Paeonia: classification and overview of peony species.
The latest and most widely accepted scientific classification (2021) was made by Hong De-Yuan.(1) It revises his previous classification(2), which was based on morphology, by adding insights from genetic analysis. Although this will be the most widely accepted classification, some scientists still disagree with some of his findings.(3) The reason for this is twofold: morphological characters in paeonia are not discrete but rather continuous and paeonia shows large genetic variability.(4)
There are two other alternative classifications also shown here. The classification by Don Smith is based largely on genetics and groups the species according to their historical relationships.(5) The last classification is our own grouping of species and includes some ‘other’ species not accepted or mentioned by Hong De-Yuan. The reason why we consider them separate species are listed with the individual species descriptions themselves. Our own classification will be a ‘dynamic’ classification as revisions deemed reasonable will be made from time to time to keep it up-to-date with new insights and/or discoveries.
Remark: still under construction!
Source: Hong, De-Yuan. “Peonies of the world. Part III: Phylogeny and evolution.” Kew Publishing: Royal Botanic Gardens Kew; 2021, pp. 215-222
| The infrageneric subdivision of paeonia consists of: 2 subgenera and 7 sections with 34 species in all. | |||||||
| 1A. Shrubs. Subgenus 1. Moutan | |||||||
| 2A. Flowers 2-4 in a cyme; disk fleshy, annular, enveloping only base of carpels; carpels always glabrous. Section 1A. Delavayanae | |||||||
| 2B. Flowers solitary and terminal; disk leathery or half-leathery, enveloping only carpels by half or entirely; carpels tomentose or glabrous. Section 1B. Moutan | |||||||
| 1B. Herbs. Subgenus 2. Paeonia | |||||||
| 3A. Flowers 2-4 in a cyme or solitary but often with one or two undeveloped flower buds in axils; roots carrot-shaped. Section 2A. Albiflorae | |||||||
| 3B. Flowers solitary and terminal; roots fusiform-thickened or carrot-shaped. | |||||||
| 4A. Leaves mostly with bristles along veins above; leaflets finely segmented; leaflets plus segments (9-)21-340 in number; roots fusiform-thickened. Section 2B. Paeonia | |||||||
| 4B. Leaves always glabrous above; roots carrot-shaped or slightly fusiform-thickened. | |||||||
| 5A. Petals nearly as long as sepals (slighly longer or shorter); stems often branched; roots slighly fusiform-thickened. Section 2E. Onaepia | |||||||
| 5B. Petals much longer (>2 times) than sepals; stems simple | |||||||
| 6A. Lower leaves always biternate, with 9 leaflets; leaflets entire; carpels mostly 2 or 3, rarely 1, 4 or 5. Section 2C. Obovatae | |||||||
| 6B. Lower leaves biternate, with 9, rarely 8 or 7 leaflets; leaflets segmented or entire; leaflets plus segments 9-95, very rarely 8 or 7; carpels 2-5, rarely 6, 1, 7-10. Section 2D Corallinae | |||||||
| Subgenus 1. Moutan | |||||||
| Shrubs. Flowers in a cyme, or solitary and terminal on a branch. Disk fleshy, annular, dentate, or leathery and enveloping carpels half to entirely. The subgenus consists of nine wild species all endemic to China and one cultivated hybrid species, Paeonia x suffruticosa, which is now cultivated almost worldwide. Two sections are easily recognized as follows. | |||||||
| Section 1A. Delavayanae | |||||||
| Shrubs up to 3.5 m tall. Flowers 2-4 in a cyme. Disk fleshy, dentate, annular, enveloping only the base of the carpels; carpels 1-5 in number. | |||||||
| Included species (2): | |||||||
| Paeonia ludlowii | |||||||
| Paeonia delavayi | |||||||
| Distribution: endemic to SW China: SE Tibet, W Sichuan and Yunnan. | |||||||
| Section 1B. Moutan | |||||||
| Shrubs up to 2 m tall. Flowers solitary on top of branches, not in a cyme. Disk leathery, enveloping carpels by half or entirely until mid-anthesis. Carpels mostly 5, less often 3 or 4 in number. | |||||||
| Included species (7): | |||||||
| Paeonia decomposita | |||||||
| Paeonia rotundiloba | |||||||
| Paeonia rockii | |||||||
| Paeonia ostii | |||||||
| Paeonia jishanensis | |||||||
| Paeonia qiui | |||||||
| Paeonia cathayana | |||||||
| Distribution: endemic to C China: N & W Sichuan, SE Gansu, Shaanxi, Hubei, Henan, SW Shanxi and C Anhui | |||||||
| Subgenus 2. Paeonia | |||||||
| Section 2A. Albiflorae | |||||||
| Roots more or less carrot-shaped. Leaves glabrous or with bristles along veins on the upper surface. Flowers usually several per stem, in a cyme, rarely solitary, or solitary but often with undeveloped (sterile) flower buds at axils; sepals mostly caudate at the apex. Almost all diploid with 2n=10 (P. emodi with a tetraploid population, 2n=20, in S Xizang (Tibet). | |||||||
| Included species (5): | |||||||
| Paeonia anomala | |||||||
| Paeonia emodi | |||||||
| Paeonia lactiflora | |||||||
| Paeonia sterniana | |||||||
| Paeonia veitchii | |||||||
| Distribution: all the species in Paeonia sect. Albiflorae are found in Asia, but with the range of P. anomala extending into north-eastern Europe (Kola of Russia) | |||||||
| Section 2B. Paeonia | |||||||
| Lateral roots fusiform or tuberous. Lower leaves biternate or triternate; leaflets nearly always segmented with leaflets/leaf segments (9-)21-340 in number; leaves mostly with bristles along veins above. Flowers solitary and terminal; sepals mostly rounded at apex. Mostly tetraploid (diploid only in P. intermedia and P. tenuifolia). | |||||||
| Included species (7) | |||||||
| Paeonia arietina | |||||||
| Paeonia intermedia | |||||||
| Paeonia officinalis | |||||||
| Paeonia parnassica | |||||||
| Paeonia peregrina | |||||||
| Paeonia saueri | |||||||
| Paeonia tenuifolia | |||||||
| Distribution: from Middle Asia to the northern Mediterranean. | |||||||
| Section 2C. Obovatae | |||||||
| Roots carrot-shaped. Lower leaves biternate; leaflets 9, entire, or segmented, forming 13-24 leaflets/segments, always glabrous above. Flowers always solitary and terminal; sepals mostly rounded at the apex, but sometimes one caudate; carpels mostly 2 or 3, rarely 1, 4 or 5, glabrous or yellow papillate to hispidulous. One diploid and one tetraploid. | |||||||
| Included species (2) | |||||||
| Paeonia obovata | |||||||
| Paeonia mairei | |||||||
| Distribution: E Asia: Jpan, Korean peninsula, the Far East of Russia, and NE, E, C and SW China, with its westernmost part in the eastern edge of the Pan-Himalaya. | |||||||
| Section 2D. Corallinae | |||||||
| Roots carrot-shaped. Lower leaves biternate; leaflets/leaf segments usually numbering 9 or more but fewer than 21 (up to 32 in P. broteri and 95 in P. clusii); leaves always glabrous above. Flowers always solitary and terminal; sepals mostly rounded at the apex. Both diploid and tetraploid with 2n=10 and 2n=20. | |||||||
| Included species (9) | |||||||
| Paeonia algeriensis | |||||||
| Paeonia broteri | |||||||
| Paeonia cambessedesii | |||||||
| Paeonia clusii | |||||||
| Paeonia coriacea | |||||||
| Paeonia corsica | |||||||
| Paeonia daurica | |||||||
| Paeonia kesrouanensis | |||||||
| Paeonia mascula | |||||||
| Distribution: S Europe, NW Africa and W Asia (From the western Mediterranean to Iran). | |||||||
| Section 2E. Onaepia | |||||||
| Perennials. Lateral roots slightly fusiform. Stems often branched. Lower leaves ternate or biternate. Flowers solitary and terminal on main stem and branches; petals nearly equal in size to, or smaller than, sepals; disk dentate, almost interrupted, enveloping the base of the carpels until mid-anthesis; carpels 2-5 in number, always glabrous. Diploid with 2n=10. | |||||||
| Included species (2) | |||||||
| Paeonia brownii | |||||||
| Paeonia californica | |||||||
| Distribution: the section is endemic to western (Pacific) North America: the USA and Mexico. | |||||||
| 1a. Shrubs; disk fleshy and annular, enveloping only base of carpels or half-leathery, enveloping half to whole carpels until mid-anthesis (Subgenus Moutan) | |||||||||
| 2a. Flowers usually 2 – 4 in a cyme, more or less pendent; disk fleshy, enveloping only base of carpels; carpels always glabrous (Section Delavayanae) | |||||||||
| 3a. Carpels nearly always single, rarely 2; follicles 4.7 – 7 cm long, 2 – 3.3 cm in diameter; petals, filaments and stigmas always purely yellow Paeonia ludlowii | |||||||||
| 3b. Carpels usually 2 – 5 (- 7); follicles less than 4 cm long, 1.5 cm in diameter; petals, filaments and stigmas often not purely yellow Paeonia delavayi | |||||||||
| 2b. Flowers solitary, upright; disk half-leathery, enveloping carpels halfway or completly until mid-anthesis; carpels tomentose or glabrous (Section Moutan) | |||||||||
| 4a. Carpels glabrous, 2 – 5 in number; disk enveloping carpels halfway or up to base of styles until mid-anthesis; lower leaves decompound with (20 -) 25 – 54 (- 71) leaflets; leaflets all lobed. | |||||||||
| 5a. Carpels 5, very occasionally 4, in number; disk enveloping ovary up to halfway at anthesis; leaflets of lower leaves (35 -) 37 – 54 (- 71) in number, terminal leaflet elliptic or narrow-rhomboid, with length/width ratio (1.46 -) 1.62 – 2.18 (- 2.55) Paeonia decomposita | |||||||||
| 5b. Carpels mostly 3, less often 4, 2 or 5 in number; disk enveloping whole ovary at anthesis; leaflets of lower leaves (20 -) 25 – 37 (-49) in number, terminal leaflet orbicular or wide-rhomboid, with length/width ratio (1.02 -) 1.03 – 1.57 (- 2.20) Paeonia rotundiloba | |||||||||
| 4b. Carpels densely lanate or tomentose, 5 (- 7) in number; disk completely enveloping carpels until mid-anthesis; lower leaves biternate, biternate-pinnate or ternate-bipinnate, with leaflets usually fewer than 20 (- 33), if more, at least some of them entire | |||||||||
| 6a. Lower leaves with 9 leaflets; leaflets ovate or ovate-orbicular, only terminal leaflets usually 3-lobed, often reddish above; petals often with a reddish blotch at base Paeonia qiui | |||||||||
| 6b. Lower leaves with more than 9 leaflets, if 9, then leaflets mostly lobed; leaflets green above; petals without a blotch, or with a large, dark purple blotch at base | |||||||||
| 7a. Lower leaves with 11-33 leaflets; leaflets usually ovate to lanceolate, mostly entire, less frequently ovate-orbicular and mostly lobed | |||||||||
| 8a. Lower leaves with 11-15 leaflets; leaflets ovate tot ovate-lanceolate, mostly entire; petals white, rarely pale pink, without a blotch Paeonia ostii | |||||||||
| 8b. Lower leaves with (17 -) 19 -33 leaflets; leaflets lanceolate or ovate-lanceolate and mostly entire, or ovate to ovate-orbicular and mostly lobed; petals white, rarely red, always with a large, dark purple blotch at base Paeonia rockii | |||||||||
| 7b. Lower leaves with 9 (occasionally 11 or 15 in P. jishanensis) leaflets; leaflets ovate, ovate or nearly orbicular, mostly or all lobed | |||||||||
| 9a. Leaflets ovate, terminal leaflets 3- or 5-cleft, with additional 1 to several lobes, lateral leaflets mostly 2- or 3-lobed, less frequently entire; lobes acute at apex; leaf blade glabrous on lower surface; sepals all caudate or mucronate Paeonia cathayana | |||||||||
| 9b. Leaflets ovate-orbicular to orbicular, all 3-cleft; segments lobed, acute to rounded at apex; leaf blade villose along veins on lower surface; sepals all rounded at apex Paeonia jishanensis | |||||||||
| 1b. Herbs; disk fleshy and short, enveloping only base of carpels (Subgenus Paeonia) | |||||||||
| 10a. Petals nearly equal in size to, or smaller than, sepals; disk dentate, almost interrupted; lower leaves ternate or biternate, with 3 or 9 leaflets; lateral root slightly fusiform. (Section Onaepia) | |||||||||
| 11a. Lower leaves biternate, with 55 – 110 leaf segments and final lobes; segment or lobes acute to rounded at apex; carpels usually 5, rarely 3, 4 or 6 in number; sepals exceeding petals Paeonia brownii | |||||||||
| 11b. Lower leaves ternate, with 30 – 78 leaf segments and final lobes; segments or lobes mostly acute at apex; carpels usually 3, rarely 2 or 4 in number; sepals slightly smaller than, or subequal to, petals Paeonia californica | |||||||||
| 10b. Petals much larger than sepals; disk annular, waved or flat; lower leaves biternate or triternate, with 9 or more leaflets; lateral roots carrot-shaped or fusiform | |||||||||
| 12a. Flowers 2 – 4 in a cyme, less frequently solitary but with 1 – 2 undeveloped flower buds at axils, rarely simply solitary; sepals mostly caudate at apex; leaf blade usually with bristles along veins above (Section Albiflorae) | |||||||||
| 13a. Leaf blade cartilaginous and denticulate on margin, pubescent along veins or glabrous beneath; carpels glabrous, very rarely hairy Paeonia lactiflora | |||||||||
| 13b. Leaf blade smooth on margin, glabrous beneath; carpels glabrous or hairy | |||||||||
| 14a. Lower leaves with 70 – 140 leaflets/leaf segments; petals red, rarely pale rose, very occasionally white | |||||||||
| 15a. Plants (60 -) 66 – 103 (- 120) cm tall; flowers usually simply solitary, less frequently solitary but with 1 or 2 undeveloped flower buds, very rarely 2; carpels mostly 5, less frequently 4, rarely 3 Paeonia anomala | |||||||||
| 15b. Plants (24 -) 34 – 59 (-71) cm tall; flowers mostly 2 or 3, less frequently 4 or solitary but with 1 or 2 undeveloped flower buds, occasionally simply solitary; carpels mostly 3 or 2, rarely 4 or 1 Paeonia veitchii | |||||||||
| 14b. Lower leaves with fewer than 30 leaflets or leaf segments; petals white, rarely pale rose | |||||||||
| 16a. Carpels 1, rarely 2 in number, mostly tomentose, less frequently glabrous; flowers often several per stem Paeonia emodi | |||||||||
| 16b. Carpels mostly 2 or 3 in number, rarely 4, always glabrous; flowers nearly always solitary, rarely 2 in number, but sometimes with 1 – 2 undeveloped flower buds at axils Paeonia sterniana | |||||||||
| 12b. Flowers always solitary and terminal; sepals mostly rounded at apex; leaf blade glabrous or with bristles along veins above | |||||||||
| 17a. Tap roots not thickened, lateral roots always fusiform; lower leaves with mostly more than 20, rarely fewer, leaflets or leaf segments; leaf blade covered with bristles along veins above, or, if glabrous above, nearly always villose beneath (Section Paeonia) | |||||||||
| 18a. Stems mostly hirsute; sepals mostly hairy on abaxial side; leaflet blade always glabrous or rarely villose at base above, more or less villose beneath | |||||||||
| 19a. Lower leaves with mostly more than 20, very rarely as few as 11, leaflets or leaf segments; leaflet blade linear-elliptic or lanceolate; sepals hispidulous or glabrous on abaxial side Paeonia officinalis | |||||||||
| 19b. Lower leaves with mostly fewer than 20, occasionally up to 32,leaflets or leaf segments; leaflet blade elliptic, oblong or ovate-lanceolate; sepals densely villose on abaxial side | |||||||||
| 20a. Flowers rose to red; anthers yellow; lower leaves with 11 – 25, occasionally up to 32, leaflets or leaf segments Paeonia arietina | |||||||||
| 20b. Flowers dark purple; anthers orange; lower leaves with 9 – 15, occasionally up to 25, leaflets or leaf segments. Paeonia parnassica | |||||||||
| 18b. Stems glabrous; sepals always glabrous; leaflet blade mostly with bristles along veins above | |||||||||
| 21a. Leaflets or leaf segments always dentate-lobed, with lobes less than 1 cm long; stigmas yellow or pale pink Paeonia peregrina | |||||||||
| 21b. Leaflets or leaf segments entire or rarely deeply lobed; stigmas red | |||||||||
| 22a. Lower leaves with 19 – 45 leaflets or leaf segments; leaf blade hispidulous beneath Paeonia saueri | |||||||||
| 22b. Lower leaves with more than 70 leaflets or leaf segments; leaf blade always glabrous beneath | |||||||||
| 23a. Leaflets or leaf segments of lower leaves 130 – 340 in number, mostly filiform, 0.5 – 8 mm wide Paeonia tenuifolia | |||||||||
| 23b. Leaflets or leaf segments of lower leaves 70 – 100 in number, linear, 4 – 18 mm wide Paeonia intermedia | |||||||||
| 17b. Tap roots carrot-shaped; lower leaves with no more than 24 (only in P. broteri up to 32 and in P. clusii 23 – 95) leaflets or leaf segments; leaf blade always glabrous above | |||||||||
| 24a. Leaflets 9 or 13 – 24 (including leaf segments); carpels mostly 2 or 3, rarely 1, 4, or 5 (Section Obovatae) | |||||||||
| 25a. Leaflets 9, entire; carpels glabrous Paeonia obovata | |||||||||
| 25b. Leaflets mostly segmented, forming 13 – 24 leaflets or segments; carpels yellow papillate to hispidulous, rarely glabrous Paeonia mairei | |||||||||
| 24b. Leaflets 9, rarely 8 or 7, or leaflets/leaf segments up to 95; carpels 2 – 5, rarely 6, 1, 7 – 10 (Section Corallinae) | |||||||||
| 26a. Leaflets plus leaf segments of lower leaves 23 – 95 in number, linear to ovate Paeonia clusii | |||||||||
| 26b. Leaflets plus leaf segments of lower leaves less than 21 (up to 32 in P. broteri), broad elliptic to obovate | |||||||||
| 27a. Lower leaves with 9 or fewer leaflets; carpels always glabrous Paeonia cambessedesii | |||||||||
| 27b. Lower leaves with usually 10 – 15, less frequently 9 or more, leaflets plus leaf segments; carpels mostly tomentose | |||||||||
| 28a. Carpels 1, less frequently 2, nearly always glabrous, very occasionally sparsely hairy; follicles columnar, 4 – 5.4 cm long Paeonia algeriensis | |||||||||
| 28b. Carpels (1 -) 2 – 4, rarely more, tomentose or glabrous; follicles long-ovoid or ellipsoid, up to 4 cm long | |||||||||
| Less indentation here for readability purposes! All that follows should be a level below 28b. | 29a. Styles 1.5 – 3.5 mm long; lower leaves with 11 – 14, rarely up to 17, leaflets or leaf segments; leaf blade densely or sparsely villose beneath; carpels always glabrous Paeonia kesrouanensis | ||||||||
| 29b. Styles usually absent (only in P. corsica 1.5 – 3 mm long); lower leaves with 9 – 20 leaflets or leaf segments; leaf blade glabrous, sparsely hispid, puberulent or villose beneath; carpels tomentose or glabrous | |||||||||
| 30a. Carpels always lanate or tomentose with hairs 2 – 3 mm long; styles absent | |||||||||
| 31a. Leaflets usually entire; lower leaves with 9, rarely 10, very occasionally 11, leaflets; leaflet blade usually ovate, rounded, nearly truncate with a mucro, or rounded, less frequently acute at apex Paeonia daurica | |||||||||
| 31b. Leaflets at least some segmented; lower leaves with usually 10 or more, very occasionally 9, leaflets or leaf segments; leaflet blade usually acute at apex | |||||||||
| 32a. Lower leaves with mostly (11 -) 15 – 21, rarely up to 32, leaflets or leaf segments; leaflets 4 – 10 (- 15) cm long, 1.5 – 5 (- 6.5) cm wide, always glabrous; carpels with hairs 2 mm long Paeonia broteri | |||||||||
| 32b. Lower leaves with mostly (9 -) 11 – 15, rarely up to 21 leaflets or leaf segments; leaflets 9 – 18 cm long, 4.5 – 9 cm wide, sparsely hispid or glabrous; carpels with hairs 3 mm long Paeonia mascula | |||||||||
| 30b. Carpels glabrous or tomentose with hairs 1.5 mm long; styles present or absent | |||||||||
| 33a. Carpels tomentose, rarely glabrous; lower leaves with usually 9, rarely up to 20 leaflets/leaf segments; leaf blade holosericeous beneath; styles 1.5 – 3 mm long Paeonia corsica | |||||||||
| 33b. Carpels glabrous, very occasionally sparsely hairy; lower leaves with 10 – 15 leaflets/leaf segments; leaflet blade glabrous, very occasionally sparsely hairy (hairy leaves always accompanied by hairy carpels); styles lacking Paeonia coriacea | |||||||||
| Source: Hong, De-Yuan. “Peonies of the world. Part III: Phylogeny and evolution.” Kew Publishing: Royal Botanic Gardens Kew; 2021, pp. 219-222 | |||||||||
| Paeonia classification | Clade 1 | Clade 2 | |||||
| Non-hybrid species | tenuifolia | intermedia | veitchii | Extinct #1 | lactiflora | Extinct #2 | mairei |
| Hybrid intra-clade species | |||||||
| peregrina | 1/4 | 2/4 | 1/4 | ||||
| officinalis | 3/8 | 2/8 | 3/8 | ||||
| arietina, humilis, parnassica | 1/2 | 1/2 | |||||
| anomala | 3/4 | 1/4 | |||||
| emodi | 1/2 | 1/2 | |||||
| Hybrid inter-clade species | |||||||
| wittmanniana | 1/4 | 1/4 | 1/4 | 1/4 | |||
| obovata | 1/4 | 1/4 | 2/4 | ||||
| clusii, mascula group | 1/2 | 1/2 | |||||
| banatica | 1/4 | 1/4 | 2/4 | ||||
| sterniana | 1/4 | 1/4 | 2/4 | ||||
| cambessedesii, corsica | 1/2 | 1/2 | |||||
We are far from concluding this, so whilst it is unfinished, here are some of the (sub)species unrecognized by Hong:
Paeonia wendelboi
Paeonia archibaldii
Paeonia flavescens
Paeonia arietina ssp arasicola
Paeonia hybrida
Paeonia sandrae
Paeonia turcica
- Hong, De-Yuan. “Peonies of the world. Part III: Phylogeny and evolution.” Kew Publishing: Royal Botanic Gardens Kew; 2021, pp. 215-222[back to text]
- Hong De-Yuan. “Peonies of the World. Polymorphism and diversity.” Royal Botanic Gardens: Kew, 2011, pp. XII-XV[back to text]
- Punina, E.O.; Machs, E.M.; Krapivskaya, E.E.; Kim, E.S.; Mordak, E.V. Myakoshina Y.A. & A.V. Rodionov. “Interspecific hybridization in the genus Paeonia (Paeoniaceae): Polymorphic sites in transcribed spacers of the 45S rRNA genes as indicators of natural and artificial peony hybrids.” In: Russian Journal of Genetics, 2012, vol 48, nr 7, pp. 684-697[back to text]
- Degtjareva, G.V.; Efimov, S.V.; Terentieva, E.I.; Samigullin, T.H. & C.M. Vallejo-Roman. “The problem of species in the genus Paeonia.” In: Skortsovia, 2020, vol 6, nr 2, pp. 26-27[back to text]
- Smith, Donald R. “A new ‘unified’ classification for the genus paeonia”. In: American Peony Society Bulletin, 2021, nr 398, pp. 25-40, and nr 399, pp. 33-37[back to text]
