Paeonia mascula(1)

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 ovoidcolumnar.

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 mascula

Chromose number: 2n=20

Growing in deciduous broad-leaved forests dominated by Quercus, Fagus, Platanus or Castanea, or in conifer forests of Pinus, Abies or Cedrus, or in thickets at the edges of forests. It is mostly found on limestone areas at altitudes from 300 to 2,200 m. Most widely distributed, covering nearly the entire range of the species, from N Spain to Iraq (absent from Sicily and Calabria of Italy, and some islands in Greece). There are some specimens collected from the island of Steep Holm, England, but according to Stern (1946) and Clapham and his co- workers (Clapham er al., 1987), the species was introduced and has been naturalised there.

Copyright Liberto Dario. "Paeonia mascula subsp. mascula on Lesvos island (Greece). Very abundant in this open chestnut forest location."

Copyright Liberto Dario. “Paeonia mascula subsp. mascula on Lesvos island (Greece). Very abundant in this open chestnut forest location.”

 

Copyright Liberto Dario. “Paeonia mascula subsp. mascula on Lesvos island (Greece).”

 

Paeonia mascula subsp russoi

Chromosome number: 2n=20 (tetraploid).

Growing in sparse forests dominated by Fagus sylvestris, Quercus sp. and Castanea sativa, or in thickets of Crataegus sp.; on limestones at altitudes from 800 to 1,650 m. Confined to Sicily and Calabria, Italy.

Paeonia mascula subsp. russoi is characterised by petals that are mostly white, or white with pink shade at the base or periphery, less frequently pink or red, and leaves that are mostly rather densely hispid, rarely glabrous beneath. The subspecies has also been reported to occur in Corsica (France), Sardinia (Italy), and the Ionian islands and Akarnania Province (Greece). However, the peonies in these regions have leaflets/leaf segments 9, rarely more in number, are nearly always villose beneath, have carpels covered with short hairs (1.5 mm long) and have chromosomes 2n=10. They actually form a separate species, Paeonia corsica Sieber ex Tausch (1828) (Hong & Wang, 2006). Thus, P. mascula subsp. russoi is confined to Sicily and Calabria.

Images below:
Paeonia mascula ssp russoi in Italy, Northern Sicily, Parco delle Madonie.  Copyright Stefan Hertel
“Paeonia mascula ssp russoi flowers earlier than Paeonia [mascula ssp] flavescens. It was mainly withered at a higher altitude.”

Paeonia mascula ssp bodurii

Chromosome number: 2n=20

Growing in sparse forests of Quercus infestoria, Pinus nigra, Tilia sp. and Castanea sativa on stony slopes. Not far away from the population we sampled was a pure and well-developed Pinus nigra forest. It is apparent that the habitat of Paeonia mascula subsp. bodurii is secondary. It was found on granites at an altitude of 400-760 m. According to N. & E. Özhatay and our observations, the subspecies is confined to a small area near Camyayla in Cannakale Province, NW Turkey.

This subspecies is characterised by white petals, and by fewer but larger leaflets/leaf segments (9-11 in number; terminal one 13-18 cm long, 7-9 cm wide) than in subsp. mascula.

 

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.

Paeonia mascula ssp hellenica on Evia island (Greece) - Copyright Liberto Dario

Copyright Liberto Dario. “Paeonia mascula subsp. hellenica on Evia island (Greece). White is the dominant color in this taxon. Apart from the mainland, this endemic subspecies is found on the islands of Evia and Andros.”

 

Paeonia mascula ssp hellenica on Evia island (Greece) - Copyright Liberto Dario

Copyright Liberto Dario. “Pink form of Paeonia mascula subsp. hellenica on Evia island (Greece).”

Footnotes:

  1. Hong, De-Yuan. “Peonies of the World. Taxonomy and phytogeography.” Kew: Royal Botanic Gardens, 2010, pp. 181-187.[]

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