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Tree peony

The tree peony (Paeonia × suffruticosa), a cultivated hybrid deciduous woody shrub in the family Paeoniaceae, native to the mountains of northwestern China and grown to heights of 1.8–2.4 meters with an erect, open habit supported by thick, brittle stems that persist through winter.^[1] Its alternate, bipinnately compound leaves are light green above and bluish below, consisting of ovate leaflets 5–10 cm long with 3–5 lobes or teeth, while its flowers—large, fragrant, and up to 30 cm across—bloom in mid- to late spring in shades from white and pink to deep red and purple, often featuring a basal blotch on the petals and numerous violet-red stamens.^[1]^[2] Unlike herbaceous peonies, tree peonies maintain their woody framework year-round, producing 75–100 blossoms per mature plant under optimal conditions, though their stems may require staking due to fragility.^[2] Originating as a hybrid primarily from P. jishanensis and P. rockii, the tree peony has been cultivated in China for over 1,600 years, initially documented as a medicinal plant around 200 BCE and later prized for its ornamental beauty, with historical records noting its abundance in regions like Luoyang by the 11th century.^[2]^[3] Introduced to Europe in the 18th century and Japan centuries earlier, it became a symbol of prosperity in Chinese culture, leading to the development of thousands of cultivars selected for flower form, color, and size; today, over 2,000 named varieties exist worldwide, with at least half originating in China.^[3] Cultivation favors full sun to partial shade, well-drained, fertile, slightly alkaline soils (pH >8.0) rich in organic matter, and USDA hardiness zones 3a–8a, though it demands consistent winter chill for flowering and is intolerant of drought or transplanting once established.^[1]^[2] Beyond its aesthetic appeal, the tree peony holds significant economic and cultural value: its seeds yield an edible oil high in unsaturated fatty acids, beneficial for health, while its roots and flowers have been used in traditional Chinese medicine for anti-inflammatory and analgesic properties since ancient times.^[3] Deer- and rabbit-resistant with attractive qualities for butterflies, it thrives as a long-lived landscape specimen but faces challenges from environmental stress, such as desiccation in its cultivated central Chinese habitats, prompting research into drought-tolerant ecotypes like 'Luo Yang Hong'.^[2]^[3]

Description

Habit and structure

Tree peonies (Paeonia × suffruticosa) are deciduous woody shrubs characterized by an erect, open habit, typically reaching heights of 1 to 2 meters with a spread of 1 to 1.5 meters, forming a multi-stemmed framework supported by persistent branches that do not die back in winter.^[4]^[1]^[5] These shrubs develop thick, woody stems with coarse branching, contributing to their long-lived structure that can persist for decades once established.^[6] The overall architecture is robust yet slow-growing, often taking several years to reach mature size, with stems that may require occasional pruning to maintain shape and remove leggy growth.^[5] The leaves are alternate, compound, and bipinnately divided into 9 to 13 ovate leaflets, each 5 to 10 cm long with 3 to 5 lobes or teeth, presenting a light green color above and a glaucous, bluish-green hue underneath that adds subtle ornamental interest throughout the growing season.^[1] These large, dissected leaves emerge in spring and provide dense foliage cover during summer, enhancing the shrub's textural appeal.^[4] The root system consists of thick, fleshy, and often tuberous roots that extend deeply and horizontally, adapted for nutrient storage and longevity, making the plants difficult to transplant once established and best suited to undisturbed sites.^[6]^[5] This subterranean structure supports the shrub's perennial nature, storing reserves to fuel annual growth without reliance on frequent division. The growth cycle begins with bud break in early spring, followed by rapid foliage development through summer, culminating in autumn leaf drop while the woody framework remains intact through winter dormancy.^[4]^[1] This deciduous pattern allows for a distinct seasonal rhythm, with new shoots emerging from persistent buds on the branches each year, contributing to the plant's enduring presence in the landscape.^[5]

Flowers and reproduction

Tree peonies produce large, showy flowers measuring up to 25 cm in diameter, featuring 5-10 broad, silky petals in shades of white, pink, red, yellow, or purple, surrounding numerous stamens and multiple carpels.^[7]^[2]^[5] In double-flowered cultivars, many stamens are transformed into petal-like structures, often rendering them infertile and reducing seed production potential.^[8] Flowers typically bloom in late spring to early summer, from April to May, often before the leaves fully expand, with terminal buds forming on the previous year's woody stems.^[5]^[9] Pollination occurs primarily through insects such as bees and beetles, which are attracted to the pollen-rich centers.^[10] Most tree peony species and cultivars exhibit self-incompatibility, necessitating cross-pollination for successful fertilization and fruit set, as self-pollination results in significantly reduced seed yield.^[11]^[12] Following pollination, fertilized carpels develop into follicles that mature in late summer, each containing several large, black seeds encased in a fleshy aril.^[6] These seeds are primarily dispersed by gravity, though ant-mediated dispersal via the aril's elaiosome may occur in natural settings. Germination is slow and requires double dormancy release through sequential stratification: initial warm conditions (15-25°C) for 8-12 weeks to promote embryo growth and radicle emergence, followed by cold stratification (0-10°C) for 10-16 weeks to break epicotyl dormancy.^[13] This process can take 1-2 years, with gibberellic acid occasionally used to enhance rates in cultivation.^[13]

Taxonomy

Classification history

Tree peonies, known in Chinese as mudan, were first documented in ancient texts during the Han Dynasty (206 BCE–220 CE), particularly in the Shennong Bencao Jing (Divine Farmer's Materia Medica), where the root bark was praised for its medicinal properties, including cooling the blood, relieving pain, and treating gynecological disorders.^[14] This early recognition highlighted their value beyond ornamentation, establishing a foundation for their cultural and botanical significance in China long before Western scientific classification.^[15] Western botanical interest emerged in the 18th century, with Carl Linnaeus establishing the genus Paeonia in Species Plantarum (1753) based on European herbaceous species like P. officinalis.^[16] The tree peony itself was formally described in 1805 by Henry C. Andrews as Paeonia suffruticosa, drawing from cultivated double-flowered specimens imported from China, which were already hybridized forms rather than wild plants.^[17] In the 19th century, botanists advanced the taxonomy by distinguishing woody forms; Augustin Pyramus de Candolle established section Moutan in 1824 to encompass the shrubby, East Asian species, separating them from the herbaceous section Paeonia based on habit and morphology.^[18] Joseph Decaisne further refined classifications in the 1830s–1840s through descriptions of variations, such as Paeonia moutan forms, contributing to early understandings of diversity within the group. In modern taxonomy, tree peonies are placed in the family Paeoniaceae, genus Paeonia L., section Moutan DC., comprising diploid shrubs endemic to China.^[6] Debates persist on species delimitation due to extensive natural and cultivated hybridization, with P. × suffruticosa representing a complex of cultivars derived from wild progenitors. Key revisions in the late 20th century, led by Hong De-yuan, addressed this complexity; his 1990s fieldwork and analyses recognized eight species (three with subspecies), including P. suffruticosa subspp., P. ostii, P. rockii subspp., P. jishanensis, P. qiui, P. decomposita subspp., P. delavayi, and P. ludlowii, emphasizing morphological and geographical distinctions amid cultivar proliferation.^[19] Hong's subsequent updates, including a 2021 classification, recognized nine wild species and subspecies across subsections Delavayanae (P. delavayi, P. ludlowii) and Vaginatae (P. cathayana, P. decomposita and subsp., P. jishanensis, P. ostii, P. qiui, P. rockii and subsp.) while underscoring hybridization's role in blurring boundaries.^[20]^[21]

Recognized species

The recognized species within Paeonia section Moutan, collectively known as tree peonies, are primarily distributed in China and include several distinct wild taxa distinguished by their floral morphology, foliage, and geographic ranges across subsections Delavayanae and Vaginatae. These species form the genetic foundation for numerous cultivated varieties, with P. × suffruticosa serving as the collective name for the hybrid cultivar group rather than a wild species.^[6]^[22] Paeonia ostii, native to central China in provinces such as Anhui and western Henan, is a deciduous shrub growing to 1-2 meters tall, characterized by its vigorous growth and early blooming period in spring. Its solitary flowers are typically white to pale pink, up to 12 cm across, with prominent yellow anthers and a basal crimson blotch on the petals, blooming earlier than many other section Moutan species. This species is rare in the wild due to habitat loss but is widely cultivated and serves as a primary seed source for hybrid tree peony production, contributing to its economic importance.^[23]^[24]^[22] Paeonia rockii, widespread in northwestern China including Shaanxi, Gansu, and Sichuan at 1,000–2,600 m, is a shrub to 2 m with glaucous leaves and large white to pale yellow flowers (12–20 cm) featuring dark basal blotches and yellow anthers, blooming in late spring; it is a key progenitor of many cultivars and shows high genetic diversity.^[25]^[26] Paeonia cathayana, found in central China (southern Shaanxi, Hubei) at 800–1,800 m in forests and thickets, grows to 1.5 m with divided leaves and white flowers (10–15 cm) with purple basal markings, flowering mid-spring; it serves as the primary maternal parent in P. × suffruticosa genetics.^[27]^[22] Paeonia decomposita is endemic to northwestern China, specifically the Dadu River valley in Gansu and Sichuan provinces, at elevations of 1500-2500 meters, where it grows in rocky slopes and scrub. This endangered species reaches 1-1.5 meters in height and is notable for its deeply divided leaves with 19-33 narrow, lanceolate leaflets that are glaucous blue-green. Its pendant flowers are single, rose-purple to pink, about 10 cm wide, with yellow anthers, blooming in early summer; populations are threatened by overcollection for medicinal use and habitat degradation.^[28]^[29] Paeonia jishanensis and Paeonia qiui represent rare endemics with highly restricted distributions in central China. Paeonia jishanensis occurs in southwestern Shanxi, northern Henan, and central Shaanxi, in secondary forests and thickets at 900-1700 meters, forming shrubs to 1.5 meters with white to pale pink flowers, 10-12 cm across, that have entire or slightly lobed leaflets and bloom in mid-spring. Paeonia qiui is confined to western Henan and Hubei, on grassy slopes and limestone cliffs at 1000-2200 meters, distinguished by its purplish-red upper leaf surfaces, entire leaflets, and pale pink single flowers with a basal red blotch, measuring 8-12 cm, flowering in late spring. Both species have limited populations, making them vulnerable to extinction, and feature unique petal markings and foliage traits that aid in their taxonomic identification.^[30]^[31]^[32]^[33] In subsection Delavayanae, Paeonia delavayi is distributed in southwestern China (Yunnan, Sichuan) at 2,500–3,500 m in open woodlands, growing to 1–1.5 m with yellow to red flowers (8–12 cm) and deeply lobed leaves, blooming early summer; it is distinct by its often multi-carpellate fruits. Paeonia ludlowii, rarer in southeastern Tibet and Bhutan at 3,000–3,700 m, reaches 2–3.5 m with single bright red flowers (10 cm) and glaucous foliage.^[34]^[35]^[36] Many cultivated tree peonies, often collectively referred to under Paeonia × suffruticosa, originate from interspecific hybridization among wild species within section Moutan, particularly involving P. cathayana, P. rockii, P. ostii, P. jishanensis, and P. qiui in subsection Vaginatae, resulting in diverse floral forms and colors.^[22]^[6]

Distribution and habitat

Native range

Tree peonies, classified within Paeonia section Moutan, are endemic to China, with wild populations distributed across temperate regions primarily in the central and western parts of the country. Their geographic range extends from the northwestern provinces of Gansu and Shaanxi westward through Sichuan and Yunnan, reaching eastward to Anhui, Hubei, Henan, and Shanxi. This distribution centers around mountainous terrains, where the species have evolved in isolation, contributing to their genetic diversity.^[37]^[16] These shrubs typically inhabit elevations between 1,000 and 2,500 meters, though some species extend slightly beyond this range, such as P. rockii up to 2,800 meters in the Qinling Mountains—a key area of concentration for multiple taxa. Specific locales like the Qinling range in Shaanxi and Gansu host dense populations of species including P. ostii and P. rockii, while southwestern sites in Sichuan and Yunnan support P. delavayi. Such highland settings provide the cool, seasonal conditions essential for their growth.^[38] In natural environments, tree peonies occupy mixed deciduous forests, open scrublands, and rocky slopes, often in association with broad-leaved trees like oaks and pines. For instance, P. delavayi thrives in sparse thickets and dry woodlands, while P. ostii favors forested slopes and thickets. Wild distributions remain confined to China, with no confirmed native disjunct populations elsewhere, though historical human introductions have established non-native stands in regions like Japan and Europe.^[38]^[39] Habitat fragmentation and loss due to deforestation, urbanization, and overharvesting for medicinal use pose severe threats, resulting in classifications of Vulnerable or Endangered under Chinese national red lists for most species; for example, P. rockii is assessed as Endangered.^[40] These pressures have reduced wild populations significantly, underscoring the need for targeted conservation in their native Chinese ranges.^[41]

Ecological adaptations

Tree peonies (Paeonia sect. Moutan) exhibit notable drought tolerance, primarily through their extensive root systems and woody structure, which enable survival in semi-arid conditions common to their native Chinese habitats. The plants develop deep, tuberous roots that access groundwater and store reserves, allowing them to endure periods of water scarcity once established.^[42] This adaptation is complemented by their preference for well-drained, loamy soils that prevent waterlogging while supporting root penetration.^[43] Additionally, the thick, woody bark on mature stems contributes to reduced transpiration and water loss, aiding resilience during dry spells.^[1] In terms of temperature extremes, tree peonies demonstrate cold hardiness across USDA zones 4 to 8, where winter chilling induces necessary bud dormancy for proper spring growth and flowering.^[42] This dormancy period, typically requiring 6 weeks of temperatures below 40°F (4°C), protects developing buds from premature emergence and subsequent damage by late spring frosts, though severe freezes can still affect exposed shoots in marginal sites.^[44] Tree peonies form symbiotic associations with arbuscular mycorrhizal fungi (AMF), such as Glomus species, which colonize their roots and enhance nutrient uptake, particularly phosphorus and nitrogen, in the nutrient-poor, rocky mountain soils of their native range.^[45] These fungi extend the root system's effective reach into low-fertility substrates, improving overall plant vigor and stress tolerance in oligotrophic environments.^[46] AMF diversity in tree peony rhizospheres varies by cultivar and location, underscoring the symbiosis's role in adapting to heterogeneous soil conditions across cultivation areas in China.^[47] For defense against herbivores, tree peonies produce bioactive compounds, including monoterpenoid glycosides like paeoniflorin in leaves and seeds, which deter feeding by insects and mammals through toxicity and bitterness.^[48] These secondary metabolites contribute to chemical barriers that reduce herbivory pressure in natural settings.^[49] Concurrently, the large, nectar-rich flowers play a key role in local pollinator networks, attracting bees and syrphid flies that facilitate cross-pollination and enhance seed set, thereby supporting biodiversity in meadow and woodland edges.^[50] Bee-mediated pollination, in particular, boosts fruit development and yield, integrating tree peonies into broader ecological interactions.^[51]

Cultivation

Historical development

The cultivation of tree peonies (Paeonia section Moutan) originated in Chinese gardens during the Tang Dynasty (618–907 CE), where they were prized for their ornamental beauty and integrated into elite landscapes.^[52] By the early 10th century, during the Later Tang dynasty, imperial cultivation had become prominent in Luoyang, with Emperor Zhuangzong commissioning the planting of over a thousand specimens in the Linfang Hall, as recorded in historical annals.^[53] This period marked the transition from wild collection to deliberate gardening, establishing tree peonies as symbols of prosperity and refinement in imperial courts. Advancements in selective breeding during the Song Dynasty (960–1279 CE) further refined tree peony traits, emphasizing larger flowers, diverse colors, and symmetrical forms through careful propagation techniques.^[54] Chinese horticultural texts from this era documented nearly 203 cultivars, highlighting the era's systematic approach to varietal improvement and solidifying the plant's status as the "king of flowers."^[54] Tree peonies reached Europe in the late 18th century via maritime trade from China, with the first living specimens arriving in England around 1788 and blooming at Kew Gardens, igniting widespread horticultural interest.^[55] American cultivation commenced in the 19th century, as plants were imported by botanists and nurserymen, expanding their presence in North American ornamental gardens.^[55] The 20th century brought innovative hybridizations, including the Rockii group, resulting from crosses such as P. rockii × P. suffruticosa, based on seeds collected by explorer Joseph F. Rock in western China during the 1920s.^[56] These efforts introduced novel traits like dark basal flares and enhanced fragrance. Ongoing breeding programs in China and Japan persist, focusing on genetic diversity and adaptability through modern selection and hybridization. Recent advances as of 2025 include molecular breeding with genetic markers for improved traits and in vitro grafting techniques to accelerate propagation of elite cultivars.^[57]^[58]^[59]

Growing conditions

Tree peonies thrive in full sun to partial shade, requiring at least six hours of direct sunlight daily for optimal flowering, though light afternoon shade can protect buds from scorching in hot climates.^[5] They prefer deep, fertile, well-drained soils that are neutral to slightly alkaline, with a pH range of 6.5 to 7.5, to support healthy root development and prevent issues like root rot.^[60] Amending planting sites with compost improves soil structure and nutrient content, ensuring excellent drainage is essential as these plants are intolerant of waterlogged conditions.^[43]^[5] During the establishment phase, provide moderate watering to keep soil consistently moist but not soggy, typically about one inch per week, transitioning to drought tolerance once mature after two to three years.^[5] Mulching around the base with organic material, such as bark or compost, helps retain moisture, suppress weeds, and regulate soil temperature.^[43] Fertilize in early spring with a low-nitrogen, balanced formula (such as 5-10-10) applied around the drip line to promote blooming without excessive foliage growth.^[60] Prune only to remove dead or damaged wood immediately after flowering, and lightly shape leggy plants in early fall to maintain an open structure and encourage bushiness.^[5] Common pests include aphids, which cluster on new growth and can transmit viruses, managed through insecticidal soap or natural predators like ladybugs.^[61] Diseases such as botrytis blight, causing gray mold on stems and buds, are prevalent in humid conditions; prevention involves ensuring good air circulation by spacing plants adequately and avoiding overhead watering.^[62] Tree peonies are generally resilient but benefit from monitoring for scale insects and cankers, treated by pruning affected parts and applying horticultural oil.^[5]

Propagation methods

Tree peonies (Paeonia sect. Moutan) can be propagated through both sexual and asexual methods, with the latter being more common for preserving specific cultivars due to the plant's slow growth and complex seed dormancy. Sexual propagation via seeds produces variable offspring, while asexual techniques like division, grafting, and micropropagation ensure clonal reproduction.^[63] Seed propagation begins with harvesting mature seeds in late summer or early fall, typically from late August to early September, when seed pods turn dark brown and begin to crack open. Fresh seeds are then subjected to a double stratification process to break dormancy: first, a warm moist period at 70–80°F (21–27°C) for about 3 months to initiate root development, followed by a cold period at 34–40°F (1–4°C) for 3 months to promote shoot emergence; this cycle may need repetition if germination does not occur. Germination often takes 1–2 years and has low success rates, frequently below 50%, owing to the physiological dormancy inherent in peony seeds.^[64]^[65] Division of roots is an asexual method suitable for own-rooted tree peonies, performed best in fall on mature plants aged 5–10 years to avoid stressing younger specimens. The process involves digging up the plant to expose the woody root system, then carefully separating natural offsets or divisions that include at least one bud (eye) and accompanying roots, using sharp tools to minimize damage to the brittle structure. Replanting immediately in well-drained soil is essential, though success depends on the cultivator's experience, as the taproot-like configuration makes clean separations challenging and recovery slow.^[66]^[67]^[63] Grafting is the predominant commercial asexual propagation technique, accelerating establishment by uniting a tree peony scion—typically a bud or short stem segment—with a robust herbaceous peony understock, such as Paeonia lactiflora roots, which serve as a nurse tissue. The procedure, often done in late winter or early spring, involves matching cambium layers via whip-and-tongue or cleft grafts, followed by healing under high humidity and warmth (around 75–80°F); the herbaceous root eventually dies as the tree peony develops its own system. This method is favored in nurseries for its reliability and speed, yielding flowering plants in 2–3 years compared to 5+ years from seeds.^[68]^[69]^[70] Micropropagation through tissue culture is employed for rare or elite cultivars, enabling mass production of disease-free plants via in vitro techniques. Meristem explants, such as shoot tips or axillary buds, are surface-sterilized and cultured on Murashige and Skoog (MS) medium supplemented with cytokinins like 6-benzylaminopurine (BA) for shoot multiplication and auxins such as indole-3-butyric acid (IBA) or naphthaleneacetic acid (NAA) for rooting. Challenges include phenolic browning, which is mitigated by antioxidants, and low acclimatization rates (often 50–70%), but advancements have improved efficiency for commercial applications.^[71]^[72]

Uses and cultural significance

Ornamental value

Tree peonies (Paeonia suffruticosa) are highly prized in ornamental horticulture for their spectacular blooms, which feature large, showy flowers in a wide array of colors including whites, pinks, yellows, and deep reds. Each individual flower typically lasts 1-2 weeks on the plant, providing extended visual interest during the late spring blooming period, while established specimens can maintain reliable flowering performance for 10-15 years or more without significant decline due to their woody, perennial nature.^[73]^[42] These long-lived shrubs offer not only floral beauty but also attractive deep green foliage throughout the growing season, turning to vibrant shades of red, orange, and gold in the fall for additional ornamental appeal.^[74] Over 1,000 cultivars of tree peonies have been developed, primarily through centuries of selective breeding in China and Japan, showcasing diverse flower forms as classified by the American Peony Society. These include singles with a central cluster of stamens surrounded by one row of petals; doubles featuring multiple layers of petals without prominent stamens; anemone-flowered types with elongated petaloids in the center resembling sea anemones; and Japanese forms characterized by ruffled, collar-like petaloids around the stamens.^[75] Among the most celebrated varieties is 'Shimadaijin', a Japanese cultivar noted for its large, semi-double, deep red flowers that exude a rich fragrance and hold up well in garden settings. In landscape design, tree peonies serve as striking specimen plants in open areas where their architectural form and bold blooms can be showcased individually, often requiring at least 5 feet of space around the base to accommodate their mature size. They integrate well into mixed borders along pathways or fences, providing mid-level structure amid taller shrubs and lower perennials, and thrive at woodland edges where dappled shade mimics their natural habitat. For enhanced year-round interest, they pair effectively with shade-tolerant perennials such as hostas, whose broad leaves complement the peony's textured foliage without competing for resources, creating layered plantings that extend visual appeal beyond the bloom season.^[76]^[77]^[78]

Medicinal applications

The root bark of the tree peony, known as Mu Dan Pi or Moutan Cortex in Traditional Chinese Medicine (TCM), has been utilized for centuries to cool the blood, alleviate fever, and address menstrual disorders such as irregular menstruation and dysmenorrhea.^[15] This herb is particularly valued for clearing heat from the blood level, reducing symptoms like nosebleeds, subcutaneous bleeding, and irritability associated with febrile conditions.^[15] Key bioactive compounds in the root bark include paeonol, a phenolic compound with anti-inflammatory properties, and paeonoside, a glycoside contributing to its therapeutic effects.^[79]^[80] Pharmacological studies have validated the anti-inflammatory and analgesic effects of Mu Dan Pi extracts, demonstrating inhibition of pro-inflammatory cytokines such as TNF-α and IL-6 via pathways like NF-κB and MAPK.^[81] These properties make it effective for managing pain and swelling in conditions like rheumatoid arthritis and neuropathic pain.^[15] Additionally, extracts have shown benefits for skin conditions, including chronic dermatitis and photoaging, by reducing inflammation and oxidative stress, with applications extending to acne management through topical formulations that soothe irritated skin.^[15]^[82] In clinical practice, Mu Dan Pi is commonly prepared as decoctions for internal use to treat blood heat, tinctures for enhanced absorption in circulatory issues, or modern capsules for standardized dosing, typically ranging from 6–12 grams daily in decoction form.^[15] However, it is contraindicated for pregnant individuals due to its emmenagogue properties, which may promote uterine contractions and menstrual flow, potentially leading to miscarriage.^[83] Caution is also advised for those with yin deficiency or excessive sweating.^[83] Modern research highlights cardiovascular benefits of tree peony extracts, particularly in regulating blood pressure and preventing atherosclerosis, attributed to flavonoids and paeonol that improve vascular function, reduce oxidative stress, and inhibit platelet aggregation.^[84] Studies in animal models have shown cardioprotective effects against ischemia-reperfusion injury, with flavonoid content enhancing antioxidant capacity and endothelial health.^[85] These findings support its traditional use for blood stasis while underscoring potential in managing hypertensive and thrombotic conditions.^[84]

Symbolism in culture

In Chinese culture, the tree peony (Paeonia suffruticosa), known as mǔdān or the "King of Flowers" (huá wáng), symbolizes wealth, prosperity, honor, and high social status.^[86] This reverence dates back centuries, with the flower frequently featured in imperial gardens, such as those in the Forbidden City during the Qing Dynasty, where it was cultivated as the state flower to represent imperial grandeur and nobility.^[87] Tree peonies also appear prominently in classical Chinese art, including paintings from the Ming Dynasty (1368–1644), where their lush blooms embodied feminine beauty, love, and opulence, as seen in works depicting multicolored peony shrubs in full bloom.^[88] In Japan, where tree peonies (botan) were introduced from China in the 8th century CE by Buddhist monks, the flower holds associations with good fortune, a noble spirit, and prosperity, often integrated into temple and court gardens.^[89] These gardens, including tea gardens employing shakkei (borrowed scenery) techniques to harmonize plantings with surrounding landscapes, use tree peonies to evoke elegance and seasonal beauty, enhancing the contemplative atmosphere.^[90] The flower's symbolism of wealth and honor is celebrated in events like the annual peony blooms at Otokuni-dera Temple in Kyoto, a site known as the "Peony Temple," where around 2,000 plants from 30 varieties draw visitors to honor its cultural legacy during late April festivals.^[91]^[92] In Western traditions, tree peonies symbolize romance, prosperity, and extravagance, reflecting their lush, showy blooms that convey opulence and affectionate bonds.^[93] During the Victorian era, in the language of flowers (floriography), peonies represented bashfulness and happy marriages, often gifted to express romantic sentiments or celebrate enduring love.^[94] In contemporary Western culture, this symbolism persists in modern tattoos, where tree peony designs depict prosperity, good fortune, and passionate romance, serving as personal emblems of beauty and abundance.^[95] The tree peony's cultural prominence is highlighted in festivals like China's Luoyang Peony Festival, held annually since 1983 and recognized as a national intangible cultural heritage since 2008, which celebrates the flower's role in heritage through exhibitions, performances, and gardens showcasing over 1,000 varieties.^[96] These events underscore the plant's enduring status as a bridge between nature and societal values across East Asia.

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[21]
Multiple species of wild tree peonies gave rise to the 'king of flowers ...
Cultivated tree peonies, collectively named Paeonia suffruticosa Andrews, were recorded in cultivation in China 1400 years ago [7]. Because of their beauty and ...
[22]
Paeonia ostii - Trees and Shrubs Online
Distribution China Wild but rare or extinct in Anhui and W Henan; widely cultivated. Habitat Deciduous forests and thickets, 300–1,600 m. USDA Hardiness ...
[23]
https://www.treesandshrubsonline.org/articles/paeonia/paeonia-ostii/
[24]
Paeonia decomposita - Trees and Shrubs Online
It is a narrow endemic, existing as a few populations threatened by habitat degradation, inbreeding and digging for medicine or local gardens (Hong 2010; Hong ...
[25]
Inbreeding and inbreeding depression of Paeonia decomposita ...
Dec 5, 2019 · Paeonia decomposita belongs to a xenogamous category and is partially self-compatible; moreover, it requires pollinators for seed production.
[26]
Paeonia jishanensis - Trees and Shrubs Online
2001; Hong 2010). Distribution China N Henan, C Shaanxi, SW Shanxi. Habitat Secondary deciduous forests and thickets; 900–1,700 m. USDA Hardiness Zone 5-9.
[27]
Paeonia jishanensis T.Hong & W.Z.Zhao | Plants of the World Online
The native range of this species is China (C. Shaanxi, SW. Shanxi, N. Henan). It is a shrub and grows primarily in the temperate biome.
[28]
Paeonia qiui - Trees and Shrubs Online
Distribution China W Henan, W Hubei; very rare but sometimes cultivated.. Habitat Deciduous forests, grassy slopes, limestone cliffs; 1000–2200 m. USDA ...
[29]
Paeonia qiui Y.L.Pei & D.Y.Hong | Plants of the World Online
The native range of this species is China (W. Hubei, W. Henan). It is a shrub and grows primarily in the temperate biome. Taxonomy · General information.
[30]
Taxonomy, Geographic Distribution and Ecological Habitats of Tree ...
Aug 7, 2025 · Wild tree peonies are distributed in Henan, Gansu, Shaanxi, Shanxi, Anhui, Hubei, Sichuan, Yunnan and Xizang (Tibet) of China. The most densely ...
[31]
What paeonia species is this? Key to the species of peony.
Paeonia ludlowii is a tall shrub that often forms large and dense clumps with dozens of stems. A single individual can have up to 105 ﬂowers (Hong, 1997). All ...Missing: classification | Show results with:classification
[32]
https://www.treesandshrubsonline.org/articles/paeonia/paeonia-qiui/
[33]
(PDF) The phylogeographic structure and conservation genetics of ...
Aug 6, 2025 · The endangered species Paeonia rockii is the most important ancestral species of the cultivated tree peonies. These well-known ornamental ...
[34]
Characterization of the DNA molecular sequence of complete plastid ...
Aug 13, 2021 · This species has been listed as endangered species in the IUCN Red List, and urgent management and conservation are required. In this study ...
[35]
Tree Peony: How to Plant and Grow and Care for Woody Peonies — Peony's Envy
### Summary of Tree Peony Care Details
[36]
How to grow tree peonies - RHS
Tree peonies are very hardy, but the foliage and flower buds can be damaged by late frosts so avoid frost pockets; Choose a sheltered position to prevent ...<|separator|>
[37]
https://www.researchgate.net/publication/225541919_Taxonomy_Geographic_Distribution_and_Ecological_Habitats_of_Tree_Peonies
[38]
Arbuscular mycorrhizal fungi associated with tree peony in 3 ...
We examined the mycorrhizal status of the rhizosphere of 14 common cultivars of tree peony (Paeonia suffruticosa) in 3 different geographic locations in China.
[39]
Efficient protocols for the micropropagation of tree peony (Paeonia ...
Arbuscular mycorrhizal fungi (AMF) are ubiquitous soil microorganisms that form biotic associations with the roots of most vascular plant species (Harley, 1989 ...
[40]
AM fungi diversity in the main tree-peony cultivation areas in China
Jul 20, 2007 · Abstract: In order to characterize arbuscular mycorrhizal (AM) fungal resources in the rhizosphere of tree-peony (Paeonia suffruticosa), we ...
[41]
Comprehensive Metabolite Profile Uncovers the Bioactive ... - NIH
Jun 25, 2023 · Our findings indicate that newly grown tree peony leaves may serve as alternative sources of antioxidant and antibacterial substances, which ...
[42]
Unveiling spatial metabolome of Paeonia suffruticosa and Paeonia ...
Apr 17, 2021 · In Paeonia suffruticosa (PS), the texture of wood core and other parts are quite different, and using the dry-coating method (sublimation) may ...
[43]
Peonies and Pollinators: A Dance in the Garden - Gardenia.net
Peonies are renowned for their enticing fragrance and striking blooms, which draw a variety of pollinators, predominantly ants and bees.Missing: networks | Show results with:networks
[44]
Bee-mediated pollination enhances fruit set and seed yield in ...
Nov 1, 2025 · Bee pollination delivers ample high-quality pollen to the stigmas of oil tree peony, enhancing seed production. Moreover, a trend high ...
[45]
A review of the ethnobotany, phytochemistry and pharmacology of ...
Tree peony also formed a unique culture in the long history of China. The peony culture is manifested through poetry and songs (Ying, 2011), paintings, flower ...
[46]
A brief history of peony cultivation
In the Tang (923-926AD) the Emperor Zhuangzong constructed Linfang Hall in Luoyang and "cultivated over a thousand peonies at the front" (Qingyi Records by Tao ...
[47]
Multiple species of wild tree peonies gave rise to the 'king of flowers ...
Dec 22, 2014 · Because of their beauty and fragrance, cultivated tree peonies were crowned the 'king of flowers' in the Chinese Tang Dynasty, symbolizing ...
[48]
Peony Timeline | Matthaei Botanical Gardens & Nichols Arboretum
The first living “tree peony” is introduced to Europe (into England) from China. Peony mania ensues after it bloooms in Kew Gardens (then a private royal garden) ...
[49]
May 2014 Plant Profile: Paeonia suffruticosa (Rockii Group)
May 6, 2014 · Tree peonies are long-lived shrubs with exquisite flowers, but they take careful placement and a lot of patience until they're well established.
[50]
Germplasm resources and genetic breeding of Paeonia - Nature
Jul 1, 2020 · In China, most of the new Paeonia cultivars were selected using this method before 1960. The largest selective breeding program for tree peony ...
[51]
[PDF] Growing & Planting Woody (Tree) Peonies
Plant woody peonies in well-drained, loose soil with at least half day sun. Plant upright, on an angle, or on their side, burying much of the plant.
[52]
https://www.sciencedirect.com/science/article/pii/S0254629918320702
[53]
[PDF] Control of Ornamental Diseases - VCE Publications - Virginia Tech
Peony (Paeonia) – Botrytis blight, Cercospora leaf spot,. Cladosporium ... Tulip Tree – aphids, borers, scale insects, weevils. Tree-of-heaven - spotted ...<|control11|><|separator|>
[54]
Woody Peony Propagation
The primary method of clonal (copying) propagation for 'woodies' is through the procedure of grafting, that is the joining of woody peony stem to herbaceous ...
[55]
Starting Peonies from Seed - Cricket Hill Garden
A good seed producing tree peony can yield over 50 seeds per pod. The herbaceous seeds are ready a bit later, around the end of August. An almost ripe ...Missing: follicles dispersal<|separator|>
[56]
None
### Summary of Indoor Method for Germinating Peony Seeds
[57]
Dividing a Tree Peony - Cricket Hill Garden
Apr 16, 2020 · Only tree peonies on their own roots can be divided. Remove leaves, dig around the drip line, and look for natural divisions with buds and ...
[58]
https://www.sciencedirect.com/science/article/abs/pii/S0254629925003655
[59]
[PDF] Grafting Woody Peonies
In the case of woody peony grafts, a scion from a woody peony is harvested and joined to a herbaceous peony's root, which serves as a nurse root.
[60]
Grafting Tree Peonies - Cricket Hill Garden
Apr 16, 2020 · Grafting tree peonies involves fusing a bud onto a root, using scions and herbaceous roots, matching cambium layers, and healing at high heat.
[61]
Grafting Tree Peonies
In tree peonies, it involves implanting a scion of a desirable tree peony variety onto a piece of root from an herbaceous peony.
[62]
Micropropagation of tree peony (Paeonia sect. Moutan): A review
Dec 6, 2019 · Browning is a common problem in the tissue culture of woody plants such as tree peony and is usually attributed to phenolic compounds being ...
[63]
Peony tissue culture
Nov 2, 2018 · Peony tissue culture, initially difficult, now enables large-scale, fast, and clean propagation, providing disease-free material, though ...
[64]
How to Grow and Care for Tree Peony - The Spruce
Sep 4, 2025 · Tree peonies (Paeonia suffruticosa) are deciduous shrubs that showcase large, fragrant flowers for about a week to 10 days in mid to late ...Missing: habit cycle
[65]
Paeonia suffruiticosa - Shrub / Tree Peony (Assorted Colors)
In northern areas (USDA zones 3-6), Peony foliage lasts all summer and turns brilliant red and orange colors in the fall. Due to its need for cold winters, ...
[66]
Awards - American Peony Society
The “Gold Medal” is awarded annually for peonies exhibiting outstanding floral qualities and in recent years has also been designated “Peony of the Year.” These ...Missing: Shimadaijin | Show results with:Shimadaijin
[67]
How to Use Peonies to Create a Beautiful Landscape - The Spruce
May 18, 2025 · A tree peony needs ample space, at least 5 feet from the base all around. Itoh peonies are also a good choice as a specimen in mixed beds.How To Add Peonies To Your... · On Borders And Hedges · Caring For Peonies
[68]
Landscape Design with Peonies - Cricket Hill Garden
Apr 17, 2020 · Apply the same principles of space and sunlight, color and form, and size of plants when planning a peony border against fences and stone walls.
[69]
https://www.treepeony.com/blogs/peonies/grafting-tree-peonies
[70]
Paeonol: pharmacological effects and mechanisms of action - PubMed
Apr 25, 2019 · Paeonol injection has been successfully applied in China for nearly 50 years for inflammation/pain-related indications.
[71]
Biological Mechanisms of Paeonoside in the Differentiation of Pre ...
Jun 27, 2021 · Paeonia suffruticosa has been used as a medicinal plant for thousands of year and has beneficial effects in the treatment of various ...
[72]
Paeonia × suffruticosa (Moutan Peony)—A Review of the Chemical ...
5.3. Anti-Inflammatory Effect. Studies conducted both in vitro and in vivo on the anti-inflammatory activity of Moutan cortex indicated that two compounds— ...
[73]
Paeonia suffruticosa Andrews root extract ameliorates photoaging ...
Mar 5, 2025 · Numerous studies have shown that it possesses anti-inflammatory, antioxidant, and anti-aging effects due to its rich content of bioactive ...Missing: analgesic | Show results with:analgesic
[74]
Mu Dan Pi (Cortex of the Peony Tree Root)
Contraindicated in pregnancy, Yin deficiency with excessive sweating, and diarrhea. Some traditional sources say this herb is not to be used with Bulbus ...
[75]
Paeonol for the Treatment of Atherosclerotic Cardiovascular Disease
Paeonia suffruticosa, a natural herb, has been used for the treatment of atherosclerotic cardiovascular disease for thousands of years in Eastern countries.
[76]
Biological Activities of Paeonol in Cardiovascular Diseases: A Review
They related the improvement in vascular function to the fact that paeonol reduced blood viscosity and oxidative stress and improved antioxidant capacity. 3.4.
[77]
China's King of the Flowers Explained: History, Symbolism, and ...
Sep 27, 2025 · The peony's story stretches back more than a thousand years. Records show it was grown in China as early as the Zhou dynasty (around 1,000 BCE), ...Missing: origins | Show results with:origins<|control11|><|separator|>
[78]
Its Peony Time in China! - Cricket Hill Garden
Tradition tells us that peonies first arrived in Luoyang in the Tang Dynasty during the reign of Empress Wu (690-705 CE).Missing: origins | Show results with:origins
[79]
Peonies | Cleveland Museum of Art
The shrubs of this tree peony are in full bloom, blossoming in various colors. Peony paintings were popular from the Southern Song period to the Ming dynasty.
[80]
Peonies in Japan - Cricket Hill Garden
Apr 17, 2012 · There is no species of tree peony native to Japan. Historians date the arrival of the tree peonies (known as Botan in Japanese) in Japan to the 8th century CE.
[81]
[PDF] Plant of the Season: Botan by Katie Croft - Japanese Garden Society
Botan (Paeonia x suffruticosa) is a tree peony, a hybrid from China, with Chinese forms being larger and bushier, and Japanese forms smaller with sparser ...Missing: Exhibition | Show results with:Exhibition
[82]
Otokuni-dera Temple | Sightseeing Spots - Another Kyoto
It is also famous as "Peony Temple". Every year around late April, about 2000 peony flowers from 30 different species are in full bloom and you can enjoy the ...
[83]
The Peony: Flower of Wealth and Honor | Kyoto National Museum
Feb 26, 2020 · The peony, originally a Chinese flower and popularly called the "King of Flowers," has traditionally been regarded as a symbol of wealth and ...
[84]
Flower Meanings: The Language of Flowers | Almanac.com
Discover the language of flowers with the Almanac's complete chart of Flower Meanings. Which flowers represent love, strength and innocence?Victorian-era etiquette · Tansy Flower Guide · Baby's Breath · MilkweedMissing: Western | Show results with:Western<|separator|>
[85]
Flower Delivery Grand Rapids (MI) Same-Day Florist - Eastern Floral
Peony Symbolism. General symbolism of the peony includes good fortune, wealth ... In the Victorian Age, peonies were synonymous with beauty and bashfulness.
[86]
Peony Tattoo Meaning - The Total Tattoo
Mar 25, 2023 · Peonies have been a long-standing symbol of wealth, good fortune and prosperity. With deep roots in Eastern and Western cultures alike.
[87]
Luoyang Peony Festival - China Discovery
Luoyang Peony Festival has been held for about 40 years and has become a namecard of Luoyang and an important gala for both Luoyang locals and tourists.