Systematic treatment of the Genus Paeonia L. with some nomenclatoric changes.

Systematické tridení rodu Paeonia s nekterými nomenklatorickými zmenami

Josef J. Halda

Sedlonov 125, CZ-51782 Sedlonov, Ceská republika

Acta Mus. Richnov. Sect. natur. 4(2):25-32 (1997). ISBN 80-86076-07-5.

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

(index JJ Halda)

Abstract. An infrageneric classification of the genus Paeonia L. is presented. The genus is divided into 3 subgenera, 6 sections and 6 subsections. Key is given to the subgenera. Two new hybrids are described and many new combinations are made.

Keywords: Paeonia, system, taxonomy.

[Paeoniaceae Rudolphi, Syst. Orb. Veg. 61 (1830); Ranunculaceae Tribus V. Paeoniaceae DC. Prodr. 1:64 (1824)].

Key to the subgenera:

Paeonia L, Spec. Plant. 539 (1753).

Perennial herbs dying down in the winter or deciduous shrubs with erect tuberous stock and fleshy roots. Leaves large, simply biternate or further divided. Flowers 6-20cm across; petals white, yellow, pink or red. Stamens numerous. Carpels 2-8, free, glabrous or pubescent. Fruit a group of horizontally spreading follicles; seeds in two rows.

Subgenus Paeonia

[sect. Paeon DC., Prodr. 1:65 (1824); sect. Albiflorae Salm-Dyck, Hort. Dyck 365,366 (1834); sect. Compactae Salm-Dyck, Hort. Dyck 365,366 (1834); sect. Corallinae Salm-Dyck, Hort. Dyck 365,366 (1834); sect. Laciniatae Salm-Dyck, Hort. Dyck 365,366 (1834); sect. Lobatae Salm-Dyck, Hort. Dyck 365,366 (1834); sect. Macrocarpae Salm-Dyck, Hort. Dyck 365,366 (1834); sect. Microcarpae Salm-Dyck, Hort. Dyck 365,366 (1834); sect. Paeonia Reichenbach, Repert. Herb. s. Norn. Gen. PI. 191 (1841); sect. Eupaeonia Baillon, Adansonia 4:56 (1863); sect. Tripaeonia Baillon, Adansonia 4:56 (1863); sect. Palearcticae & Herbaceae Huth in Engler, Bot. Jahrb. 14:265 (1891); subgen. Paeonia prop. Baker in Gard. Chron. N. Ser. 21:779 (1884); subgen. Paeon (DC.) Seringe, H. Jard. 3:193 (1849)].

Type species: P. Officinalis L

Stems herbaceous, annual, dying down at end of each growing season. Petals thin, much larger than sepals; disc not produced as conspicuous outgrowths.

sect. Paeonia

[Paeonia subsect. Dissectifoliae Stern, A study of the genus Paeonia 1 (1946); Paeonia sect. Sternia Kem.- Nath., Trudy Tbil. Bot. Inst. 21:35 (1961)].

Leaves biternate or triternate; leaf-segments deeply cut to the lanceolate or lineate leaflets;, petioles grooveform with interrupted ring of sclerenchym. Typus: P. officinalis L

subsect. Paeonia

[Paeonia subsect. Dissectifoliae gr. Officinalis et gr. Peregrina Stern, A study of the genus Paeonia 1 (1946); Paeonia sect. Paeonia ser. Officinalis Kem. - Nath. ex Uspensk. in Bjull. Mosk. Obsch. Ispyt. Prir. otd. Biol. 92(3):84 (1987); Paeonia sect. Sternia ser. Officinalis et ser. Peregrina Kem. - Nath. in Trudy Tbil. Bot. Inst. 21:35 (1961)].

Leaf-segments dentate. Typus: P. Officinalis L

subsect. Anomalae (Kem. - Nath. ex Uspensk.) J. J. Halda stat. nov.

[Paeonia sect. Paeonia ser. Anomalae Kem. - Nath. ex Uspensk. in Bjull. Mosk. Ispyt. Prir. otd. Biol. 92(3):84 (1987); Paeonia sect. Sternia ser. Anomalae et ser. Hybridae Kem. - Nath. in Trudy Tbil. Bot. Inst. 21:35 (1961), nom. nud.; Paeonia subsect. Dissectifoliae gr. Anomala Stern, A study of the genus Paeonia 112 (1946), nom. nud.].

Leaf segments laciniate; leaflets lanceolate. Typus: P. anomala L.

subsect. Tenuifoliae (Stern ex Uspensk.) J. J. Halda stat. nov.

[Paeonia sect. Paeonia ser. Tenuifoliae Stern ex Uspensk. in Bjull. Mosk. Ispyt. Prir. otd. Biol. 92(3):84(1987); Paeonia subsect. Dissectifoliae gr. Tenuifolia et gr. Anomala Stern, A study of the genus Paeoma110(1946), nom. nud.; Paeonia ser. Fissae Komarov in Fl. SSSR 7:31 (1937), nom. inval.].

Leaf-segments pinnatisect. Typus: P. tenuifolia L.

sect. Palearcticae Huth emend. Uspensk. in Bjull. Mosk. Obsch. Ispyt. Prir. otd. Biol. 92(3):83 (1987).

[Paeonia sect. Palearcticae Herbaceae Huth in Bot. Jahrb. 14:256 (1891), p. p.; Paeonia sect. Paeonia DC., Prodr. 1:56 (1824), p. p.; Paeonia sect. Flavonia Kem. - Nath. in Trudy Tbil. Bot. Inst. 21:14 (1961), nom. nud,; Paeonia sect. Paeon Kem.- Nath. I. c. 21:14 (1961); Paeonia subsect. Foliolatae Stern, A study of the gen. Paeonia 1 (1946)].

Leaves biternatisect; leaf-segments entire, rarely bilaciniate or trilaciniate; petioles cylindric with continuous ring of sclerenchym. Typus: P. mascula Mill.

subsect. Flavonia (Kem.- Nath.) Uspensk. in Bjull. Mosk. Obsch. Ispyt. Prir. otd. Biol. 92(3):83.

[Paeonia sect. Flavonia Kem.-Nath. et ser. Macrophyllae Kem. - Nath. et ser. Wittmannianae Stern ex Kem. - Nath. in Trudy Tbil. Bot. Inst. 21:18,22 (1961); Paeonaser. Corallinae Komarov in Fl. SSSR 7:28 (1937); Paeonia subsect. Foliolatae group Wittmanniana Stern, A study of the genus Paeonia 53 (1946)].

Flowers yellow, creamy or whitish-yellow. Leaf-segments entire. Typus: P. wittmanniana Hartwiss ex Lindl.

subsect. Foliolatae Stern, A study of the genus Paeonia 1 (1946).

[Paeonia sect. Paeon ser. Broteri Kem. - Nath. in Trudy Tbil. Bot. Inst. 21:14 (1961), nom. nud.; Paeonia sect. Paeon ser. Arietinae Kem. - Nath. I. c. 21:14 (1961), nom. nud.; Paeonia sect. Paeon ser. Masculae Kem.- Nath. I. c. 21:14 (1961); Paeonasect. Paeon ser. Mairei Kem.- Nath. I. c. 21:14 (1961), nom. nud.; Paeonia sect. Russi Kem. - Nath. I. c. 21:18 (1961), nom. nud.; Paeonia sect. Palearcticae Huth. emend. Uspensk. subsect. Foliolatae Stern ser. Masculae F. C. Stern ex Uspensk. et ser. Arietinae Stern ex Uspensk. in Bjull. Mosk. Obch. Prir. otd. Biol. 92(3): 83 (1987); Paeonia gr. Mascula et gr. Coriaceae et gr. Russi et gr. Arietanae et gr. Broteri et gr. Mairei Stern, A study of the genus Paeonia 67-86 (1946); Paeonia ser. Masculae et ser. Arietinae Kem. - Nath. in Trudy Tbil. Bot. Inst. 21:28 (1961); Paeonia ser. Corallinae Komarov in Fl. SSSR 7:28 (1937); Paeona sect. Corallinae Salm-Dyck in Hort. Dyck. 365 (1834)].

Flowers red; leaf- segments entire or deeply cut to 2-3 leaflets. Typus: P. mascula Mill.

subsect. Obovatae Komarov ex Uspensk. in Bjull. Mosk. Obsch. Ispyt. Prir. otd. Biol. 92(3):84 (1987).

[Paeonia ser. Obovatae Komarov in H. SSSR 7:26 (1937), nom. inval.; Paeonia gr. Obovata Stern, A study of the genus Paeonia 21 (1946)].

Flowers rose or white; leaf-segments entire, orbiculate or oval. Typus: P. obovata Maxim.

sect. Albiflorae Salm - Dyck, Hort. Dyck. 365,366 (1834).

Leaf-edge cartilaginous, more or less dentate, gradually acute towards the apex. Typus: P. lactiflora Pallas.

Subgenus Onaepia (Lindl) Lynch in Journ. Roy. Hort. Soc., London, N. Ser. 12:431,433 (1890).

sect. Onaepia

[sect. Onaepia Lindl. in Bot. Reg. 25, subt.30 (1839); sect. Nearcticae Huth in Bot. Jahrb. 14:273 (1891)].

Type species: P. brownii Douglas ex Hooker.

Stems herbaceous, annual, dying down at end of each growing season. Petals coriaceous, not larger or only slightly larger than sepals; disc produced as fleshy lobes at the base of carpels.

Subgenus Moutan (DC.) Seringe, Fl. Jard. 3:187 (1849).

[section Moutan DC., Prodr. 1:65 (1824); sect. Sufrutiosae Salm-Dyck, Hort. Dyck, 365,366 (1834); sect. Moutania Reichenbach, Report. Herb. s. Nom. Gen. R. 191 (1841); sect. Palearcticae subsect. Fruticosae Huth in Engler, Bot. Jahrb. 14:272 (1891); sect. Mutan Ascherson & Graebner, Syn. Mitteleur. Fl. 5 (2):558 (1923); Moutan (DC.) Reichb. in Moessler, Handb. Gewächsk. Ed. 2 (1):55 (1827), pro gen.].

Typus: P. suffruticosa Andrews.

Stems woody, perennial; petals thin, much longer than sepals.

sect. Moutan

[sect. Moutan DC. subsect. Vaginatae Stern inA Study of the Genus Paeonia p.1 (1946); Paeonia sect. Moutan ser. Suffruticosae Kem. - Nath. in Trudy Tbil. Bot. Inst. 21:14 (1961), nom. inval.].

Disc produced as a thin leathery sheath which at first completely envelops the carpels; lower leaves bipinnate with some hairs on lower surface; pinnae entire or more often palmately 2-3-lobed.

Typus: P. suffruticosa Andrews.

sect. Delavayanae (F. C. Stern) J. J. Halda stat. nov.

[sect. Moutan DC. subsect. Delavayanae Stern in A Study of the Genus Paeonia p.1 (1946); Paeonia sect. Flavoniasef. Lutea Kem. - Nath. in Trudy Tbil. Bot. Inst. 21:14 (1961), nom. inval.; Paeonia sect. Paeon ser. Delavayanae Kem. - Nath. I. c. 21:14 (1961)].

Disc produced as conspicuous fleshy lobes around the base of carpels; lower leaves biternate, glabrous; leaflets pinnatipartite, with pinnae deeply lobed or toothed. Typus: P. delavayi Franch.

New combinations and descriptions:

P. mascula (L.) Miller subsp. kesrouanensis (J. Thiebaut) J. J. Halda stat. nov.

[bas: P. kesrouanensis J. Thiebaut in Mem. Inst. Egypte 31:37 (1936)].

Differs from subsp. mascula in having glabrous carpels and long stigma, coiled at the extreme apex. Syria, Lebanon, Turkey.

P. daurica Andrews subsp. lagodechiana (Kem.-Nath) J. J. Halda stat. nov.

[bas: P. lagodechiana Kem.- Nath. in Trudy Tbil. Bot. Inst. 21:33 (1961)].

Differs from subsp. daurica in having glaucous narrowly elliptic leaflets and broadly open pink flowers up to 150mm across. Gruzia: Kachetia.

P. tenuifolia L. subsp. biebersteiniana (Ruprecht) J. J. Halda stat. nov.

[bas: P. biebersteiniana Ruprecht, Flora Cauc. 47 (1869)].

Differs from subsp. tenuifolia in having leaflets much broader (up to 15mm wide). Caucasus: Stavropol.

P. anomala L subsp. hybrida (Pallas) J. J. Halda stat. nov.

[bas: P. hybrida Pallas, Fl. Ross. 1(2):94 (1788)].

Differs from subsp. anomala in having tomentose carpels. Finnish Lapponia, N. Russia, Altai, Turkestan.

P. veitchii Lynch subsp. altaica (K. M. Dai) J. J. Halda stat. nov.

[bas: P. altaica K. M. Dai & T. H. Ying in Bull. Bot. Res. 10(4):33 (1990)].

Differs from subsp. veitchii in having broader leaflets, bigger (100-180mm across) emarginate flowers and subglobose, smaller pollen grains. China: Xinjiang.

P. emodi Wall. ex Royle subsp. sterniana (H. R. Fletcher) J. J. Halda stat. nov.

[bas: P. sterniana H. R. Fletcher in Journ. Roy. Hort. Soc. Lond. 134:327 (1959)].

Differs from subsp. emodi in having smaller solitary flowers (up to 80mm across) and deeply lobed and toothed leaflets. SE Tibet: Kongbo.

P. wittmanniana Hartwiss ex Lindley subsp. macrophylla (Albow) J. J. Halda stat. nov.

[bas: P. corallina Retz. var. wittmanniana forma macrophylla N. Albow, Prodr. R. Colchid. 15 (1895)].

Differs from subsp. wittmanniana in having larger leaflets and glabrous carpels. Gruzia, Adzharia, W Caucasus.

P. obovata Maximowicz subsp. japonica (Makino) J. J. Halda stat. nov.

[bas: P. obovata Maximowicz var. japonica Makino In Bot. Mag. Tokyo 12:302 (1898)].

Differs from subsp. obovata in having glabrous leaves and white flowers with short stigmas and being diploid (2n=10). Sachalin, Japan.

P. suffruticosa Andr. subsp. rockii S. G. Haw & L. A. Lauener var. linyanshanii (T. Hong et G. L. Osti) J. J. Halda stat. nov.

[bas: P. rockii (S. G. Haw & L. A. Lauener) T. Hong et J. J. Li subsp. linyanshanii T. Hong et G. L Osti in Bull. Bot. Res. 14(3):237 (1994)].

Differs from var. rockii having folioles lanceolate or narrowly ovate, entire. Gansu, Hubei.

P. suffruticosa Andr. subsp. rockii S. G. Haw & L. A. Lauener var. yananensis (T. Hong et M. R. Li) J. J. Halda stat. nov.

[bas: P. yananensis T. Hong et M. R. Li in Bull. Bot. Res. 12(3):226 (1992)].

Differs from var. rockii having only few leaflets (up to 11); petals smaller, sometimes purplish and reddish purple stigma and disc. Shaanxi.

P. suffruticosa Andr. subsp. ostii (J. Hong et J. X. Zhang) J. J. Halda stat. nov.

[bas: P. ostii T. Hong et J. X. Zhang in Bull. Bot. Res. 12(3):223 (1992)].

Differs from subsp. rockii having narrowly ovate-lanceolate, entire leaflets, which are glabrous beneath; terminal leaflets 1-3 lobed. Petals mostly white, without purple basal blotch; filaments, discs and styles purplish red. Gansu, Henan, Hunan, Shaanxi.

P. suffruticosa Andr. subsp. spontanea (Rehder) S. G. Haw & L. A. Lauener var. jishanensis (T. Hong et W. Z. Zhao) J. J. Halda stat. nov.

[bas: P. jishanensis T. Hong et W. Z. Zhao in Bull. Bot. Res. 12(3):225 (1992)]. Differs from var. spontanea having white petals without petaloid stamens. Shanxi.

P. suffruticosa Andr. subsp. spontanea (Rehder) S. G. Haw & L. A.. Lauener var. qiui (Y. L. Pei et Hong) J. J. Halda stat. nov.

[bas: P. qiui Y. L. Pei et Hong in Acta Phytotaxonomica Sinica 33(1):91 (1995)].

Differs from var. spontanea having folioles ovate or ovate-rotundate, entire; the terminal one trifide or dentate. Hubei.

P. x francheti J. J. Halda hybr. nov.

[P. lutea Franch. x P. potaninii Komarov].

Descriptio: Frutex 0.8-1 metralis, trunco 5mm diam., cortice bruneo-griseo in lamellas satis tenues soluto; ramuli teretes, viridiscentes, basi squamis persistentibus obtectis. Folia alterna, chartacea, 120-180mm longa, biternata; folia lanceolata vel oblongo-lanceolata, basi cuneata, dissecta; petioli 120-160mm longi. Flos 60-70mm diametro; bracteae 5-7, virides; petala cuprea, orbiculata, emarginata; stamina numerosa, antheris flavis, linearibus; carpella 3-4, glabra, flavo-viridia, conica, stylis brevibus, stigmate piano, conico; discus lobulatus. Folliculi viridescentes, bracteis et sepalis persistentibus suffulti.

Holotypus: China austro-occidentalis, provincia Sechuan: montes Haba Shan supra Daju; inclivis dumetosis meridionalis cca 2600m supra mare.

Coll. J. J. Halda 20.6.1990 (JJH9006422). (PR).

P. x handel-mazzettii J. J. Halda hybr. nov.

[P. delavayi Franchet x P. lutea Franchet].

Descriptio: Frutex 1-1.5 metralis, trunco 8mm diam., cortice nigro-griseo in lamellass satis tenues soluto; ramuli teretes, viridiscentes, basi squamis persistentibus obtectis. Folia alterna, chartacea, 140-200mm longa, biternata; foliola lanceolata vel oblongo-lanceolata, basi cuneata, superne 3-lobata; petioli 150-200mm longi. Flos 80-90mm diametro: bracteae 5-6, virides; petala cuprea, obovata, crenato-marginata; stamina numerosa, antheris flavis, linearibus, filamentis albis, linearibus; carpella 3-4, flavo-viridia, conica, glabra, stylis brevibus, stigmate piano, recurvo; discus lobulatus. Folliculi viridescentes, bracteis et sepalis persistentibus suffulti.

Holotypus: China austro-occidentalis, provincia Yunnan: montes Haba Shan prope Daju; in clivis dumetosis meridionalis cca 3400m supra mare.

Coll. J. J. Halda 12.6.1990 (JJH9006315). (PR)

P. brownii Douglas ex Hooker subsp. californica (Nutt. ex Torrey & Gray) J.J. Halda stat. nov.

[bas: P.californica Nutt. ex Torrey & Gray , FI. N. Amer. 1:41 (1838); P.brownii Douglas ex Hooker var. californica (Nutt. ex Torrey & Gray) Lynch in Journ. Roy. Hort. Soc. 12:433 (1890)].

Differs from subsp. brownii in having leaves less divided and less lobed; leaf-segments and lobes acute at apex; petals 17-20mm long, as long as or slightly longer than the inner sepals, which are 15-19mm long. DISTRIBUTION: USA [Central and Southern California].

Bibliography:

Anderson, G. A. (1816): A monograph of the genus Paeonia in Trans. Linn. Soc. Lond. 1:1.

De Candolle, A. P. (1824): Prodromus systematis naturalis regni vegetabilis vol.1. Paris. Kemularija-Natadze, L. A. (1961): Kavkazskie predstaviteli roda Paeonia L. in Trudy Tbil. Bot. Inst. 21:3-52.

Pimenov, M. G. (1968): Sistematiceskaja gruppirovka vidov roda Paeonia L. SSSR po koefficientam schodstva. Bjull. Mosk. Isp. Prir., otd. biol. 73(1).

Salm-Dyck, J. (1834): Hortus Dyckensis. Paris.

Seringe, N. C. (1849): Flore des jardins et des grandes culturas ou description des plantes. Vol. 3. Paris.

Stern, F. C. (1946): A study of the genus Paeonia. The Royal Horticultural Society. London. Uspenskaja, M. S. (1987): Dopolnenie k sisteme roda Paeonia L. in: Bjull. Mosk. Isp. Prir. otd. biol. 92(3)79-85.