The Caucasian Representatives of the Genus Paeonia L.

L.M. Kemularia-Nathadse, Trudy Tiflis. Botan. Sada 1961

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Chapter III

Classification of the genus Paeonia L. by a table for defining section.

Dividing of the Genus Paeonia L. into sections has been firstly brought about by De Candolle (Prodr. 1), who distributed all known to him 16 species between two sections: Mountan DC. and Paeon DC. According to him, the first section comprised only one species P. Moutan Sims., which is a shrubby species with a peculiar, thin-walled staminodal outgrowth in form of a pitcher, which is covering a gynoecium nearly completely, and the second section included all other species presented by perennial peonies with a flat staminodal disk-shaped outgrowth: moreover, all the latter species have been divided by him into two groups in accordance with the absence or presence of pubescence upon the inferior side of leaf-blades. However. De Candolle did not ascribe any taxonomic purport to these groups.

In 1827. Reichenbach (Moesseler,.Handb, Gewächs, 2, 1 ) has declared the section Moutan DC. to be a self-dependent genus.

In 1834. Salm-Dyck (Hort-Dyck) has renamed the section Moutan DC, into the section Suffruticosae Salm-Dyck, and the section Paeon DC, has been divided by him into 6 very small sections, which have not been acknowledged by other botanists afterwards and which pertained to the groups of Stern or the serieses of V. Komarov in some or other way.

In 1893, Lingley (in Bot. Reg. 25, 30) has established a new section for the north-american peonies, giving it a name Sect. Onaepia Lindl.

In 1849, Seringe (in Fl. Jard. 3. 187) announced both section of De Candolle to be subgenuses, such as the subgenus Moutan (DC) Seringe and the subgenus Paeon (DC) Seringe.

In 1863, Baillon (in Adansonia 4, 56) has divided the section Paeon DC. into two self-dependent sections, one of which was called by him the sect. Eupaeonia Baillon, which included red-blossommed peonies with broad-leaf lobes, and the second section was called by him the sect. Tripaeonia Baillon, which comprised all other peonies.

In 1890, Lynch (in Journ. Roy. Hort. Soc. Lond. Ser. 12, 431, 433) added to the subgenuses Seringe one subgenus more that was named Onaepia (Lindl.) Lynch.

In 1892, Huth (in Engler. Bot. Jahrb. 14, 265), in his monograph, rejected all classifications by previous authors and offered his own one, based on the geographical principle, according to which all peonies growing in the Old World related to the sect. Palearcticeae Huth. The first section has been divided by him into two sections, as follows: the subsect. Fruticosae Huth comprising shrubby species and the subsect. Herbaceae Huth comprising perennial species.

The classification, proposed by Huth has not been acknowledged by botanists in future, excepting Schipczinsky (1921), and the most of them abided by the classification by De Candolle, as for example, N. A. Busch (1901) adhering to it in his study of Caucasian representatives of the genus Paeonia L.

In 1937, the academician Komarov ("Fl. USSR", v. VII) rejected the sectional division of the genus Paeonia L. ancTalFspecies growing within the boundaries of the USSR were arranged by him into the evolutional serieses, as follows: Ser. Chinensis Kom., Sre. Obovatae Kom., Corallinae Kom., Ser. Dentatae Kom. and Ser. Fissae Kom.

In 1946, in his monograph (in the Royal Horticulture Society), Stern proceeded to the division into three sections, such as the sect. Moutan DC., sect. Paeon DC, and sect. Onaepia Lindl. The first two sections have been divided by Stern into subsections, which in their turn have been divided into the groups of species, which are mostly i analogous with the evolutional series by Komarov. The section Moutan DC. is comprehended by Stern more widely than by De Candolle, as the former has included in it not only the shrubby species with a pitcher-shaped staminodal outgrowth, but also the perennial peonies with a lobed staminodal disk, excepting the North American species. Moreover, for the first species he has established the subsection Vaginatae Stern with the type P. suffruticosa Donn. and for the second species he has introduced the subsection Delavayanae Stern with the type P. Delavayi Franchet established by him.

Stern includes all peonies, which can not be related with any of the sections Moutan DC and Onaepia Lindl., in the section Paeon DC. Moreover, the section Paeon is divided by Stern into two subsections, such as the subsect. Foliolatae Stern, which comprises species with broad smooth-edged lobes (leaflets) of double and triple ternate leaves and with the type P. mascula Mill., and the subsect. Dissectifoliae Stern, which is characterized with narrow lance-shaped or linear lobes of double triple dissected or pinnately parted leaves or which lobes are deeply incised or toothed, and with the type P. officinalis L. emened Wilid chosen by him. The subsections introduced by Stern are divided by him into groups, in most cases equivalent to the serieses by Komarov. The subsections Foliolatae and Dissectifolia Stern are divided by Stern into 8 and 3 species groups, respectively. The monograph by Stern is to be regarded as newest and most complete nowadays. The system expounded by him and based on the classification of the De Candolle may be considered more advanced in comparison with all previous systems. Nevertheless, we can't accept it as a whole.

It is difficult to justify the expanding of the volume of the section Moutan DC., as there are no convincing arguments for joining the east-asian perennial species with it. So, De Candolle was quite right when he included in this section only the shrubby species with a peculiar pitcher-shaped staminodal disk, covering a gynoecium almost completely. Even if the east-asian perennial species, included by Stern in the above-mentioned section, have a clearly distinct lobed or toothed staminodal disk, nevertheless, it surrounds only the base of a gynoecium. Such a staminodal disk like this is availed by some species from the section Paeon DC, also. Therefore, we transfer the subsection Delavayanae Stern to the section Paeon DC, excluding from it the yellow-flowered species, which we include in our new section. As to the subsection Vaginatae Stern with its characteristic features, we suppose it to be a synonyme of the section Moutan DC. The latter section is very old and does not retain division into sections.

The section Paeon DC. is comprehended by us in somewhat other way than by De Candolle and all other subsequent authors. Diminishing its volume, we exclude from it all species with yellow and cream blossoms and almost all species with white blossoms, which petals get yellow after got dried out. This change of colour is explained by presence of the yellow pigment flavone in petals, which is the initial pigment of all pigments related with anthocyan (see See chapter "Morphology of peonies"). The above mentioned group of species, which includes yellow-flowered peonies from the subsection Delavayanae Stern, forms a natural group with a complex of characteristic morphological features and with a definite area of its distribution (Caucasus, the East Asia). We consider the above-mentioned group a self-dependant section of the genus Paeonia L., giving it a name defined by the pigment characteristic for all representatives included in this group.

Our section, just as the section Moutan, is very old. It exists in montane tertiary forests of Caucasus and the East Asia (China and Japan) till present.

From the section Paeon DC. we exclude the subsection Dissectifolia Stern and determine it a self-depent section with a name the section Sternia Kem.-Nath. in honour of the botanist Stern who has firstly found this group as a subsection. This section is divided into the evolutional serieses of species. As to the subsection Foliolatae Stern, we consider it a synonym of the section Paeon DC.

Some changes are introduced by us in the interpreting of Stern's groups volume and positions. First of all, from the section Moutan we transfer into the section Paeon DC. the Delavayanae group Stern, from which we exclude P.lutea Delavay, which suits to our new section. For this species we establish a new series named the series Luteae Kem. -Nath. and place it at the beginning of our section.

Besides, we consider the Officinalis group Stern, artificial and divide it into 3 series, as follows: series Officinalis Stern, series Hybridae Kem.-Nath. and series Fissea Kom.

So, for the genus Paeonia L. we accept 5 sections, such as the sect. Moutan DC., sect. Flavonia Kem.-Nath., sect. Onaepia Lindl., sect. Paeon DC. and sect. Sternia Kem.-Nath.

Species of every section are arranged by us in evolutional series , according to the conception of V. L. Komarov. Moreover, the "sect. Moutan DC. and sect. Onaepia Lindl. comprise only one evolutional series for each of them, which are the serieses Suffiuticosae Kem.-Nath. and Brownii Stern, respectively.

The representatives of the sect., Flavonia Kem.-Nath. are arranged, in 4 serieses as follows: the series Luteae Kem.-Nath., series Lactiflorae Stern, series Macrophyllae Kem.-Nath., ser. Wittmannianae Stern.

The sect. Paeon DC. is represented by 7 serieses, as follows: the ser. Delavayanae Stern, ser. Russii Stern, ser. Masculae Stern, ser. Obovatae Stern, ser. Mairei Stern, ser. Arietinae Stern, ser. Broteri Stern. The sect. Sternia Kem.-Nath. is consisted of 5 serieses, as follows: the ser. Officinalis Stern, ser. Peregrinae Stern, ser. Anomalae Stern, ser. Hybridae Kem.-Nath. and ser. Fissae Kom.

Our classification of the Genus Paeonia L. is graphically represented by the Scheme 1



Table for defining sections of the genus Paeonia L.

1. Shrubbery. The blossoms are very large, pinkish-yellowish, polypetalous. The staminodal, pitcher-urn-shaped disk is covering a gynoecium almost completely. The area of growing is the East Asia (China, Japan) ............. Sect. Moutan DC.

- Large perennials. The blossoms are less large, variating in colour. The staminodal disk with fleshy toothed edges surrounds the base of a gynoecium, or is practically unseen .. 2

2. Flowers yellow, whitish-yellowish, greenish-yellowish, in rare cases yellowish-pinkish, but always containing a yellow pigment. The leaves are with smooth-edged lobes. The plants grow in the montane forests of Caucasus, the North Iran and East Asia ........ Sect. Flavonia Kem.-Ndth.

- Blossoms purple, red, brown, pink or white, that means they always contain the pigment anthocyan, or are without it. The leaves lobes are variating in form, being smooth-edged, toothed, incised or dissected ......... 3

3. Leaves fleshy, lobes deeply and roundly incised. The blossoms are dark-purple-brownish. The plants grow in the South-West States of USA ......... Sect. Onaepia Lindley

- Leaves not fleshy, their lobes are broad and smooth-edged or incised, or cut up into narrow lance-shaped or linear lacinules. The blossoms are of different colour ......... 4

4. Leaves double-triple ternate, with broad smooth-edged lobes. The fruits are long, oblong-ovate, significantly spreading or pendent. The plants grow in montane forests of Caucasus, the Minor Asia, South Europe and Asia ......... Sect. Paeon DC.

- Leaves triternately pinnate, with pinnately incised or dissected into numerous narrow lance-shaped narrow-linear or even nearly filiform lacinule lobes. The fruits are short, broad-ovate or ovate-elliptical, straight or spreading. The plants grow in steppes and turned-into-steppes areas of Caucasus, the South and Middle Europe, Middle Asia, China, in rare cases in forests of Siberia ............. Sect. Stemia