecular formula C30H48; they may also be thought of as consisting of six isoprene units. Animals, plants and fungi all produce triterpenes, including squale ...
s,
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,
paeonol Paeonol is a phenolic compound found in peonies such as ''Paeonia suffruticosa'' (moutan cortex), in ''Arisaema erubescens'', and in ''Dioscorea japonica''. It is a chemical compound found in some traditional Chinese medicines. Biological effec ...
s, and
phenols In organic chemistry, phenols, sometimes called phenolics, are a class of chemical compounds consisting of one or more hydroxyl groups (— O H) bonded directly to an aromatic hydrocarbon group. The simplest is phenol, . Phenolic compounds are c ...
. ''
In vitro ''In vitro'' (meaning in glass, or ''in the glass'') studies are performed with microorganisms, cells, or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in biology an ...
'' biological activities include antioxidant, antitumor, antipathogenic, immunomodulative, cardiovascular-system-protective activities and central-nervous-system activities. Paeoniaceae are dependent on
C3 carbon fixation carbon fixation is the most common of three metabolic pathways for carbon fixation in photosynthesis, along with and CAM. This process converts carbon dioxide and ribulose bisphosphate (RuBP, a 5-carbon sugar) into two molecules of 3-phosph ...
. They contain
ellagic acid Ellagic acid is a polyphenol found in numerous fruits and vegetables. It is the dilactone of hexahydroxydiphenic acid. Name The name comes from the French term ''acide ellagique'', from the word ''galle'' spelled backwards because it can be ob ...
,
myricetin Myricetin is a member of the flavonoid class of polyphenolic compounds, with antioxidant properties. Common dietary sources include vegetables (including tomatoes), fruits (including oranges), nuts, berries, tea, and red wine. Myricetin is structu ...
,
ethereal oil An essential oil is a concentrated hydrophobic liquid containing volatile (easily evaporated at normal temperatures) chemical compounds from plants. Essential oils are also known as volatile oils, ethereal oils, aetheroleum, or simply as the o ...
s 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 Palmitic acid (hexadecanoic acid in IUPAC nomenclature) is a fatty acid with a 16-carbon chain. It is the most common saturated fatty acid found in animals, plants and microorganisms.Gunstone, F. D., John L. Harwood, and Albert J. Dijkstra. The Li ...
).


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 Friedrich Gottlieb Bartling (December 9, 1798 – November 20, 1875) was a German botanist who was a native of Hanover. He studied natural sciences at the University of Göttingen, and in 1818 took a botanical journey through Hungary and Croatia ...
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 Ranunculales is an order of flowering plants. Of necessity it contains the family Ranunculaceae, the buttercup family, because the name of the order is based on the name of a genus in that family. Ranunculales belongs to a paraphyletic group kno ...
. 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 Saxifragales (saxifrages) are an order of flowering plants (Angiosperms). They are an extremely diverse group of plants which include trees, shrubs, perennial herbs, succulent and aquatic plants. The degree of diversity in terms of vege ...
. 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'' 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 Reticulate evolution, or network evolution is the origination of a lineage through the partial merging of two ancestor lineages, leading to relationships better described by a phylogenetic network than a bifurcating tree. Reticulate patterns ca ...
. 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 #REDIRECT Hybrid name In botanical nomenclature, a hybrid may be given a hybrid name, which is a special kind of botanical name, but there is no requirement that a hybrid name should be created for plants that are believed to be of hybrid origin. ...
. 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 In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
family Paeoniaceae to a group of families with woody species in the order
Saxifragales The Saxifragales (saxifrages) are an order of flowering plants (Angiosperms). They are an extremely diverse group of plants which include trees, shrubs, perennial herbs, succulent and aquatic plants. The degree of diversity in terms of vege ...
. 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 In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
(about 30 species) **''
Paeonia algeriensis ''Paeonia algeriensis'' is a herbaceous species of peony that naturally occurs in the coastal mountain range of Algeria (Kabylie). It has solitary flowers with pink to magenta petals and one or two carpels per flower, that develop into follicles ...
'' **''
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 arietina'', also known as the ram's horn peony, is a species of flowering plant within the family Paeoniaceae. Description ''Paeonia arietina'' is a perennial species. Stems can range from 30 to 70 cm tall and usually possess g ...
'' **''
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 ''Paeonia brownii'' is a low to medium height, Herbaceous plant, herbaceous Perennial plant, perennial flowering plant in the family Paeoniaceae.Sierra Nevada Wildflowers, Karen Wiese, 2nd ed., 2013, p. 102 It has compound, steely-gray, somewhat ...
'' (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 ''Paeonia cambessedesii'' is a Perennial plant, perennial Herbaceous plant, herbaceous species of peony about 45 cm high. It has pink flowers. The stems, major veins and undersides of the leaves remain purple red, while the upper surface of ...
'' (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 daurica ''Paeonia daurica'' is a perennial herbaceous plant belonging to the peony family. It has slender carrot-shaped roots, leaves mostly consisting of nine leaflets, with one flower per stem. The flower is subtended by none to two leafy bracts, and ...
'' ***subsp. ''coriifolia'' ***subsp. ''daurica'' ***subsp. '' macrophylla'' ***subsp. '' mlokosewitschii'' ***subsp. ''tomentosa'' ***subsp. ''velebitensis'' ***subsp. '' wittmanniana'' **''
Paeonia emodi ''Paeonia emodi'' is a robust herbaceous perennial plant that winters with buds underground (a so-called hemicryptophyte), has large white flowers and large deeply incised leaves, and belongs to the family Paeoniaceae. Its local vernacular name ...
'' **'' Paeonia intermedia'' **''
Paeonia kesrouanensis ''Paeonia kesrouanensis'' is a species of peony native to Syria, Turkey and Lebanon Lebanon ( , ar, لُبْنَان, translit=lubnān, ), officially the Republic of Lebanon () or the Lebanese Republic, is a country in Western Asia. It i ...
'' ( 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 Chi ...
'' (
Chinese Chinese can refer to: * Something related to China * Chinese people, people of Chinese nationality, citizenship, and/or ethnicity **''Zhonghua minzu'', the supra-ethnic concept of the Chinese nation ** List of ethnic groups in China, people of va ...
or common
garden A garden is a planned space, usually outdoors, set aside for the cultivation, display, and enjoyment of plants and other forms of nature. The single feature identifying even the wildest wild garden is ''control''. The garden can incorporate both ...
peony) **''
Paeonia mairei ''Paeonia mairei'' is a species of peony, that is endemic to the mountains of central China. Its vernacular name in China is 美丽芍药 (mei li shao yao) meaning "beautiful peony". The plant may be between 45 and 100 cm high and has mostly ...
'' **''
Paeonia mascula ''Paeonia mascula'' is a species of peony. It is a herbaceous perennial tall, with leaves that are divided into three segments, and large red flowers in late spring and early summer. Native to Syria, Spain, France, Italy, Croatia, Bosnia and Her ...
'' (
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 ''Paeonia officinalis'', the common peony, or garden peony, is a species of flowering plant in the family Paeoniaceae, native to mainly mountainous areas of Southern Europe and introduced in Central and Western Europe and North America. ''Paeon ...
'' (
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
an or common peony, type species) **''
Paeonia parnassica ''Paeonia parnassica'', the Greek peony, is a plant that is native to the mountains of south-central Greece. The flowers are produced in late spring with a deep maroon red colouring on 65 cm stems. The blooms are large, up to 12 cm in ...
'' (
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
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 sterniana'' is a perennial, herbaceous peony of approximately 45 cm high in cultivation, with white or sometimes pinkish flowers. It grows in the wild in southeastern Tibet. This peony is very rare in cultivation. It produces blue ...
'' **''
Paeonia tenuifolia ''Paeonia tenuifolia'' is a herbaceous species of peony that is called the steppe peony or the fern leaf peony. It is native to the Caucasus Mountains of Russia and the Black Sea coast of Ukraine, spreading westward into Bulgaria, Romania and S ...
'' (Steppe peony) **'' Paeonia veitchii'' (Veitch's peony) * Woody species (about 8 species) **''
Paeonia decomposita Paeonia or Paionia may refer to: * The genus ''Paeonia'', which comprises all peony plants * Paeonia (kingdom), an ancient state occupying roughly the same area as the present-day Republic of North Macedonia * Paionia (municipality) Paionia ( e ...
'' **''
Paeonia delavayi ''Paeonia delavayi'' is a low woody shrub belonging to the peonies, that is endemic to China. The vernacular name in China is 滇牡丹 (diān mǔdan). In English it is called Delavay's tree peony, Delavay peony, Dian peony, and dian mu dan. It ...
'' (Delavay's
tree peony 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 ...
) **''
Paeonia jishanensis ''Paeonia jishanensis'' is a species of peony 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. Scientists differ on ...
'' (Jishan peony) **''
Paeonia ludlowii ''Paeonia ludlowii'', is a medium high, deciduous shrub, belonging to the peonies, that is endemic to southeast Tibet. In Tibet it is known as lumaidao meaning "God’s flower". The vernacular name in Chinese is 大花黄牡丹 (da hua huang mu ...
'' (Ludlow's tree peony) **''
Paeonia ostii ''Paeonia ostii'' is a hardy shrub in the peony family, Paeoniaceae. It can be found in the Gansu, Anhui, Shaanxi and Henan provinces of China. It can reach heights of 1.5 m with grey-brown bark and lance shaped leaflets. Flowers are produced i ...
'' (Osti's peony) **''
Paeonia qiui ''Paeonia qiui'' is a species of peony very similar to '' P. jishanensis'', but with more divided foliage. It can reach up to 1.2m in height. Identification characteristics: petals with a red blotch at base; leaflets mostly entire Entire may re ...
'' (Qiu's peony) **''
Paeonia rockii ''Paeonia rockii'', or Rock's peony, is a woody species of tree peony that was named after Joseph Rock. It is one of several species given the vernacular name tree peony, and is native to the mountains of Gansu and adjoining provinces in Chin ...
'' (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 Paeonia or Paionia may refer to: * The genus ''Paeonia'', which comprises all peony plants * Paeonia (kingdom), an ancient state occupying roughly the same area as the present-day Republic of North Macedonia * Paionia (municipality) Paionia ( e ...
'' 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 Asclepius (; grc-gre, Ἀσκληπιός ''Asklēpiós'' ; la, Aesculapius) is a hero and god of medicine in ancient Religion in ancient Greece, Greek religion and Greek mythology, mythology. He is the son of Apollo and Coronis (lover of ...
, the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
god In monotheism, monotheistic thought, God is usually viewed as the supreme being, creator deity, creator, and principal object of Faith#Religious views, faith.Richard Swinburne, Swinburne, R.G. "God" in Ted Honderich, Honderich, Ted. (ed)''The Ox ...
of medicine and healing. In Greek mythology, when Asclepius became jealous of his pupil,
Zeus Zeus or , , ; grc, Δῐός, ''Diós'', label=Genitive case, genitive Aeolic Greek, Boeotian Aeolic and Doric Greek#Laconian, Laconian grc-dor, Δεύς, Deús ; grc, Δέος, ''Déos'', label=Genitive case, genitive el, Δίας, ''D ...
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'', 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 sjd, Куэлнэгк нёа̄ррк , image_name= Kola peninsula.png , image_caption= Kola Peninsula as a part of Murmansk Oblast , image_size= 300px , image_alt= , map_image= Murmansk in Russia.svg , map_caption = Location of Murmansk Oblas ...
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 The Tian Shan,, , otk, 𐰴𐰣 𐱅𐰭𐰼𐰃, , tr, Tanrı Dağı, mn, Тэнгэр уул, , ug, تەڭرىتاغ, , , kk, Тәңіртауы / Алатау, , , ky, Теңир-Тоо / Ала-Тоо, , , uz, Tyan-Shan / Tangritog‘ ...
Mountains of Kazakhstan. The rest of the section concentrates around the Mediterranean, and in Asia. The species around the Mediterranean include ''
Paeonia algeriensis ''Paeonia algeriensis'' is a herbaceous species of peony that naturally occurs in the coastal mountain range of Algeria (Kabylie). It has solitary flowers with pink to magenta petals and one or two carpels per flower, that develop into follicles ...
'' that is an
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
of the coastal mountains of Algeria, '' P. coriacea'' in the
Rif Mountains The Rif or Riff (, ), also called Rif Mountains, is a geographic region in northern Morocco. This mountainous and fertile area is bordered by Cape Spartel and Tangier to the west, by Berkane and the Moulouya River to the east, by the Mediterrane ...
and
Andalusia Andalusia (, ; es, Andalucía ) is the southernmost Autonomous communities of Spain, autonomous community in Peninsular Spain. It is the most populous and the second-largest autonomous community in the country. It is officially recognised as a ...
, '' 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 Sardinia ( ; it, Sardegna, label=Italian, Corsican and Tabarchino ; sc, Sardigna , sdc, Sardhigna; french: Sardaigne; sdn, Saldigna; ca, Sardenya, label=Algherese and Catalan) is the second-largest island in the Mediterranean Sea, after ...
and
Sicilly Sicily ( it, Sicilia , ) is the list of islands in the Mediterranean, largest island in the Mediterranean Sea and one of the 20 regions of Italy, regions of Italy. The Strait of Messina divides it from the region of Calabria in Southern Italy. I ...
, '' P. corsica'' on Corsica, Sardinia, the Ionian islands and in western Greece'', P. clusii'' subsp. ''clusii'' on
Crete Crete ( el, Κρήτη, translit=, Modern: , Ancient: ) is the largest and most populous of the Greek islands, the 88th largest island in the world and the fifth largest island in the Mediterranean Sea, after Sicily, Sardinia, Cyprus, and ...
and
Karpathos Karpathos ( el, Κάρπαθος, ), also Carpathos, is the second largest of the Greek Dodecanese islands, in the southeastern Aegean Sea. Together with the neighboring smaller Saria Island it forms the municipality of Karpathos, which is part o ...
, and subsp. ''rhodia'' on
Rhodes Rhodes (; el, Ρόδος , translit=Ródos ) is the largest and the historical capital of the Dodecanese islands of Greece. Administratively, the island forms a separate municipality within the Rhodes regional unit, which is part of the So ...
, '' P. kesrouanensis'' in the Western
Taurus Mountains The Taurus Mountains ( Turkish: ''Toros Dağları'' or ''Toroslar'') are a mountain complex in southern Turkey, separating the Mediterranean coastal region from the central Anatolian Plateau. The system extends along a curve from Lake Eğirdir ...
, '' P. arietina'' from the Middle Taurus Mountains, '' P. broteri'' in Andalucia, '' P. humilis'' from Andalucia to the
Provence Provence (, , , , ; oc, Provença or ''Prouvènço'' , ) is a geographical region and historical province of southeastern France, which extends from the left bank of the lower Rhône to the west to the Italian border to the east; it is bor ...
, '' 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 Catalonia (; ca, Catalunya ; Aranese Occitan: ''Catalonha'' ; es, Cataluña ) is an autonomous community of Spain, designated as a ''nationality'' by its Statute of Autonomy. Most of the territory (except the Val d'Aran) lies on the north ...
and southern France to Israel and Turkey. Between the two concentrations, the subspecies of ''
Paeonia daurica ''Paeonia daurica'' is a perennial herbaceous plant belonging to the peony family. It has slender carrot-shaped roots, leaves mostly consisting of nine leaflets, with one flower per stem. The flower is subtended by none to two leafy bracts, and ...
'' 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 The Caucasus () or Caucasia (), is a region between the Black Sea and the Caspian Sea, mainly comprising Armenia, Azerbaijan, Georgia, and parts of Southern Russia. The Caucasus Mountains, including the Greater Caucasus range, have historically ...
, Kaçkar and
Alborz The Alborz ( fa, البرز) range, also spelled as Alburz, Elburz or Elborz, is a mountain range in northern Iran that stretches from the border of Azerbaijan along the western and entire southern coast of the Caspian Sea and finally runs nort ...
Mountains. ''
Paeonia emodi ''Paeonia emodi'' is a robust herbaceous perennial plant that winters with buds underground (a so-called hemicryptophyte), has large white flowers and large deeply incised leaves, and belongs to the family Paeoniaceae. Its local vernacular name ...
'' 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 Qinghai (; alternately romanized as Tsinghai, Ch'inghai), also known as Kokonor, is a landlocked province in the northwest of the People's Republic of China. It is the fourth largest province of China by area and has the third smallest po ...
,
Ningxia Ningxia (,; , ; alternately romanized as Ninghsia), officially the Ningxia Hui Autonomous Region (NHAR), is an autonomous region in the northwest of the People's Republic of China. Formerly a province, Ningxia was incorporated into Gansu in ...
,
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 Shaanxi (alternatively Shensi, see #Name, § Name) is a landlocked Provinces of China, province of China. Officially part of Northwest China, it borders the province-level divisions of Shanxi (NE, E), Henan (E), Hubei (SE), Chongqing (S), Sichu ...
,
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 Sichuan (; zh, c=, labels=no, ; zh, p=Sìchuān; alternatively romanized as Szechuan or Szechwan; formerly also referred to as "West China" or "Western China" by Protestant missions) is a province in Southwest China occupying most of the ...
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 Manchuria is an exonym (derived from the endo demonym " Manchu") for a historical and geographic region in Northeast Asia encompassing the entirety of present-day Northeast China (Inner Manchuria) and parts of the Russian Far East (Outer Manc ...
, Korea, Japan, Far Eastern Russia (
Primorsky Krai Primorsky Krai (russian: Приморский край, r=Primorsky kray, p=prʲɪˈmorskʲɪj kraj), informally known as Primorye (, ), is a federal subject (a krai) of Russia, located in the Far East region of the country and is a part of the ...
) and on
Sakhalin Sakhalin ( rus, Сахали́н, r=Sakhalín, p=səxɐˈlʲin; ja, 樺太 ''Karafuto''; zh, c=, p=Kùyèdǎo, s=库页岛, t=庫頁島; Manchu: ᠰᠠᡥᠠᠯᡳᠶᠠᠨ, ''Sahaliyan''; Orok: Бугата на̄, ''Bugata nā''; Nivkh: ...
, 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 A crossbreed is an organism with purebred parents of two different breeds, varieties, or populations. ''Crossbreeding'', sometimes called "designer crossbreeding", is the process of breeding such an organism, While crossbreeding is used to main ...
of species currently limited to eastern Asia. The large distance between the ranges of the parent species and the
nothospecies #REDIRECT Hybrid name In botanical nomenclature, a hybrid may be given a hybrid name, which is a special kind of botanical name, but there is no requirement that a hybrid name should be created for plants that are believed to be of hybrid origin. ...
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 Confucius ( ; zh, s=, p=Kǒng Fūzǐ, "Master Kǒng"; or commonly zh, s=, p=Kǒngzǐ, labels=no; – ) was a Chinese philosopher and politician of the Spring and Autumn period who is traditionally considered the paragon of Chinese sages. C ...
(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 The Tang dynasty (, ; zh, t= ), or Tang Empire, was an Dynasties in Chinese history, imperial dynasty of China that ruled from 618 to 907 AD, with an Zhou dynasty (690–705), interregnum between 690 and 705. It was preceded by the Sui dyn ...
, 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 The Song dynasty (; ; 960–1279) was an imperial dynasty of China that began in 960 and lasted until 1279. The dynasty was founded by Emperor Taizu of Song following his usurpation of the throne of the Later Zhou. The Song conquered the rest ...
,
Luoyang Luoyang is a city located in the confluence area of Luo River (Henan), Luo River and Yellow River in the west of Henan province. Governed as a prefecture-level city, it borders the provincial capital of Zhengzhou to the east, Pingdingshan to the ...
, was the centre for its cultivation, a position it still holds today. A second centre for peony cultivation developed during the
Qing dynasty The Qing dynasty ( ), officially the Great Qing,, was a Manchu-led imperial dynasty of China and the last orthodox dynasty in Chinese history. It emerged from the Later Jin dynasty founded by the Jianzhou Jurchens, a Tungusic-speak ...
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
crossbreed A crossbreed is an organism with purebred parents of two different breeds, varieties, or populations. ''Crossbreeding'', sometimes called "designer crossbreeding", is the process of breeding such an organism, While crossbreeding is used to main ...
ing, 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 period Shōwa may refer to: * Hirohito (1901–1989), the 124th Emperor of Japan, known posthumously as Emperor Shōwa * Showa Corporation, a Japanese suspension and shock manufacturer, affiliated with the Honda keiretsu Japanese eras * Jōwa (Heian ...
s). 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 Kew Gardens is a botanical garden, botanic garden in southwest London that houses the "largest and most diverse botany, botanical and mycology, mycological collections in the world". Founded in 1840, from the exotic garden at Kew Park, its li ...
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 accomplishm ...
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 A primordium (; plural: primordia; synonym: anlage) in embryology, is an Organ (anatomy), organ or tissue in its earliest recognizable stage of development. Cell (biology), Cells of the primordium are called primordial cells. A primordium is the ...
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
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...
s have gained the
Royal Horticultural Society The Royal Horticultural Society (RHS), founded in 1804 as the Horticultural Society of London, is the UK's leading gardening charity. The RHS promotes horticulture through its five gardens at Wisley (Surrey), Hyde Hall (Essex), Harlow Carr (Nort ...
'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 medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
s of
Korea Korea ( ko, 한국, or , ) is a peninsular region in East Asia. Since 1945, it has been divided at or near the 38th parallel, with North Korea (Democratic People's Republic of Korea) comprising its northern half and South Korea (Republic o ...
,
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 Chi ...
'' used to be called ''ebisugusuri'' ("foreign medicine"). Pronunciation of 牡丹 (peony) in Japan is "botan." In ''
kampo , often known simply as , is the study of traditional Chinese medicine in Japan following its introduction, beginning in the 7th century. It was adapted and modified to suit Japanese culture and traditions. Traditional Japanese medicine use ...
'', the Japanese adaptation of
Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of action ...
, its root was used as a treatment for convulsions. It is also cultivated as a
garden plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...
. In Japan ''
Paeonia suffruticosa Paeonia or Paionia may refer to: * The genus ''Paeonia'', which comprises all peony plants * Paeonia (kingdom), an ancient state occupying roughly the same area as the present-day Republic of North Macedonia * Paionia (municipality) Paionia ( e ...
'' 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 Chi ...
'' 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 Infusion is the process of extracting chemical compounds or flavors from plant material in a solvent such as water, oil or alcohol, by allowing the material to remain suspended in the solvent over time (a process often called steeping). An inf ...
of peony petals, was used for drinking in the
Middle Ages In the history of Europe, the Middle Ages or medieval period lasted approximately from the late 5th to the late 15th centuries, similar to the post-classical period of global history. It began with the fall of the Western Roman Empire a ...
. The petals may be added to salads or to punches and lemonades. Peonies are also extensively grown as
ornamental plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...
s 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 ''Prunus mume'' is an East Asian and Southeast Asian tree species classified in the ''Armeniaca'' section of the genus ''Prunus'' subgenus ''Prunus''. Its common names include Chinese plum, Japanese plum, and Japanese apricot. The flower, long ...
, it is a traditional floral symbol of China, where the ''
Paeonia suffruticosa Paeonia or Paionia may refer to: * The genus ''Paeonia'', which comprises all peony plants * Paeonia (kingdom), an ancient state occupying roughly the same area as the present-day Republic of North Macedonia * Paionia (municipality) Paionia ( e ...
'' 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 The Qing dynasty ( ), officially the Great Qing,, was a Manchu-led imperial dynasty of China and the last orthodox dynasty in Chinese history. It emerged from the Later Jin dynasty founded by the Jianzhou Jurchens, a Tungusic-speak ...
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 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 nort ...
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 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 ...
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 The National People's Congress of the People's Republic of China (NPC; ), or simply the National People's Congress, is constitutionally the supreme state authority and the national legislature of the People's Republic of China. With 2, ...
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 Luoyang is a city located in the confluence area of Luo River (Henan), Luo River and Yellow River in the west of Henan province. Governed as a prefecture-level city, it borders the provincial capital of Zhengzhou to the east, Pingdingshan to the ...
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 In the history of Europe, the Middle Ages or medieval period lasted approximately from the late 5th to the late 15th centuries, similar to the post-classical period of global history. It began with the fall of the Western Roman Empire a ...
, 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 Woodpeckers are part of the bird family Picidae, which also includes the piculets, wrynecks, and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar, and the extreme polar regions. ...
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
Serbia Serbia (, ; Serbian language, Serbian: , , ), officially the Republic of Serbia (Serbian language, Serbian: , , ), is a landlocked country in Southeast Europe, Southeastern and Central Europe, situated at the crossroads of the Pannonian Bas ...
n folklore. Known as
Kosovo Kosovo ( sq, Kosova or ; sr-Cyrl, Косово ), officially the Republic of Kosovo ( sq, Republika e Kosovës, links=no; sr, Република Косово, Republika Kosovo, links=no), is a partially recognised state in Southeast Euro ...
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 The Indiana General Assembly is the state legislature, or legislative branch, of the state of Indiana. It is a bicameral legislature that consists of a lower house, the Indiana House of Representatives, and an upper house, the Indiana Senate. ...
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 Indiana () is a U.S. state in the Midwestern United States. It is the 38th-largest by area and the 17th-most populous of the 50 States. Its capital and largest city is Indianapolis. Indiana was admitted to the United States as the 19th s ...
, a title which it holds to this day. It replaced the
zinnia ''Zinnia'' is a genus of plants of the tribe Heliantheae within the family Asteraceae. They are native to scrub and dry grassland in an area stretching from the Southwestern United States to South America, with a centre of diversity in Mexico. ...
, 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 Floriography (language of flowers) is a means of cryptological communication through the use or arrangement of flowers. Meaning has been attributed to flowers for thousands of years, and some form of floriography has been practiced in tradition ...
. 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
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s 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 (also spelled or sometimes ) is the Japanese word for tattoo, and is used in English to refer to a distinctive style of Japanese tattooing, though it is also used as a blanket term to describe a number of tattoo styles originating in Japan, inc ...
was inspired by the
ukiyo-e Ukiyo-e is a genre of Japanese art which flourished from the 17th through 19th centuries. Its artists produced woodblock prints and paintings Painting is the practice of applying paint, pigment, color or other medium to a solid surfac ...
artist
Utagawa Kuniyoshi Utagawa Kuniyoshi ( ja, 歌川 国芳, ; January 1, 1798 – April 14, 1861) was one of the last great masters of the Japanese ukiyo-e style of woodblock prints and painting.Nussbaum, Louis Frédéric ''et al'' (2005). "Kuniyoshi" in He was a ...
'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 Conrad Gessner (; la, Conradus Gesnerus 26 March 1516 – 13 December 1565) was a Swiss physician, naturalist, bibliographer, and philologist. Born into a poor family in Zürich, Switzerland, his father and teachers quickly realised his tale ...
(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 Martin Schongauer (c. 1450–53, Colmar – 2 February 1491, Breisach), also known as Martin Schön ("Martin beautiful") or Hübsch Martin ("pretty Martin") by his contemporaries, was an Alsatian engraver and painter. He was the most important ...
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 The Qing dynasty ( ), officially the Great Qing,, was a Manchu-led imperial dynasty of China and the last orthodox dynasty in Chinese history. It emerged from the Later Jin dynasty founded by the Jianzhou Jurchens, a Tungusic-speak ...
, 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