Paeonia daurica1

Perennials. Roots carrot—shaped, attenuate downwards, up to 4.6 cm in diameter. Lower leaves biternate with 9 leaflets, occasionally 1-2 leaflets segmented, and thus leaflets/leaf segments usually 9, rarely 10 or 11, in number, entire, broad—obovate, oblong, rarely wide—elliptic, sometimes undulate, truncate or rounded, rarely acute or even short—acuminate at the apex, 8—17 cm long, 4.8—11.5 cm wide, glabrous above, glabrous or sparsely to densely villose, or sparsely to rather densely puberulous beneath. Flowers solitary and terminal; involucrate bracts 0—2 in number, leaf-like; sepals mostly 3, less frequently 2 in number, green but with purple periphery or entirely purple, orbicular or ovateorbicular, up to 3.5 cm long, all rounded at the apex; petals 5—8 in number, usually red or rose, less frequently yellow, pale yellow, white, or yellow but with a red spot at the base or with red periphery; filaments purple; disk c. 1 mm high, waved, tomentose above; carpels 1-5 but mostly 2 or 3 in number, mostly tomentose, less frequently glabrous, hairs 2.5-3 mm long; stigmas nearly sessile, red, 1.5—2 mm wide.

Chromosome number: 2n = 10 and 20 (see each subspecies for detail).

Three lowland subspecies, subsp. daurica, subsp. coriifolia and subsp. mlokosewitschii are diploid, whereas three alpine or subalpine subspecies, subsp. macrophylla, subsp. tomentosa and subsp. wittmanniana are tetraploids. No chromosome information is yet available for subsp. velebitensis.

Growing in deciduous broad—leaved or mixed forests, or at the edges of forests, on various media and at altitudes from 350 to 2,740 m. Widely distributed from Croatia in the west to N Iran in the east via Turkey and the Caucasus. Within the range of this species, the populations in the Caucasus (including the Transcaucasus), the Talish Mountains and the Elburz Mountains are diverse and polytypical, both in morphology and chromosome number. They have been treated as five allopatric subspecies with subsp. coriifolia (diploid) in the lowlands of Georgia, subsp. mlokosewitschii (diploid) in E Georgia, NW Azerbaijan and Dagestan in Russia, subsp. wittmanniana (tetraploid) in the highlands of Abchasia (Georgia) and adjacent regions, subsp. macrophylla (tetraploid) in the western Transcaucasus (Armenia, Georgia and NE Turkey), and subsp. tomentosa (tetraploid) in the Talish Mountains (Azerbaijan and Iran) and the Elburz Mountains (Iran) (Hong & Zhou, 2003)

In Paeonia daurica, the number of leaflets/ leaf segments of lower leaves is mostly 9, rarely 10, very occasionally 11, whereas in P. mascula this ranges from 11 to 22, rarely 10, very occasionally 9. This character is rather stable within populations. ln the population D. Y. Hong et al. H02215 (Mt Amanos, Hatay, Turkey), for example, we observed 37 individuals that all had 9 leaflets/leaf segments. On this mountain, there were hundreds of P. daurica individuals at altitudes from 1,300 to 1,550 m, but very few of them had more than 9 leaflets/leaf segments. The length of the terminal leaflets varied from 5.5 to 15.2 cm (9.2 ± 1.9 cm) in P. daurica, compared with 7.7—16.7 cm (11.8 ± 2.5 cm) in P. mascula. The ratio of the length to width ranged from 1.01 to 1.82 (1.45 ± 0.23) in P. daurica, and from 1.33 to 2.18 (1.79 ± 0.22) in P. mascula. The apex of terminal leaflets was mostly truncate, broad- rounded or rounded in P. daurica, but mostly acute, cuspidate or rounded—cuspidate in P. mascula. The widest point of the terminal leaflets also differed between the two species. Although the widest point was above the middle of the terminal leaflets in both entities, it was much above the middle, about halfway between the top and the middle in P. daurica, but not much above the middle in P. mascula. Thus, the terminal leaflets were broad-obovate or nearly orbicular in P. daurica, but obovate, oblong or ovate in P. mascula. As indicated by Hong et al. (2006), P. daurica was clearly, though not distantly, differentiated from P. mascula in morphology. They were not intermingled with each other, even in S Turkey where the two entities were sympatric. Morphologically, they differed in number of leaflets/ leaf segments of lower leaves, and also in the shape of the terminal leaflets. The great majority of individuals of these two species could be distinguished.

Stearn and Davis (1984) stated that P. daurica (= P. mascula subsp. triternata) and P. mascula geographically overlapped in the eastern Aegean islands (Lesvos and Samos), and that they showed some morphological overlap in Anatolia. We critically examined all the specimens of this group from the regions mentioned above at the herbaria ATH, E, G, LD and UPA, but found that the specimens from Lesvos and Samos, including E. Stamatiadou 2666 (ATH) determined by P. H. Davis as P. mascula subsp. triternata, all belonged to P. mascula subsp. mascula. Thus, no subsp. triternata was present on these two islands. In Anatolia, subsp. triternata and subsp. mascula did coexist in some areas, e.g. in Hatay Province, Turkey, but they were not found to overlap morphologically. Instead, they were rather distinct. Their morphological differentiation can be easily understood when their difference in chromosome number is considered. No hybrids between them have been discovered.

Seven subspecies are recognised and keyed out below.

1a. Sepals often villose on the abaxial side; leaves rather densely villose on the lower side: subsp. velebitensis D. Y. Hong

1b. Sepals glabrous; leaves sparsely villose or puberulous, less frequently densely villose on the lower side or glabrous.

     2a. Carpels glabrous or nearly glabrous; petals yellow.

          3a. Leaflets/leaf segments densely villose and thus greyish beneath: subsp. macrophylla (Albov) D. Y. Hong

          3b. Leaflets/leaf segments usually sparsely villose beneath: subsp. wittmanniana (Hartwiss ex Lindl.) D. Y. Hong

     2b. Carpels tomentose; petals red, rose, white or yellow.

          4a. Leaflets/leaf segments puberulous or glabrous beneath, obovate, apex rounded or obtuse, often with a short mucro: subsp. mlokosewitschii (Lomakin) D. Y. Hong

          4b. Leaflets/leaf segments villose or glabrous beneath, obovate, oblong or wide-elliptic, apex rounded to short—acuminate.

               5a. Petals red or rose; leaflets/leaf segments glabrous or sparsely villose beneath.

                    6a. Leaflets/leaf segments broad-obovate, truncate to rounded at apex: subsp. daurica

                    6b. Leaflets/leaf segments obovate to oblong, rounded to acute at apex: subsp. coriifolia (Rupr.) D. Y. Hong

               5b. Petals yellow or yellowish white, but sometimes red at periphery or with a red spot at base; leaflets or segments villose beneath.

                    7a. Leaflets/leaf segments mostly densely villose and thus greyish beneath: subsp. tomentosa (Lomakin) D. Y. Hong

                    7b. Leaflets/leaf segments usually sparsely villose beneath: subsp. wittmanniana (Hartwiss ex Lindl.) D. Y. Hong


Paeonia daurica subsp velebitensis

The present author examined 11 collections with 12 sheets in the Hungarian Natural History Museum (BP), the Conservatoire et Jardin botaniques de la Ville de Geneve (G), the University of Vienna (WU) and the Botanic Garden in Munich (M). In these collections, all lower leaves are biternate, with 9 leaflets, which are obovate, rarely oblong-elliptic, rounded to acute at the apex, rather densely villose beneath. The carpels are 2 or 3 in number, tomentose, with hairs 3 mm long. These characters indicate the position of these collections within Paeonia daurica Andrews. The character states of 9 leaflets on lower leaves with obovate shape, mostly rounded apex, and rather densely villose hairs beneath make them distinctly different from P. mascula (leaflets/leaf segments of lower leaves more than 9, ovate or oblong-elliptic, acute at the apex, glabrous or sparsely hispid beneath).

The peony in Velebit of Croatia is characterised by villose calyx and rather dense villose hairs on the lower surface of leaves. It is similar to subsp. tomentosa (Lomakin) D. Y. Hong, which is found in S Azerbaijan and NE lran (the Talish Mountains and Elburz Mountains), but subsp. tomentosa has yellow petals. The sepals of subsp. velebitensis are often villose, and thus different from all other subspecies of Paeonia daurica. Therefore, the peony in the Velebit Mountains is rather distinct in morphology from the other six subspecies. Nevertheless, we place it here as a subspecies because its leaf pattern and shape are very similar to those of the typical subspecies, and because we have limited knowledge relating to this peony. The subspecies is confined to the Velebit Mountains in Croatia, and is found at an altitude of 900—1,150 m


Paeonia daurica subsp macrophylla

Chromosome number: 2n=20

Growing at altitudes from (800)—1,160—2,400 m, in deciduous forests or mixed coniferous and deciduous broad-leaved forests, and also in glades, with no specific soil preference. Confined to high mountainous areas in Armenia, SW Georgia (Adjaria, Guria, S Imereti, SW Kartli and Meskheti) and NE Turkey.

This subspecies was established as a new taxon by Albov (1895) and raised to specific rank by Lomakin (1897) on the basis of its large leaflets. The entity is characterised by larger leaflets and glabrous carpels. Glabrous carpels also occur, however, in subsp. wittmanniana and occasionally in subsp. tomentosa, and we have shown that the larger leaflets are within the continuous variation of leaflet size in Paeonia daurica. We have not found any other character that distinguishes subsp. macrophylla clearly from the other subspecies of P. daurica (Hong & Zhou, 2003).


Paeonia daurica subsp wittmanniana

Chromosome number: 2n=20

Growing in deciduous forests and alpine or subalpine meadows at altitudes of (800)—1,500—2,300 m. All the available records of herbarium specimens show that it is confined to limestone areas. Confined to NW Georgia (Abchasia, lmereti, Megrelia, Ratsha— Letskhumi and Svaneti) and the adjacent region of Russia (upper reaches of the Mzymta River).

We were unable to visit the localities of this subspecies when we were in Georgia in 1999 because of security considerations. Nevertheless, we have examined a large number of herbarium specimens and live individuals in the Tbilisi and Bakuriani Botanical Gardens in Georgia. These showed that the leaflets of this subspecies varied from glabrous to sparsely villose beneath, carpels were 1-3 in number and varied from glabrous or sparsely villose to tomentose, and petals varied from yellow to yellow with a pink spot at the base. The entity is not sufficiently distinct from its closest relatives to allow specific status.


Paeonia daurica subsp mlokosewitschii

Chromosome number: 2n= 10

Growing in deciduous forests dominated by Fagus, Quercus, Ulmus, Castanea and Acer, at an altitude of 960—1,060 m. To date, found only in E Georgia, NW Azerbaijan and Dagestan in Russia.

The type specimens of Paeonia mlokosewitschii and P. lagodechiana both came from the same population, which we visited and observed during our expedition in 1999. This population (D. Y. Hong & S. L. Zhou H99035) was extremely polymorphic in petal colour and leaflet indumentum.

Among the seven subspecies of Paeonia daurica, subsp. mlokosewitschii is relatively distinct; its leaflets are usually obovate, rounded and mucronate at the apex, and mostly sparsely or rather densely puberulous but sometimes glabrous beneath. Saunders and Stebbins (1938) made a cross between Paeonia triternata from SE Europe (= P. daurica subsp. daurica) and this entity, which showed that they were interfertile. This suggests that P. mlokosewitschii might be better treated as a subspecies within P. daurica.


Paeonia daurica subsp daurica

Chromosome number: 2n = 10 (diploid)

Growing in Woods at altitudes from 350 to 1,550 m. Distributed in Bosnia-Herzegovina, Croatia, Serbia, Macedonia, Romania, Bulgaria, Ukraine (Crimea), Greece (Drama), Turkey, Syria and Lebanon


Paeonia daurica subsp coriifolia

Chromosome number: 2n=10

Found at altitudes below 1,100 m in deciduous forests, dominated by Quercus, Fagus, Ulmus, Acer and Fraxinus, or in mixed deciduous broad—leaved and coniferous forests, dominated by Picea, Fagus and Quercus. On a wide range of media: sandstones, limestones and volcanic rocks. Confined to W and NW Caucasus.

Paeonia daurica subsp. coriifolia differs from the other subspecies in having red petals and leaflets that are glabrous or sparsely villose beneath.



Paeonia daurica subsp tomentosa

Chromosome number: 2n=20

Growing in deciduous forests with sand soils, at altitudes from 1,170 to 2,740 m. We also found this subspecies growing on a pasture with scattered trees or shrubs. Confined to the Talish Mountains of Azerbaijan and N Iran, and Elburz Mountains in N Iran.

The subspecies is characterised by leaflets nearly always densely covered with villose hairs on the lower surface, carpels mostly densely, rarely sparsely, tomentose, or occasionally glabrous, and petals usually pale yellow, rarely pale yellow but reddish at the base. It mostly resembles subsp. wittmanniana, but differs in having leaflets mostly densely villose on the lower surface and carpels more-or-less tomentose. By contrast, subsp. wittmanniana possesses leaflets that are mostly sparsely villose and carpels that are often glabrous. The two subspecies are geographically isolated by a considerable distance.

  1. Hong, De-Yuan. “Peonies of the World. Taxonomy and phytogeography.” Kew: Royal Botanic Gardens, 2010, pp. 168-171.[]

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