Paeonia daurica
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
ovate—
orbicular, 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.
The Peony Society also has a page about this species here.If you need more info about paeonia species, click here.
This species overview and synonyms on Kew Plants of the World Online:
hereSource: Hong, De-Yuan. “Peonies of the World. Taxonomy and phytogeography.” Kew: Royal Botanic Gardens, 2010, p. 168-171.