Peony

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For other uses, see Peony (disambiguation).

Peony
Paeonia suffruticosa
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Saxifragales
Family: Paeoniaceae
Raf.[1]
Genus: Paeonia
L.
Type species
Paeonia officinalis
L.
Sections
  • Moutan
  • Onaepia
  • Paeoniae

and for lower taxa see text

The range of Paeonia.

The peony or paeony (/ˈpiːəni/)[2][3] is a flowering plant in the genus Paeonia,[4] the only genus in the family Paeoniaceae. Peonies 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,[5][6] although the current consensus describes 33 known species.[7] The relationships between the species need to be further clarified.[8]

Most are

perennial plants 0.25–1 metre (1–3 ft) tall, but some are woody shrubs 0.25–3.5 metres (1–11 ft) tall. They have compound
, deeply lobed leaves and large, often fragrant flowers, in colors ranging from purple and pink to red, white or yellow, in late spring and early summer. The flowers have a short blooming season, usually only 7–10 days.

Peonies are popular garden plants in temperate regions. Herbaceous peonies are also sold as cut flowers on a large scale, although generally only available in late spring and early summer.[9]

Description

Morphology

All Paeoniaceae are

carpels, which have a very short style and a decurrent stigma. Each of these develops into a dry fruit (which is called a follicle), which opens with a lengthwise suture and each of which contains one or a few large fleshy seeds. The annual growth is predetermined: if the growing tip of a shoot is removed, no new buds will develop that season.[9][10][11]

  • Paeonia suffruticosa, Spring buds
    Paeonia suffruticosa,
    Spring buds
  • P. ludlowii Spring shoot
    P. ludlowii Spring shoot
  • young growth of a tree peony
    young growth of
    a tree peony
  • Paeonia veitchii, leaf
    Paeonia veitchii,
    leaf
  • Paeonia tenuifolia, leaves and flower buds
    Paeonia tenuifolia,
    leaves and flower buds
  • Paeonia suffruticosa, showing the disk that encloses the carpels
    Paeonia suffruticosa, showing the disk that encloses the carpels
  • Paeonia wittmanniana, ripe follicles with seeds
    Paeonia wittmanniana,
    ripe follicles with seeds
  • Paeonia anomala, seeds
    Paeonia anomala,
    seeds

Phytochemistry

Over 262 compounds have been obtained so far from the plants of Paeoniaceae. These include

triterpenoids, steroids, paeonols, and phenols. In vitro biological activities include antioxidant, antitumor, antipathogenic, immunomodulative, cardiovascular-system-protective activities and central-nervous-system activities.[12]

Paeoniaceae are dependent on

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).[10][11]

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.[10]

Taxonomy

The family name "Paeoniaceae" was first used by Friedrich K.L. Rudolphi in 1830, following a suggestion by

P. suffruticosa is a cultivated hybrid swarm, not a naturally occurring species.[17]

The remainder of the species belongs to the section Paeonia, which is characterised by a complicated

nothospecies. Known diploid taxa in the Paeonia-section are P. anomala, P. lactiflora, P. veitchii, P. tenuifolia, P. emodi, P. broteri, P. cambedessedesii, P. clusii, P. rhodia, P. daurica subsps. coriifolia, daurica, macrophylla and mlokosewitschii. Tetraploid taxa are P. arietina, P. officinalis, P. parnassica, P. banatica, P. russi, P. peregrina, P. coriacea, P. mascula subsps. hellenica and mascula, and P. daurica subsps. tomentosa and wittmanniana. Species that have both diploid and tetraploid populations include P. clusii, P. mairei and P. obovata. P. anomala was proven to be a hybrid of P. lactiflora and P. veitchii, although being a diploid with 10 chromosomes. P. emodi and P. sterniana are diploid hybrids of P. lactiflora and P. veitchii too, and radically different in appearance. P. russi is the tetraploid hybrid of diploid P. lactiflora and P. mairei, while P. cambedessedesii is the diploid hybrid of P. lactiflora, likely P. mairei, but possibly also P. obovata. P. peregrina is the tetraploid hybrid of P. anomala and either P. arietina, P. humilis, P. officinalis, P. parnassica or less likely P. tenuifolia, or one of their (now extinct) common ancestors. P. banatica is the tetraploid hybrid of P. mairei and one of this same group. P. broteri, P. coriacea, P. clusii, P. rhodia, P. daurica subsp. mlokosewitschi, P. mascula subsp. hellenica and ssp. mascula, and P. daurica subsp. wittmanniana are all descendants of hybrids of P. lactiflora and P. obovata.[17][18]

Phylogeny

Recent genetic analyses relate the monogeneric family Paeoniaceae to a group of families with woody species in the order Saxifragales. This results in the following relationship tree.[16] 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.[19][20]

Saxifragales

Peridiscaceae

 woody clade 

 core Saxifragales 

genus Paeonia
section Paeoniae

all Eurasian herbaceous peonies

section Moutan
subsection Vaginatae

all other tree peonies

subsection Delavayanae
section Onaepia

Species

  • Paeonia anomala
    Paeonia anomala
  • Paeonia broteri
    Paeonia broteri
  • Paeonia brownii
    Paeonia brownii
  • Paeonia californica
    Paeonia californica
  • Paeonia cambessedesii
    Paeonia cambessedesii
  • Paeonia cambessedesii
    Paeonia cambessedesii
  • Paeonia daurica subsp. mlokosewitschii
    Paeonia daurica subsp. mlokosewitschii
  • Paeonia delavayi
    Paeonia delavayi
  • Paeonia emodi
    Paeonia emodi
  • Paeonia intermedia
    Paeonia intermedia
  • Paeonia ludlowii
    Paeonia ludlowii
  • Paeonia mairei
    Paeonia mairei
  • Paeonia mascula
    Paeonia mascula
  • Paeonia mascula subsp. russoi
    Paeonia mascula subsp. russoi
  • Paeonia peregrina
    Paeonia peregrina
  • Paeonia obovata subsp. japonica
    Paeonia obovata subsp. japonica
  • Paeonia officinalis
    Paeonia officinalis
  • Paeonia ostii
    Paeonia ostii
  • Paeonia rockii
    Paeonia rockii
  • Paeonia tenuifolia
    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 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,

Tien Shan
Mountains of Kazakhstan. The rest of the section concentrates around the Mediterranean, and in Asia.

The species around the Mediterranean include

Sicilly, P. corsica on Corsica, Sardinia, the Ionian islands and in western Greece, P. clusii subsp. clusii on Crete and Karpathos, and subsp. rhodia on Rhodes, P. kesrouanensis in the Western Taurus Mountains, P. arietina from the Middle Taurus Mountains, P. broteri in Andalucia, P. humilis from Andalucia to the Provence, P. officinalis from the South of France, through Switzerland to the Middle of Italy, P. banatica in western Romania, northern Serbia and Slovenia and in southern Hungary, P. peregrina in Albania, western Bulgaria, northern Greece, western Romania, Serbia, Montenegro and Bosnia, while P. mascula has a large distribution from Catalonia
and southern France to Israel and Turkey.

Between the two concentrations, the subspecies of Paeonia daurica occur, with subspecies velebitensis in Croatia, and daurica in the Balkans and Crimea, while the other subspecies coriifolia, macrophylla, mlokosewitschii, tomentosa and wittmanniana are known from the Caucasus, Kaçkar and Alborz Mountains.

Paeonia emodi occurs in the western Himalayas between Pakistan and western Nepal, P. sterniana is an endemic of southeastern Tibet, P. veitchii grows in Central China (Qinghai, Ningxia, Gansu, Shaanxi, Shanxi, 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).[18]

Distributional history

The species of the section Paeonia have a

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.[18]

Distribution maps of the species of Paeonia in Europe and Western Asia.

Cultivation

Paeonia 'Sarah Bernhardt'

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, when they were grown in the imperial gardens. In the tenth century the cultivation of peonies spread through China, and the seat of the Song dynasty, Luoyang, was the centre for its cultivation, a position it still holds today.

A second centre for peony cultivation developed during the

Shōwa periods). During the 1940s Toichi Itoh succeeded in crossing tree peonies and herbaceous peonies and so created a new class of so-called intersectional hybrids. Although P. officinalis and its cultivars were grown in Europe from the fifteenth century on, originally also for medicinal purposes, intensive breeding started only in the nineteenth century when P. lactiflora was introduced from its native China to Europe. The tree peony was introduced in Europe and planted in Kew Gardens in 1789. The main centre of peony breeding in Europe has been in the United Kingdom, and particularly France. Here, breeders like Victor Lemoine 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.[9] 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.[21]

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.[22]

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.[23] 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.[24]

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.[9]
  • 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".[25]

Flower types

Time lapse of a peony flower blooming.

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.[9]
  • Paeonia ×arendsii 'Claire de Lune', single flowered
    Paeonia ×arendsii 'Claire de Lune',
    single flowered
  • Paeonia 'Walter Mains', Japanese flowered
    Paeonia 'Walter Mains',
    Japanese flowered
  • Paeonia lactiflora 'Bowl Of Beauty', anemone flowered
    Paeonia lactiflora 'Bowl Of Beauty',
    anemone flowered
  • Paeonia lactiflora 'James Kelway', semi-double flowered
    Paeonia lactiflora 'James Kelway',
    semi-double flowered
  • Paeonia 'Ruth Clay', bomb flowered
    Paeonia 'Ruth Clay',
    bomb flowered
  • Paeonia lactiflora 'Da Fu Gui', double flowered
    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.[26][27]

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.[28]

Cultivars

The numerous peony hybrids and cultivars have gained the Royal Horticultural Society's Award of Garden Merit,[29] including:

  • 'Bartzella', a double yellow-flowered Itoh (intersectional) peony[30]
  • 'Coral Charm', a semi-double salmon-pink-flowered herbaceous peony[31]
  • Paeonia × festiva 'Rubra Plena', a bomb red-flowered herbaceous peony[32]
  • Paeonia × lemoinei 'High Noon', a semi-double yellow-flowered tree peony[33]

The American Peony Society is the International Cultivar Registration Authority for the genus, and accepts over 7,000 registered cultivars.[34]

Uses

The herb known as Paeonia, in particular the root of P. lactiflora (Bai Shao, Radix Paeoniae Lactiflorae), has been used frequently in

Paeonia suffruticosa is called the "King of Flowers" and Paeonia lactiflora is called the "Prime Minister of Flowers."[35]

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.[36]

Culture

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 centre 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

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.[37]

In 1903, 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

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.[38]

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.[39] 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.[40]

The red flowers of the species Paeonia peregrina are important in Serbian folklore. Known as Kosovo peonies (Serbian: косовски божур, kosovski božur), they are said to represent the blood of Serbian warriors who died in the Battle of Kosovo.[41]

In 1957, 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.[42]

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.[22]

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 consequence.[citation needed
]

Famous painters of peonies have included

Giuseppe Castiglione (1688-1766), who worked at the court of the Qianlong Emperor in the Qing dynasty, also painted peonies.[39]

References

  1. hdl:10654/18083
    . Retrieved 6 July 2013.
  2. ^ The Concise Oxford English Dictionary (12th ed., 2011) lists paeony as a variant spelling of peony.
  3. ^ "paeony". Dictionary.com Unabridged (Online). n.d.
  4. ^ "Paeonia". Merriam-Webster.com Dictionary.
  5. ^ a b c Halda, Josef J.; Waddick, James W. (2004). The genus Paeonia. Oregon, USA: Timber Press.
  6. ^
    Springer-Verlag
    . pp. 265–269.
  7. doi:10.11646/phytotaxa.261.3.1
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  8. ^ Deyuan, Hong; Kaiyu, Pan; Turland, Nicholas J. (2001). Flora of China (PDF). Vol. 6. pp. 127–132. Retrieved 10 May 2016.
  9. ^ a b c d e Kamenetsky, Rina; Dole, John (2012). "Herbaceous Peony (Paeonia): Genetics, Physiology and Cut Flower Production" (PDF). Floriculture and Ornamental Biotechnology. 6 (Special Issue 1): 62–77. Retrieved 29 April 2016.
  10. ^ a b c "Saxifragales". ANGIOSPERM PHYLOGENY WEBSITE. Retrieved 26 April 2016.
  11. ^ a b L. Watson; M.J. Dallwitz. "Paeoniaceae Rudolphi". The families of flowering plants. Retrieved 26 April 2016.
  12. S2CID 205544231
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  13. ^ 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).
  14. ^ David J. Mabberley. 2008. Mabberley's Plant-Book.Cambridge University Press: UK.
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  19. ^ Sang, Tao (1996). Phylogeny and Biogeography of Paeonia (PAEONIACEAE), dissertation. The Ohio State University. cited on "Tao Sang, DISSERTATION, chapter 4". Paeon. Retrieved 19 May 2016.
  20. PMID 9335140
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  21. ^ "'The industry's about to explode': Peony market flourishes in Alaska". Anchorage Daily News. 5 August 2018. Retrieved 10 January 2019.
  22. ^ a b The Language of Flowers, edited by Sheila Pickles, 1990
  23. ^ "HPS Frequently Asked Questions: Ants on Peonies". Heartland Peony Society. Retrieved 2 May 2010.
  24. ^ "Ants on Peony Flowers: An Example of Biological Mutualism". Integrated Pest Management - University of Missouri.
  25. ISBN 978-9020965070
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  26. ^ Coit, J. Eliot (1908). "Peony". Bulletin. 259: 104.
  27. ^ "Peony". Royal Horticultural Society.
  28. ^ "Paeonia suffruticosa".
  29. ^ The Award of Garden Merit lists: Ornamentals list. The Royal Horticultural Society. Updated April 2023. Retrieved 15 June 2023.
  30. ^ "Paeonia 'Bartzella'". RHS. Retrieved 15 June 2023.
  31. ^ "Paeonia 'Coral Charm'". RHS. Retrieved 15 June 2023.
  32. ^ "Paeonia × festiva 'Rubra Plena'". RHS. Retrieved 15 June 2023.
  33. ^ "Paeonia × lemoinei 'High Noon'". RHS. Retrieved 15 June 2023.
  34. ^ "Peony Registry". The American Peony Society. Retrieved 12 October 2023.
  35. ^ Sasaki, Sanmi. 2005. Chado: The Way of Tea: A Japanese Tea Master's Almanac. Translated from the Japanese by Shaun McCabe and Iwasaki Satoko. Boston: Tuttle. Page 247.
  36. ^ "Getting to know peonies' symbolism - part 2". Bloomthis.
  37. ^ Terese Tse Bartholomew, Hidden Meanings in Chinese Art (San Francisco: Asian Art Museum/Chong-Moon Lee Center for Asian Art and Culture), 2006.
  38. ^ Whysall, Steve (14 May 2015). "Into China's peony paradise". Vancouver Sun. Archived from the original on 27 September 2021. Retrieved 16 March 2023.
  39. ^ a b Walter Good. "The World of the Peony". Archived from the original on 13 March 2016.
  40. ^ Sowerby, James & John Edward. 1899. Ranunculaceae to Cruciferae. London: George Bell & Sons. Page 69.
  41. ^ Natalia. "A Pilgrimage to Kosovo Today". Serbian Orthodox Church. Retrieved 21 April 2018.
  42. ^ "Peony". State Symbols USA. 25 May 2014. Retrieved 21 April 2018.

Bibliography

External links

Wikispecies has information related to Paeonia.
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