Notes on the observations upon the structure of the Paeonia seeds, fruit and roots.

Josef J. Halda

Sedlonov 125, CZ-517 82 Sedlonov, Ceska republika

Halda, J. J.: Notes on the observations upon the structure of the Paeonia seeds, fruit and roots. Acta Mus. Richnov. 5(1):1-10 (1998). ISBN 80-86076-08-3.

Publication on with kind permission of the author. 1. Oct. 2000


Seeds, fruit and roots of Paeonia L. are revised: a new subgenus Albiflora, sections Emodi and Tenuifotiaeare stated; section Flavonia is renowed; a sample of seeds, fruit and roots of each subgenus is illustrated: Key of subgenera, sections, subsections and species is made.

1. Seeds.

27 taxa of seeds were observed under a binocular microscope. Five groups of shape are described:

1) cylindric [from large ones of P. lutea (18x10mm) to small cylindric ones of P. tenuifolia (7-8 x 2-3mm)].

2) quadrangular-cylindric [P. suffruticosa subsp. rockii (15 x 10mm)],

3) regularly ovate [P. emodi (10x9mm)],

4) irregularly ovate [P. lactiflora (8-10 x 6-7mm)] and

5) almost globular [P. mlokosewitschi or P. wittmanniana (6-7mm across)].

All observed seeds were from wild collections.

List of observed seeds:

[Note: There is a correlation between the number of seeds in the follicle and the size and shape of seeds - a great number of seeds are mostly much smaller and irregular. For example when P. suffruticosa-follicles are very full, almost all seeds are nearly regularly quadrangular, but when there are only a few seeds, they are regularly cylindric. The same P. lutea, P. delavayi, P. potanini, P. brownii, P. lactiflora, P. veitchii and P. anomala].



P. lutea


P. obovata








P. brownii


P. tenuifolia








P. lactiflora


P. corsica








P. emodi


P. anomala








P. mlokosewitschi


P. veitchii


2. Fruit.

Four main groups of follicles are described.

1) slim erect semi-open follicles ["Suffruticosa-type" - P. suffruticosa, P. decomposita, P. lactiflora].

2) thick erect semiopen follicles with thick walls [ "Delavayi-type" - P. delavayi. P. lutea, P. potanini ],

3) thick erect half-open follicles with thin walls [ "Brownii-type" – P. brownii], which are always pendent and

4) widely open follicles of most herbaceous species ["Mascula-type"].

P. brownii


P. suffruticosa








P. brownii


P. delavayi








P. lutea


P. officinalis








P. emodi




P. emodi








P. lactiflora


P. mlokosewitschi


P. veitchii





3. Roots.

Seven main root-types are described.

1) vertical primary (main) root with many or several secondary roots ["Suffruticosa-type" - all "woody" peonies].

2) more or less regularly fasciculate main roots with a "head-like" top ["Lactiflora-type" - P. lactiflora], with many secondary roots.

3) irregularly fasciculate main roots ["Brownii-type" - P. brownii] with many secondary roots.

4) tapering or carrot-like main root ["Mascula-type" - P. mascula-group].

5) fusiform or "digitate"-form - from the rootstock on a string-like attachements grow corm-like swollen roots, tapering into thin roots. ["Officinalis-type" - subsect. Paeonia].

6) fusiform with a thick irregular main root with swollen corm-like roots on string-like attachments ["Anomala-type" - subsect. Anomala and Obovata].

7) fusiform with a thin rootstockand many subglobular swollen bulblets (as ashort "coral-chain") ["Tenuitolia-type". - P. tenuifolia].

P. suffruticosa


P. mascula








P. lactiflora


P. anomala








P. officinalis


P. tenuifolia








P. brownii





After a revaluation of all those characters were done some changes; sect. Albiflora, [which does not fit in subgenus Paeonia] was changed to the subgenus -rank, because together with P. brownii they create a special group with similar seeds, follicles and roots. Seeds of P. emodi are much closer to P. lactiflora that to the others herbaceous species. Seeds and roots of P. tenuifolia are exceptional and need a special status. Yellow Caucasian P. mlokosewitschii and P. wittmanniana with fleshy lobate staminodial disc and small globose seeds are presented as section Favonia. Species of the "Mascula-group" are presented as sect. Paeonia subsect Masculae - their main roots are tapering or carrot-like and leaf-segments mostly integrous.

subgenus Albiflora (Salm-Dyck) J. J. Halda stat. nov.

[bas: Paeonia sect. Albifora Salm-Dyck, Hort. Dyck. 366(1834); Paeonia sect. Albiflora Salm-Dyck ex Uspensk. in Bjull. Mosk. O-va Ispyt. Prir. otd. biol. 92(3):83 (1987)].

Roots fasciculate, tapering, more or less vertical; leaves chartaceous, margin cartilaginous, scabrid, irregularly dentate-serrate in two rows; stem with several flowers; carpels 3-5, erect, slim; seeds pale brown, irregularly cylindric, often compressed in regular rows. Type species: P. lactiflora Pall.

sect. Emodi J. J. Halda sect. nov.

Flos solitarius vel 2-4, pendulus; petala alba; semina ovoideo-globosa, brunneo-nigra.

Flower solitary or more, pendent; petals white; seeds large, ovate-globose, dull brownish-black. Type species: P. emodi Wall. ex Royle.

sect. Tenuifoliae (Stern) J. J. Halda stat. nov.

[bas: Paeonia sect. Paeon subsect. Dissectifoliae group Tenuifolia Stern, Paeonia 110 (1946), nom. nud.; Paeonia ser. Fissae Komarov in Fl. SSSR 7:31 (1937), nom. inval.; Paeonia sect. Paeonia ser. Tenuifolia Stern ex Uspensk. in Bjull. Mosk. O-va Ispyt. Prir. Otd. Biol. 92 (3):84 (1987); Paeonia subg. Paeonia sect. Palearcticae subsect. Tenuifoliae (Stern ex Uspensk.) J. J. Halda in Acta Mus. Richnov. 4(2):27 (1997)].

Roots with short chains of coraliform globose or shortly cylindric bulblets; leaf-segments pinnatisect; flower single, erect; petals red; carpels 3 (rarely 1-2), densely coarse-tomentose; seeds shiny pale brown, narrowly cylindric. Type species: P. tenuifolia L.

sect. Flavonia Kem.-Nath. in Trudy Tbil. Bot. Inst. 21:18 (1961).

[Paeonia sect. Palearcticae Huth emend. Uspensk. subsect. Flavonia (Kem.-Nath.) Uspensk. in Bjull. Mosk. O-va Ispyt. Prir. Otd. Biol. 92(3):83 (1987)].

Flowers single, yellow or creamy; staminodial disc fleshy, lobate, at the base of carpel; seeds globose, shiny black. Type species: P. wittmanniana Hartwiss.

sect. Paeonia subsect. Masculae (F. C. Stern ex Uspensk.) J. J. Halda stat. nov

[bas: Paeonia sect. Palearcticae ser. Masculae F. C. Stern ex Uspensk. in Bjul. Mosk. O-va Ispyt. Prir. Otd. Biol. 92(3):83 (1987); Paeonia sect. Palearcticae group Mascula F. C. Stem, Paeonia 67 (1946). nom. nud.; Paeonia ser. Masculae Kem.-Nath. in Tr. Tbil. Bot. Inst. 21:28 (1961), nom. nud.; Paeonia sect. Palearcticae groups Coriaceae & Russii Stern, Paeonia 88 (1946), nom. nud.; Paeonia ser Corallinae Komarow in Fl. SSSR 7:28 (1937), nom. nud.; Paeonia sect. Corallinae Salm-Dyck, Hort. Dyck. 365.366 (1834)].

Roots tapering of carrot-like; leaf-segments more or less integrous. Type species: P. mascula Mill.

Key to the subgenera:

1a Roots succulent, not woody, stems herbaceous, deciduous

..... 2

1b Roots woody, stems woody, persistent

..... subg. Moutan

2a Leaves thin or coriaceous, margin entire, smooth

..... 3

2b Leaves chartaceous, margin scabridi cartilaginous with rows of irregular serrate

teeth, [single species: P. lactiflora Pall],

..... subg. Albiflora

3a Staminodial disc reaches more than 1/3 of gynoeceum: petals coriaceous, slightly

larger than sepals [single species: P. browni Douglas ex Hooker],

..... subg. Onaepia

3b Staminodial disc small or poorly developed; petals thin, much larger than sepals.

..... subg. Paeonia

Key to the subgenus Moutan:

1a Staminodial disc as a thin leathery sheath, covering whole carpels in full bloom: lower leaves bipinnate [sect. Moutan].

..... 2

1 b Staminodial disc as conspicuous flashy lobes around the base of carpels, reaching up to 1/3 of carpels; lower leaves biternate, deeply lobed or toothed, [sect. Delavayanae].

..... 3

2a Carpels completely enveloped by stammodial disc

..... P. suffruticosa

2b Carpels enveloped only up to 1/2- 2/3

..... P. decomposita

3a Calyx of 5 leathery greenish sepals with a conspicuous involucre of 8-12 green bracts; flowers red

..... P. delavayi

3b Calyx without an involucre; bracts and sepals 5-7, the outer 1-4 more or less foliaceous, the innermost rounded and sepaloid

..... 4

4a Segments and lobes of leaves 17-30 mm wide; flowers up 90mm across yellow or yellow with reddish blotch at base

..... P. lutea

4b Segments and lobes of leaves 5-10 mm wide, upper segments wider; flowers smaller, red, white or yellow

..... P. potanini

Key to the subgenus Paeonia:

1a Flowers yellow or creamy; Staminodial disc fleshy, lobate; seeds globose, shiny-black.

sect. Flavonia

1 b Flowers not yellow or creamy; staminodial disc imperceptible; seeds ovate or cylindric

..... 2

2a Flowers pendent, white; seeds ovate-globose, dull brownish-black [single species: P. emodi Wall. ex Royle]

...sect. Emodi

2b Flowers more or less erect; carpels 3 (2) or more; seeds cylindric or ovate

..... 3

3a Roots tapering, fusiform to subglobose; segments of lower leaves less than 40, each segment more than 5mm wide.

sect Paeonia

3b Roots with coraliform-globose or shortly cylindric bulblets; segments of lower leaves more than 40, each segment less than 5mm wide [single species: P. tenuifolia L.].

...... sect. Tenuifoliae

sect. Flavonia

1a Staminodial disc fleshy, lobate, ca 4mm; diploid.

..... P. mlokosewitschi

1b Staminodial disc tiny, ca 1mm; tetraploid.

...... P. wittmanniana

sect. Paeonia

1a Roots fusiform or subglobose

..... 2

1b Roots tapering, more or less carrot-like.[subsect. Masculae]

..... 10

2a Carpels narrowly conical, attenuated to the relatively large curved stigma; lower leaves alternate [subsect. Obovatae]

..... 9

2b Carpels not narrowed, not attenuated

..... 3

3a Leaf-segments dentate [subsect. Paeonia]

..... 4

3b Leaf-segments laciniate, leaflets lanceolate [subsect. Anomalae]

..... 8

4a Flowers white

..... 5

4b Flowers red or violet

..... 6
5a Lower leaves with 40-80 leaflets and lobes, the leaflets usually 3-5-fid to 1/2 or 2/3 into oblong-lanceolate lobes

..... P. clusii

5b Lower leaves with 9-29 leaflets and segments, mostly ovate to oblong-elliptic or lanceolate

..... P.rhodia

6a Flowers red

..... 7

6b Flowers dark violet

..... P. parnassica

7 a Flowers half-open; leaf-segments broadly triangular, spreading, the leaflet appearing serrate at the apex

..... P. peregrina

7b Flowers widely open; leaflets very narrowly elliptical to lanceolate

..... P. officinalis

8a Flower single

..... P. anomala

8b Flowers 2-6

..... P. veitchii

9a Leaflets unequal, terminal usually obovate, laterals broadly oval or oblong, all shortly acuminate at the apex, cuneate or widely cuneate at the base

..... P. obovata

9b Lateral leaflets sometimes bifurcate; leaflets elliptic or obovate-elliptic, cuneate or long-cuneate at the base, long-acuminate to caudate at the apex

..... P. mairei

10a Most leaflets divided, the lower leaves with 17-30 narrowly elliptical segments

..... P. broteroi

10b Most leaflets undivided, the lower leaves with 9-16 elliptical, ovate or orbicular segments

..... 11

11a Leaflets orbicular; margins undulate

..... P. daurica

11b Leaflets generally narrower; margins not undulate

..... 12

12a Follicles 5-8, ca 6cm, glabrous, purple

..... P. Corsica

12b Follicles less than 5, less than 6cm, glabrous or pubescent, never purple, sometimes reddish

..... 13

13a Follicles (3.5-) 4.5cm, glabrous, with apex attenuate into stigma

..... P. coriacea

13b Follicles 2.5-4cm, usually pubescent, with obtuse or rounded apex and sessile stigma.

14a Leaves glabrous beneath; leaflets 2-4cm wide, narrowly elliptical

..... P. broteroi

14b Leaves usually pubescent beneath; leaflets 5-10cm wide, elliptical

..... P. mascula


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