The peony or paeony () is any

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...

in the

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

''Paeonia'', the only genus in the

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

Paeoniaceae. Peonies are native to

Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

, and Western

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

. Scientists differ on the number of

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

that can be distinguished, ranging from 25 to 40, although the current consensus describes 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 ...

perennial plant In horticulture, the term perennial (''wikt:per-#Prefix, per-'' + ''wikt:-ennial#Suffix, -ennial'', "through the year") is used to differentiate a plant from shorter-lived annual plant, annuals and biennial plant, biennials. It has thus been d ...

s tall, but some are woody

shrub A shrub or 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 by their multiple ...

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 struc ...

, deeply lobed leaves and large, often fragrant flowers, in colors ranging from purple and pink to red, white or yellow, in late spring and early summer. The flowers have a short blooming season, usually lasting for only 7–10 days. Peonies are popular garden plants in temperate regions. Herbaceous peonies are also sold as

cut flowers Cut flowers are flowers and flower buds (often with some Plant stem, stem and leaf) that have been cut from the plant bearing it. It is removed from the plant for decorative use. Cut greens are leaves with or without stems added to the cut flow ...

on a large scale, although they generally are only available in late spring and early summer.

Description

Morphology

All Paeoniaceae are

perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...

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 Leaf, 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 ...

(tufted), because the crown produces adventitious buds, while others have

stolon In biology, a stolon ( from Latin ''wikt:stolo, stolō'', genitive ''stolōnis'' – "branch"), also known as a runner, is a horizontal connection between parts of an organism. It may be part of the organism, or of its skeleton. Typically, animal ...

s. They have rather large compound leaves without

gland A gland is a Cell (biology), cell or an Organ (biology), organ in an animal's body that produces and secretes different substances that the organism needs, either into the bloodstream or into a body cavity or outer surface. A gland may also funct ...

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 (botany), petiole). They are primarily found among dicots and rare among monocots. Stipules are considered part ...

s, and with anomocytic

stoma In botany, a stoma (: stomata, from Greek language, Greek ''στόμα'', "mouth"), also called a stomate (: stomates), is a pore found in the Epidermis (botany), epidermis of leaves, stems, and other organs, that controls the rate of gas exc ...

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, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also look ...

, 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 (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...

s, with

anther The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...

s fixed at their base to the filaments, and are sagittate in shape, open with longitudinal slits at the outer side and free

pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...

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 Disc or disk may refer to: * Disk (mathematics), a two dimensional shape, the interior of a circle * Disk storage * Optical disc * Floppy disk Music * Disc (band), an American experimental music band * ''Disk'' (album), a 1995 EP by Moby Other ...

, which is presumed not to excrete

nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...

. Within the disk is a varying number (1-15) of separate

carpel Gynoecium (; ; : gynoecia) 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 ...

s, which have a very short

style Style, or styles may refer to: Film and television * ''Style'' (2001 film), a Hindi film starring Sharman Joshi, Riya Sen, Sahil Khan and Shilpi Mudgal * ''Style'' (2002 film), a Tamil drama film * ''Style'' (2004 film), a Burmese film * '' ...

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'',
spring buds File:Paeonia ludlowii Spring bud elongation lateral view.jpg, ''P. ludlowii'',
spring shoot File:Paeonia vetchii feuille.jpg, ''Paeonia veitchii'',
leaf File:Paeonia 2015-04-16 183.jpg, ''Paeonia tenuifolia'',
leaves and flower buds File:Paeonia wittmanniana Fruits BOGA.jpg, ''Paeonia wittmanniana'',
ripe follicles with seeds File:Peony anomil.jpg, ''Paeonia anomala'',
seeds File:Paeonia.jpg, Young growth of a tree peony

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 chemically 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. Th ...

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 ...

tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and Precipitation (chemistry), precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' is widel ...

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 Chalconoid, chalcones. Most stilbenoids ...

triterpenoid Triterpenes are a class of terpenes composed of six isoprene units with the molecular formula C30H48; they may also be thought of as consisting of three terpene units. Animals, plants and fungi all produce triterpenes, including squalene, the pre ...

steroid A steroid is an organic compound with four fused compound, fused rings (designated A, B, C, and D) arranged in a specific molecular configuration. Steroids have two principal biological functions: as important components of cell membranes t ...

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 eff ...

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 ar ...

. ''

In vitro ''In vitro'' (meaning ''in glass'', or ''in the glass'') Research, studies are performed with Cell (biology), cells or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in ...

'' 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, the other two being and CAM. This process converts carbon dioxide and ribulose bisphosphate (RuBP, a 5-carbon sugar) into two molecules of ...

. 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 backward because it can be o ...

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 stru ...

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 substituted derivatives, ...

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 or . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydr ...

. The wax tubules that are formed primarily consist of palmitone (the

ketone In organic chemistry, a ketone is an organic compound with the structure , where R and R' can be a variety of carbon-containing substituents. Ketones contain a carbonyl group (a carbon-oxygen double bond C=O). The simplest ketone is acetone ( ...

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 ...

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 k ...

. 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: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

Saxifragales Saxifragales is an order (biology), order of flowering plants in the Superrosids, superrosid clade of the eudicots. It contains 15 Families (biology), families and around 100 genera, with nearly 2,500 species. Well-known and economically import ...

. 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. californica''. The section ''Moutan'' is divided into ''P. delavayi'' and ''P. ludlowii'', together making up the subsection ''Delavayanae'', and ''P. cathayana'', ''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 can ...

. 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 "unispe ...

family Paeoniaceae to a group of families with woody species in the order

. 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 ...

species (about 30 species) **''

Paeonia algeriensis ''Paeonia algeriensis'', also known as the ''Algerian peony'', 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 p ...

'' (Algerian peony) **'' Paeonia anomala'' **''

Paeonia turcica ''Paeonia turcica'', the Turkish peony, is a flowering plant in the family Paeoniaceae. It is also called the "bear rose" in Turkey. It is native to the mountainous regions of south-western Turkey, particularly in Caria and Lycia. This rare per ...

'' (Turkish peony) **'' Paeonia arietina'' (ram's horn peony) **'' Paeonia broteri'' (Brotero's peony) **'' Paeonia brownii'' (Brown's peony, native peony, or western 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 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 the leaves turns into a metallic b ...

'' (Majorcan peony or Balearic peony) **'' Paeonia clusii'' ***subsp. ''clusii'' ***subsp. ''rhodia'' **'' Paeonia coriacea'' (Andalusian peony) **'' Paeonia corsica'' (Corsican peony) **''

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 ...

'' (Crimean peony) ***subsp. ''coriifolia'' ***subsp. ''daurica'' ***subsp. '' macrophylla'' ***subsp. '' mlokosewitschii'' (Caucasian peony or golden peony) ***subsp. ''tomentosa'' ***subsp. ''velebitensis'' ***subsp. '' wittmanniana'' (Wittmann's peony) **'' Paeonia emodi'' (Himalayan peony) **'' Paeonia intermedia'' (Intermediate peony or Altay peony) **''

Paeonia kesrouanensis ''Paeonia kesrouanensis'' is a species of peony native to Syria, Turkey and Lebanon Lebanon, officially the Republic of Lebanon, is a country in the Levant region of West Asia. Situated at the crossroads of the Mediterranean Basin and th ...

'' (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 ...

'' (Chinese peony, Chinese herbaceous peony, or common garden 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 peony, wild peony, or male peony) ***subsp. ''mascula'' ***subsp. ''bodurii'' ***subsp. ''hellenica'' ***subsp. ''russoi'' **''

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 woodla ...

'' (woodland peony) ***subsp. ''willmottiae'' **''

Paeonia officinalis ''Paeonia officinalis'', the common peony, or garden peony, is a species of flowering plant in the family (biology), family Paeoniaceae, native plant, native to mainly mountainous areas of Southern Europe and introduced in Central Europe, Centra ...

'' (European peony, common peony, or garden peony; type species) **''

Paeonia parnassica ''Paeonia parnassica'', the Greek peony, is a flowering plant in the family Paeoniaceae. It is native to the mountains of south-central Greece. This endangered perennial species is characterized by its dark, almost blackish-red flowers measurin ...

'' (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 ...

'' (Kosovar peony or Romanian peony) **'' Paeonia sterniana'' **''

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, with large fields found in Vashlivani National Park in Georgia and the Black Sea coast of Ukr ...

'' (steppe peony or fern leaf peony) **'' Paeonia veitchii'' (Veitch's peony) * Woody species, tree peonies (about 8 species) **'' Paeonia decomposita'' **''

Paeonia delavayi ''Paeonia delavayi'' is a low woody shrub belonging to the peony family, and 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. ...

'' (Delavay's tree peony or Dian peony) **'' Paeonia jishanensis'' (Jishan peony) **''

Paeonia ludlowii ''Paeonia ludlowii'' is a deciduous shrub of medium height, belonging to the tree peony section ''Moutan'' of the genus Paeonia, and endemic to southeast Tibet. In Tibet it is known as ''≠'lumaidao meaning "God's flower". The vernacular name ...

'' (Tibetan tree peony or Ludlow's tree peony) **'' Paeonia ostii'' (Osti's peony) **'' Paeonia qiui'' (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 China. ...

'' (Rock's peony;

synonym A synonym is a word, morpheme, or phrase that means precisely 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 a ...

Paeonia suffruticosa Paeonia or Paionia may refer to: * The genus ''Paeonia'', which comprises all peony The peony or paeony () is any flowering plant in the genus ''Paeonia'', the only genus in the family Paeoniaceae. Peonies are native to Asia, Europe, and Wes ...

'' 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'' subsp. ''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 mascula'' subsp. ''russoi'' File:Paeonia peregrina kz01.jpg, ''Paeonia peregrina'' File:Paeonia japonica.JPG, ''Paeonia obovata'' subsp. ''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''

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 The Sierra Nevada ( ) is a mountain range in the Western United States, between the Central Valley of California and the Great Basin. The vast majority of the range lies in the state of California, although the Carson Range spur lies primari ...

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 The Kola Peninsula (; ) is a peninsula in the extreme northwest of Russia, and one of the largest peninsulas of Europe. Constituting the bulk of the territory of Murmansk Oblast, it lies almost completely inside the Arctic Circle and is border ...

in North-West Russia, to

Lake Baikal Lake Baikal is a rift lake and the deepest lake in the world. It is situated in southern Siberia, Russia between the Federal subjects of Russia, federal subjects of Irkutsk Oblast, Irkutsk Oblasts of Russia, Oblast to the northwest and the Repu ...

in Siberia and South to the

Tien Shan The Tian Shan, also known as the Tengri Tagh or Tengir-Too, meaning the "Mountains of God/Heaven", is a large system of mountain ranges in Central Asia. The highest peak is Jengish Chokusu at high and located in Kyrgyzstan. Its lowest point is ...

Mountains of Kazakhstan. The rest of the section concentrates around the Mediterranean, and in Asia. The species around the Mediterranean include ''

'' that is an

endemic Endemism is the state of a species being found only 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 foun ...

of the coastal mountains of Algeria, ''P. coriacea'' in the

Rif Mountains The Rif (, ), also called Rif Mountains, is a geographic region in northern Morocco. It is bordered on the north by the Mediterranean Sea and Spain and on the west by the Atlantic Ocean, and is the homeland of the Rifians and the Jebala people. ...

and

Andalusia Andalusia ( , ; , ) is the southernmost autonomous communities of Spain, autonomous community in Peninsular Spain, located in the south of the Iberian Peninsula, in southwestern Europe. It is the most populous and the second-largest autonomou ...

, ''P. cambessedesii'' on

Majorca Mallorca, or Majorca, is the largest of the Balearic Islands, which are part of Spain, and the List of islands in the Mediterranean#By area, seventh largest island in the Mediterranean Sea. The capital of the island, Palma, Majorca, Palma, i ...

, ''P. russoi'' on

Corsica Corsica ( , , ; ; ) is an island in the Mediterranean Sea and one of the Regions of France, 18 regions of France. It is the List of islands in the Mediterranean#By area, fourth-largest island in the Mediterranean and lies southeast of the Metro ...

Sardinia Sardinia ( ; ; ) is the Mediterranean islands#By area, second-largest island in the Mediterranean Sea, after Sicily, and one of the Regions of Italy, twenty regions of Italy. It is located west of the Italian Peninsula, north of Tunisia an ...

and

Sicily Sicily (Italian language, Italian and ), officially the Sicilian Region (), is an island in the central Mediterranean Sea, south of the Italian Peninsula in continental Europe and is one of the 20 regions of Italy, regions of Italy. With 4. ...

, ''P. corsica'' on Corsica, Sardinia, the

Ionian islands The Ionian Islands (Modern Greek: , ; Ancient Greek, Katharevousa: , ) are a archipelago, group of islands in the Ionian Sea, west of mainland Greece. They are traditionally called the Heptanese ("Seven Islands"; , ''Heptanēsa'' or , ''Heptanē ...

and in western Greece, ''P. clusii'' subsp. ''clusii'' on

Crete Crete ( ; , Modern Greek, Modern: , Ancient Greek, Ancient: ) is the largest and most populous of the Greek islands, the List of islands by area, 88th largest island in the world and the List of islands in the Mediterranean#By area, fifth la ...

and

Karpathos Karpathos (, ), 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 of the regional unit ...

, and subsp. ''rhodia'' on

Rhodes Rhodes (; ) is the largest of the Dodecanese islands of Greece and is their historical capital; it is the List of islands in the Mediterranean#By area, ninth largest island in the Mediterranean Sea. Administratively, the island forms a separ ...

, ''P. kesrouanensis'' in the Western

Taurus Mountains The Taurus Mountains (Turkish language, Turkish: ''Toros Dağları'' or ''Toroslar,'' Greek language, Greek'':'' Ταύρος) are a mountain range, mountain complex in southern Turkey, separating the Mediterranean Sea, Mediterranean coastal reg ...

, ''P. arietina'' from the Middle Taurus Mountains, ''P. broteri'' in Andalucia, ''P. humilis'' from Andalucia to the

Provence Provence is a geographical region and historical province of southeastern France, which stretches from the left bank of the lower Rhône to the west to the France–Italy border, Italian border to the east; it is bordered by the Mediterrane ...

, ''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 is an autonomous community of Spain, designated as a ''nationalities and regions of Spain, nationality'' by its Statute of Autonomy of Catalonia of 2006, Statute of Autonomy. Most of its territory (except the Val d'Aran) is situate ...

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 The Caucasus () or Caucasia (), is a region spanning Eastern Europe and Western Asia. It is situated between the Black Sea and the Caspian Sea, comprising parts of Southern Russia, Georgia, Armenia, and Azerbaijan. The Caucasus Mountains, i ...

, Kaçkar and

Alborz The Alborz ( ) 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 northeast and merge ...

Mountains. ''Paeonia emodi'' occurs in the western

Himalayas The Himalayas, or Himalaya ( ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the Earth's highest peaks, including the highest, Mount Everest. More than list of h ...

between Pakistan and western Nepal, ''P. sterniana'' is an endemic of southeastern Tibet, ''P. veitchii'' grows in Central China (

Qinghai Qinghai is an inland Provinces of China, province in Northwestern China. It is the largest provinces of China, province of China (excluding autonomous regions) by area and has the third smallest population. Its capital and largest city is Xin ...

Ningxia Ningxia, officially the Ningxia Hui Autonomous Region, is an autonomous region in Northwestern China. Formerly a province, Ningxia was incorporated into Gansu in 1954 but was later separated from Gansu in 1958 and reconstituted as an autonomous ...

Gansu Gansu is a provinces of China, province in Northwestern China. Its capital and largest city is Lanzhou, in the southeastern part of the province. The seventh-largest administrative district by area at , Gansu lies between the Tibetan Plateau, Ti ...

Shaanxi Shaanxi is a Provinces of China, province in north Northwestern China. It borders the province-level divisions of Inner Mongolia to the north; Shanxi and Henan to the east; Hubei, Chongqing, and Sichuan to the south; and Gansu and Ningxia to t ...

Shanxi Shanxi; Chinese postal romanization, formerly romanised as Shansi is a Provinces of China, province in North China. Its capital and largest city of the province is Taiyuan, while its next most populated prefecture-level cities are Changzhi a ...

Sichuan Sichuan is a province in Southwestern China, occupying the Sichuan Basin and Tibetan Plateau—between the Jinsha River to the west, the Daba Mountains to the north, and the Yunnan–Guizhou Plateau to the south. Its capital city is Cheng ...

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 a historical region in northeast Asia encompassing the entirety of present-day northeast China and parts of the modern-day Russian Far East south of the Uda (Khabarovsk Krai), Uda River and the Tukuringra-Dzhagdy Ranges. The exact ...

, Korea, Japan, Far Eastern Russia (

Primorsky Krai Primorsky Krai, informally known as Primorye, is a federal subjects of Russia, federal subject (a krais of Russia, krai) of Russia, part of the Far Eastern Federal District in the Russian Far East. The types of inhabited localities in Russia, ...

) and on

Sakhalin Sakhalin ( rus, Сахали́н, p=səxɐˈlʲin) is an island in Northeast Asia. Its north coast lies off the southeastern coast of Khabarovsk Krai in Russia, while its southern tip lies north of the Japanese island of Hokkaido. An islan ...

, 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. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...

or tetraploid hybrids that resulted from the

crossbreeding A crossbreed is an organism with purebred parents of two different breeds, varieties, or populations. A domestic animal of unknown ancestry, where the breed status of only one parent or grandparent is known, may also be called a crossbreed though ...

of species currently limited to eastern Asia. The large distance between the ranges of the parent species and the

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 closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...

and so produced the Himalayan nothospecies ''P. emodi'' and ''P. sterniana''.

Cultivation

Ancient Chinese texts mention the peony was used for flavoring food. Peonies have been used and cultivated in China since early history. Ornamental cultivars were created from plants cultivated for medicine in China as of the sixth and seventh century. Peonies became particularly popular during the

Tang dynasty The Tang dynasty (, ; zh, c=唐朝), or the Tang Empire, was an Dynasties of China, imperial dynasty of China that ruled from 618 to 907, with an Wu Zhou, interregnum between 690 and 705. It was preceded by the Sui dynasty and followed ...

, 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 ( ) was an Dynasties of China, imperial dynasty of China that ruled from 960 to 1279. The dynasty was founded by Emperor Taizu of Song, who usurped the throne of the Later Zhou dynasty and went on to conquer the rest of the Fiv ...

Luoyang Luoyang ( zh, s=洛阳, t=洛陽, p=Luòyáng) is a city located in the confluence area of the Luo River and the Yellow River in the west of Henan province, China. Governed as a prefecture-level city, it borders the provincial capital of Zheng ...

, 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 Dynasties of China, imperial dynasty of China and an early modern empire in East Asia. The last imperial dynasty in Chinese history, the Qing dynasty was preceded by the ...

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. A domestic animal of unknown ancestry, where the breed status of only one parent or grandparent is known, may also be called a crossbreed though ...

ing, particularly during the eighteenth to twentieth centuries (middle

Edo Edo (), also romanized as Jedo, Yedo or Yeddo, is the former name of Tokyo. Edo, formerly a (castle town) centered on Edo Castle located in Musashi Province, became the '' de facto'' capital of Japan from 1603 as the seat of the Tokugawa shogu ...

to early

Shōwa period Shōwa most commonly refers to: * Hirohito (1901–1989), the 124th Emperor of Japan, known posthumously as Emperor Shōwa ** Shōwa era (昭和), the era of Hirohito from 1926 to 1989 * Showa Corporation, a Japanese suspension and shock manufactu ...

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 1759, 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 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

While the peony takes several years to re-establish itself when moved, it blooms annually for decades once it has done so.The Language of Flowers, edited by Sheila Pickles, 1990 Peony bud and ants

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. 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 peony, 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

Seven 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 staminodes, 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. * triple: the flower consists of triple row of broad petals that broaden and overlap each other. * 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.

Cultivars

The numerous peony hybrids and cultivars have gained the Royal Horticultural Society's Award of Garden Merit, including: *'Bartzella', a double yellow-flowered Itoh (intersectional) peony *'Coral Charm', a #Semi-double, semi-double salmon-pink-flowered herbaceous peony *''Paeonia'' × ''festiva'' 'Rubra Plena', a #Bomb, bomb red-flowered herbaceous peony *''Paeonia'' × ''lemoinei'' 'High Noon', a semi-double yellow-flowered tree peony The American Peony Society is the International Cultivar Registration Authority for the genus, and accepts over 7,000 registered cultivars.

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 and Japan. In Japan, ''

'' used to be called ''ebisugusuri'' ("foreign medicine"). Pronunciation of 牡丹 (peony) in Japan is "botan." In ''kampo'', the Japanese adaptation of traditional Chinese medicine, Chinese medicine, its root was used as a treatment for convulsions. It is also cultivated as a garden plant. In Japan ''

'' is called the "King of Flowers" and ''

'' 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.

Culture

The peony is among the longest-used flowers in Eastern culture. Along with the Ume, plum blossom, it is a traditional floral symbol of China, where the ''

'' 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

declared the peony as the national flower. Currently, the Republic of China government in Taiwan designates the plum blossom as the national flower, 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

has a reputation as a cultivation centre for the peonies. Throughout Chinese history, peonies in Luoyang have been said to be the finest in the country. Dozens of peony exhibitions and shows are still held there annually. The Greek doctor Dioscorides named aglaophotis, a herb supposedly capable of warding off certain evils, as a member of the peony family. 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 ''

'' are important in Serbian folklore. Known as Kosovo peonies (, ), they are said to represent the blood of Serbian warriors who died in the Battle of Kosovo. In 1957, the Indiana General Assembly passed a law to make the peony the list of state flowers, state flower 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, Language of Flowers. 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 ''Water Margin, Suikoden'', 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 in 1879. ''Paeonia officinalis'' can be found in the altar picture of ''Maria im Rosenhag'' by Schongauer in the former Dominican Church in Colmar. The Italian Jesuit, painter and architect Giuseppe Castiglione (Jesuit), Giuseppe Castiglione (1688-1766), who worked at the court of the Qianlong Emperor in the

, also 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 Peonies, Symbols of Indiana