from: Tutin et al. : Flora Europaea 2nd Ed. 1995

62. PAEONIACEAE

EDIT. D. A. WEBB

Perennial herbs or shrubs. Leaves alternate, exstipulate. Flowers usually solitary, hermaphrodite, actinomorphic, hypogynous. Sepals 5, free. Petals 5-10(-13), free. Stamens numerous. Carpels 2-8, free, borne on a fleshy disc. Fruit a group of 2-8 follicles, each with several seeds.


1. Paeonia L.

J. Cullen & V. H. Heywood (Edition 1) Revised by J. R. Akeroyd (Edition 2)

Robust perennials, with erect, tuberous stock and fleshy roots. Leaves large, simply biternate or further divided. Flowers 6—14 cm in diameter; petals white, pink or red in the European species. Stamens numerous. Follicles spreading horizontally; seeds in 2 A number of species and hybrids are widely cultivated for ornament.

Literature: F. C. Stern, A Study of the Genus Paeonia. London 1946. W. T. Stearn & P. H. Davis, Peonies of Greece. Kifissia. 1984


1

 

 

Segments of lower leaves more than 40, each segment less than 5 mm wide

 

 

 

1. tenuifolia

1

 

 

Segments of lower leaves fewer than 40, each segment more than 5 mm wide

2

 

 

Petals white, occasionally flushed with pink

 

 

 

6. clusii

2

 

 

Petals red or deep pink

3

 

 

Most leaflets divided; lower leaves with narrowly elliptical to lanceolate segments

 

4

 

Leaves without minute bristles along the veins of upper surface

 

 

5

Segments of lower leaves fewer than 15; petals blackish-red

 

 

 

4. parnassica

 

 

5

Segments of lower leaves usually more than 15; petals pink to red

 

 

6

Leaves green or brownish and pubescent beneath; filaments red

 

 

 

5. officinalis

 

 

6

Leaves glaucous and glabrous beneath; filaments greenish-white to yellow

 

 

 

7. broteroi

 

4

 

Leaves with minute bristles along the veins of upper surface

 

 

7

Ultimate leaf-segments narrowly triangular, deltate or subulate; filaments yellow

 

 

 

2. anomala

 

 

7

Ultimate leaf-segments broadly triangular; filaments pink or red

 

 

 

3. peregrina

3

 

 

Most leaflets undivided; lower leaves with lanceolate, elliptical, ovate or orbicular segments

 

 

8

Leaflets ovate to orbicular; margins undulate

 

 

 

8. mascula

 

 

8

Leaflets lanceolate to ovate; margins not undulate

 

 

9

Follicles 5-8, c. 6 cm, glabrous, purple

 

 

 

10. cambessedesii

 

 

9

Follicles fewer than 5, less than 6 cm, glabrous or pubescent, sometimes reddish but not purple

 

 

10

Follicles (3-5-)4-5 cm, glabrous, with apex attenuate into the stigma

 

 

 

9. coriacea

 

 

10

Follicles 2-5—4 cm, usually pubescent, with obtuse or rounded apex and sessile stigma

 

 

11

Leaves glabrous beneath; leaflets 2—4 cm wide

 

 

 

7. broteroi

 

 

11

Leaflets usually pubescent beneath; leaflets 5—10 cm wide

 

 

 

8. mascula


1. P.tenuifolia L., Syst. Nat. ed. 10, 2: 1079 (1759). Stems 20-40 cm. Leaves divided into numerous linear segments, each less than 5 mm wide, glabrous above, pubescent beneath. Flowers 6-8 cm in diameter, red, appearing to rest on the leaves. Filaments yellow. Follicles 2 or 3, c. 2 cm, tomentose. 2n = 10. S.E. Europe. Bu Ju Rm Rs (C, W, K, E).


2. P. anomala L., Mantissa Alt. 247 (1771). Stems 50-100 cm. Leaf-segments numerous, the ultimate segments deltate or subulate, more than 5 mm wide, glabrous beneath, with minute bristles along the main veins on the upper surface. Flowers 7—13 cm in diameter, red. Filaments yellow. Follicles 3-5 , c. 2 cm. 2n = 10. N.E. Russia. Rs(N, C). (N. Asia.)


Most European plants of this species belong to var. intermedia (C. A. Meyer) B. Fedtsch., with villous follicles, and are regarded by some Russian authors as var. intermedia (C. A. Meyer) Krylov of P. hybrida Pallas. Var. anomala, with glabrous follicles, is restricted in Europe to parts of Ural.


3. P. peregrina Miller, Gard. Dict. ed. 8, no. 3 (1768) (P. decora G. Anderson). Stems 35-50 cm, glabrous. Lower leaves divided into 17-30 narrowly elliptical segments, the ultimate segments short, broadly triangular, spreading, giving the apex of the leaflet a serrate appearance, sparsely villous to glabrous beneath, with minute bristles along the main veins above. Flowers 7-13 cm in diameter, dark red, less widely open than in other species. Filaments pink or red. Follicles 2 or 3, 2-3-5 cm, tomentose. 2n = 10, 20. Balkan peninsula, E. Romania, Moldavia; one station in C. Italy. Al Bu Gr It Ju Rm Rs(W) Tu.


4. P. parnassica Tzanoudakis, Kytt. Meleti Gen. Paeonia Ell. 43 (1977) (P. arietina auct., non G. Anderson). Stems 30-65 cm pubescent. Lower leaves divided into 9—13 obovate to narrowly elliptical or lanceolate segments, densely pilose beneath, giving a greyish-green appearance; ultimate segments acute or acuminate entire. Flowers 8—12 cm in diameter, blackish-red. Filaments purplish. Follicles 2 or 3, tomentose. 2n = 20. Mountains of S.C. Greece. Gr.


5. P. officinalis L., Sp. Pl. 530 (1753). Lower leaves divided into 17-30 segments, pubescent beneath, glabrous above; petioles deeply channelled on the upper surface. Flowers 7-13 cm in diameter, red. Filaments red. Follicles 2 or 3, 2-3-5 cm, usually tomentose, sometimes glabrous. S. & S.C. Europe. Al Ga He Hi Hu It Ju Lu Rm [Ge].


1

 

 

At least the central leaflet divided almost to base

 

2

 

Most of the leaflets divided almost to base

 

 

 

(a) subsp. officinalis

 

2

 

Only the central leaflet divided almost to the base

 

 

 

(b) subsp. banatica

1

 

 

Leaflets divided not more than 1/3 of the distance to base

 

 

3

Follicles pubescent; stems and petioles floccose

 

 

 

(c) subsp. villosa

 

 

3

Follicles glabrous; stems and petioles pubescent

 

 

 

(d) subsp. microcarpa


(a) Subsp. officinalis: Stems and petioles floccose. Most leaflets divided almost to base. Follicles tomentose. 2n = 20. From S.E. France to Albania.


(b) Subsp. banatica (Rochel) Soó, Növényföldrajz 146 (1945) (P. banatica Rochel). Like subsp. (a) but only the central leaflet deeply divided. 2n = 20. Hungary, Jugoslavia, Romania.


(c) Subsp. villosa (Huth) Cullen & Heywood, Feddes Report. 69; 34 (1964): Stems and petioles floccose. Leaflets divided to not more than 1/3 of the distance to base. Follicles tomentose. From S. France to C. Italy.


(d) Subsp. microcarpa (Boiss. & Reuter) Nyman, Consp. 22 (1878) (P. microcarpa Boiss. & Reuter, P. humilis Retz., P. officinalis subsp. humilis (Retz.) Cullen & Heywood, P. paradoxa G. Anderson): Stems and petioles pubescent. Leaflets divided to not more than 1/3 of the distance to base. Follicles glabrous. S, W. Europe.


6. P.clusii F.C. Stern, Bot. Mag. 162 :t. 9594 (1940) (P. officinalis var. glabra (Boiss.) Hayek). Stems 20-30 cm. Lower leaves with 30 or more narrowly oblong to narrowly elliptical, acute segments, glabrous above and beneath, or slightly pubescent beneath. Flowers 7—10 cm in diameter, white, rarely flushed with pink. Filaments pink. Follicles 2—5, c. 3 cm, densely tomentose. 2n = 10, 20. Kriti and Karpathos. Cr.


Outside Europe, known only from the E. Aegean islands, where the plants have been distinguished as P. rhodia Stearn, Gard. Chron., ser. 3, 110: 159 (1941) (subsp. rhodia (Stearn) Tzan.).


7. P. broteroi Boiss. & Reuter, Diagn. Pl. Nov. Hisp. 4 (1842) (P. lusitanica auct., non Miller). Stems up to 50 cm. Lower leaves divided into (9-) 17-20 narrowly elliptical, glabrous segments. Flowers 8-13 cm in diameter, purplish-pink. Filaments greenish- white to yellow. Follicles 2-4, 3-4 cm, densely tomentose. 2n = 10. S., C. & IT. Spain; Portugal. Hs Lu.


8. P. mascula (L.) Miller, Gard. Dict. ed. 8, no. 1 (1768) (P. corallina Retz., P. caucasica (Schipcz.) Schipcz.; incl. P. banatica auct., non Rochel). Stems 20-80 cm. Leaves simply biternate, or with a few leaflets divided; segments 9—16 (—21), narrowly to broadly elliptical to ovate or orbicular, glabrous or pubescent beneath. Flowers 8—14 cm in diameter, red or rarely white. Follicles (2)3-5(6), 2—4 cm, usually villous, rounded at apex, with a sessile stigma. S. Europe, extending northwards to N.C. France and Austria. Al Bu Co Ga Gr Hs It Ju Rm Rs(K) Sa Si [Au Br].


1

 

 

Flowers white or pinkish-white

 

 

 

(d) subsp. hellenica

1

 

 

Flowers pink to red

2

 

 

Lower leaves with 9 or 10(11) broadly elliptical to orbicular leaflets

 

3

 

Leaflets broadly ovate to orbicular, with undulate margins, glabrous beneath

 

 

 

(b) subsp. triternata

 

3

 

Leaflets elliptical to broadly elliptical with plane margins, pubescent beneath

 

 

 

(c) subsp. russoi

2

 

 

Lower leaves usually with more than 12 narrowly elliptical to ovate leaflets

 

 

4

At least stem and petioles without hairs; leaflets elliptical to ovate

 

 

 

(a) subsp. mascula

 

 

4

Stem and petioles and lower surface of leaves pubescent; leaflets narrowly elliptical

 

 

 

(e) subsp. arietina


(a) Subsp. mascula: Leaves glabrous or pubescent beneath; lower leaves with 9-16(-21) elliptical to ovate leaflets. Flowers red. Filaments purple. 2n = 20. From C. France to E. Greece.


(b) Subsp. triternata (Pallas ex DC.) Stearn & P. H. Davis, Peonies of Greece 107 (1984) (P. triternata Pallas ex DC., P. daurica Andrews, P. taurica auct.): Leaves glabrous beneath; leaflets usually 9, broadly ovate to suborbicular, with undulate margins. Flowers pale purplish-red. Filaments yellow. Follicles 2 or 3. Romania; Krym.


(c) Subsp. russoi (Biv.) Cullen & Heywood, Feddes Report. 69: 35 (1964) (P. russi Biv., P. ovatifolia Rouy & Fouc., P. corallina var. pubescens Moris, P. Corsica Sieber): Leaves sparsely pubescent beneath; lower leaves with 9 or 10 broadly elliptical leaflets. Flowers purplish-pink. Filaments white or pink. 2n = 10. Mediterranean region, from C. Spain to W. Greece, mainly in the islands,


(d) Subsp. hellenica Tzanoudakis, Kytt. Meleti Gen. Paeonia Ell. 36 (1977) (P. flavescens C. Presl): Leaves glabrous or pubescent beneath; lower leaves with 9—13 broadly ovate or elliptical leaflets. Flowers white. Filaments purplish. 2n = 20. S. & S.E. Greece.


(e) Subsp. arietina (G. Anderson) Cullen & Heywood, Feddes Report. 69: 35 (1964) (P. arietina G. Anderson): Leaves pubescent beneath; lower leaves with 12—15 narrowly elliptical leaflets. Flowers red. Filaments purple. S.E. Europe.


9. P. coriacea Boiss., Elenchus 7 (1838). Like 8(a) but stems 50-100 cm; leaves coriaceous; follicles 2(3), somewhat longer (3-5-4-5 cm), glabrous, with the apex attenuate into the stigma. S. Spain; Corse, Sardegna. Co Hs Sa.


10. P. cambessedesii (Willk.) Willk. in Willk. & Lange, Prodr Fl. Hisp. 3: 976 (1880), in obs., et Willk., Ill. Fl. Hisp. 1: 104, t. 65 (1883). Stems 20-50 cm. Lower leaves biternate; leaflets lanceolate to ovate, occasionally elliptical, glabrous above and beneath flushed with purple beneath. Flowers 6-10 cm in diameter, red. Follicles 5-8, c. 6 cm, glabrous, purplish. 2n = 10. Islas Baleares. Bl.