and for lower taxa see text
The range of Paeonia.
The peony or paeony is a flowering plant in the genus Paeonia,
the only genus in the family Paeoniaceae. They are native to Asia,
Europe and Western North America. Scientists differ on the number of
species that can be distinguished ranging from 25 to 40,
although the current consensus is 33 known species. The
relationships between the species need to be further clarified.
Most are herbaceous perennial plants 0.25–1 metre
(0.82–3.28 ft) tall, but some are woody shrubs 0.25–3.5
metres (0.82–11.48 ft) tall. They have compound, deeply lobed
leaves and large, often fragrant flowers, in colors ranging from
purple red to white or yellow, in late spring and early summer.
Peonies are among the most popular garden plants in temperate regions.
Herbaceous peonies are also sold as cut flower on a large scale,
although generally only available in late spring and early summer.
3.1 Distributional history
4 Chemistry and biological activity
Plant growth habits
5.2 Flower types
9 External links
All Paeoniaceae are deciduous perennial herbs or shrubs, with thick
storage roots and thin roots for gathering water and minerals. Some
species are caespitose, because the crown produces adventitous buds,
while others have stolons. They have rather large compound leaves
without glands and stipules, and with anomocytic stomata. 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, 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 stamens, with anthers fixed at their base
to the filaments, and are sagittate in shape, open with longitudal
slits at the outer side and free pollen 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. Within the disk is a varying number (1-15) of separate
carpels, which have a very short style and a decurrent stigma. Each of
these develops into a dry fruit (which is called a follicle), which
opens with a lengthwise suture and each of which contains one or a few
large fleshy seeds. The annual growth is predetermined: if the growing
tip of a shoot is removed, no new buds will develop that
young growth of
a tree peony
leaves and flower buds
Paeonia suffruticosa, showing the disk that encloses the carpels
ripe follicles with seeds
Paeoniaceae are dependent on C3 carbon fixation. They contain ellagic
acid, myricetin, ethereal oils and flavones, as well as crystals of
calcium oxalate. The wax tubules that are formed primarily consist of
palmitone (the ketone of palmitic acid).
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
2017-06-25 1655 Peony
The family name "Paeoniaceae" was first used by Friedrich K.L.
Rudolphi in 1830, following a suggestion by Friedrich Gottlieb
Bartling that same year. The family had been given other names a
few years earlier. The composition of the family has varied, but
it has always consisted of Paeonia and one or more genera that are now
placed in Ranunculales. It has been widely believed that Paeonia is
closest to Glaucidium, and this idea has been followed in some recent
Molecular phylogenetic studies, however, have
demonstrated conclusively that Glaucidium belongs in the Ranunculaceae
Ranunculales order, but that Paeonia belongs in the
unrelated order Saxifragales. The genus Paeonia consists of about
35 species, assigned to three sections: Moutan, Onaepia and Paeoniae.
The section Onaepia only includes P. brownii and P. californicum. The
section Moutan is divided into P. delavayi and P. ludlowii, together
making up the subsection Delavayanae, and P. catayana, P. decomposita,
P. jishanensis, P. osti, P. qiui and P. rockii which constitute the
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. Only about half
of the (sub)species is diploid, the other half tetraploid, while some
species both have diploid and tetraploid populations. In addition to
the tetraploids, are some diploid species also likely the result of
hybridisation, or nothospecies. Known diploid taxa in the
Paeonia-section are P. anomala, P. lactiflora, P. veitchii, P.
tenuifolia, P. emodi, P. broteri, P. cambedessedesii, P. clusii, P.
rhodia, P. daurica subsps. coriifolia, daurica, macrophylla and
mlokosewitschii. Tetraploid taxa are P. arietina, P. officinalis, P.
parnassica, P. banatica, P. russi, P. peregrina, P. coriacea, P.
mascula subsps. hellenica and mascula, and P. daurica subsps.
tomentosa and wittmanniana.
Species that have both diploid and
tetraploid populations include P. clusii, P. mairei and P. obovata. P.
anomala was proven to be a hybrid of P. lactiflora and P. veitchii,
although being a diploid with 10 chromosomes. P. emodi and P.
sterniana are diploid hybrids of P. lactiflora and P. veitchii too,
and radically different in appearance. P. russi is the tetraploid
hybrid of diploid P. lactiflora and P. mairei, while P.
cambedessedesii is the diploid hybrid of P. lactiflora, likely P.
mairei, but possibly also P. obovata. P. peregrina is the tetraploid
hybrid of P. anomala and either P. arietina, P. humilis, P.
officinalis, P. parnassica or less likely P. tenuifolia, or one of
their (now extinct) common ancestors. P. banatica is the tetraploid
hybrid of P. mairei and one of this same group. P. broteri, P.
coriacea, P. clusii, P. rhodia, P. daurica subsp. mlokosewitschi, P.
mascula subsp. hellenica and ssp. mascula, and P. daurica subsp.
wittmanniana are all descendants of hybrids of P. lactiflora and P.
According to recent genetic analyses, the monogeneric family
Paeoniaceae is related to a group of families with woody species in
the order Saxifragales. This results in the following relationship
tree. One dissertation suggests the section Onaepia branches off
earliest. 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
all Eurasian herbaceous peonies
all other tree peonies
Herbaceous species (about 30 species)
Paeonia brownii (Brown's peony)
Paeonia californica (California peony or wild peony)
Paeonia cambessedesii (Majorcan peony)
Paeonia kesrouanensis (Keserwan peony)
Paeonia lactiflora (Chinese or common garden peony)
Paeonia mascula (Balkan, wild or male peony)
Paeonia officinalis (European or common peony, type species)
Paeonia parnassica (Greek peony)
Paeonia tenuifolia (Steppe peony)
Paeonia veitchii (Veitch's peony)
Woody species (about 8 species)
Paeonia delavayi (Delavay's tree peony)
Paeonia jishanensis (Jishan peony)
Paeonia ludlowii (Ludlow's tree peony)
Paeonia ostii (Osti's peony)
Paeonia qiui (Qiu's peony)
Paeonia rockii (Rock's peony or tree peony; synonym Paeonia
suffruticosa subsp. rockii (Chinese tree peony, known as "moutan
(moutan peony)" in China))
Paeonia daurica mlokosewitschii
Paeonia obovata japonica
The peony is named after Paeon (also spelled Paean), a student of
Asclepius, the Greek god of medicine and healing. When Asclepius
became jealous of his pupil,
Zeus saved Paeon from the wrath of
Asclepius by turning him into the peony flower.
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 to Japan. One species
of the section Paeonia, P. anomala, has by far the largest
distribution, which is also north of the distribution of the other
species: from the
Kola peninsula in North-West Russia, to Lake Baikal
in Siberia and South to the
Tien Shan Mountains of Kazakhstan. The
rest of the section concentrates around the Mediterranean, and in
The species around the Mediterranean include
Paeonia algeriensis that
is an endemic of the coastal mountains of Algeria, P. coriacea in the
Rif Mountains and Andalucia, P. cambessedesii on Majorca,
P. russoi on Corsica,
Sardinia and Sicilly, P. corsica on
Corsica, Sardinia, the Ionian islands and in western Greece,
P. clusii subsp. clusii on
Crete and Karpathos, and subsp. rhodia
on Rhodes, P. kesrouanensis in the Western Taurus Mountains,
P. arietina from the Middle Taurus Mountains, P. broteri in
Andalucia, P. humilis from
Andalucia to the Provence,
P. officinalis from the South of France, through Switzerland to
the Middle of Italy, P. banatica in western Romania, northern
Serbia and Slovenia and in southern Hungary, P. peregrina in
Albania, western Bulgaria, northern Greece, western Romania, Serbia,
Montenegro and Bosnia, while P. mascula has a large distribution
Catalonia and southern France to Israel and Turkey.
Between the two concentrations, the subspecies of Paeonia daurica
occur, with subspecies velebitensis in Croatia, and daurica in the
Balkans and Crimea, while the other subspecies coriifolia,
macrophylla, mlokosewitschii, tomentosa and wittmanniana are known
from the Caucasus, Kaçkar and
Paeonia emodi occurs in the western Himalayas between Pakistan and
western Nepal, P. sterniana is an endemic of southeastern Tibet,
P. veitchii grows in Central
China (Qinghai, Ningxia, Gansu,
Sichuan and the eastern rim of Tibet), like
P. mairei (Gansu, Guizhou, Hubei, Shaanxi, Sichuan, and Yunnan),
while P. obovata grows in warm-temperate to cold China, including
Manchuria, Korea, Japan, Far Eastern Russia (Primorsky Krai) and on
Sakhalin, and P. lactiflora occurs in Northern China, including
Manchuria, Japan, Korea, Mongolia, Russia (Far East and Siberia).
The species of the section Paeonia have a disjunct distribution, with
most of the species occurring in the Mediterranean, while many others
occur in eastern Asia. Genetic analysis has shown that all
Mediterranean species are either diploid or tetraploid hybrids that
resulted from the crossbreeding of species currently limited to
eastern Asia. The large distance between the ranges of the parent
species and the nothospecies suggest that hybridisation already
occurred relatively long ago. It is likely that the parent species
occurred in the same region when the hybrids arose, and were later
exterminated by successive Pleistocene glaciations, while the
nothospecies remained in refugia to the South of Europe. During their
retreat P. lactiflora and P. mairei likely became sympatric and so
produced the Himalayan nothospecies P. emodi and P. sterniana.
Chemistry and biological activity
Over 262 compounds have been obtained so far from the plants of
Paeoniaceae. These include monoterpenoid glucosides, flavonoids,
tannins, stilbenoids, triterpenoids, steroids, paeonols, and phenols.
In vitro biological activities include antioxidant, antitumor,
antipathogenic, immune-system-modulation activities,
cardiovascular-system-protective activities and central-nervous-system
Paeonia 'Sarah Bernhardt'
Ancient Chinese texts mention the peony was used for flavoring food.
Confucius (551–479 BC) is quoted to have said: "I eat nothing
without its sauce. I enjoy it very much, because of its flavor."
Peonies have been used and cultivated in
China since early history.
Ornamental cultivars were created from plants cultivated for medicine
China as of the sixth and seventh century. Peonies became
particularly popular during the Tang dynasty, when they were grown in
the imperial gardens. In the tenth century the cultivation of peonies
spread through China, and the seat of the Sung dynasty, Luoyang, was
the centre for its cultivation, a position it still holds today. A
second centre for peony cultivation developed during the Qing dynasty
in Cáozhōu, now known as He Ze. Both cities still host annual peony
exhibitions and state-funded peony research facilities. Before the
tenth century, P. lactiflora was introduced in Japan, and over time
many varieties were developed both by self fertilisation and
crossbreeding, particularly during the eightienth to twentieth
centuries (middle Edo to early Shōwa periods). During the 1940s
Toichi Itoh succeeded in crossing tree peonies and herbaceous peonies
and so created a new class of so-called intersectional hybrids.
Although P. officinalis and its cultivars were grown in
the fifteenth century on, originally also for medicinal purposes,
intensive breeding started only in the nineteenth century when P.
lactiflora was introduced from its native
China to Europe. The tree
peony was introduced in
Europe and planted in
Kew Gardens in 1789. The
main centre of peony breeding in
Europe has been in the United
Kingdom, and particularly France. Here, breeders like Victor Lemoine
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.
Plant growth habits
Peony species come in two distinct growth habits, while hybrid
cultivars in addition may occupy an intermediate habit.
herbaceous: During summer, renewal buds develop on the underground
stem (the "crown"), particularly at the foot of the current season's
annual shoots. These renewal buds come in various sizes. Large buds
will grow into stems the following growing season, but smaller buds
remain dormant. The primordia for the leaves can already be found in
June, but the flower only starts differentiating in October, as the
annual shoots die down, completing its development in December, when
sepals, petals, stamens and pistils are all recognisable.
tree: During the summer, large buds develop at the tip of the annual
growth and near its foot. In the autumn, the leaves are shed, and the
new stems become woody and are perennial.
Itoh (or "Intersectional"): In 1948 horticulturist Toichi Itoh from
Tokyo used pollen from the yellow tree peony "Alice Harding" to
fertilize the herbaceous P. lactiflora "Katoden", which resulted in a
new category of peonies, the Itoh or intersectional cultivars. These
are herbaceous, have leaves like tree peonies, with many large flowers
from late spring to early autumn, and good peony wilt resistance. Some
of the early Itoh cultivars are "Yellow Crown", "Yellow Dream",
"Yellow Emperor" and "Yellow Heaven".
Six types of flower are generally distinguished in cultivars of
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
anemone: a single or double row of broad petals encircle narrow
incurved petal-like staminodes; fertile stamens are absent, carpels
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
double: the flower consists of many broad petals only, including those
which likely are altered stamens and carpels.
Paeonia ×arendsii "Claire de Lune",
Paeonia "Walter Mains",
Paeonia lactiflora "Bowl Of Beauty",
Paeonia lactiflora "James Kelway",
Paeonia "Ruth Clay",
Paeonia lactiflora "Da Fu Gui",
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
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.
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, Paeonia
lactiflora used to be called ebisugusuri ("foreign medicine").
Pronunciation of 牡丹 (peony) in
Japan is "botan." In kampo (the
Japanese adaptation of Chinese medicine), its root was used as a
treatment for convulsions. It is also cultivated as a garden plant. In
Paeonia suffruticosa is called the "King of Flowers" and Paeonia
lactiflora is called the "Prime Minister of Flowers."
In China, the fallen petals of
Paeonia lactiflora are parboiled and
sweetened as a tea-time delicacy.
Peony water, an infusion of peony
petals, was used for drinking in the Middle Ages. The petals may be
added to salads or to punches and lemonades.
Peonies are also extensively grown as ornamental plants for their very
large, often scented flowers.
In this gold-engraved lacquerware food tray from the Song dynasty
(960–1279), the two long-tailed birds represent longevity, and the
peony seen at the top center represents prosperity
Peony, by Chinese artist Wang Qian,
Yuan dynasty (1271–1368)
Portrait of a peony by Chinese artist Yun Shouping, 17th century
The peony is among the longest-used flowers in Eastern culture. Along
with the plum blossom, it is a traditional floral symbol of China,
Paeonia suffruticosa is called 牡丹 (mǔdān). It is also
known as 富貴花 (fùguìhuā) "flower of riches and honour" or
花王 (huawang) "king of the flowers", and is used symbolically in
Chinese art. In 1903, the
Qing dynasty declared the peony as the
national flower. Currently, the Republic of
China government in Taiwan
designates the plum blossom as the national flower, while the People's
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
National People's Congress failed to ratify
the selection. In 2003, another selection process has begun, but to
date, no choice has been made.
The ancient Chinese city
Luoyang 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.
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.
In 1957, the
Indiana General Assembly
Indiana General Assembly passed a law to make the peony
the 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. While the peony takes several years to re-establish itself
when moved, it blooms annually for decades once it has done so.
Peonies tend to attract ants to the flower buds. This is due to the
nectar that forms on the outside of the flower buds, and is not
required for the plants' own pollination or other growth.
Peonies are a common subject in tattoos, often used along with
koi-fish. The popular use of peonies in Japanese tattoo was inspired
by the ukiyo-e artist Utagawa Kuniyoshi's illustrations of Suikoden, 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
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 (1688-1766), who worked at the court of the
Qianlong Emperor in the Qing dynasty, painted peonies.
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Wikispecies has information related to Paeonia
Wikimedia Commons has media related to Paeonia.
Family and Suprafamilial Names At: James L. Reveal
Paeoniaceae in Topwalks
Flora Europaea: Paeonia
Ornamental Plants from Russia: Paeonia
Peony Society (UK) (defunct as of 2106)
Carsten Burkhardt's Open Source
China Daily article on the 2003 national flower selection process
Rockii Tree Peony
Blossoming of the