Herbs perennial up to 80 cm tall. Roots always carrot—shaped. Stems usually green, sometimes purple, always glabrous, up to 1 cm in diameter. Lower leaves biternate with some leaflets segmented; leaflets/leaf segments usually 11-18, rarely less or more, up to 21 in number, ovate, obovate, wide-elliptic or oblong, cuneate or rounded-cuneate at the base, acute at the apex, 4.5-18 cm long, 3-9 cm wide, glabrous above, mostly glabrous, less frequently sparsely, rarely rather densely hispid beneath. Flowers solitary and terminal; involucrate bracts 1-2 in number, leaf-like; sepals 3-4 in number, all rounded or only one acute at the apex, purple or purple at the periphery, glabrous, 1.5-3 cm long, 1-2.5 cm wide; petals 7-9 in number, pink, red, white or white with pink shade at the base or periphery, obovate, rounded at the apex, 5-7 cm long, 2.5-4 cm wide; ﬁlaments pink to purple; anthers yellow; disk waved, 1 mm high, tomentose; carpels mostly 3-4, rarely 1-2, occasionally more in number, always lanate, hairs 3 mm long; styles absent or 1 mm long; stigmas 2-2.5 mm wide, red. Follicles ovoid—columnar.
Chromosome number: 2n = 20 (see each subspecies for detail).
Relatively widely distributed, from N Spain (Cantabria and Soria) to Iraq via France, Italy, the Balkans, Cyprus and Turkey.
The circumscription of Paeonia mascula has changed greatly. Cosson (1887) adopted a very broad concept of P. corallina (=P. mascula), including P. broteri, P. cambessedesii, P. coriacea, P. russoi and P. coriacea var. atlantica (= P. algeriensis Chabert). Gürke (1903) used the name P. mascula (L.) Desf. (Tabl. école bot., 126 (1804)), but his concept is similar to Cosson’s (1887). However, Lynch’s (1890) P. corallina excluded P. broteri, P. coriacea, P. triternata (= P. daurica) and P. russoi as separate species. Huth (1891) held a broad concept following Cosson (1887), and thus broad concepts were prevalent until Stern’s (1946) monograph. Fiori (1898) developed an even broader concept for this group, treating P. corallina as a subspecies of P. officinalis, and P. mascula as a variety of this subspecies, i.e. P. officinalis subsp. corallina var. mascula, but his treatment has never been accepted. Stern (1943, 1946) gave P. mascula a narrow sense, recognising the Arietina group (including P. arietina and P. bakeri), the Broteri group, the Coriacea group (P. coriacea and P. coriacea var. atlantica) and the Russoi group (P. russoi and P. cambessedesii) as groups allied to the Mascula group (P. mascula, P. daurica, P. banatica and P. kesrouanensis).
Cullen and Heywood (1964) revised the taxonomy of the genus Paeonia in Europe, treating P. russoi Biv. as a subspecies in P. mascula, and merging P. arietina G. Anderson for the first time into P. mascula as a subspecies, P. mascula subsp. arietina (G. Anderson) Cullen & Heywood. Thus, the species then had three subspecies. This circumscription was accepted by Davis and Cullen (1965) in ‘Flora of Turkey’. Bolòs and Vigo (1974) adopted Cosson’s (1887) viewpoint, treating P. cambessedesii as a subspecies within P. mascula but no one has subsequently followed this treatment. Tzanoudakis (1977, 1983) adopted Cullen and Heywood’s (1964) concept and described two new subspecies within P. mascula, i.e. subsp. hellenica and subsp. icarica (Tzanoudakis, 1977). Greuter and Burdet (1982) treated the plant from Algeria as P. mascula subsp. atlantica (Coss.) Greuter and Burdet (= P. algeriensis Chabert, = P. corallina var. atlantica Coss. = P. coriacea var. atlantica (Coss.) Stern). Stearn and Davis (1984) basically adopted Cullen and Heywood’s (1964) concept, but treated P. daurica Andrews (1807) as a subspecies within P. mascula, P. mascula subsp. triternata (Pall. ex DC.) Stearn & P. H. Davis. Muñoz-Garmendia and Navarro (1993) recorded P. mascula subsp. mascula in the Iberian Peninsula for the first time. Akeroyd (1993) basically followed Cullen and Heywood (1964), Tzanoudakis (1977), and Stearn and Davis (1984), in recognising five subspecies: subsp. mascula, subsp. arietina, subsp. hellenica, subsp. russoi and subsp. triternata. Paeonia mascula subsp. bodurii N. Özhatay was described as new from NW Turkey (Canakkale Province) by N. & E. Özhatay (1995). Schmitt (1997) recognised five subspecies in P. mascula: subsp. mascula, subsp. atlantica, subsp. coriacea (Boiss.) Malagarriga (1975), subsp. hellenica and subsp. russoi, failing to mention subsp. arietina, subsp. bodurii and subsp. triternata. Halda (1997, 2004) gave no reasons when he merged P. kesrouanensis Thiébaut into P. mascula as a subspecies.
It is clear from the above historic review of taxonomic treatments of P. mascula and its allies that there were 10 valid subspecific names before Hong et al. (2005, 2007), subsp. mascula, subsp. arietina, subsp. atlantica, subsp. bodurii, subsp. coriacea, subsp. hellenica, subsp. kesrouanensis, subsp. orientalis, subsp. russoi and subsp. triternata.
Our observations of sampled populations and herbarium specimens, and subsequent analysis, show that P. daurica Andrews (= P. mascula subsp. triternata) is an independent species (Hong et al., 2007). It has mostly nine leaflets or segments and is a diploid, with the exception of three subalpine or alpine subspecies in the Caucasus (Hong & Zhou, 2003). Paeonia algeriensis Chabert (1889) is a distinct species with mostly single and nearly always glabrous carpels, large follicles and always villose lower leaf surfaces (see that species for a detailed analysis). Paeonia arietina has its closest relatives in the P. officinalis group rather than the P. mascula group because it has tuberous roots and densely hirsute stems, leaves and sepals (Hong et al., 2008). Hong and Wang (2006) demonstrated that the peonies in Corsica, in Sardinia and on the Ionian Islands and the adjacent mainland (Akarnania Province of Greece), which have been identified as Paeonia subsp. russoi, do not belong to this subspecies but form an independent species, P. corsica Sieber ex Tausch (1828). According to their concept, subsp. russoi is confined to Sicily and Calabria in Italy. Paeonia coriacea Boiss. is distinct and can be readily distinguished from P. mascula as it is glabrous throughout, its styles 1.5—3 mm long, and its carpels mostly 2 in number. Paeonia kesrouanensis (Thiébaut) Thiébaut differs distinctly from P. mascula in always having glabrous carpels, in having leaves that are rather densely villose beneath, and in having styles and stigmas that are 5-7 mm long in total, and recurved at the apex. Paeonia kesrouanensis and P. mascula meet in S Turkey (Hatay Province), but we found no hybrids there during our field observations in 2002, indicating that the two species are indeed distinct (Hong et al., 2005).
Paeonia mascula here circumscripted is a variable species. According to our observations and statistical analysis, four subspecies could be recognised in its entire range.
1a. Flowers red or pink (entire range, except Sicily, Calabria, Qanakkale Province of Turkey and some Aegean islands): subsp. mascula
1b. Flowers mostly white, rarely red or pink.
2a. Leaves nearly always hispid, very rarely glabrous beneath (Sicily and Calabria of Italy): subsp. russoi
2b. Leaves mostly glabrous, rarely very sparsely hispid beneath.
3a. Leaflets/leaf segments of lower leaves 9—11 in number, 13-17.5 x 7-9 cm (Qanakkale in NW Turkey): subsp. bodurii
3b. Leaflets/leaf segments of lower leaves (9—)11—14(—21) in number, 9—13 x 4.5—7.5 cm (S Greece and Aegean islands): subsp. hellenica
Paeonia mascula subsp hellenica
Chromosome number: 2n = 20 (tetraploid)
Found mostly in forests dominated by Abies and Arbutus, or in sparse Crataegus thickets with scattered trees of Castanea sativa, Platanus orientalis, etc.; on limestones at an altitude of 500—2,000 m. Conﬁned to S Greece and the Aegean islands.
Flowers of this subspecies are mostly white, rarely red or pink. According to Dr Tzanoudakis at the University of Patras, Greece (on the evidence of his slides of a population from Euboea), 90% of individuals have white petals. It is closely related to subsp. russoi, from which it differs only in having leaves mostly glabrous, rarely sparsely hispid on the lower surface.Footnotes:
- Hong, De-Yuan. “Peonies of the World. Taxonomy and phytogeography.” Kew: Royal Botanic Gardens, 2010, pp. 181-187.